422 *♦ ^ 0Fc °. § C -M, t o ^r E s of y .** NOAA Technical Report NMFS Circular 422 A Revision of the Catsharks, Family Scyliorhinidae Stewart Springer April 1979 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and toestablish levels for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coasial waters, and the development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyzes, and publishes statistics on various phases of the industry. The NOAA Technical Report NMFS Circular series continues a series that has been in existence since 1941. The Circulars are technical publications of general interest intended to aid conservation and management. Publications that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management in- vestigations appear in the Circular series. NOAA Technical Report NMFS Circulars are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the marine sciences. Individual copies may be obtained (unless otherwise noted) from D825, Technical Information Division, Environmental Science Information Center, NOAA, Washington, D.C. 20235. Re- cent Circulars are: 365. Processing EASTROPAC STD data and the construction of ver- tical temperature and salinity sections by computer. By Forrest R. Miller and Kenneth A. Bliss. February 1972. iv + 17 p., 8 figs., 3 app. figs. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice, Washington. D.C. 20402. 366. Key to field identification of anadromous juvenile salmonids in the Pacific Northwest. By Robert J. MacConnell and George R. Snyder. January 1972. iv + 6 p., 4 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 367. Engineering economic model for fish protein concentration processes. Bv K. K Almenas, L. C. Durilla, R. C. Ernst, J. W. Gentry, M. B. Hale, and J. M. Marchello. October 1972, iii + 175 p., 6 figs., 6 tables. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 368. Cooperative Gulf of Mexico estuarine inventory and study, Florida: Phase I, area description. By J. Kneeland McNulty, William N. Lindall, Jr., and James E. Sykes. November 1972, vii + 126 p., 46 figs., 62 tables. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 369. Field guide to the anglefishes (Pomacanthidae) in the western Atlantic. By Henry A. Feddern. November 1972, iii + 10 p., 17 figs. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice. Washington, D.C. 20402. 370. Collecting and processing data on fish eggs and larvae in the California Current region. By David Kramer. Mary J. Kalin, Elizabeth G. Stevens, James R. Thrailkill, and James R. Zweifel. November 1972, iv + 38 p., 38 figs., 2 tables. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C. 20402. 371. Ocean fishery management: Discussion and research. By Adam A. Sokoloski (editor). (17 papers, 24 authors.) April 1973, vi + 173 p., 38 figs.. 32 tables, 7 app. tables. 372. Fishery publications, calendar year 1971: Lists and indexes. By Thomas A. Manar. October 1972. iv + 24 p.. 1 fcg. For sale by the Superintendent of Documents, U.F. Government Printing Office, Washington, D.C. 20402. 374. Marine flora and fauna of the northeastern United States. Annelida: Oligochaeta. By David G. Cook and Ralph O. Brinkhurst. May 1973, iii + 23 p., 82 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 375. New Polychaeta from Beaufort, with a key to all species recorded from North Carolina. By John H. Day. July 1973, xiii + 140 p., 18 figs., 1 table. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 376. Bottom-water temperatures on the continental shelf. Nova Scotia to New Jersey. By John B. Colton, Jr. and Ruth R. Stoddard. June 1973, iii + 55 p., 15 figs., 12 app. tables. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C. 20402. 377. Fishery publications, calendar year 1970: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. December 1972, iv + 34 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 378. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. By Arthur C. Borror. September 1973, iii + 62 p., 5 figs. For sale bv the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 379. Fishery publications, calendar year 1969: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. April 1973, iv + 31 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 380. Fishery publications, calendar year 1968: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. May 1973, iv + 24 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Of- fice. Washington, D.C. 20402 381. Fishery publications, calendar year 1967: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv + 22 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 382. Fishery publications, calendar year 1966: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. July 1973, iv + 19 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Of- fice, Washington, D.C. 20402. 383. Fishery publications, calendar year 1965: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. .July 1973, iv + 12 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 384. Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe. By Edwin T. Moul. September 1973, iii + 60 p., 109 figs. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, D.C. 20402. 385. Fisherv publications, calendar year 1972: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. November 1973, iv + 23 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice, Washington, D.C. 20402. 386. Marine flora and fauna of the northeastern United States. Pyc- nogonida. By Lawrence R. McCloskey. September 1973, iii + 12 p., 1 fig. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 387. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda. By Raymond B. Manning. February 1974, iii + 6 p., 10 figs. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, D.C. 20402. Continued on inside back cover r ^FNl Ctf NOAA Technical Report NMFS Circular 422 A Revision of the Catsharks, Family Scyliorhinidae Stewart Springer April 1979 U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, Secretary § National Oceanic and Atmospheric Administration Richard A. Frank, Administrator 5 Terry L. Leitzell, Assistant Administrator for Fisheries % National Marine Fisheries Service For Sale by the Superintendent of Documents, U.S. Government Printing Offic Washington, D.C. 20402 Stock No. O03-02O-OO147-S The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. CONTENTS Page Introduction 1 Materials and methods 2 Family Scyliorhinidae 3 Some characters used for the identification of genera and species 5 Key to genera of the Scyliorhinidae 9 Apristurus 11 Key to species of Apristurus 12 Apristurus atlanticus 14 Apristurus brunneus 15 Apristurus canutus 16 Apristurus herklotsi 18 Apristurus indicus 19 Apristurus investigatoris 20 Apristurus japonicus 20 Apristurus kampae 20 Apristurus laurussonii 21 Apristurus longicephalus 22 Apristurus macrorhynchus 22 Apristurus maderensis 23 Apristurus microps 23 Apristurus nasutus 24 Apristurus parvipinnis 25 Apristurus platyrhynchus 26 Apristurus prof undorum 27 Apristurus riveri 27 Apristurus saldanha 29 Apristurus sibogae 30 Apristurus verweyi 30 Asymbolus 30 Asymbolus analis 31 Atelomycterus 32 Atelomycterus macleayi 33 Atelomycterus marmoratus 35 Aulohalaelurus 35 Aulohalaelurus labiosus 36 Cephaloscyllium 37 Key to species of Cephaloscyllium 38 Cephaloscyllium fasciatum 38 Cephaloscyllium isabella 39 Cephaloscyllium laticeps 41 Cephaloscyllium sufflans 41 Cephaloscyllium ventriosum 42 Cephalurus 45 Cephalurus cephalus 45 Galeus 47 Key to adult Galeus 49 Galeus arae arae 51 Galeus arae antillensis 55 Galeus arae cadenati 58 Galeus boardmani 58 Galeus eastmani 59 Galeus melastomus melastomus 60 Galeus melastomus murinus 61 Galeus nipponensis 62 Galeus piperatus 64 Galeus polli 65 iii Galeus sauteri 66 Galeus schultzi 67 Halaelurus 68 Key to adult and half-grown Halaelurus 69 Halaelurus boesemani 70 Halaelurus buergeri 72 Halaelurus canescens 74 Halaelurus dawsoni 76 Halaelurus hispidus 77 Halaelurus lineatus 79 Halaelurus lutarius 82 Halaelurus natalensis 83 Halaelurus quagga 86 Haploblepharus 88 Key to adults of species of Haploblepharus 88 Haploblepharus edwardsii 88 Haploblepharus fuscus 89 Haploblepharus pictus 91 Holohalaelurus 92 Key to species of Holohalaelurus 92 Holohalaelurus punctatus 92 Holohalaelurus regani 93 Juncrus 95 Juncrus vincenti 96 Parapristurus 97 Parapristurus spongiceps 98 Parmaturus 99 Key to species of Parmaturus 99 Subgenus Dichichthys 100 Parmaturus campechiensis 100 Parmaturus melano bronchus 101 Subgenus Campagnoia 102 Parmaturus manis 102 Parmaturus stenseni 104 Subgenus Parmaturus 106 Parmaturus pilosus 106 Parmaturus xaniurus 108 Pentanchus 110 Pentanchus profundicolus 110 Poroderma 112 Key to Poroderma 112 Poroderma africanum 112 Poroderma marleyi 114 Poroderma pant her inum 114 Schroederichthys 115 Key to species of Schroederichthys 116 Schroederichthys biuius 116 Schroederichthys chilensis 118 Schroederichthys maculatus 120 Schroederichthys tenuis 122 Scyliorhinus 123 Key to species of Scyliorhinus 124 Scyliorhinus besnardi 126 Scyliorhinus boa 128 Scyliorhinus canicula 129 Scyliorhinus capensis 132 Scyliorhinus cervigoni 133 Scyliorhinus garmani 135 Scyliorhinus haeckelii 135 Scyliorhinus hesperius 137 iv Scyliorhinus meadi . . Scyliorhinus rotifer . . Scyliorhinus stellaris . Scyiiorhinus torazame Scyliorhinus torrei . . . Acknowledgments Literature cited 139 141 143 144 146 147 147 A Revision of the Catsharks, Family Scyliorhinidae STEWART SPRINGER 1 ABSTRACT The family includes 86 species in 17 genera. Species are characterized and most of them are illustrated. Summaries of biological studies pertinent to classification are included. Keys to genera and species are provided. Six new species and one new subspecies are described: Parmaturus manis of the western North Atlantic; P. stensenl of the Pacific continental slope off Panama; P. campechiensis from the Gulf of Mexico; Galeus schultzi from Philippine seas; G. arae antillensis from the West Indies; and two species of Apristurus from the western Atlantic, A. canutus Springer and Heemstra and A. parvipinnis Springer and Heemstra. Two populations close to Galeus arae and one population close to G. melastomus are treated here as sub- species; they are G. a. antillensis (new subspecies) and G. a. cadenati of the Caribbean region and G. m. murinus of northeastern Atlantic island slopes. Treatment emphasizes descriptive accounts thought to be useful for species identification. INTRODUCTION My interest in the small sharks of the family Scylio- rhinidae developed incidentally from exploratory fishing on the continental slopes and the deeper parts of the con- tinental shelves. Small sharks were found numerous enough to be ecologically significant in the rich and varied fauna of the upper slopes of the Gulf of Mexico. My first objective was to find names for deepwater sharks and attempts to identify and characterize species remained the core of my interest. Excepting a few species, the scyliorhinids are poorly known with only one or a few specimens of many species available in study collections. These collections will not be expanded easily because a majority of the species can be caught only with trawls or dredges such as are operated by scientific survey ships on continental slopes or in areas not exploited by fisheries. Although many scyliorhinids are poorly known, the number of nominal species has increased greatly in the past and probably will continue to grow. Muller and Henle (1841) recog- nized 12 species and Regan (1908b) 30 species in the group now comprising the family. Garman (1913) placed 37 species among 10 genera. Since then the number of nominal species has more than doubled and additional genera have been proposed. In spite of the lack of material, I believe that it is important now for me to organize the data I have gathered on the family so that others may refine and reconstruct parts or all of my revi- sion. For many species the inadequate data base has cur- tailed treatment, especially in some of the keys where one poorly known species is compared with another poorly known species with respect to a single character. The user of keys in this review is cautioned to be suspi- cious of oversimplification of key diagnoses. 'Systematics Laboratory, National Marine Fisheries Service, NOAA, U.S. National Museum, Washington, DC 20560; present address; Mote Marine Laboratory, Sarasota, Fla. I have recognized species and have included accounts of them if one character was found, either in the original description or in specimens examined, that would separate the species from all others. This brings an in- complete quality to this review that I could have cor- rected only by making subjective or arbitrary decisions to discard some names of populations that I found repre- sented by one or a few specimens. Thus I have deferred to future workers by assuming from little evidence that several species are valid because I had no positive reasons for rejection. In this way I have avoided burying some significant observations of earlier writers. For ex- ample, I recognize Apristurus maderensis, a species known from only one adult female. It is distinguished from all other Atlantic specimens that I have seen by having pointed prolongations of the gill covers. Changes in color pattern, in body proportions, and in denticle and tooth shapes accompany growth in many scyliorhinid species. Juveniles are identifiable usually to genus but not always to species. My descriptive ac- counts and diagnoses emphasize characteristics in adult specimens if adults were available to me for study. The most important weakness of my study probably comes from the small data base for most species which prevented uniform treatment of species, or, in extreme instances, prevented good comparison of species charac- teristics. The inconsistencies of treatment, however, are due not only to lack of material but also to the long period of data assembly. When I began to gather data on scyliorhinids more than 25 yr ago, I omitted recording many details that I found later would have been useful but which were difficult or impossible to recover. Furthermore, my earlier studies did not include either dissection of specimens or thorough review of anatomical works on sharks. For the serious student of elasmo- branchs, it is relevant that my interests are concen- trated in the area of field studies about distribution, behavior, and functional anatomy so that these interests have contributed variously some bias or some strength to this review. My own field work, including examination of freshly caught specimens, was carried out off the Atlantic and Pacific coasts of North America, especially in the Gulf of Mexico and Caribbean Sea, and off the east coast of Africa. All scyliorhinids are marine. Their fin shapes and fin positions suggest that they are weak swimmers and none are known to migrate over great distances. Most species are bottom-dwellers or near-bottom dwellers but a few, notably some species of Parmaturus, are part-time in- habitants of midwater (Lee 1969), far from the bottom. The majority of species live in the cool waters of the up- per continental slopes, and specimens of the most speciose genus, Apristurus, are taken typically in depths greater than 500 m. A few species of the Australian region and of Indian Ocean coasts, however, are known only from comparatively warm, shallow waters, and some of the better known forms, such as Scyliorhinus stellaris of the eastern North Atlantic, Cephaloscyllium uentriosum of the southern California coast, and Schroederichthys bivius of the Straits of Magellan region, frequent cool and shallow waters near shore. A few scyliorhinid species attain lengths of about 1.5 m but most are smaller. Cephaloscyllium is said to reach a length of 2.4 m (Graham 1956) in New Zealand waters, but the commonly recorded length for adult New Zealand Cephaloscyllium is 1 to 1.2 m. Scyliorhinids feed chiefly on invertebrates and small fishes. They exhibit little variety in tooth and jaw struc- tures that would suggest specialized feeding habits. Only a few species are taken by commercial fishermen and are of minor importance as food fish. Scyliorhinus canicula of the northeastern Atlantic and Mediterranean is an im- portant experimental animal. MATERIALS AND METHODS Representative specimens of 77 of the 86 scyliorhinid species recognized here were examined during the study but large series, including adults of both sexes as well as immature specimens, were available for only about 25 species. Species not seen were Apristurus atlanticus, A. inuestigatoris, A. maderensis, A. longicephalus, A. sal- danha, A. microps, A. kampae, Galeus boardmani, and Poroderma marleyi. Many of the specimens examined are designated by museum numbers. The museum or col- lections are abbreviated as follows: AMS AMNH ANSP BMNH DM FMNH HUMZ LACM Australian Museum, Sydney, Australia American Museum of Natural History, New York, N.Y. Academy of Natural Sciences, Philadelphia, Pa. British Museum (Natural History), London, England National Museum (Dominion Museum), Wel- lington, New Zealand. Field Museum of Natural History, Chicago, 111. Hokkaido University Museum of Zoology, Hakodate, Japan. Los Angeles County Museum of Natural His- tory, Los Angeles, Calif. MCZ Museum of Comparative Zoology (Harvard), Cambridge, Mass. NHMR Natural History Museum, Reykjavik, Iceland. ORI Oceanographic Research Institute, Durban, South Africa. SAM South African Museum, Cape Town, South Africa. SPIO Sao Paulo University Institute of Oceanography, Sao Paulo, Brazil. SU Stanford University Museum, Stanford, Calif, (fish collection now at California Academy of Sciences, San Francisco, Calif.). UA University of Arizona, Department of Biology, Tucson, Ariz. UBC University of British Columbia, Vancouver, British Columbia, Canada. UMMZ University of Michigan Museum of Zoology, Ann Arbor, Mich. UNCIMS University of North Carolina Institute of Marine Science, Morehead City, N.C. USNM U.S. National Museum, Washington, D.C. UTM University of Tokyo Museum, Tokyo, Japan. VFC Vanderbilt Foundation Collection, now at Cali- fornia Academy of Sciences, San Francisco, Calif. WAM Western Australian Museum, Perth, Western Australia. ZMB Zoologisches Museum an der Humbolt-Uni- versitat zu Berlin, East Berlin, Germany. ZMK Universitetets Zoologiske Museum, Copen- hagen, Denmark. ZMO Zoological Museum, Oslo, Norway. ZMUB Zoologisk Museum Universitetet I Bergen, Bergen, Norway. ZSI Zoological Survey of India (Indian Museum), Calcutta, India. Measurements of specimens recorded here were made with calipers or with a fish measuring board, and record- ed entries refer to dimensions taken in the manner exem- plified in Figures 1 and 2. Vertebral counts were made from radiographs. In this paper most of the measurements entered as length of shark or total length are unadjusted, projected distances from the tip of the snout to the tail tip with the shark placed in a straight position as near as possible. Longitudinal measurements given as tip of snout to points along the body are also projections, usually made on a V-shaped fishboard. Good measurements could not be made, however, if specimens were stiff and brittle and figures given for such measurements are estimates. All other measurements, such as fin base lengths, were made with calipers. The origin of any fin, except the upper caudal lobe, is here considered to be the midpoint of the curve beginning at the divergence of the fin from the body. The point of origin of the upper caudal lobe is obscure due to lack of elevation of the caudal axis so I have considered the upper fin lobe to originate at the beginning of the caudal crest, if the shark has one, and, if not, at the point where some elevation of the dorsal edge .. Length snout < Height of f~ first dorsal (fin) i Length inner margin second dorsal (fin) Figure 1. — Generalized outline of a scyliorhinid shark to illustrate terminology and methods of measurement. Nasal flap Least distance between nasal apertures Length mouth Figure 2.— Outline of ventral surface of head of a scyli- orhinid shark. of the fin becomes apparent. The eye length, and also the length of the orbit as used here, measures the horizontal length of the opening in the skin surrounding the eye, not the bony orbit. If I were to start again in the assembly of measure- ments of scyliorhinids, I would record additionally the greatest width of the head and the least distance across the top of the cranium between the bony orbits as well as the least distance across the top of the head between the upper eyelids. I would also be sure to record least dis- tances between bases of adjacent fins. At the beginning of each species account I have in- cluded a brief series of literature references. The first refers to the original species description in most in- stances but the rest were selected to list the more fre- quently used synonyms or were selected because the ac- count of the species or the accompanying illustrations seemed especially informative. I have omitted citing many sources either because I was unable to accept the species identification as correct or because the contained information could be found in other papers. FAMILY SCYLIORHINIDAE Family diagnosis. Small galeoid sharks, maximum length about 1.5 m (for possible exception see under Cephaloscy Ilium isabella); five gill slits, the 5th or 4th and 5th over the pectoral fin base; anal fin present; usually two dorsal fins (Pentanchus profundicolus has only one dorsal fin); dorsal fins without spines, either lobate or semilobate, the inner posterior tips sometimes sharply angled but not extended as a salient point; pos- terior end of base of first dorsal fin (or only dorsal fin) posterior to a vertical through the origin of the pelvic fin base; length of base of first dorsal less than length of caudal fin; caudal axis not elevated or little elevated; no keels or pits on caudal peduncle ; "nasal apertures of most species not connected to the mouth by channels crossing a depressed segment of the upper lip, but if channels pre- sent (as in a few species of the family), then no barbels extending from nasal flaps and no barbels elsewhere on head or body; pectoral fins usually short and broad, not falcate; eye openings longer than high, their anterior cor- ners usually close to a vertical through the anterior edge of the upper lip; a fold of skin below eye (secondary lower eyelid), which in many species forms the lower border of a denticle-lined gutter below the primary lower eyelid; species not connected to the mouth by channels crossing moderately or strongly arched, mouth opening nearly confluent with curved arc of jaws; teeth small, numerous, their arrangement alternate with several transverse series functional; teeth of upper and lower jaws not great- ly different in shape, always multicuspid in immature specimens and multicuspid in adults of most species but monocuspid in adult male Apristurus riueri and Schroe- derichthys bivius; dermal denticles of dorsal and lateral surfaces of adults typically imbricate with moderately broad blades on slender pedicles, the blades usually with ridges extending to three posteriorly directed points, the central point longest and strongest; lateral points and ridges often obsolete on denticles of ventral surfaces; denticles of juveniles typically narrower, more erect, sometimes spikelike; vertebral column moderately slen- der and weak, the transition from long monospondylous vertebrae to short diplospondylous vertebrae in pelvic region abrupt. The foregoing definition omits reference to most of the internal anatomical characters that are needed to place the family within a general classification of elasmo- branchs and to indicate relationships with other shark families. Relationships and evolutionary history. Major changes in classification of the higher categories of living elasmobranchs have been proposed several times. In the most recent and well-reasoned, that of Compagno (1973), the Scyliorhinidae fall within the class Chondrichthyes, subclass Elasmobranchii, cohort Euselachii, and super- order Galeomorphii. The superorder Galeomorphii is divided into four orders: Heterodontiformes, Orectolobi- formes, Lamniformes, and Carcharhiniformes, and the scyliorhinids constitute one of the eight families of the Carcharhiniformes. The characters of the greatest significance in Compagno's proposed classification are internal and an adequate discussion of them is beyond the scope of this account. Skeletal characters and relationships of some scylio- rhinid genera are discussed and illustrated in Nakaya's (1975) account of Japanese scyliorhinids. Many sharks can be placed in the appropriate family by reference to obvious external features. Some scylio- rhinids are similar in size, fin shape, fin position, and general appearance to the smaller orectolobids and without close examination might be confused with them. Differences in the head, and especially in the mouth, that can be seen without dissection distinguish the two families although important internal differences in the head cannot be seen without dissection. Some of the in- ternal differences between scyliorhinids and orecto- lobids are shown in diagrams in Compagno's (1973, fig. 4B, C) paper on interrelationships in living elasmo- branchs. The contrasting external appearances are also shown (Compagno 1973, plate 2C, D). Separation of scyliorhinids from orectolobids is not dif- ficult. The mouth of scyliorhinids is moderately or strongly arched with the mouth opening approximately superimposed on the line of occlusion of the upper and lower jaws, whereas the mouth of orectolobids is nearly transverse, not arched or little arched with the mouth opening not closely approximating the curve of the jaws. In most species of the Scyliorhinidae the upper lips do not have depressed areas forming channels between the nasal cavities and the mouth beneath enlarged nasal flaps, but in the few species with such a structure, barbels and barbellike structures are absent around the mouth or elsewhere on the head or body. Channels between the nasal cavities and the mouth, more or less protected or covered by structures of the lips, are present in all orectolobids and barbels are present either near the mouth or elsewhere on the head or body. The lips of scyliorhinid sharks are structurally simple, thin, and closely adherent to the jaws in contrast with the lips of orectolobids which are complex in form, relatively thick, and externally do not appear closely adherent to the jaws. In a synopsis of the Scyliorhinidae, Regan (1908a) noted the absence of oronasal grooves but did not describe the structures. In his revision of the Orecto- lobidae, Regan (1908b) pointed out that the oronasal grooves in sharks merely correspond in position to the embryonic oronasal grooves of amniote vertebrates, and that they are specialized structures which have arisen independently in different families and do not approxi- mate the condition in ancestral higher vertebrates. Nevertheless, Regan continued to use the term for those structures in the orectolobids and further confused ter- minology by stating that in three scyliorhinids, Scylio- rhinus canicula, S. edwardsii (= Haploblepharus edward- sii), and S. marmoratus ( = Atelomycterus marmoratus) , the nasal cavities are so near the mouth that the large nasal flaps overlie the upper lip and, hence, those species have no oronasal grooves. After Regan's series of papers on shark classification in 1908 and earlier, Smith and Radcliffe in Smith (1913) described Cirrhoscyllium ex- politum, a species that clearly belongs in the Orectolo- bidae but has no deep grooves between the nasal cavities and the mouth. In summary, it may be said that the channels connecting the nasal cavities with the mouth are not necessarily homologous with oronasal grooves of amniote vertebrates and even in sharks may have arisen independently through convergence in orectolobids and scyliorhinids. The relationships of scyliorhinids to those shark families placed by Compagno (1973) in the order Carcharhiniformes (Proscylliidae, Pseudotriakidae, Lep- tochariidae, Triakidae, Hemigaleidae, Carcharhinidae, and Sphyrnidae) are clear enough and seem to follow ap- proximately the traditional lines indicated by White (1937) and others and later diagrammed by Nakaya (1975, fig. 41). The relationships of scyliorhinids to galeoid sharks placed by Compagno in the orders Hetero- dontiformes and Lamniformes are not clear to me but at least seem to show no close affinities. Schaeffer (1967) summarized evidence from fossil studies on the broad aspects of elasmobranch evolution and concluded that the three major groups of modern elasmo- branchs — galeoids, squaloids, and batoids — belong to the same organizational level but that distinctive char- acters in each group pose unsolved problems of origin and mutual relationships. The earliest fossils sometimes tentatively assigned to the Scyliorhinidae (Glikman 1964; Romer 1966) appear in the Upper Jurassic, but whether any such fossils should be placed in the family as herein defined for extant species is open to question. The modern Scyliorhinidae, Proscylliidae, Pseudo- triakidae, Leptochariidae, Triakidae, Carcharhinidae, and Sphyrnidae form a series of families with basic simi- larities but increasing specialization. In a broad sense a step by step trend appears in the series: in reproduction, from oviparity as in most of the Scyliorhinidae to vivi- parity and the formation of pseudoplacentas in the more advanced of the other families; from small average species size in the Scyliorhinidae to medium size in the Triakidae to large size in some species of the Carcha- rhinidae and Sphyrnidae; from small, numerous multi- cuspid teeth in the young of the Scyliorhinidae to larger, less numerous, usually unicuspid teeth in some of the other families; from relatively few, long, and often poorly calcified vertebrae in some of the Scyliorhinidae to some- what greater numbers of shorter and more extensively calcified vertebrae in the other families; from unpro- tected eyes in some species of the Scyliorhinidae to eyes protected by a unique closing lower eyelid in the more specialized Carcharhinidae and Sphyrnidae; from flex- ible, usually small, and hydrodynamically inefficient, lobelike fins in the Scyliorhinidae to stiffer, larger, and hydrodynamically useful fins in the higher families; from a posteriorly placed and weak first dorsal fin suited only to eellike swimming in the Scyliohinidae to larger, stiffer, first dorsal fins that are located farther forward in the other families and provide a stabilizing plane against which the tail can push in the manner of a stern sculling oar. Reproduction. Many of the better-known species of the Scyliorhinidae are oviparous. A single large-yolked egg is enclosed within a leathery capsule by nidamental glands when it passes through the anterior part of either of the two oviducts. Typically an egg capsule is extruded after a very brief period within the oviduct and usually becomes attached to an object on the sea bottom by means of slender tendrils projecting from each of the four corners of the capsule. The extruded egg capsules of scyliorhinids have a distinctive shape (Fig. 3), the pos- terior half with a somewhat greater diameter then the anterior half. Although as recently as 1948 Bigelow and Schroeder (1948:196) stated that in scyliorhinids development is oviparous as far as known, it was later found that some species were ovoviviparous. Apparently it was Poll (1951) who first noted that not all scyliorhinids were oviparous. Later, Cadenat (1959) described the ovoviviparous form that Poll had seen as a new species, Galeus polli. I can confirm from my own observations the reports that Galeus polli is ovoviviparous and add that Cepha- lurus cephalus should be so characterized. It is possible that a few other scyliorhinids are ovoviviparous also since details of the modes of reproduction are known for less than half the scyliorhinid species. Examinations of female reproductive organs do not show obvious, gross differences between oviparous and ovoviviparous species. The egg capsules of ovoviviparous species are mem- branous and may be difficult to detect if, indeed, they are not absorbed when embryos are well developed. Kudo (1959) called attention to a variant form of oviparity in which several of the tough, leathery egg capsules are retained in oviducts until the embryos are partially developed. Nakaya (1975) called this mode of development in Halaelurus buergeri "multiple oviparity" to distinguish it from "single oviparity" in which one egg capsule at a time may occupy an oviduct and then for a brief period. Springer and D'Aubrey (1972) observed multiple oviparity in Halaelurus boese- mani Bass (1973), and Bass et al. (1975) found it in H. lineatus and H. natalensis. Probably multiple oviparity characterizes H. quagga, but H. lutarius and H. canescens both have one capsule at a time in each oviduct, and multiple oviparity is not yet known from other scyliorhinid genera. SOME CHARACTERS USED FOR THE IDENTIFICATION OF GENERA AND SPECIES Supraorbital shelf and eyes. In six scyliorhinid genera the cartilage of the chondrocranium extends out- ward above the orbits forming narrow shelves (Fig. 4A) that usually can be detected by touch without dis- section. In the 11 remaining genera the supraorbital shelf is absent (Fig. 4B). The anatomical term for the supra- orbital shelf is supraorbital crest. As a general rule the species with a supraorbital crest have a firmer body, thicker skin, and stronger (more extensively calcified) vertebral column than other species. These characters suggest the possibility of dividing the family into two subfamilies. Although the eyes of all scyliorhinids are dorsolateral, the species with a supraorbital shelf appear to have their dorsal vision somewhat restricted by presence of the shelf but may have a better lateral field of vision. The eyes in most scyliorhinids are proportionally large with the eye opening longer than high. Usually the only eye dimension practical to measure is the length of the opening because in preservation the lower eyelid may nearly close the eye, making the opening an elongate slit. A fold of skin, variously strong or weak in different species, sometimes not visible at all due to distortion of preservation, extends below the eye approximately parallel to, and separated from the lower eyelid by a Figure 3. — Egg capsules of Scyliorhinus stellaris (BMNH specimens from Dungeness, England). The larger capsule is 115 mm long, not including horns or tendrils; its greatest width is about 45 mm. The lateral margins of S. stellaris capsules are notably thick and strong. Figure 4. — Diagram of cross section of head through middle of eye (cartilage stippled). A. Scyliorhinus canicula, supraorbital shelf present. B. Galeus arae, supraorbital shelf absent. For detailed illus- trations of cranium and visceral skeleton of S. canicula see Parker (1878, plates 36-38). depressed channel sometimes called a subocular pouch but here called a subocular gutter. In the Scylio- rhinidae, the subocular gutter is lined with denticles, ex- cept in some species with sparse denticle arrangement or in some juveniles. The subocular gutter is less obvious in some species than others but comparisons are difficult because its prominence depends considerably on whether or not the eye is fully open. Nevertheless, some com- parisons are valid; for example, the subocular gutter is poorly or not at all developed in Apristurus but strongly developed in Atelomycterus. The lower eyelids of the scyliorhinids appear to be capable of partly or fully clos- ing the slitlike openings to protect the eyes and even in those species with the more strongly developed subocular gutters the fold of skin below the eye does not come in contact with the eyeball or become a true secondary lower eyelid. In the more advanced species of the Car- charhinidae protection for the eye is afforded by a partly or fully closing lower eyelid sometimes called the nic- titans (for discussion of structure and terminology of the carcharhinid eyelid see Gilbert and Oren 1964; Com- pagno 1970; Compagno and Springer 1971). Small spiracles, one located close to the posterior corner of each eye opening, are present in all scyliorhinids. In a few species, the spiracle lies within the posterior part of the subocular gutter which is as strongly developed poste- riorly as anteriorly. Mouth and labial furrows. In all scyliorhinids the mouth opening is moderately large and either broadly or steeply arched. The width of the mouth opening measured as the projected distance between mouth cor- ners is occasionally useful in identification. The presence or absence of labial furrows, whether or not the furrows are continuous around the mouth cor- ners, and their comparative lengths along the upper and lower jaws are traditionally among the most used char- acters for the identification of scyliorhinids. I have found the character reliable except in Halaelurus buergeri, in which labial furrows may be either very short or some- times absent at one or both mouth corners. Species of Scyliorhinus have an inconspicuous tab of the upper lip usually projecting slightly over the lower lip at each mouth corner. Its outer margin in some instances may look like a short upper labial furrow. Close examination will show, however, that it is not continuous with the lower labial furrow in Scyliorhinus. Snout, pore arrangement, and nasal flaps. The snout as defined in this study is that part of the head anterior to the upper lip, and its length is measured as a projec- tion from its tip to the edge of the upper lip. In scylio- rhinids this distance is nearly the same as the distance similarly measured in front of the anterior eye corners, but these distances may differ substantially in other families. Pores arranged in patterns on both the upper and lower sides of the snout are conspicuous and more or less characteristic of species in some scyliorhinids. The arrangements appear to be somewhat variable within a species and hence I have referred to pore arrangements in only a few species diagnoses. The nasal apertures of all sharks are partly divided into two openings by an internal septum extending from the midpoint of the nasal cavity wall. The anterior and more lateral opening is generally without any covering but the posterior and medial opening may be entirely hidden, partly covered, or not covered at all by a flap of skin that may extend from the anterior margin of the nasal aperture. In this study this structure is called the nasal flap. Regardless of its shape or function it is not here referred to as a cirrus or as a valve. In some scylio- rhinids, a small and inconspicious, denticle-free fold of membranous skin along the posterior margin of the pos- terior nasal opening may function, together with the nasal flap, as a valve to direct or reduce the outflow of water. Morphometries. I have avoided the use of propor- tional measurement differences for species diagnosis whenever alternative characters could be found because proportions may change during growth and because preservation may variously affect measurements of specimens. Bass (1973) proposed methods of description and analysis of proportional dimensions with mathe- matical expressions of growth patterns for eight scylio- rhinid species, the pattern for each species based on measurements of 14 to 102 specimens. His illustration (1973, fig. 9) of Holohalaelurus regani shows the remarkable difference in that species between a 129-mm juvenile and a 393-mm adult. Scyliorhinids, like other sharks, are subject to changes in proportions during growth. As pointed out by Garrick (1960) in a discussion of New Zealand Squalidae, the rate of growth in parts of the head or tail differs from the rate in midsections of the body. In some species the body cavity of females may in- crease disproportionally in length at sexual maturity. Proportional measurements are further complicated because specimens may either stretch or shrink in pre- servative. In soft-bodied scyliorhinids, the bulky mass of tissues of the trunk frequently shrink and pull the weak vertebral column out of normal alignment, often, but not always, producing an unnatural hump in the back. In the same specimen the vertebral column of the caudal sec- tion may be stiff and strong enough to prevent marked distortion. Dermal denticles. Two scyliorhinid genera, Galeus and Parmaturus, differ from all others in having a caudal crest of modified denticles, somewhat larger and different (Fig. 5) in shape from the denticles of the lateral surfaces of the tail and separated from them by a narrow strip of naked skin. The denticles of juveniles of most species are narrow, spikelike or needlelike, and are not especially close together. The replacement denticles that appear during growth in most species are progressively larger and wider, sometimes with three points (Fig. 6), and usually over- lapping. Teeth. Teeth of all scyliorhinid sharks are small and numerous with several transverse series functional. Upper and lower jaw teeth are similar in general shape. In all known juvenile scyliorhinids the teeth have three, five, or sometimes more, sharply pointed cusps. Typical- ly the teeth of the central part of the jaws are symmet- rical with the middle cusp much the longest, but teeth in Figure 5. — Caudal crest denticles of Galeus arae cadenati. Crest den tides of marginal rows are usually asymmetrical at all ages in Gale us but only in adults of some Parmaturus. Figure 6. — Dermal denticles of dorsolateral surfaces from a series of female Seyliorhinus retifer from the Gulf of Mexico. Left: from a 175-mm specimen. Center: from a 295-mm specimen. Right: from one 4(15 mm long. Camera lucida drawings to the scale indicated. the lateral parts of the jaws are progressively less sym- metrical and have shorter and more numerous cusps. Usually upper jaw teeth are slightly larger than lower jaw teeth but in adult male Aulohalaelurus (females and young not examined) the lower jaw teeth are substan- tially larger. Females and immature males of Apristurus riveri have multicuspid teeth typical of the family, but adult males have unicuspid teeth more than twice as large as the teeth of females of comparable size (Fig. 7). Other species of Apristurus in which both adult males and females are known do not exhibit such sexual dimor- phism. A transition from multicuspid teeth to unicuspid teeth at maturity also occurs in Schroederichthys bivius. Tooth dimorphism of small magnitude may be charac- teristic of some other scyliorhinids but has not yet been noted. The number of vertical rows of teeth (a row is defined here as a file of teeth crossing from the lingual to the labial side of the jaw approximately perpendicular to the jaw axis) remains constant during growth in many sharks, the more advanced carcharhinids for example. In those sharks the number of tooth rows may be charac- teristic of a species. I have not found evidence that the number of tooth rows changes with growth in scylio- rhinids but I have not adequately investigated that possi- bility. In some batoids the number of tooth rows may in- crease greatly during growth (Templeman 1965; Springer and Collette 1971). The teeth of scyliorhinids (Fig. 8) begin development in a germinal area on the lingual side of the jaws as in other modern sharks (Landholt 1947; James 1953), and, as new tooth buds form in the germinal area the devel- oping teeth move toward the labial side of the jaws and into functional positions as they become fully formed and hard. Finally the teeth move to the outer transverse Figure 7. — Camera lucida outlines of teeth of a 405-mm adult female Apristurus riveri (left) compared with same scale outlines of teeth of a 430-mm adult male (right), both specimens from 860 to 914 m depth off the Caribbean coast of Panama. series and are shed. Applegate (1965) found that in Odontaspis taurus (Odontaspididae) the size of the teeth was in approximate linear correlation with the size of the shark. The rate of tooth development and shedding in scyliorhinids has not been determined but evidence from species of other families (Ifft and Zinn 1948; Moss 1967) points to rapid replacement rates, especially for im- mature sharks. Tooth arrangement in the Scyliorhinidae, following Strasburg's (1963) classification, is chiefly in the alter- nate overlap pattern which precludes shedding teeth in bands as in some squaloids. The number of vertical tooth rows is often difficult to determine, not only because of M 1 trim D L d* V*>. l mm T i 1 mir. Figure 8. — Camera lucida drawings of typical flank denticles of some western Atlantic scyliorhinids: A. Scyliorhinus meadi, immature male, 26-1 mm; B. Apristurus profundorum, immature female, 380 mm; C. Scyliorhinus torrei, adult male, 243 mm; D. Scyliorhinus hesperius, imma- ture male. 2(l(> mm; E. Apristurus riveri, adult female, 405 mm; F. Schr oederichthys maculatus, adult female, 295 mm; G. Scyliorhinus retifer, adult male. 110 mm: H. Galeus arae, immature female, 2N7 mm. the small, crowded, and somewhat irregular ar- rangement of teeth but also because of the greater promi- nence of diagonal files of teeth in some species. The number of diagonal files is always substantially less than the number of rows perpendicular to the jaw axis. Thus, Saemundsson's (1922) count of tooth rows for the holotype of Scyllium laurussonii is less than mine because we used different definitions of a row. Because of the difficulties, I do not make substantial use of tooth counts for separating species. Vertebral numbers. As pointed out by Springer and Garrick (1964), vertebral numbers are important char- acters in systematic studies of carcharhinid sharks and might also be important in the study of other families. In their comprehensive survey these authors reported numbers of both precaudal and caudal vertebrae. In the Scyliorhinidae, however, the caudal axis is not elevated and no pits or other reference points conveniently es- tablish an exact point in separating precaudals from caudals. Furthermore, the terminal caudal vertebrae of some young scyliorhinids may not be visible on radio- graphs. I have, therefore, indicated the number of mono- spondylous vertebrae for all species that were available to me for X-ray examination. Total numbers of vertebrae are also given for some species. In scyliorhinid sharks the vertebral column has com- paratively long vertebrae in the trunk region with a dis- tinct and visibly abrupt transition to shorter vertebrae above the pelvic fins. The long vertebrae of the anterior region are here called monospondylous and the shorter vertebrae of the posterior region diplospondylous. I do not intend by the use of these terms to imply that I either noted or looked for morphological differences in the vertebrae other than length. Goodrich (1930) called the transition from monospondyly to diplospondyly in Scyl- lium (= Scyliorhinus) short but gradual. It is true that the last monospondylous vertebra is somewhat shorter than the preceding one, but the length decrease between the last monospondylous and the first diplospondylous is so marked that a precise point of change from one form to the other can be seen easily. KEY TO GENERA OF THE SCYLIORHINIDAE (In the text following this key the arrangement of genera is alphabetical.) la. Supraorbital crest present (Fig. 4A) 2 lb. Supraorbital crest absent (Fig. 4B) 7 2a. Labial furrows present, at least along lower jaw 3 2b. No labial furrows along either jaw C ephaloscy ilium - 5 species 3a. No upper labial furrows present, and upper labial furrows not continued around each mouth corner; upper lip usually with a small projecting tab which overlaps lower lip at each mouth corner 4 3b. Both upper and lower labial furrows present; labial furrows continuous around each mouth corner, no upper lip tabs 5 4a. Nasal flap barbels absent or represented only by swollen areas in the nasal flap; tips of these swollen areas sometimes projecting posteriorly past margins of nasal flaps for a distance not greater than the diameter of the spiracle Scyliorhinus - 13 species 4b. Nasal flap barbels present; longer than the diameter of the spiracle, and longer than the diam- eter of the bases of the barbels which extend from the margins of the nasal flaps Poroderma - 3 species 5a. Nasal flaps of various lengths but none reaching past edge of upper lip; no depressed areas of upper lip forming broad channels between nasal cavities and mouth 6 5b. Nasal flaps large and very long, their posterior tips reaching past edge of upper lip; upper lip with broad depressed areas forming broad channels between nasal cavities and mouth Atelomycterus - 2 species 6a. Nasal flaps large and long, their tips not quite reaching edge of upper lip; labial furrows very long, the upper extending about two-thirds the distance toward the upper jaw symphysis .... Aulohalaelums labiosus 6b. Nasal flaps moderately large, but their tips short of edge of upper lip; labial furrows short or moderately long, the upper extending less than half the distance toward the jaw symphysis . . Schroederichthys - 4 species 7a. Two dorsal fins present 8 7b. Only one dorsal fin present Pentanchus profundicolus 8a. Caudal crest of specialized denticles in three or many rows extending along the proximal half of upper edge of caudal fin, the crest denticles of some of them different in shape from denticles of the lateral surfaces of the caudal fin and separated from them by a narrow strip of naked skin 9 8b. No caudal crest of modified denticles 10 9a. Caudal crest nearly flat across its top, its marginal denticles strongly asymmetrical in young as well as in adults, small median denticles of crest usually in less than 5 rows; dorsal surfaces either marked with a variegated color pattern or dorsal surfaces darker than ventral surfaces; pectoral fins remarkably broad, their greatest widths measured straight across the fins near the convex distal margins usually greater than the width of the mouth Galeus - 12 species 9b. Caudal crest rounded across its top, its marginal denticles strongly asymmetrical only in adults or half-grown and symmetrical in juveniles, small median denticles of crest usually in more than five rows; dorsal surfaces not variegated and usually unmarked, color contrast between dorsal and ventral surfaces slight; pectoral fins not remarkably broad, their greatest widths usually less than or equal to the width of the mouth Parmaturus - 6 species 10a. Nasal flaps large, overlapping upper lip and hiding broad, shallow channels that connect nasal apertures with mouth Haploblepharus - 3 species 10b. Nasal flaps small or moderate in size, not overlapping upper lip; no depressed areas or channels across upper lip 11 11a. Labial furrows present along both upper and lower jaws and around mouth corners (except in some specimens of Halaelurus buergeri) 12 lib. No labial furrows Holohalaelurus - 2 species 12a. Adults tadpole-shaped as viewed from above, tapering from anterior part of head to origin of caudal fin; head length from tip of snout to fifth gill slit more than one-fourth total length . . Cephalurus cephalus 12b. Adults not tadpole-shaped as viewed from above, not tapering regularly from anterior part of head to origin of caudal fin; head length from tip of snout to fifth gill slit less than one-fourth total length 13 13a. Throat with transverse wrinkles formed by rows of dermal denticles alternating with vary narrow bands of naked skin Parapristurus spongiceps 13b. Throat without transverse wrinkles 14 14a. Species with moderately firm bodies and thick skins covered by well-hardened denticles; color usually not uniform, most species with a pattern of darker spots, bars, or dorsal saddle blotches; angle of view of eyes more lateral than dorsal; snout not notably long or spatulate 15 14b. Soft-bodied species with thin skins and weakly stiffened dermal denticles; color usually uniform with markings absent except for darker areas around mouth, gill slits, and fin margins; angle of view of eyes more dorsal than lateral; snout typically flattened, elongate, and spatulate with sharply angled edges Apristurus - 21 species 15a. Basal half or third of pelvic fins of adult males united, forming an apron between the claspers and the pelvic trunk 16 15b. Adult males without an apron Halaelurus - 9 species 10 16a. Claspers of adult males notably long and slender, extending for at least half their lengths past pelvic fin tips Juncrus vincenti 16b. Claspers of adult males not notably long and slender, extending less than half their lengths past pelvic fin tips Asymbolus analis APRISTURUS GARMAN Apristurus Garman 1913:96 (type-species, Scylliorhinus indicus Brauer, by original designation). Diagnosis. Species of Apristurus are somewhat compressed, soft-bodied scyliorhinids with two dorsal fins, much flattened heads, and long, spatulate snouts having conspicuous pores (ampullae of Lorenzini) above and below, the pores usually arranged in a pattern. The anterior and lateral edges of the snout are usually thin and sharp. The eyes are located dorsally so that the shark's angle of view is as much dorsal as lateral. Apristurus has prominent labial furrows that are continuous around the mouth corners. The nasal apertures are large and oblique and nasal flaps are reduced to narrow and usually short points that do not cover the posterior half of the nasal apertures and often do not extend across them. All Apristurus lack a supraorbital crest (shelf) of the chondrocranium; they lack depressions or channels in the upper lip connecting the nasal apertures with the mouth; they lack a crest of modified denticles on the proximal half of the up- per edge of the caudal fin; and they lack a series of transverse gular wrinkles that are formed by narrow bands of denticle- covered skin alternating with narrow bands of naked skin. Most Apristurus are black, dark brown, or dark gray without markings except darker areas around the mouth, gill slits, and on the leading edges of fins. All species lack dark saddle blotches and darker or lighter spots or bars. The dorsal and ventral surfaces are either uniform in color or the ventral surface may be slightly darker. One species, Apristurus sibogae, known only from the immature holotype, was described as reddish-white. The teeth of Apristurus species are small and numerous, usually similar in the upper and lower jaws, and with several transverse series functional. Symphyseal teeth are absent and considerable space may separate teeth on the right half from the left half of each jaw, but the amount of separation is variable and may be due only to the degree of relaxation of symphyseal ligaments. Typically the teeth are moderately flattened, multicuspid, the central cusp somewhat larger than the lateral cusps. The number of vertical rows of teeth is not easily counted and counts given in the literature may sometimes represent the number of diagonal rows instead of vertical rows. In one species, A. riveri, the teeth of adult males become enlarged and monocuspid through replacement (Fig. 7) but such sexual dimorphism has not been observ- ed in other species of Apristurus. The dermal denticles are small, usually with a three-pointed blade in adults. Young Apristurus may have slender, nee- dlelike denticles which may be replaced by broad-bladed denticles as the shark grows larger. The denticles are sometimes rather flexible and the degree of flexibility may determine whether the skin feels rough or velvety to touch. The soft denticle points of some specimens may be an artifact of preservation. The number of monospondylous vertebrae is comparatively small in Apristurus and these vertebrae are long, thin, and have weakly calcified centra. From X-ray examination it can be seen that vertebral centra are often broken, possibly during capture. The number of vertebrae in species of Apristurus shown in Table 1 may be useful in systematics as a character of secondary importance. For a discussion of vertebral numbers in the Scyliorhinidae see under Galeus. Discussion. Twenty-one species of Apristurus have been named including the two new species described here. Although some of the nominal species are of doubtful validity it seems probable that some remain undiscovered. Such a view is supported, for example, by three specimens collected by the RV Albatross in 1888 near the Galapagos Islands. Each specimen, USNM 135363, 133206, and 133207, was given a manuscript name many years ago but no description was published. My own examination of the specimens indicated that A. brunneus, A. nasutus, and A. kampae, the three species hitherto described from the eastern Pacific, were not represented, but more specimens would have to be available to give an adequate basis for description of a new species. Few museum specimens are available from many parts of the world. Some type-specimens are missing and some others are in very bad condition. To add to the difficulties of study, all of the species are undistinguished by prominent markings and outstanding structural differences. Members of the genus are all characterized by loose, thin skins and by weakly supported, soft bodies that preclude precise and reproduceable measurements, especially after long preservation. Apristurus specimens hitherto reported from African coasts include the holotype of A. atlanticus from the vicinity of the Canary Islands, the three type-specimens of A. indicus from the coast of Somalia and the Gulf of Aden, the holotypes of A. microps and A. saldanha from South Africa, three specimens from the vicinity of Saldanha Bay, South Africa, referred to A. saldanha by Norman (1935) and tentatively by Bass et al. (1975), and finally, one specimen from Southwest Africa reported by Karrer (1973) as Apristurus sp. Eleven additional specimens not yet in published reports were taken by research vessels of the U.S.S.R. (Georgio Golovan, P. P. Shirshov Institute of Oceanology, Acad. Sci. 11 Table 1.— Numbers of monospondylous vertebrae in species of Apristurus. Number of vertebrae ( monospondylous) 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 A. herklotsi A. riueri A. longicephalus 1 A. profundorum A. canutus A. kampae A. uerweyi A. sibogae A. nasutus A. platyrhynchus A. parvipinnis A. macrorhynchus A. laurussonii A. indicus A. brunneus A. japonicus 1 2 9 2 18 15 11 2 1 3 2 2 3 1 1 1 2 3 2 4 3 'Data from Nakaya (1975). U.S.S.R., Moscow, pers. commun.). As provisionally identified by Golovan, four fit A. profundorum, three A. microps, and the remaining four remain unidentified but are of great interest and may better establish the characters by which A. atlanticus and A. saldanha may be recognized (see discussions under those species). Apristurus species are not known from the Mediterranean and have not been taken from the eastern North Atlantic except peripherally from Iceland, Madeira, and Canary Island slopes. A large number of Apristurus specimen records of capture were available to me from exploratory vessels of the National Marine Fisheries Service (formerly Bureau of Commerical Fisheries). These indicate that for the Gulf of Mexico-Caribbean region, Apristurus is demersal and sparsely distributed on mud or soft-bottom localities at depths usually greater than 500 m. There, Apristurus was taken in the greatest numbers from 700 to 1,200 m but the deeper limits of distribution were poorly defined. Apristurus was not taken in midwater in the Gulf of Mexico-Caribbean area but Apristurus brunneus was taken off central California (J. A. F. Compagno, pers. commun.) at several hundred meters above the bottom. I suspect that the midwater occurrence is not really a rarity, especially in some localities, although most records suggest strictly demersal habitats. Publications by Chirichigno (1976) and Golovan (1976) which recorded Apristurus from the west coast of South America and from the west coast of Africa, respectively, were not seen until too late for adequate consideration for this paper. The well-developed ampullae of Lorenzini that are so conspicuous on the snout of species of Apristurus undoubtedly play an important role in the location of prey for these sharks. The ampullae of Lorenzini detect electrical fields generated by animals in their proximity. The ability of some sharks to locate hidden prey through electrosensitivity has been experimentally demonstrated for Scyliorhinus canicula (Kalmijn 1966, 1971). Such a means of locating food must be relatively more important for Apristurus because of its habitat in deep water where sunlight is nil and food is com- paratively scarce. I may speculate further that the spatulate snout packed with canals of the ampullae system is well adapted for food seeking in soft mud. Key to Species of Apristurus la. Gill covers without a median pointed lobe, or if present, the lobe not projecting for a distance as great as 2X spiracle diameter 2 lb. Gill covers with a median pointed lobe projecting posteriorly for a distance about equal to 2X spiracle diameter A. maderensis 2a. Anal fin long, its rear tip separated from origin of lower caudal lobe by a distance less than half the length of the eye opening 3 2b. Anal fin shorter, its rear tip separated from origin of lower caudal lobe by a distance greater than half the length of the eye opening A. kampae 12 3a. Distance between dorsal fin bases greater than length of base of second dorsal fin 4 3b. Distance between dorsal fin bases less than length of base of second dorsal fin A. microps 4a. Origin of first dorsal fin over or in advance of rear end of pelvic fin base 9 4b. Origin of first dorsal fin posterior to rear end of pelvic fin base 5 5a. Color black, brown, or gray, not white or reddish-white 6 5b. Color white or reddish-white A. sibogae 6a. Distance between dorsal fin bases less than distance from tip of snout to eye 7 6b. Distance between dorsal fin bases greater than distance from tip of snout to eye A. platyrhynchus 7a. Rear end of base of second dorsal fin in advance of rear end of base of anal fin 8 7b. Rear end of base of second dorsal fin over or behind rear end of base of anal fin A. verweyi 8a. Distance between pectoral and pelvic fin bases short, 6.0 to 8.5% total length (TL); anal fin base long, 18.0 to 22.0% TL; monospondylous vertebrae 33-36 A. canutus 8b. Distance between pectoral and pelvic fin bases long, 10.0 to 14.0% TL; anal fin base relatively shorter, 16.0 to 18.0% TL; monospondylous vertebrae 37-41 A. parvipinnis 9a. Eye not very large, its horizontal diameter less than 4.2% TL 10 9b. Eye very large, its horizontal diameter more than 4.2% TL A. atlanticus 10a. First dorsal fin much smaller than second dorsal fin, first dorsal area less than half area of second dorsal, length base first dorsal less than three-fourths length base second dorsal 11 10b. First dorsal fin equal to, or only a little smaller than second dorsal fin, first dorsal area more than half area of second dorsal, length base of first dorsal three-fourths or more than three- fourths length base of second dorsal 13 11a. Origin of first dorsal fin in advance of rear end of pelvic fin base 12 lib. Origin of first dorsal fin over rear end of pelvic fin base A. herklotsi 12a. Length of anal fin base less than distance from tip of snout to spiracle A. riveri 12b. Length of anal fin base greater than distance from tip of snout to spiracle A. investigatoris 13a. Interdorsal space not very long, distance between first and second dorsal fin bases less than distance from tip of snout to spiracle 14 13b. Interdorsal space very long, distance between first and second dorsal fin bases about equal to distance from tip of snout to spiracle A. saldanha 13 14a. Distance between bases of pectoral and pelvic fins less than distance from tip of snout to first gill slit 15 14b. Distance between bases of pectoral and pelvic fins greater than distance from tip of snout to first gill slit A.japonicus 15a. Snout (in front of mouth) short or only moderately long, its length 6.6 to 10.5% TL; abdominal trunk moderately long, distance from base of pectoral fin (pectoral axilla) to base of pelvic fin (pelvic fin origin) usually more than 8.0% TL 16 15b. Snout very long, its length about 12% TL; abdominal trunk short, distance from base of pectoral fin to base of pelvic fin about 7% TL A. longicephalus 16a. Distance between pectoral and pelvic fin bases greater than distance from tip of snout to spiracle 17 16b. Distance between pectoral and pelvic fin bases less than distance from tip of snout to spiracle A. laurussonii 17a. Snout comparatively short, its length in front of mouth 6.6 to 8.2% TL 18 17b. Snout moderately long, its length in front of mouth 8.5 to 10.4% TL 19 18a. Distance between dorsal fin bases usually about equal to length of snout in front of mouth A. brunneus 18b. Distance between dorsal fin bases usually greater than length of snout in front of mouth A. nasutus 19a. Length of base of anal fin long, 15. 7 to 19.1% TL 20 19b. Length of base of anal fin short, 13.9% TL A. profundorum 20a. Origin of first dorsal fin over middle of base of pelvic fins A. indicus 20b. Origin of first dorsal fin over posterior end of base of pelvic fins A. macrorhynchus Apristurus atlanticus (Kofoed) Scylliorhinus atlanticus Kofoed 1927:18, pi. IV, No. 3 (from vicinity of eastern Canary Islands). Not Apristurus atlanticus: Bigelow et al. 1953:217, fig. 1. Material examined. Not seen, Kofoed's holotype, ZMO, is an immature male about 250 mm long, taken from 1,365 m at Michael Sars station no. 41,235 at lat. 28°08'N, long. 13°35'W, yellow mud bottom. Although I did not see the type, Alwyne Wheeler (in correspondence) gave me his recent measurements of the specimen. Western Atlantic specimens referred to this species by Bigelow and Schroeder (1948) and by Bigelow et al. (1953), are here regarded as specimens of Apristurus laurussonii. Diagnosis. Kofoed's measurement of eye length, 4.9% TL, indicates a proportionally larger eye than in any other species of Apristurus. His measurements show that the base of the anal fin, 14.4% TL, is slightly greater in proportion than that of the holotype of A. profundorum, 13.9% TL, and slightly less than that of two of the types of A. indicus, 15.7 and 16.2% TL. Kofoed's figure for the holotype of A. atlanticus indicates that the two dorsal fins are nearly equal in size, the first dor- sal origin over the posterior end of the pelvic fin base; length of snout in front of mouth about equal to the distance 14 between the dorsal fin bases and to the distance from the pectoral axilla to the pelvic fin origin; posterior tip of anal fin reaches origin of lower caudal fin and extends only a little posterior to the second dorsal fin base. Discussion. Bigelow and Schroeder (1944:21-22) called attention to the strongly curved (concave) anterior edges of the gill slits in their account of Apristurus and noted that the curved gill slit margins were illustrated by Kofoed (1927, pi. 4) from the type of atlanticus. Bigelow and Schroeder believed that the genus might be divided on the basis of this character, with atlanticus, riveri, and others having curved gill slits and brunneus and other species having more or less straight ones. Examination of larger series, however, shows the character to be variable and unreliable. The larger specimens of some species often have gill slit edges straight or nearly so and young specimens have curved gill slit edges. The absence of denticles around the gill openings makes an oval patch of bare black skin. It appears to be frequent in im- mature examples but infrequent in adults of several species. As with several species of Apristurus known from single specimens, the case for recognition of A. atlanticus is not strong but it is also not clear that it belongs in synonymy. Eastern Atlantic specimens of Apristurus are few in com- parison with numbers taken in the western North Atlantic but whether this reflects a real scarcity or is due to differences in collecting effort is not clear. Apristurus brunneus (Gilbert) Figure 9 Figure 9. — Apristurus brunneus, 330-mm immature female from coast of Washington. Drawing by Mary Wagner. Catulus brunneus Gilbert 1892 (from off La Jolla, Calif.). Apristurus brunneus: Garman 1913:99; DeLacy and Chapman 1935:63; Roedel and Ripley 1950:49, fig. 33; Beebe and Tee-Van 1941:99; Clemens and Wilby 1961:78; Cox 1963:281, fig. 9. Material examined. USNM 51708, holotype, adult (ad.) $, 50 cm Albatross stn. 2396, lat. 32°49'N, long. 117°29'W, in 556 m; ad. S, 655, 605 mm; imm. 3, 545 mm; ?, 567, 545, 355, 352, 265 mm, coast of Washington in 730-790 m; young S 65 mm (probably taken from egg capsule before hatching), Santa Barbara Channel, Calif. Diagnosis. In A. brunneus the area of the first dorsal fin is usually somewhat less than the area of the second dorsal although the size difference between the two fins is not great and is somewhat variable; the distance between dorsal fin bases is about equal to the snout length; the anal fin is long, the length of its base about equal to the distance from the snout tip to the spiracle, the anal fin tip reaching past the origin of the lower caudal lobe. Apristurus brunneus is brown in life and fades somewhat in preservative. It is important to note for purposes of identification that A. brunneus, like all other Apristurus, lacks a caudal crest. The lack of a caudal crest distinguishes it from Parmaturus xaniurus, which may be found in the southern part of its range. Two other Apristurus have thus far been described from the eastern Pacific. Apristurus kampae from the Gulf of California, here regarded as known only from the holotype, has a much shorter anal fin than A. brunneus, its posterior tip not reaching the lower lobe of the caudal fin. Apristurus nasutus, described from specimens taken off the coast of Chile, has a slightly shorter snout and a longer interspace between dorsal fin bases than A. brunneus from the coasts of Washington and British Columbia. The southward range of A. brunneus from southern California along the coast of Central America to Panama is suggested by some recent collections made in Panama Bay (C. E. Dawson, pers. com- mun.) which indicate the need to reexamine the status of A. nasutus (see discussion under that species). 15 Description. The following proportional measurements given as percentages of total length show something of the extent of variation with size in proportions. In each group of proportional measurements, the first is from a 655-mm adult male, the second from a 567-mm subadult female, the third from a 265-mm young female, and the last from a 65- mm hatchling male. Tip of snout to: front of mouth, 6.0, 9.0, 9.4, 6.6; eye, 7.3, 9.7, 10.1, 6.5; first gill slit, 17.4, 19.8, 20.8, 20.6; origin pectoral fin, 21.7, 23.3, 26.4, 23.3; origin first dorsal fin, 51.1, 52.9, 46.0, 47.7; origin pelvic fins, 44.6, 45.9, 45.3, 45.3; origin anal fin, 53.4, 58.6, 52.8, 51.0; origin upper caudal fin lobe, 69.0, 70.5, 67.9, 72.4. Orbit: horizontal diameter, 2.7, 3.5, 3.0, 4.0. Mouth: width, 6.7, 7.9, 9.1, 9.9; length, 3.7, 4.2, 3.8, 5.4; length lower labial furrow, 2.6, 3.2, 2.6, 2.0; length upper labial furrow, 3.1, 3.4, 2.6, 2.5 Nostrils: least distance between (= internasal), 2.4, 3.2, 3.4, 3.5. Gill slits: height of first, 1.5, 2.1, 2.6, 2.6; height of fifth, 1.7, 1.6, 1.5, 1.7. Length of fin bases: first dorsal, 5.8, 6.0, 6.8, 7.7; second dorsal, 6.7, 7.4, 7.5, 7.7; anal, 13.7, 12.0, 13.6, 13.9. Distance between fin bases: first and second dorsal fins, 7.6, 7.1, 6.4, 3.9. Anterior margin of fins: pectoral, 10.7, 10.4, 9.8, 10.5; first dorsal, 10.2, 7.9, 10.6, 9.2; second dorsal, 8.1, 10.2, 11.3, 9.2; anal, 7.2, 9.2, 6.4, 7.2. The teeth are mostly five-cusped, the middle cusp much the largest, upper and lower teeth are similar except that the lower jaw teeth may be somewhat more slender and less erect. Upper teeth are generally in 29 to 31 vertical rows, lower teeth in 30 to 34 rows. Dermal denticles are relatively small and densely distributed, nearly erect, with three posterior points. Monospondylous vertebrae were 39 to 41 in five specimens. Discussion. Apristurus brunneus is not uncommon at depths from 137 to 347 m off British Columbia and occurs as far north as the Strait of Georgia (Clemens and Wilby 1961). I saw moderately large numbers taken in some trawl catches at depths from 455 to 790 m off the coast of Washington. Apristurus brunneus has been taken in small numbers off Oregon and California, but has not been collected south of the Mexican border. The locality for the holotype was neither the Santa Barbara Islands nor the Gulf of California as suggested by the title of the paper in which Gilbert (1892) described A. brunneus (see Beebe and Tee- Van 1941:99). Apristurus brunneus egg capsules were described by DeLacy and Chapman (1935) from four capsules, two taken from a 452-mm female caught in about 82 m in Washington coastal waters and two trawled from the same area. One capsule was attached by its tendrils to the stalk of a sea whip. The authors also gave proportional measurements for 22 specimens 368 to 500 mm long taken from Saratoga Passage, Wash. The egg capsule of A. brunneus is the smallest one produced by California scyliorhinids (Cox 1963). The capsules are usually described as brown, translucent or transparent, a little more than 50 mm long and a little less than 25 mm wide, slightly constricted at one end. Apristurus canutus Springer and Heemstra, n. sp. Figures 10, 11, 12, 13; Table 1 Figure 10.— Apristurus canutus, 428-mm adult male paratype, Leeward Islands. Wrinkles of skin that characterize most Apristurus specimens preserved in alcohol not shown. Drawing by Mildred Carrington. 16 Figure 12. — Dermal denticles from flank of Apristurus canutus, den- ticles about 0.5 mm long. Surface reflections partly obscure shingle- like structure of the denticle blades. Figure 1 1 . — Ventral view of head of 455 mm female holotype of Apris- turus canutus to show pattern of prominent pores. Figure 13. — Photograph of typical alcohol-preserved specimen of Apristurus (A. canutus) showing wrinkles usually following preservation of scyliorhinids with loose skins (Apristurus, Parmaturus, Cephalurus, Parapristurus, and Pentanchus). Holotype: USNM 206176, ad. ?, 455 mm, RV Oregon II stn. 10834, Leeward Islands near Anguilla, lat. 18°18'N, long. 63°23"W, in 687 m. Paratypes: USNM 206180, ad. Islands near Antigua, lat. 16°53'N, ' 428, 395 mm, imm. S, 388 mm, ad. long. 61°53'W, in 750-840 m. 395 mm, RV Oregon I stn. 6703, Leeward Diagnosis. Second dorsal fin 2 to 3 times larger in area than first dorsal; origin of first dorsal at vertical between posterior end of pelvic fin bases and origin of anal fin; eye rather large, length of orbit (eye opening) about 2.9 to 3.5% TL, about equal to nostril length and three-fourths of least distance between nostrils; eye opening length contained 2.9 to 3.5 times in length of snout to front of mouth (edge of upper lip), 6.5 to 8 times in length of head to fifth gill opening; snout length 8.6 to 10.0% TL; distance between dorsal fin bases less than distance from tip of snout to level of anterior ends of orbits; width of snout at anterior ends of nostrils considerably greater than width of mouth; anal fin base long, 18 to 22% TL; distance between pectoral fin base and origin of pelvic fin small, 6.0 to 8.5% TL; monospondylous vertebrae 33 to 36. Apristurus canutus belongs with the group of species A. riveri and A. parvipinnis in the Atlantic, having the second dorsal fin twice as large as the first dorsal. It differs from A. riveri in having the origin of the first dorsal fin posterior to the rear end of the pelvic fin base. The distance between the pectoral fin base and the pelvic fin base in A. canutus is 7.2 to 8.5% TL but in A. parvipinnis is 11.3 to 13.7% TL. The length of the anal fin base is 18.0 to 21.6% TL in A. canutus but in A. parvipinnis it is 16.4 to 17.9% TL. 17 Description. Proportional measurements as percentages of the total length are given below for the 455-mm holotype with the range among the type series, holotype, and five paratypes, 319 to 455 mm, in parentheses. Tip of snout to: front of mouth, 9.9 (8.6-9.9); eye, 11.0 (9.7-11.0); spiracle, 15.0 (13.9-15.0); first gill slit, 18.7 (17.7- 18.7); fifth gill slit, 22.9 (21.8-22.9); origin pectoral fin, 21.1 (21.1-21.9); origin first dorsal fin, 49.1 (47.6-50.3); origin pelvic fins, 38.5 (36.0-38.5); origin second dorsal fin, 61.6 (61.6-63.5); origin upper caudal fin, 71.9 (69.6-71.9); anterior end cloacal opening, 43.3 (40.5-43.3). Greatest width of trunk: at pectoral origin, 10.6 (7.7-11.1); at origin pelvics, 7.3 (4.4-9.6); at origin caudal fin, 1.8 (1.8-1.9). Greatest depth of trunk: at pectoral origin, 8.1 (7.8-9.3); at origin pelvics, 10.1 (6.1-10.1); at origin caudal fin, 4.6 (2.0-4.6). Orbit: horizontal diameter of opening, 3.3 (3.3-3.8); vertical diameter of opening, 1.1 (1.0-1.3). Spiracles: greatest diameter, 0.4 (0.4-0.6); distance from eye, 0.7 (0.5-0.8). Mouth: width, 6.4 (6.4-7.3); length, 3.1 (3.1-4.4); length upper labial furrow, 3.1 (3.1-3.5); length lower labial furrow, 2.0 (1.8-2.1). Nostrils: least distance between, 4.4 (3.7-4.4). Gill slits: height of first, 1.1 (1.0-1.8); height of fifth, 1.9 (1.0-1.9). First dorsal fin: length base, 3.7 (3.2-3.8); length posterior tip, (free inner margin), 2.4 (2.0-2.8); height, 1.5 (1.5- 2.6); length anterior margin, 6.4 (5.4-6.4). Second dorsal fin: length base, 5.9 (5.0-6.1); length posterior tip, 2.4 (2.3-3.4); height, 2.4 (2.4-2.8); length anterior margin, 8.8 (7.5-9.3). Anal fin: length base, 19.1 (18.0-21.6); length posterior tip, 0.9 (0.5-0.9); height, 3.7 (2.5-3.7); length anterior margin, 8.8 (7.8-9.4). Pectoral fins: width base, 8.8 (7.2-8.8); anterior margin, 14.1 (11.2-14.1); greatest width, 10.3 (7.8-10.3). Pelvic fins: overall length, origin to posterior tip, 12.3 (9.1-12.3). Caudal fin: upper margin, 27.7; anterior margin lower lobe, 9.0 (8.9-9.8). Distance between fin bases: first and second dorsal fins, 8.4 (7.7-9.8); pectoral and pelvic, 7.3 (7.2-8.5); pelvic and anal, 3.1 (3.1-4.2); anal to lower caudal, 1.3 (0.0-1.3); second dorsal to upper caudal, 4.8 (4.3-5.8). Body moderately compressed; head and snout flattened; snout long with prominent pattern of pores medially on the lower side of the snout (Fig. 11), maxillary pores prominent; nasal apertures large, the upper nasal flap pointed, reaching across nostril; labial grooves prominent along both upper and lower jaws, upper grooves somewhat longer than lower, length of upper about half as great as distance from corner of mouth to symphysis of upper jaw; eyes large, angle of view as much dorsal as lateral. Fins moderately large; their distal margins generally frayed, not notably rounded; base of anal more than 2 times base of second dorsal; tip of appressed pectoral reaching or nearly reaching origin of pelvics. Teeth generally with three to five cusps, in more than 50 vertical rows in both jaws; no difference in tooth shapes in males and females. Dermal denticles of flanks, three-pointed, with a weak central ridge, surface of blades sculptured (Fig. 12). Type series with 33 to 36 monospondylous vertebrae; holotype with 34 monospondylous vertebrae, 74 precaudal, and about 55 caudal vertebrae. Color dark gray with minute white spots underneath denticles, ventral surfaces slightly darker than dorsal surfaces; leading edges of fins, except pelvics and lower caudal, narrowly edged with black; narrow, denticle-free areas around lips and gill slits black; inside of mouth black. Apristurus herklotsi (Fowler) Pentanchus herklotsi Fowler 1934:238, fig. 3 (southern Philippines near Jolo). Material examined. USNM 93134, holotype, v, 312 mm, RV Albatross stn. 5424. Diagnosis. Apristurus herklotsi is one of the species having the first dorsal fin much smaller than the second dorsal; the origin of the first dorsal is about over the rear end of the anal fin base. The caudal fin is long, its length from origin of the lower caudal lobe more than one-third the total length. The eye is comparatively small, its length (length of opening) less than the least distance between the nasal apertures. The anal fin is long, the length of its base much greater than the distance between dorsal fin bases and about equal to the distance from the tip of the snout to the first gill slit. In com- bination, the preceding characters set off herklotsi reasonably well from all other species of Apristurus. The holotype of A. herklotsi has 28 monospondylous vertebrae, fewer than any other specimen of Apristurus examined for this study (Table 1). 18 Discussion. The holotype of herklotsi is not now in good condition, especially in the head region, and measurements cannot be made accurately for some of the structures. In my opinion, however, the original description and illustration are consistent with the type specimen and appear reliable in all details. I do not follow Fowler's (1934) emphasis on the presence or absence of nasal cirri as useful characters for the separation of Apristurus species. The structures are sometimes very small and easily damaged or distorted in preservation. Furthermore, I suspect they are variably developed within species. Apristurus indicus (Brauer) Table 1 Scylliorhinus indicus Brauer 1906:8, pi. 14, fig. 1 (Indian Ocean off Somalia and Gulf of Aden). Pentanchus indicus: Fowler 1941:60. Apristurus indicus: Garman 1913:97; Bigelow and Schroeder 1948:221. Material examined. ZMB 22424, 336-mm, $, here designated the lectotype, Valdivia station 259, off coast of Somalia, lat. 02°59'N, long. 47°06'E, 1,289 m; ZMB 17411, 325-mm S, paralectotype, same station as lectotype. Brauer described the species from three specimens but the smallest, 132 mm, from the Gulf of Aden in 1,840 m probably was not preserved (Christine Karrer, pers. comraun.). Diagnosis. Second dorsal fin somewhat larger than the first dorsal fin in area; origin of first dorsal over middle of the pelvic fin bases; eye small, length of opening 2.4 to 2.8% TL, two-thirds nostril length, one-half to two-thirds internarial distance, contained about 3.8 to 4.2 times in length of snout; length of snout to front of mouth about 9.5 to 10.5% TL; dis- tance between dorsal fin bases less than distance from tip of snout to eye opening; snout width at anterior ends of nostrils greater than width of mouth; teeth numerous, in about 60 to 62 vertical rows in upper jaw, most teeth with five cusps; color brown-black or black. Description. The following descriptive account and proportional measurements are based solely on the two extant syntypes. The lectotype, a female, ZMB 22424, now measures about 345 mm TL and the smaller paralectotype, also a female ZMB 17411, 327 mm. The following measurements are in percentages of total length, the first figure in each pair is for the lectotype. Dashes indicate measurements were not made. Tip of snout to: front of mouth, 10.1, 9.2; eye, 11.6, 10.4; spiracle, 15.7, 14.0; first gill slit, 20.9, 20.5; fifth gill slit, 24.1, 23.2; origin pectoral fin, 23.8, 22.9; origin first dorsal fin, 47.0, 49.8; origin pelvic fins, 40.6, 41.6; origin second dor- sal fin, 58.0, 64.2; origin anal fin, 49.3, 54.1; origin upper caudal lobe, 69.6, 71.2; anus, 48.6, 47.8. Greatest width of: trunk at pectorals, — , 11.9; trunk at pelvics, 6.4, 6.4; trunk at caudal fin origin, 2.0, 2.4. Greatest height of: trunk at pectorals, — , 8.6; trunk at pelvics, 9.3, 9.5; trunk at origin caudal, 4.6, 4.9. Orbit: horizontal diameter of opening, 2.8, 2.4; vertical diameter, 0.3, 0.7. Spiracles: greatest diameter, 0.3, 0.8; least distance from eye opening, 1.0, 0.9. Mouth: width, 9.0, 8.9; length, 4.3, 3.4; length upper labial furrow, 3.8, 3.4; length lower labial furrow, 2.9, 2.8. Nostrils: least distance between, 4.6, 4.9. Gill slits: height of first, 1.3, 1.5; height of fifth, 1.3, 1.2. First dorsal fin: length base, 6.4, 7.0; length posterior tip along free inner margin, 3.2, 3.4; height, 3.5, 3.4; length anterior margin, 9.6, 9.2. Pelvic fins: overall length, origin to posterior tip, 12.2, 13.8. Second dorsal fin: length base, 7.2, 7.6; length posterior tip, 3.5, 1.5; height, 3.8, 3.4; length anterior margin, 10.7, 9.8. Anal fin: length base, 15.7, 16.2; length posterior tip, 1.4, 0.9; height 5.2, 5.8; length anterior margin, 9.0, 7.6. Caudal fin: upper margin, 29.9, 29.4; anterior margin lower lobe, 10.1, 10.1; tip caudal to notch, 4.3, 4.6. Distance between fin bases: first and second dorsals, 9.0, 8.6; pectoral and pelvic, 8.1, 10.7; pelvic and anal, 1.4, 1.5; anal and lower caudal, 0.0, 0.3; second dorsal and upper caudal, 1.7, 1.5. Teeth of upper jaw in both specimens in about 62 vertical rows, the lower jaw teeth in slightly fewer rows but more irregular. Most teeth with four or five cusps, the median cusp longest. Denticles with three posteriorly directed points, the middle point longest and with a weak ridge. First dorsal fin origin in both specimens over the middle of the pelvic fin base. In the lectotype the second dorsal is only slightly larger in area than the first dorsal, but in the paralectotype the second dorsal is about V/ 3 times that of the first dorsal. The two syntypes had 39 and 40 monospondylous vertebrae, respectively. 19 Discussion. The two larger of Brauer's (1906) three type-specimens were loaned to me by the Zoological Museum of Humbolt University of Berlin and are in remarkably good condition for Apristurus preserved for so long a time. Brauer's own measurements were obviously taken from the 132-mm juvenile specimen which was not preserved and most of the differences between Brauer's descriptive account and mine can be attributed to differences between juveniles and half-grown or adult specimens. Juveniles of Apristurus species may differ greatly in proportions from half-grown or adult specimens. My own identification of Gulf of Mexico specimens of A. parvipinnis as A. indicus (Springer 1966) followed my mistakes in understanding Brauer's account of the species. My error evidently was responsible for the reference to A. indicus by Bass et al. (1975:7) as having the origin of the first dorsal behind the pelvic axil. In A. indicus the origin of the first dorsal fin is well in advance of the posterior end of the bases of the pelvic fins. Apristurus investigatoris (Misra) Pentanchus (Parapristurus) investigatoris Misra 1962:636, pi. 1 (Andaman Sea). Material examined. No specimens seen. The holotype and only specimen mentioned by Misra is ZSI Fl, 627/2, collected at HMS Investigator station in the Andaman Sea, lat. 11°46'N, long. 93°10'E, from 1,040 m, bottom green mud, globigerina ooze, bottom temperature 4.4°C. The holotype is a female 260 mm long. Diagnosis. The illustration with Misra's description of A. investigatoris shows the first dorsal fin to be about half the area of the second dorsal fin, the first dorsal origin about over the middle of the pelvic fin bases, and the distance between the dorsal fin bases about 75% of the length of the snout measured from the tip to the midpoint of the upper lip. Misra's diagnosis separates investigatoris from herklotsi by several differences in proportional measurement. The most striking difference, indicated by comparison of illustrations, is that the caudal fin in investigatoris is only 1.5 times the head length but is 1.9 times the head length in herklotsi. Apristurus japonicus Nakaya 1975 Apristurus japonicus Nakaya 1975:24, figs. 10, 11 (off Cape Daito, Japan). Material examined. None. Types. Holotype: HUMZ 40082, 697 mm ad. S, off Cape Daito Japan. Paratypes: HUMZ 40075, 626 mm ad. ?; HUMZ 39961, 690 mm ad. '; HUMZ 40076-40081, 654-711 mm ad. £; all paratypes from same locality as holotype. Diagnosis. Nakaya ( 1975:24) stated that A. japonicus differed from other Apristurus in having the distance between the origins of the pectoral and pelvic fins greater than the distance from the snout tip to the origin of the pectoral fin, but shorter than that from the snout tip to the end of the pectoral base. Nakaya noted that interbranchial septa sometimes had slight prolongations. Nakaya's measurements of A. japonicus indicate a very short snout in adults, shorter than the snouts of any of the specimens of Apristurus that I have examined. Also, A. japonicus has a greater number of monospondylous vertebrae, 43 to 46, than any other Apristurus. Description. Nakaya (1975:24) gave a comprehensive description which will not be repeated here. Points that might be emphasized are that A. japonicus is a moderately large species of Apristurus, adults 626 to 711 mm. The first dorsal is slightly smaller in area than the second dorsal and its origin is slightly in advance of the posterior end of the pelvic fin bases. The interspace between dorsal fin bases is greater than the length of the snout. Discussion. The description of A. japonicus should reduce the confusion over identification of Apristurus species from Japan. The proportions, long trunk sector and short snout, may differ in young specimens but Nakaya's (1975) descriptive account provides a good starting reference for comparisons of Japanese Apristurus. Apristurus kampae Taylor Apristurus kampae Taylor 1972:71, figs. 1, 3A, 4, 5 (from Gulf of California). Material examined. None. The species is treated here as being known only from the holotype. Taylor's second specimen may belong to this species but differs in several important features (see under discussion below). The holotype, SIO 70-278, a 335-mm i from lat. 27°22.4'N, long. 111°20.5'W to lat. 27°10.1'N, long. 111°29.6'W in 1,830 to 1,888 m, is the sole basis for the following diagnosis and description, here modified from Taylor (1972). 20 Diagnosis. Apristurus kampae is distinguished from all other species of the genus by its comparatively short anal fin, its base only about twice as long as the base of either the first or second dorsal, the posterior end of its base separated from the lower caudal lobe by a distance about as great as the length of the base of the second dorsal fin. The anal fin is also very high. Additional characters useful in comparison with characters noted in this paper for other species and prac- tical for comparison with most original descriptions are: caudal fin moderately long, its length about equal to the dis- tance from the tip of the snout to the origin of the pectoral; first dorsal fin origin in advance of posterior end of pelvic fin base, its area somewhat less than area of second dorsal but not as little as half the area of the second dorsal; length of snout in front of mouth and distance between dorsal fin bases about equal, and about three-fourths length of anal fin base; gill slits long, their length about 3.4 to 4.8% TL; eye moderately large, length of eye opening about 3.5% TL. Description. Taylor's (1972) description of the holotype of A. kampae is comprehensive and details will not be repeated here except to note that the holotype has 34 monospondylous vertebrae and that it is 335 mm long, at which size it appears nearly mature. Apristurus laurussonii (Saemundsson) Figure 14 Figure 14.— Apristurus laurussonii, 540-mm female, MCZ 38406, collected off New England. Drawing by Mary Wagner. Scyllium laurussonii Saemundsson 1922:73, pi. 4, fig. 2 and pi. 5, fig. 4, Saemundsson's text references to figures for Scyllium laurussonii and Pristiurus jenseni are reversed. (Near Vestmannaeyjar Island off southern Iceland.) Apristurus profundorum: Bigelow et al. 1953:214-217, part; Bigelow and Schroeder 1954:40, part. Apristurus laurussonii: Springer 1966:615, fig. 24B; in Monod and Hureau 1973:20. Material examined: NHMR, holotype, ?, 673 mm, from 560 m off southern Iceland; MCZ 38406, $, 540 mm, coast of New England; 28, 660-675 mm, from 1,024 m and 1,462 m off north coast of Hispaniola; 4 $, 466-550 mm, from 840 to 1,115 m off Alabama; 2 $, 4 $, 280-510 mm, from 1,097 m off Tamaulipas, Mexico; S, 536 mm, from 1,100 m, Gulf of Campeche. Diagnosis. Dorsal fins nearly equal in area; origin of first dorsal fin over or slightly behind middle of pelvic fin base; body sector (tip snout to cloaca) slightly shorter than tail sector; anal fin long, extending from tips of pelvic fin nearly or completely to origin of lower caudal lobe, anal fin base equal to or greater than distance from tip of snout to first gill slit and greater than distance between dorsal fin bases; length of eye opening less than least distance between nostrils; snout relatively short, less than 8.0% TL; monospondylous vertebrae 38 to 43. Description. Proportional measurements as percentages of total length are given below, first figure for 673 mm female holotype from Iceland, second figure for 675 mm adult male from Hispaniola. Tip of snout to: front of mouth, 7.6, 6.9; eye, 8.5, 7.5; first gill slit, 17.2, 15.6; fifth gill slit, 20.4, 20.0; origin pectoral, 19.9, 18.8; origin first dorsal fin, 50.4, 49.6; origin pelvic fins, 42.3, 44.0; origin anal fin, 58.1, 55.8; origin second dorsal fin, 65.6, 65.1; origin upper caudal lobe, 72.3, 75.8; anterior margin cloaca, 49.3, 45.1. Orbit: horizontal diameter of opening, 3.2, 2.5. Spiracles: diameter, 0.6, 0.4; least distance from eye, 0.8, 1.2. Mouth: width, 7.2, 7.0; length, 3.2, 3.1; length upper labial furrow, 2.7, 2.7; length lower labial furrow, 2.0, 2.4. Nostrils: least distance between, 3.8, 3.7. Gill slits: height of first, 1.5, 1.5; height of fifth, 1.4, 1.0. 21 First dorsal fin: length base, 7.0, 6.5; height, 2.1, 3.3; anterior margin, 10.3, 11.1. Second dorsal fin: length base, 5.8, 5.8; height, 2.4, 3.4; anterior margin, 9.7, 10.8. Anal fin: length base, 14.6, 16.9; height, 4.6, 5.0; anterior margin, 10.0, 8.6. Pectoral fins: anterior margin, 11.9, 13.9; greatest width, 8.2, 8.9. Caudal fins: length upper caudal lobe, 27.7, 24.2. Distance between fin bases: first and second dorsals, 9.1, 10.5; pectoral to pelvic, 14.8, 17.6. Teeth: mostly tricuspid, small, in about 40 to 50/38 to 44 diagonal rows; in about 70 to 90/54 to 64 vertical rows; no indication of sexual dimorphism. Dermal denticles: small, about 0.4 mm long and 0.3 mm wide, three-pointed, central ridge moderately strong, lateral ridges weaker; denticles of male slightly larger. Vertebrae: monospondylous, 43, 41, (range in 16 specimens, 38-43). Discussion. Apristurus laurussonii was taken in 560 mm off Iceland, but specimens from the Gulf of Mexico latitudes are apparently restricted to nearly twice that depth. Saemundsson's (1922) original description may have been overlooked because it was in Icelandic and the captions for laurussonii and Pristiurus jenseni were reversed. The holotype in the Museum at Reykjavik was in good condition when I examined it in 1964. Apristurus longicephalus Nakaya Apristurus longicephalus Nakaya 1975:32, figs. 15, 16 (Tosa Bay, Shikoku, Japan). Material examined. None. Diagnosis. Nakaya's diagnosis was based only on the holotype, HUMZ 42399, an immature male 375 mm TL. He stated: distance between pectoral and pelvic fin origins equal to distance from snout tip to eye; first dorsal fin origin over posterior part of pelvic fin base; snout about 2X interorbital distance and tapering abruptly in advance of nostrils. The snout length, 12.4% TL, is greater than in any other species of Apristurus that I have examined. Nakaya found 32 monospondylous vertebrae in the holotype. Discussion. Nakaya's account of A. longicephalus is thorough and details will not be repeated here. His description is, nevertheless, based on only one immature specimen which seems to be closely similar to A. herklotsi (Fowler 1934) and to A. investigatoris (Misra 1962). These also are one-specimen species, each known only from an immature holotype. Fowler's holotype of A. herklotsi, USNM 93134, is in poor condition and not satisfactory now for detailed measurements. I have not seen the holotype of A. investigatoris. All three species, A. herklotsi, A. investigatoris, and A. longicephalus, were illustrated and the illustrations suggest general similarity in body proportions. All three have proportionally long tails, their lengths from the origin of the lower caudal lobe as measured on the illustrations are 38.9^ TLfor.A. herklotsi, 35.6% TL for A. longicephalus, and 29.6% for A. investigatoris. Because proportional tail lengths and snout lengths in scyliorhinids may change during growth, the differences noted above do not set A. longicephalus apart from other Apristurus species in an entirely satisfactory way. The small number of monospondylous vertebrae, 32, in the holotype of A. longicephalus confirms its distinction from other species of Apristurus from Japanese waters. Apristurus herklotsi has 28 monospondylous vertebrae. I have no vertebral count for A. investigatoris. Apristurus macrorhynchus (Tanaka) Scyliorhinus macrorhynchus Tanaka 1909:1 (neighborhood of Misaki, Japan), type, No. 2153, Imperial University Collection. Apristurus macrorhynchus: Garman 1913:97; Nakaya 1975:36, figs. 17-19. Pentanchus macrorhynchus: Fowler 1941 :56, fig. 5. Note that fig. 5 is captioned platyrhynchus but depicts macrorhynchus. Material examined. USNM 22623, ?, 660 mm, Japan. Diagnosis. In A. macrorhynchus the two dorsal fins are nearly equal in area; the snout is long and somewhat tapered, its length nearly equal to the distance between the dorsal fin bases; the origin of the first dorsal is slightly in advance of the posterior end of the pelvic fin base; and the posterior end of the second dorsal fin base is in advance of the posterior end of the anal fin base by a distance nearly half as great as the horizontal diameter of the eye opening. In combination, these characters separate macrorhynchus from all other species of the genus. 22 Description. Tanaka's description of macrorhynchus designated a type, No. 2153 in the Imperial University collection, and included some measurements of the type specimen. The only specimen I have seen that agrees well with Tanaka's description of macrorhynchus is USNM 22623. The specimen was evidently the subject for an illustration (fig. 5) in Fowler's 1941 monograph but was mislabeled as A. platyrhynchus. I have included Tanaka's measurements of the type, a 470-mm young male, revised here as percentages of total length, with similar proportional measurements of the 660 mm female from the USNM. In the following series, the first figure in each pair refers to the type, the second to USNM 22623; dashes are used where data are not available, in this instance due to the poor condition of the USNM specimen. Tip of snout to: front of mouth, 9.4, 9.7; eye, 10.6, — ; first gill slit, 17.2, 16.7; fifth gill slit, — , 21.2; origin pectoral fin, 22.8, 20.0; origin first dorsal fin, 46.8, 48.5; origin pelvic fins, 38.3, 36.4; origin second dorsal fin, 61.7, 61.2; origin anal fin, — , 53.0; origin upper caudal fin, — , 70.9; origin lower caudal fin lobe, 69.2, — . Orbit: horizontal diameter of opening, 3.2, — . Nostrils: least distance between, 3.0, — . First dorsal fin: length base, 4.7, 4.5. Second dorsal fin: length base, 4.7, 5.3. Anal fin: length base, 15.3, 16.9. Distance between dorsal fin bases: 10.2, 10.2. Tanaka (1909) specifically compared macrorhynchus and platyrhynchus. Apristurus macrorhynchus was said to have small dermal denticles but not as small as those of A. playyrhynchus. Discussion. (See discussion under A. platyrhynchus.) Apristurus maderensis Cadenat and Maul Apristurus maderensis Cadenat and Maul 1966:769, figs. 1, 2, 3 (Madeira); Springer 1973:20. Material examined. None. The holotype, an adult female, 668 mm, is No. 18750, Museu Municipal do Funchal, is the only known specimen. Diagnosis. Apristurus maderensis is unique in the genus in having median, posteriorly projecting lobes extending from the gill covers (interbranchial septa) across the second to the fifth gill slits. It is one of the species having the second dorsal fin larger than the first dorsal in area and in having the origin of the first dorsal over or slightly in advance of the posterior end of the pelvic fin base. Discussion. Cadenat and Maul's description and measurements of the holotype are comprehensive and will not be repeated here. The holotype of A. maderensis had an egg capsule with more than 20 longitudinal striations. The illustration (Cadenat and Maul 1966, fig. 3) showed the striations appreciably higher than in other Apristurus egg capsules identified during the source of this study. Apristurus riveri and A. parvipinnis had egg capsules without prominent striations. Nakaya (1975) observed fine striations on egg capsules of A. platyrhynchus and A. macrorhynchus. The egg capsules of A. brunneus as illustrated by Cox (1963) appear smooth or very finely striated. The egg capsule of A. maderensis greatly resembles the egg capsule of Halaelurus canescens. Apristurus microps (Gilchrist) Scylliorhinus microps Gilchrist 1922:46, pi. VII, fig. 1 (coast of South Africa). Pentanchus microps: Smith 1949:54, fig. 41, part. Material examined. None. Gilchrist's holotype was not located. Diagnosis. According to Gilchrist's brief description and figure, A. microps has the smallest eye of any species of the genus. The very short distance separating the dorsal fin bases, less than the length of the first dorsal fin base, is also unique among species of Apristurus. Gilchrist did not give the size of the only specimen he had, and the specimen was not seen by Barnard (1925) when he wrote his monograph on South African fishes. A specimen in the British Museum labeled microps does not fit Gilchrist's description. 23 Discussion. If Gilchrist's illustration showing a remarkably small distance between the dorsal fin bases is accurate, this species must be recognized. No type has been found, however, and it is possible that Gilchrist's illustrations were not good representations of proportions of parts. Three specimens, 325 to 465 mm, collected by research vessels of the U.S.S.R. between lat. 26°S and 33°S off West Africa in 800 to 1,000 m were identified as Apristurus microps (Georgio Golovan, Acad. Sci. U.S.S.R., Moscow, pers. commun.). Golovan had a larger collection of Apristurus from African coasts than hitherto reported and seems to have been able to make identifications consistent with original descriptions. Apristurus nasutus DeBuen Apristurus nasutus DeBuen 1959:176 (vicinity of Valparaiso, Chile, in 400 m); Kato et al. 1967:25; Chirichigno 1974:30, fig. 17. Material examined. ZMK - Galathea 739, ad. &, 503 mm, Gulf of Panama, lat. 07°22'N, long. 79°32'W, 915-925 m; USNM, S, 465 mm, off Valparaiso, Chile, lat. 35°26'S, long. 73°01'W, 290-450 m; USNM ad. $, 520 mm, off Guaya- quil, Ecuador, lat. 03°57'S, long. 81°20'W, 600-750 m. DeBuen's holotype, S, 594 mm, EBMCh 10.184 was not seen. Diagnosis. Apristurus nasutus is very much like A. brunneus but differs in having a greater interspace between dorsal fin bases and in being dark gray or gray-black instead of brown or brownish. Adults of A. nasutus, A. brunneus, and A. japonicus usually have shorter snouts (measured from front of mouth) than other species of Apristurus. In A. nasutus the length of the orbit (3.4-3.8) is greater than the length of the orbit in A. japonicus (2.4-3.0). Apristurus nasutus belongs to the group of Apristurus species having the first and second dorsal fins about equal in area, and this serves to separate it from the Pacific species A. platyrhynchus, A. longicephalus, A. verweyi, and A. herklotsi. The very long snout of A. longicephalus and the long snout of A. macrorhynchus set those species off from A. nasutus. Description. In the following proportional measurements expressed as percentages of total length, the first figure in each group refers to the adult male, 503 mm, the second figure to the 465-mm female, and the third to a second adult male, 520 mm long. Tip of snout to: front of mouth, 6.6, 7.7, 6.7; eye, 8.0, 9.0, 7.1; spiracle, 12.1, 12.3, 10.9; first gill slit, 17.7, 18.1, 16.1; fifth gill slit, 24.4, 22.1, 24.2; origin pectoral fin, 23.9, 23.4, 22.3; origin first dorsal fin, 50.7, 51.6, 49.9; origin pelvic fins, 43.4, 44.1, 45.7; origin second dorsal fin, 67.5, 66.7, 66.2; origin anal fin, 56.7, 59.1, 59.1; origin upper caudal lobe, 75.2, 77.4, 73.5; anterior end cloacal opening, 48.8, 51.6, 50.3. Orbit: horizontal diameter of opening, 3.4, 3.4, 3.8; height opening, 1.0, 1.7, 1.2. Spiracles: diameter, 0.3, 0.6, 1.0; least distance from eye, 1.0, 0.9, 1.0. Mouth: width, 7.2, 6.7, 9.0; length, 5.2, 4.5, 4.6; length upper labial furrow, 2.8, 3.2, 5.0; lower labial furrow, 2.1, 1.7, 2.5. Nostrils: least distance between, 3.4, 3.7, 2.9. Gill slits: height of first, 1.8, 1.9, 1.7; height of fifth, 1.2, 1.5, 1.5. First dorsal fin: length base, 5.8, 4.9, 6.0; length free inner margin, 3.2, 3.0, 3.3; height, 2.8, 2.8, 2.5; length anterior margin, 8.6, 8.6, 8.1. Second dorsal fin: length base, 5.0, 5.2, 5.4; length free inner margin, 3.2, 3.2, 3.3; height, 4.2, 2.8, 4.4; length anterior margin, 8.4, 9.5, 8.3. Anal fin: length base, 13.7, 14.8, 12.1; length free inner margin, 1.4, 1.1, 1.3; height, 4.2, 4.7, 4.6; length anterior margin, 8.0, 7.3, 7.7. Pectoral fins: width base, 6.0, 8.0, 7.7; length anterior margin, 11.7, 11.8, 12.9; greatest width, 6.4, 8.0, 5.8. Distance between fin bases: first to second dorsal, 11.9, 10.8, 9.8. Teeth in all specimens mostly tridentate, but some with five cusps near angles of jaws. Upper and lower teeth similar, small, about 75 vertical rows in both jaws. Monospondylous vertebrae 36 in three specimens. Color dark-gray or gray-brown without prominent markings. DeBuen (1959) noted: color in Formalin uniform gray with posterior borders of dorsal, pectoral, anal, and caudal tip colorless. Discussion. The specimens of Apristurus nasutus examined for this study were from the coasts of Chile, Ecuador, and Panama, and those specimens were compared with examples of A. brunneus from the coast of Washington. Separa- tion of the two species may be impossible on the basis of the dorsal interspace measurements. A larger collection of specimens is needed to determine whether or not A. nasutus should be recognized as a valid species. 24 Apristurus parvipinnis Springer and Heemstra, n.sp. Figure 15, Table 1 Figure 15.— Apristurus parvipinnis, 395-mm female, taken off Pensacola, Fla. Drawing by Mary Wagner. Apristurus indicus: Springer 1966:613 part, and fig. 22B. Holotype: USNM 206178, S, 476 mm, Gulf of Mexico off Pensacola, Fla., in 1,115 m. Paratypes: USNM 206179, 5, 465 mm, same locality as holotype. USNM, ?, 395 mm, from MV Oregon stn. 3586, off the Caribbean coast of western Panama from 875 to 930 m; S, 268 mm, Gulf of Campeche from 895 m; ?, 512 mm, Gulf of Campeche from 1,097 m; ?, 485 mm, off west coast Florida from 722 m; 2 $, 386-521 mm off French Guiana, 636 m; ?, 415 mm, off French Guiana, 676 m; S, 340 mm, ?, 459 mm, off Pensacola, Fla., 914 m; 2 S, 425, 470 mm, off Caribbean coast of Colombia. Diagnosis. Second dorsal fin 2 to 4 times larger than first dorsal; origin of first dorsal at a vertical between posterior end of pelvic fin base and origin of anal fin; orbit moderately long, 2.9 to 3.6% TL, its length about equal to nostril length and somewhat less than the least distance between nasal apertures; snout long, 8.5 to 10.7% TL, its length usually somewhat greater than distance between dorsal fin bases; anal fin base 16.1 to 18.6% TL; distance between pectoral fin base and pelvic fin base about 11 to 14% TL; monospondylous vertebrae 37 to 41. Apristurus parvipinnis and A, canutus differ from other species of Apristurus in having, in combination, the second dorsal fin 2 to 4 times as large in area as the first dorsal fin and in having the first dorsal origin posterior to the end of the pelvic fin base. Apristurus parvipinnis has a long trunk segment, the distance between pectoral and pelvic fin bases about 11 to 14% TL as compared with 6.0 to 8.5% TL in A. canutus. The longer prepelvic trunk of parvipinnis is also reflected by its greater monospondylous vertebrae count, 37 to 40 as compared with 33 to 36 in canutus. Description. Proportional measurements as percentages of total length of the holotype (first figure), paratype (second figure), and range in 12 specimens 268 to 512 mm long (third figures), are given as follows: Tip of snout to: front of mouth, 8.8, 9.2, 8.6-10.4; eye, 10.1, 10.5, 9.6-11.2; spiracle, 14.5, 14.8, 13.7-15.7; first gill slit, 18.7, 18.9, 17.4-19.4; fifth gill slit, 22.5, 22.4, 20.9-23.5; origin pectoral fin, 21.4, 20.6, 20.0-22.5; origin first dorsal fin, 51.5, 51.6, 49.6-52.8; origin pelvic fins, 42.0, 41.1, 39.4-43.0; origin second dorsal fin, 63.0, 64.5, 60.4-65.0; origin anal fin, 54.6, 53.3, 51.3-54.7; origin upper caudal lobe, 72.2, 73.1, 69.4-73.8; anterior end cloacal opening, 48.3, 47.5, 45.1-48.3. Greatest width of trunk at: pectoral fin origin, 9.5, 9.5, 8.9-10.4; pelvic fin origin, 5.9, 5.8, 4.7-6.9; caudal base, 2.1, 2.4, 1.6-2.4. Greatest height of trunk at: pectoral fin origin, 9.0, 9.7, 5.1-9.8; pelvic fin origin, 10.9, 11.0, 7.8-11.0; caudal base, 4.6, 4.3, 4.0-4.9. Orbit: horizontal diameter of opening, 3.1, 3.0, 2.9-3.5; vertical diameter opening, 1.2, 0.9, 0.7-1.4. Spiracles: greatest diameter, 0.4, — , 0.4-0.6 in four specimens; least distance from eye opening, 1.1, 1.3, 0.7-1.3. Mouth: width, 7.1, 7.5, 7.0-9.2; length, 3.6, 3.9, 2.8-3.9; length upper labial furrow, 2.9, 2.8, 2.6-3.4; length lower labial furrow, 1.7, 2.2, 1.7-2.2. Nostrils: least distance between, 3.8, 3.8, 3.6-4.5. Gill slits: first, 1.7, 1.9, 1.3-2.2; fifth, 1.5, 1.5, 1.0-1.7. First dorsal fin: length base, 3.8, 4.1, 2.6-5.0; length posterior tip (inner posterior margin), 2.3, 2.2, 1.4-2.6; height, 3.4, 1.5, 1.3-3.4; length anterior margin, 6.1, 6.0, 5.5-7.6. Second dorsal fin: length base, 5.7, 5.8, 5.3-7.3; posterior tip, 2.9, 2.4, 2.3-3.2; height, 2.5, 2.4, 2.0-3.2; anterior margin, 9.2, 9.0, 8.3-9.7. Pectoral fins: width base, 8.0, 7.7, 7.1-9.8; length anterior margin, 11.6, 10.1, 10.1-14.6; greatest width, 8.2, 7.6, 7.2-8.5. Pelvic fins: distance origin to posterior tip, 12.6, 11.0, 10.1-12.6. Caudal fin: upper margin, 27.3, 26.2, 26.2-30.9. 25 Distance between fin bases: first to second dorsals, 8.0, 8.8, 6.7-9.4; pectoral to pelvic, 13.7, 13.5, 11.3-13.7; pelvic to anal, 3.8, 2.4, 2.4-3.9; second dorsal to caudal, 4.0, 3.4, 2.2-5.2; anal to lower caudal origin, 0.0, 0.0, 0.0-1.0. Both of the types have 37 monospondylous vertebrae but the numbers in 44 other specimens are from 38 to 40. Teeth small, crowded, with more than 50 somewhat irregular vertical rows in each jaw; teeth with three to six cusps, but those with four cusps most numerous; tooth surfaces with one to several vertical ridges, the ridges extending nearly to tooth tips; teeth of upper and lower jaws similar; no indication of sexual dimorphism of tooth shape in the specimens available. Apristurus paruipinnis specimens are usually black without prominent markings. The flattened snout appears broad in comparison with species such as A. riueri and A. macrorhynchus which have somewhat narrower snouts slightly taper- ing toward the tip. Discussion. Apristurus paruipinnis is one of the more commonly collected Apristurus in the Gulf of Mexico- Caribbean area along with A. laurussonii. Apristurus platyrhynchus (Tanaka) Scyliorhinus platyrhynchus Tanaka 1909:4 (from Japanese waters), holotype No. 2154, Imperial University Collection. Apristurus platyrhynchus: Garman 1913:98; Nakaya 1975:28, figs. 12-14. Pentanchus platyrhynchus: Fowler 1941, part, not fig. 5, see under A. macrorhynchus. Material examined. FMNH 74130, imm. 3, 525 mm (Sagami Sea) Owston collection No. 1019. Diagnosis. First dorsal about half as large in area as second dorsal, the length of its base about two-thirds length of second dorsal base, its origin posterior to posterior end of base of pelvics; snout moderately short, its length from tip of snout to upper lip less than distance between dorsal fin bases; anal fin long, its base as long or longer than distance from the tip of snout to first gill slit; eye moderately large, the length of its opening about equal to least distance between nasal openings; dermal denticles very small, smooth and velvety to touch; the holotype, an 800-mm male, presumably adult, is one of the largest Apristurus specimens recorded. Description. Tanaka's measurements of the 800-mm male, here revised as percentages of total length, follow together with proportional measurements of a 525 mm, immature male, FMNH 74130. The first measurement in each pair of figures represents Tanaka's holotype, the second represents the immature male. Dashes are used where data are absent. Tip of snout to: upper lip, 7.1, 8.4; eye, 8.5, 8.8; spiracle, — , 11.8; first gill slit, 15.6, 15.2; fifth gill slit — , 20.4; origin pectoral fin, 20.0, 19.6; origin first dorsal fin, 50.0, 46.7; origin pelvic fins, 33.6, 37.5; origin second dorsal fin, 63.8, 61.5; origin anal fin, 50.0, 50.5; origin lower caudal lobe, 70.0, — ; origin upper caudal fin, — , 69.2; anterior end of anus, — , 41.9. Orbit: horizontal diameter of opening, 3.0, 2.9. Nostrils: least distance between, 2.9, 2.9; greatest distance between, — , 6.4. Mouth: width, 6.5, 7.0; length, — , 3.0. Spiracle: diameter, 0.5, 0.2; distance from eye, — , 0.6; distance between spiracles, 6.5, — . First dorsal fin: length base, 2.8, 4.2; height, — , 1.9; length anterior margin, — , 6.7. Second dorsal fin: length base, 4.3, 5.3; height, — , 2.5; length anterior margin, — , 7.8. Anal fin: length base, 18.0, 15.0. Pectoral fin: width base, 6.8, 7.4; length anterior margin, — , 12.4. Distance between fin bases: first and second dorsals, 11.0, 11.0; pectoral and pelvic, — , 9.1; pelvic and anal, — , 4.8; anal and lower caudal, — , 0.0; second dorsal and upper caudal, — , 5.7. Tanaka (1909) stated that the holotype had very small scales, much smaller than in macrorhynchus. The larger flank denticles of FMNH 74130 are about 0.4 mm long, with three points, the middle point longest. In the original description the teeth are described as tricuspid, the median cusp largest, set at wider intervals than in macrorhynchus. The longest teeth in FMNH 74130 are only about 1.2 mm. Discussion. Before Nakaya's (1975) work on Japanese catsharks appeared, I had examined 11 specimens of Apristurus from Japanese waters but could identify with confidence only one as A. platyrhynchus and one as A. 26 macrorhynchus, before 1975 the only species of Apristurus known from Japan. It is true that many of the specimens I at- tempted to identify were immature but now with Nakaya's study it would be possible to do more. There may still be identification problems with young Apristurus from Japan as elsewhere either because species remain undescribed or because young specimens are not known or are poorly known. Apristurus profundorutn (Goode and Bean) Scylliorhinus profundorum Goode and Bean 1896:17, pi. 5, fig. 16 (off Delaware Bay in 1,492 m). Apristurus profundorum: Garman 1913:99; Bigelow and Schroeder 1948:222, fig. 38A; not of Springer 1966:612, figs. 22, 23 (see Parmaturus manis). Material examined. Only specimen seen certainly identified as A. profundorum was the holotype, USNM 35646. The holotype is in very bad condition and the following descriptive account is based chiefly on the description and illustra- tion of the holotype by Bigelow and Schroeder (1948:222-223, fig. 38, 38A). Diagnosis. First dorsal fin not much smaller in area than second dorsal; origin of first dorsal fin somewhat in advance of the posterior end of the base of the pelvic fin; eye length (length of opening) about 2.7% TL, eye length contained about 10 times in head (tip of snout to fifth gill slit); anal fin base 13.9% TL, about equal to distance from tip of snout to spiracle; distance between dorsal fin bases 8.2% TL, only a little less than the width of the mouth; snout (in front of mouth) about 8.9% TL, abruptly narrowed in advance of nostrils, tapering, the tip broadly rounded; little if any inter- space between the posterior end of the anal fin base and the origin of the lower caudal fin lobe; anal fin moderately high, 4.3% TL. The holotype has 33 monospondylous vertebrae. Discussion. The holotype of A. profundorum, a male 510 mm long, USNM 35646, taken off Delaware Bay in 1,492 m, seems to have deteriorated greatly between 1948 and 1964. At present it is impossible to determine much about morphometries from the specimen so the measurements made by Bigelow and Schroeder (1948:222) are assumed to apply. In June and July 1952, collections of fishes were made under the direction of W. C. Schroeder at depths from 722 to 969 m off the coasts of Nova Scotia and New England. Several specimens identified as Apristurus profundorum were ob- tained and were reported with some detail in a series of publications (Bigelow et al. 1953; Bigelow and Schroeder 1954; Schroeder 1955; Springer 1966). I reexamined the same series of specimens for this study and found that I had misiden- tified the specimens and that the series contained no A. profundorum, but instead those described and figured in greatest detail (Springer 1966:612, figs. 22A, 23, and 25 left) were the forms described in later pages as Parmaturus manis and some others of the series were Apristurus laurussonii. Parmaturus manis has a well-defined caudal crest and does not belong in Apristurus. Whether A. laurussonii should be considered a synonym of A. profundorum remains uncertain. The fact that the holotype of laurussonii has 10 more monospondylous vertebrae than the holotype of profundorum supports separation of the two forms. Four specimens, 243-290 mm, taken off the west coast of Africa south of the Canary Islands at depths from 1,200 to 1,500 m were identified by Georgio Golovan as A. profundorum (pers. commun.). Clarification of the problems of identi- fication of nominal Apristurus from the African coasts, A. indicus, A. microps, A. atlanticus, and a specimen recorded by Karrer (1973), will have to await comparisons of larger collections from the area than have been available to me. Apristurus riveri Bigelow and Schroeder Figures 7, 8, 16, 17, 18; Table 1 Figure 16. — Apristurus riveri, adult female, 400 mm, taken off Caribbean coast of Panama. Drawing by Mary Wagner. 27 Figure 17. -Mouth of adult male Apristurus riveri, 430 mm, showing comparatively large monocuspid teeth. Figure 18. — Apristurus riveri, immature male, 350 mm, western Florida, showing tricuspid teeth found in all females and in imma- ture males. Apristurus riveri Bigelow and Schroeder 1944:23, pi. 7 (off north coast of Cuba, RV Atlantis stn. 2993, in 1,060 m); Bigelow and Schroeder 1948:225, fig. 39; Springer 1966:613, figs. 4, 5, 7. Material examined. MCZ 36092,$, 407 mm, holotype, north coast of Cuba; 3, 460 mm, ?, 338 mm, west of Dry Tortugas, Florida, 914 m; ?, 400 mm, 3, 250 mm, off Mississippi coast, 622 m; $, 430 mm, 3 ?, 400, 400, 405 mm, off Caribbean coast of Panama, 860-914 m. Diagnosis. Apristurus riveri is best recognized by its long narrow snout, narrower than in other species, with a narrower pore pattern on the lower side of the snout, the number of longitudinal rows of prominent pores extending from the level of the posterior end of the nasal aperture forward not more than four as compared with about eight or more in most other species of Apristurus. The rows of pores are somewhat irregular but I have found the character reliable for half-grown and adult riveri. Sexual dimorphism in tooth size and shape is greater for A. riveri than for any other species examined for this study. Adult males have monocuspid, cone-shaped teeth that are much larger than the teeth of the adult females or of the im- mature males (Fig. 7). The teeth of immature males have the same shape as the teeth of females. Apristurus riveri belongs to the group of Apristurus species having the second dorsal fin much larger than the first dor- sal in area and in having the origin of the first dorsal well in advance of the posterior end of the pelvic fin base. Some A. riveri have denticle-free black skin around the gill slits and the gill filament tips may extend past the gill covers. I have noted this condition chiefly in young specimens of both A. riveri and A. parvipinnis and do not regard it as reliable in diagnosis. The gill region of some A. riveri is well covered with denticles. Counts of monospondylous vertebrae are 30 to 32 in 16 specimens of A. riveri, a somewhat smaller number than observed for other Apristurus (Table 1) except A. herklotsi. Mature specimens examined were from 390 to 475 mm long. Description. Proportional measurements of the holotype were given by Bigelow and Schroeder (1948:223). Following are comparable measurements expressed as percentages of the total length for two adult males, 430 and 460 mm, with the range in three adult females, 400, 405, and 405 mm, in parentheses. Tip of snout to: front of mouth, 8.1, 7.4, (9.4-10.0); eye, 10.7, 10.0, (10.4-11.5); first gill slit, 19.8, 18.7, (18.3-21.0) fifth gill slit, 25.3, 23.9, (22.2-24.5); origin first dorsal fin, 48.9, 48.9, (47.5-48.6); origin pelvic fins, 41.9, 40.9, (40.2-42.7) origin anal fin, 53.6, 52.4, (51.8-52.3); second dorsal fin, 62.8, 61.3, (59.3-60.2); upper caudal lobe, 71.6, 71.1, (68.3-70.4) anterior end cloacal opening, 45.3, 43.5, (44.9-47.7). Orbit: horizontal diameter of opening, 3.5, 3.3, (3.3-3.5). Nostrils: least distance between, 4.0, 3.9, (3.7-4.3). 28 Mouth: width, 5.3, 6.7, (6.9-7.3); length, 4.6, 5.2, (3.2-4.3); length upper labial furrow, 2.3, 2.2, (2.0-2.5); length lower labial furrow, 2.8, 2.2, (2.5-3.0). First dorsal fin: length base, 4.0, 3.3, (4.2-4.3). Second dorsal fin: length base, 6.3, 5.4, (6.1-6.5). Anal fin: length base, 14.2, 13.0, (14.2-15.8). Pectoral fin: length anterior margin, 11.2, 9.8, (10.0-11.6). Distance between fin bases: pectorals and pelvics, 8.6, 9.8, (9.9-11.8); first and second dorsals, 7.6, 8.9, (6.9-7.4). Tip of oppressed pectoral to origin pelvic fin: 4.2, 5.4, (5.0-6.9). In A. riveri, as in other species of Apristurus, it is difficult to count the number of vertical tooth rows because of a tendency to arrangement in diagonal files (Fig. 8). Counts of vertical rows in the specimens examined were from 24 to 29 rows on each half of the upper jaw and 19 to 22 rows on each half of the lower jaw. Apristurus riveri is not as large as some of the other species. I estimate length of females at maturity as about 390 mm but males may be slightly larger. The longest specimen measured was 475 mm. Egg capsules of A. riveri are about 50 to 60 mm long by 12 to 15 mm wide. The capsules are smooth-surfaced, greenish, somewhat translucent, and have some indistinct longitudinal bands of darker color. Discussion. Apristurus riveri was taken less frequently by exploratory fishing vessels operating in the Gulf of Mexico- Caribbean area than A. parvipinnis and A. laurussonii. One possible explanation is that the depth range of A. riveri is in somewhat deeper water than other Apristurus of the region and fewer trawl hauls were made in deeper water. In an earlier paper (Springer 1966), A. riveri was the only species of the genus that was adequately and correctly defined but additional collections have confirmed earlier notes on riveri and add a little to the earlier account. New material shows that immature males have teeth similar to those in females, and it is evident that the transition from tricuspid to monocuspid teeth must be abrupt. Proof can only come with the capture of a male with both kinds of teeth, developing cone-shaped teeth near the germinal series and multicuspid teeth in the functional series. Since no published explanation of shark tooth dimorphism has yet appeared, I offer the following hypothesis to ac- count for it in A. riveri. In the few known observations of shark copulation, one species of the family Scyliorhinidae, Scyliorhinus canicula, has been observed. The essential preliminary is that the male seizes one of the female's pectoral fins with its jaws and then twists into position to permit copulation. Multicuspid teeth, especially those that are more or less flattened, tend to cut rather than pierce. Cone-shaped piercing teeth, as in the adult males of A. riveri, are less likely to cut through the thin pectoral fin of the female, and not only do less damage to the female, but also provide a firmer grip. This may be most important to the survival of the species in a sparsely distributed deepwater population. It was previously reported (Springer 1966) that female A. riveri with egg capsules in the oviducts had a flattened ring of white tissue around the cloaca. Subsequent collections of both A. riveri and A. parvipinnis exhibited a similar condi- tion but only when egg capsules were present in the oviducts and had capsule tendrils extending outside the oviducts. It is here postulated that the pad of white tissue, which is comparatively firm, is used to press and rub the tendrils against a relatively hard object. This may cause the tendril to bend and coil around the object so that the egg capsule can be pul- led out of the oviduct. It is possible that some external pull is necessary for deepwater species such as A. riveri in which the abdominal walls are weak and perhaps too poorly muscled to expel egg capsules. Apristurus saldanha (Barnard) Scylliorhinus saldanha Barnard 1925:44 (off Saldanha Bay, South Africa in 914 m). Pentanchus saldanha: Fowler 1941:59. Apristurus saldanha: Bigelow and Schroeder 1948:221 (key only); not of Norman 1939:36; not of Bass et al. 1975:8, fig. 5. Material examined. None. Barnard's holotype, now possibly lost, was an 810-mm male and the largest example of any species of Apristurus yet recorded. Barnard's description of the claspers shows the holotype to be an adult male. Diagnosis. In Apristurus saldanha the distance between the two dorsal fin bases is about as great as the distance between the tip of the snout and the spiracle, greater than in any other species of Apristurus except Apristurus platyrhynchus (Tanaka 1909) from Japanese waters. Also, as calculated from Barnard's description, A. saldanha has: first dorsal fin slightly smaller (in area) than second dorsal fin; length of anal fin base almost equal to distance from tip of snout to first gill slit; length of body before vent about one-half total length; length of head to fifth gill slit about one- fifth total length; eye (there is some question as to how Barnard measured the eye because he states length of eye [not the orbit as indicated by scaleless skin]) three in snout — this is calculated as 2.6% TL; first dorsal fin origin about over mid- point of pelvic fin base; snout length about 7.2% TL; teeth tridentate; dermal denticles tridentate, lateral points much smaller than central point, obscure; claspers reaching anal fin origin. 29 Discussion. In 1969, I examined three specimens (BMNH 1935.5.2, 56-58) which appeared to be those identified by Norman (1935, 1937) as A. saldanha and were taken off Saldanha, South Africa, in 735-1,000 m at RV Discovery Station J. When I saw the specimens, two were labeled A. saldanha and one A. microps. All appeared to be immature, the two larger, a male 430 mm and a female 440 mm, differed from the smallest one in having somewhat larger eyes and a propor- tionally longer trunk. None of the specimens had wide-bladed denticles that usually characterize most mature Apristurus. Bass et al. (1975:7, 8, figs. 5, 20A), tentatively and, I think, reluctantly, referred the specimens to A. saldanha and provided an important service by including a description and illustrations in their report. I do not agree that the specimens should be called A. saldanha chiefly because of their differences from Barnard's (1925) holotype in the proportions of dorsal fin interspaces. The differences seem to me to be too great to attribute to differential growth. I cannot identify the specimens, however, as belonging either to an African species or to a species known from other areas. Additional material may be reported soon for 1 of 11 Apristurus collected off the west coast of Africa. A 560-mm female Apristurus taken off Saldanha Bay by a research vessel of the U.S.S.R. had an interdorsal distance nearly equal to the distance from the tip of the snout to the spiracle (Georgio Golovan, Acad. Sci. U.S.S.R., Moscow, pers. commun.). Although this specimen does not fit exactly Barnard's description, its differences may be attributable to its small size. Apristurus sibogae (Weber) Scyliorhinus sibogae Weber 1913:595 (Makassar Strait in 655 m). Pentanchus sibogae: Fowler 1941:60. Material examined. ZMA 111.076, imm. 2, 210 mm, Indonesia, Makassar Strait, lat. 00°32'S; long. 119°39'E, in 665 m, the holotype and only specimen known. Diagnosis. First dorsal fin about one-third the area of the second dorsal, its base about one-half the length of the second dorsal base, its origin posterior to the rear end of the pelvic fin base and over the anal fin origin; eye opening very small, its length about one-half the least distance between nasal apertures and less than one-third the length of the snout to upper lip; length of snout about equal to distance between dorsal fin bases; color reddish-white. Discussion. Apristurus specimens are usually black, dark gray, or dark brown but the holotype of sibogae is now pinkish-white and was described as reddish-white by Weber in 1913. Bright light or sunlight may cause the dark color of Apristurus specimens to fade. One of Brauer's (1906) syntypes of indicus kept on exhibition for many years had faded to grayish-white by 1970 (Christine Karrer, pers. commun.); but except for this syntype of indicus and the holotype of sibogae, I have seen no light colored Apristurus. The holotype of sibogae was pinkish when examined by me in 1970 and I do not believe that the color could have been the result of fading. The specimen was in good condition but very fragile and, although I handled it carefully, it broke in two pieces. Apristurus verweyi (Fowler) Pentanchus verweyi Fowler 1934:237, fig. 2 (vicinity of Sibuko Bay, Borneo). Material examined. USNM 93135, imm. S, 295 mm, the holotype, Sibuko Bay, Borneo. Diagnosis. In Apristurus verweyi the first dorsal fin is much smaller than the second dorsal and its origin is posterior to the rear end of the pelvic fin base. It is thus similar to A. platyrhynchus but differs from that species in having the rear end of the second dorsal fin base over or slightly behind the rear end of the anal fin base. Discussion. The holotype and only known specimen of A. verweyi seen is not in good condition. The separation of this species from platyrhynchus seems tenuous. ASYMBOLUS WHITLEY Asymbolus Whitley 1939:229 (type-species, Scy Ilium anale Ogilby, by monotypy). Diagnosis. Asymbolus lacks a supraorbital crest that extends as a narrow shelf of the chondrocranium above the orbits, it lacks a crest of modified denticles along the upper edge of the caudal fin, it lacks barbels, and the posterior margins of the nasal flaps do not reach the mouth and do not cover a depression or channel connecting the nasal cavity with the mouth. Asymbolus has two dorsal fins. It has prominent but short labial furrows continuous around the mouth corners and extending along the upper and lower jaws for a distance not greater than the least distance between the nasal apertures. The pelvic fins of males are not united directly by a union of their inner margins (as in Juncrus) but are con- 30 nected by a membrane that forms a partial apron beneath the claspers. The eyes of Asymbolus are comparatively large with a prominent secondary lower eyelid which is separated from the primary lower eyelid and eye by a deep, but fully scaled, subocular gutter. The gutter extends from a little before the anterior corner of the eye, under the full length of the eye to the spiracle which is in the subocular gutter at its posterior end. In Asymbolus the membrane connecting the pelvics originates from the dorsal side of one fin near, but not at, the inner margin and passes under the claspers to the other fin. This arrangement differs from that forming an apron or partial apron in Scyliorhinus and Juncrus which have proximal inner margins of pelvics directly united for varying lengths. It is not unique in the family, however, since an arrangement similar to that in Asymbolus is present in Poroderma. Among scyliorhinid genera known to be represented in Australian waters, Asymbolus differs from Aulohalaelurus, Atelomycterus, and Cephaloscy Ilium in that Asymbolus lacks a supraorbital crest of the chondrocranium. In having a partial apron beneath the claspers, Asymbolus differs from all genera known from Australian waters except Juncrus, in which the partial apron is formed by the direct union of part of the inner margins of the pelvics as in Scyliorhinus. In Halaelurus membranes may be noted between the pelvic fins and claspers and between the two claspers at their bases but not extending under the claspers to connect the pelvics. Asymbolus analis (Ogilby) Scyllium anale Ogilby 1885:445 (type locality, Port Jackson Harbor, N.S.W., Australia). Scyliorhinus analis: Regan 1908a:460. Halaelurus analis: Garman 1913:85; McKay 1966:68. Asymbolus analis: Whitley 1940:89, figs. 29, 78, 81. Material examined. USNM 40016, ?, 565 mm, Sydney, Australia; USNM 40027, imm. S, 475 mm, Port Jackson, Australia (these two specimens must have been incorrectly measured by Fowler who reported (1941:49) lengths of 612 and 515 mm; the difference from present lengths seems too great to be accounted for by shrinkage); BMNH 1890.9.23.273, imm. 3, 470 mm, Australia; BMNH 1915.4.20.1-2, ad. $, 550 mm, New South Wales; BMNH 1937.9.21.1, stn. 113, lat. 42°40'S, long. 148°27.5'E, off Tasmania. Type. Ogilby designated as a holotype an adult male 22.5 in (ca. 570 mm) long, giving the register number AMS B.8447. J. A. F. Garrick recently examined the holotype for me (pers. commun.) and noted that it was ca. 560 mm long. Based on comparison of original measurements with remeasurements made after many years of storage in alcohol a shrinkage in length of 1.75% as here, or 1 to 3%, is to be expected. Diagnosis. Asymbolus analis should be readily recognized by the characters given in the diagnosis for Asymbolus. All of the specimens that I have seen (long in preservative) are light or dark brown above with scattered darker spots about the size of the dark-adapted pupil, and light tan or yellowish below without spots. Dorsal saddle blotches may also be present for these are indicated in Whitley's (1940, fig. 81) illustration and are mentioned in McKay's (1966) account. Asymbolus analis apparently never has white spots such as characterize Juncrus vincenti. Description. Measurements expressed as percentages of the total length follow. In each series of figures the first refers to an immature male 475 mm long, from Port Jackson, Australia, USNM 40027; the second figure to a female, pos- sibly mature, 560 mm long, from Sydney, Australia, USNM 40016; and the third to a 365-mm adult male from Tasmania, BMNH 1937.9.21.1. (see discussion of Tasmanian specimens as "island" forms of Australian scyliorhinids). Tip of snout to: front of mouth, 3.6, 3.8, 5.0; eye, 3.8, 4.1, 5.2; spiracle, 7.8, 8.0, 9.6; first gill slit, 12.2, 12.5, 12.3; fifth gill slit, 17.1, 16.6, 17.0; origin pectoral fin, 15.4, 15.9, 16.0; origin first dorsal, 47.6, 47.3, 41.1; origin pelvics, 37.9, 40.7, 35.6; origin second dorsal, 70.7, 66.1, 63.0; origin anal, 66.9, 57.1, 56.2; origin upper caudal lobe, 77.7, 79.5, 76.7; anterior end anal opening, 42.1, 43.4, 39.5. Eyes: horizontal diameter opening, 3.2, 3.2, 4.1; vertical diameter opening, 1.1, 0.7, 1.4. Spiracles: greatest diameter, 1.1, 0.4, 0.3; distance eye to spiracle, 0.8, 0.9, 1.1. Mouth: width, 5.5, 5.7, 7.1; length, 3.6, 3.4, 3.8; length upper labial furrow, 0.8, 0.8, 1.1; lower labial furrow, 1.3, 1.2, 1.6. Nasal apertures: minimum distance between, 1.6, 1.8, 1.9. Gill slits: height of first, 1.5, 2.0, 1.6; height of fifth, 1.1, 1.1, 0.8. First dorsal fin: length base, 6.1, 6.3, 5.8; length free inner margin, 2.1, 2.1, 2.2; height, 3.6, 3.9, 4.7; length anterior margin, 8.4, 9.6, 7.9. Second dorsal fin: length base, 6.9, 7.0, 6.3; length free inner margin, 2.3, 2.1, 2.2; height, 3.8, 3.9, 4.4; length anterior margin, 9.1, 9.6, 7.9. Anal fin: length base, 10.9, 11.6, 12.3; height, 3.4, 3.2, 3.3; length anterior margin, 7.4, 8.2, 6.6. 31 Pectoral fin: width base, 4.4, 4.8, 5.5; length anterior margin, 10.1, 11.3, 9.9; greatest width, 8.0, 8.8, 9.6. Pelvic fin: origin to rear tip, 11.8, 11.8, 11.5. Caudal fin: upper margin, 22.3, 20.5, 24.7. Distance between fin bases: first and second dorsals, 11.8, 12.0, 15.9; pectoral and pelvic, 18.1, — , — ; pelvic and anal, 11.8, — , — ; anal and caudal, 7.4, — , — ; second dorsal and caudal, 5.9, — , — . Asymbolus analis is slender and elongate, the head moderately flattened and the posterior trunk moderately compressed. The head is relatively short, less than one-fifth total length (less than three-sixteenths total length in specimens examined) and the tail, measured from origin of upper caudal, is from one-fourth to one-fifth total length. The mouth is large with a high rounded arch, the length of the mouth nearly as long as the snout. Labial furrows are short but moderately prominent and continuous around the mouth corners, the lower longer than the upper. Eye large, its length a little less than length of snout; length of subocular gutter including spiracle is a little greater than snout length. The nasal apertures are large, separated from one another by a distance about one-half the snout length and have broad outer nasal flaps entirely covering posterior nasal apertures but not reaching upper lips. The second dorsal fin is slightly larger than the first and the interdorsal distance is about twice the length of base of the second dorsal. The first dorsal originates behind the posterior end of the pelvic base and the second dorsal originates over the posterior third of the anal base. The lower caudal fin lobe originates considerably in advance of the origin of the upper lobe. The pectorals are moderate in length and wide with rounded corners. The teeth are small, in about 60/62 rows, those in the central part of the upper jaw and in most of the lower jaw are tricuspid with middle cusps longest; lateral upper jaw teeth mostly have five cusps; upper and lower jaw teeth are generally similar except lateral lower jaw teeth which are much smaller, flattened, with low cusps. The modification of the lateral lower jaw teeth appears to adapt them to a crushing function and the arrangement of teeth in a more or less shinglelike array is similar to the condition in lateral lower jaw teeth of some Atelomycterus. The dermal denticles are tridentate, their blades imbricate. The blades have a wide, but low, and inconspicuous ridge extending posteriorly from the central point. Pigment may be present within the denticles of those areas which have darker brown spots. The denticles are similar in shape over most external body surfaces. Small denticles are present on the anterior margin of the tongue and on the gill bars. The specimens examined were all light brown above with the roundish darker spots about the size of the dark-adapted pupil somewhat irregularly distributed over dorsal and lateral surfaces. Lower surfaces were lighter and unmarked. In two specimens vertebral numbers were: monospondylous, 39, 39; precaudal, 86, 86; caudal, ca. 42, ca. 56; total, ca. 128, ca. 142. Discussion. Asymbolus analis is recorded from New South Wales, Tasmania, and the southern coast of Australia west at least as far as Albany, Western Australia. It has been taken on the continental shelf and to depths of about 175 m. This species and other scyliorhinids of southern Australia and Tasmania are described by McKay (1966) as mostly in- habiting shallow coastal waters, in contrast to scyliorhinids of the subtropical and tropical Atlantic species which in- habit continental slopes at depths usually greater than 175 m. Asymbolus is oviparous. Its egg cases are described and illustrated by Whitley (1940:40, fig. 29). Three males, 470, 475, and 510 mm, are immature but a 550-mm male is mature. All of these specimens are from the Australian mainland shelf. The only Tasmanian A. analis seen was a 365-mm adult male. It seems possible that large series would show differences between mainland and Tasmanian populations. In the Australian-Tasmanian genus Juncrus, however, the larger form seems to be the Tasmanian one. ATELOMYCTERUS GARMAN Atelomycterus Garman 1913:100 (type-species Scyllium marmoratum Bennett 1830, by monotypy). Diagnosis. Atelomycterus has two dorsal fins nearly equal in area, the first originating over the middle of the pelvic base or over the posterior end of the pelvic base. It has a well-developed supraorbital crest on the chondrocranium pro- jecting as a narrow shelf above the orbits. The nasal flaps are long and usually extend to or past the edge of the upper lip, each covering a depressed segment in the upper lip forming channels in the upper lip connecting the mouth with the nasal cavities. Prominent and long labial furrows are continuous around the mouth corners, extending along the upper jaw for a distance slightly less than length of the orbit and along the lower jaw for a somewhat greater distance. Adults have the trunk vertebrae of adults calcified in a "Maltese-cross" pattern (White 1937). However, the type of calcification has not been determined either in young Atelomycterus or in many of the less common scyliorhinid species such as Aulohalaelurus labiosus. Discussion. Atelomycterus has a slender, almost eellike form tapering from the posterior end of the jaws. A slight constriction in the area of the first gill slit gives Atelomycterus a somewhat reptilian appearance, but otherwise it resem- 32 bles Scyliorhinus canicula in shape. According to White (1936b), Atelomycterus is found among coral reefs where its eel- like body is adapted to the sinuous movements needed for swimming into holes and around sharp corners. Atelomycterus probably gains some flexibility from its comparatively large number of vertebrae. It not only has a larger number than any other scyliorhinid, up to 172 in some males, but the trunk vertebrae are numerous with less abrupt and a less con- spicuous transition from the longer monospondylous vertebrae to the shorter diplospondylous vertebrae than in other scyliorhinids. White (1936a) included a family, the Atelomycteridae, in a brief synoptic list of families of galeoid sharks and later (1936b:19) gave a definition for the family, emphasizing the vertebral calcification pattern, in a paper entitled "Some transitional elasmobranchs connecting the Catuloidea with the Carcharinoidea." Although White treated only A. mar- moratus and six other species in that paper, she specifically stated that A. marmoratus was not a transitional shark but one adapted to a restricted environment. I provisionally recognize two species of Atelomycterus, A. marmoratus and A. macleayi. I am not certain, however, that A. macleayi warrants recognition or that A. marmoratus (as used here) represents a single species. Atelomycterus macleayi Whitley Atelomycterus macleayi Whitley 1939:230, fig. 3 (type-locality Melville Island, Northern Territory, Australia); Whitley 1940:92, figs. 86, 87; McKay 1966:67 (reference only); Taylor 1964:54. Material examined. USNM 174070, ad. $, Alb mm, 6 mi NNW of Darwin, Northern Territory, depth about 0.5 m, 20 March 1948; USNM 174071, ad. ?, 510 mm, 7 mi north of Darwin, depth about 3.5 m. The holotype, AMS 1.5269, was not examined. Diagnosis. Whitley's description of A. macleayi merely states that it differs from A. marmoratus in color, in relative position of the fins, and in the shape of the egg capsule which lacks tendrils at the anterior end. I did not make direct comparisons of specimens of A. marmoratus with the two specimens of Atelomycterus in the U.S. National Museum collection and my notes are insufficient to show differences other than small differences in the relative positions of first dorsal and pelvic fins. Atelomycterus needs further study and, as an aid to such a study, I have included a comprehensive description of A. macleayi based on one adult male and one adult female. Description. Body elongate, subcylindrical; head short, less than one-fifth total length; tail from origin of upper caudal fin about one-fifth total length; distance from tip of snout to anterior end of cloaca slightly more than two-fifths total length. Head and snout tapering from posterior end of jaw cartilages forward to a blunt, rounded point; head moderately flattened; snout in front of mouth short, its length a little more than one-half width of mouth; mouth moderately arched, its width somewhat less than half its length. Nasal flaps large and long, entirely covering posterior openings of nasal apertures and extending past upper lip, cover- ing depressed areas or channels in upper lip between nasal apertures and mouth; right and left nasal flaps confluent with upper lip and separated from one another by a distance about equal to length of fifth gill slit. Eyes moderately large, lateral, not visible from below, openings elongate, length more than two-thirds snout length; a prominent subocular gutter fully lined with denticles originating somewhat in advance of anterior eye corner and extend- ing under full length of eye to spiracle which is within the gutter at its posterior end; subocular gutter U-shaped in cross section and about as deep as wide; spiracles small, separated from eye opening by a distance slightly greater than spiracle diameter; gill slits moderate, the height of the first somewhat less than length of orbit, the first four subequal, the fifth somewhat shorter. Pectoral fins short, their distal margins broadly rounded, origins under or slightly posterior to fourth gill slit; first and second dorsal fins about equal in area, the base of the first shorter than the base of the second, their bases separated from one another by about twice the length of first dorsal base, both dorsal fins broadly rounded at the apex, their distal margins concave and their inner posterior tips angular; first dorsal origin over the posterior end of the pelvic base; second dorsal origin over or slightly in advance of the midpoint of the anal base; anal fin much lower than either dorsal fin, its area little more than one-half area of either dorsal; caudal fin narrow, no salient extension near origin of lower caudal, notch very strong. Proportional dimensions are given here as percentages of total length of the 475-mm adult male followed in parentheses by dimensions of the 510-mm female. Tip of snout to: front of mouth, 3.4, (2.9); eye, 4.0, (4.1); spiracle, 7.6, (7.1); first gill opening, 12.6, (12.0); fifth gill opening, 17.9, (17.6); origin pectoral, 16.8, (16.7); origin first dorsal, 44.2, (45.1); origin pelvics, 38.3, (39.2); origin second dorsal, 64.2, (64.9); origin anal, 62.1, (63.3); origin upper caudal lobe, 80.0, (80.0); cloacal opening, 41.3, (41.2). 33 Greatest width of: head, 9.3, (9.8); trunk at pectorals, 8.2, (8.8); trunk at pelvics, 6.5, (6.7); caudal peduncle, 2.7, (2.5). Greatest height of: head, 6.5, (6.5); trunk at pectorals, 6.9, (8.0); trunk at pelvics, 6.9, (8.2); caudal peduncle, 4.2, (3.9). Eyes: horizontal diameter, 2.5, (2.5); vertical diameter, 0.9, (0.6). Spiracles: greatest diameter, 0.4, (0.4); least distance from eye, 0.5, (0.5). Mouth: width, 5.7, (5.7); length mouth, 2.9, (2.2); length upper labial furrow, 2.5, (2.5); length lower labial furrow, 3.4, (3.1). Nasal apertures: distance between nasal flaps at inner posterior corner, 1.5, (1.5). Gill slits: height of first, 2.1, (2.0); height of fifth, 1.5, (1.6). First dorsal fin: length base, 6.7, (6.1); length inner posterior margin, 3.2, (2.7); height, 6.3, (2.9); note, distal margin of fin of female seems to have been nibbled away. Second dorsal: length base, 8.4, (7.8); length inner posterior margin, 3.2, (2.7); height, 6.3, (6.7). Anal fin: length base, 8.6, (7.8); length inner posterior margin, 2.5, (2.2); height, 3.2, (3.5). Caudal fin: length upper margin, 20.0, (20.0). Distance between fin bases: first and second dorsal, 13.7, (13.7); pectoral and pelvic, 18.7, (20.4); pelvic and anal, 18.1, (17.3); anal and lower caudal, 7.4, (7.3); second dorsal and upper caudal, 6.5, (7.5). Teeth very small, in about 70/70 rows, no indication of sexual dimorphism; teeth similar in upper and lower jaws, tridentate, the middle cusps much the longest (crowns of largest teeth in 475 mm male about 0.7 mm high), basal ridges of tooth crowns moderately strong; lateral 15 (more or less) rows on each side less than half size of central teeth. Dermal denticles close together, arranged almost in pavement, with thick denticle blades mostly with only one posteriorly directed point (in adults), but a weak lateral point on one or both sides of central point present on some den- ticles; points slightly elevated, rough to touch when rubbed toward head, smooth when rubbed toward tail; dorsal and lateral denticles with one to seven low ridges on anterior part of blade, ridges reduced or obsolete on ventral denticles, blade surfaces of denticles of snout and leading fin edges smooth; inside of mouth, tongue, and gill bars covered with very small, nearly erect, cone-shaped denticles separated from one another by distances greater than the diameter of their bases. No gill rakers present. Total number vertebrae, 172 in male, 167 in female; monospondylous 44 in both; precaudal 112 in both; caudal 60 in male, 55 in female. Claspers of 475-mm male (apparently an adult with well-calcified clasper cartilages) moderately robust, not notably long, extending 24 mm past tips of pelvic fins; no apron formed beneath claspers by partial union of inner margins of pelvics or by membranous connection of pelvics. Female of 510 mm with only right ovary functional, containing nine large eggs about 8 by 8 mm to 11 by 13 mm, the eggs arranged linearly so that the ovary extends nearly the full length of the body cavity; two oviducts, the left oviduct containing one egg capsule with enclosed egg; embryo, if present, not sufficiently developed to detect; the egg capsule about 66 by 21 mm, a lateral constriction at both anterior and posterior ends of capsule [as shown in Whitley (1939, fig. 3; 1940, fig. 87) J; anterior end of capsule without horny tendrils, posterior end with relatively short horny tendrils; liver not large or oily, confined to anterior half of body cavity; valvular intestine with about 18 turns. Color tan dorsally with numerous brown spots which coalesce dorsally to form short bars and posteriorly to form saddles; spots in irregular longitudinal lines on sides and extend onto all fins of female and all fins, except anal and lower caudal, of male. Ventral surfaces of body unspotted. Discussion. The adult male was collected where water depth was said to be IV2 ft (45 cm) and water temperature 95°F (35°C); the female was caught over a sand bottom with a few rocks from a depth not greater than 12 ft (3.7 m) and where the water temperature was 90°F (32.2°C). These records (Taylor 1964) are of much higher water temperatures than I have found for any other scyliorhinid and indicate that Atelomycterus must be adapted to an unusually warm and shallow-water environment. Apparently such environments are occupied by scyliorhinids only in the tropical Indo- Pacific area. One other scyliorhinid, Aulohalaelurus, of Western Australia and recorded from shallow water in the Arnhem Land area, also lives in warm, shallow water. While Aulohalaelurus has a relatively small eye, usual in shallow water, diurnal sharks, the eye of Atelomycterus is not as small as might be expected. The upper eyelid in Atelomycterus, however, appears to be capable of shading the eye making it less exposed to strong light than the eye of Aulohalaelurus. It seems likely that a study of the clasper structure might be useful in determining species of Atelomycterus. The clasper of Scy ilium marmoratum (A. marmoratus) illustrated by Leigh-Sharpe (1926, fig. 8) differs greatly from the clasper of A. macleayi not only in length and shape but also in the number and distribution of component elements. 34 Atelomycterus marmoratus (Bennett) Figure 19 J.1L* •» Figure 19.— Atelomycterus marmoratus, 437-mm female. Gulf of Thailand. Drawing by Mary Wagner. Scy Ilium marmoratum Bennett 1830:693 (type-locality, Sumatra); Giinther 1870:401. Atelomycterus marmoratus: Garman 1913:100; White 1936b:6; Fowler 1941:62, fig. 6; Teng 1962:53, Fig. 13; Chen 1963:32; McKay 1966:66, fig. 1; Besednov 1969:31, fig. 16. Material examined. VFC 516, ?, 437 mm, Gulf of Thailand, lat. 12°11'N, long. 100°19'E; SU 13562, ad. $, 615 mm, Philippines; SU 13659, 8, 475 mm, Philippines; SU 13689, S, 466 mm, Philippines. Holotype not seen, presumed lost. According to Giinther (1870) it is the skin of a half-grown specimen that was presented to BMNH by T. S. Raffles. Diagnosis. Atelomycterus marmoratus is readily separable from scyliorhinids of other genera by the characters noted under the generic diagnosis. The provisional distinction used here to separate A. marmoratus from A. macleayi is that in specimens of A. marmoratus the origin of the first dorsal fin is over the middle of the pelvic fin base whereas in A. macleayi it is over the posterior quarter of the pelvic fin base. Description. Following are measurements expressed as percentages of the total length for a 437-mm female A. marmoratus from the Gulf of Thailand. Tip of snout to: front of mouth, 3.2; eye, 4.6; spiracle, 7.8; first gill slit, 12.8; fifth gill slit, 18.1; origin pectoral fin, 15.6; origin first dorsal, 43.2; origin pelvics, 39.4; origin second dorsal, 66.4; origin anal, 62.2; origin upper caudal fin lobe, 80.5; anterior end cloacal opening, 41.2. Greatest width of trunk: at pectoral origin, 8.9. Greatest height of trunk: at pectoral origin, 6.9. Orbit: horizontal diameter, 3.0; vertical diameter, 0.7. Mouth: width, 5.7; length upper labial furrow, 2.5; length lower labial furrow, 2.7. Gill slits: height of first, 2.1; height of fifth, 1.6. First dorsal fin: length base, 7.3; length free inner margin, 3.0; height, 6.9. Second dorsal fin: length base, 8.2; length free inner margin, 2.3; height, 5.7. Anal fin: length base, 7.6; length free inner margin, 1.6; height, 3.4. Caudal fin: length upper margin, 20.6; anterior margin lower lobe, 7.3. Distance between fin bases: first and second dorsals, 15.1; pectoral and pelvic, 17.6; second dorsal and caudal, 6.4; pelvic and lower caudal, 34.8. Teeth small, in about 74/74 rows, lateral 10 to 12 similar in upper and lower jaws, lateral 10 or 12 rows on each side with very short cusps not readily counted or seen, remainder of teeth tridentate, the middle cusp much the longest. Vertebral numbers in four specimens examined were: monospondylous 44 to 47; total, 157-162. Discussion. The color pattern in A. marmoratus is remarkably variable. The extent of variation in pattern can be seen by comparing the drawing here (Fig. 19) of a specimen from the Gulf of Thailand; three specimens shown by Fowler (1941, fig. 6), possibly from the Philippines; two specimens from Western Australia (McKay 1966, fig. 1); and a specimen illustrated by Besednov (1969, fig. 16), from the Gulf of Tonkin. A ULOHALAELURUS FOWLER, 1934 Aulohalaelurus Fowler 1934:237 (type-species Catulus labiosus Waite, by original designation). 35 Diagnosis. Aulohalaelurus has a well-developed supraorbital crest of the chondrocranium that extends as a narrow shelf above the orbit. It resembles A telomycterus in this as well as in its elongate, somewhat cylindrical body form, in its fin positions, in its conspicuous, long, labial furrows, and in the absence of an apron formed by the partial union of the pelvic fins beneath the claspers of males. In Aulohalaelurus, however, the nasal flaps do not quite reach the mouth and Aulohalaelurus has no depressed sections of the upper lip that connect the nasal cavity with the mouth as in Atelomycterus. Aulohalaelurus also differs from Atelomycterus in having a smaller eye with much less development of a subocular gutter and a secondary lower eyelid. In direct comparison of an adult male Aulohalaelurus labiosus with an adult male Atelomycterus macleayi the follow- ing additional differences were noted. In Aulohalaelurus the spiracle was proportionally larger and the teeth were proportionally much larger. The lower jaw teeth were larger than the upper jaw teeth in Aulohalaelurus; the reverse is true of Atelomycterus and other scyliorhinids except Schroederichthys bivius adult males. Aulohalaelurus has scattered black spots on the belly whereas Atelomycterus does not. Aulohalaelurus labiosus (Waite 1905) Catulus labiosus Waite 1905:57, fig. 23, Freemantle, Australia (holotype, a 620 mm ' WAM). Halaelurus labiosus: Garman 1913:88; Fowler 1941:51; McKay 1966:68. Aulohalaelurus labiosus: Whitley 1940:89, figs. 78, 80. Material examined. WAM P. 12020, ad. J, 540 mm, Point Peron, Freemantle, Western Australia, 3.7 m, 1 March 1965. Diagnosis. Aulohalaelurus labiosus is most easily separated from other scyliorhinids by the characters given under the diagnosis of the genus. Description. The following proportional measurements are in percentages of total length of a 540-mm adult male . Tip of snout to: posterior end of nasal flap, 3.06; front of mouth, 3.14; eye, 5.1; spiracle, 8.0; first gill slit, 13.9; fifth gill slit, 18.5; origin pectoral, 17.8; origin first dorsal, 49.3; origin pelvics, 43.5; origin second dorsal, 66.9; origin anal, 63.2; origin upper caudal lobe, 79.6; anus, 46.3. Eyes: horizontal diameter, 1.9; distance between rims of upper eyelids, 5.7. Spiracles: greatest diameter, 0.7; least distance from eye, 1.0; distance between, 6.5. Mouth: width, 7.4; length upper labial furrow, 3.3; length lower labial furrow, 3.7. Nasal apertures: level of anterior opening to tip of snout, 2.0; level of posterior opening to tip of snout, 3.1; least distance between, 2.3. Gill slits: height of first, 2.2; height of fifth, 1.1. First dorsal fin: length base, 6.9; inner posterior margin, 3.1; height, 5.6. Second dorsal: length base, 8.3; inner posterior margin, 2.8; height, 5.2. Anal: length base, 8.1; inner posterior margin, 2.0; height, 3.0. Pectorals: width base, 4.3; anterior margin, 11.3; greatest width, 8.1. Pelvics: length (origin to rear tip), 10.0. Distance between bases: first and second dorsals, 10.4; pectoral and pelvic, 20.4; pelvic and anal, 13.5; anal and lower caudal origin, 6.9; second dorsal and upper caudal, 4.8. Greatest width of: head, 11.5; trunk at pectorals, 10.0; trunk at pelvics, 7.6; trunk at caudal base, 2.6. Greatest height of: head at spiracles, 6.9; trunk at pectorals, 9.4; trunk at pelvics, 8.7; trunk at caudal base, 3.9. Aulohalaelurus labiosus is a species of moderate size, the holotype was 620 mm, the largest specimen recorded is a 673- mm female, WAM P. 5690 (McKay 1966), and the specimen on which this description is based, a 540-mm mature male. Body elongate, subcylindrical; head somewhat flattened, its length less than one-fourth total length; snout short with a rounded point; nasal flaps large, completely covering posterior nasal openings, posterior edges nearly reaching upper lip; visceral cavity moderately long; pectorals short, their tips rounded; dorsal fins nearly equal in area, origin of the first over posterior end of pelvic base, origin of second over middle of anal base; anal base about as long as second dorsal base; caudal fin short, its length from upper caudal fin origin about one-fifth total length, lower caudal fin low, with projecting lobe. Mouth proportionally large with long and prominent labial furrows, their lengths more than 1 l /s times the length of the eye. Color of 540-mm male brownish-gray on dorsal and lateral surfaces, yellowish-white below head and on belly, with roundish black spots 5 to 10 mm in diameter in generally random arrangement over all surfaces except the gular area; and with a few light-colored, small, and somewhat indistinct spots over dorsal surfaces, more notable posteriorly, small white areas at tips of dorsal and caudal fins. 36 Teeth of upper and lower jaws generally similar in shape, mostly tricuspid in the central part of the jaws, the middle cusp much the longest; teeth of the central part of the lower jaw about twice as high as corresponding teeth of the upper jaw; extreme lateral teeth of both jaws very small, with five or more short cusps, and about five series functional; tooth formula, about 50/45; highest tooth of lower jaw about 3.4 mm, highest tooth of upper jaw about 1.9 mm. Dermal denticles over most dorsal surfaces imbricate, with three posteriorly directed points, the middle point largest; blades with a heavy central ridge, occasionally double, but no lateral ridges; denticles near midline of back somewhat more erect than on flanks; belly denticles with lateral points reduced or absent; length of blades of larger denticles about 0.7 to 1.0 mm. Claspers of mature male project less than half their length past the tips of the pelvic fins. The pelvic fins are not united to form an apron or partial apron beneath the claspers. Discussion. Aulohalaelurus labiosus appears to be authentically recorded only from Western Australia (see McKay 1966). The specimen described here was collected in 2 fathoms (3.6 m), unusually shallow water for a scyliorhinid in most parts of the world. The eyes of A. labiosus are small suggesting also a well-lighted (shallow-water) habitat. It has not been determined whether the species is oviparous, and egg capsules, if formed, are not known. Aulohalaelurus labiosus was illustrated by Whitley (1940:78 and 80) and by Waite (1905, fig. 23). CEPHALOSCYLLIUM GILL Cephaloscyllium Gill 1862:407, 408 (type-species, Scyllium laticeps Dumeril 1853, by original designation). Diagnosis. The presence of a supraorbital crest of the chondrocranium in combination with the lack of either upper or lower labial furrows distinguishes Cephaloscyllium from all other scyliorhinid genera. Species of Cephaloscyllium are robust with wide heads and short snouts. They are unique among sharks in having the ability to inflate their stomachs with water (or air) to give a balloonlike shape to the body. Discussion. Cephaloscyllium changes greatly in body proportions during growth. For example, in a young specimen of C. ventriosum at hand, the distance from the tip of the snout to the cloaca is 45% TL, but in an adult the distance is 62% TL. Measurements given by Scott (1963) for C. isabella laticeps also indicate great proportional differences between young and adults; eye length, for example, is 4.7 to 6.2% TL in six adults but 7.4% in a foetus 116 mm long. Jordan and Fowler (1903) characterized C. umbratile from Japan as having the space between the first and second dorsals much greater than the length of the first dorsal base. In eight immature C. umbratile that I examined, the interdorsal space is only slightly longer than the length of the first dorsal base. Immature specimens reported by Chen (1963) and Besednov (1969) were also characterized in this way. Garman (1913) stated that the length of the first dorsal base is a little longer than the space between the dorsal fins in C. umbratile, but his key to the genus on a preceding page contradicts the statement. The type of C. umbratile was noted by Jordan and Fowler (1903) to be a dried skin, SU 12693, and specific mention was not made of another C. umbratile. Their illustration of C. umbratile appears from the clasper length to be an adult and shows the interdorsal distance slightly less than the length of the first dorsal base. In summary, the preceding details of measurement, and others from the literature not mentioned here, suggest that in immature ex- amples of all species of Cephaloscyllium the interdorsal distance is usually greater than length of the first dorsal base, but that in adults the length of the first dorsal base is usually equal to or slightly longer than the interdorsal distance. In addition, the small number of specimens that I have seen suggest that body proportions vary considerably even within a single size group. The dermal denticles of Cephaloscyllium species also change in shape and number during growth. Typical flank den- ticles are erect and needlelike in young specimens but broader and not very erect in adults. Whitley's (1940) descriptive account and illustration of C. laticeps of Australia was obviously based on young specimens, "usually about one foot," and his description of New Zealand C. isabella was based on adults. The number of tooth rows in either upper or lower jaws of specimens that I have examined varied from about 55 to 70. 1 have no explanation for Garman's (1913:81) count of 118/124 for C. umbratile. The adults of various species of Cephaloscyllium appear to reach maturity at lengths of about 450 to 900 mm depend- ing on both species and sex. New Zealand specimens, as already noted, are said to reach a length of 8 ft (2.45 m). I have not been able to verify the existence of any specimen longer than about 1 V2 m. I here recognize five species. One, C. fasciatum, is highly distinctive in color pattern and readily separable from the other four species by other characters. The remaining four species are very near one another in body proportions and all appear either to have been originally described because of geographic separation or on the basis of size or sex differences. On the basis of color differences and some other minor differences in the following key I recognize these five species as probably distinct. Adult specimens have not been available for my study in satisfactory numbers, and it is likely that I have failed to find or observe some characters that would take C. umbratile out of the synonymy of C. isabella or to find some character that would better distinguish C. laticeps and C. isabella than the structure of the egg capsule. 37 The inflation of the body by swallowing great quantities of water probably enables Cephaloscy ilium to wedge itself in crevasses or small rocky caves and thus avoid some predators. When taken from the water, specimens may gulp air. The noise reported by fishermen to be like a barking dog (Whitley 1940), is the result of expulsion of air. Key to Species of Cephaloscyllium la. No definite color pattern of dorsal spots, lines, or blotches either in young or adults, but faintly indicated darker smudges with indefinite outlines occasionally visible in position of dorsal saddle blotches C. sufflans lb. A definite color pattern present, usually more prominent or regular in young specimens 2 2a. A prominent pattern of dark lines outlining quadrilateral, dorsal, saddle areas; sometimes also a few irregularly scattered dark spots C. fasciatum 2b. A pattern of spots or blotches but no dark lines 3 3a. Posterior margins of nasal flaps extending to edge of upper lip C. ventriosum 3b. Posterior margins of nasal flaps not reaching edge of upper lips 4 4a. Egg capsules with strong transverse ridges C. laticeps 4b. Egg capsules without transverse ridges C. Isabella Cephaloscyllium fasciatum Chan Figures 20, 21 r~ * h **$/* Figure 20. — Cephaloscyllium fasciatum, newly hatched female about 1 2H mm, BMNH 1925.9.18, Philippines. Figure 21 .—Cephaloscyllium fasciatum, egg capsules about N.'S mm long by .'!.! mm wide, found attached to the Hong Kong-Manila cable in .'!l(l m near Manila. 38 Cephaloscyllium fasciatum Chan 1966:232, figs. 6 and 7 (off Cape Bantagan, Vietnam). Material examined. BMNH 1965.8.11.1, ? (maturing), 422 mm, holotype, 35 mi ESE of Cape Bantagan, Vietnam, in 219-314 m on soft mud bottom; 2 S, 235, 245 mm, paratypes, same locality as holotype; BMNH 1925.9.18.1, ?, 128 mm (hatchling), lat. 14°24'N, long. 120°15'E, with two egg capsules, attached to Hong Kong-Manila cable in about 310 m, Figs. 20, 21. Diagnosis. Cephaloscyllium fasciatum was distinguished in the original description by the presence of a lobelike extension of the nasal flap, its inner margin with a deep notch. That character may not always be sufficient since one C. umbratile of several specimens examined had a lobed nasal flap. The color pattern, however, is distinctive, and the presence of dark bordered saddle blotches or spots should readily distinguish C. fasciatum from all scyliorhinids except Scyliorhinus retifer of the western Atlantic. Scyliorhinus retifer has lower labial furrows and so differs from any species of Cephaloscyllium. Discussion. Chan's (1966:232, table 1) comprehensive description of the species is accompanied by proportional measurements and details will not be repeated here. His account did not mention the presence of two rows of enlarged denticles on the backs of the smaller specimens of the type-series. The additional specimen that I examined, a hatchling 128 mm long, did not have the enlarged denticles so I assume they are not present on the young of this species. Such den- ticles are present, although perhaps only briefly, at hatching size on the young of C. Isabella, C. umbratile, and C. ventriosum. I do not know whether young C. suf flans have these denticles. Possibly the absence of the enlarged denticles on C. fasciatum has a connection with a deeper habitat and the thinner walled egg capsules of C. fasciatum. For discus- sions of this see under C. ventriosum and also under Halaelurus boesemani. Cephaloscyllium isabella (Bonnaterre) Squalus isabella Bonnaterre 1788:6 (La mer du sud). Cephaloscyllium umbratile Jordan and Fowler 1903:602, fig. 1 (Nagasaki, Japan); Garman 1913:80; Fowler 1941:32; Chen 1963:29, fig. 9; Lindberg and Legeza 1959:44, fig. 24; Chan 1966:229, figs. 4, 5, 7b, 7d; Besednov 1969:27, figs. 9, 10. Cephaloscyllium laticeps (not of Dumeril): Waite 1909:136, pis. xiv - 1, xx - 1. Cephaloscyllium isabellum: Garman 1913:79; Fowler 1941:31; 1967:358. Cephaloscyllium formosanum Teng 1962:48, fig. 11. Cephaloscyllium isabella nascione Whitley 1932:323, fig. 2, no. 2. Cephaloscyllium isabella: Whitley 1940:91, fig. 84. Material examined. USNM 176779, subad. $, 630 mm, Auckland, New Zealand; USNM 176795, ad. i, 690 mm, Cook Strait, N.Z.; USNM 176801, ad. v, 855 mm, Cook Strait, N.Z.; FMNH-Owston No. 1138, imm. ?, imm. i, 465, 382, 335 mm, Boshu, Japan; FMNH-Owston No. 836, imm. ?, 335 mm., Sagami Sea, Japan; FMNH-Owston No. 932, imm. $, 303 mm, imm. ?, 268 mm, Idzu, Japan; FMNH-Owston 1351, imm. ?, 365 mm, imm. S, 313, 326 mm, Yokahama Market, Japan. Types. The holotype of Squalus isabella Bonnaterre was not located. The holotype of Cephaloscyllium umbratile Jordan and Fowler, SU 12693, a dried skin, was not found after search and is presumed lost. I did not see the holotype of Cephaloscyllium formosanum Teng, a 655-mm female, TFI 4339, taken off Tungkang, Formosa (Taiwan), in about 374 m, or the holotype of Cephaloscyllium isabella nascione Whitley, AMS - IA 2829. Diagnosis. Cephaloscyllium isabella has a color pattern of irregularly arranged darker spots and saddle blotches, none of which have the black marginal lines characteristic of C. fasciatum. Also, C. isabella has broader and usually shorter nasal flaps than C. fasciatum, and egg capsules with thicker walls. Young isabella up to at least 226 mm long may have two rows of enlarged denticles on the back, but such denticles apparently do not appear on young C. fasciatum. Separation of C. isabella from other species of Cephaloscyllium is much more difficult on the basis of characters observed in the few specimens that I examined. The South African Cephaloscyllium suf flans lacked the prominent color markings of isabella but I have not been able to find other differences. The length of the snout (in front of mouth) is somewhat shorter and the eye somewhat smaller in the adult C. isabella from New Zealand than in C. sufflans. The eastern Pacific C. ventriosum is usually distinguished from isabella by nasal flaps that reach to or past the edge of the upper lip. Although nasal flaps in isabella were somewhat variable in shape, in the specimens I examined they were all short, and their edges did not reach the upper lip margin. Cephaloscyllium isabella lays smooth-surfaced egg capsules but otherwise differs little from C. laticeps which lays egg capsules with strong transverse ridges. 39 Description. Proportional dimensions as percentages of the total length are given following: first, for a 690-mm adult male, USNM 176795; second, for an adult female, USNM 176801; and third in each series of figures for an immature female, FMNH-Owston 1138. Tip of snout to: front of mouth, 3.5, 3.9, 3.4; eye, 6.4, 6.1, 5.6; spiracle, 11.0, 11.0, 9.7; first gill slit, 18.1, 17.3, 13.8; last gill slit, 22.2, 21.3, 17.6; origin pectoral fin, 19.1, 18.0, 16.6; origin first dorsal, 50.7, 51.5, 49.5; origin pelvics, 44.9, 46.2, 45.2; origin second dorsal, 65.2, 66.7, 64.7; origin anal, 63.7, 64.3, 63.7; anterior end cloacal opening, 49.3, 51.5, 48.4. Greatest width of: trunk at pectoral origin, 16.5, 18.1, 14.6; trunk at pelvic origin, 9.1, 9.8, 6.5; trunk at caudal origin, 2.2, 2.2, 2.4. Greatest height of: trunk at pectoral origin, 9.9, 14.0, 10.8; trunk at pelvic origin, 9.9, 9.4, 7.7; trunk at caudal base, 3.3, 2.8, 3.0. Eyes: horizontal diameter of opening, 3.5, 3.4, 3.2; height of opening, 8.4, 0.6, 0.9. Spiracles: greatest diameter, 0.9, 0.7, 0.2; least distance from eye opening, 1.2, 1.1, 1.3. Mouth: width, 11.7, 11.8, 10.5; length, 6.8, 4.8, 4.5. Nasal apertures: level of anterior ends to tip of snout, 2.5, 2.6, 1.9; level of posterior ends to front of mouth, 0.3, 0.3, 0.2; minimum distance between, 3.2, 3.5, 2.4. Gill slits: height of first, 2.2, 2.5, 1.7; height of fifth, 1.7, 2.0,0.5. First dorsal fin: length base, 8.7, 9.6, 8.0; length posterior tip, 2.6, 2.3, 2.6; height, 5.5, 5.8, 4.5; length anterior margin, 10.1, 11.8, 9.0. Second dorsal fin: length base, 6.1, 6.4, 4.3; length posterior tip, 2.5, 2.3, 2.6; height, 3.3, 3.2, 4.5. Anal fin: length base, 7.0, 7.4, 6.7; length posterior tip, 2.9, 2.5, 2.6; height, 4.3, 4.1, 3.0; length anterior margin, 9.0, 8.3, 6.5. Pectoral fins: width base, 10.1, 11.2, 6.9; length anterior margin, 15.8, 17.7, 13.3; greatest width, 14.1, 13.6, 10.5. Pelvic fins: origin to rear tip, 14.2, 12.3, 9.7. Caudal fin: length upper margin, 22.0, 21.3, 22.4. Distance between fin bases: first and second dorsals, 7.8, 7.5, 8.6; pectorals and pelvics, 14.8, 17.4, — ; pelvics and anal, 11.4, 9.2, — ; anal and lower caudal, 4.6, 4.6, — ; second dorsal and upper caudal, 6.5, 5.4, — . Teeth are small, highest tooth in 465 mm female 1.2 mm, similar in upper and lower jaws, some teeth near the middle of each jaw three-cusped but most lateral teeth four- or five-cusped, their bases with ridges with a few having ridges ex- tending nearly to tooth tips. Number of tooth rows (rows perpendicular to jaw axis) about 50 to 70 in the upper jaw and about 45 to 65 in the lower jaw. Dermal denticles erect and narrow in young becoming wider with blades heavy and parallel to skin surface in adults. Hatchlings with two rows of enlarged denticles on back probably similar in structure and function to those described un- der C. ventriosum. Vertebral numbers counted by X-ray were: USNM 176795, total 120, monospondylous 47, precaudal 79, caudal 41; USNM 176801, total about 116, monospondylous 45, precaudal 76, caudal about 40; USNM 176779, total about 118, monospondylous 48, precaudal 78, caudal about 40. The egg capsule of C. Isabella from New Zealand illustrated by Waite (1909, pi. xxi, fig. 1) has a smooth surface and is similar in shape to the egg capsule of C. ventriosum (Fig. 26). Discussion. Except for C. fasciatum, which has highly distinctive color markings, the species of Cephaloscy Ilium appear similar and are consequently distinguished by minor differences. These differences have not been checked with large numbers of specimens, however, and the treatment here should be regarded as tentative. In earlier accounts of species of Cephaloscy Ilium it seems that insufficient attention was given to changes in body and fin proportions and to changes in dermal denticle structure during growth. Earlier keys generally separate Isabella, laticeps, and umbratile from one another by attributing juvenile characters to some and adult characters to others. This should be easily cor- rected but will require a greater amount of data than have been available to me. I can find no confirmation of the existence of very large New Zealand specimens of Cephaloscyllium and assume that lengths to 8 ft attributed to the genus are based on misidentifications. Waite (1909:136) and McCulloch (1911:6) referred to Cephaloscyllium as a "carpet shark," a common name often applied to larger species of the shark family Orec- tolobidae in Australia. It seems obvious that Whitley's account and illustration of C. Isabella (1940:91, fig. 84) are based on large specimens, mature or nearly so, whereas his description and illustration of C. laticeps (1940:92, fig. 85) are taken from small and immature specimens. Whitley (1932:323) seems to have been first to note that egg cases of Cephaloscyl- lium from southeastern coasts of Australia were "flanged, not smooth" but other characters he used to characterize the species were based on juvenile specimens. It appears from Whitley's account (1932:323) that both C. isabella and C. laticeps may occur in Australian waters with C. laticeps restricted to the southeastern coast and being more common off Tasmania. 40 Cephaloscyllium laticeps (Dumeril) Figure 22 Figure 22.— Cephaloscyllium laticeps, BMNH 1849.11.2.8-9, George- town, Tasmania. The function of the transverse ridges is unknown but a possible explanation for heavy-walled egg capsules is discussed under C. ventriosum. Scy Ilium laticeps Dumeril 1853:84 (coasts of New Holland = Tasmania); Giinther 1870:404 (Tasmania). Cephaloscyllium laticeps: Whitley 1940, part 92, fig. 85, no. la (Tasmanian form), also fig. 30, no. 14. Cephaloscyllium isabella laticeps: Whitley 1932:323, fig. 2, no. 1; Scott 1963:3, fig. 1 (Tasmania). Material examined. None except dried egg capsules: BMNH 1849.11.2.8,9, Georgetown (probably Georgetown, Tasmania); similar egg capsules in BMNH marked unknown locality. Diagnosis. I am unable to find differences in accounts of Cephaloscyllium morphology between C. laticeps and C. isabella that are consistent with all of the material and data that are available to me. The two forms are separated here solely upon the differences in egg capsule form (Fig. 22). Cephaloscyllium laticeps produces egg capsules that are flanged (Whitley 1932:323) or having about 31 to 33 strong transverse ridges in Tasmanian waters (Scott 1963:5). Egg capsules from Tasmania in the British Museum have about 25 to 28 transverse ridges and no doubt the number of ridges varies considerably. Whitley (1940:43, fig. 30, no. 14) illustrated a similar "laminated" egg capsule from Victoria and referred to several from the Bass Strait area without species identification. What is known of the distribution of egg capsules sug- gests that C. laticeps is restricted to Tasmanian waters and to the nearby Australian shores. Cephaloscyllium isabella, C. ventriosum, and C. fasciatum all have smooth-surfaced egg capsules. I have not found a description of egg capsules of C. sufflans. Discussion. Egg capsules of Cephaloscyllium species are discussed in more detail under C. ventriosum. Several authors have indicated that New Zealand Cephaloscyllium grow to a larger size than do Tasmanian Cephaloscyllium . I am not able to verify this and suspect that the larger size attributed to some New Zealand specimens is in error. Scott (1963) included a table of measurements of seven Cephaloscyllium 885 to 961 mm and a 116-mm foetus from Tasmania. His illustration of the young one before hatching shows the two rows of enlarged denticles on the back as in hatchlings of other species of Cephaloscyllium except C. fasciatum. Cephaloscyllium sufflans (Regan) Scyliorhinus sufflans Regan 1921:413 (coast of Natal in 219 to 238 m). Scylliorhinus sufflans: Barnard 1925:41. Cephaloscyllium sufflans: Smith 1949:52, fig. 34; Bass et al. 1975:9, fig. 6. Material examined. USNM 201910, imm. 3, 345 mm, off Tugels River, South Africa; USNM SS 610, imm. S, 200 mm, off Kenya, east Africa, lat. 02°50'S, long. 40°31'E, in 290 m, 8 November 1964. 41 Diagnosis. The lack of dark markings may serve to separate C. sufflans from other species of Cephaloscyllium at all sizes. The two small specimens of C. sufflans examined show no trace of markings and small specimens of other species have prominent markings. Regan's (1921) description of the 750-mm holotype of C. sufflans includes the statement, "pectoral . . . , extending not quite half the distance from its origin to that of the pelvics." In the New Zealand C. Isabella, 630 to 855 mm, at hand the pectoral extends considerably more than half the distance. Description. Regan's (1921:413) description of C. sufflans, based on his only specimen, the 750-mm holotype, includes no information that I find useful to separate it from other species except that he stated the specimen was without distinct spots or markings. All other Cephaloscyllium species have markings or spots although they sometimes may be diffuse. Both C. sufflans specimens that I examined were immature without distinct markings. Both had slender and more or less erect denticles that gave the specimens a somewhat shaggy appearance. Vertebral numbers in the 345-mm male, USNM 201910, as shown by X-ray were: total number about 125; monospon- dylous 49; precaudal 85; caudal about 40. Bass et al. (1975:9, table 2) provided a thorough analysis of variation in proportional dimensions of C. sufflans based on measurements of 41 males and 61 females following methods proposed by Bass (1973). Comparisons of these measure- ments of C. sufflans with measurements, similarly treated, of large series of C. isabella subspecies and regional popula- tions of C. isabella should reveal morphometric and growth characters to separate these forms if any such characters ex- ist. Cephaloscyllium ventriosum (Garman) Figures 23, 24, 25 y -"-■■'•■ Figure 2.'!.— Cephaloscyllium ventriosum, 400-mm male, California, SU L9090. Drawing by Mary Wagner. Figure 24. — Cephaloscyllium ventriosum, newly hatched male, 15N mm, Monterey, Calif. 42 Figure 25.-Egg capsule of Cephaloscyllium ventriosum, with embryo. Photograph courtesy of Marineland of the Pacific, Palos Verdes, Calif. 43 Scy Ilium uentriosum Garman 1880:167 (Valparaiso, Chile). Catulus uter Jordan and Gilbert in Jordan and Evermann 1896:25, pi. 3, fig. 12 (Santa Barbara Channel, Calif.). Scyliorhinus ventriosus. Regan 1908a:458. Cephaloscyllium uentriosum: Garman 1913:80, pi. 9, figs. 6-9; Kato et al. 1967:24, fig. 36; Nelson and Johnson 1970:732; Grover 1972a: 191; 1974:359, figs. 1-3. Cephaloscyllium uter: Roedel and Ripley 1950:50, fig. 34; Cox 1963:283, fig. 11. Material examined. USNM 52853. ad. S, 820 mm, California, 1895; USNM uncat.. imm. 6, 158 mm, Monterey Bay, Calif., 46 m, 4 February 1964; USNM 196142, imm. 3, 550 mm, Avila Bay, Calif.; USNM 25084, 26866, ad. 8, ca. 850 mm, Santa Barbara, Calif. Holotype not seen. Diagnosis. The posterior margins of the nasal flaps of C. uentriosum reach to or past the upper lip and the species thus differs from all other species of Cephaloscyllium (see insert, Fig. 23). Description. A robust species reaching a length of about a meter and like other species of Cephaloscyllium, remarkable in its ability to distend the stomach by swallowing water (or air), swelling the elastic skin of the trunk into a nearly spherical shape. Head flattened, very broad, its length about one-fifth total length, its breadth nearly as great as its length to first gill slit; mouth very wide, its width greater than length of first dorsal base; snout very short, its length from front of mouth not much greater than least distance between nasal apertures; eyes small, their horizontal diameter in adults about equal to least distance between nasal apertures, with a moderately prominent subocular gutter under the full length of the eye but not reaching small spiracle close behind eye; gill slits short, the fifth somewhat shorter than the first. Fins increasing relatively in size with growth; pectorals much larger than second dorsal and anal fins in area; distance between first and second dorsal fin bases slightly greater than length of first dorsal base in young, less than length of first dorsal base in adults; origin of first dorsal over or in advance of midpoint of pelvic base; origin of second dorsal over or slightly posterior to anal origin; lower caudal fin moderately broad but lacking an anterior salient point; caudal notch prominent. Teeth relatively small, numerous in about 55/55 to 60/60 rows, tricuspid, a few with additional small cusps, the middle cusp much the strongest, similar in upper and lower jaws, no great reduction in tooth size toward jaw angles. Proportional dimensions of an 820-mm adult male in percentages of total length are given here, each followed in parentheses by similar dimensions for a newly hatched 158-mm male. Tip of snout to: posterior margin nasal flap, 3.2, (3.2); front of mouth, 3.0 (3.2); eye, 5.0, (5.7); first gill opening, 16.3, (15.8); fifth gill opening, 22.2, (20.3); origin pectoral, 21.0, (20.3); origin first dorsal, 52.4, (46.2); pelvic origin, 57.3, (41.5); anus, 61.6, (45.3); second dorsal origin upper caudal fin, 78.7, (72.2). Eye: length orbit, 2.8, (4.1). Mouth: width, 9.9, (10.1); length 4.9, (4.1). Gill slits: height of first, 1.7, (2.2); height of fifth, 1.5, (1.9). First dorsal fin: length base, 8.8, (7.0); length posterior inner margin, 3.8, (3.2); height, 7.1, (5.7). Second dorsal fin: length base, 5.9, (5.7); length posterior inner margin, 2.8, (2.2); height, 3.4, (3.2). Anal fin: length base, 7.8, (8.2); height, 4.5, (3.2). Pectoral fin: width base, 8.7, (7.6); anterior margin, 17.1, (13.3); greatest width, 12.2, (9.8). Distance between fin bases: first and second dorsal, 8.0, (7.6); pectoral and pelvic, 17.9, (15.5); pelvic and anal, 11.8, (9.5); anal and lower caudal, 5.1, (5.1); second dorsal and upper caudal, 5.4, (4.4). Dermal denticles of sides of body and over most surfaces erect and needle-shaped with no lateral expansion of a blade in juveniles; denticles of adults with a stout blade, more or less triangular with a single posterior point; denticle blades in adults nearly parallel to skin surface in ventral areas, more erect to angle of about 45° in dorsal areas; ventral denticles with smooth surfaces without ridges, dorsal denticles with one or several ridges; dark spots on adults apparently due chiefly to pigment incorporated within denticles. (Forms of dermal denticles of adults illustrated in Garman 1913, pi. 9, fig. 9.) Newly hatched young have two rows of much enlarged denticles extending along each side of the middorsal line from the level of the first gill slit to the level of the origin of the first dorsal fin. Each row consists of about 28 uniformly spaced denticles. These large denticles have stout, paddle-shaped and sharp-edged blades on very heavy and broad-based pedicles. Apparently these denticles are shed soon after hatching. Vertebral numbers for 820 mm adult male: total 111, monospondylous 39, precaudal 70, caudal 41. Color in newly hatched specimen, yellowish-white with seven or eight light-brown, saddlelike areas across dorsal and lateral surfaces, and numerous nearly round, dark-brown spots about the size of the eye, more or less randomly dis- tributed over body surfaces and fins. Color in adults similar but saddles and spots less well defined, and all surfaces darker. 44 Discussion. All of the specimens of C. ventriosum I have seen were from California where the species is regularly taken. The type-locality of C. ventriosum is Valparaiso, Chile, suggesting that the species has either an antitropical dis- tribution or that the absence of records from the tropical eastern Pacific is due to lack of collecting effort in that area. If C. ventriosum usually spends daylight hours in crevices of rocky reefs as indicated by Grover (1972a) and moves about feeding mostly at night (Nelson and Johnson 1970), it can easily have been missed in collections depending on daylight trawling or daytime handline fishing. Nelson and Johnson (1970) observed C. ventriosum commonly at depths from 9 to 30 m around Santa Catalina Island, Calif., but noted that others had reported California specimens to depths of 300 m. In their observations, Nelson and Johnson found C. ventriosum to be bottom dwellers preferring the rocky, algae-covered slopes of kelp-bed communities, and active only at night. Their experimental study found C. ventriosum to exhibit an endogenous rhythm compared with an exogenous rhythm indicated for the horn shark, Heterodontus francisci, studied in the same experimental series. Cephaloscy Ilium ventriosum is oviparous laying eggs in capsules which, according to Cox (1963:283, fig. 11), range from 90 to 125 mm in length and 28 to 55 mm in width. The capsules have a moderately heavy wall (Fig. 26) with smooth surfaces but with some thickening for reinforcement along each lateral margin. The young are 140 to 150 mm at hatching (Grover 1974:360) and at that time have two rows of enlarged denticles along the back. These denticles, accord- ing to Grover's detailed observations, are employed by the hatchling to escape from the relatively stiff-walled egg cap- sule using a ratchet and pawl mechanical arrangement to pull itself through the compressing walls of the egg capsule (see Grover 1974, fig. 3). In an earlier paper, Grover (1972b:871) reported evidence that predators, both mollusks and teleosts, might cut through egg capsule walls to destroy eggs or young of C. ventriosum and noted that this supported the view that egg cap- sules in elasmobranchs evolved for protection rather than for more effective osmoregulation. My overview of Cephaloscy Ilium species and other scyliorhinids seems to support the idea that protection is the primary function of egg capsules. Grover (1972a: 191) found differences in the egg capsule, tendril length, the size of egg capsules, and the relative fin sizes between the C. ventriosum population surrounding Santa Catalina Island and the population inhabiting shores of nearby California. He had sufficient material to show that these differences were significant. While my data are inade- quate to be conclusive, the separation of the largely Tasmanian form, C. laticeps, from the more extensively distributed C. isabella is similar in being based on egg capsule differences. The Cephaloscyllium species that are known to me to have egg capsules with thick walls, C. isabella, C. laticeps, and C. ventriosum, also have enlarged dorsal denticles during the hatching stage, presumably in each instance to assist the hatchling to escape from the somewhat inflexible walls of the egg capsule. Cephaloscyllium fasciatum, on the other hand, has egg capsules with thin, weak walls and apparently does not develop enlarged dorsal denticles during the hatching stage. CEPHALURUS BIGELOW AND SCHROEDER Bigelow and Schroeder 1941:73 (type-species, Catulus cephalus Gilbert 1892, by original designation). Diagnosis. Cephalurus has a wide, flattened head with a short, broadly rounded snout and a very wide mouth. The branchial region is especially long and wide, with widely spaced gill slits. Cephalurus has a short abdominal trunk which tapers to a slender and moderately long tail. As viewed from above, the shark has a tadpolelike outline that readily dis- tinguishes it from all other scyliorhinids. It is also the only genus of the Scyliorhinidae with the first dorsal fin origin in advance of the origin of the pelvic fins. The skin is loosely adherent to the body and the body is soft as in many sharks in- habiting moderately deep continental slope waters. Juvenile Cephalurus have denticles of the upper edge of the caudal fin near its origin arranged much as in juvenile Parmaturus xaniurus, but development of a prominent caudal crest of somewhat enlarged and modified denticles does not occur in Cephalurus as it does in Parmaturus. Discussion. The young of Cephalurus develop in very thin-walled egg capsules within the oviducts of the mother. Probably the egg capsules are ruptured or absorbed and free-swimming young are born at a length somewhat less than 100 mm. Adults reach a length of about 300 mm. Cephalurus cephalus (Gilbert) Figure 26 Catulus cephalus Gilbert 1892:541 (from vicinity of Clarion Island, Revillagigedo Islands); Jordan and Evermann 1896: 24; Garman 1913:78. Scyliorhinus cephalus: Regan 1908a:460. Cephalurus cephalus: Bigelow and Schroeder 1941:73; Kato et al. 1967:23; Mathews and Ruiz 1974:556. 45 Figure 2(i. — Cephalurus cephalus, immature male, 195 mm, SU 293, Gulf of California. Drawing by Mary Wag- ner. Material examined. USNM 125094, ad. ', 240 mm, holotype, Albatross stn. 2992, lat. 18°17'30"N, long. 114°43'15"W, 841 m, near Clarion I.; SU293, imm. 3 195 mm, Albatross stn. 3007, lat. 25°27'30"N, long. 110°50'30"W, 662 m, near Santa Catalina I., Gulf of California; USNM, 3 juv. i, 82-89 mm, Albatross stn. 3007; USNM 87557, marked Albatross but no other data, probably stn. 3007; UA 67-63-1, gravid 5, 235 mm, 40 mm, embryos, Te Vega cruise 16, lat. 28°27'N, long. 112°34.5'W, 600-800 m, south of Tiburon I., Gulf of California, 9 October 1967; USNM, imm. $, 224 mm; imm. ?, 200 mm; 11 ad. 5, 260-320 mm, Anton Bruun cruise 18A, lat. 15°4.5'S, long. 75°45'W, 275 m, off Peru; USNM, 5 imm. i, 160-190 mm, lat. 11°24.5'S, long. 78°05'W, 380 m, off Peru; USNM 201525, I, 180 mm, lat. 07°48'S, long. 80°23'W, 365 m off Peru; USNM 3 juv. 3, 92-96 mm; juv. ?, 96 mm, lat. 23°41'S, long. 70°34'W, 250-400 m, off Chile. Diagnosis. As only one species of Cephalurus has been described, the generic diagnosis adequately distinguishes C. cephalus from other species. Description of adults. Head very wide and flattened with a very long branchial region; abdominal sector short, wider than high at pectoral, higher than wide at pelvic origin; tail sector moderately compressed, its length to caudal tip from midpoint of cloacal opening about one-half total length; caudal fin a little more than one-fifth total length; caudal axis not elevated; caudal peduncle higher than wide. Snout short, its length from front of mouth less than one-half mouth width; snout tip broadly rounded; pores of midsection on upper and lower sides of snout usually not conspicuous, arranged in single file lines, those of lower side of snout forming goblet-shaped pattern. Nasal apertures very large, oblique, nearer mouth than snout tip, not connected with mouth by a groove, least dis- tance between nasal apertures much less than horizontal diameter of eye; posterior inner opening of each aperture partly or completely covered by a short, broad flap, the outer corner of flap projecting to partly separate anterior and posterior openings. Mouth very wide and broadly arched, its width greater than the length of head in front of spiracles; short labial furrows present, continuous around mouth corners, lower furrow slightly longer, its length less than least distance between nasal apertures; inside of mouth with numerous short papillae on roof and tongue; gill rakers represented only by low mounds of tissue. Eyes moderately large; openings elongate, notched at both ends; eyes not visible from below but their angle of view more lateral than dorsal; no distinct secondary lower eyelid or subocular gutter; spiracles moderately small, diameter less than one-fifth eye length, behind and slightly below eye, separated from eye by distance slightly more than spiracle diameter. Gill slits large and very far apart, the fourth slightly in advance of the pectoral origin. Teeth small, generally somewhat similar in upper and lower jaws, those of central part of jaws generally tricuspid with the middle cusp much the longest; away from symphyses teeth become progressively slightly smaller toward the jaw angles, have proportionally shorter central cusps, and a greater number of cusps per tooth to about seven; teeth of cen- tral part of upper jaw of adult male holotype about l'/ 2 times as high as teeth of gravid female of same total length; three to five series of teeth functional, their arrangement alternate; 28 + 27/27+27 rows in adult male holotype, 33+33/34+34 in adult female from Gulf of California, 37 + 36/36+36 in one adult female from Peru; nosymphyseal teeth, varying space separates right and left halves of each jaw, from no space to space enough for 5 rows of teeth suggesting considerable elasticity of symphyseal ligaments. Dermal denticles over most of body surfaces widely spaced, somewhat needlelike, with only slight if any lateral expan- sion to form blades; most denticles nearly erect with moderate curve to direct points posteriorly, secondary points weak if developed at all on blades; denticles more closely spaced and broader near snout, around mouth, as a patch at the origin of the upper caudal fin, and, in some specimens along midline of back; denticles absent from roof of mouth, tongue, and gill bars. Two dorsal fins about equal in area; the first dorsal origin slightly in advance of pelvic origin; anal fin slightly larger than either dorsal fin and approximately under the second dorsal; pectoral fin origins under fourth gill slit or very slight- ly posterior to it; fin positions and sizes unusually variable in the available material except origin of first dorsal slightly in advance of pelvic origins in all specimens. 46 Skeletal elements not greatly mineralized; vertebrae, except for a few terminal, caudal vertebrae of smaller specimens, well defined on radiographs; radiographs made for vertebral counts show jaw cartilage outlines but do not show girdle and fin elements or chondrocranium outline; total number of vertebrae about 100 (95 to 106 in 18 specimens, mean 100.2); number of monospondylous vertebrae about 30 (28 to 35 in 27 specimens, mean 30.1); supraorbital crest of chon- drocranium absent. Color somewhat variable perhaps due to different preservation times and methods; generally dark gray, dark brown, or blackish above and somewhat lighter below, without prominent markings. Proportional dimensions in percentages of total length for the 237-mm adult male holotype from near Clarion Island, for a 235-mm adult female from the Gulf of California, and for a 290-mm adult female from Peru are as follows. Tip of snout to: front of mouth, 5.5, 5.1, 5.2; eye, 5.5, 5.5, 5.2; spiracle, 11.0, 9.9, 9.5; first gill opening, 21.9, 17.9, 20.4; fifth gill opening, 31.7, 30.6, 29.7; origin pectoral, 30.4, 27.2, 28.3; origin first dorsal, 44.7, 44.7, 45.5; origin pelvics, 46.0, 46.4, 46.6; origin second dorsal, 60.3, 62.6, 62.1; origin anal, 59.6, 61.7, 61.1; origin upper caudal lobe, 73.0, 76.6, 74.2; anterior end cloacal opening, 47.7, 47.2, 49.0. Orbit: horizontal diameter, 5.1, 4.9, 3.3; vertical diameter, 1.9, 1.7, 1.0. Spiracle: greatest diameter, 1.3, 0.9, 0.7; least distance from eye, 1.5, 0.9, 1.4. Mouth: width, 13.1, 12.8, 11.0; length, 6.3, 6.0, 4.1; length upper labial furrow, 0.8, 1.1, 0.7; lower labial furrow, 1.7, 1.4, 1.4. Nostrils: least distance between openings, 1.7, 2.9, 2.6. Gill slits: height of first, 3.8, 3.8, 2.8; height of fifth, 3.4, 3.8, 3.3. First dorsal fin: length base, 9.5, 9.2, 7.9; length posterior tip, 3.4, 1.9, 3.8; height, 3.4, 3.8, 3.8; length anterior margin, 10.1, 7.7, 7.9. Second dorsal fin: length base, 8.4, 7.0, 8.3; length posterior tip, 3.0, 2.1, 4.0; height, 4.2, 3.8, 3.5; length anterior margin, 10.1, 6.8, 7.9. Anal fin: length base, 9.3, 8.1, 10.2; length posterior tip, 3.0, 1.3, 3.6; height, 4.6, 4.0, 4.1; length anterior margin, 9.3, 8.1, 8.3. Pectoral fin: width base, 5.5, 5.1, 6.6; length anterior margin, 10.6, 8.9, 10.4. Caudal fin: upper margin, 27.0, 23.4, 25.9; anterior margin lower caudal lobe, 11.0, 10.2, 9.0; tip caudal to notch, 5.5, 6.4, 4.3. Distance between fin bases: first and second dorsal, 6.8, 9.5, 7.2; pectoral to pelvic, 10.6, 12.3, 13.1; pelvic and anal, 8.0, 7.4, 5.2; anal and lower caudal lobe origin, 5.5, 4.7, 3.5; second dorsal and origin upper caudal, 5.1, 6.4, 3.8. Claspers of male holotype project well beyond pelvic fin tip and are stiffened by clasper cartilage calcification; claspers have small fleshy projections at tip much as in some species of Parmaturus and Galeus; no clasper apron present either by union of inner margins of pelvic fins or membranous connection between pelvic fins. Three large females collected 25 August 1966 from lat. 15°04.5'S off Peru were partially dissected. All three specimens had large livers with the posterior tips of both right and left lobes extending beyond the cloaca; valvular intestine with five or six turns. Discussion. Only one species of Cephalurus is recognized here but additional material and further study may show that specimens taken off Peru and Chile represent a species distinct from those of the Lower California area. The darker color and the somewhat smaller eye notable in most of the specimens from Peru and Chile are regarded here as due either to circumstances of preservation or to intraspecific variation. The inside of the mouth of the specimens from Peru and Chile is dusky with numerous melanophores. The inside of the mouth in the Lower California specimens is white or yellowish. Cephalurus has been taken from nearly lat. 29°N to nearly lat. 24°S by trawling in depths from 250 to 841 m. The bot- tom temperature for Albatross station 2992 was 5.4°C and at station 3007, 7.0°C (Townsend 1901). Cephalurus cephalus is ovoviviparous retaining egg capsules with developing embryos within the oviducts. The 235- mm female from the Gulf of California contained embryos about 40 mm long. The three large females from Peru that were dissected each had one egg capsule in each oviduct. The egg capsules were transparent, very thin-walled, and fragile. Each capsule contained one large egg yolk about 15 by 30 mm, but embryos, if present, were not found. The egg capsules were shaped like egg capsules of oviparous scyliorhinids at the lower end but tendrils were represented only by short hooks and the upper (inner) end of the capsule was formless. Only the right ovaries were developed. The nidamental glands were small and each thin-walled oviduct was about 45 mm long. GALEUS RAFINESQUE Galeus Rafinesque 1810:13 (type-species, Galeus melastomus Rafinesque, designated by Fowler 1908:53). 47 ■ Pristiurus Bonaparte 1834, fasc. VII (type-species, Galeus melastomus Rafinesque, by monotypy). Figaro Whitley 1929:238 (type-species, Pristiurus (Figaro) boardmani Whitley, by monotypy). Diagnosis. Galeus species lack supraorbital crests of the chondrocranium that extend as narrow shelves above the orbits; they lack depressions in the upper lip or channels connecting the nasal apertures with the mouth; and they lack barbels. Galeus species have prominent labial furrows that are continuous around the mouth corners and extend for short distances along both upper and lower jaws; they have well-developed nasal flaps that cross or cover the posterior nasal openings; they have moderately long snouts broadly rounded at the tips and not greatly flattened; they have shallow subocular gutters that extend under the eyes for nearly their full lengths. Galeus differs from other scyliorhinid genera (except Parmaturus) in having a caudal crest of dermal denticles along the upper edge of the proximal half (or more) of the caudal fin. The caudal crest in Galeus consists of several rows of denticles, the central ones not much larger than denticles of body surfaces, but the marginal row of denticles on each side are much larger than other denticles and are strongly asymmetrical (Fig. 27). The top of the crest in Galeus is flattened and the marginal denticles project on the sides almost hiding a narrow band of naked skin that separates the caudal crest from denticles of the lateral surfaces. Furthermore, in species of Galeus known from juveniles as well as adults, the crest is well differentiated from the time of birth or hatching. In some species of Parmaturus, notably P. xaniurus, the crest is not fully differentiated in juveniles and lateral denticles of the crest are not asymmetrical as they are in adult P. xaniurus. Parmaturus species of all sizes have the top of the crest rounded, not flat as in Galeus, and in comparable sizes have more longitudinal rows of denticles in the crest than Galeus species. Dermal denticles of dorsolateral surfaces of juvenile Galeus are sparsely distributed, nearly erect, and without lateral blade points. Replacement denticles are progressively wider and less erect. Dorsolateral surfaces of adults have im- bricate denticles. Galeus has large and broad pectoral fins, their greatest width usually greater than the width of the mouth (about equal in G. piperatus) whereas Parmaturus has narrower pectorals, their greatest width about equal to or less than the mouth width (see discussion). Species of Galeus are either oviparous (as in G. melastomus) or ovoviviparous (as in G. polli). Claspers of adult males are moderately long and slender or very long and slender (in G. nipponensis) with remarkably complex terminal struc- tures but no hooks. Aprons are partially developed in some species. All species of Galeus are darker above than below and several species have elaborate patterns of markings on the dor- sal surfaces that diverge considerably from the pattern of saddle blotches present on many scyliorhinid species. Par- maturus species have few color markings and both the ventral and dorsal surfaces are uniformly dark. Discussion. The livers of Galeus are small and short in the species that I have examined and have low oil content with little or no squalene. The broad pectorals of Galeus may be an adaptation to assist in providing the lift needed to offset a high sinking factor. The other genus with a caudal crest, Parmaturus, is presumably aided in moving more often into midwater by greater liver oil buoyancy and does not require such a large pectoral fin as Galeus. Species of Galeus occur in the eastern Atlantic from the coasts of Iceland and Norway nearly to Angola. They are pres- ent in the Mediterranean. They are found in the tropical and subtropical western North Atlantic. They are present in the Gulf of California and elsewhere in the Pacific in waters around Japan, Formosa, the Philippines, and Australia. Figure 27. — Enlarged denticles of the caudal crest of Galeus arae arae. Drawing by Mary Wagner. 48 Galeus has not been taken off the coasts of South Africa, an area having more species of scyliorhinids than any other, and is unknown from the Indian Ocean. In some regions where Galeus is unknown, the lack of records may merely reflect the lack of deepwater trawling. On the Atlantic and Caribbean coasts of the Americas, however, the known geographical range of Galeus ends abruptly at its southern end off the coast of Colombia and at its northern end off the coast of Georgia. Galeus is absent from the Gulf of Mexico west of the Mississippi River mouth and from the Gulf of Campeche. Variation is great in color pattern and in morphometries even among adults within a single species. In the following key, two species, G. piperatus and G. schultzi, key out in three places and G. arae antillensis keys out in two places. Vertebral numbers are given for most of the species (Table 2) and these may be of some assistance in identification. Hatchlings and the smaller postjuvenile specimens of Galeus may not be identifiable to species and their iden- tifications are unlikely to be helped by the following key. Known hatchlings or newborn Galeus have the seven or eight dorsal saddle blotches that occur among juveniles of several scyliorhinid genera. Twelve forms of Galeus are recognized, two of them described as new. Three of the 12 are regarded as subspecies, a category that I consider useful in this genus. I have been unable to make an entirely satisfactory classification for some West Indian populations, here grouped as Galeus arae antillensis and the few available specimens suggest the possibility that slopes of each island may have populations distinct from others (see Fig. 31). Juveniles are poorly known and are remarkably rare in collections. I am unable to suggest a reason other than probable loss through the large meshes of nets commonly used in deepwater trawling. Species of Galeus are oviparous and some are ovoviviparous. Substantial evidence shows that G. melastomus melastomus is oviparous and that G. polli is ovoviviparous. Observations indicate that G. arae arae is probably ovoviviparous and that G. arae antillensis and G. piperatus are probably oviparous. Gross anatomical differences between the oviparous species and the ovoviviparous species have not been observed. I have placed Figaro Whitley (1929) in the synonymy of Galeus. Whitley's Figaro boardmani is one of the few catsharks that I have not seen. It is described (Whitley 1940) as differing from Galeus in having modified denticles similar to those of the caudal crest at the base of the lower caudal fin lobe. Other species of Galeus do not have modified denticles at the base and on the leading edge of the lower caudal lobe but such denticles are present on some large specimens of Parmaturus pilosus. Table 2. — Numbers of monospondylous vertebrae in species of Galeus. Species of Galeus 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 G. arae arae G. arae antillensis G. arae cadenati G. melastomus melastomus (Norway) G. melastomus melastomus (Mediterranean) G. polli G. nipponensis 1 G. eastmani 1 G. sauteri 1 G. schultzi G. piperatus 2 12 43 23 8 1 8 21 7 3 5 5 6 8 2 1 1 - 17 5 1 5 2 16 1 33 4 18 7 2 3 7 2 2 7 1 6 3 3 3 1 3 •Data added from Nakaya (1975). 2 Data added from Hubbs and Taylor (1969). Key to Adult Galeus la. Denticle arrangement similar to caudal crest present on lower side of caudal peduncle, continu- ing on to leading edge of lower caudal fin lobe G. boardmani lb. Denticle arrangement similar to caudal crest not present on lower side of caudal peduncle and not on leading edge of lower caudal fin lobe 2 2a. Dorsolateral body surfaces brown or gray-black, either uniform in color or with weakly contrast- ing saddle blotches or spots darker than the ground color but with poorly defined margins; not variegated 3 2b. Dorsolateral body surfaces brown or gray-black variegated with saddle blotches, spots, or lines of darker or lighter color, the markings usually with sharply defined margins; variegated 10 49 3a. Dorsal and lateral body surfaces with poorly defined darker markings, their margins indefinite sometimes recognizable as saddle blotches 4 3b. Dorsal and lateral body surfaces uniformly colored, without markings 8 4a. Lining of mouth white 5 4b. Lining of mouth black 6 5a. Distance from tip of snout to nearest rim of nasal opening less than horizontal diameter of eye . . G. eastmani 5b . Distance from tip of snout to nearest rim of nasal opening greater than horizontal diameter of eye . . G. nipponensis 6a. Length of snout in front of mouth comparatively short, 5.4 to 5.9% TL some G . schultzi 6b. Length of snout in front of mouth comparatively long, 5.8 to 6.6% TL 7 7a. Length of base of anal fin comparatively short, usually less than 11% TL some G. arae antillensis 7b. Length of base of anal fin comparatively long, usually more than 11% TL some G. piperatus 8a. Both dorsal fins and sometimes upper and lower caudal fin lobes with a dark spot or dark smudge near their upper terminal margins (or lower tip on lower lobe of caudal fin) and usually not narrowly edged with black G. sauteri 8b. Both dorsal fins and caudal fins not spotted or smudged, fins either uniformly colored, or edged with black, or marked with a narrow terminal colorless band that contrast with the color of the rest of the fin 9 9a. Adults 500 mm or more TL G. melastomus murinus 9b. Adults less than 400 mm TL G. piperatus 10a. Highly variable color pattern on dorsolateral surfaces, usually made up of dark markings with white borders or white areas over or near the dorsal midline, markings often in longitudinal series or forming stripes G. arae antillensis 10b. Color pattern or dorsolateral surfaces made up of series of dark spots or saddle blotches or both 11 11a. Distance between bases of dorsal fins greater than length of base of anal fin; Pacific species 12 lib. Distance between bases of dorsal fins less than length of base of anal fin; Atlantic species 13 12a. Mouth width comparatively small, 6.5 to 6.7% TL; mouth arch comparatively high G. schultzi 12b. Mouth width comparatively great, 8.3 to 13.2% TL; mouth arch comparatively low and broad some G. piperatus 50 13a. Base of anal fin comparatively short, 10.3 to 13.6% TL; its length usually less than the distance between bases of the two dorsal fins G. arae arae 13b. Base of anal fin comparatively long, 13.2 to 17.0% TL, its length greater than the distance between bases of the two dorsal fins 14 14a. Dorsal ground color and markings in shades of gray or gray-black (before storage in alcohol that may release brown-staining pigments from livers); length of snout in front of mouth comparatively great, 7.4 to 8.5% TL G. polli 14b. Dorsal ground color and markings in shades of brown; length of snout in front of mouth 6.0 to7.8%TL 15 15a. Tip of anal fin reaching a little past the origin of lower caudal fin lobe G. melastomus melastomus 15b. Tip of anal fin not reaching origin of lower caudal fin lobe G. arae cadenati Galeus arae arae (Nichols) Figures 27, 28A, 29, 30 *£P> Figure 28.— A. Galeus arae arae, 290-mm female, coast of Florida. B. Galeus arae cadenati, 300-mm female, Caribbean coast of Panama. C. Galeus arae antillensis, 320-mm female, coast of Puerto Rico (see Fig. 32 for other color patterns of G. a. antillensis). 51 Pristiurus arae Nichols 1927:1 (off Miami Beach, Fla.). Galeus arae: Bigelow and Schroeder 1948:216, figs. 36, 37; Springer 1966:608, figs. 20A, 21A, 27F; Bullis 1967:141, figs. 8-3 to 8-6. Material examined. USNM, 3 ad. $, 272-290 mm, 1 imm. S, 190 mm, 1 imm. S, 125 mm, 1 juv. 3, 90 mm, 3 ad. ?, 255-331 mm, 3 imm. ?, 196-228 mm, all from the coast of Florida. Examined by X-ray: 48 males, 37 females, coast of Florida; 72 males, 38 females, Caribbean coasts of Nicaragua, Honduras, and Costa Rica; 86 other specimens from Florida and Honduras coasts. A print-out from data cards on file at the National Marine Fisheries Service Labo- ratory at Pascagoula, Miss., on 213 trawling stations where G. arae arae were taken in the Gulf of Mexico, the Caribbean Sea, and adjacent waters by exploratory fishing vessels. Diagnosis. Galeus arae arae is the smallest Galeus of the Atlantic, females rarely reaching 360 mm and males 340 mm. Galeus arae cadenati may be only a little larger but is not well enough known for a reliable size estimate. Galeus arae antillensis is larger, some specimens reaching more than 400 mm. The Galeus of the eastern Atlantic all may reach a length of more than 400 mm. Galeus arae arae is readily separable by its complex color pattern (Fig. 28A) from the plain colored or weakly marked species of the western Pacific: G. sauteri, G. eastmani, and G. nipponensis. It has a shorter snout than G. polli, its length in front of mouth 5.3 to 7.1% TL for G. arae arae and 7.4 to 8.5% TL for G. polli. Galeus arae arae is similar to G. schultzi of Philippine seas in morphometries, but has a more strongly marked color pattern with more spots on the top of the head and anterior trunk than G. schultzi. Galeus arae arae and G. melastomus melastomus both have complex color patterns but G. arae arae is not only a smaller form but also has a shorter anal fin base, 10.3 to 13.6% TL compared with about 15.0 to 16.0% TL for G. melastomus melastomus. Also in adult G. arae arae the posterior tip of the anal fin falls appreciably short of the origin of the lower lobe of the caudal fin, but reaches it in both G. melastomus melastomus and G. polli. In G. arae arae the length of the snout in front of the mouth is 5.3 to 7.1% TL but in G. piperatus it is 6.3 to 7.8% TL; in G. arae arae the distance between bases of the two dorsal fins is 10.5 to 12.2% TL, but in G. piperatus it is 15.2 to 17.3% TL. Galeus arae antillensis may differ from G. arae arae in any of the following ways: it may reach a larger size at maturity; it may have a shorter anal fin; it may have more monospondylous vertebrae; it may have either a much reduced and less distinct color pattern (Fig. 28C) or a stronger but different pattern (Fig. 31). Throughout its range as defined below, G. arae arae of half-grown to adult size have a distinctive and complex color pattern with little variation (Fig. 28A). Small and large spots among the usual saddle blotches of brown or tan are in a bilaterally symmetrical arrangement over dorsal and lateral surfaces. Ventral surfaces are light colored without spots. Very small examples, under 150 mm, have only the saddle blotches. The pattern is not much different from that of G. melastomus melastomus. The complexity of the pattern and distinctness of spots is somewhat reduced in some other species of Galeus. Egg capsules certainly referable to G. arae arae are unknown, but eggs not enclosed in capsules were taken from ovi- ducts of specimens collected off Honduras (Bullis 1967). Very small specimens, 100 mm or less, are known from only a few specimens. A 90-mm male, presumably newborn, collected 9 November 1960 off Ft. Lauderdale, Fla., from 376 m had indistinct saddle blotches. The denticles were needlelike, as in many young scyliorhinids, and were arranged in longitudinal rows on the flanks. Some denticles around the mouth had a narrow blade and were three-pointed. The teeth were similar in shape and number to the teeth of the adult. The caudal crest was well organized with the lateral scales asymmetrical as in adults. The crest had fewer scales in the central rows than adults. A 125-mm male collected 30 April 1961, off St. Augustine, Fla., from 302 m had more numerous denticles with only a faint tendency to form longitudinal rows. Most of the denticles were needlelike. Specimens 150 mm or more had mostly tridentate denticles not arranged in rows. Description (morphometries) . The range of measurements expressed as percentages of total lengths for 10 Galeus arae arae 190 to 321 mm long, all from the coast of Florida, follow. Tip of snout to: front of mouth, 5.3-7.1; eye, 5.9-7.8; first gill slit, 13.8-16.0; fifth gill slit, 16.3-19.8; origin pectoral, 15.3-19.3; origin first dorsal, 42.4-47.3; origin pelvics, 37.2-42.0; origin anal, 50.9-56.9; origin second dorsal, 59.0-67.7; upper caudal fin from origin of caudal crest, 65.5-72.0; anterior end cloacal opening, 39.8-46.3. Eye: length, 3.4-3.9; distance eye to spiracle, 0.8-1.5. Mouth: width, 6.2-7.5; length, 3.2-4.3; length upper labial furrow, 1.0-1.5; length lower labial furrow, 1.4-2.2. Gillslits: height of first, 1.3-2.1; height of fifth, 0.6-1.3. First dorsal fin: length base, 4.2-5.9; length anterior margin, 7.2-8.3. Second dorsal fin: length base, 4.4-5.7; length anterior margin, 7.0-8.0. Anal fin: length base, 10.3-13.6; length anterior margin, 5.5-8.0. Distance between: first and second dorsal fin bases, 10.3-13.6; posterior inner ends of nostril, 2.5-3.3. Pectoral fin: length anterior margin, 10.5-12.2. 52 Discussion (Geographical and Vertical Ranges). Galeus arae arae is found from the coast of Georgia southward on the continental slope around the Dry Tortugas and northward in the Gulf of Mexico as far as the Mississippi River delta. It is rare west of Pensacola and absent from the western Gulf of Mexico and Gulf of Campeche. It occurs off Honduras, Nicaragua, and Costa Rica on the continental slope and also off neighboring islands and oceanic banks including Pedro Bank south of Jamaica. It is replaced along the Caribbean coastal slope of Panama and Colombia by G. arae cadenati. The very sparse population of Galeus on the northeastern coast of Cuba, the Santaren Channel, and Mona Passage seems to be made up of intergrades of G. arae arae and G. arae antillensis. The geographical range of G. arae arae is restricted to a narrow band along the slopes well below the edge of the conti- nental shelf. The species was recorded from several hundred stations of research vessels of the National Marine Fisheries Service at depths from 292 to 732 m (160 to 400 fathoms). The water temperature taken at the bottom by reversing thermometer was recorded from 56 stations where G. arae arae was taken and ranged from 5.6° to 11.1°C. The mean temperature at 44 stations off the Florida coast was 9.2°C and at 12 stations off Central America it was 8.75°C. We have too few records of depth or temperature to make firm conclusions but it is interesting that the only temperature record at the point of capture of G. arae cadenati was 4.6°C and the shallowest record for a Galeus in the western Atlantic was for a G. arae antillensis taken from 146 m. Although G. arae arae must be common within most of its range, it seems to be irregularly distributed. The largest trawl haul contained 180 specimens and more than 10 were frequently taken in single hauls. Nevertheless, many hauls made in areas expected to have G. arae arae produced no specimens. The object of the trawling was chiefly to define areas of occurrence of red shrimp, Hymenopeneus robustus Smith, a commercially valuable species on which Galeus sometimes feeds. Stomach examinations showed that small Hymenopeneus and a smaller shrimp, Penaeopsis megalops (Smith) made up the greatest bulk of the food of the G. arae arae examined. Bullis (1967) found adults but very few immature specimens in depths of more than 450 m. Although immature exam- ples were most common in less than 450 m, adults of both sexes were also common there. In view of the large numbers of trawling stations providing records of the presence or absence of G. arae arae, the limits of the geographical and depth ranges would be well established except for the unexplained paucity of data on very small specimens (under 150 mm) and on gravid females. Reproduction. Bullis (1967) found eggs without egg capsules in the oviducts posterior to the shell glands in G. arae arae taken off Honduras. This constitutes the only direct evidence about the site of embryonic development of G. arae arae. Springer (1966) had assumed the species to be ovoviviparous but only because of failure to find egg capsules. The smallest young example seen was the 90-mm specimen mentioned in the foregoing description and a few, probably not more than a dozen specimens, less than 150 mm long were seen. If, as seems certain, G. arae arae gives birth to fully developed young, gravid females should have been found among the large number of mature females seen. Except for those reported by Bullis, no others have been observed. The only plausible explanation for this is that gravid females congregate in nursery areas of very rough bottom that precludes trawling. One nursery area for scyliorhinid sharks is known for Scyliorhinus retifer, an egg laying species. The unsatisfactory state of knowledge about the development of G. arae arae is accentuated by the fact that in G. arae antillensis, for which relatively few specimens are available, one has been found with encapsulated eggs, one egg in each oviduct. Vertebral numbers. The comprehensive survey of vertebral numbers in sharks by Springer and Garrick (1964) outlined some of the tendencies toward special family characteristics in vertebral number. They discussed differences between monospondylous and diplospondylous vertebrae but did not report numbers of each in their tabular summary. The Scyliorhinidae represent a special case because, on the average, they are smaller than sharks of other families and their vertebrae are more difficult to count for that reason alone. The Scyliorhinidae, also on the average, are species of deeper waters where strength and rigidity of the vertebral column are not so important as in shallow waters, hence it is not surprising that scyliorhinid vertebrae are comparatively poorly calcified. There are few exceptions to the rule that the long monospondylous vertebrae in the Scyliorhinidae end abruptly in the pelvic region to be replaced in the post- pelvic region by obviously shorter diplospondylous vertebrae. The point of change is easily recognized. The point at which caudal vertebrae begin in the Scyliorhinidae is not easily established. The caudal axis is either little or not at all elevated and there is no visible difference in vertebrae in the part of the vertebral column near the base of the tail. In two genera, Galeus and Parmaturus, the beginning of the caudal sector may be determined as opposite the point of origin of the caudal crest. In other scyliorhinid genera the point of origin of the upper caudal fin lobe may be used as a reference point. It is not a good one, however, because the upper fin lobe rises gradually and its exact point of origin is subjective. From some early counts of vertebrae of several species of Galeus, it appeared that males might have more vertebrae than females and that species (or populations) with a northerly habitat might have more vertebrae compared with those from a more southerly habitat. A test procedure was arranged using all Galeus arae arae and all G. polli available at that time. The specimens were all radiographed and vertebrae were counted by a student technician, John Asarian, following a uniform procedure. The G. arae arae were divided into two groups: one group of 85 from the coast of Florida north of lat. 23°30'N, and the other group of 110 from the Caribbean coasts of Honduras-Nicaragua-Costa Rica south of lat. 53 17°40'N. The sharks of the two groups could not be distinguished from each other by color pattern or external morphological features that separate the species and subspecies now described in the genus Galeus. The test series was further subdivided by sex and also by size into three length categories: small specimens 150-227 mm, medium specimens 228-253 mm, and large specimens over 253 mm. The G. polli were grouped only by sex and area of origin; northern specimens from lat. 5° to 8°N off Sierra Leone and Liberia, and southern specimens from lat. 3° to 4°S off Gabon. The results of examinations are summarized here in two Hubbs-Perlmutter graphs as modified by Hubbs and Hubbs (1953) and show our data for monospondylous vertebrae (Fig. 29) and for total vertebral numbers (Fig. 30). The graphs permit a rough estimate of significance in that the lack of overlap between any two dark bars indicates a significant difference. The length of the line underneath the dark and light bars shows the range of variation in the sample. Thus for G. arae arae it can be seen at a glance that differences in total length within sex and area groupings are not significant, that differences between northern females and southern males are not significant, and that differences between monospondylous vertebral numbers between northern and southern females are not significant. It is evident that the great range of variation in total numbers of vertebrae, 130 to 142 in the sample of G. arae arae and 126 to 137 in the G. polli sample, makes the use of these counts of little use in identification. The variation is less in monospondylous vertebrae, in the samples used here, 34 to 38 for G. arae arae and 32 to 35 for G. polli. The analysis shows only that, in Galeus arae arae, a small but significant difference in numbers of vertebrae exists between males and females and that both males and females may have a significantly greater number of vertebrae in one geographical area than in another. Smell northern moles f 24) Medium northern moles' E Lorqe northern moles 19) Northern moles combined Smoll northern females fie) Medium northern females (101 _ Large northern females (91 Northern females combined (37). Smoll southern moles (21) Medium southern moles I 61 Large southern males (35) Southern moles combined "21 Smoll southern females '12) Medium southern females (13) Large southern females (131 Southern females combined i38 3 Northern moles 1221 Northern females 120] Southern males [6] Soufhi BG £fcl 2. .« females [101 35 _i_ 36 NUMBER OF MONOSPONDYLOUS VERTEBRAE Figure 2ft. — Number of monospondylous vertebrae in samples of Galeus arae arae and G. polli shown in a Hubbs-Perlmutter graph. The samples of the northern population of G. a. arae were taken between lat. 2.')° and 2!I°N and the samples of the southern population between lat. 9° and 18°N. The northern samples of G. polli were taken be- tween lat. 3° and H°N and the southern samples between lat. .'!° and 4°S. From the same data, t values were calculated for G. a. arae as follows: between northern males and northern females t = 3.6205 (difference significant at 99.9% level); between south- ern males and southern females t = 4.4010 (significant at 99.9%); between northern males and southern males t = 2.9149 (significant at 99.5%); between northern females and southern females t = 1.0X22 (not significant at Xlfvr level). 54 ■ H 129 Small northern males (24) Medium northern males 115) Large northern males (9) Northern males combined (48) Small northern females (18) Medium northern females (10) Large northern females (9) Northern females combined (37) Small southern moles (21) Medium southern males 06} Large southern moles (35) Southern males combined (72) Small southern females (12) Medium southern females (13) Large southern females (13) Southern females combined (38) Northern moles (22) Northern females (20) Southern moles (6) Southern females (10) 138 >9 TOTAL NUMBER OF VERTEBRAE Figure 30. — Total number of vertebrae in same samples (as in Fig. 29) of Gateus arae arae and G. polli. The f values calculated for G. a. arae are as follows: between northern males and northern females t = 3.1(>87 (difference significant at the 99.57c level); between southern males and southern females f = 4.0844 (significant at 99.9%); between northern and southern males t = 4.8680 (significant at 99.9 r /t); between northern and southern females t = 4.7338 (significant at 99.9%). Galeus arae antillensis New Subspecies Figures 28C, 31, 32 Holotype: USNM 214178, a 347-mm ?, near St. Kitts, Leeward Islands, W.I., from 550 to 585 m, 18 May 1967, RV Oregon stn. 6695, water temperature at point of capture 8.9°C. Other material examined. USNM-ORE 5417, 2 imm. S, 270, 255 mm, near La Tortue Island, Windward Passage, W.I., 658 m; USNM-ORE 6701, 3 $, 325, 328, 345 mm, Leeward Islands, W.I., 640-676 m; USNM-ORE 6722, 2 imm. S, Leeward Islands, 612-698 m; USNM-ORE 2646, imm. S, 160 mm, Leeward Islands, 384 m; USNM-ORE 10862, 2 S, 250, 430 mm, Cal Sal Bank, Santaren Channel, 457 m; USNM-SB5181, imm. S, 270 mm, $, 240 mm, Mona Passage, W.I., 548 m; USNM-ORE 10840, 2 ?, 237, 410 mm, Leeward Islands, 642 m; USNM-ORE 10841, ?, 200 mm, Leeward Islands, 643 m; USNM-ORE 10842, imm. S, 260 mm, 4 ?, 130, 235, 350, 370 mm, Leeward Islands, 579 m; USNM-ORE 10843, imm. S, 320 mm, 3 5, 290, 320, 435 mm, Leeward Islands, 589 m; USNM-ORE 10207, imm. 8, 200 mm, 2 ?, 150-200 mm, Leeward Islands, 658 m; USNM-ORE 5427, ad. 8, 310 mm, off Hispaniola, 512-576 m; USNM-ORE 6721, ?, 340 mm, Leeward Islands, 622-695 m; USNM-ORE 6695, ad. 8, 335 mm, 3 ?, 345, 355, 425 mm, all nongravid adults, Leeward Islands, 550-585 m. Diagnosis. Galeus arae antillensis can usually be recognized by the characters used in the foregoing key to species but it is remarkably variable. It reaches a length of 435 mm or more, but some specimens, the holotype, for example, are mature at less than 350 mm. It is a larger form, however, than the other western Atlantic Galeus. The color pattern diverges only slightly from G. arae arae on the eastern and southern sides of the Straits of Florida, but specimens taken off Hispaniola and Puerto Rico have obscure (Fig. 28C) and dull markings. In the vicinity of the Leeward Islands, patterns appear to vary greatly from obscure markings to vivid patterns of brown, black, and white (Fig. 31). The color patterns differ from the pattern of the continental form in being either stronger, weaker, or only slightly different in general arrangement. 55 -i-,;.\ Figure 31. — Galeus arae antillensis. Two color patterns from immature specimens from the vicinity of Antigua, West Indies. Drawings by Mil- dred Carrington. Figure 32. — X-ray of pelvic area and claspers of adult male Galeus arae antillensis, slope off northwestern Peninsula of Haiti. This shows a thin layer of mineral that is deposited at the surface of the principal clasper cartilage (stem) immediately following maturation of the testes. Appreciable growth in length of all male sharks stops abruptly or becomes negligible at the time of clasper calcification (stiffening). The heavily calcified terminal cartilages are more com- plex in Galeus than in most other scyliorhinids. 56 The subspecies has a shorter anal fin base and a longer distance between the two dorsal fin bases than other Atlantic species except possibly the little known G. melastomus murinus. Too few specimens scattered over a very large geographical area were available to use for a statistical study. The mean number of monospondylous vertebrae in Galeus arae antillensis was not much different than the mean number for a much larger sample of G. arae arae but the range in G. arae antillensis was twice as great. Description. The dimensions of the 347-mm holotype, a sexually mature female with ovarian eggs about 10 mm in diameter in the right ovary, are given below as percentages of the total length. Tip of snout to: front of mouth (upper lip), 6.6; eye, 6.9; spiracle, 12.7; first gill slit, 17.0; fifth gill slit, 20.7; origin pectoral, 20.1; first dorsal origin, 46.9; origin pelvics, 41.7; origin second dorsal, 64.4; origin anal, 57.5; origin caudal crest, 73.9; anterior end cloacal opening, 44.0. Eye: length orbit, 4.3; height, 1.7; distance between the two upper eyelids, 7.8. Spiracle: greatest diameter 0.6; least distance from eye, 1.2; distance between the two spiracles, 8.3. Mouth: width, 7.5; length, 4.0; length upper labial furrow, 1.7; length lower labial furrow, 2.0. Nasal apertures: level of anterior ends to tip of snout (projection), 3.7; level of posterior ends to level of middle of upper lip, 0.8; least distance between, 2.9. Gill slits: height of first, 2.0; height of fifth, 1.2. First dorsal fin: length of base, 5.2; length of free inner margin, 2.6; height, 4.0; length of anterior margin, 6.9. Second dorsal fin: length of base, 4.9; length of free inner margin, 2.9; height, 2.9; length of anterior margin, 6.6. Anal fin: length of base, 10.9; length of free inner margin, 1.4; height, 2.2; length of anterior margin, 6.3. Pectoral fin: width of base, 6.3; length of anterior margin, 10.9; greatest width of fin, 8.3; distance between the two pectoral axils, 6.3. Pelvic fins: overall length, origin to tip, 13.5. Caudal, fin: upper margin from origin of caudal crest, 28.2; anterior margin of lower caudal lobe, 7.8; tip of second dorsal to origin caudal crest, 1.7; tip of anal to origin of lower caudal lobe, 5.2. Distance between fin bases: first and second dorsals, 14.1; pectoral and pelvic, 15.5; pelvic and anal, 7.5; anal and lower caudal origin, 6.0; second dorsal and origin of crest, 5.2. Body dimensions: greatest width head, 11.2; trunk width at pectoral origin, 10.1; trunk width at pelvic origin, 6.3; width at origin caudal crest, 2.0; trunk height at pectoral origins, 7.5; trunk height at pelvic origin, 9.5; caudal peduncle height at origin caudal crest, 3.5. Teeth small and crowded, estimated number of vertical tooth rows 28+28/26+26; the largest about 0.7 mm high above the enamel line; about 3 or 4 rows nearest symphysis with three cusps, the central cusp much the longest, most of the teeth with four, five, or six cusps, number of cusps increasing laterally but irregularly. Lateral 8 to 10 rows of teeth in lower jaw with 5 to 7 low subequal-length cusps forming a comblike structure, otherwise teeth of upper and lower jaws similar. Dermal denticles of dorsolateral surfaces small, imbricate, with three lateral points, the middle one much the longest, the larger flank denticles about 0.2 to 0.3 mm long, transparent. Caudal crest of holotype about 64 mm long, the longest asymmetrical marginal denticles about 1.2 mm long, number of longitudinal rows of small, median, symmetrical den- ticles increasing from zero at origin to about four near posterior terminus. Inside of mouth with moderate numbers of denticles on anterior mouth roof, estimated about 0.05 mm long, denticles not detectable elsewhere in mouth or on gill bars. Gill bars with one to three low and broad tubercles but no gill rakers. Color of inside of mouth dusky, probably black in life, ground color after about 8 yr in preservative light yellowish white, dorsolateral surfaces with tan markings; top of head and dorsal trunk with somewhat irregular large tan saddle blotches, one at first dorsal, one at second dorsal, and two across tail. Liver short, its lobes reaching less than halfway toward posterior end of body cavity, no oil detected by touch after long preservation. Peritoneum black. Right ovary with 3+ yolks about 10 mm in diameter. Discussion. Galeus arae antillensis, as defined here, represents the island form of Galeus arae. The geographical range of G. arae antillensis extends from the eastern and southern slopes of the Straits of Florida east-southeastward along the slopes off the northern coasts of Cuba, Hispaniola, and Puerto Rico through slopes off many of the Leeward Islands. Specimens have not been seen from slopes off the southern coasts of Cuba, Hispaniola, or Puerto Rico, or off Jamaica or southward from Martinique but the absences could be due to lack of fishing effort there. With respect to color pattern, G. arae antillensis is nearest to G. arae arae in the adjacent parts of the ranges of the two, but diverges greatly in the Leeward Islands area. The average number of monospondylous vertebrae in 50 G. arae antillensis was about the same (35.76) as from a com- posite sample of 86 continental slope G. arae arae (35.31), but the range of variation was greater (33 to 39) in G. arae an- tillensis as compared with that (34 to 37) in G. arae arae. 57 One 345-mm female from the Leeward Islands near Antigua had one egg capsule in each oviduct. The capsules measured approximately 38 by 16.5 by 10 mm. The evidence for oviparity in G. arae antillensis is scant as is the evidence for ovoviviparity in G. arae arae. It is not apparent why as much material as is available does not give much more infor- mation about prenatal development. The holotype was trawled where the water temperature within a meter of the bottom was 8.9°C and within the temperature range at which G. arae arae has been taken. As with G. arae arae, the specimens of antillensis were taken mostly from 300 to 700 m, but a few antillensis were recorded in RV Oregon notes as being collected at lesser depths near Puerto Rico. Galeus arae cadenati Springer Figure 28B Galeus cadenati Springer 1966:609, fig 20 (Caribbean coast of Panama). Material examined. USNM 260468-F1, 2, 303 mm, holotype, Caribbean coast of Panama, 439 m, 30 May 1962; USNM-ORE 3600, 9, 314 mm, Caribbean coast of Panama, 548 m; USNM-ORE 5722, imm. -\ 290 mm, 2 1, 270, 300 mm, Caribbean coast of Panama, 512 m. Diagnosis. Galeus arae cadenati has a longer anal fin base than the other western Atlantic subspecies of Galeus arae, the anal fin base length in G. arae cadenati being about as long or longer than the distance between dorsal fin bases. The few specimens of G. arae cadenati seen indicate a somewhat greater size than G. arae arae. Discussion. Galeus cadenati was described (Springer 1966) from a small series of female specimens from the Caribbean coast of Panama. Only a few additional specimens were collected and these included some from the coast of Colombia (Harvey R. Bullis, Jr., pers. commun.). The separation of the large population of typical Galeus arae arae of the Caribbean coast northward from Panama from cadenati is not clear and abrupt. Provisionally all the specimens seen from the coasts of Panama and Colombia are regarded here as Galeus arae cadenati. Galeus boardmani (Whitley) Pristiurus (Figaro) boardmani Whitley 1928, pi. 18, fig. 3 (Montague Island, New South Wales, 128-146 m). Figaro boardmani socius Whitley 1939:230 (Great Australian Bight off western South Australia, and eastern Western Australia). Figaro boardmani: Whitley 1940:90, fig. 83. Galeus boardmani: Fowler 1941:28. Material examined. None Diagnosis. Galeus boardmani differs from other species of the genus as here defined in having a caudal crest structure on the lower side of the caudal peduncle and extending for some distance on the leading edge of the lower caudal fin lobe. Galeus boardmani shares this character with some individuals of Parmaturus pilosus and juveniles of P. melano bronchus and P. campechiensis. It may be distinguished, however, from species of Parmaturus by its wider and larger pectoral fins; the maximum width of the pectoral is greater than the mouth width in Galeus but less than mouth width in Parmaturus. Whitley's (1940, fig. 83) illustration of G. boardmani shows a shark with the typical Galeus-Mke seven or eight saddle blotches with an additional eight or more slightly smaller blotches interspersed. Discussion. The holotype of Galeus (Figaro) boardmani is a 540-mm male, AMS IA 2483, trawled from the vicinity of Montague Island, southern New South Wales in July 1925 from 70-80 fathoms (128-146 m). Whitley's (1928) description mentioned para types trawled from slightly deeper water nearby in September. Whitley (1928) placed his new species boardmani in a new subgenus, Figaro, on the basis of the presence of modified scales on the ventral edge of the caudal peduncle and later (1939, 1941) treated Figaro as a genus. According to Whitley's accounts (1928:238; 1939:230) many specimens were in the Endeavor collections trawled from the eastern edge of Bass Strait in 100-200 fathoms. He also reported the collection of specimens from the Great Australian Bight (off South Australia and Western Australia) in depths of 70 to 450 fathoms (128-823 m). He noted that all were males less than 16 in (about 400 mm) long. Specimens from the Bight were described as having fainter color bands (dorsal saddles) by Whitley and given subspecific rank as Figaro boardmani socius. Although I have not recognized Figaro here, I should point out that Whitley's (1940, fig. 83) figure of the holotype shows a shark with a larger spiracle than I would expect in Galeus and a pointed clasper. If the holotype is a mature male, the figure suggests that Figaro might be separated from Galeus by clasper characters. 58 I suspect that the hump in the back mentioned by Whitley as a G. boardmani character is the result of fixation with strong preservative which, for deepwater scyliorhinids with weak vertebrae, tends to distort the trunk and often produces a hump. Galeus eastmani (Jordan and Snyder) Pristiurus eastmani Jordan and Snyder 1904:230, pi. 60 (off Izu, Japan); Garman 1913:93. Galeus eastmani: Fowler 1941:26; Besednov 1969:28, fig. 11; Nakaya 1975:23-25, 46. Material examined. SU 7740, ?, 337 mm, holotype (originally measured 345 mm), off Izu, Japan; FMNH, Owston 678, ad. S, 347 mm, Japan (Misaki); FMNH, Owston 705, subad. $, 315 mm, Sagami Sea, Japan. Diagnosis. In the original description (Jordan and Snyder 1904:230) it was stated, "Teeth each with seven acutely pointed cusps . . ." and was repeated by Garman (1913) and Fowler (1941). My examination of the holotype revealed that only the extreme lateral teeth have seven cusps. The majority of the teeth have four or five cusps as shown in Nakaya's illustration (1975, fig. 24) and are very small, those in the holotype reach at most about 1 mm high from the enamel line. The teeth of G. eastmani are smaller than the teeth of G. sauteri. They are crowded with more than one series in function. The dermal denticles are small; typical three-pointed flank denticles of the 347-mm male are 0.3 to 0.4 mm long compared with 0.7 for flank denticles of a male G. sauteri of similar size. Galeus eastmani is a slender species of medium length, smaller than G. nipponensis but larger than G. schultzi. One G. eastmani that I examined had been in preservative for more than 50 yr and its color was uniform brownish without markings. Nakaya (1975) found fresh specimens gray or brownish-gray on dorsolateral surfaces and gray on the upper surfaces of the pectoral fins and vertical fins with saddles and blotches of darker color without definite edging. In the three G. eastmani examined, the distance between bases of the dorsal fins was 15.3 to 16.2% TL compared with 13.2 to 14.5% TL for two specimens of G. nipponensis, 12.7 to 13.7% TL for six G. sauteri, and 11.0 to 14.2% TL for four specimens of G. schultzi. For larger series of fresher specimens, Nakaya (1975) found (for the distance between dorsal fin bases) in G. eastmani 14.2 to 15.9% TL, for G. nipponensis 13.5 to 15.6% TL, and for G. sauteri 11.7 to 13.8% TL. Description. The following description is of the 347-mm adult male, FMNH-Owston 678. Body slender; fins relatively small; pectorals broad but short, distal margins nearly straight, greatest width (along distal margin) much greater than mouth width, tips of appressed pectorals not reaching more than halfway to the origin of the pelvics; origin of first dorsal fin over posterior part of pelvic base; distance between first and second dorsal fin bases somewhat greater than length of head to first gill slit; first and second dorsals about equal in area but the length of base of first dorsal somewhat longer; anal fin long and moderately low, its base somewhat longer than lengths of first and second dorsals combined; basal half of pelvics united to form an apron beneath the fully calcified claspers. Head narrow with moderately pointed tapering snout; mouth moderately large and regularly arched, its width about equal to the length of the snout in front of mouth; distance between inner posterior ends of nostrils about half the longitudinal diameter of the eye opening; nasal flap triangular without an extended point. Teeth very small, the largest about 1 mm high; teeth of upper and lower jaws similar in shape, the upper jaw teeth slightly larger; teeth near symphysis tricuspid, but laterally the teeth have more cusps, to five or seven cusps in extreme lateral teeth; teeth crowded and difficult to count, 20-25 rows in each half of upper jaw, 20-22 rows in each half of lower jaw. Dermal denticles small, the largest flank denticles about 0.3 to 0.4 mm long, imbricate, with three posterior points, the central one much the longest. Caudal crest denticles extending along upper edge of tail for about 65% of its length. Caudal crest near its origin consists of three central rows of symmetrical denticles between the two marginal rows of larger asymmetrical denticles. Except for a trace of lighter color near the tips of the two dorsal fins the specimen has no markings. Measurements of series. The following measurements are expressed as percentages of the total length. The first measurement in each set of three refers to the 337-mm holotype, the second to the 347-mm adult male, and the third to a 315-mm immature male. Tip of snout to: front of mouth, 6.2, 5.5, 5.4; eye, 5.6, 5.8, 5.7; spiracle, 11.6, 11.2, 10.2; first gill slit, 14.8, 13.8, 14.0; fifth gill slit, 18.4, 17.9, 18.1; origin pectoral, 17.5, 17.3, 17.1; first dorsal fin, 42.1, 43.2, 43.2; origin pelvics, 35.6, 37.2, 33.3; origin second dorsal fin, 62.3, 61.9, 63.2; anal fin, 50.7, 53.9, 52.4; origin caudal crest, 71.2, 70.6, 71.4; anterior end cloacal opening, 38.0, 38.9, 38.1. Greatest width of: trunk at pectorals, 7.4, 7.8, 7.6; trunk at pelvics, 4.5, 4.0, 4.4; caudal base, 1.8, 2.2, 2.3. Greatest height of: trunk at pectorals, 8.6, 7.5, 7.0; trunk at pelvics, 7.4, 7.5, 6.6; caudal base, 3.6, 3.2, 3.5. Eyes: horizontal diameter of opening, 4.2, 3.7, 4.4; vertical diameter, 1.2, 1.2, 1.0. Spiracles: greatest diameter, 0.6, 0.4, 0.5; least distance from eye, 0.6, 0.9, 0.7. 59 Mouth: width, 6.2, 5.8, 6.4; length mouth, 3.7, 4.0, 3.2; length upper labial furrow, 1.2, 1.4, 1.1; lower labial furrow, 1.5, 1.4, 1.3. Nasal apertures: minimum distance between, 2.1, 1.9, 1.9. Gill slits: height of first, 1.5, 1.4, 1.3; height of fifth, 0.9, 0.9, 0.6. First dorsal fin: length base, 4.7, 4.6, 4.1; length posterior tip, 2.4, 1.7, 1.7; height, 3.9, 3.2, 3.2; length anterior margin, 7.4, 6.6, 6.4. Second dorsal fin: length base, 5.6, 5.0, 5.1; length posterior tip, 2.1, 1.8, 1.9; height, 3.9, 3.3, 3.2; anterior margin, 7.4, 6.9, 6.9. Anal fin: length base, 11.9, 11.5, 12.4; length posterior tip, 1.5, 1.7, 1.3; height, 3.0, 2.3, 2.9; length anterior margin, 5.9, 5.8, 6.7. Pelvic fins: overall length, 11.0, 11.2, 9.8. Claspers: reach past pelvic fin tips, 0.0, 3.5, 2.5. Pectoral fins: width base, 4.5, 4.6, 5.1; anterior margin, 9.5, 10.0, 9.5; greatest width, 8.0, 8.3, 7.3. Distance between fin bases: first and second dorsals, 15.4, 15.3, 16.2. Discussion. Nakaya (1975:50, fig. 25) found yellow, smooth-surfaced egg capsules about 60 mm long and 16 mm wide in G. eastmani; mode of reproduction single oviparity, that is, egg capsules not retained in the oviducts for more than a very short part of the development period; no evidence of sexual dimorphism in teeth; females mature at about 360 to 370 mm. Galeus melastomus melastomus Rafinesque Galeus melastomus Rafinesque 1810:13 (occurrence in Sicilian waters implied). Pristiurus melanostomus Bonaparte 1834, fasc. VII; Muller and Henle 1841:15; Collett 1905:95; Regan 1908a:463; Lozano Rey 1928:315; Nobre 1935:418; Andriashev 1954:35; Albuquerque 1954-1956:95; Azouz and Capape 1971:123; Quignard and Capape 1971a:132, 1971b:160. Pristiurus atlanticus: Vaillant 1888:59, pi. 1. Pristiurus melastomus: Garman 1913:92; Maurin and Bonnet 1970:130, fig. 4. Galeus melastomus: Bigelow and Schroeder 1948:215; Dollfus 1955:82; Tortonese 1956:133, figs. 70-71; Cadenat 1959, figs. 5, 6; Bini 1967:61; Krefft 1968:40; Wheeler 1969:46; Springer 1973:11.3.1. Material examined. USNM 23029, !, 635 mm, Norway; Bergen Museum, ad. S, 612 mm, Norway; USNM 17494, imm. S, 407 mm; USNM 48275, imm. S, 337 mm; USNM 48276, imm. ?, 313 mm; SU 20604, imm. ?, 377 mm, Naples; ZMK Thor stn. 92, juv. S, 149 mm, west coast of Scotland from 550 m; about 100 specimens, coasts of Tunisia, Algeria, Morocco; BMNH 1934, 8.8. 3a, 4a, egg capsules, off S.W. Ireland; BMNH 1928, 9.18. 13, 16, egg capsules off S.W. Ireland, 292 m. Holotype not found. Diagnosis. Galeus melastomus melastomus occurs along continental slopes from the vicinity of Trondheim, Norway, southward around the British Isles and into the Mediterranean and Adriatic Seas. Its range overlaps that of Galeus polli in the vicinity of Tangier and off the Atlantic coast of Morocco. Two characters by which G. m. melastomus may be dis- tinguished, size at maturity and number of vertebrae, are greater in the northern part of its range than in the southern sector. Galeus m. melastomus differs from G. m. murinus in having a longer anal fin and in having much more prominent dorsolateral markings. A juvenile G. m. melastomus from the coast of Scotland has the base of the anal fin 18.1% TL as compared with the juvenile holotype of G. m. murinus which has the base of the anal fin 11.8% TL. Galeus m. melastomus differs from G. polli in producing its eggs in leathery capsules. It is a larger species reaching more than 600 mm as compared with G. polli which reaches a length of about 450 mm. It has smaller and more numerous dorsolateral markings than G. polli. Its anal fin base length is more than 15.0% TL as compared with less than 14.0% TL for G. arae arae. Its strongly marked dorsolateral surfaces serve to mark it off from plain colored species such as G. sauteri or from species with diffuse markings such as G. nipponensis. Krefft (1968:40) noted additionally that the tip of the second dorsal fin extended rearward past the origin of the caudal crest and posterior tip of the anal fin in melastomus but not in polli. He also noted greater numbers of vertebrae in North Sea specimens of melastomus than in African coast specimens of polli and observed that melastomus usually had 15 to 18 saddle blotches as compared with 9 or 10 for polli. Maurin and Bonnet (1970, fig. 4) illustrated for comparison a 410-mm male melastomus and a 410-mm male polli, presumably both from the northwest coast of Africa. The illustration shows a larger and wider band for melastomus; a less arched mouth; a longer pectoral; a greater distance between the nostrils; and a somewhat lesser distance between the axis of the pectorals. Description. Galeus melastomus melastomus reaches a length of 900 mm (Krefft 1968) but that length may be unusual. Among specimens examined for this study a male 612 mm long from Norway was certainly an adult and a 60 female 635 mm long, also from Norway, probably was an adult. Individual variation in proportional measurements is substantial in G. m. melastomus as it is for many scyliorhinid species. Variation in vertebral number is great but, in part, appears to be geographical. Following are measurements expressed as percentages of the total length. The first figure in each series represents a 612-mm adult male from Norway; the second represents a 635-mm female from Norway; the paired figures in parentheses give the range in four half-grown specimens, two males and two females 313 to 407 mm long from Naples; the final figure in each series represents a juvenile male 149 mm long from the west coast of Scotland. Tip of snout to: front of mouth, 7.8, 6.3, (6.4-8.6), 8.2; eye, 7.2, 6.3, (6.4-8.3), 7.2; spiracle, — , 10.6, ( ), 13.4; first gill slit, 15.0, 12.6, (15.4-17.5), 17.4; last gill slit, 21.2, 17.2, (18.8-23.0), 21.2; origin pectoral, 21.6, 16.7, (18.1-22.1), 21.6; origin first dorsal, 48.5, 46.1, (44.9-48.2), 40.9; origin pelvics, 41.5, 39.1, (36.9-39.6), 34.9; origin second dorsal, 66.7, 63.9, (61.5-63.5), 55.7; origin anal, 58.0, 53.9, (52.1-53.6), 46.3; origin upper caudal lobe (= origin caudal crest), 76.0, 72.1, (70.0-72.5), 65.1; anterior end anal opening, 43.0, 41.9, (40.3-43.1), 39.6. Eyes: length eye opening, 4.2, 3.6, (4.2-4.5), 5.4. Spiracles: greatest diameter, 0.7, — , ( ), 0.4; least distance from eye, 0.8, 0.8, (0.7-1.3), 0.4. Mouth: width, 7.8, 7.9, (6.2-7.9), 6.7; length, 3.9, 3.3, (3.4-5.4), 4.0; length upper labial furrow, 1.5, 1.4, (1.0-1.6), 1.3; lower labial furrow, 1.6, 1.6, (1.3-1.9), 2.0. Nasal apertures: least distance between, 2.5, 2.4, (2.9-3.5), 2.7. Gill slits: height of first, 2.1, 2.8, (2.0-2.7), 1.3; height of fifth, 1.1, 1.4, (0.9-1.3), 0.8. First dorsal fin: length base, 5.7, 4.9, (4.4-5.1), 6.0; length anterior margin, 8.7, 7.9, (7.7-7.9), — . Second dorsal fin: length base, 5.4, 4.6, (4.8-5.4), 6.7; length anterior margin, 8.3, 7.6, (8.0-8.6), — . Anal fin: length base, 15.4, 15.3, (12.8-15.5), 18.1; length anterior margin, 8.2, 7.2, (6.4-8.1), — . Pectoral fins: greatest width, 11.4, — , ( ), 8.7; length anterior margin, 11.4, 13.2, (11.9-12.8), 11.4. Pelvic fins: overall length, origin to posterior tip, 11.4, — , ( ), 10.7; reach of claspers posterior to pelvic fin tip, 6.0, — , ( ), -4.0. Distance between fin bases: first and second dorsals, 13.2, 14.0, (10.9-12.3), 9.4; pectoral and pelvic, 11.8, 15.3, (9.6-11.7), — . Galeus melastomus melastomus has high numbers of vertebrae compared with other species of Galeus for which vertebral counts were made. The two specimens from Norway had 40 and 43 monospondylous vertebrae and 28 Mediter- ranean specimens, mostly from the vicinity of Tangier, had 37 to 40 vertebrae. Quignard and Capape (1971b) reported 39 and 40 monospondylous vertebrae from specimens taken off Tunis. Galeus m. melanostomus is strongly marked with a pattern of dark saddle blotches over the dorsolateral surfaces. Ventral surface is uniformly light colored. Juveniles have 7 or 8 saddle blotches, but the number increases during growth to as many as 19 in very large specimens. Discussion. Egg capsules about 58 by 21 by 14 mm, not including horns or tentacles, taken off the southwest coast of Ireland from about 290 m presumably belong to G. m. melastomus. Azouz and Capape (1971) reported partially digested crustaceans and cephalopods from the stomachs of the species taken off Tunis. The name melastomus refers to the black color of the inside of the mouth. Black mouths seem to be usual among species of Galeus although the color is rather quickly lost after preservation. Galeus melastomus murinus (Collett) Pristiurus murinus Collett 1904:4 (Faroe Channel from 1,200 m). Pristiurus jenseni Saemundsson 1922:169, pi. 4, fig. 2, pi. 5, fig. 4; not pi. 4, fig. 1 and pi. 5, fig. 3 as indicated by Saemundsson. (Vestmannaeyjar off southern coast of Iceland.) Pristiurus murinus: Regan 1908a:463; Garman 1913:93. Galeus murinus: Wheeler (lx, in list of species); Springer 1973:11.3.2. Material examined. ZMO, Michael Sars stn. 76, imm. ?, 222 mm, holotype (specimen examined at Zoological Museum, Bergen). Diagnosis. In the juvenile 222-mm female holotype of Galeus marinus, the length of the base of the anal fin is 11.8% TL but in a 149-mm juvenile of Galeus melastomus from the west coast of Scotland the length of the base of the anal fin is 18.1% TL. Galeus murinus was originally described as being uniformly brown above, lighter below without the variegated pattern that characterizes G. melastomus. Saemundsson's holotype of Pristiurus jenseni, a 630-mm male, as measured by Saemundsson (1922:171) differs only moderately in proportional dimensions from the holotype of murinus. 61 For example, its eye length is 3.4% TL and the length of its anal fin base is 13.5% TL, dimensions that set it off somewhat from murinus but in the direction of melastomus characteristics. Description. After describing murinus in 1904, Collett (1905) prepared a more thorough description of the species but again referred to only one specimen. Following are proportional measurements I made in 1969 when the total length of Collett's holotype was 222 mm. Figures are percentages of total length. Tip of snout to: front of mouth, 8.1; eye, 8.6; spiracle, 13.6; first gill slit, 16.8; fifth gill slit, 20.5; origin pectoral, 19.5; origin first dorsal, 44.1; origin pelvics, 36.8; origin second dorsal, 55.9; origin anal, 48.6; origin caudal crest, 66.4; anterior end cloacal opening, 42.7. Eyes: horizontal diameter, 4.1; vertical diameter, 1.4. Spiracles: greatest diameter, 0.6; least distance from eye, 0.8. Mouth: width, 7.7; length, 3.6; length upper labial furrow, 1.7; length lower labial furrow, 2.1. Gill slits: height of first, 1.1; height of fifth, 0.9. First dorsal fin: length base, 5.2; length free inner margin, 2.5; height, 2.5; length anterior margin, 8.0. Second dorsal fin: length base, 5.9; length free inner margin, 3.2; height, 2.5; length anterior margin, 9.1. Anal fin: length base, 11.8; length free inner margin, 0.9; height, 4.1; length anterior margin, 9.5. Pectoral fin: width pectoral base, 5.9; length anterior margin, 11.4; greatest width, 7.7. Pelvic fins: origin to rear tip, 10.5. Caudal fin: length from origin caudal crest, 32.3. Distance between fin bases: first and second dorsals, 8.6; anal to lower caudal lobe, 0.9. Discussion. As pointed out by Blacker (1962:269), most "rare" fish are common somewhere. Blacker was referring to occasional, surprisingly large catches of Galeus melastomus made by research trawlers although that species had been considered uncommon in commercial trawler catches. In conversation with a group of Grimsby, England, trawlermen I learned that Galeus was indeed taken in large numbers although not regularly by "distant water" trawlers fishing in the direction of Iceland. Also, I learned that the black-mouths (Galeus) were variable in color, not so spotted and grayer in color toward Iceland. Galeus nipponensis Nakaya Figures 33, 34 Galeus nipponensis Nakaya 1975:51, figs. 26-28 (Mimase, Kochi Prefecture, Japan). Material examined. SU 13887, S, 592 mm, Japan; BMNH 1937.7.7.7, subad. S, 550 mm, Sagami Channel, Japan. Diagnosis. Nakaya's diagnosis, directed chiefly toward separating G. nipponensis from other Galeus of Japanese waters is as follows: body with dark blotches; origin of anal fin below middle or posterior half of interdorsals; inside of mouth white; snout in front of anterior nasal aperture longer than horizontal diameter of eye. Additionally I note that G. nipponensis has remarkably wide and short pectoral fins, their greatest widths, measured parallel to distal fin margin, considerably greater than the width of the mouth or length of the snout in front of mouth, and equal to or slightly greater than the length of the base of the anal fin; the pectoral fins short, their lengths equal to or not much greater than their widths. The first dorsal fin in G. nipponensis is slightly larger than the second dorsal fin in Figure 33. — Galeus nipponensis, female, 5!)2 mm. Lower jaw damaged and absence of lower labial furrow probably due to injury. Drawing by Mildred Carrington. 62 Figure 34. — Galeus nipponensis, subadult male, 550 mm, ventral side of head, from BMNH 1937.7.7. all dimensions and this differentiates it from many Galeus species. The snout in front of mouth is tapering and about as long as the width of the mouth. Description. The specimens that I examined had been in preservative for 30 yr or more and no markings were visible on their uniform brown dorsal surfaces or somewhat lighter ventral surfaces. The male had a darker smudge on the upper part of each dorsal fin. The 550-mm male has very long but uncalcified claspers, their tips reaching the origin of the anal fin and extending past the tips of the pelvic fins for a distance greater than the length of the snout. In this specimen the extension of the claspers past the tips of the pelvic fins is 6.9% TL. Only Galeus schultzi and Juncrus vincenti, among adult male scyliorhinids, have proportionally longer claspers. The claspers of adult male G. sauteri reach nearly to the base of the anal fin, but in the series of five adult male G. sauteri examined for this study the claspers extend beyond the tips of the pelvic fins by only 2.8 to 3.6% TL. The claspers of the 347-mm adult male G. eastmani examined extended past the pelvic tips only about halfway to the anal fin origin, 3.5% TL. Dermal denticles of the flank of the 592-mm female are imbricate, tridentate, and about 0.3 to 0.4 mm long. Thus, the denticles are about the same size as those on G. eastmani but smaller than the flank denticles of the 0.7-mm denticles that appear typical of G. sauteri. Following are measurements expressed as percentages of the total length. Measurements for the 592-mm female are represented by the first figure in each pair; the 550-mm male's proportions are shown by the second figure. Tip of snout to: front of mouth, 6.4, 6.9; eye, 6.4, 7.3; spiracle, 11.1, 10.7; first gill slit, 15.5, 15.3; fifth gill slit, 19.9, 19.1; origin pectoral fin, 19.4, 18.4; origin first dorsal, 43.1, 43.6; origin pelvic fins, 38.5, 37.8; origin second dorsal, 62.7, 65.4; origin anal fin, 57.6, 60.0; origin caudal crest, 71.6, 71.8; anterior end cloacal opening, 40.7, 40.9. Eyes: horizontal diameter, 3.9, 3.5; vertical diameter, 1.5, 1.1. Spiracles: greatest diameter, 1.0, 0.9; least distance from eye opening, 0.8, 0.5. Mouth: width, 6.4, 6.9; length, 3.9, 3.5; length upper labial furrow, 2.0, 1.6; length lower labial furrow, — , 2.0. Nasal apertures: minimum distance between, 2.1, 2.0. Gillslits: height of first, 1.5, 1.5; height of fifth, 0.9, 1.0. First dorsal fin: length base, 6.1, 6.2; length lower posterior margin, 2.2, 2.2; height, 4.4, 4.5; length anterior margin, 8.8, 8.0. Second dorsal fin: length base, 5.9, 5.8; length lower posterior margin, 2.0, 1.8; height, 3.9, 3.5; anterior margin, 7.6, 7.6. Anal fin: length base, 8.8, 7.9; length lower posterior margin, 1.5, 1.8; height, 3.5, 3.3; anterior margin, 5.6, 6.4. Pectoral fins: width base, 5.2, 6.0; anterior margin, 10.1, 9.6; greatest width, 11.3, 9.1. Pelvic fins: Overall length, 11.0, 14.5. Claspers: reach past pelvic fin tips, — , 6.9. Distance between fin bases: first and second dorsals, 13.2, 14.5. Discussion. Both the specimens that I examined had previously been identified variously as G. eastmani or G. 63 hertwigi. Galeus nipponensis is nearly as large as Parmaturus pilosus (= G. hertwigi) but has a sharper, more tapered, and shorter snout as well as much wider pectoral fins. Nakaya, (1975) reported that G. nipponensis attains sexual maturity at about 530 to 550 mm; sexual dimorphism in teeth of adults is minor; the egg capsule is finely striated longitudinally, 89 by 20 mm, rather thick-walled and strong, brown, opaque; and a hatched embryo is 133 mm TL. Galeus piperatus Springer and Wagner Figure 35 Figure 35. — Galeus piperatus, female, 302 mm, holotype, Gulf of California. The plain color of this specimen may be unusual. The few other specimens now available have a color patttern approaching that of G. arae arae. Drawing by Mary Wagner. Galeus piperatus Springer and Wagner 1966:1, figs. 1, 2 (Gulf of California); Hubbs and Taylor 1969:310, figs. 1-7; Brewer 1973:5. Galeus sp. Lavenberg and Fitch 1966:96, 98. Material examined. LACM 7552, ad. ?, 302 mm, holotype; LACM 8818, imm. ?, 256 mm, paratype; USNM 200413, ad. ?, 296 mm, paratype. All types from RV Alaska stn. 64A2-16, lat. 28°55'N, long. 112°50.5'W, south of Isla de la Guar- dia in 402-412 m, Gulf of California. Diagnosis. When described from the type-material listed above, Galeus piperatus was stated to lack well-defined dorsal blotches as in the holotype or to have indistinct mottling as in the other two specimens. Hubbs and Taylor (1969) examined six additional specimens and found moderately conspicuous markings set off by a light subhexagonal reticulum approaching that figured by Springer (1966, fig. 20) for a specimen of G. arae of comparable size. The lack of markings on the holotype (as in Fig. 35) may have been due in part to the method of preservation as suggested by Hubbs and Taylor. I have not seen a preserved Galeus arae arae nearly or entirely lacking markings, however, among hundreds that I have examined and I suspect that G. piperatus is a more variable species than G. arae arae. The separation of G. piperatus from G. arae arae seems a lesser problem than its separation from G. arae cadenati, and I doubt that such separations can be made with confidence except on the basis of large series of both populations. Some confirmation that G. piperatus differs from G. arae arae and G. arae cadenati lies in the vertebral count (Table 2) but overlap is present for both subspecies. Description. Measurements of the three type-specimens are given by Springer and Wagner (1966) and also by Hubbs and Taylor (1969) for the same specimens with an additional five specimens. These measurements are not repeated here. As noted several times in the present study, scyliorhinid sharks shrink, swell, and twist in a variety of ways depending on the manner of their death and preservation. Furthermore, during storage in alcohol they show a fairly consistent tendency to shrink in total length but such shrinkage does not appear to be uniform for all dimensions. In addition, small differences in method of measurement make small differences in results, and differences in defini- tion of whatever is measured may produce some remarkably divergent figures. It is instructive to compare the measurements made by Springer and Wagner with those made by Hubbs and Taylor. The agreement is only approximate for many measurements. For one measurement it is obvious that different things were being measured and reported under similar but confusing names. Springer and Wagner referred (1966:3) to "inter- space between first and second dorsal" and (in table 1) to "distance between dorsal fins." In both instances the space be- ing measured was the distance between the posterior end of the first dorsal fin base and the origin of the second dorsal fin base. Hubbs and Taylor (1969) referred to (table 2) "interdorsal space" and the measurement reported was the distance from the appressed posterior tip of the first dorsal fin to the anterior end of the base of the second dorsal. 64 Hubbs and Taylor (1969) reported a scyliorhinid egg capsule 35 mm long believed to belong to G. piperatus for several reasons that seem valid. They also found one juvenile, 85 mm in life but 80 mm when preserved. Their illustra- tion (1969, fig. 2) shows the usual color of juveniles of several Galeus species. Galeus polli Cadenat Galeus polli Cadenat 1959:395, 18 figs, (coast of Senegal); Krefft 1968:40. Galeus melastomus: Fowler 1936:40; Poll 1951:22, figs. 6, 7, pi. XII, fig. 1. Pristiurus polli: Maurin and Bonnet 1970:131, fig. 4. Material examined. USNM, 58 uncatalogued specimens, west coast of Africa between lat. 10°N and 5°S; USNM, 17 uncatalogued specimens, Liberia to Angola; USNM, 2 S, 360-385 mm, 1 2, 320 mm, from 420 to 450 m at lat. 35°41'N, long. 04°55'W, near Ceuta, Spanish Morocco; Guinean Trawling Survey, lat. 09°32'N to 05°17'N, 10 ?, 210 to 299 mm, 6 S, 205 to 228 mm, west coast of Africa. Diagnosis. Galeus polli has a longer anal fin than G. arae arae, G. arae antillensis, and probably longer than some G. arae cadenati, G. melastomus murinus, and some G. melastomus melastomus. Its longer snout (in front of mouth) separates it from western Atlantic species. Its color pattern distinguishes it from G. melastomus murinus and from the unmarked or weakly marked Pacific species. Galeus polli and G. melastomus melastomus have overlapping ranges off the coast of Spanish Morocco and probably for some distance southward off the Atlantic coast of Morocco and off Senegal. Distinguishing G. polli and G. m. melastomus is not always possible in the area of overlap on the basis of single morphometric characters. Adult G. polli reach lengths up to 430 mm, but G. m. melastomus grows to a much larger size, especially in the north- ern part of its range (Cadenat 1959; Krefft 1968; Maurin and Bonnet 1970). Galeus polli adults have from 9 to 11 dorsal saddle blotches as compared with 15 to 18 for G. m. melastomus; G. polli has smaller denticles in the caudal crest than G. m. melastomus; the tip of the anal fin overlaps the origin of the lower caudal fin lobe in G. m. melastomus but barely reaches it in G. polli (Krefft 1968). The least distance between nostrils in G. polli is half or less than half the distance between the pectoral axils in G. polli but about equal to that distance in G. m. melastomus (Maurin and Bonnet 1970, fig. 4). Krefft (1968) noted differences between the number of trunk vertebrae, 33 to 35 for polli and 40-41 for melastomus from the northern part of its range. The difference decreases somewhat toward the area of overlap. Quignard and Capape (1971b) found 39 and 40 monospondylous vertebrae (= trunk vertebrae) for melastomus from the coast of Tunis. The three polli noted in the material examined section above from Spanish Morocco had 33, 34, and 34 monospondylous vertebrae. Among specimens that I have seen, G. polli has stronger markings than other species of Galeus and the dorsolateral color and saddle blotches were gray and gray-black rather than brown. Description. The following dimensions expressed as percentages of total length were taken from Cadenat (1959:396); the series of figures for each dimension refer first to a 390-mm male, second to a 350-mm male, third to a 410-mm female, and last to a 415-mm female. Tip of snout to: front of mouth, 8.2, 7.4, 8.5, 7.4; origin first dorsal, 46.1, 44.2, 45.1, 43.3; origin second dorsal, 62.8, 62.8, 62.1, 57.8; origin anal, 53.8; 54.2, 51.2, 48.0; pectoral, 19.2, 20.0, 19.5, 19.2; pelvic, 38.4, 38.5, 37.8, 34.9. First dorsal fin: length of base, 5.1, 6.5, 7.3, 4.8; height, 2.8, 2.8, 3.1, 2.4. Second dorsal fin: length of base, 5.1, 5.6, 6.0, 6.0; height, 3.0, 3.1, 3.4, 2.8. Anal fin: length of base, 14.0, 15.1, 17.0, 15.6; height, 3.8, 4.2, 4.2, 4.3. Length upper caudal border: 28.2, 28.5, 28.0, 25.3. Greatest length pectoral fin: 14.6, 12.8, 13.4, 43.3. Width of mouth: 7.2, 7.4, 7.8, 7.4. Distance between fin bases: first and second dorsals, 11.5, 12.0, 9.7, 9.6; second dorsal to upper caudal, 3.8, 2.8, 3.6, 3.6. Distance between fin origins: pectoral and pelvic, 19.2, 18.5, 18.2, 18.0; pelvic and anal, 15.3, 14.2, 13.4, 13.2. Body comparatively slender; length of snout in front of mouth about 2'/ 2 in head; head about 20% TL; tail from origin of caudal crest about 25 to 30% TL; orbit about 3.0 to 4.0% TL. Origin of first dorsal over posterior part of base of pelvic fin; dorsal fins not greatly different in size, the second slightly larger; anal fin long, much longer than distance between bases of dorsal fins, its tip barely reaching origin of lower lobe of caudal fin. Dermal denticles and teeth similar to those of other species of Galeus. 65 Discussion. Galeus polli eggs, as found in the oviducts before appreciable embryonic development (Cadenat 1959, figs. 8-10), have transparent and apparently very thin shells which soon disappear. Galeus polli gives birth to living, fully formed young. Egg capsules are not heavy-walled, substantial structures and appear to be absorbed in early stages of development of the embryos in the oviducts. Cadenat reported that the females produce from 4 to 10 young at a time, the sexes in nearly equal numbers among 51 foetuses counted. Young apparently are born at a length of about 115 mm. Poll (1951) had already reported on the capture of 77 specimens taken between lat. 6°08'S and 19°52'S on the west coast of Africa and had noted that the species was ovoviviparous "contrairement a l'opinion generale adnaise que les Scyliorhinidae sont ovipares." That Poll used the name Galeus melastomus for these sharks is not so surprising to me as is Poll's apparent willingness to discard opinions of dozens of zoologists that G. melastomus lays eggs in leathery cases. It is nevertheless true that the literature on the development of G. melastomus is scanty and some of its conclusions may be based on assumptions. Cadenat (1959:398) included a table with lengths and weights of males and females more than 300 mm long. The table shows that females 355 to 415 mm long weigh from 128 to 220 g and that males 305 to 390 mm long weigh from 70 to 140 g. Cadenat (1959:400) determined the liver weight in a series of 16 adult G. polli to be from 3.6 to 8.1% of total weight (average 5.47% total weight). I do not know of another report on the weights of scyliorhinid sharks. Galeus sauteri (Jordan and Richardson) Pristiurus sauteri Jordan and Richardson 1909:160, pi. 63, fig. 1 (Takao, Formosa). Galeus sauteri: Fowler 1941:28; Teng 1962:43, fig. 9; Chen 1963:27, fig. 8; Nakaya 1975:41, figs. 20-22. Material examined. SU 21261, 4 ad. J, 360-365 mm, syntypes, Takao, Formosa, coll. Hans Sauter, 1906; USNM 191185, ad. 5, 380 mm, 2 ad. ?, 415-430 mm, Tamshui, Taipei, Taiwan; USNM, juv. 5, 138 mm, Ta-chi, Taiwan, 23 November 1969. Diagnosis. Galeus sauteri is distinctively marked with the upper edges of the first and second dorsal fins and caudal fin with a black spot near the tips; the pectorals and pelvics are unmarked except for a moderately distinct band of lighter color along the distal margins; the dorsal and lateral surfaces are plain gray or brown; ventral surfaces are white or yellowish, the light color extends somewhat on the lateral surfaces, especially posterior to the pelvics; the color of the juvenile specimen examined is similar to color of adults except the apex of the anal fin and the apex of the lower caudal lobe are black tipped and the dark and light areas are in stronger contrast. Galeus sauteri adul,t males, at about 360 to 385 mm TL, are in contrast to G. schultzi in which the adult male holotype is about 260 mm. Galeus sauteri is smaller than nipponensis. In the latter, a 550-mm male is not quite sexually mature. The claspers of mature male G. sauteri are proportionally about as long as those of other species of Galeus except nip- ponensis and schultzi. In G. sauteri, the tips of the claspers reach approximately to origin of the anal fin but reach past the tips of the pelvics for a distance a little more than half the length of the snout. In nipponensis, the claspers reach past the pelvic fin tips by a distance about equal to the snout length. In the study series of G. sauteri, the anal fin base is longer at 11.7 to 15.0% TL than nipponensis for which the propor- tion is 8.2 to 8.8% TL. The proportion of the distance between dorsal fin bases in sauteri is less than that of eastmani and nipponensis and about equal to that of schultzi. Description. Although the dermal denticles are imbricate they are somewhat rough to touch. Flank denticles are about 0.7 mm long with three points and one strong central ridge. The caudal crest denticles of the central rows are about 1.0 mm long and the lateral crest denticles about 2.0 mm long. The jaws are strongly arched. The teeth are small, crowded, each tooth with 3 to 5 points. Several symphyseal teeth are present. The claspers reach past the tips of the pelvic fins in the mature males for a distance about equal to the length of the eye. The pelvic fins are united beneath the claspers for about half their length forming a partial apron. Measurements expressed as percentages of the total length follow. The first pair of figures under each category gives the range in the four syntypes, all mature males, 360 to 365 mm long. The second pair of figures represents the range in two females 420 and 440 mm long. The final single figure represents a 385-mm adult male. Tip of snout to: front of mouth, 6.1-6.7, 6.4-6.6, 6.2; eye, 6.3-7.2, 6.7-6.8, 6.8; first gill slit, 15.6-16.7, 15.7-16.0, 15.3; fifth gill slit, 18.6-19.7, 19.8-20.7, 20.0; origin pectoral, 17.5-18.4; 19.0-20.0, 19.5; origin first dorsal, 44.5-45.9, 43.3-43.6, 44.1; origin pelvic fins, 36.1-39.5, 38.4-39.8, 37.4; origin second dorsal, 63.3-63.9, 62.4-62.5, 60.3; origin anal, 53.9-54.8, 52.3-53.6, 53.2; origin caudal crest, 72.2-75.1, 68.9-73.8, 71.2; anterior end cloacal opening, 40.6-43.6, 40.0-42.9, 39.7. Eyes: horizontal length, 3.6-4.2, 3.8-4.1, 3.9. 66 Spiracles: greatest diameter, 0.8-0.8, 0.7-0.9, 0.8; distance from eye, 0.8-0.8, 0.7-1.1, 0.8. Mouth: width, 6.0-7.0, 6.6-6.7, 6.0. Nasal apertures: least distance between, 2.2-3.0, 2.4-2.5, 2.3. Gillslits: height of first, 1.1-1.4, 1.6-1.7, 1.3; height of fifth, 0.8-0.8, 1.0-1.1, 0.8. First dorsal fin: length of base, 4.2-5.3, 5.2-5.9, 4.7; length anterior margin, 6.7-7.5, 7.6-8.2, 6.0. Second dorsal fin: 4.2-5.0, 4.3-4.5, 4.2; length anterior margin, 5.5-6.7, 6.4-6.7, 6.0. Anal fin: length base, 11.7-13.9, 15.0-15.0, 12.5; length anterior margin, 5.3-6.4, 4.5-6.7, 5.7. Pectoral fin: length anterior margin, 10.1-11.1, 10.0-10.4, 8.8; greatest width, 8.5-8.9, 10.2-10.2, 8.6. Caudal fin: length from origin caudal crest, 26.7-29.5, 26.0-28.2, 28.6. Distance between fin bases: first and second dorsals, 13.1-13.7, 12.7-12.9, 12.7. Discussion. Galeus sauteri is common around Taiwan (Formosa) and according to Nakaya (1975:46) may occur around the southernmost islands of Japan. A specimen from the Philippines in the California Academy of Sciences collection may belong to this species (L. J. V. Compagno, pers. commun.). Nakaya (1975) described the egg capsules of G. sauteri as transparent, yellow, and about 35.5 mm long by 15 mm wide. The egg capsule surfaces were smooth without longitudinal streaks and had long tendrils. Galeus schultzi New Species Holotype: USNM 122312 (Albatross Stn. 4693), ad. 5, 297 mm, Balayan Bay, Luzon, Philippines, from 329 m, 20 February 1909. Paratypes: USNM 122307, ad. S, 254 mm, China Sea off southern Luzon from 431 m, 16 January 1908; USNM 122311, ad. $, 268 mm, Balayan Bay, Luzon, from 391 m, 22 February 1909. Other material examined: USNM, imm. 3, 220 mm, Philippines. Diagnosis. Galeus schultzi is one of the smallest species of Galeus. It is sexually mature at less than 300 mm. The holotype is the largest specimen of those available, only 297 mm TL. The specimens had been in preservative for more than 50 yr before I saw them and presumably had faded somewhat and had assumed the light tan color common to preserved sharks. Four saddle blotches were visible on the holotype, one at the first dorsal, one at the second dorsal, and two on the tail. The first and second dorsal fins are nearly equal in area. The anal fin base length is greater than that of G. nipponensis, but is variable and overlaps anal base lengths of G. eastmani and G. sauteri. The claspers of the 254-mm adult male are longer proportionally than other Galeus except possibly G. nipponensis. The small size (length at sexual maturity) is probably the most reliable character for the separation of G. schultzi from other Galeus now known from the Pacific. Galeus schultzi resembles G. piperatus in more ways than any other Galeus species. Following is a direct comparison of the 297-mm holotype of G. schultzi with a 293-mm paratype of G. piperatus. The head of G. schultzi is narrower, the eye slightly smaller, the nasal apertures somewhat less oblique, the mouth smaller with a higher arch, gill slits only about half as long, posterior tip of pelvics to origin of anal longer, caudal crest much stronger, outer rows of crest den- ticles larger, fewer middle crest denticles, and caudal crest flatter in schultzi than in piperatus. Description. The measurements of the type-series made 50 yr after preservation are given below expressed as percentages of the total length. The first figure in each series represents the 297-mm female holotype, the second the 268- mm female, and the third the 254-mm mature male. Tip of snout to: front of mouth, 5.9, 5.4, 5.5; eye, 6.1, 5.2, 6.3; spiracle, 10.8, 10.4, 10.4; first gill slit, 16.5, 15.7, 14.6; fifth gill opening, 21.2, 19.4, 19.7; origin pectoral fin, 20.2, 17.9, 18.5; origin first dorsal, 45.4, 44.7, 46.3; origin pelvics, 39.4, 38.0, 37.0; origin second dorsal, 62.6, 61.9, 64.6; origin anal, 55.6, 54.1, 57.9; origin caudal crest, 70.7, 70.9, 74.4; anterior end cloacal opening, 41.1, 43.3, 41.9. Eyes: length of opening, 3.7, 2.4, 4.1. Spiracles: greatest diameter, 0.7, 1.0, 0.9; least distance from eye, 1.3, 1.1, 1.2. Mouth: width, 6.6, 6.5, 6.7; length, 4.0, 3.7, 4.7; length upper labial furrow, 0.7, 0.4, 0.6; length lower labial furrow, 1.2, 1.1, 1.2. Nasal apertures: least distance between, 2.4, 2.4, 2.8. Gillslits: height of first, 1.5, 1.0, 0.9; height of fifth, 1.0, 0.7, 0.6. First dorsal fin: length base, 5.1, 6.3, 4.7; length free inner margin, 2.7, 2.6, 1.6; height, 4.0, 3.4, 3.5; length anterior margin, 8.4, 9.7, 5.9. Second dorsal fin: length base, 5.6, 6.2, 4.7; length free inner margin, 2.8, 2.8, 1.9; height, 3.0, 3.4, 3.0; length anterior margin, 8.1, 8.6, 4.7. 67 Anal fin: length base, 11.8, 11.2, 11.8; length free inner margin, 1.7, 1.7, 1.4; height, 3.7, 3.7, 2.6; length anterior margin, 6.7, 6.3, 4.7. Pectoral fins: width base, 4.7, 4.5, 5.1; anterior margin, 12.6, 12.3, 12.6; greatest width, 9.4, 10.2, 9.1. Pelvic fins: overall length, 12.1, 11.9, 9.5; reach of claspers past posterior tips of pelvics, — , — , 8.3. Distance between fin bases: first and second dorsals, 12.1, 11.6, 14.2; pectoral and pelvic, 13.8, 14.5, 12.6; pelvic and anal, 8.8, 7.1, 11.8; anal and lower caudal lobe, 5.4, 4.1, 4.3; second dorsal and origin caudal crest, 2.7, 3.0, 3.1. Teeth of the holotype, small, crowded, in about 24+24 rows in the upper jaw, probably not quite so many in the lower jaw but number of rows not readily counted. About three rows near the symphyses in each jaw tricuspid, other teeth with five cusps or a few with six or seven cusps near corners in lower jaw. Dermal denticles moderately small, largest flank denticles about 0.4 to 0.5 mm long, with three points, the central one much the longest. Asymmetrical marginal caudal crest denticles 1.0 to 1.2 mm long. The specimens had been in preservative for about 60 yr when examined and it seems probable that most of the obscure markings may once have been moderately strong saddle blotches. Those at the first and second dorsal fins and two over the caudal appeared most definite. The inside of the mouth was dusky and some scattered small denticles could be felt on the roof of the mouth. Discussion. As in some other Galeus species, individual variation may account for the differences in measurements in the type-series. The specimens were first recognized and set aside in the USNM collection by Henry W. Fowler but evidently were forgotten in the press of other work. Fowler had labeled the specimens Galeus schultzi, the specific name given to honor Leonard P. Schultz, long Curator of Fishes at the National Museum. I am pleased to follow Fowler in this as in many of his actions in shark classification. HALAELURUS GILL Halaelurus Gill 1862:407 (type-species, Scyllium biirgeri Miiller and Henle, by monotypy). Diagnosis. Species of Halaelurus lack a supraorbital crest or shelf above the orbits. Labial furrows are present and extend as a continuous groove around the mouth corners, except that in Halaelurus buergeri the furrows may be weak, very short, incomplete, or even absent on one side. Probably in exceptional examples of H. buergeri labial furrows may not be detected at all but in all the specimens I have examined furrows were present. Nasal flaps are present in varying size but do not project posteriorly past the upper lip. Species of Halaelurus have no groove or depression in the upper lip connecting the nasal apertures with the mouth. They lack a caudal crest of modified dermal denticles. The pelvic fins of males may be united near their bases but the condition is notable only in juveniles, the fins do not unite to form an apron beneath the claspers as in some other genera. The claspers of males are not notably long at sexual maturity and do not extend for more than half their length past the pelvic fin tips. The angle of view of the eyes is more lateral than dorsal and the snout in Halaelurus is not notably flattened and spatulate. Discussion. At one time or another 17 or more species have been placed in the genus Halaelurus. In the treatment here, the genus includes only nine species. Because none of the other species have been assigned to the synonymy within Halaelurus, it may reduce reader confusion to know where they went. Halaelurus garmani Fowler (1934) has earlier been regarded as a species of Scyliorhinus (Springer and Garrick 1964) and is so considered here not only because the holotype has the overlapping upper lip corner found in Scyliorhinus but also because it has a supraorbital crest. Scyllium bivium A. Smith (1837) and Scyllium chilense Guichenot (1848) are also excluded from Halaelurus because they have supraorbital crests and are placed in Schroederichthys because of their development characteristics and other features. The distinctive Catulus labiosus Waite (1905) has a supraorbital crest and too many other notable features to go in any other scyliorhinid genus than Aulohalaelurus. Two Australian species, Scyllium vincenti Zietz (1908) and Scyllium anale Ogilby (1885), like Halaelurus, do not have supraorbital crests but for other reasons are referred here to Juncrus and Asymbolus, respectively. Two South African species, Scylliorhinus regani Gilchrist (1922) and Scylliorhinus punctatus Gilchrist (1914), lack supraorbital crests but have no trace of labial furrows. They are referred to Holohalaelurus . One specimen identified by Alcock (1899) as Scyllium canescens Giinther (1878) and later described as Halaelurus alcockii Garman (1913), I have not seen and am unable to identify from descriptive accounts. It had Indian Museum, Calcutta, number 68/1. Alcock's description does not fit clearly the description of any scyliorhinid that I know and I prefer to leave it in limbo. 68 The nine species of Halaelurus that I recognize fall readily into two groups (Table 3). One group includes buergeri, boesemani, quagga, lineatus, and natalensis. They are all relatively firm-bodied, strongly marked, small sharks of the outer continental shelves and island terraces occuring from the Sea of Japan through waters off China, the East Indies, Australia, the Indian subcontinent, and the eastern coast of Africa and perhaps rarely to Saldanha Bay north and west of the Cape of Good Hope. The other group, canescens, dawsoni, hispidus, and lutarius, consists of relatively soft-bodied, drably and weakly marked species, often in deeper water of the upper continental and island slopes off the west coast of South America, New Zealand, the Andaman Islands, the Gulf of Mannar, the Gulf of Aden, and Mozambique. Halaelurus boesemani has the most extensive geographical range among species of the genus occurring from the Gulf of Tonkin to the coast of Somalia. It is present in the East Indies and also on the coast of Western Australia. Table 3. — Numbers of monospondylous vertebrae in species of Halaelurus. Species of Halaelurus 31 32 33 34 35 36 37 38 39 40 41 42 43 H. buergeri 4 2 H. boesemani 16 1 H. quagga 2 1 H. lineatus 1 6 H. natalensis 13 2 H. hispidus 14 5 1 H. dawsoni 2 4 2 H. lutarius 16 8 11 H. canescens 2 11 3 2 1 Key to Adult and Half-grown Halaelurus la. Body relatively firm, skin tightly adherent and well covered by imbricate denticles, its surface moderately hard and smooth. A color pattern of darker saddle blotches, transverse bars, and spots or some combination of these 2 lb. Body usually somewhat soft, skin either somewhat loose or at least not hard and smooth, flank denticles not closely imbricate, prickly or velvety to touch. No strong color pattern, appearance usually drab, sometimes with rather poorly defined saddle blotches, white spots, or white patches 6 2a. Snout obtusely pointed, its tip knoblike and turned upward 3 2b. Snout not pointed, its tip rounded and not turned upward 4 3a. Tan dorsally with about 26 narrow dark bars across back and extending on sides; numerous brown spots and vermiculate marks between bars; the darkest bars show tendency to occur in pairs with a single and somewhat lighter colored bar between each pair; mouth relatively small, its width about 7% or less of total length, its length 2.1 to 2.6% TL H. lineatus 3b. Tan dorsally with about 10 darker saddle blotches, each blotch made up of a pair of dark bars enclosing a somewhat lighter colored area; definite spots or vermiculations between saddle blotches absent or rare; mouth relatively large, its width 7.3 to 8.1% TL, its length 2.8 to 3.5% TL H. natalensis 4a. Color pattern made up primarily of transverse dark bars with few spots; the spots, if present, usually arranged in one or more transverse rows between bars H. quagga 4b. Color pattern made up primarily of spots; transverse markings or saddle blotches, if present, usually weak and without definite margins 5 69 5a. Dark spots not remarkably numerous and usually moderately large, usually much larger than the spiracle, often 4 to 5 mm in diameter in adults, sometimes arranged along borders of barely visible saddle blotches or sometimes arranged in clusters of two to five spots; labial furrows very short and weak, longest in adult specimens seen 2 mm, sometimes visible only on one side of mouth H. buergeri 5b. Dark spots very numerous and small, usually not appreciably larger than spiracle, usually less than 2 mm in diameter in adults, often arranged in clusters of two to five spots and usually dis- tributed over and between weakly evident saddle blotches; labial furrows moderately strong, the lower limb extending usually 5 mm or more along lower jaw in adults H. boesemani 6a. Length of base of anal fin as long or longer than the distance between dorsal fin bases; nasal flaps lobelike, not triangular, sides of lobe nearly parallel H. dawsoni 6b. Length of base of anal fin shorter than the distance between the dorsal fin bases; nasal flaps essentially triangular 7 7a. Length of anal fin base less than 1.3 times the length of the second dorsal fin base; adults 440 to 690 mm long; color uniform dark above and from uniform dark or somewhat lighter below; usually no markings of any kind but sometimes with white fin tips, especially on young specimens H. canescens 7b. Length of anal fin base 1.5 or more times the length of the second dorsal fin base; adults less than 400 mm long; color uniform gray or brown in adults or with darker saddle blotches often poorly defined or limited to the region of the first dorsal fin and posterior to it; young may have white markings 8 8a. Roof of mouth with numerous small papillae; eye length in adults less than 14 times in distance from tip of snout to first dorsal origin; adults 240 to 285 mm H. hispidus 8b. Roof of mouth without papillae; eye length in adults 14 or more than 14 times in distance from tip of snout to first dorsal origin; adults 300 to 345 mm H. lutarius Halaelurus boesemani Springer and D' Aubrey Figures 36, 37 Figure 36.— Halaelurus boesemani, adult male 447 mm, from coast of Somalia. The arrangementof spots variesjin some specimens the spots are clustered to a greater extent. Drawing by Mildred Carrington. Scyllium biirgeri (not of Mviller and Henle 1841), Bleeker 1856:69, one specimen, Amboina; Giinther 1870:404, part, adult from Amboina only. Halaelurus biirgeri (not of Muller and Henle 1841), McKay 1966:68, Western Australia, Besednov 1969:30, Gulf of Tonkin. Scyliorhinus quagga (not of Alcock 1899), Norman 1939:8, Gulf of Aden. Halaelurus boesemani Springer and D'Aubrey 1972:11, coast of Somalia. 70 L 1 mm Figure 37. — Halaelurus boesemani, upper jaw teeth and der- mal denticles. Left, tooth from 436-mm female, 4th row from symphysis. Center, dermal denticles from flank below first dorsal fin. Right, tooth from 432-mm adult male, 4th row from symphysis. Material examined. USNM 205136, ad. 8, 430 mm, holotype, coast of Somalia, 67-72 m; 3 ad. S, 416-447 mm, 3 ad. ?, 432-467 mm, damaged but identifiable female with four egg capsules, 1 juv. ?, 72 mm, all from same locality as holotype; BMNH 1939.5.24.1, ad. 8, 460 mm, Gulf of Aden; BMNH 1858.4.128, marked Frank's collection, ad. 8, 477 mm, Amboina; WAM-P14836, ad. 8, 430 mm, Western Australia. Diagnosis. Halaelurus boesemani has moderately small but readily visible labial furrows extending for a distance of 2 or more times the greatest diameter of the spiracle along the lower jaws and continuous around the mouth corner for a shorter distance along the upper jaw. Halaelurus boesemani is a firm-bodied species with imbricate denticles, smooth to touch when rubbed in the head to tail direction. It has a strong pattern of very small dark nearly round spots, the spots usually not much larger than the spiracle and sometimes, although not always, arranged in irregular clusters of from two to five spots. The spots usually overlie weakly defined saddle blotches. Halaelurus boesemani differs from H. buergeri in having smaller spots and in having much stronger labial furrows. It differs from H. quagga in not having transverse and narrow dark bars or in not having spots arranged in transverse series. Description. The teeth are small in adults, the largest about 1 mm high. The upper and lower jaw teeth are similar. The middle cusp is by far the longest and accessory cusps, usually one or two on each side of the principal cusp, are so small as to be barely visible. As in most scyliorhinids, the middle cusps are somewhat shorter toward the angles of the jaws. Vertical striations are present near the tooth bases. There are usually one to three rows of small symphyseal teeth. In H. boesemani the teeth of an adult male were larger than the teeth of a female of about the same length (Fig. 37). The dermal denticles (Fig. 37) are small, about 0.5 mm long, with a strong median ridge. The denticles are imbricate so that the skin is smooth to touch when rubbed in a head to tail direction. In eight specimens the numbers of monospondylous vertebrae were 33 to 35, the total number of vertebrae 135 to 139. Proportional measurements as percentages of the total length follow, the first figure in each pair refers to the 430-mm adult male holotype, the second to a 467-mm adult female, both from the coast of Somalia. The range of some of the measurements are given in Springer and D'Aubrey (1972, table 3) for nine specimens including one from Western Australia and one from Amboina. Tip of snout to: front of mouth, 4.0, 4.1; eye, 4.9, 4.4; spiracle, 8.6, 7.9; first gill slit, 13.3, 13.7; fifth gill slit, 17.9, 19.9; origin first dorsal, 41.9, 43.7; origin pelvic fins, 36.3, 38.8; second dorsal, 67.0, 66.2; origin anal, 59.3, 58.4; upper caudal fin, 81.4, 80.3; anterior end cloacal opening, 38.6, 41.3. Greatest width of: trunk at pectorals, 12.1, 11.8; trunk at pelvics, 7.2, 7.9; trunk at caudal origin (origin of upper caudal lobe), 2.1, 1.7. Greatest height of: trunk at pectoral, 8.1, 9.0; trunk at pelvics, 7.9, 9.0; trunk at caudal origin, 2.8, 3.0. Eyes: length of eye opening, 3.3, 3.0; height of opening 0.9, 0.9. Spiracles: greatest diameter, 0.7, 0.6; least distance from eye, 0.5, 0.4. Mouth: width, 7.9, 5.9; length, 3.5, 3.2; length upper labial furrow, 0.5, 0.4; lower labial furrow, 1.5, 1.0. Nasal apertures: least distance between, 1.9, 1.9. Gillslits: height of first, 1.9, 1.3; height of fifth, 0.9, 0.6. First dorsal fin: length of base, 7.0, 6.0; length free inner margin, 1.6, 1.9; height, 5.1, 4.7; length anterior margin, 8.6, 7.9. Second dorsal fin: length of base, 8.1, 6.4; length free inner margin, 1.9, 1.7; height, 4.0, 4.5; length anterior margin, 8.6, 7.5. Anal fin: length of base, 8.4, 8.0; length free inner margin, 1.9, 1.7; height, 3.0, 3.0; length anterior margin, 7.2, 8.1. Pectoral fin: width base, 5.3, 5.4; anterior margin, 10.5, 10.9; greatest width, 8.8, 8.6. Caudal fin: length upper margin, 18.6, 19.7. Distance between fin bases: first and second dorsals, 17.1, 15.7; pectoral and pelvic, 14.0, 13.3; pelvic and anal, 15.6, 13.3; anal and lower caudal lobe, 12.3, 14.3; second dorsal and upper caudal lobe, 7.9, 8.1. Discussion. Halaelurus boesemani on the coast of Somalia retains egg capsules in the oviducts for at least long enough for embryos to reach 49 to 54 mm and to reach the external gill filament stage of development. A 72-mm juvenile 71 with yolk stalk attached was collected at about the time the female with developing young was taken in mid-December 1964. Several egg capsules, as many as four in one oviduct, were firmly bound together by a sticky, fibrous material (Springer and D'Aubrey 1972:15, fig. 4). This method of development, in which several egg capsules with leathery shells are retained in each oviduct for a part or all of the embryonic development period, is the condition of "multiple oviparity" described by Nakaya (1975:83). Halaelurus buergeri (Miiller and Henle) Figures 38, 39 Figure 38. — Halaelurus buergeri, 442-mm adult male, Nagasaki, Japan, SU 26791. Drawing by Mildred Carrington. Figure 39. — Halaelurus buergeri, dorsal view of 442-mm adult male from Japan. Scy ilium biirgeri Miiller and Henle 1841:8, second unnumbered plate of dorsal view of whole shark (Japan), not ventral view of head; Temminck and Schlegel 1850:301; Gunther 1870:404 a, b, and c, Japan and Formosa, not d and e. Halaelurus biirgeri: Gill 1862:407 (name only); Jordan and Fowler 1903:601 (part); White 1936b, fig. 1C; Garman 1913:83 (part); Teng 1962:51, fig. 12; Chen 1963:31, fig. 10; Nakaya 1975:61, figs. 30-32. Scyliorhinus buergeri: Regan 1908a:461 (part). Halaelurus buergeri: Springer and D'Aubrey 1972:10, fig. 1C, tables 1-3. Material examined. SU 26791, 2 ad. S, 442, 428 mm, Nagasaki, Japan; BMNH 1862.11.1.38, ad. ?, 485 mm, Japan; BMNH 1862.12.26, subad. S, 404 mm, ?, 430 mm, Swintoe's collection, Formosa; BMNH 1869.11.28, 2, purchased of Bleeker (locality not stated but believed to be Japan); ZMK 16.2.1918 (1), imm. 8, 325 mm, Nagasaki, Japan; ZMK 16.2.1918 (2), imm. ?, Nagasaki, Japan, 1911 coll. Jordan. Diagnosis. Halaelurus buergeri has extremely short labial furrows or perhaps none at all in some individuals. It may have perceptible labial furrows at one corner of the mouth but not at the other. In some of the specimens examined the lower labial furrow, when present, was the longer but a turn around the mouth corner into an upper labial furrow could be seen in most instances. When visible, the lower labial furrow was 2 mm or less and the upper labial continuation of the furrow only a fraction of a millimeter long. Identification of H. buergeri should not be made from the labial furrow character alone. Halaelurus buergeri does not have a supraorbital shelf and so is easily distinguished from Scyliorhinus torazame and Cephaloscyllium isabella, species that occur in the same geographical areas as H. buergeri and have occasionally been confused with it. Halaelurus buergeri is evidently related closely to H. boesemani and the few specimens examined of each species do not show overlapping geographical ranges. In specimens examined, the labial furrows of H. buergeri were much shorter and weaker than those of H. boesemani and the rounded dark spots in the color pattern of if. buergeri were much larger. Description. Proportional measurements of two adult males, SU 26791, from Nagasaki, Japan, are given here as percentages of total length, the first figure refers to the 442-mm male, the second to the 428-mm male. 72 Tip of snout to: front of mouth, 4.1, 4.0; eye, 5.0, 4.9; spiracle, 8.1, 8.6; first gill slit, 13.1, 13.1; fifth gill slit, 16.7, 16.8; origin pectoral fin, 16.1, 16.1; origin first dorsal, 42.5, 40.2; origin pelvics, 34.2, 33.6; origin second dorsal, 63.8, 64.7; origin anal, 58.1, 56.5; origin upper caudal fin lobe, 79.6, 77.8; anterior end cloacal opening, 37.3, 37.1. Greatest width of: trunk at pectorals, 10.2, 10.3; trunk at pelvics, 6.3, 5.6; trunk at caudal origin, 2.3, 2.1. Greatest height of: trunk at pectorals, 8.1, 8.2; trunk at pelvics, 7.5, 6.8; trunk at caudal origin, 2.7, 2.8. Eyes: length eye opening, 2.9, 2.8; height eye opening, 0.9, 0.9. Spiracles: greatest diameter, 0.8, 1 .2; least distance from eye, 0.9, 0.5. Mouth: width, 6.6, 7.0; length, 4.1, 3.0; length upper labial furrow, barely visible, 0.05; lower labial furrow, absent, 0.5. Nasal apertures: minimum distance between, 2.0, 2.1. Gillslits: height of first, 1.1, 1.2; height of fifth, 0.7, 0.7. First dorsal fin: length base, 6.3, 6.8; length free inner margin, 2.3, 4.2; height, 3.6, 4.9; length anterior margin, 8.1, 8.6. Second dorsal fin: length base, 6.1, 7.2; length free inner margin, 2.0, 1.9; height, 3.6, 3.5; length anterior margin, 7.5, 8.2. Anal fin: length base, 6.8, 7.5; length free inner margin, 1.8, 1.9; height, 2.7, 2.3; length anterior margin, 6.3, 6.7. Pelvic fins: overall length, 11.8, 11.7; claspers reach past tip of pelvic fins, 3.6, 4.9. Caudal fin: length from origin upper caudal lobe, 20.4, 21.7. Distance between fin bases: first and second dorsals, 15.2, 16.4; pectoral and pelvic, 13.8, 13.3; pelvic and anal, 15.4, 15.7; anal and lower caudal lobe origin, 12.0, 13.2; second dorsal and upper caudal lobe origin, 8.8, 7.2. The teeth are tricuspid but the lateral cusps of teeth near the middle of the jaws are much smaller than the central cusp. The teeth of the upper and lower jaws are similar, those of the lower jaw slightly smaller; teeth in about 24 to 28 + 2 + 24 to 28 rows in the upper jaw, but somewhat fewer in the lower jaw. The largest teeth are about 1 mm high. Dermal denticles of the dorsolateral surfaces of the trunk have three points, the central one much the longest, the den- ticles about 0.5 to 0.6 mm long. Typical denticles have one strong median ridge, wide or double near the anterior end of the blade. In four specimens of H. buergeri the number of monospondylous vertebrae was 33 to 34, total number of vertebrae 132 to 134. The adult males examined had no apron formed by union of the inner margins of the pelvic fins but the mature male, 325 mm, had a partial apron, the pelvic fins being united about halfway from base to tips. Discussion. Springer and D'Aubrey (1972:10, 11) included Hong Kong in the geographical range of H. buergeri but listed no specimens from that area. A possible source of the reference to Hong Kong may be a 446-mm gravid female now in the California Academy of Sciences collection which is definitely H. buergeri, not H. boesemani (L. J. V. Compagno, pers. commun.). It is VFC, HK98 and was from Shan Island about 160 km southwest of Hong Kong. Specimen records available to me indicate that H. buergeri is replaced by H. boesemani southward from Hong Kong although possibly with some overlap. Boeseman (1947:215) designated as lectotype a specimen collected by Burger in Japan. It is a stuffed specimen, No. D2593 in the Rijksmuseum van Natuurlijke Historie, Leiden. Muller and Henle's (1841) description was based on a specimen in the Leiden Museum. Their description included the statement that there are almost no labial furrows ("Fast Keine Mundwinkelfalten"), but their illustration insert depicting the lower side of the head shows longer and stronger labial furrows than I have seen. Their illustration of the whole shark (dorsal view) shows the color pattern of moderately large spots that characterize the eight specimens of H. buergeri that I have seen. Evidently H. buergeri is rarely caught and preserved. I have seen only eight specimens, all taken more than 60 yr ago. Jordan and Fowler's treatment (1903) was based on a dried skin from Nagasaki. Carman's (1913) brief description seems to have been based only on the literature. The only correctly labeled specimens in United States collections are the two from the Stanford University collection noted above, and one in the Vanderbilt Foundation Collection (VFC), all three now at the California Academy of Sciences. Perhaps because of its rarity or perhaps from reluctance by ichthyologists to accept color pattern as an important character of a species, H. buergeri has sometimes been confused with the common scyliorhinid Scyliorhinus torazame of Japanese waters. Fowler's (1941:45) account of H. buergeri was obviously drawn from specimens of S. torazame. Schmidt (1930a) found that specimens collected in Tokyo in 1885 and placed in the Zoological Museum, Academy of Science, U.S.S.R., conformed to descriptions of Scyliorhinus torazame. Schmidt assumed incorrectly that the species he had should be called Halaelurus torazame instead of Scyliorhinus torazame. In torazame, as in other species of Scyliorhinus, the upper lip has a very small tab that extends down and over the lower lip when the mouth is closed, so that without close examination it may appear that a labial furrow is continuous around the mouth corner (see under Scyliorhinus torazame). Nakaya (1975:66, fig. 32) reported that H. buergeri has translucent, yellow, and smooth-surfaced egg capsules about 51 mm long and 21 mm wide with long, weak tentacles extending from each end. Several of the egg capsules may be 73 retained in each oviduct for at least a part of the embryo's development period. This method of development was called multiple oviparity by Nakaya (1975:83). It occurs also in H. boesemani (Springer and D'Aubrey 1972:14, fig. 4) and in both H. lineatus and H. natalensis (Bass et al. 1975:14, 15). Halaelurus canescens (Gunther, 1878) Figures 40, 41 Figure 40. — Halaelurus canescens, immature female, 242 mm, coast of Chile, SU 22689. Figure 41. — Halaelurus canescens, egg capsule from oviduct of (ifiO-mm specimen from vicinity of Corral, Chile (lat. 39°57'S) in 250-300 m. Scyllium canescens Gunther 1878:18 (Pacific coast of South America); type not seen but according to Garman a female, about 275 mm. Probably in British Museum. Not of Alcock 1891:310. Scyliorhinus canescens: Garman 1899:26; Regan 1908a:461. Halaelurus canescens: Garman 1913:87; DeBuen 1959:175; Kato et al. 1967:25, fig. 38; Chirichigno 1974:30. 74 ■ ft Material examined. SU 22689, imm. ?, 220 mm, off Chile, Albatross stn. 2780; USNM, 2 9, gravid, 645, 660 mm, near Corral, lat. 39°57'S, in 250-300 m, longline, mud and rock bottom, egg capsules from same area; 3 ad. 3, 550, 630, 640 mm, coast of Chile, lat. 35°26'S, in 290-450 m; 1 ad. 9, 600 mm, coast of Chile, lat. 33°22'S, in 260-280 m; 1 ad. ?, 590 mm, coast of Chile, lat. 32°08'S, in 960 m; 7 imm. $, 280-435 mm, off Chile, lat. 32°22'S, to lat. 18°10'S off Peru, 260-280 m; 3 imm. 2, 197-390 mm, off Chile, lat. 33°22'S, in 260-280 m; 1 imm. ?, 340 mm, lat. 03°57'S, off Ecuador and northern Peru in 600-700 m. Diagnosis. Halaelurus canescens is one of the drab soft-bodied species of Halaelurus. It has no spots, stripes, or saddle blotches. The dorsal surfaces are blackish and the ventral surfaces are variously dusky white to blackish. The dor- sal fins of some but not all immature specimens have whitish tips in moderate contrast to the generally dark color. It is the largest species of the genus with adults 450 to 660 mm long. Halaelurus canescens may be most closely allied to H. dawsoni of New Zealand waters but differs in having triangular nasal flaps instead of the lobelike flaps that characterize H. dawsoni. Halaelurus canescens is also different from H. dawsoni in having the anal fin base length shorter than the distance between dorsal fin bases in all but juvenile specimens. In H. canescens the length of the anal fin base in adults is less than 1.3 times the length of the base of the second dorsal fin whereas it is more than 1.5 times that length in H. hispidus and H. lutarius. Of scyliorhinids from the Pacific coast of South America, H. canescens may be confused occasionally with Schroederichthys chilensis, but among other differences H. canescens lacks the supraorbital crest that is prominent in species of Schroederichthys. There are superficial resemblances between H. canescens and Parmaturus xaniurus, a species that may in the future be reported from the equatorial eastern Pacific. In H. canescens the posterior tip of the second dorsal fin extends far past the anal tip but little, if any, past it in P. xaniurus. Description. Halaelurus canescens is a robust scyliorhinid when full grown. It has a wide head with comparatively large nostrils and a wide mouth that is only moderately arched. The snout is moderately long, that is appreciably longer than the snout in Cephaloscyllium, but definitely shorter than the snout in Apristurus. The tip of the snout varies in shape from bluntly pointed to broadly rounded. The series of specimens examined shows H. canescens to be a variable species in many respects but without indication of consistent sex or age differences in tooth form or in morphometries. The teeth are small, numerous, and difficult to count because of crowding and because they form both vertical and diagonal rows. The largest tooth in a 660-mm specimen was 1.6 mm high above the enamel line. Upper and lower jaw teeth are similar in size and shape in immature specimens, but in adults the teeth of the lower jaw appear to be slightly smaller than upper jaw teeth and teeth of females slightly smaller than teeth of males. Midjaw teeth are tricuspid but lateral teeth may have up to seven points. Dermal denticles of juveniles are needlelike and nearly erect as in many other scyliorhinids and show little develop- ment of lateral points. Denticles of dorsolateral surfaces of adults are wider, usually less erect, and sometimes have points on their lateral blade expansions. Denticles of dorsolateral surfaces of a 660-mm female were 0.8 to 1.0 mm long without lateral points; denticles from fin surfaces were 0.4 to 0.5 and were tridentate. Gill rakers about 3 mm high, three to six somewhat irregularly placed on both sides of gill bars. Liver large, posterior tips extending to or past cloaca; one functional ovary (right); one female 660 mm had one egg capsule with strong longitudinal ridges (see Fig. 41) in each oviduct. Each egg capsule contained one egg about 22 mm in diameter. The egg capsules were about 66 by 32 mm. These and several other egg capsules from the coast of Chile had from 12 to 18 longitudinal ridges, the ridges about 2 mm high. Valvular intestines in several specimens dissected had seven turns. Proportional measurements expressed as percentages of the total length follow. The first figure in each measurement category refers to an adult male 640 mm long, the second figure to a 660-mm adult female, and the third to a 197-mm juvenile female. Tip of snout to: front of mouth, 4.5, 5.1, 5.8; eye, 5.1, 5.5, 5.8; spiracle, 10.0, 11.5, 11.2; first gill slit, 16.4, 16.1, 17.3; fifth gill slit, 21.1, 20.3, 20.8; origin pectoral, 20.3, 18.6, 20.3; origin first dorsal fin, 46.9, 48,0, 41.6; pelvic fins, 43.8, 46.1, 39.1; origin second dorsal fin, 64.1, 64.2, 56.9; origin anal fin, 59.4, 58.3, 49.2; upper caudal lobe, 77.3, 76.8, 72.1. Greatest width of: trunk at pectoral origin, 11.1, 15.0, 10.7; trunk at pelvic origin, 5.8, 7.9, 4.1; trunk at caudal origin, 1.6, 1.9, 1.8. Greatest height of: trunk at pectoral origin, 8.4, 10.5, 7.6; trunk at pelvic origin, 7.7, 9.1, 6.1; trunk at caudal origin, 3.1, 3.0, 3.0. Eyes: horizontal diameter, 3.9, 5.0, 5.3; vertical diameter, 1.1, 1.5, 1.5. Spiracles: greatest diameter, 0.8, 1.1, 0.6; least distance from eye, 0.9, 0.8, 0.7. Mouth: width, 8.4, 10.3, 7.6; length, 4.5, 4.8, 4.3; length upper labial furrow, 0.8, 0.8, 0.5; length lower labial furrow, 1.5, 2.0, 1.5. Nostrils: least distance between, 1.9, 2.0, 2.3. 75 Gill slits: height of first, 2.7, 3.3, 1.8; height of fifth, 2.0, 2.3, 0.8. First dorsal fin: length base, 5.6, 6.1, 6.3; length posterior tip (= free inner margin), 3.0, 4.2, 2.3; height, 5.0, 5.2, 4.6; length anterior margin, 8.9, 10.8, 9.1. Second dorsal fin: length base, 6.9, 7.3, 8.1; length posterior tip, 2.3, 3.6, 2.5; height, 4.2, 5.0, 4.1. Anal fin: length base, 7.5, 9.2, 10.2; height, 3.4, 5.2, 2.5. Pectoral fins: width base, 5.6, 6.8, 4.3; anterior margin, 10.9, 13.5, 10.7; greatest width, 10.5, 11.7, 8.4. Pelvic fins: overall length, 10.8, 13.3, 9.4; projection of clasper past posterior fin tips, 5.0, — , — . Distance between fin bases: first and second dorsal, 9.1, 9.5, 11.6; pectoral to pelvic, 13.5, 21.2, 17.2; pelvic to anal, 6.1, 6.2, 8.9. The vertebral numbers in 19 specimens that were collected over a great range in latitude 3° to 35 °S were 39 to 42 monospondylous and 117 to 131 in total number. Discussion. Halaelurus canescens is an egg-laying and highly variable species, rather common south of the Equator to about lat. 35° S. The stomach contents of a few specimens that were dissected were empty or contained remains that were not iden- tifiable but appeared to be mostly invertebrates. Most of the specimens seen were trawled from mud bottom, but a few were collected from rock and mud areas on bottom longlines. Halaelurus dawsoni Springer, 1971 Figure 42 Figure 42. — Halaelurus dawsoni, immature female, 18G mm, from 371-375 m near Auckland Islands, New Zealand. The row of light-colored spots on the dorsolateral surface is not present on the few other known specimens, all of which are larger. Drawing by Mildred Carrington. Halaelurus dowsoni Springer 1971:235, fig. 1 (west of southern South Island, New Zealand). Material examined. DMW 5188.1, subad. S, 349 mm, holotype, trawled from 389-420 m west of southern South Island, New Zealand, lat. 44°32.5'S, long. 166°41'E, 20 November 1970; DMW 5523, 3 ?, 2 imm. S, 241 to 250 mm from same haul as holotype; USNM 206096, 1 imm. S, 324 mm, same haul as holotype, paratypes; USNM 205537, 1 imm. 5, 186 mm, trawled from 371-375 m near Auckland Islands, N.Z., lat. 50°01'S, long. 167°49'E; N.Z. Oceanographic Inst. Stn. F89, P.187, 1 ?, 242 mm, from Auckland Islands location. Diagnosis. Halaelurus dawsoni belongs with H. canescens, H. Lutarius, and H. hispidus, that is, with the drab- colored and more or less soft-bodied species of the genus. The specimen shown in the accompanying illustration (Fig. 42) was the only one of the type-series with the rows of white dorsal spots but all had broad bands or areas of white at the fin tips. Halaelurus dawsoni probably reaches 350 to 450 mm as suggested by the 349-mm subadult male holotype which has large but flexible claspers. It is thus a larger species than H. hispidus but a smaller species than H. canescens. The nasal flaps in H. dawsoni are lobelike and have nearly parallel lateral margins. The nasal flaps of other Halaelurus species are triangular or nearly so. The length of the base of the anal fin in H. dawsoni is about as great or greater than the distance between dorsal fin bases. In all other Halaelurus, except H. hispidus and H. canescens, the distance between dorsal bases is much the greater. In H. hispidus and H. canescens the interdorsal distance usually is greater. 76 Description. Halaelurus dawsoni has moderately large fins, the dorsals lobelike, the second somewhat greater in area than the first. The pectorals are broad and short and the anal fin is low on a long base, the ends of its base separated respectively from the base of the pelvics and the origin of the lower caudal lobe by distances less than the horizontal diameter of the eye. The origin of the first dorsal is over or in advance of the middle of the pelvic base and the second dorsal originates above the middle of the pelvic base. The body is relatively soft with somewhat loose skin. The gill slits in the specimens seen are short and the spiracle small, its greatest diameter about equal to its least distance from the eye. The teeth are small and crowded, and most of them have five cusps, the central cusp longest in the midsections of the jaws, the largest about 1 mm high in the 349-mm specimen, in about 64/70 rows. The dermal denticles of specimens under 300 mm long were somewhat needlelike and erect, mostly with a single posterior point and only slight lateral expansions of the blades, the expansions with or without points. In the largest specimen, the 349-mm male, the denticle blades were wider with lateral points more or less strongly developed. Vertebral numbers: monospondylous vertebrae in holotype 36 (range in eight others 36 to 38); precaudal vertebrae 67 (range 67-73), caudal vertebrae about 57 (range about 55 to 62); total number 124 (range 124 to 129). Measurements of three specimens are given below as percentages of their total lengths. The first figure in each series refers to the 349-mm subadult male holotype, the second to a 334-mm female, and the third to a 186-mm female. Tip of snout to: front of mouth, 5.4, 5.4, 6.7; eye, 5.4, 5.7, 7.0; spiracle, 10.9, 11.1, 12.6; first gill slit, 14.6, 14.4, 17.2; fifth gill slit, 18.9, 17.7, 19.9; origin first dorsal, 45.8, 44.9, 41.9; origin second dorsal, 60.5, 60.8, 57.5; origin pelvics, 39.5, 41.9, 40.3; origin anal fin, 54.7, 54.2, 50.5; origin upper caudal fin, 71.6, 71.3, 68.8; anterior end cloacal opening, 45.3, 44.9, 42.5. Greatest width of: trunk at pectoral origin, 12.3, 12.0, 10.8; trunk at pelvic origin, 7.2, 6.0, 5.4; caudal peduncle, 2.3, 2.1, 1.9. Greatest height of: trunk at pectoral origin, 7.2, 8.1, 8.6; trunk at origin pelvics, 8.3, 6.6, 7.5; caudal peduncle, 3.7, 3.6, 3.2. Eyes: horizontal diameter, 4.9, 4.0, 3.8; vertical diameter, 0.6, 0.7, 1.1. Spiracles: greatest diameter, 1.1, 0.9, 0.4; least distance from eye, 1.1, 0.9, 0.8. Mouth: width, 8.0, 7.5, 7.5; length, 3.4, 3.3, 3.3; length upper labial furrow, 0.9, 0.7, 0.4; length lower labial furrow, 2.0, 1.8, 1.8. Nasal openings: minimum distance between, 2.9, 2.4, 2.7. Gill slits: height of first, 1.6, 2.4, 1.2; height of fifth, 0.9, 0.9, 1.1. First dorsal fin: length base, 6.9, 6.9, 7.2; length free inner margin, 2.6, 2.1, 2.7; height, 5.4, 4.8, 3.2; anterior margin, 10.3, 10.2, 9.7. Second dorsal fin: length base, 7.7, 8.1, 8.6; length free inner margin, 2.9, 2.1, 2.7; height, 5.2, 5.1, 3.2; anterior margin, 11.5, 11.4, 11.3. Anal fin: length base, 11.5, 10.8, 11.6; length free inner margin, 2.0, 2.4, 2.7; height, 4.3, 4.5, 4.3; length anterior margin, 8.3, 8.1, 8.6. Pectoral fins: width base, 6.3, 6.6, 5.8; length anterior margin, 12.9, 14.4, 12.4; greatest width, 9.2, 10.2, 9.7. Pelvic fins: overall length, 11.2, 12.3, 8.6; inner margin claspers, 9.7, — , — ,; reach claspers past fin tip, 2.9, — , — . Distance between fin bases: first and second dorsal, 9.5, 9.6, 10.2; pectoral and pelvic, 17.2, 20.1, 13.4; pelvic and anal, 6.6, 5.1, 4.3; anal and origin lower caudal lobe, 4.0, 2.4, 3.8; second dorsal and origin upper caudal lobe, 2.6, 2.7, 1.9. Discussion. The four soft-bodied, drab-colored species of Halaelurus now known are from widely separated areas: lutarius from the east coast of Africa, hispidus from India and the Andaman Islands, dawsoni from New Zealand waters, and canescens from the west coast of South America. It seems likely that more exploration of moderately deep water on continental or island slopes between these areas would reveal more species. Halaelurus dawsoni ranges farther south than the other species insofar as ranges are known. No information on reproductive habits of H. dawsoni is now available. Halelurus hispidus (Alcock) Figure 43 Scyllium hispidum Alcock 1891:21 (Andaman Sea, Investigator stn. 115, 344-402 m); Alcock 1899:15 (Andaman Sea, 344-766 m); Alcock 1900, figs. 3, 3a, pi. 8 (pi. 8 dated 1894). Scyliorhinus hispidus: Regan 1908a:460 (Andaman Sea). Halaelurus hispidus: Garman 1913:85 (Andaman Sea); Springer and D'Aubrey 1972:8, figs. IB and 2B (Andaman Sea); Nair and Lai Mohan 1973:71, fig. 1 (Gulf of Mannar). 77 Figure 43. — Halaelurus hispidus, a 245-mm adult male from Andaman Sea. Material examined. BMNH 98.7.13.21 (227/1), ad. S, about 237 mm, Andaman Sea, 344-402 m, holotype; USNM, 6 imm. 8, 195 to 220 mm, 3 ad. S, 240 to 260 mm, 1 ad. ?, 285 m, all from Andaman Sea, in 293 m at lat. 10°39'N, long. 97°06'E. Diagnosis. Halaelurus hispidus is the smallest species of the genus. Alcock's largest specimen in a series of six was 280 mm; the largest USNM specimen was 285 mm; and of more than 100 reported by Nair and Lai Mohan from the Gulf of Mannar the largest was 265 mm. Halaelurus hispidus belongs with canescens, dawsoni, and lutarius, species of Halaelurus without a characteristic pattern of dark spots, bars, or saddle blotches as adults. Nair and Lai Mohan (1973) described the first hispidus taken outside the Andaman Sea and also reported on smaller specimens than had previously been collected. They observed great variation in color among 130 Gulf of Mannar specimens, mostly immature, from a single haul. Some were pale brown with gray crossbands, some had white markings, and a few were almost white with scattered patches of darker pigmentation. Alcock's first description (1891) of H. hispidus noted that the walls of the buccal cavity and the surface of the tongue are covered with small papillae. These papillae were not mentioned in Alcock's second account of the species (1899) nor did Nair and Lai Mohan (1973) mention their presence. Papillae were present on the tongue and over most of the mouth lining in the specimens I examined but were absent in lutarius; dawsoni has only a few papillae anteriorly on the roof of the mouth and on the tongue; canescens has numerous and rather large mouth papillae. Halaelurus hispidus has nearly triangular and moderately large nasal flaps, quite different from the lobelike nasal flaps of dawsoni. Halaelurus hispidus is nearest to H. lutarius but is a little smaller and has proportionally a somewhat larger eye than lutarius. Description. Following are measurements expressed as percentages of the total length. All three specimens are from the Andaman Sea. The first figure in each series represents a 255-mm adult male, the second a 285-mm adult female, and the third an immature male 193 mm long. Tip snout to: front of mouth, 5.3, 5.3, 5.9; eye, 5.3, 5.6, 6.7; spiracle, 10.4, 10.0, 10.6; first gill slit, 16.1, 15.4, 16.6; fifth gill slit, 20.6, 21.4, 22.8; origin pectoral fin, 19.6, 20.7, 21.8; origin first dorsal, 46.3, 47.7, 44.6; origin pelvic fins, 42.0, 43.5, 40.9; origin second dorsal, 65.1, 65.3, 62.2; origin anal fin, 60.4, 60.7, 57.0; origin upper caudal fin lobe, 75.3, 77.6, 74.1; anterior end cloacal opening, 44.3, 46.0, 42.5. Greatest width at: origin pectorals, 10.2, 10.5, 10.4; origin pelvics, 5.1, 5.3, 5.7; caudal base, 1.2, 1.4, 2.1. Greatest height at: origin pectorals, 6.7, 6.7, 8.3; origin pelvics, 5.9, 6.7, 6.5; caudal base, 2.7, 2.8, 2.9. Eyes: horizontal diameter, 3.9, 3.7, 3.4; vertical diameter, 1.6, 0.7, 1.3. Spiracles: greatest diameter, 0.6, 0.7, 0.5; least distance from eye, 0.8, 0.8, 0.8. Mouth: width, 8.8, 8.4, 9.8; length, 4.3, 5.3, 3.6; length upper labial furrow, 0.8, 1.0, 0.8; lower labial furrow, 1.6, 1.8, 1.3. Nasal openings: least distance between, 2.7, 2.8, 2.6. Gill slits: height of first, 1.6, 1.1, 1.3; height of fifth, 1.2, 0.9, 0.9. First dorsal fin: length base, 5.5, 6.0, 4.9; length free inner margin, 1.6, 1.9, 1.6; height, 4.7, 3.9, 4.4; anterior margin, 7.8, 7.7, 7.8. Second dorsal fin: length base, 5.5, 6.7, 6.2; length free inner margin, 1.8, 1.8, 1.8; height, 2.5, 2.5, 3.4; length anterior margin, 6.7, 7.4, 7.2. Anal fin: length base, 9.8, 10.5, 10.4; length free inner margin, 1.6, 1.4, 1.3; height, 3.1, 2.1, 3.1; length anterior margin, 7.1, 6.3, 5.7. Pectoral fins: width base, 5.1, 4.9, 5.2; greatest width fin, 9.0, 8.4, 7.8; length anterior margin, 11.8, 12.3, 10.4. Pelvic fins: overall length exclusive of claspers, 9.8, 10.5, 8.8; length inner margin claspers, 8.6, — , 4.4; extent claspers beyond posterior pelvic fin tip, 3.1, — , 1.0. 78 Least distance between fin bases: pelvics and anal, 9.4, 11.9, 9.6. first and second dorsals, 10.6, 11.2, 11.4; pectoral and pelvic, 15.7, 17.6, 14.5; In 11 H. hispidus the number of monospondylous vertebrae ranged from 35 to 38 and the total number of vertebrae from 123 to 131. The teeth of upper and lower jaws similar, those in the central part of the upper jaw with a high central cusp and a low lateral cusp on each side; some teeth near the corners of the mouth with four or more cusps, the cusp lengths nearly equal; teeth numerous and crowded with number of vertical rows difficult to count. In one adult male I counted 34+36/42+41 vertical rows. Dermal denticles spikelike in juveniles and tridentate in large specimens, the blades rather wide in adults. Inside of mouth with numerous papillae; gill rakers present only as low humps on gill bars, intermediate in relative size between those of H. lutarius and H. canescens. Liver of moderate size, not notably oil-filled, valvular intestine with 8.5 to 9.0 turns. A 216-mm female had many small eggs 2 to 4 mm in diameter in its functional right ovary, but was considered im- mature because its oviducts were very slender and appeared underdeveloped for an adult. A 285-mm female had only one egg, 15 by 21 mm in diameter, in its ovary. The oviducts were empty but well developed. Relative to the size of the mother the ovarian egg was remarkably large. Discussion. Both Halaelurus hispidus and H. lutarius are variable but are generally similar in many characters. As presently known, however, I believe their separation as distinct species is justified. The known geographical range of H. hispidus extends only from the Gulf of Mannar at the southern tip of India partly across the Bay of Bengal to the Andaman Sea. It is thus separated by very large distances from the geographical areas of its relatives, H. lutarius on the east coast of Africa and H. dawsoni near New Zealand. Presumably additional continen- tal and island slope collecting would extend the range of H. hispidus. The name hispidus presumably comes from the hairlike papillae in the mouth of the Andaman Sea specimens. It would be useful to know whether Gulf of Mannar specimens also have papillose mouths. The separation of hispidus and lutarius depends strongly on the presence or absence of papillae, especially because morphometric differences and differences in vertebral number in scyliorhinids may often reflect only differing conditions at different latitudes. Halaelurus lineatus Bass, D' Aubrey, and Kistnasamy Figures 44, 45, 46 Figure 44. — Halaelurus lineatus, 330-mm immature female taken off Durban, South Africa, in 68-70 m. Halaelurus lineatus Bass, D'Aubrey, and Kistnasamy 1975:12, fig. 8A (Durban, Natal, South Africa, from shore). Material examined. USNM 214224, imm. S, 340 mm, cruise 8, HOE stn. 1948, from shrimp trawl haul in 68-70 m off Durban, South Africa, lat. 29°27'S, long. 31°31'E, 25 September 1964; USNM, 2 imm. S, 313, 357 mm, 3 imm. 2, 313, 333, 351 mm, from same haul as preceding; USNM, ad. $, 475 mm, off Durban; SAM 10556, ad. ?, 457 mm, off Natal. Diagnosis. Halaelurus lineatus is similar to H. natalensis in many respects and both differ from other Halaelurus species and from all other scyliorhinids also in having a pointed and upturned snout tip. This character had not been noted in accounts earlier than the 1972 report of Springer and D'Aubrey even as a feature of H. natalensis probably because small sharks preserved in glass containers often have snouts deformed by being pushed into a too small space. The turned up snout tip of H. lineatus and H. natalensis, is easily seen when the sharks are alive. It is a little more prominent in H. natalensis than in H. lineatus. 79 Figure 45. — Left, two Halaelurus natalensis, Algoa Bay, South Afri- ca. Right, 475-mm female H. lineatus taken off Durban. Figure 46. — Halaelurus lineatus, 457-mm female, Natal, South Africa, SAM 10556. Halaelurus lineatus is easily distinguished from H. natalensis by its color pattern (see Figs. 44 and 45). Characteristically H. lineatus has many small dark spots, as small or smaller than the spiracle, between transverse dark bars that cross the dorsal surface. Halaelurus natalensis has no such spots. In H. lineatus some specimens have a single transverse row of spots between some of the bars as does H. quagga. Halaelurus quagga does not have an upturned snout tip. Some H. lineatus have small spots on the two dorsal fins and near the caudal fin tip, but one specimen, SAM 10556, has no fin spots. Small spots were not seen on any of the specimens of H. natalensis that were examined. Few differences in body and fin proportions seen among specimens of H. lineatus and H. natalensis were sufficiently great to appear significant. The fins of H. lineatus were slightly smaller. The distances between dorsal fin bases in all specimens of H. lineatus seen were more than 75?o TL but were less than 15% TL in specimens of H. natalensis. 80 Skin surfaces of H. lineatus feel appreciably smoother than those of H. natalensis apparently because H. natalensis denticles have their points directed upward as well as posteriorly. Description. A medium sized but slender cat shark, males 337 and 357 mm not mature, a male 475 mm and a female 457 mm probably mature; whether young hatch from egg capsules or carried in oviducts until birth is not known. Abdominal trunk short as in other Halaelurus with firm bodies, distance from tip of snout to anterior end of cloacal opening about 40% TL or less, both abdominal and caudal sections of trunk more or less cylindrical; head moderately short and flattened ; snout pointed but not long, its length to front of mouth considerably less than width of mouth, tip of snout a rounded point slightly turned upward, somewhat knoblike, dermal denticles of snout tip whitish; lateral margins of snout with moderately acute edges. Mouth rather wide, its arch low; labial furrows continuous around the mouth corners, upper furrow about one-fourth as long as lower furrow, lower furrow about half as long as eye; inner posterior ends of nasal apertures close together, separated by a distance about equal to half of eye length; nasal flaps broadly triangular, their sides meeting at about 90° to form flap tips; flaps entirely covering posterior inner part of nasal aperture, their posterior margins close to anterior upper lip, separated from it by a distance less than diameter of spiracle. Eyes moderate, a little more than 5% TL in half-grown or adult individuals; no supraocular shelf of cranium but angle of view of eyes more lateral than dorsal; subocular gutter fully scaled, well developed, about as long as length of eye; spiracle moderately large, opening elongate, its length about half eye length, its position very near posterior eye corner, either near, in, or continuous with, subocular gutter, its distance from eye less than half its greatest diameter. Gill slits short, progressively shorter from first to fifth, the last two over the pectoral base. Fins rather small; pectorals short and broad, their greatest widths only a little less than their greatest lengths, appressed pectoral tips to origin of pelvics only a little more than snouth length, distance from pectoral base (axil) to origin pelvics 12 to 15% TL in specimens examined, probably greatest in mature females; pelvic fins rather long with moderate lateral projection; claspers not notably large or long, clasper tips of adult males not reaching origin of anal fin; dorsal fins of nearly equal size, the first usually a little greater in area, the distance between dorsal fin bases 16 to 18% TL in specimens examined, first dorsal origin a little posterior to midpoint of pelvic fin base, second dorsal origin a little anterior to posterior end of anal fin base; anal fin relatively smaller in area than other fins, lower than dorsal fins, the length of its base only a little greater than length of either dorsal fin base. Teeth small, crowded, tricuspid near middle of jaws, five to seven cusps at angles, in about 60 vertical rows in upper jaw, 55 in lower, the longest in the holotype about 0.5 mm, symphyseals not well defined, upper teeth very slightly larger than lower but not greatly different in shape. Dermal denticles imbricate and small posterior blades of dorsolateral denticles with three points, the points posterior- ly at angle not greater than 40° above body surface, all surfaces when rubbed from head toward tail smooth and firm. In- side of mouth covered generally with very small denticles; papillae in mouth confined to fringe of jaw area; no gill rakers, except a lump less than 1 mm high on each gill bar. Ventral surfaces whitish without spots or markings on body or fins; dorsolateral surfaces brown with a pattern of darker transverse bars and between bars small darker brown spots, the small spots generally as small or smaller than spiracle but tending to coalesce to form irregular vermiculations. Inside of mouth whitish. Following are measurements expressed as percentages of total length. The first figure in each series represents the 340- mm subadult male holotype, the second a 330-mm immature female, and the third the 457-mm adult female. Tip of snout to: front of mouth, 5.3, 5.3, 5.3; eye, 5.6, 5.8, 5.3; spiracle, 9.1, 8.9, 7.9; first gill slit, 13.6, 13.6, 12.5; fifth gill slit, 17.7, 17.9, 16.0; origin pectoral, 16.2, 17.3, 15.5; origin first dorsal, 41.6, 41.2, 42.0; origin pelvics, 35.4, 35.1, 34.6; origin second dorsal, 64.9, 63.3, 64.5; origin anal fin, 54.6, 56.4, 56.2; origin upper caudal fin lobe, 79.1, 80.3, 81.0; anterior end cloacal opening, 39.2, 38.8, 38.9. Greatest width of: trunk at pectoral origin, 11.5, 9.7, 10.5; trunk at pelvic origin, 6.5, 6.7, 7.7; caudal peduncle, 2.4, 2.7, 2.6. Greatest height of: trunk at pectoral origin, 8.5, 9.1, 7.7; trunk at origin pelvic fins, 7.9, 8.2, 8.6; caudal peduncle, 2.9, 3.0, 3.3. Eyes: horizontal diameter, 3.5, 3.0, 2.8; vertical diameter, 1.2, 0.9, 0.7. Spiracles: greatest diameter, 1.5, 1.5, 1.3; least distance from eye, 0.9, 0.9, 0.4. Mouth: width, 7.7, 7.0, 5.9; length, 3.0, 2.1, 2.2; length upper labial furrow, 0.9, 0.6, 0.7; lower labial furrow, 1.8, 1.5, 1.3. Nasal apertures: least distance apart, 1.9, 1.7, 2.2. Gill slits: height of first, 1.8, 2.0, 1.8; height of fifth, 0.9, 0.8, 0.9. First dorsal fin: length base, 5.3, 6.5, 5.9; length posterior inner margin, 2.7, 2.1, 1.8; height, 5.0, 4.5, 5.0; length anterior margin, 8.0, 7.6, 7.2. Second dorsal fin: length base, 5.0, 6.5, 5.7; length posterior inner margin, 2.1, 2.1, 1.8; height, 4.1, 3.9, 3.9; length anterior margin, 6.8, 7.6, 7.2. 81 Anal fin: length base, 7.4, 7.0, 7.0; length posterior inner margin, 3.0, 2.6, 2.2; height, 3.5, 2.9, 2.8; length anterior margin, 6.5, 6.4, 6.3. Pectoral fin: width base, 4.7, 4.8, 4.6; length anterior margin, 11.21, 10.6, 10.9; greatest width, 8.9, 9.1, 9.0. Pelvic fin: overall length, 12.9, 12.1, 12.0; inner margin clasper, 11.2, — , — ; reach clasper past posterior tip pelvic fin, 1.8, — , — . Least distance between fin bases: first and second dorsals, 16.8, 17.3, 17.1; pectoral and pelvic, 15.3, 12.1, 13.8; pelvic and anal, 13.2, — , 13.8; anal and origin lower caudal lobe, 13.2, — , 14.7; second dorsal and origin upper caudal lobe, 9.1, 11.2, 9.8. Vertebrae numbers in 457-mm adult female were, monospondylous 34, percaudal 92, caudal 47, total vertebra 139. Discussion. Halaelurus lineatus probably shares more characters in common with H. natalensis than with other species of the genus and seems to be in some ways intermediate between H. quagga and H. natalensis. The turned up snout tip is absent on H. quagga, present but weak on H. lineatus, and strong on H. natalensis. The pattern of transverse bars with transverse rows of dots or small spots is present on H. quagga; it is present but confused by additional markings on H. lineatus; and it is transformed to a pattern of dorsal saddles on H. natalensis. Bass et al. (1975:13, 14) described H. lineatus partly from the 500-mm holotype that was caught on a hook and line from shore at Durban. The capture of a scyliorhinid by hook and line from shore underlines the littoral to upper con- tinental slope distribution of many of the scyliorhinids of the southeastern coast of Africa in contrast to the restricted slope distribution of scyliorhinids on the eastern continental slope of North America. One of the H. lineatus paratypes was trawled from 290 m off Durban. The known range of H. lineatus extends from the Natal coast to the vicinity of Beira, Mozambique. Bass et al. (1975:13, 14) summarized breeding data for H. lineatus in the following statement. "Pregnant females of this oviparous species are caught in the surf in Natal during late winter (July to September). The egg-cases measure about 40 by 20mm and have long but very thin weak tendrils. They are covered with a coating of short sticky hairs which help them to adhere to the substrate. The embryos are fairly advanced, measuring 30 to 40 cm in length, when the egg- cases are laid. Females carry up to eight egg-cases in each uterus simultaneously. A series of egg-cases laid in tanks at the Durban Aquarium hatched out between 23 and 36 days after deposition (at a water temperature averaging between 19 and 20°C)." Halaelurus lutarius Springer and D' Aubrey Figure 47 Figure 47. — Halaelurus lutarius, 313-mm adult male, Delagoa Bay, Mozambique. Halaelurus lutarius Springer and D'Aubrey 1972:6, figs. 1A, 2A (off Delgoa Bay, Mozambique); Smith 1975:13. Scylliorhinus hispidus (not of Alcock 1891) Brauer 1906:7 (coast of Somalia). Material examined. USNM 205135, ad. S, 326 mm, the holotype, trawled from 450-455 m off Delgoa Bay, East Africa, lat. 25°32'S, long. 33°24'E, 2 ad. S, 310, 317 mm and 14 ad. ?, 310 to 367 mm, same haul as holotype. Diagnosis. Halaelurus lutarius differs from others of the genus except H. canescens, H. dawsoni, and H. hispidus by its lack of prominent dorsal spots or crossbands of lighter or darker color. Halaelurus lutarius may be distinguished from the larger and more robust H. canescens (adults 500 to 650 mm) by its relatively longer anal fin base, 1.5 times the length of the second dorsal base in lutarius but less than 1.3 times the length in canescens. Halaelurus lutarius lacks papillae on the roof of the mouth such as characterize H. hispidus. The nasal flap of lutarius is roughly triangular, quite unlike the lobelike nasal flap in H. dawsoni. Halaelurus lutarius and H. hispidus resemble each other in general appearance. Halaelurus lutarius is larger, adults 310 to 367 mm for lutarius and about 230 to 285 mm for hispidus. Halaelurus lutarius lacks the small papillae that are 82 distributed in moderate density over the roof of the mouth and tongue of hispidus. In adult specimens examined the eye length in lutarius is less relative to total length than the eye length in hispidus. Description. Following are measurements expressed as percentages of the total length. The first number in each series refers to the 323-mm adult male holotype, the second to a 367-mm adult female, and the third to a 310-mm adult female. Tip of snout to: front of mouth, 5.1, 6.3, 6.1; eye, 6.2, 6.5, 6.8; spiracle, 10.8, 10.6, 11.9; first gill slit, 16.1, 16.4, 16.1; fifth gill slit, 20.7, 20.7, 20.1; origin pectoral fin, 20.4, 19.6, 19.7; origin first dorsal fin, 48.0, 50.1, 49.7; origin pelvic fins, 40.9, 44.1, 45.2; origin second dorsal fin, 65.6, 68.4, 69.0; origin anal fin, 60.4, 64.6, 61.6; origin upper caudal fin lobe, 74.6, 77.9, 78.1; anterior end cloacal opening, 45.2, 46.3, 46.1. Greatest width of: trunk at pectoral origin, 9.3, 10.9, 10.3; trunk at pelvic origin, 5.0, 6.5, 5.5; caudal base, 1.5, 1.9, 1.9. Greatest height of: trunk at pectoral origin, 6.2, 8.2, 8.7; trunk at pelvic origin, 7.1, 8.2, 9.4; caudal base, 3.1, 3.3, 3.2. Eyes: horizontal diameter, 2.8, 3.3, 3.2; vertical diameter, 0.9, 1.1, 1.0. Spiracles: greatest diameter, 0.3, 0.5, 0.5; least distance from eye, 0.9, 0.7, 0.8. Mouth: width, 6.2, 8.4, 7.1; length, 4.0, 3.8, 3.9; length upper labial furrow, 1.2, 1.1, 1.5; length lower labial furrow, 1.5, 2.0, 1.6. Nasal apertures: least distance between, 2.5, 3.0, 3.2. Gillslits: height of first, 1.1, 1.4, 1.2; height of fifth, 0.9, 1.1, 0.6. First dorsal fin: length of base, 5.3, 4.8, 5.8; length free inner margin, 2.5, 3.0, 2.3; height, 3.1, 4.1, 3.5; anterior margin, 7.1, 7.6, 7.4. Second dorsal fin: length base, 5.3, 5.0, 5.3; length free inner margin, 1.9, 2.5, 2.3; height, 2.2, 2.5, 2.4; length anterior margin, 6.2, 6.4, 6.4. Anal fin: length base, 11.1, 10.2, 10.3; length free inner margin, 1.5, 1.4, 1.5; length anterior margin, 7.1, 5.7, 6.5. Pectoral fins: width base, 5.0, 5.5, 4.5; greatest width fin, 7.1, 8.2, 7.7; length anterior margin, 10.8, 12.5, 11.0. Pelvic fins: overall length without claspers, 9.6, 11.0, 10.0; length inner margin clasper, 9.6, — , — ; extent clasper past posterior tip pelvic fin, 3.4, — , — . Distance between fin bases: first and second dorsals, 12.7, 12.0, 13.2; pectorals and pelvics, 17.0, 16.6, 17.1; pelvics and anal, 13.3, 12.5, 11.0. Of the 17 H. lutarius examined, the number of monospondylous vertebrae varied from 38 to 42 and the total number of vertebrae from 129 to 139. The teeth of H. lutarius are tricuspid near the jaw symphyses; the central cusp of each tooth is much the longest. The teeth in the 12 to 15 rows near the mouth corners have lower cusps than those of the midjaw area and usually have about five cusps to each tooth. The teeth of the upper and lower jaws are similar, only slightly smaller and lower in the lower jaw. The larger teeth are coarsely marked with vertical striations at their bases. The teeth are crowded into about 76/86 vertical rows. The dermal denticles are tridentate over the dorsolateral surfaces, their blades slightly elevated posteriorly. Mouth papillae generally absent, no distinct gill rakers. The liver is comparatively small and is confined to the anterior third of the body cavity. The valvular intestine has about 10 turns. The specimens examined (all adults) were gray-brown above and lighter below without well-defined or prominent spots or bands. Some of the specimens have ill-defined saddle blotch areas, strongest in the second dorsal fin region. Discussion. Halaelurus lutarius is known to occur off the east coast of Africa from the southern coast of Mozambique to the coast of Somalia in 338 to 766 m. The species is ovoviviparous. The young are retained in the oviducts until they are a little more than 100 mm long. Egg capsules in H. lutarius are thin-walled and very fragile bags. Such a capsule or bag would not furnish appreciable protec- tion outside an oviduct to an incompletely formed embryo or to an embryo made somewhat inactive by attachment to a large yolk sac. Bass et al. (1975:12) described an egg capsule of H. lutarius as a thin, fragile bag about 50 by 15 mm. Halaelurus lutarius adults are about 310 to 390 mm long. Halaelurus natalensis (Regan) Figures 48, 49, 50 Scy Ilium natalense Regan 1904:128 (coast of Natal). Scyliorhinus natalensis: Regan 1908a:461 (Natal and Cape Colony). 83 Figure 48. — Halaelurus natalensis, 334-mm immature female, Algoa Bay, South Africa. Figure 49. — Halaelurus natalensis, 425-mm adult male, Saldanha Bay, South Africa. Figure 50.— Halaelurus natalensis, head region of holotype or para- type, 425-mm specimen, BMNH. 84 Scylliorhinus natalensis: Barnard 1925:43 (Algoa Bay to Natal). Halaelurus natalensis: Garman 1913:84 (coast of Natal); Smith 1949:50, pi. 2, no. 40 (Knysna to Natal). Material examined. BMNH 1904.6.28.29, ad. 3, 425 mm, holotype or paratype, see discussion, one of two specimens received from J. F. Quekett, coast of Natal only locality given; BMNH 1905.6.8.34, 2 ad. S, 416, 430 mm, 16 mi northeast Bird Island in 40 fathoms, Algoa Bay; USNM 119224, 3 imm. S, 233, 317, 322 mm, Algoa Bay, South Africa; USNM, imm. ?, 334 mm, Algoa Bay; SAM 22992, 1 ad. $, 425 mm, Saldanha Bay, South Africa. Diagnosis. The most important character to set off Halaelurus natalensis from all other sharks, except the newly described H. lineatus, is its upturned snout tip. Regan (1904) called the snout "elliptical with obtusely pointed tip." I suppose he meant that the cross section of the snout was elliptical. The tip of the snout, in all the specimens noted above and several more that I examined but did not record, had a very small knoblike tip, denticle covered, usually colorless, and having a somewhat worn appearance. The knob is present on specimens of both natalensis and lineatus but is more prominent on natalensis. Regan's (1904) original description of natalensis noted that it was allied to H. quagga. There are general similarities but only natalensis and lineatus have obtusely pointed snouts. Halaelurus quagga has a broadly rounded snout without a trace of a knoblike tip. The difference in color pattern between H. natalensis and H. lineatus can be seen best by comparison of illustrations (H. natalensis Figs. 48, 49, 50, with H. lineatus Figs. 44, 46). In natalensis, the darkest bars partly surround areas of lighter brown forming discrete saddle blotches of two shades of brown or brownish-black. The dorsolateral surfaces ofH. natalensis thus have three shades of brown in a complex pattern. Within this pattern additional fainter markings may be present to varying degrees. In Figure 50 showing the head region of the holotype or paratype the pattern described in Regan's (1904) species description is clearly evident. "The second band is represented by two oval patches at the level of the gill openings, nearly meeting in the middle line." Figure 49 shows another pattern arrangement of the second saddle blotch in which the two "oval patches" noted by Regan have merged. The specimens of H. natalensis examined have no single body proportion that clearly separates them from specimens of H. lineatus. It is possible that differences could be found by comparison of a larger series of both species. Both species are presumably variable but in a general way the material available shows that H. natalensis has a more robust form, has slightly larger fins, has a slightly shorter but more pointed snout with a more prominent knoblike tip, has lesser dis- tances between fin bases, and feels somewhat rougher to touch than H. lineatus. Description. For an account of proportional measurements I have chosen a 425-mm adult male from Saldanha Bay and a 334-mm immature female from Algoa Bay. Measurements are given as percentages of total length and in the following the first figure in each pair represents the adult male, the second figure the immature female. Tip of snout to: front of mouth, 4.5, 4.5; eye, 5.9, 5.1; spiracle, 9.2, 9.0; first gill slit, 13.6, 13.2; fifth gill slit, 17.9, 17.4; origin pectoral fin, 16.2, 15.6; origin first dorsal fin, 42.8, 41.0; origin pelvic fins, 36.0, 35.6; origin second dorsal fin, 63.8, 62.6; origin anal fin, 56.5, 54.5; origin upper caudal fin lobe, 80.0, 78.8; anterior end cloacal opening, 41.9, 40.4. Greatest width of: trunk at pectoral origin, 11.3, 12.0; trunk at pelvic origin, 6.8, 8.1; caudal base, 2.1, 2.4. Greatest height of: trunk at pectoral origin, 7.3, 6.0; trunk at pelvic origin, 8.2, 7.2; caudal base, 3.1, 3.0. Eyes: horizontal diameter, 3.1, 3.6; vertical diameter, 0.5, 0.9. Spiracles: greatest diameter, 0.9, 1.2; least distance from eye, 0.3, 0.6. Mouth: width, 7.3, 7.5; length, 3.5, 3.0; length upper labial furrow, 1.1, 1.5; length lower labial furrow, 1.8, 1.8. Nasal apertures: least distance between, 2.1, 1.8. Gill slits: height of first, 2.1, 1.9; height of fifth, 0.9, 1.0. First dorsal fin: length base, 6.1, 6.3; length free inner margin, 2.4, 2.1; height, 4.5, 5.7; length anterior margin, 8.7, 8.7. Second dorsal fin: length base, 7.3, 6.4; length free inner margin, 2.1, 2.1; height, 4.0, 7.2; length anterior margin, 8.4, 7.5. Anal fin: length base, 9.4, 7.5; length free inner margin, 2.4, 2.4; height, 3.5, 3.9; anterior margin, 7.1, 6.6. Pectoral fins: width base, 5.4, 5.4; greatest width fin, 9.4, 10.2; anterior margin, 11.8, 11.7. Pelvic fins: overall length not including clasper, 12.7, 12.6; inner margin claspers, 12.7, — ; extent clasper past fin tips, 4.2, — . Distance between fin bases: first and second dorsals, 14.1, 14.7; pectorals and pelvics, 15.3, — ; pelvics and anal, 12.5, — ; anal and lower caudal lobe, 11.5, — ; second dorsal and upper caudal lobe, 8.7, — . The 425-mm male from Saldanha Bay had 34 monospondylous vertebrae, 85 precaudal vertebrae, and 45 caudal vertebrae. The total number was 130. An interesting summary of proportional dimensions of 21 males and 20 females of Halaelurus natalensis compared with series of H. lutarius is given by Bass et al. (1975:43). 85 The dentition was similar to that in most other species of Halaelurus, that is, small, crowded teeth, tridentate in the middle of the jaws with some increase in the number of cusps near the mouth corners. In the 322-mm female from Algoa Bay the highest teeth reached from 0.6 to 0.7 mm above the enamel line; the number of vertical rows of teeth was 26 + 1 + 27/20 + 1 + 20 ; and the lateral cusps of many of the larger teeth of the middle part of the jaws were scarcely visible . The dermal denticles of dorsolateral surfaces were tridentate, the denticle points more inclined upward than in H. lineatus. The inside of the mouths of most of the specimens had some papillae anteriorly and some scattered denticles of very small size on the tongue and gill bars. The snout of all the specimens seen had the terminal knob turned upward. Discussion. The locality given by Regan (1904) for two specimens on which he based his description was "coast of Natal." All of the specimens, except the type that I have seen, have been taken between Algoa Bay and Saldanha Bay, south and west of the coast of Natal. All the specimens of H. lineatus that I have seen have been labeled either as from the coast of Natal or more specifically from the vicinity of Durban which is centrally located on the coast of Natal accord- ing to current maps. My material therefore does not clearly indiate that the geographical ranges of natalensis and lineatus overlap. The British Museum (Natural History) has only one of the specimens remaining of the two on which Regan based his description. Regan mentioned only one dimension, 325 mm. The present measurement of the single remaining specimen is 420 mm TL. Bass et al. (1975:15) reported that H, natalensis is oviparous and has egg cases about 40 by 15 mm, the egg cases with fairly thick, robust tendrils in contrast with those of H. lineatus which are thin and weak. They stated that female H. natalensis may carry up to 11, but usually 6 to 9, egg cases in each oviduct at a time. Halaelurus quagga (Alcock) Figures 51, 52 Figure 51 .—Halaelurus quagga, 354-mm adult male, coast of Somalia in 59-61 m. Drawing by Mildred Carrington. Figure 52.— Halaelurus quagga. Redrawn by Mildred Carrington from Alcock's (1899) illustration of the holotype, an adult male, 280 mm, from Malabar Coast. Scy Ilium quagga Alcock 1899:17 (off Malabar coast, India, in 186 m). Scyliorhinus quagga: Regan 1908a:461 (compiled). Halaelurus quagga: Garman 1913:84 (compiled); Springer and D'Aubrey 1972:15, fig. IE (coast of Somalia). Scyliorhinus (Halaelurus) silasi Talwar 1972:779, fig. 1 (off Quilon, southwest coast of India, Malabar coast, in 300 m). Material examined. ZSI, ad. S, 333 mm, lat. 10°00'N, long. 51°15'E, coast of Somalia, presented to Indian Museum for comparison with holotype; USNM, 2 ad. S, 342, 354 mm, same locality as preceding; USNM, juv. S, 79.5 mm, lat. 09°41'N, long. 51°15'E. All specimens in 59-61 m. Diagnosis. The holotype, ZSI 751/1, is an adult male about 280 mm TL. The specimens listed above are the only specimens that I have seen. They were almost identical in appearance with a pattern of transverse bars as shown in Figure 51. One of these was sent to the Indian Museum and was reported to be similar in all essential features to the holotype (A. G. K. Menon, pers. commun.). The illustration, Figure 52, redrawn from Alcock's plate, may be a better representation of some specimens from the coast of India than of specimens from the coast of Somalia, Figure 51. The status of H. quagga and of Scyliorhinus (Halaelurus) silasi Talwar remains uncertain because it has not been determined whether the holotypes of either lack supraorbital crests. The reported absence of labial furrows in silasi brings to mind past difficulties in the determination of distinctions between Halaelurus buergeri and Scyliorhinus torazame without reference to other characteristics such as the presence or absence of a supraorbital crest. None of the specimens ofH. quagga have the knoblike structure at the tip of the snout such as is present on H. lineatus and H. natalensis. The adult males of H. quagga lack the numerous small black spots irregularly scattered between transverse dark bars or saddle blotches as in H. boesemani. When spots are present on H. quagga they tend to be arranged in transverse rows. The reticular marbling that is faintly present between transverse bars of H. lineatus is absent from specimens of H. quagga. Description. Proportional measurements expressed as percentages of the total length follow. The first figure refers to a 354-mm adult male, the second to a 342-mm adult male, and the third to a juvenile male 79.5 mm long that is probably newly hatched or born. The proportions of juveniles differs considerably from proportions of adults in most shark species. Tip of snout to: front of mouth, 4.5, 4.7, 5.9; eye, 5.1, 5.3, 5.3; spiracle, 8.8, 9.4, 10.0; first gill slit, 13.6, 14.0, 16.4; fifth gill slit, 19.2, 18.7, 19.5; origin pectoral fin, 18.1, 17.5, 19.9; origin first dorsal fin, 42.9, 43.6, 38.4; pelvic fins, 37.3, 36.3, 33.5; origin second dorsal fin, 65.0, 64.0, 57.9; origin anal fin, 56.5, 56.7, 48.4; origin upper caudal fin lobe, 78.5, 80.1, 75.0; anterior end cloacal opening, 39.3, 40.9, 37.2. Eyes: horizontal diameter, 3.7, 3.5, 4.4; vertical diameter, 0.8, 1.2, 0.8. Spiracles: greatest diameter, 0.7, 0.6, 1.4; least distance from eye, 0.6, 0.6, 0.5. Mouth: width, 6.2, 6.4, 8.7; length, 3.7, 3.5, 3.5; length upper labial furrow, 0.4, 0.5, — ; length lower labial furrow, 1.1, 1.5, undeveloped ? . Nasal apertures: least distance between, 2.0, 2.0, 2.0. Gillslits: height first, 1.6, 1.2, 1.5; height fifth, 1.4, 1.0,0.8. First dorsal fin: length base, 5.1, 6.4, 6.5; length free inner margin, 1.8, 1.5, 2.9; height, 4.0, 4.1, 4.0; length anterior margin, 7.6, 7.9, 9.4. Second dorsal fin: length base, 6.2, 6.4, 7.5; length free inner margin, 1.8, 1.5, 2.3; length anterior margin, 7.1, 7.3, 10.3. Anal fin: length base, 7.5, 7.3, 10.1; length free inner margin, 2.3, 2.9, 2.5; length anterior margin, 5.1, 5.6, 7.5. Pectoral fins: width base, 4.5, 5.0, 6.3; greatest width fin, 7.6, 7.9, 7.5; length anterior margin, 9.6, 9.4, 10.3. Pelvic fins: overall length exclusive of claspers, 10.4, 11.1, 11.6; length inner margin claspers, 11.0, 11.1, 2.9; reach claspers past pelvic fin tip, 3.7, 3.2, (-3.3). Distance between fin bases: first and second dorsals, 16.9, 14.6, 11.3; pectorals and pelvics, 13.8, 15.2, 8.9; pelvics and anal, 13.3, 10.8, 6.5; anal and lower caudal fin lobe, 12.1, 14.3, 11.3; second dorsal and upper caudal lobe, 9.3, 10.2, 8.2. In the three adult males from the coast of Somalia the number of monospondylous vertebrae was 31 to 32; the total number of vertebrae from 123 to 125. The teeth in adult H. quagga were tridentate and similar in upper and lower jaws, the longest about 0.8 to 0.9 mm above the enamel line. The number of vertical rows of teeth were 26+3+26/27 + 27 and 28+3+28/27+1+27 in two of the adult males. Dermal denticles of the dorsolateral surfaces were tridentate from 0.3 to 0.6 mm long, with a double central ridge. The inside of the mouth had no prominent papillae, and denticles of very small size were scattered over the anterior mouth roof and on the tongue and gill bars. 87 Discussion. Halaelurus quagga and H. boesemani were taken in trawl hauls made only a few miles apart on the continental shelf of Somalia at depths of 60 to 70 m. No indications that the two forms occupied different habitats were observed. HAPLOBLEPHARUS GARMAN Haploblepharus Garman 1913:101 (type-species, Scyllium edwardsii Cuvier 1817; by monotypy). Diagnosis. Haploblepharus lacks a supraorbital crest or narrow shelf of cartilage above the eye and thus differs from Atelomycterus. Of the genera without a supraorbital crest, Haploblepharus differs in having in combination, large con- fluent nasal flaps that reach past the edge of the upper lip, prominent labial furrows that extend around the mouth cor- ners as a continuous groove, and depressed areas of the upper lip that make a connection between the nostrils and the mouth. It also lacks a caudal crest of modified and enlarged denticles. The three species recognized here are robust sharks with wide and moderately depressed heads, short snouts, and wide mouths. They have moderately small eyes with poorly developed subocular gutters below the eyes, the gutters not reaching the spiracles which are very close to the posterior eye corners. Color and markings vary and the significance of color differences for systematics in this genus has not been establish- ed. Discussion. Specimens of Haploblepharus are not comonly available in systematic collections outside of South Africa and, until the recent publication by Bass et al. (1975), the species were poorly known. I have examined only four specimens divided among three species. As a consequence, much of my treatment of the genus is based on data from Bass et al. (1975) who examined 44 specimens. Since their morphometric data are presented only as a summary of an analysis of variation in proportional dimensions (see also Bass 1973), I have included here proportional dimensions of specific specimens of two of the species. The three species of Haploblepharus are not easily distinguished from one another but seem more reliably separated by color characters than by morphometric differences. All three are known from shore waters of continental shelf waters of the coasts of southern Africa and have somewhat different but probably overlapping geographical ranges. Two leathery egg capsules 60 by 30 mm were laid by a female identified as Haploblepharus edwardsii by Bonde (1945). One of these hatched in 104 days. Bass et al. (1975) regard it as likely that the species was H. pictus. All three species are probably oviparous. Haplobleparus has moderate sexual dimorphism in the teeth, the upper jaw teeth of adult males having somewhat longer middle cusps than the teeth of females or immature males. This difference appears greater in H. fuscus than in the other two species. Key to Adults of Species of Haploblepharus la. Origin of first dorsal fin in advance of posterior end of base of pelvic fin; dorsolateral surfaces gen- erally dark with a few large saddle blotches extending onto paired fins, dorsolateral surfaces also with scattered spots of lighter color, the spots generally larger than the spiracle but smaller than the eye H. pictus lb. Origin of first dorsal fin over or posterior to posterior end of base of pelvic fin; dorsolateral sur- faces either dark or light, sometimes tan, reddish, or brown, with or without scattered spots of lighter color, the light colored spots, if present, generally smaller than the eye 2 2a. Dorsolateral surfaces of adults with distinct saddle blotches, sometimes bordered at front and back with noticeably darker margins; many small pale spots not much larger than spiracles . H. edwardsii 2b. Dorsolateral surfaces of adults either plain colored or with indistinct bands or narrow saddle blotches; small pale spots either absent or present H. fuscus Haploblepharus edwardsii (Muller and Henle) Figure 53 Scyllium edwardsii Muller and Henle 1841:4, first unnumbered plate (Cape of Good Hope). Haploblepharus edwardsii: Garman 1913:102; Smith 1950:879, fig. 1; Bass et al. 1975:17, fig. 11. 88 4 i Figure 53. — Haploblepharus edwardsi, adult male, esti- mated length 450 mm (part of tail missing), taken off Simontown, South Africa, in 18 m, SAM 22021. Material examined. SAM 22021, S, estimated length 450 mm, part of tail missing, Simontown, South Africa, 18 m; USNM 201809, 5, 470 mm, Algoa Bay, South Africa. Diagnosis. In Haploblepharus edwardsii the origin of the first dorsal fin is over or posterior to the rear end of the base of the pelvic fin, not in advance of it as in H. pictus. The separation of H. edwardsii and H. fuscus is based on color pattern as described in the key. The color pattern appears to be variable, especially in H. fuscus, and separation on color alone may be tenuous. Although the accompanying illustration of if. edwardsii (Fig. 53) shows a strong pattern of saddle blotches that in one form or another is usually present on H. edwardsii, the illustration oiH. fuscus here (Fig. 54) shows just one of several patterns for that species. Another color phase of H. fuscus was illustrated by Bass et al. (1975, fig. 12). Discussion. Bass et al. (1975:16) observed a separate cartilage in Haploblepharus between the two halves of the lower jaw. That structure was illustrated for H. edwardsii (1975, fig. 10). I made no observations of Haploblepharus jaw struc- tures but have seen somewhat similar intermediate or symphyseal cartilages in some Heptranchias perlo (Family Hep- tranchidae). Unlike Haploblepharus which has a remarkably wide mouth with a low arch, Heptranchias has a very narrow mouth with a very high arch. Haploblepharus fuscus Smith Figure 54 Haploblepharus fuscus Smith 1950:883, fig. 2 (East London and Knysna coast of South Africa) ; Bass et al. 1975:19, fig. 12. Material examined. SAM 24545, ad. 92-mm adult male, Bredas- dorp, South Africa, SAM 24545. Description. The 692-mm male H. fuscus was brown dorsally, paler ventrally, with faint darker dorsal crossbands (5 or 6) and numerous white spots about spiracle size on dorsolateral surfaces. Following are proportional measurements of the 692-mm adult male expressed as percentages of the total length. Tip of snout to: front of mouth, 3.9; eye, 4.6; spiracle, 7.8; first gill slit, 14.5; fifth gill slit, 18.4; origin pectoral fin, 17.5; origin first dorsal fin, 43.7; origin pelvic fins, 37.9; origin second dorsal fin, 65.1; origin anal fin, 56.4; origin upper caudal fin lobe, 81.8; anterior end cloacal opening, 42.5. Greatest width of trunk: at pectoral origin, 12.9; at pelvic origin, 8.1; at origin caudal fin, 3.0. Greatest height of trunk: at pectoral origin, 9.5; at pelvic origin, 8.8; at caudal origin, 3.6. Eyes: length of eye opening, 2.5. Spiracles: greatest diameter, 0.7; least distance from eye, 0.3. Mouth: width, 7.4; length, 2.0; length upper labial furrow, 1.7; length lower labial furrow, 2.2. Gill slits: height of first, 2.7; height of fifth, 1.2. First dorsal fin: length base, 7.4; length free inner margin, 2.6; height, 5.6; length anterior margin, 9.3. Second dorsal fin: length base, 8.7; length free inner margin, 2.6; height, 5.5; length anterior margin, 9.7. Anal fin: length base, 11.9; length free inner margin, 2.5; height, 5.1; length anterior margin, 9.7. Pectoral fins: width base, 5.9; greatest width of fin, 10.4; length anterior margin, 12.7. Pelvic fins: overall length exclusive of claspers, 11.6; inner margin claspers, 12.0; reach of claspers past tip of pelvic fins, 3.8. Distance between fin bases: first and second dorsals, 12.7; pectorals and pelvics, 14.5; pelvics and anal, 12.1; anal and lower caudal origin, 10.7; second dorsal and upper caudal origin, 8.1. Vertebral numbers for the 692-mm adult male are: total 134, monospondylous 36, precaudal 90, caudal 40. Discussion. Comparative examinations of larger series of specimens are needed to test the validity of the species. The illustration of the clasper of H. edwardsii in Bonde's (1945) account of external development and my sketches of the clasper or the 692-mm H. fuscus do not differ appreciably. 90 Haploblepharus pictus (Miiller and Henle) Figure 55 Figure 55. — Haploblepharus pictus, 410-mm female, Liideritz Bay, Southwest Afri- ca, SAM 24345. Haploblepharus pictus Miiller and Henle 1841:4 (Cape of Good Hope); Smith 1975:13; Bass et al. 1975:21, fig. 13. Material examined. SAM 24345, ?, 410 mm, Luderitzbucht, Southwest Africa. Diagnosis. The more anterior position of the origin of the first dorsal fin, its origin in advance of the posterior end of the pelvic base (over or posterior to it in H. edwardsii and H. fuscus) should distinguish H. pictus at all ages. Differential growth usually affects relative position of fins near the midpoint in the shark's length less than the extremes. The larger spots noted by Miiller and Henle (1841) and present in SAM 24345 may be a practical way to identify H. pictus if it turns out to be a reasonably constant character. Description. The color pattern of the 410-mm female is best shown in Figure 55. Following are measurements of the 410-mm female expressed as percentages of the total length. Tip of snout to: front of mouth, 3.7; eye, 5.1; spiracle, 8.3; first gill slit, 14.9; fifth gill slit, 20.0; origin pectoral fin, 17.1; origin first dorsal, 44.7; origin pelvic fins, 39.0; origin second dorsal fin, 63.7; origin anal fin, 56.9; origin upper caudal fin, 80.0; anterior end cloacal opening, 41.5. Greatest width at: pectoral origin, 14.9; pelvic fin origin, 9.8; caudal base, 3.2. Greatest height at: pectoral origin, 11.2; pelvic fin origin, 10.5; caudal base, 3.7. Eyes: length, 2.4; height opening, 0.7. Spiracles: greatest diameter, 1.1; least distance from eye, 0.7. Mouth: width, 9.8; length, 2.4; length upper labial furrow, 2.0; length lower labial furrow, 2.9. Gill slits: height of first, 3.2; height of fifth, 1.5. First dorsal fin: length base, 7.8; length free inner margin, 3.2; height, 5.9; length anterior margin, 10.7. Second dorsal fin: length base, 8.8; length free inner margin, 2.9; height, 5.9; length anterior margin, 10.7. Anal fin: length base, 10.7; length free inner margin, 2.9; height, 4.6; length anterior margin, 8.5. Pectoral fins: width base, 6.6; greatest fin width, 11.5; length anterior margin, 11.5. Pelvic fins: overall length, 12.4. Distance between fin bases: first and second dorsals, 12.7; pectorals and pelvics, 15.6; pelvics and anal, 8.8; anal and lower caudal fin, 9.3; second dorsal and upper caudal fin, 7.1. 91 In the female examined the teeth were tricuspid in the middle part of the jaws and the lateral cusps were more than half as long as the central cusps. Vertebral numbers for the 410-mm female are: total 129, monospondylous 35, precaudal 82, caudal 47. Discussion. Possibly H. pictus prefers cooler waters than the other two species of Haploblepharus; at least its presence on the Atlantic side of southern Africa suggests this. HOLOHALAELURUS FOWLER Holohalaelurus Fowler 1934 (type-species, Scylliorhinus regard Gilchrist, by original designation). Diagnosis. Holohalaelurus species have no supraorbital crests of the chondrocranium and no labial furrows; they have a partial apron formed beneath the claspers of males by a union of the basal third or half of the pelvic fins, but the skin uniting the pelvics is denticle free as in Juncrus (not denticle covered as in Scyliorhinus and Poroderma); the skin between the gill slits and immediately adjacent to the gill slits is denticle free. Species of Holohalaelurus are relatively small scyliorhinids with broad, flattened heads, and distinctively marked with a color pattern of variegated dark spots or sometimes with a few white spots or more complex ocellalike markings. The eyes are on the dorsal side of the head, not visible from below but their angle of view is more lateral than dorsal. Species of Holohalaelurus have a poorly developed, shallow, subocular gutter either denticle lined or naked extending below the eye for about two-thirds its length. They have denticle-free gill covers and denticles are absent from the skin immediately adjacent to the gill slits. In Holohalaelurus substantial changes in body proportions accompany growth. Also, the adult color pattern differs greatly from that of juveniles (Bass et al. 1975, fig. 15). Even the color patterns of adults vary considerably and show some geographical variation. Adult males are larger than adult females whereas the reverse is the more general condition among sharks. Holohalaelurus is restricted to the east coast waters of Africa and is known from about 200 to 740 m. I follow Bass et al. (1975:23) in the recognition of two of the four nominal species. Key to Species of Holohalaelurus from Bass et al. (1975:23) la. Anal base equal to or less than 3V2 times anal height; denticles on back and top of head uniform in size H.punctatus lb. Anal base more than 3V& times anal height; a series of relatively large denticles on back and top of head H. regani Holohalaelurus punctatus (Gilchrist) Figure 56 y Figure 56. — Holohalaelurus punctatus, ventral view of head, 315-mm adult male showing shape of mouth characteristic of adult males of genus. 92 Scyliorhinus punctatus Gilchrist 1914:129 (off Cape Point, South Africa); Barnard 1925:43. Scyliorhinus polystigma Regan 1921:413 (15 to 22 mi off Umvoti River, Natal); Norman 1939:10 (as synonym of S. punctatus Gilchrist). Halaelurus punctatus: Fowler 1935:361, fig. 1. Scyliorhinus melanostigma Norman 1939: 9 (part), fig. 2A (Zanzibar; syntype BMNH 1939.5.24.5 only). Holohalaelurus punctatus: Smith 1949:55, pi. 2, fig. 42; Bass et al. 1975:23, fig. 14. Material examined. 219 to 238 m). BMNH 1921:3.1.1, ad. S, 315 mm, holotype of Scyliorhinus polystigma Regan 1921 (Natal in Diagnosis. The characters given in the preceding key probably are sufficient to separate both adult and immature H. punctatus from the larger sizes of H. regani. Additionally, in adult H. punctatus the length of the base of the anal fin is usually shorter than the distance between the bases of the dorsal fins but nearly or quite as long as the interdorsal dis- tance in H. regani. Also in adult H. punctatus the area of the first dorsal fin is nearly the same as the area of the second dorsal, but in H. regani the first dorsal usually has the larger area. Discussion. A 205-mm immature male that I saw in fresh condition, when it was hauled up in a trawl off the coast of Tanzania, was one of the most beautiful of all sharks. The dorsal background color was orange-brown becoming cream colored ventrally. The entire dorsolateral surface including top of head and upper side of pectorals and pelvics was covered rather uniformly with dark brown spots, each spot a little larger than the spiracles. Each dorsal fin was marked by a small and short brown bar bordered by bright yellow. A white spot about pupil size was present on each side just above the pectoral axilla. The only marks on the cream colored ventral surface were brown spots on the distal halves of the pectoral and pelvic fins. Holohalaelurus punctatus is somewhat variable in color as shown by the description of Bass et al. (1975:23). According to those authors, H. punctatus is oviparous forming egg capsules about 17 by 10 mm, much smaller than any shark egg capsule that I have seen. Holohalaelurus regani (Gilchrist) Figures 57, 58 Figure o7. — Holohalaelurus regani, 576-mm adult male, SAM 24408. 93 % / -, Figure 58. — Holohalaelurus regani, 576-mm adult male, ventral view to show markings. Similar markings are present on Raja laevis, the barndoor skate. Scyliorhinus regani Gilchrist 1922:45 (Cape Seas, 174-320 m). Holohalaelurus regani: Smith 1949:55, fig. 43; Bass et al. 1975:25, fig. 15. Material examined. SAM 24408, ad. S, 576 mm, South Africa off Dassen I.; BMNH 1935.5.2.55, subad. ', 540 mm, Discovery Stn. A., Cape of Good Hope; USNM 201751, ?, 360 mm, off Mozambique, lat. 22°25'S, long. 35°54'E, trawled from 740 m, 1 October 1964. Diagnosis. Holohalaelurus regani, with males reaching maturity at about 575 mm, is a larger species than H. punctatus in which adult males are about 315 mm or a little more. Probably most specimens of H. regani can be iden- tified by the characters in the preceding key. The color pattern of H. regani adults differs greatly from that of juveniles as shown in illustrations in Bass et al. (1975, fig. 15). As adults both H. regani and H. punctatus vary somewhat in color pattern, particularly in the shape of the spots that cover the dorsolateral surfaces. Both species occur usually in shades of brown. Ventral punctate spots appear on ventral surfaces of adults (Fig. 58). The punctate spots may be absent on some immature specimens. Description. Holohalaelurus regani is a medium size scyliorhinid, males reaching maturity at about 575 mm; head and anterior trunk short, flattened, and wide tapering to tail, head about one-sixth total length; body cavity short, dis- tance from tip of snout to cloaca less than two-fifths total length; depth of head at spiracles less than one-half head width; snout in front of mouth short, its length about equal to length of first dorsal base; gill slits very small, length of longest less than horizontal diameter of orbit which is less than length of snout; pectoral fins very large, broad, the length of the anterior margin about equal to distance from tip of snout to first gill slit; first dorsal fin somewhat smaller than second, its origin slightly in advance of posterior end of pelvic base; second dorsal origin over posterior third of anal base; anal long and low, the length of its base twice as long or nearly twice as long as second dorsal base; caudal fin narrow without a salient point on anterior lower lobe. Eyes of medium size; a shallow subocular gutter extending below the eye for about two-thirds eye length, its deepest part not covered with denticles; gutter not reaching small spiracles which are moderately large in adult male and very close to posterior eye corner. Gill slits very small in immature female, proportionally much larger in adult male, skin between gill slits and im- mediately adjacent area denticle free. 94 Nasal apertures long, oblique, the inner posterior nasal opening covered by a broad triangular flap without salient point; posterior margin of nasal flap not reaching upper lip. Inside of mouth without denticles except for padlike groups of denticles at upper end of each gill bar; papillalike gill rakers irregularly distributed on gill bars, heights not greater than width of bases. Basal half of inner margins of pelvics united by denticle-free skin forming an apron beneath the claspers; tips of claspers of mature male just reaching tips of pelvic fins, clasper tips slender, flexible. Teeth in immature female small, in about 60/66 rows, mostly with five cusps, teeth of central parts of jaws with longer middle cusps, difference in cusp length less notable in lateral parts of jaws; teeth of upper and lower jaws generally similar; lateral teeth small, low, with all five cusps low. Teeth of adult male with about seven rows on each side of symphysis of lower jaw, larger than opposite upper jaw teeth and lacking accessory cusps; rest of teeth of adult male generally similar to teeth of immature female but with somewhat higher central cusps. Dorsal and lateral dermal denticles nearly erect with narrow blades; ventral dermal denticles somewhat less erect with broader blades; denticles generally with a strong high central ridge, usually double, leading to a posterior point, no lateral ridges and lateral points weak if present; dermal denticles of a broad dorsal area extending from the level of the eyes to the level of the second dorsal fin of two sizes, the larger denticles widely spaced in several irregular longitudinal series among the much more numerous smaller denticles; large denticles about twice the height of the small denticles. Color in the specimens I examined was light gray, dark gray, and black but probably this was due to changes after preservation. According to Bass et al. (1975:25, 26) the overall color is brown, except in juveniles in which the ground color is black. Punctate black spots on the ventral surface were regarded by Bass et al. (1975:24) as characteristic of the genus and are shown in accompanying Figure 58. Vertebral numbers of adult male with numbers for immature female following in parentheses are: total number 130 (124), monospondylous 31 (29), precaudal 80 (79), caudal 50 (45). Proportional dimensions as percentages of total length are given below for the 576-mm adult male followed in parentheses for dimensions of 360-mm immature female. Tip of snout to: front of mouth, 3.8, (4.7); eye, 5.0, (6.3); spiracle, 8.7, (10.0); first gill slit, 13.4, (14.7); fifth gill slit, 17.0, (17.2); origin first dorsal, 42.6, (40.3); pelvics, 32.7, (34.2); second dorsal, 60.8, (59.2); anal, 51.2, (50.0); origin up- per caudal fin, 76.4, (76.5); anterior end cloacal opening, 36.0, (35.6). Greatest width of: head, 12.7, (12.2); trunk at pectorals, 13.0, (12.2); trunk at pelvics, 7.6, (7.5); trunk at caudal origin, 1.9, (2.4). Greatest depth of: head at spiracles, 6.1, (5.6); trunk at pectorals, 6.9, (6.1); trunk at pelvics, 7.5, (5.8); trunk at caudal origin, 2.8, (3.1). Eyes: horizontal diameter, 3.3, (3.5). Spiracles: greatest diameter, 0.8, (0.6); least distance from eye, 0.5, (0.7). Mouth: width, 7.5, (8.3); length, 3.5, (4.4), (labial furrows absent). Gillslits: height of first, 2.1, (0.6); height of fifth, 0.7, (0.4). First dorsal fin: length base, 4.9, (4.6); length inner (posterior) margin, 2.8, (2.5); height, 4.2, (3.9). Second dorsal fin: length base, 6.8, (7.0); length inner (posterior) margin, 3.3, (3.3); height, 5.0, (4.4). Anal fin: length base, 13.2, (13.1); length free inner margin, 2.3, (1.7); height, 3.6, (2.8); length anterior margin, 6.8, (6.7). Pectoral fins: width base, 8.9, (7.2); greatest width fin, 10.9, (8.9); anterior margin 14.2, (14.2). Pelvic fins: overall length, 13.9, (11.7); inner margin claspers, 10.1, ( — ). Distance between fin bases: first and second dorsals, 15.5, (15.0); pectorals and pelvics, 8.7, (10.8); pelvics and anal, 9.2, (9.7); anal and lower caudal origin, 9.9, (9.7); second dorsal and upper caudal origin, 9.2, (9.7). Discussion. All of the following data on H. regani is from Bass et al. (1975:26). Egg capsules measured 35 by 15 mm and all of 14 pregnant females carried one egg capsule in each oviduct. This is taken to mean that the species is oviparous and in the condition described by Nakaya (1975:82) as single oviparity. The smallest specimen recorded was 130 mm long. Off the Natal coast males were mature at 500 to 550 mm, but 570 mm in males from the Cape region. Mature females were smaller — 380 to 440 mm — on the coast of Natal. JUNCRUS WHITLEY Juncrus Whitley 1939:229 (type-species, Scy Ilium uincenti Zietz, by monotypy). Diagnosis. Only one species is known. 95 Juncrus lacks a supraorbital crest or narrow shelf extending from the chondrocranium above each orbit. It lacks a caudal crest of modified dermal denticles, and does not have nasal flaps extending posterior to or past the edge of the up- per lip. Juncrus has labial furrows of moderate length extending around the mouth corners but less than half the dis- tance toward the jaw symphyses. Juncrus has a partial apron beneath the claspers in males formed by the union of the denticle covered pelvic fins near their bases and by this character is separable from Halaelurus. In Juncrus the partial apron beneath the claspers (between the claspers and the caudal part of the trunk) is formed by the direct union of the denticle-covered inner margins of the pelvic fins near their bases, whereas in Asymbolus the connection between the pelvic fins is made by a denticle-free membrane that is attached to each pelvic fin near its inner margin on the dorsal side of the fin. The claspers of adult male Juncrus are remarkably long and slender and extend past the tips of the pelvic fins for half the total clasper length. In this respect Juncrus differs from all other scyliorhinids except Atelomycterus and the two species of Galeus, G. nipponensis and G. schultzi. Juncrus vincenti (Zietz) Scyllium vincenti Zietz 1908:287 (type-locality, Kangaroo Island, South Australia). Scyliorhinus vincenti: McCulloch 1911:4, pi. 2. Halaelurus vincenti: Fowler 1941:50; McKay 1966:68. Juncrus vincenti: Whitley 1940:90, fig. 82. Study material. WAM P. 3777, subad. $, about 350 mm, Esperance, Western Australia; BCIF 56-406, ad. $, about 365 mm, Adelaide area, South Australia; USNM, 2 ad. $, 490, 505 mm, 4?, 455, 458, 458, 520 mm, Tamar River mouth, Tasmania. Type not seen, type-series in South Australian Museum (McCulloch 1911:4). Diagnosis. Juncrus vincenti is distinguishable from other scyliorhinids by characters used in the key to genera and in the diagnosis for Juncrus. Juncrus vincenti has a color pattern including many light colored, small spots (generally smaller than the dark adapted pupil) and in this differs from other Australian scyliorhinids. Description. Juncrus vincenti is a slender, elongate scyliorhinid with the head and trunk only moderately higher than wide, and the postpelvic trunk only moderately wider than high; head blunt, snout rounded, its length in front of mouth a little more than 2 times the horizontal diameter of the orbit; eyes moderate, length of orbit as great or greater than least distance between nasal apertures; secondary lower eyelid prominent, extending below eye for its entire length, subocular gutter deep, fully covered by denticles, spiracle moderately small, very near posterior corner of orbit; mouth with a high arch, labial furrows continous around each mouth corner, lower somewhat longer than upper, both shorter than least distance between nasal apertures; nasal flaps entirely covering posterior nasal openings, posterior margins somewhat sinuous or slightly notched, not reaching upper lip; gill slits short, the first usually longest, its length less than least distance between nasal apertures; pectoral fins small, their corners rounded, their origins under fourth gill slits; dorsal fins nearly equal in area, the second usually with a somewhat longer base, the origin of the first over or posterior to the end of the pelvic base; anal fin usually long, length of its base greater than base length of either dorsal; caudal fin about one-fifth total length, caudal axis not elevated, the lower caudal fin without a projecting lobe. Teeth numerous in about 60/56 rows, with three to five cusps, the middle one longest, similar in upper and lower jaws, crown bases with vertical ridges not extended on cusps, no indication of sexual dimorphism in teeth. Dermal denticles with tridentate blades on high, slender pedicles; blades imbricate, all three blade points strong, the central one longest; blades with a central ridge and a lateral ridge on each side but ridges not prominent and obsolete on denticles of some parts of body; denticles mostly transparent, microstructure visible but not prominent; some denticles of specimens from Tasmania with some diffuse pigment and small dark pigment spots (pigment not evident in two specimens, WAM P. 3777 and BCIF 56-406, both of which have been in preservative for a comparatively long time); very small denticles in a patch anteriorly on roof of mouth, denticles also present on gill bars. No gill rakers. Generally mottled with darker color (gray or brown) on dorsal surfaces with numerous light-colored spots randomly distributed, the spots not larger than the dark-adapted pupil and separated from one another by a distance about equal to or more than spot diameter; seven or eight ill-defined saddle blotches of darker color; ventral surfaces whitish with some diffused darker color on some specimens. Total number of vertebrae 130 to 139 in seven specimens (caudal vertebrae in one specimen partly fused and could not be counted); monospondylous vertebrae, one specimen with 34, five with 35, one with 36, one with 37; precaudal vertebrae 80 to 85. Following are proportional measurements expressed as percentages of total length. The first figure in each series refers to a 505-mm adult male from Tasmania, the second figure to a 520-mm adult female from Tasmania, and the third figure to a 350-mm adult male from Esperance, Western Australia. 96 Tip of snout to: front of mouth, 4.4, 4.6, 3.4; eye, 4.8, 5.2, 5.0; spiracle, 8.1, 8.5, 8.3; first gill slit, 11.9, 13.1, 12.9; fifth gill slit, 17.0, 17.7, 15.7; origin pectoral, 16.2, 17.3, 14.9; origin first dorsal fin, 48.5, 50.0, 42.9; origin pelvics, 37.0, 37.5, 34.3; second dorsal fin, 67.5, 69.6, 65.7; origin anal fin, 61.4, 59.2, 54.3; origin upper caudal fin, 79.2, 78.8, 80.0; anterior end cloacal opening, 41.0, 40.8, 40.0. Greatest width of: trunk at pectoral origin, 7.5, 10.3, 8.3; trunk at pelvic origin, 6.9, 8.4, 2.6; at caudal origin, 2.8, 3.3, 2.6. Height of trunk at: pectoral origin, 6.7, 8.4, — ; pelvic origin, 7.9, 8.6, 7.1; caudal origin, 3.8, 4.4, 4.3. Eyes: length, 2.4, 2.5, 2.9; height, 0.6, 1.1, 0.7. Spiracles: greatest diameter, 0.3, 1.0, 0.9; least distance from eye, 0.5, 0.6, 0.6. Mouth: width, 5.9, 6.0, 6.0; length, 3.8, 3.1, 2.7; length upper labial furrow, 1.2, 1.0, 1.0; lower labial furrow, 1.6, 1.3, 1.6. Nasal apertures: least distance between, 1.8, 2.0, 2.3. Gillslits: height of first, 1.0, 1.7, 1.7; height of fifth, 0.8, 0.8, 0.6. First dorsal fin: length base, 6.3, 6.2, 8.0; length free inner margin, 2.6, 2.9, 2.3; height, 4.4, 4.0, 3.7; length anterior margin, 8.7, 7.9, 10.0. Second dorsal fin: length base, 7.1, 7.7, 8.6; length free inner margin, 2.8, 2.5, 2.6; height, 3.8, 4.2, 3.7; length anterior margin, 9.1, 9.6, 10.0. Anal fin: length base, 10.5, 11.5, 9.1; length free inner margin, 2.0, 1.9, 1.7; height, 3.8, 3.8, 3.4; length anterior margin, 7.7, 8.1, 7.1. Pectoral fins: width base, 4.3, 4.6, 4.9; greatest fin width, 7.5, 7.9, 7.4; length anterior margin, 10.5, 11.2, 10.0. Pelvic fins: overall length exclusive of claspers, 12.1, 13.1, 14.3; inner margin claspers, 15.8, — , 16.6; extent claspers past pelvic fin tips, 9.5, — , 8.0. Distance between fin bases: first and second dorsals, 13.9, 14.0, 13.4; pectorals and pelvics, 16.2, 15.9, 15.7; pelvics and anal, 15.6, 13.5, 10.9; anal and lower caudal, 6.1, 5.2, 6.6; second dorsal and upper caudal, 5.3, 4.2, 7.4. Discussion. McCulloch (1911) compared a specimen taken by the research vessel Endeavour from 37 m off the mouth of the Murray River, South Australia, with one of Zietz's (1908) specimens that he called a co-type and noted several differences in body proportions and fin positions between the two. Similar variations were found in the eight specimens that I examined. Differences among specimens examined found in accounts by others are random and I can only conclude that J. vincenti is variable in color, relative positions and size of fins, and in size at maturity. In the series at hand the distance from the tip of snout to the origin of the first dorsal fin varies from 42.9 to 50.0% TL. The first dorsal origin is either over the end of the pelvic base or considerably posterior to it. The length of the anal fin base varies from 9.0 to 13.2% TL. In one specimen the anal fin base length is approximately 2 times the first dorsal base length but in another it is only 1 l /s times the first dorsal base. The distance between dorsal fin bases varies from 11.8 to 16.4% TL. The USNM specimens from Tasmania resemble Whitley's (1940, fig. 82) illustration of the species in color pattern more closely than the other two specimens in the present study series. More specimens are needed to determine whether the treatment here of all specimens as one variable species is correct. Juncrus vincenti is oviparous. The following descriptive account of the egg capsules is taken from Hale (1935). The egg capsule is quadrangular in shape, somewhat flattened, 52 mm long by 21 mm wide by 10.5 mm in greatest thickness. At the posterior end of the capsule each corner is produced, tapering to form a single filament which is about 250 mm long. When the capsule was removed from the oviduct and placed in saline solution (seawater?) the posterior filaments (ten- drils) "corkscrewed." Along the lateral margins of the egg capsule, silky filaments emerge and extend to 30 or 40 mm. At the anterior corners much longer filaments composed of fine threads are 350 mm long. Hale speculated that when the egg capsule was extruded the posterior tendrils twisted rapidly around some weed or other support and that the anterior threads tend to cling to any object touched. PARAPRISTURUS FOWLER, 1934 Parapristurus Fowler 1934:237 (type-species, Catulus spongiceps Gilbert, by original designation). Diagnosis. Parapristurus has a series of transverse folds or wrinkles in a denticle-free patch of skin extending across the throat between the ends of the anterior gill slits. Such a patch of wrinkles is not present in other scyliorhinid sharks but is present in some species of the squaloid sharks of the genera Centrophorus and Deania. In Parapristurus the wrinkle patch is associated with basihyals that are comparatively large and heavy for a scyliorhinid shark and may func- tion to permit sudden expansion of the branchial cavity to draw in prey. Parapristurus is similar to Apristurus in having a somewhat flattened and spatulate snout, strong labial furrows ex- tending around the mouth corners, lobe-shaped fins, generally uniform dark color on all surfaces without notable markings, and is like most Apristurus species in having a long anal fin with a much larger area than the two dorsal fins combined. Like Apristurus, Parapristurus has a domelike tectum (roof) of the chondrocranium and no supraorbital 97 crests but differs somewhat from Apristurus in having postorbital processes extending across the posterior corners of the orbits. In Parapristurus the eyes are more lateral than dorsal with thick upper lids that project slightly above the eye and with a definite pouch under the anterior half of the eye. In Apristurus the eyes are more dorsal than lateral with thin upper lids and at most a shallow and inconspicuous gutter below the eye. Discussion. Only one species of Parapristurus has been described; P. spongiceps is known only from the 500-mm adult female holotype from the vicinity of Hawaii and a 105-mm juvenile reported by Weber (1913) from the Banda Sea. Parapristurus spongiceps (Gilbert) Catulus spongiceps Gilbert 1905:579. Scyliorhinus spongiceps: Regan 1908a:459. Pristiurus spongiceps: Garman 1913:94. Pentanchus spongiceps: Fowler 1941:53, fig. 2. Apristurus spongiceps: Bigelow and Schroeder 1948:220. Material examined. USNM 51590, ad. ?, 500 mm, near Bird Island, WNW of Hawaii in 572-1,462 m, holotype. Diagnosis. The generic characters, especially the presence of a patch of transverse small folds or wrinkles across the throat, serve to distinguish Parapristurus spongiceps from all other scyliorhinid sharks. Description. The following proportional measurements of the holotype in percentages of total length, except as indicated, do not differ greatly from measurements included in the original description (Gilbert 1905) but are more com- prehensive. The holotype is in moderately good condition but undoubtedly has been somewhat distorted during storage. Tip of snout to: front of mouth, 8.6; eye, 8.9; first gill slit, 20.4; fifth gill slit, 25.6; origin pectoral, 24.8; origin first dorsal, 49.0; pelvics, 43.6; second dorsal, 63.4; anal, 55.2; upper caudal lobe origin, 74.0; anus, 49.0. Greatest width of: head, 12.7; trunk at pectorals, 10.6; trunk at pelvics, 9.0; trunk at caudal origin, 2.0. Greatest height of: head at spiracles, 10.4; trunk at pectorals, 15.0; trunk at pelvics, 15.0; trunk at caudal origin, 4.4. Eyes: horizontal diameter, 3.2; vertical diameter, 1.2; distance between upper eyelid rims, 9.6. Spiracles: greatest diameter, 0.5; least distance from eye, 1.2; distance between, 10.4. Mouth: width, 9.0; length, 4.8; length upper labial furrow, 2.8; length lower labial furrow, 3.6. Nasal apertures: level of anterior ends to tip of snout, 3.8; level of posterior ends to tip of snout, 7.0; minimum distance between, 4.2; greatest length (diagonal), 3.2. Gill slits: height of first, 1.4; height of fifth, 1.0. First dorsal fin: length base, 6.8; inner posterior margin, 2.4; height, 4.0. Second dorsal fin: length base, 6.6; inner posterior margin, 2.0; height, 4.0. Anal fin: length base, 13.0; inner posterior margin, 1.8; height, 5.0. Pelvic fins: length (origin to rear tip), 11.2. Caudal fin: upper margin, 26.0; tip to notch, 5.6; anterior margin lower lobe, 10.0. Distance between fin bases: first and second dorsal, 8.6; pectoral and pelvic, 10.6; pelvic and anal, 2.2; anal and lower caudal, 0.0; second dorsal and upper caudal, 5.0. Teeth: arrangement alternate; separation at symphyses, 2 mm; number of tooth rows, counted as number perpendicular to line of occlusion of jaws 25/22, counted as oblique rows 18/18. Vertebrae: total number about 106; monospondylous 34, precaudal 61, caudal about 45. Body moderately compressed, deep, stout, visceral cavity short, head about one-fourth total length, tail about one- fourth total length; fins lobelike, dorsal fins nearly equal in area, the origin of the first over midpoint of pelvic base, the origin of the second slightly in advance of midpoint of anal base; anal fin origin slightly in advance of pelvic tips; poste- rior end anal base separated from lower caudal lobe only by a complete notch. Snout broadly rounded (collapsed in holotype); two double rows of prominent pores on lower side of snout, about 12 in longitudinal series; nasal openings large, oblique, nasal flaps developed only as points, not covering either anterior or posterior openings; mouth moderately arched, large, with conspicuous labial furrows around corners; gular area denticle free, with 10-25 transverse small folds; denticles not detected inside mouth, no gill rakers, a few scattered small papillae present. Teeth similar in upper and lower jaws, mostly with five cusps, the central cusp much the longest. Dermal denticles generally similar over all surfaces, spikelike, nearly erect, slightly curved posteriorly. 98 Color of preserved holotype uniform brown. Parapristurus spongiceps was illustrated by Fowler (1941, fig. 2). Discussion. The holotype has a fully formed egg capsule in the right oviduct, length not including tentacles 52 mm, width 23 mm. The surface has fine longitudinal striations, but otherwise is unmarked. The egg capsule is firm as would be expected in an oviparous species. PARMATURUS GARMAN Parmaturus Garman 1906:203 (type-species, Parmaturus pilosus Carman, designated by Jordan 1917-1920:518). Diagnosis. Parmaturus lacks a supraorbital crest (a narrow shelf of the chondrocranium that extends over the orbits). The nasal flaps do not reach as far back as the upper lip. The nostrils are not connected to the mouth by a groove or a depression of the upper lip near each nostril. The gular area in most species is covered with denticles and the area has no conspicuous transverse wrinkles. Species of Parmaturus lack the color pattern of darker saddle blotches or spots that characterize many scyliorhinid species including most Galeus species. Parmaturus has two dorsal fins, labial furrows of moderate length that extend around the mouth corners, and a caudal crest, the crest made up of modified and sometimes enlarged dermal denticles on the proximal half or more of the upper edge of the caudal fin, the crest separated from the denticles of the lateral surfaces of the caudal fin by a narrow strip of naked skin on each side. Parmaturus and Galeus are distinguished from all other scyliorhinids by the presence of the caudal crest. Parmaturus differs from Galeus in the following respects: Parmaturus lacks saddle blotches whereas Galeus usually has them; the pectoral fins of Parmaturus are not notably large or broad, their greatest width (measured parallel to the distal margin of the pectoral) less than the width of the mouth, whereas in Galeus the pectorals are broad, usually as wide or wider than the width of the mouth; Parmaturus species are comparatively soft-bodied scyliorhinids, their dermal denticles not closely imbricate and their denticle points not notably stiff so that Parmaturus feels more velvety than prickly to touch whereas Galeus species are moderately hard-bodied, their dermal denticles imbricate in adults, and their denticle points rather stiff so that Galeus feels more prickly than velvety to touch. Discussion. Species of Parmaturus may have different and less complex clasper structures than Galeus species. Parmaturus xaniurus has striking external differences in clasper structure from G. arae, G. sauteri, and G. eastmani (L. J. V. Compagno, pers. commun.). I have not used clasper structure in the classification of scyliorhinids to the extent that may be warranted. For Parmaturus at least, there is the obvious excuse that I have seen an adult male of only one of the six species. As defined here, Parmaturus includes six somewhat diverse species, four of which are known only from immature specimens. One of the species here placed in Parmaturus was originally described by Chan (1966) as Dichichthys melanobranchus based only on the immature holotype from the South China Sea. Chan regarded his species as tran- sitional between the Galeus-Parmaturus "complex" and Apristurus. I now add three new species no better known than melanobranchus and each is transitional in one way or another within the Galeus-Parmaturus- Apristurus group of genera. Furthermore, I have found that juveniles of both Cephalurus cephalus and Halaelurus canescens have incipient caudal crests that disappear in later growth stages. One Parmaturus pilosus that I examined had a short section of the ventral edge of the caudal peduncle with denticles modified in about the same way as caudal crest denticles are modified. Parmaturus melanobranchus was also described as having a crest of enlarged denticles on the ventral edge of the caudal peduncle and on the front portion of the lower caudal lobe (Chan 1966:223). Although the species of Parmaturus are loosely tied together by the presence of a caudal crest on the upper edge of the caudal fin, I suspect that an equally important similarity, but hypothetical for most species, is their ability to live in midwater without excessive energy expenditure. Parmaturus pilosus and P. xaniurus are the only scyliorhinid species known to have an abundant supply of liver oil that is chiefly squalene (see discussion under P. xaniurus). My solution to the treatment of the diverse group is to make subgeneric distinctions which reflect my present opinion on relationships. Key to Species of Parmaturus la. Nasal flaps either triangular and large enough to nearly cover posterior nasal apertures or lobe- like and long enough to reach across nasal apertures 2 lb. Nasal flaps very small, pointed, and neither large enough to cover the posterior nasal apertures nor long enough to reach across nasal apertures 3 99 2a. Nasal flaps triangular; origin of second dorsal fin nearly over origin of anal fin; posterior end of base of second dorsal fin well in advance of posterior end of base of anal fin . . . P. (Parmaturus) xaniurus 2b. Nasal flaps lobelike; origin of second dorsal fin over or slightly in advance of midpoint of base of anal fin ; posterior end of base of second dorsal fin about over posterior end of base of anal fin . . P. (Parmaturus) pilosus 3a. Eye relatively large, its length less than 2 times in length of snout in front of mouth 4 3b. Eye relatively small, its length more than 2 times in length of snout in front of mouth 5 4a. Origin of first dorsal fin over or slightly in advance of origin of pelvic fins P. (Dichichthys) campechiensis 4b. Origin of first dorsal fin over posterior end of base of pelvic fins P. (Dichichthys) melanobranchus 5a. Snout tapering, somewhat cone-shaped, its tip a blunt point P. (Campagnoia) stenseni 5b. Snout moderately flattened, its tip broadly rounded P. (Campagnoia) manis Subgenus Dichichthys Chan First dorsal fin smaller than second dorsal fin; eyes large, their lengths less than 2 times in length of snout in front of mouth; eyes more lateral than dorsal; adults not known. Parmaturus campechiensis n. sp. Figure 59 Figure 59. — Parmaturus campechiensis, 157-mm female holotype from Gulf of Campeche in 1,097 m. Drawing by Keiko Hiratsuka Moore. Holotype. USNM 206184, juv. ?, 157 mm, northwestern Bay of Campeche, Gulf of Mexico, lat. 21°33'N, long. 96°48'W, 1,097 m, trawled by RV Oregon II, stn. 10956, 3 June 1970. Only specimen known. Diagnosis. Parmaturus campechiensis differs from other species of the genus in having the first dorsal fin origin over or slightly in advance of the pelvic origin, whereas it is slightly behind the pelvic origin in P. pilosus and P. xaniurus and farther back in other species. Parmaturus campechiensis has many characters in common with P. melanobranchus of the South China Sea and P. pilosus of Japanese waters. In P. campechiensis, the first dorsal is appreciably smaller than the second as in P. melanobranchus; the two fins are nearly equal in P. pilosus. In both P. campechiensis and P. melanobranchus, the gill slits are small and especially close together but are well separated in P. pilosus. The forward position of the first dorsal fin in P. campechiensis sets it off distinctly from P. melanobranchus which has the first dorsal origin over the posterior fourth of the pelvic base. 100 In direct comparison with specimens of P. xaniurus 135, 170, and 183 mm long, the 157-mm holotype has a smaller, narrower head with a smaller mouth, shorter gill slits spaced much closer together, a small eye, the first dorsal fin smaller than the second, the posterior tip of the second dorsal fin extending well past the tip of the anal fin (tips about opposite in P. xaniurus), and much smaller nasal flaps. The more anterior position of the first dorsal sets P. campechiensis apart from the two other new species P. stenseni and P. manis described here. In addition, the eyes in the two latter species are more dorsal in position with their angle of view more dorsal than lateral, and both have longer snouts, somewhat pointed in P. stenseni, rounded in P. manis. Also, in those species the anal fins are comparatively long. Description. A species of Parmaturus with a caudal crest of modified denticles; color grayish, somewhat darker on belly, around gill slits, and on outer parts of fins; first dorsal fin somewhat smaller than second, its origin very slightly in advance of origin of pelvics; second dorsal fin origin a little in advance of midpoint of anal base, posterior end of second dorsal base posterior to end of anal base, tip of second dorsal overlapping origin of caudal crest; areas of second dorsal and anal nearly equal; anal base longer than second dorsal base and longer than distance between dorsal fin bases; pec- toral fins short, not broad; caudal fin not wide, its lower extension narrow. Caudal crest present with crest denticles larger and different in shape from denticles of lateral surface of tail but marginal denticles of crest not asymmetrical as in P. melanobranchus. Snout rounded, its length in front of mouth less than width of mouth; nasal apertures oblique, the nasal flap little developed, not covering posterior opening; eyes small, their lengths nearly equal to the least distance between nasal apertures; spiracles small, distance from eye about four times spiracle diameter; gill slits short, the fifth less than half the length of the first, close together. Teeth mostly tridentate, only a few with four cusps, not differing greatly in upper and lower jaws, in somewhat more than 60 vertical rows. Dermal denticles over most of body needlelike, somewhat curved toward the tips; denticles of anterior part of head with narrow tridentate blades; caudal crest denticles in only about three to five irregular rows well separated from den- ticles of the lateral surface of the tail by a band of naked skin, the caudal crest denticles with a high central ridge reaching a slender point, blades moderately widened basally with a weak posteriorly directed point on each side; caudal crest originates in advance of tip of second dorsal and continues along proximal half of caudal fin edge; a similar arrange- ment of modified denticles begins ventrally under the anal fin tip but remains differentiated and separated from lateral surface denticles by a band of naked skin only to origin of the lower caudal fin lobe, a distance about equal to length of base of second dorsal fin. Following are dimensions in percentages of total length for the 157-mm juvenile holotype. The proportions in larger specimens may be expected to be somewhat different. Furthermore the figures should be considered as approximations because they were made from a soft-bodied species. Tip of snout to: front of mouth, 5.7; eye, 7.0; first gill slit, 15.6; fifth gill slit, 18.5; origin first dorsal, 38.5; pelvics, 39.2; origin second dorsal, 54.8; anal, 51.0; origin caudal crest, 65.9; anus, 42.7. Eye: horizontal diameter, 3.2; vertical diameter, 2.5. Spiracle: diameter, 0.5; least distance from eye, 2.0. Mouth: width, 7.3; length, 3.8; length upper labial furrow, 1.4; lower labial furrow, 1.3. Gill slits: height first, 2.9; height fifth, 1.0. First dorsal fin: length base, 7.0; length posterior inner margin, 3.5; height, 3.2. Second dorsal fin: length base, 8.6; length posterior inner margin, 3.5; height, 4.5. Anal fin: length base, 10.8; length posterior inner margin, 1.3; height, 3.8. Pectoral fin: width base, 6.4; anterior margin, 8.9; greatest width, 5.7. Distance between fin bases: first and second dorsal, 8.9; pectoral and pelvic, 17.2; pelvic and anal, 5.1; anal lower caudal, 3.2; second dorsal and lower caudal, 0.0. Vertebral numbers are: monospondylous 39, precaudal 75, caudal 36, total 111. Discussion. The juvenile holotype of Parmaturus campechiensis and the immature specimens forming the type- series of P. manis are the first Parmaturus reported from the Atlantic. If these species are restricted to midwater at depths usually greater than 200 m they would be taken only accidentally by trawls or dredges designed to operate at the sea bottom. Such trawls do occasionally catch fishes during travel to and from fishing depths but are not efficient then and probably are easily avoided by all but the smallest sharks. Parmaturus melanobranchus (Chan) Dichichthys melanobranchus Chan 1966:226, figs. 2, 3 (South China Sea). 101 ■ Material examined. BMNH 1965.8.11.6, 235 mm ?, RV Cape St. Mary, cruise 4/64, stn. 119, SSE of Hong Kong, lat. 20°05'N, long. 115°03'E, trawled from 548 m, 22 August 1964, the holotype and only specimen known. Diagnosis. Parmaturus melanobranchus is readily separable from all other species here allocated to the genus by the posterior position of its first dorsal fin, its origin over the posterior one-fourth of the base of the pelvics. Parmaturus melanobranchus is obviously close to P. campechiensis. In addition to the difference in the position of the first dorsal fin, however, P. melanobranchus differs in a number of proportional measurements from P. campechiensis and also in having a somewhat more highly differentiated caudal crest. The total number of vertebrae, 140 for P. melanobranchus as compared with 111 for P. campechiensis, shows a far greater difference than can be accounted for by species variation. Discussion. Chan's description, illustrations, and measurements of the species are comprehensive and accurately cover the essential features of this shark and will not be repeated here. Parmaturus melanobranchus has modified denticles (as in the caudal crest) on the ventral side of the caudal peduncle and along the leading edge of the lower caudal fin lobe. Subgenus Campagnoia New Subgenus Type-species, Parmaturus (Campagnoia) manis. Eyes more dorsal than lateral; eyes of juveniles and young small, the horizontal diameter of orbit more than two in snout length; adults not known. Parmaturus manis n. sp. Figures 60, 61, 62, 63 Figure (>().— Parmaturus manis, 32X-mm female holotype from lat. .'!9°.)2'N, long. 70°50'W, (off Nantucket, Mass.) in 731-841 m. Drawing by Mary Wagner. Figure 61 . — Parmaturus manis, holotype, lateral view of upper lobe of tail showing caudal crest of denticles at top. 102 r"^ 1 m m Figure 62. — Parmaturus manis, Upper left, denticles from top of head in advance of eyes. Upper middle, flank denticles from below first dorsal fin. Upper right, denticle from lateral surface of tail. Lower, caudal crest denticles. 1 mm Figure 63. — Parmaturus manis, holotype, teeth from upper and lower jaws taken from 4th vertical rows, counting from sym- physes. Apristurus profundorum (not of Goode and Bean 1896). Springer 1966:612, figs. 22A, 23 (part, except holotype of profundorum). Holotype. MCZ 38299, imm. ?, 328 mm, southwest of Nantucket, Mass., lat. 39°52'N, long. 70°50'W, 731-841 m, W. C. Schroeder, stn. 24, 23 June 1952. Paratypes. MCZ 37512, imm. S, from same haul as holotype; MCZ 37535, imm. ?, 225 mm, lat. 39°52'N, long. 69°38'W, 658-768 m; MCZ 37416, 2 imm. S, 240, 245 mm, lat. 39°52'N, long. 70°43'W, 759-804 m. Material examined. Known only from the type-series. Etymology. From Latin manis, ghost or shade of the departed, in reference to its grayish-white color. Diagnosis. Parmaturus manis most closely resembles species of Apristurus except for the presence of a well-defined caudal crest of modified denticles. Parmaturus manis has a long, flattened snout with a broadly rounded tip as in Apristurus and its eyes are nearly dorsal so that the shark's angle of view is more dorsal than lateral, also as in Apristurus. Of the species here placed in Parmaturus, P. manis most closely resembles P. stenseni but differs from it in having a broadly rounded snout instead of a snout tapering to a blunt point. Parmaturus manis not only has a longer snout than P. campechiensis, P. melanobranchus, P. pilosus, and P. xaniurus, about 10% TL in P. manis, but also lacks an appreciable interspace between the posterior end of the base of its long anal fin and the origin of the lower caudal lobe. Description. Parmaturus manis is a soft-bodied scyliorhinid with a flattened head and a long flattened snout, the snout about 10% TL; snout spatulate, its tip broadly rounded; nasal apertures large, oblique; nasal flaps reduced to small pointed projections; no depressed area of upper lip forming a channel between nasal apertures and mouth; labial furrows prominent, upper and lower continuous around mouth corners; eyes small, dorsal, making shark's angle of view more dorsal than lateral; no supraorbital crest; gill slits short; spiracle small; fins lobelike, their corners rounded except posterior tips of pelvics and anal; first dorsal origin over middle of pelvic base; second dorsal larger than first, its origin over middle of anal base; anal fin long, length of its base more than 2 times length of first dorsal base. Teeth small, mostly with three or five cusps, the middle cusp much the longest; arrangement alternate, a small space at the symphysis of each jaw; teeth in about 29+30/26+26 vertical rows. Dermal denticles of lateral surfaces of tail sparsely and randomly distributed, needlelike, nearly erect, the point slightly curved posteriorly; forward from the tail the denticles become tridentate except on the top of the head where they are leaf-shaped with only one apical point and with a single very high central ridge; proximal half of upper edge of 103 Ml ^m caudal fin with a caudal crest made up of imbricate, tridentate denticles, their blades about as wide as long; crest den- ticles separated from denticles of lateral surfaces of tail by a band of naked skin (Fig. 61). Color generally light gray somewhat irregularly suffused with darker gray but not definitely lighter above or below, tips of some fins lighter. Proportional dimensions in percentages of total length are given here for the holotype with the range in dimensions for four paratypes following in parentheses. Tip of snout to: front of mouth, 10.7, (9.3-10.7); first gill slit, 22.7, (19.6-22.7); fifth gill slit, 27.7, (23.6-27.7); origin pectoral, 26.4, (23.1-26.4); origin first dorsal, 46.7, (40.4-46.9); pelvic, 40.0, (36.4-42.9); origin second dorsal, 58.7, (52.2- 58.7); origin anal, 50.7, (46.7-53.8); origin upper caudal crest, 69.3, (64.5-70.8); anus, 44.0, (40.9-45.7). Eyes: horizontal diameter, 2.4, (2.4-2.7). Spiracles: greatest diameter, 0.3, (0.3-0.6); least distance from eye, 1.1, (1.1-1.3). Mouth: width, 7.5, (5.7-7.5); length, 3.7, (3.7-4.9); upper labial furrow, 3.2, (2.2-3.3); lower labial furrow, 2.7, (1.3- 2.7). Nasal apertures: level of anterior ends to snout tip, 6.9, (5.8-6.9); level of posterior ends to snout tip, 8.5, (8.3-9.4); minimum distance between, 4.3, (3.7-4.7); greatest length, diagonal, 3.5, (3.1-3.7). Gillslits: height of first, 1.3, (0.9-1.6); height of fifth, 1.3, (0.7-1.5). First dorsal fin: length base, 6.1, (5.8-6.5); length inner posterior margin, 4.0, (3.7-4.4); height, 3.2, (2.2-3.2). Second dorsal fin: length base, 8.0, (6.0-8.0); length inner posterior margin, 4.5, (3.8-4.5); height, 2.7, (2.7-3.1). Anal fin: length base, 15.5, (13.3-15.5); height, 5.1, (2.9-5.1). Pectoral fin: width base, — , (5.3-6.7); length outer margin, 10.7, (8.4-10.7). Pelvic fins: length, origin to rear tip, 10.9, (8.9-10.9). Distance between fin bases: first and second dorsal, 7.5, (5.3-8.9). Vertebral numbers: monospondylous 35, (34-35); precaudal 64, (62-65); caudal about 56, (about 48-57); total vertebrae about 120, (about 110-121). Gill rakers not present; mouth denticles not present; inside of mouth with small papillae. Color light gray with irregular darker areas; distal parts of some fins white. Discussion. In an earlier paper (Springer 1966:612, figs. 22A, 23) I incorrectly based my account of Apristurus profundorum chiefly on the specimens here made the type-series of Parmaturus manis. The series was originally part of a collection of 16 Apristurus -like specimens caught by the RV Caryn and the fishing vessel Cap'n Bill II in the summers of 1952 and 1953. Eleven of the specimens were in fact Apristurus and rather nice-looking specimens. The other five were smaller and looked very much like sick, damaged, or long-dead Apristurus. These five are the specimens here described as Parmaturus manis. The entire collection of 16 specimens was reported as Apristurus profundorum at various times (Bigelow et al. 1953; Bigelow and Schroeder 1954; Schroeder 1955). Parmaturus stenseni n. sp. Figures 64, 65, 66 Figure 64.— Parmaturus stenseni, immature male holotype, 185 mm, from 915 to 975 m. Gulf of Panama. Drawing by Mildred Carrington. Holotype. ZMK-Galathea 739-1, imm. 8, 185 mm, Gulf of Panama, lat. 07°22'N, long. 79°32'W, 915-975 m, herring otter trawl, 15 May 1952. Paratypes. ZMK-Galathea 739-2 to 41, 40 specimens, 94 to 230 mm, same haul as holotype except one specimen, which one not known, from Galathea station number 745, Panama Bay. Condition of material examined. The only specimens known are those of the type-series. They are delicate sharks all damaged during capture in one way or another but all are complete enough for identification as belonging in the species 104 Figure 65. — Parmaturus stenseni, upper jaw teeth, longest tooth about 0.2 mm. Scanning electron photomicrograph by U.S. Na- tional Museum staff. Figure 66. — Parmaturus stenseni, dorsal and ventral views of head of 203 mm female. Drawing by Mildred Carrington. described here. The internal structures are generally in better condition than external ones. The series was fixed in For- malin and preserved in alcohol. Two specimens were cleared and stained, and are preserved in glycerin. Etymology. Parmaturus stenseni is named in honor of Niels Stensen (Steno), 1638-84, in recognition of his important studies of elasmobranch anatomy, which, because of their scientific accuracy, were influential in the begin- nings of elasmobranch systematics in the pre-Linnaean period and later, even to the time of the appearance of Muller and Henle's "Plagiostomia" in 1841. Diagnosis. Parmaturus stenseni is a soft-bodied scyliorhinid shark with a caudal crest of modified denticles. Parmaturus stenseni and P. manis differ from other species of the genus in having longer, flattened, or somewhat conical snouts with eyes set in a dorsal position (Fig. 66) such that the shark's angle of view is more dorsal than lateral. Both species have large nasal apertures with nasal flaps reduced to small points. Parmaturus stenseni differs from P. manis in having the flattened snout tapering to a blunt point, its tip not broadly rounded as in P. manis. Description. The description is based primarily upon two males and three females, the 185-mm holotype and four others, 203 to 230 mm, all immature. 105 ^^m Soft-bodied sharks with thin skins, head flattened, one-fifth to one-fourth total length; body moderately compressed; tail from origin of caudal crest about three-tenths TL, its axis not elevated, upper and lower caudal lobes narrow, poorly developed, caudal notch present but inconspicuous; body cavity short; eyes small, orbit length less than one-half snout length; nasal apertures moderately large, nearer mouth than tip of snout, with small, short, and pointed nasal flaps; mouth with a moderately high arch, about twice as wide as high; labial furrows moderately prominent and continuous around mouth corners; fins not large, pectorals and anal largest; first dorsal origin about over middle of pelvic base, separated from second dorsal by a distance less than length of anal base; second dorsal slightly larger than first, its origin over posterior part of anal base, posterior end of its base posterior to rear tip of anal; anal base long, its origin under end of base of first dorsal fin. Teeth not proportionally very small or numerous, in about 28 to 32 rows in each jaw, delicate and easily broken (see Fig. 65), with three or more cusps, the middle cusp longest, cusps somewhat lanceolate in type-series (all immature). Dermal denticles over entire body surface except caudal crest needlelike on four-pronged bases; denticles very widely spaced; dermal denticles of caudal crest with three slender points, denticle blades nearly as wide as long, imbricate or nearly imbricate; a naked band of skin separating caudal crest from denticles of lateral surfaces of tail prominent in some specimens but very narrow in others; denticles inside of mouth not observed except on gill bars, gill bar denticles more or less erect with paddle-shaped denticle blades. Livers not notably large but posterior tips reach level of cloaca in some specimens; valvular intestine with 10 or more turns. Vertebral calcification moderately strong in trunk sector; monospondylous vertebrae in 10 specimens; 33 in three, 34 in six, 35 in one; estimated total number from radiographs 100 to 110. Claspers of largest male, 208 mm, bent strongly outward, claspers moderately stiff but calcification not evident. Dimensions as percentages of total length are given below for the 185-mm holotype and in parentheses the range in the holotype and four other larger specimens. Tip of snout to: level of anterior ends nasal apertures, 4.3, (2.9-5.8); level of posterior ends nasal apertures, 5.9, (3.8-8.0); front of mouth, 6.5, (5.3-8.7); eye, 8.6, (8.6-12.2); spiracle, 11.4, (11.4-16.4); first gill slit, 18.4, (18.4-20.4); fifth gill slit, 23.2, (20.0-27.6); origin first dorsal, 47.0, (42.3-48.9); origin pelvics, 41.1, (39.4-42.2); origin second dorsal, 59.5, (59.1-62.2); anal, 54.1, (49.8-54.3); origin caudal crest, 70.0, (70.0-75.6); anus, 44.3, (41.8-47.6). Eyes: horizontal diameter, 2.7, (2.4-2.7). Spiracles: greatest diameter, 1.1, (0.7-1.1); least distance from eye, 0.6, (0.6-1.0). Mouth: width, 7.0, (7.0-8.9); length upper labial furrow, 1.6, (1.6-3.6); lower labial furrow, 2.2, (2.2-4.4). Gill slits: height first, 3.0, (1.9-3.6); height fifth, 2.7, (1.5-2.7). First dorsal fin: length of base, 4.9, (3.9-5.1); height, — , (2.4-3.5). Second dorsal fin: length of base, 5.4, (5.3-6.7); height, — , (3.0-4.4). Anal fin: length base, 10.8, (10.3-14.8); height, — , (2.5-3.6). Pectoral fins: width base, 5.9, (3.4-5.9); length anterior margin, — , (5.8-13.3). Distance between fin bases: first and second dorsal, 7.8, (7.8-10.1); pectoral and pelvic, 13.0, (8.9-13.0); pelvic and anal, 7.0, (3.0-7.0). Color, belly, lower snout, branchial region, and distal half of fins black or blackish; other external surfaces brown; inside of mouth, black; peritoneum black. Discussion. Parmaturus stenseni seems to be far more delicate than any other shark. Even the teeth and some of the long slender denticles are somewhat flexible but were very easily broken or displaced. In the largest male (208 mm), claspers were strongly bent laterally outward but I could not determine whether there was any calcification. In the largest female (230 mm), the ovary could be seen but no distinct ovarian eggs were recogniz- ed. The peritoneum was black ventrally and laterally but not on the dorsal side of the body cavity. The inside of the mouth was black. Subgenus Parmaturus Garman First and second dorsal fins subequal or the first slightly larger than the second; eyes large, the horizontal diameter of orbits of adults and juveniles less than IV2 in snout; eyes more lateral than dorsal; adults with high proportions of squalene in liver oil. Parmaturus pilosus Garman Parmaturus pilosus Garman 1906:204 (Sagami Bay, Japan, 786 m); Garman 1913:89, pi. 8, figs. 7-10. Pristiurus hertwigi Engelhardt 1912:643. 106 Material examined. MCZ 1107, imm. 8, ca. 544 mm, holotype, Sagami Bay, Japan, lat. 34°59'N, long. 139°31'E, 786 m, Owston; SU 13899, imm. S, 563 mm, Sagami Bay, Owston; FMNH 74133, ?, 640 mm, Sagami Sea, Owston; FMNH 74132, ?, 590 mm, Boshu, Japan, Owston; SU 35478, ?, 460 mm, probably Sagami Bay, Owston; FMNH 74168, 2, 300 mm, Sagami Sea, Owston; FMNH 74134, imm. 3, 293 mm, Okinose, Japan, Owston. Diagnosis. Parmaturus pilosus should be identifiable readily by use of the foregoing key. As already noted, the distinctions between Galeus and Parmaturus adults are not very sharp and Galeus eastmani is similar to P. pilosus in many respects. Parmaturus pilosus, however, has smaller and narrower pectoral fins and a somewhat longer body cavity than G. eastmani so that the distance from the appressed pectoral tip to the origin of the pelvics in P. pilosus is greater than the length of the anterior margin of the pectoral. In G. eastmani the reverse applies. All of the specimens of P. pilosus that I have seen have long been in preservative and the color has become more or less uniform brown. In three of the large specimens a series of small light-colored dots can still be seen along the lateral line. The presence of a series of white dots was emphasized by Englehardt (1912) as one of the ways to distinguish his species, Pristiurus hertwigi, from eastmani. He further separated the two by noting that hertwigi has a shorter snout and short pectoral fins, characters that apply to P. pilosus. Evidently Englehardt overlooked German's (1906) earlier description of P. pilosus. The type-series of Englehardt's Pristiurus hertwigi was lost by bomb damage in World War II and I cannot find differences other than color in Englehardt's description to separate hertwigi from pilosus. A 563-mm male was found to be immature and none of the other specimens seen gave any indication of the size at maturity. Nothing is known of the reproductive pattern. Parmaturus pilosus is larger than P. xaniurus which seems to be mature at about 450 mm. Description. The illustration of an immature male, its teeth and denticles, in Garman's Plagiostomia (1913, pi. 8, figs. 7-10) is excellent and should be consulted for additional details. Following are measurements expressed as percentages of total length for a 563-mm immature male, first figure in each pair, and a 590-mm female, second figure in each pair. Tip of snout to: front of mouth, 5.3, 4.6; eye, 5.7, 4.4; spiracle, 10.7, 9.8; first gill slit, 15.1, 13.6; fifth gill slit, 19.4, 18.3; origin pectoral fin, 18.6, 18.0; origin first dorsal fin, 45.5, 45.4; origin pelvic fins, 44.8, 44.1; origin second dorsal fin, 63.6, 63.4; origin anal fin, 58.6, 57.6; origin upper caudal fin, 71.0, 71.2; anterior end cloacal opening, 47.4, 46.1. Greatest width of: trunk at pectoral origin, 9.1, 11.0; trunk at pelvic origin, 6.2, 5.6; caudal peduncle, 1.8, 1.9. Greatest height of: trunk at pectoral origin, 11.1, 11.0; trunk at pelvic origin, 10.1, 8.8; caudal peduncle, 3.9, 3.9. Eyes: length, 4.4, 3.9; height of opening, 1.2, 1.4. Spiracles: greatest diameter, 0.7, 0.7; least distance from eye, 1.1, 1.0. Mouth: width, 7.8, 7.6; length, 4.8, 3.7; length upper labial furrow, 1.1, 1.2; length lower labial furrow, 1.7, 1.5. Nasal apertures: least distance between, 2.1, 2.2. Gillslits: height of first, 2.3, 2.0; height of fifth, 1.1, 1.2. First dorsal fin: length base, 6.6, 6.4; length free inner margin, 2.1, 1.9; height, 5.9, 5.4; length anterior margin, 9.8, 9.8. Second dorsal fin: length base, 6.7, 6.4; length free inner margin, 1.4, 2.4; height, 4.6, 4.6; length anterior margin, 9.8, 9.7. Anal fin: length base, 10.1, 11.5; length free inner margin, 1.1, 1.0; height, 4.3, 4.7; length anterior margin, 7.1, 8.0. Pectoral fins: width base, 4.1, 5.1; greatest width of fin, 7.3, 7.0; length anterior margin, 8.0, 7.6. Pelvic fins: overall length exclusive of claspers, 8.9, 9.8; length inner margin claspers, 7.6, — ; extent claspers past pelvic fin tips, 3.0, — . Distance between fin bases: first and second dorsals, 13.0, 12.2. Among the specimens examined the smaller ones had narrow, nearly erect, and needlelike dermal denticles, the larger examples had tridentate denticles more or less erect. The teeth, however, had more cusps in the smaller specimens, usually five or more cusps whereas the larger specimens had only three cusps on the larger teeth of the middle parts of the jaws. The snout shape was more pointed or conical in the larger specimens than in the smaller ones. If specimens referred to P. pilosus represent only one species it appears that marked changes in tooth and denticle form are to be expected. The number of monospondylous vertebrae was 42 in each of three specimens, the total number of vertebrae 130 to 135. Discussion. The 590-mm female P. pilosus had a short (about 15 mm) section on the ventral edge of the caudal peduncle with denticles modified in the same way as in caudal crests. Parmaturus pilosus has a high concentration of squalene in its liver oil (Tsujimoto 1920). Since squalene is a low densi- ty hydrocarbon, it seems probable that its function is to give near neutral buoyancy to the shark and to eliminate much 107 of the energy requirement for a midwater habitat. I find no records of either depth of capture or of habitat observations for P. pilosus and implications of midwater habitat are all assumed and extrapolated from observations on P. xaniurus. According to Tsujimoto (1920) one of the Japanese names for P. pilosus is imori-zame, another shusu-zame, implying that P. pilosus was a reasonably well-known shark to some Japanese fishermen. Tsujimoto found no squalene in any other scyliorhinid that he examined. I have great confidence in Tsujimoto's determination not only because of his ex- cellent reputation as a chemist but also because he apparently had his sharks identified by Shigeho Tanaka whose work with Japanese sharks was outstanding in accuracy. The range of Parmaturus pilosus has to be given here as moderately deep waters of the coasts of Japan. It should be pointed out however, that 60 to 300 yr ago, deepwater sharks were caught by Japanese and Portuguese fishermen and no others except a few on elaborately and expensively equipped expeditions. Thus, the known ranges of deepwater sharks and especially of those sharks with midwater habitats may be determined by the distribution of highly skilled fishermen whose art is now lost because of economic pressures. Parmaturus xaniurus (Gilbert) Figure 67 Figure <>7. — Parmaturus xaniurus, 480-mm adult female filetail shark from 148 m off San Pedro. Calif. Drawing by Mary Wagner. Catulus xaniurus Gilbert 1892:540 (coast of southern California). Parmaturus xaniurus: Garman 1913:90, pi. 9; Roedel and Ripley 1950:48, fig. 32. Material examined. USNM 46719, ?, 550 mm, here designated lectotype, off southern California, lat. 33°55.5'N, long. 128°28'W, in 687 m, January 1889; USNM 46718, 4 i, 141-184 mm, 3 S, 140-163 mm, off southern California, paralectotypes; USNM 12059, 6$, 146-224 mm, 7 ?, 125-191 mm; USNM, imm. 5, 410 mm, 2 J, 428, 499 mm, off San Pedro, Calif., lat. 34°16'N, long. 120°30'W, 448 m; USNM, juv. 8, 135 mm, 2 juv. ?, 170-185 mm, off southern California, lat. 34°12.5'N, long. 130°03.4'W, 91 m. Diagnosis. Parmaturus xaniurus has a comparatively short snout and large eye. In specimens near adult size the length of the snout in front of the mouth is about equal to the horizontal diameter of the orbit. In P. manis and P. stenseni the snout is more than 2 times the length of the eye. In P. xaniurus the origin of the second dorsal fin is nearly opposite the origin of the anal fin, and the posterior tip of the second dorsal and anal fins are about opposite whereas in P. pilosus, P. melanobranchus, and P. campechiensis the tip of the second dorsal extends well past the anal tip. Parmaturus xaniurus has comparatively small pectoral fins as do others of the genus but the proportional size is ap- preciably greater in large adults than in young. Description. Parmaturus xaniurus is a soft-bodied shark without conspicuous markings. It is grayish-black, not much lighter below. The fins of some individuals may have darker spots, darker tips, or darker edges but these are not consistently present. The origin of the first dorsal fin is usually in advance of the midpoint of the pelvic fin base, the first dorsal is slightly larger than the second dorsal and the distance between their bases is about equal to the length of the base of the anal fin. Following are measurements of an adult female 467 mm long, first figure in each pair, and an immature male 135 mm long, second figure of each pair. Tip of snout to: front of mouth, 4.3, 4.1; eye, 4.9, 3.7; spiracle, 11.1, 8.5; first gill slit, 16.1, 17.8; fifth gill slit, 22.7, 23.7; origin pectoral fin, 21.0, 22.2; origin first dorsal fin, 46.7, 43.7; origin pelvic fins, 45.0, 41.1; origin second dorsal fin, 61.7, 59.7; origin anal fin, 61.7, 58.3; origin upper caudal fin, 71.1, 67.1; anterior end cloacal opening, 47.1, 44.5. 108 Greatest width of trunk at: pectoral origin, 13.3, 11.1; pelvic fin origin, 5.6, 6.4; caudal origin, 1.7, 2.2. Greatest height of trunk at: pectoral fin origin, 9.6, 9.0; pelvic fin origin, 7.4, 10.7; caudal fin origin, 3.7, 3.2. Eyes: length, 4.3, 4.3; height of opening, 1.1, 1.7. Spiracles: greatest diameter, 0.6, 0.6; least distance from eye, 1.1, 1.2. Mouth: width, 9.0, 11.9; length, 5.1, 4.1; length upper labial furrow, 0.9, 1.3; length lower labial furrow, 1.7, 2.1. Gill slits: height of first, 2.4, 3.7; height of fifth, 2.6, 2.2. First dorsal fin: length base, 7.3, 7.4; length free inner margin, 3.4, 2.6; height, 4.3, 3.2; length anterior margin, 10.7, 9.6. Second dorsal fin: length base, 6.4, 6.7; length free inner margin, 3.2, 3.0; height, 3.4, 3.0; length anterior margin, 9.0, 8.9. Anal fin: length base, 9.4, 8.9; length free inner margin, 1.9, 1.3; height, 4.3, 2.4; length anterior margin, 8.1, 4.8. Pectoral fins: width base, 5.1, 4.4; greatest width fin, 8.1, 4.6; length anterior margin, 10.7, 7.4. Pelvic fins: overall length, 12.4, 10.7. Distance between fin bases: first and second dorsals, 9.6, 8.9; pectoral and pelvic, 18.0, — ; pelvic and anal, 8.4, — ; anal and lower caudal, 2.1, — ; second dorsal and upper caudal, 4.3, — . The teeth of the 467-mm female P. xaniurus were in approximately 41+45/42+42 vertical rows, the upper and lowers similar in shape, the longest upper tooth about 0.8 mm from the enamel line, the longest lower tooth only about 0.4 mm high. A few of the central rows of upper jaw teeth with three cusps, most of the uppers with four cusps, and a few near mouth corners with five cusps. Most of the lower jaw teeth with four cusps. Dermal denticles needlelike in young, dermal denticles of adults with three points, much as in other scyliorhinids. Ovary (only right ovary developed) of 467-mm female with 10 large eggs 8 to 13 mm in diameter. Oviducts mature (stretched about 7 mm in diameter); liver large reaching level of cloaca; spiral valve with seven turns. Number of monospondylous vertebrae 38 in two specimens, 39 in five specimens examined. Total number of verte- brae in seven specimens 109 to 121. Discussion. Parmaturus xaniurus is the only species of the genus known from more than a dozen specimens and is probably the only scyliorhinid to have been observed in moderately deep water. In California, P. xaniurus is well-known to ichthyologists and to a few deepwater trawl fishermen as the filetail shark. Walford (1935) noted that it occurred from southern California southward to about Port San Bartholome but was rarely taken by fishermen. Roedel (1951) reported the capture of 15 specimens taken off Santa Barbara as noteworthy. Kato et al. (1967) gave the range as central Califor- nia southward to the Gulf of California. Mathews and Ruiz (1974) recorded the species as probably collected, but not certainly identified, from the northern part of the Gulf of California. The best clue to the distribution of Parmaturus xaniurus, and by inference to all other species of Parmaturus as well, was furnished by collections made by the General Motors research vessel Swan in the Santa Barbara and Santa Cruz basins. Hauls with an Isaacs-Kidd self-closing midwater trawl caught P. xaniurus in 43 hauls made from 10 to 490 m above the bottom where water depths in the Santa Barbara basin were 500 m or more (Lee 1969). No other small sharks were collected although Apristurus brunneus and Cephaloscyllium venustum are known from the region that includes the Santa Barbara basin. On a dive in the research submarine Dowb several small sharks were seen at the bottom in the deepest part of the Santa Barbara basin (William Aron, pers. commun.). The sharks were estimated to be less than 600 mm long and were noted especially by Aron because the water there is known to be low in oxygen (Emery 1960). With the exception of the sharks, Aron observed no benthic vertebrates except some apparently moribund myctophid fishes. Several myctophids were seen lying on the bottom or motionless a short distance above the bottom. One shark, observed briefly, had a myc- tophid in its mouth. The general impression was that the sharks were feeding on myctophids that had accidentally mov- ed into water too low in oxygen to support their activity. Although the kinds of sharks most often seen in aquariums seem to need moderate to high levels of dissolved oxygen for survival, some others are known to thrive in regions of low oxygen such as in the deeper parts of the Red Sea (Marshall and Bourne 1964) or to be taken in situations of very low oxygen (Compagno and Springer 1971). A question of species identity for the Santa Barbara sharks remained so Phillip Heemstra, who was well acquainted with the silhouettes of small sharks, made a later dive aboard the Dowb. Sharks were again observed and Heemstra identified them positively as Parmaturus xaniurus. Feeding was not observed. Livers and fresh liver oil from three immature female P. xaniurus were obtained from Santa Barbara fishermen by R. S. Lee and examined by Mary H. Thompson, National Marine Fisheries Service, Miami. The liver oil was 54 to 59°c squalene and the whole oil had a specific gravity of 0.886 to 0.891 at 20°C. Except for P. pilosus and P. xaniurus, liver oil squalene has not been found in any other scyliorhinid shark although it is found in some squaloid sharks, in Chlamydoselachus, in Cetorhinus maximus, and in some odontaspids of oceanic habitats. One squaloid, Squalus acanthias, the spiny dogfish, has been shown experimentally to be able to adjust its sinking factor by shifts in the ratios of concentrations of diacyl glyceryl ethers to triglycerides in the liver (Malins and Barone 1970). It seems, therefore, not 109 unlikely that many sharks can make some minor or fine adjustments of hydrostatic balance by metabolic means. Such changes presumably would take hours or minutes to become effective. Large livers, with their content of light-weight oil, greatly reduce the energy requirements for slow swimming and are important as hydrostatic organs (Baldridge 1970, 1972) in many shark species. This seems always to be only a coarse ad- justment. The substitution of the lower specific gravity squalene for other liver oils, such as is the condition in most midwater sharks, may still be only a coarse adjustment to full hydrostatic balance. The method of fine adjustment or the attainment of neutral buoyancy that is presumed here as necessary for the shark's mastery of the midwater environment, is still largely unknown but metabolic changes of liver components, such as found by Malins and Barone (1970), may indicate one general method. Cox (1963) described the egg capsule of Parmaturus xaniurus as being 74 to 110 mm by 28.5 to 36 mm, as being relatively slender, and as having a smooth surface. His illustration (fig. 10) showed wide lateral flanges which in the text were noted to be T-shaped in cross-section. This is sufficiently unusual to distinguish P. xaniurus egg capsules from others known from the California coast. One egg capsule collected on 29 September 1966 in about 450 m had been attached to a black coral "tree" (Antipathes) along with about 184 other capsules (Bill Bradley, pers. commun.). Bradley's log noted that the water temperature was 6°C and that no filetail sharks were seen there on the day the egg capsule was recovered but that filetails had been seen among large rocks in the locality (lat. 32°38.8'N, long. 117°29.3'W) on the previous day. The recovered egg capsule was placed in a home refrigerator and apparently was alive with an active embryo about "half developed" when it died about a year later. PENTANCHUS SMITH AND RADCLIFFE Pentanchus Smith and Radcliffe in Smith 1912:490 (type-species, Pentanchus profundicolus Smith and Radcliffe, by original designation). Diagnosis. A scyliorhinid shark with only one dorsal fin, its origin over the middle of a long anal fin; body strongly compressed, head moderately flattened with a long, somewhat spatulate snout abruptly narrowed in advance of the nostrils and tapering to a rounded point; nasal apertures large, oblique; nasal flaps represented only by small points, not large enough to cover outer nasal opening; no crest of modified denticles along upper edge of caudal fin; eyes more dorsal than lateral; five gill slits. Discussion. Pentanchus is known only from the holotype of P. profundicolus. It was first described as a notidanoid shark in a new family, the Pentanchidae, because of its single dorsal fin. Pentanchus is similar to Apristurus species in so many respects that Fowler (1941) regarded the name Pentanchus as superceding Apristurus. I follow Garman (1913) and Bigelow and Schroeder (1948), however, in retaining Apristurus as distinct from Pentanchus at the generic level. Pentanchus profundicolus Smith and Radcliffe Pentanchus profundicolus Smith and Radcliffe in Smith 1912:490, pi. 42; Garman 1913:95; Bigelow and Schroeder 1948:196. Material examined. USNM 70260, ad. 3, 495 mm, Albatross Stn. 5486, lat. 10°02'N, long. 125°19'20"E, Sea of Mindanao, Philippines, 1,069 m, 31 July 1909, holotype and only known specimen. Condition of the holotype. The holotype (USNM 70260) is in fair condition. The original description gave the length as 508 mm and it now measures 495 mm, but the difference is undoubtedly due to normal shrinkage of scyliorhinid specimens that have been preserved in alcohol for long periods. At some time in the past, possibly when the specimen was first examined, the skin of the top of the head was peeled back and the upper part of the chondrocranium was remov- ed and evidently discarded. It is consequently impossible to determine whether or not cranial crests projected above the orbits. Also cuts were made at a short distance away from the dorsal midline and a few trunk vertebrae and a few vertebrae at the base of the caudal fin were removed and evidently discarded. The cuts did not damage the midline where a dorsal fin might be expected and the specimen now shows no indication at all of fin components or of scar tissue from accidental damage. The trunk section of the specimen is still firm and the number of trunk vertebrae removed can be estimated (7'/2 removed) with confidence. The number of tail vertebrae removed can be determined from the rem- nants of the neural arches still apparent in radiographs. X-ray examination of the specimen revealed one interesting and unexpected condition for which no explanation is offered. The degree of calcification is as great as or greater than in any other scyliorhinid, not only in the vertebrae but also in fin radials, components of visceral arches, and even in rostral cartilages. 110 Diagnosis. Pentanchus profundicolus may be distinguished from other scyliorhinids by the presence of only one dorsal fin but otherwise has most of the characteristics of Apristurus. In addition, however, P. profundicolus differs from species of Apristurus in having a very short visceral cavity with broad and long pectoral fins so that the tips of the ap- pressed pectorals nearly reach the origin of the pelvic fins. Perhaps the body cavity is somewhat longer in adult female P. profundicolus, but it seems more likely that the body cavity in both sexes of P. profundicolus will be found to be un- usually short. In P. profundicolus the snout is more strongly narrowed in advance of the nasal apertures and tapers to a rounded point rather than being broadly rounded as in species of Apristurus. The base of the anal fin in P. profundicolus is somewhat longer than in specimens of Apristurus that I have measured. Jordan and Hubbs (1925) reported on a stuffed scyliorhinid shark with only one dorsal fin observed in a Japanese museum. They did not identify the shark as to species or genus but their account of it, noting small pectoral fins, large pelvic fins, and a short caudal fin, would not apply to P. profundicolus. The stuffed specimen seems to have been lost (Nakaya 1975). Description. Proportional measurements as percentages of the total length in the 495-mm holotype of P. profundicolus follow. Tip of snout to: level of anterior end of nasal apertures, 6.7; level of posterior end of nasal apertures, 8.9; front of mouth, 10.0; eye, 10.7; first gill slit, 20.6; last gill slit, 25.3; origin pectoral, 23.8; origin dorsal fin, 60.6; origin pelvics, 38.2; origin anal, 49.5; upper caudal fin origin, 71.7; anus, 41.0. Greatest width of: head, 11.5; trunk at pectoral origin, 9.3; trunk at pelvics, 5.1; tail at caudal origin, 1.6. Greatest height of: head at spiracles, 6.9; trunk at pectoral origin, 8.9; trunk at pelvics, 8.9; tail at caudal origin, 5.1. Eyes: horizontal diameter, 3.2. Spiracles: greatest diameter, 0.4; least distance from eye, 0.8; distance between, 6.5. Mouth: width, 6.9; length, 2.6; length upper labial furrow, 3.4; length lower labial furrow, 2.6. Nasal apertures: minimum distance between, 3.2; greatest diagonal length, 3.6. Gill slits: height of first, 0.6; height of fifth, 1.1. Dorsal fin: length base, 7.3; length free inner margin, 2.8; height, 3.0. Anal fin: length base, 20.2; height, 3.4. Pectoral fins: width base, 7.5; length anterior margin, 14.6; greatest width, 10.5. Pelvic fins: length (origin to rear tip), 9.5. Distance between fin bases: pectoral and pelvic, 5.9; pelvic and anal, 2.6; anal and origin lower caudal, 0.0; dorsal and origin upper caudal, 3.8. The teeth are generally similar in the upper and lower jaws but the upper teeth have slightly higher cusps. Upper teeth in the central part of the jaw may have only three cusps, the central cusp much the longest, but most of the teeth have five cusps and the central cusps are lower toward the angles of the jaws. The teeth have striations near their bases. Smith and Radcliffe in Smith (1912, pi. 42) showed the main tooth cusps somewhat lanceolate, constricted at the bases. This seems to be an error. I have found the lanceolate form of tooth cusps in some juvenile Apristurus and in some juveniles of related genera, but cusps of the teeth of the holotype taper regularly from their bases to near their tips. The largest teeth are about 1.5 mm high. The teeth are numerous, alternate in both jaws, and those in the lower jaw form diagonal rows. I did not attempt to count the number of rows since counting would have required cutting one jaw corner. Whether or not gill rakers are present was not determined. The dermal denticles are tridentate, small, the blades of the largest about 0.5 to 0.7 mm long, imbricate, with a single median ridge. Denticles are similar in shape over most of the body surfaces but lateral and ventral denticles have reduc- ed lateral points somewhat less prominent than those shown in Smith and Radcliffe's figure (in Smith 1912, pi. 42). No caudal crest of modified denticles is present. The general appearance of P. profundicolus in its lateral aspect is well shown in Smith and Radcliffe's illustration. The first gill slit is shorter than the fifth as in the illustration. It appears that the angle of view of the eyes may be less dorsal than in species of Apristurus but the condition of the head of the specimen does not allow any assurance on that question. Another difference is that the patch of pores on the under side of the snout is not divided along the midline as it is in Apristurus. The viscera of the holotype are not at present in good condition but it does appear that the liver is unusually small in comparison with species of Apristurus, its lobes not reaching posteriorly for more than half the length of the short body cavity. The color of the holotype is now as described by Smith and Radcliffe, uniform dark brown. Springer and Garrick (1964:86) estimated the number of vertebrae (because of several centra missing) as 74 precaudal, 54 caudal. Ill PORODERMA A. SMITH Poroderma A. Smith 1837:85 (type-species, Squalus africanus Gmelin, 1789:1494, designated by Fowler 1908:53). Conoporoderma Fowler 1934:234 (type-species, Scyllium pantherinum Muller and Henle, by original designation). Diagnosis. Species of Poroderma are the only scyliorhinid sharks with cone-shaped barbels that extend from the nasal flaps for a distance at least as great as the diameter of the barbel base. Adult Poroderma are conspicuously marked either by dark longitudinal stripes, by round solid spots, by crescent-shaped markings that may be arranged in pairs to form interrupted circles, or by short bars. Poroderma closely resembles Scyliorhinus in many respects. In both genera the supraorbital crests of the chon- drocranium extend above the orbits as narrow shelves, lower labial furrows are present but labial furrows are not con- tinuous around the mouth corners and a true upper labial furrow is not present, the upper lip is very slightly expanded or enlarged at each mouth corner and more or less overlaps the lower lip as an inconspicuous lobe. The margin of the mouth corner lobe of the upper lip curves laterally and then anteriorly, especially in Poroderma and may then look superficially like an upper labial furrow. It can be seen easily, however, that the upper lip margin is entirely distinct from the lower labial furrow and is not connected with it. A subocular gutter appears to be somewhat less distinct in Poroderma than in Scyliorhinus and an apron formed beneath the claspers by union of the basal section of the inner margins of the pelvic fins is less developed in Poroderma than in Scyliorhinus. Discussion. Bigelow and Schroeder (1948:197) rejected the name Poroderma for the genus because its type-species, Squalus africanus Gmelin, designated by Fowler, "lacks barbels." It is true that Gmelin's (1789:1494) description of Squalus africanus includes no reference to barbels but it does not specifically indicate their absence. Gmelin's descrip- tion does state that Squalus africanus has seven parallel longitudinal lines of darker color, a striking color pattern that is found on only one described species of shark, a species with barbels. Andrew Smith (1837) listed four species, all from the Cape of Good Hope region, as belonging to Poroderma: P. africanum, P. pantherinum, P. variegatum, and P. submaculatum. Johannus Muller was present at the session of the Zoological Society of London at which Smith "demonstrated" his species. Muller and Henle (1841) published accounts of the first three species, crediting Smith. Subsequent authors have recognized variously one to three species. Regan (1908a), Garman (1913), and Barnard (1925) included only africanum and pantherinum in their accounts of Poroderma. Fowler (1934) added a species, P. marleyi, based on a 225-mm specimen from the coast of Natal. Fowler's (1941) later treatment of Poroderma and Smith's (1949) account have keys to the three species africanum, pantherinum, and marleyi. I have had only five specimens for study (see under material examined, P. africanum and P. pantherinum). Although my treatment of Poroderma here is based on my own observations and on literature references from 1949 and earlier, it is not essentially in conflict with the much more comprehensive account of Bass et al. (1975:27-32). Key to Poroderma, Adapted from J. L. B. Smith (1949) la. Three to seven black longitudinal stripes on body; adults large, more than 700 mm P. africanum lb. Black spots on body, either randomly distributed or organized into longitudinal rows 2 2a. Spots most often with light colored centers, arranged as pairs of crescent shaped marks, or irreg- ularly formed into various shaped marks, sometimes arranged in longitudinal bands; adults mature at somewhat smaller size than africanum P. pantherinum 2b. Spots of solid color and random distribution; a small species reaching about 450 mm marleyi Poroderma africanum (Gmelin) Figure 68 Squalus africanus Gmelin 1789:1494 (African seas). Scyllium africanum: Muller and Henle 1841:12; Gunther 1870:405 (part). Scyliorhinus africanus: Regan 1908a:456. Poroderma africanum: Garman 1913:70; Fowler 1941:39; Bonde 1948:465, pis. 13-16; Smith 1949:53, fig. 37; Bass et al. 1975:28, table 5, fig. 16. 112 Figure HH.— Poroderma africanum, adult male about 750 mm, Cape Peninsula, off Slangkop, South Africa, USNM 203468. Material examined. USNM 203468, ad. $, about 750 mm, Cape Peninsula, off Slangkop, South Africa; USNM, imm. S, 570 mm, same locality. Diagnosis. Poroderma africanum differs from P. pantherinum and P. marleyi as well as from all other scyliorhinids in having three to seven dark stripes (most often five) extending from the head longitudinally along the back toward the tail. The middle three or five dark stripes are usually continuous, not broken into spots anterior to the first dorsal fin. The background between dark stripes and the ventral surface is cream colored. The fins of P. africanum are unspotted whereas in P. pantherinum and P. marleyi all fins are spotted with darker color. Description. Poroderma africanum is one of the larger catsharks, reaching a length of 950 mm (Barnard 1925). Bass et al. (1975) reported that specimens were sexually mature between 580 and 760 mm for males and between 650 and 720 mm for females; they observed no indication of sexual dimorphism in tooth shape. The specimens I examined were strongly curved in preservation and not suitable for measurement, but Bass et al. (1975, table 5) provided an analytical summary of measurements of nine males and seven females. Vertebral counts in the two specimens I examined were: total 117, 120; monospondylous 44, 45; precaudal 82, 85; caudal 35, 35. Discussion. According to Bonde's account (1948), Poroderma africanum is an oviparous species and lays egg capsules about 103 by 50 mm, the capsules with long tendrils at each corner. Of two egg capsules laid at the Sea Point Aquarium by an 850-mm P. africanum, one was not viable, but the other hatched a young one 145 mm long on the 164th day. The egg capsule was transparent and smooth-surfaced, although barely visible striations indicated the structure of the cap- sule walls. Bass et al. (1975) reported that P. africanum is abundant in shallow waters off the southwestern Cape; that its range off the east coast of Africa does not reach Natal; and that the extent of its range off the southwestern coast of Africa is undetermined. They observed P. africanum lying among the rocks and in caves at depths of about 10 m in Flase Bay and quoted a report that it is nocturnal in captivity. 113 Poroderma marleyi Fowler Poroderma marleyi Fowler 1934:234 (Natal coast in 37 m); Smith 1949:53, pi. II, fig. 35; Bass et al. 1975:29, fig. 17. Material examined. None. Diagnosis. Randomly distributed round, dark spots of solid color, none much larger than the eye, distinguish P. marleyi from other Poroderma. The holotype of the species, ANSP 53427, was about 225 mm long and Smith's illustra- tion (1949, fig. 35) showed a specimen of about that size. Bass et al. (1975:29) had a 580-mm mature male from the coast of Natal. Poroderma pantherinum (Miiller and Henle) Figures 69, 70, 71 Figure 69. — Poroderma pantherinum, 610 mm female. Cape Blaize, South Africa. Figure 70. — Poroderma pantherinum. ventral side of head of (ill) mm female. Scy Ilium pantherinum Miiller and Henle 1841:13 (Cape of Good Hope). Scyllium africanum Var. y. pantherina: Giinther 1870:406. Poroderma pantherinum: Fowler 1941:37; Smith 1949:53, fig. 36; Bass et al. 1975:30, fig. 18. 114 Figure 71. — Poroderma pantherinum. Color pattern of dorsal side of tail of 397-mm immature male. Material examined. USNM, imm. 8, 430 mm, Cape Peninsula off Slangkop, South Africa; BMNH 1900.11.6.19, imm. S, 397 mm, 2, 610 mm, off Cape Blaize, South Africa in 69 m. Diagnosis. Some earlier authors cited the greater length of the barbels, barbels reaching mouth, as a way to distin- guish the present species from P. africanum in which the barbels are said not to reach the mouth. In my material, how- ever, the barbels are somewhat slenderer and proportionally longer than those of P. africanum, but do not reach the mouth in a position that appears normal, and, conversely, barbels of P. africanum may reach as far back as the mouth if pushed toward the middle of the jaws. Specimens of P. pantherinum are clearly distinct from P. africanum in color pattern in having a color pattern of spots in various shapes and configurations. Poroderma pantherinum also has spots on fins, but fins of P. africanum are free from dark markings. Description. An analytical summary of measurements of five males and five females was included in the treatment of Poroderma pantherinum by Bass et al. (1975, table 5). The color pattern and its variation in P. pantherinum is best understood by reference to illustrations (see Bass et al. 1975, fig. 18; Smith 1949, fig. 36; and Figs. 69 and 71 in this study). Vertebral numbers in a 430-mm male are: total 114, monospondylous 41, precaudal 80, caudal 34. Discussion. According to Giinther (1870) the type of Scy Ilium pantherinum is an Algoa Bay, stuffed specimen, 27 in long, from the collection of A. Smith. In the original description, Midler and Henle (1841) mentioned such a specimen among several others as belonging to this species and gave Cape of Good Hope as the locality for all. Bass et al. (1975:31) recorded lengths from 540 to 740 mm for mature males and 580 to 710 mm for mature females. They did not have data on hatching size or on egg capsules. SCHROEDERICHTHYS SPRINGER Schroederichthys Springer 1966:604 (type-species, Schroederichthys maculatus Springer, by original designation). Diagnosis. The tail section of Schroederichthys is longer than the head-trunk section, the distance from the anterior end of the cloacal opening to the tail tip about three-fifths or more than three-fifths the total length in adults and as much as two-thirds the total length of juveniles of some species. Members of the genus have supraorbital crests extending as narrow shelves above the orbits; they have labial furrows continuous around the mouth corners along both upper and lower jaws. They lack a crest of modified denticles on the upper edge of the caudal fin and the pelvic fins of males are not united along their inner margins to form an apron above the claspers. Discussion. Schroederichthys belongs with the group of genera including Scyliorhinus, Poroderma, Cephaloscyllium, Atelomycterus, and Aulohalaelurus, all having supraorbital crests. The four species of Schroederich- thys are known only from Central American or South American coasts. The type-locality of one, S. bivius, I believe to have been recorded originally in error as the Cape of Good Hope, South Africa (see discussion under S. bivius). 115 Key to Species of Schroederichthys la. Nasal flaps nearly triangular, their margins far from parallel, tip or sides of broadly based trian- gular flaps sometimes with a very short lobelike extension at the apex of the triangle 2 lb. Nasal flaps lobelike, the lobes narrow and usually long, their lateral margins parallel or nearly so 3 2a. Dorsal and lateral surfaces tan or brown with numerous randomly distributed, nearly round, light-colored spots about the size of the expanded pupil or larger, usually also with six to nine saddle blotches slightly darker than ground color, indistinct or absent on head and trunk in most adults; small sharks, maximum length about 350 mm S. maculatus 2b. Dorsal and lateral surfaces variously light brown to dark gray or brownish-black and profusely marked with black; markings include many black spots, a few round black spots sometimes present on ventral surfaces; six to nine saddle blotches, sometimes including black spots, some- times all black, and in some specimens with a few white spots; moderately large scyliorhinids, largest seen an adult male 630 mm S. chilensis 3a. Dorsal surfaces typically gray-brown usually with about seven or eight saddle blotches some- what poorly defined; usually also some irregularly scattered spots of white, smaller than dark- adapted pupil but usually highly contrasted with ground color; juveniles (three specimens seen) with large black saddle blotches of solid color and contrasting white patches between blotches; a large scyliorhinid, adults to 700 mm . S. bivius 3b. Dorsal surfaces (of two immature type-specimens) brown with numerous small darker brown spots, mostly somewhat smaller than expanded pupil and some arranged in rows outlining sad- dle blotches; no white spots; a large scyliorhinid reaching 700 mm or more S. tenuis Schroederichthys bivius (Muller and Henle) Figures 72, 73 Figure 72.— Schroederichthys bivius, female, 440 mm, southern Chile, USNM 114725. Drawing by Mary Wagner. Figure 73.— Schroederichthys bivius, juvenile male about 110 mm long, Straits of Magellan. Drawing by Mildred Carrington. 116 H Scy Ilium bivium A. Smith 1837:85 (name only). Scyllium bivium: Miiller and Henle 1841:8 (Cape of Good Hope, probably in error); Giinther 1870:405. Scyliorhinus biuius: Regan 1908a:462; Norman 1937:8; Petit and Budker 1937:120. Scylium bivium: Lahille 1929:302, figs. 3, 4, 5. Halaelurus bivius: Garman 1913:86; Springer 1966:618, fig. 26; Kato et al. 1967:26, fig. 39; Gosztonyi 1973:317, pi. 1 and fig. 1. Material examined. BMNH 1936.8.26.12.14, 2 juv. 8, about 110 mm, juv. 5, about 155 mm, near Punta Arenas, Chile, lat. 53°39'S, long. 70°54'W; USNM 114731, 5 $, 552-625 mm, Lin Bay, Chile, lat. 41°54'S, long. 73°06'W; USNM 114725, ?, 436 mm, Castro, Chile, lat. 42°29'S, long. 72°46'W; USNM 114727, 2 ?, 437-447 mm, Puerto Montt, Chile; USNM 42061, ad. S, 700 mm, Straits of Magellan; BMNH 1879.5.14. 406, 2 egg capsules with well developed embryos, Straits of Magellan, January 1876; BMNH 1849.11.2.2, egg capsule. Diagnosis. Schroederichthys bivius and S. chilensis both vary considerably in color pattern and in the proportions that are often used in the diagnosis of scyliorhinids. I recommend that first attention in identification be given to the shape of the nasal flap. It does not vary greatly in shape in either species. In S. bivius it is moderately long and slender and definitely not broadly triangular as it is in S. chilensis. Identification of specimens from the region south of lat. 50°S is made difficult because both S. bivius and S. chilensis may be found there and exhibit secondary sexual dimorphism in mouth and tooth shape. In S. bivius, and possibly also in S. chilensis, proportional differences between young and adults are great. Possible differences in total length at sexual maturity in individuals taken from areas far apart should also be taken into account in future comparative studies of the two species. In side by side comparison of an adult male 625 mm S. bivius with an adult male 620 mm S. chilensis, it was noted that S. bivius has a somewhat more pointed snout, a narrower head, a longer and narrower mouth, somewhat larger eyes, much smaller spiracles, and smaller dermal denticles with less variety in denticle shape on the head region. Most impor- tant, however, the nasal flaps were narrow and long lobes on S. bivius but broad-based and relatively short on S. chilen- sis. The lobelike, narrow and long nasal flaps of half-grown S. bivius among specimens I have seen distinguished that species from S. chilensis. I have not myself, seen either adult female S. bivius or newly hatched S. chilensis. Description. Newly hatched S. bivius (Fig. 73) from the Straits of Magellan have very long tail sectors, their lengths from the anterior end of the cloacal opening to the tip of the tail two-thirds or more of total length. The tail section was shorter in the large specimens that I examined but was near three-fifths or more than three-fifths of total length. The S. bivius, that appeared from yolk scars to be newly hatched, were about 110 mm long. Norman (1937:8) said of S. bivius that no enlarged tubercles were present on the back. On the newly hatched S. bivius two rows of much enlarged denticles are present on the back but are represented partly by scars on the 155-mm specimen and are entirely absent on the half-grown specimens seen. The enlarged denticles on S. chilensis are of a different shape and apparently persist for a longer period but are frequently absent even on half-grown individuals. The series of enlarged denticles noted as present on juvenile S. bivius are the "dents cutanees jumelees" discussed in detail by Petit and Budker (1937). Petit and Budker pointed out the structural differences between the "twin denticles" of juvenile S. bivius and the dorsal tubercles with surrounding rosettes of smaller denticles on S. chilensis. The dorsal tubercles of S. chilensis evidently persist longer (or appear later) than the enlarged denticles of S. bivius. I found the S. bivius and S. chilensis material available for examination incomplete and somewhat baffling as did most earlier authors. This difficulty is now partly removed by the account of Gosztonyi (1973) on a series of S. bivius including adults of both sexes and egg capsules. Gosztonyi (1973) described for S. bivius the secondary sexual dimorphism in both mouth and tooth morphology from a study of a series of late embryos, juveniles, and adults of both sexes collected in an estuary of the Desado River (about lat. 48°S), Argentina. In adult male S. bivius, the lower jaw teeth are definitely larger than the upper and the teeth lack accessory cusps in both jaws. The adult male's teeth are at least twice as high as the teeth of females of comparable size. The teeth of females and young are multicuspid, mostly tricuspid near the middle of the jaw but with five or more cusps near the jaw corners. The mouth of the male at maturity develops a higher arch that becomes V-shaped with the midpor- tion of the jaws occluding approximately on a transverse line; the arch of the jaws not evenly rounded in a V-shape as in females and immature males. In Gosztonyi's series, males were mature at 530 mm. Some records of lengths show S. biv- ius may reach a length of 700 mm in the Straits of Magellan. This disparity in lengths of adults seems unusual but it should be noted that the geographical range of S. bivius extends as a narrow strip over a great distance and presumably has a large range of temperature differences. The following measurements are based entirely on notes that I made years ago on two adult males collected in Lin Bay, coast of Chile, at lat. 41°54'S, long. 73°06'W, 20 January 1945. Measurements are expressed as percentages of the total length, the first figure in each pair refers to a 625-mm adult male and the second to a 555-mm adult male. 117 Tip of snout to: front of mouth, 3.5, 3.8; first gill opening, 16.0, 15.1; fifth gill opening, 20.5, 18.4; origin pectoral fin, 19.2, 18.0; origin first dorsal fin, 43.8, 40.5; anterior end cloacal opening, 40.0, 37.9; origin upper lobe caudal fin, 81.4, 78.0. Eye: horizontal diameter orbit, 2.7, 2.9; vertical diameter, 1.0, 0.7. Nasal apertures: least distance between, 2.7, 2.3. Mouth: width, 6.4, 5.8; length, 5.6, 5.0; length upper labial furrow, 3.2, 2.7; length lower labial furrow, 2.7, 2.2. Gillslits: height of first, 1.6, 1.4; height of fifth, 1.0, 0.7. First dorsal fin: length base, 6.2, 5.6; length inner posterior margin, 3.2, 2.9; anterior margin, 10.2, 9.0. Second dorsal fin: length base, 6.7, 8.3; length inner posterior margin, 3.2, 2.9; length anterior margin, 10.9, 12.6. Anal fin: length base, 9.9, 9.4; length inner posterior margin, 2.4, 2.2; length anterior margin, 7.5, 7.4. Pectoral fins: length anterior margin, 12.2, 11.7; greatest width, 10.4, 8.6. Distance between fin bases: first and second dorsals, 16.0, 16.2. Length claspers: inner margin, 9.6, 9.4; extent past pelvic fin tips, 6.4, 4.9. Pectoral fins: width base, 4.8, 4.5; length anterior margin, 12.2, 11.7; greatest width, 10.4,8.6. Distance between fin bases: first and second dorsals, 16.0, 16.2. Greatest length clasper: inner margin, 9.6, 9.4; extent past tip of pelvic fins, 6.4, 4.9. No notably enlarged dorsolateral denticles are present except on juveniles. The dorsolateral denticles are moderately heavy with three points, the middle one strongest. The denticles of ventral surfaces generally with only one point and im- bricate. Dorsal surfaces grayish, with darker gray or black saddle blotches of irregular shapes. Some small white spots about 5 mm in diameter among some darker spots of same size. First dorsal fin origin about over posterior end of pelvic fin base; distance between dorsal fin bases about equal to dis- tance from tip of snout to first gill slit. Discussion. Miiller and Henle (1841) credited their description of Scyllium bivium to Andrew Smith and indicated that it was based on a dry specimen from the Cape of Good Hope. In a footnote, Miiller and Henle (1841:8) indicated that the species was established by Smith in the South African Quarterly Journal, Oct. 1831, No. 5, and in the Govern- ment Journal of the Cape, 1 November 1828, but stated that they did not see these references. P. A. Hulley of the South African Museum looked for the articles in the South African Public Library and the Library of the Houses of Parliament (South Africa). He found (pers. commun.) no publication under the title "Government Journal of the Cape," and found no mention of a fish by Smith in issues of the "Government Gazette of the Colony of Cape of Good Hope" for 27 October 1828 and 7 November 1828; there was no entry for 1 November. The 1831 article by Smith in the South African Quarterly Journal had no reference to a shark, only to three percid fishes. Both Andrew Smith and Johannes Miiller were present at the 1837 meeting of the Zoological Society of London at which Smith gave an untitled paper which was published as a part of the Proceedings for 1837 and also, under the title "On the necessity for a revision of the groups included in the Linnaean genus Squalus" in the Annals and Magazine of Natural History in its first (1838) volume. These published accounts included Scyllium bivium only by name. Data for the Miiller and Henle description were probably obtained at this time from Smith's specimen and apparently their de- scription in 1841 was the first and was based entirely on a single stuffed specimen that can be considered the holotype. The 27-in stuffed specimen later came to the British Museum (Natural History) and is noted by Giinther (1870:405) as the type of Scyllium bivium. I suspect that Giinther did not know that Miiller had already seen the specimen and may have taken detailed notes with descriptive data for he stated in a footnote (1870:405) that, "I do not understand how Miiller and Henle obtained their notes on the coloration, as they had only one example for examination, the same from which I have made the diagnosis, and the only one known to exist in collections." Victor G. Springer examined the specimen and sent me photographs of it. The specimen is now catalogued as BMNH 1857.10.20. I have no doubts that this specimen is the holotype of Scyllium bivium. I do have strong doubts about the type-locality given by Miiller and Henle as "Am Cap" (Cape of Good Hope). In view of the great interest in stuffed natural history objects during the 19th century it seems possible at least, that Smith bought a specimen in South Africa that had been collected near Cape Horn by a sailor and stuffed for sale by the sailor by the time the sailor reached South Africa. I am indebted to G. S. Myers for this suggestion. Another possibility is that some specimens or labels were mixed since it was customary then to exhibit specimens of new species at scientific meetings, evidently before they were catalogued or formally deposited in a museum. In this in- stance the holotype, as shown by its number, was catalogued in 1857, long after Muller and Henle described it and even longer after it was obtained and exhibited bv Andrew Smith. Schroederichthys chilensis (Guichenot) Figure 74 Scyllium chilense Guichenot 1848:362 (Chile); Giinther 1870:405. 118 Figure 74. — Schroederichthys chilensis, immature female, .'ii)0 mm, coast of Peru. Drawing by Mary Wagner. Scyliorhinus chilensis: Regan 1908a:462. Halaelurus chilensis: Garman 1913:83; Springer 1966:618, fig. 26; Kato et al. 1967:26, fig. 40. Schroederichthys chilensis: Chirichigno 1974:29, fig. 21. Material examined. USNM, ad. 8, 620 mm, 5, 530 mm, Corral, Chile, lat. 39°42'S, long. 73°27'W, 36-45 mm; USNM 77305, ad. S, 565 mm, Tome, Chile; USNM, ad. 8, 555 mm, Valparaiso, Chile; USNM 127764, 2, 480 mm, Independencia Bay, Peru; USNM 77738, ad. 8, Mollenco, Peru; USNM 114728, 2 imm. 8, 400-410 mm, Coquimbo Bay, Chile; USNM 103770, imm. 8, 325 mm, I. of Quinquina, Chile; USNM, imm. 2, 385 mm, imm. 8, 400 mm, Valparaiso, Chile; USNM, imm. ?, 255 mm, San Juan, Peru. Diagnosis. The characters in the key should separate S. chilensis from S. bivius in most instances. Additional differences between the two forms were noted above under S. bivius. It has never been stated clearly and comprehen- sively how to distinguish all S. chilensis from all S. bivius. Certainly this diagnosis is not complete and may not be free from error. Such a situation probably arises because both species are highly variable, both undergo great changes during growth, and both have sexually dimorphic adults. Furthermore, if convergence in diagnostic characters occurs in the areas of geographical range overlap as it seems to do, a full diagnostic account would become remarkably long. I believe, however, that in most instances S. chilensis can be distinguished by its broadly triangular nasal flap. Some caution should be observed because both S. chilensis and Halaelurus canescens may occur in the same locality, and both have triangular nasal flaps. Large, melanistic S. chilensis may thus superficially resemble H. canescens. Description. The measurements I have for Schroederichthys chilensis are for two immature males from the coast of Chile at about lat. 30°S and two immature females from the coast of Peru at about lat. 15°S. The following proportional measurements are expressed as percentages of the total length, the first pair of figures in each group refer to 365 and 392 mm males from Chile and the second pair in parentheses to 460 and 455 mm females from Peru. Tip of snout to: front of mouth, 3.3, 3.3, (3.3, 3.3); first gill slit, 15.2, 16.6, (16.1, 15.4) fifth gill slit, 18.3, 21.4, (22.2, 20.2); origin first dorsal fin, 38.4, 38.6, (42.4, 42.6); anterior end cloacal opening, 37.0, 38.2, (39.1, 41.1); upper caudal fin, 80.3, 74.7, (81.6, 81.4). Eye: horizontal length, 2.7, 2.8, (2.6, 2.6). Nostrils: least distance between, 2.2, 2.0, (2.1, 2.2). Mouth: width, 7.4, 7.7, (8.7, 7.7); length upper labial furrow, 2.2, 2.6, (2.4, 2.4); lower labial furrow, 2.2, 2.6, (2.6, 2.4). Least distance between bases: first and second dorsal fins, 16.4, 16.6, (13.7, 15.6). Gill slits: height of first, 1.6-2.0, (2.1, 2.0); height of fifth, 1.1, 1.3, (1.5, 1.5). First dorsal fin: length base, 7.4, 7.7, (7.3, 8.4); length free inner margin, 3.6, 3.1, (3.7, 2.9); length anterior margin, 11.0, 10.4, (11.5, 12.3). Second dorsal fin: length base, 8.8, 9.2, (9.3, 9.2); length free inner margin, 4.1, 3.1, (3.7, 3.1); length anterior margin, 12.3, 12.0, (13.0, 11.7). Anal fin: length base, 8.9, 8.9, (7.6, 7.7); length free inner margin, 2.7, 2.8, (3.3, 2.9); length anterior margin, 7.4, 8.2, (7.6, 7.7). Pectoral fins: width base, 5.2, 5.1, (4.6, 4.8); length anterior margin, 12.3, 12.0, (11.1, 11.2). Pelvic fins: length anterior margin, 8.2, 7.1, (7.8, 7.5); length inner posterior margin, 8.7, 7.5, (5.2, 5.4); length outer posterior margin, 7.3, 7.0, (5.8, 6.4). 119 Base of second dorsal fin slightly longer than base of first dorsal but areas of the two fins nearly equal. Pectoral fin tips rounded, the fins broad and short. First dorsal origin about over middle of pelvic fin base. Second dorsal origin about over middle of anal fin base. Teeth in specimens examined varying from 23 to 33/22 to 33 vertical rows with four to six series apparently functional. Tooth arrangement alternate. Teeth of immature of both sexes and of adult females multicuspid. Teeth of adult males unicuspid. The dermal denticles of half-grown S. chilensis conform to the usual pattern in scyliorhinids, that is, the dorsolateral denticles have three points, the middle point the longest, they may have somewhat erect blades and may be sparsely dis- tributed; the ventral denticles have only one point and are usually closely imbricate. In half-grown S. chilensis, a.few large denticles are arranged in two series of about 12 to 20 denticles, each series extending along the dorosolateral surface from near the level of the spiracles to the level of the origin of the first dorsal fin. Each of the enlarged denticles is sur- rounded by a rosette of about 12 to 18 smaller denticles. The enlarged denticles persist at least sometimes on adults but may be unequal and irregularly arranged. The various descriptions in the literature suggest that juvenile S. chilensis have enlarged denticles but I am uncertain that newly hatched specimens have been examined and doubt that comparisons of young S. chilensis with young S. bivius have yet been made. Structural differences between enlarged denticles of young S. bivius and S. chilensis of un- specified age or size have been described by Petit and Budker (1937). They use the term tubercle to describe the enlarged denticles of S. chilensis. Between half- grown and adult stages in S. chilensis there appears to be a trend toward occurrence of proportionally larger denticles on dorsolateral surfaces especially on the head and anterior trunk. These denticles also tend to lose points or any specific shape or surface sculpturing. The head becomes well armored with a pavementlike assortment of smooth denticles and is presumably well adapted to probing among rocks, corals, or armed invertebrates. Schroederichthys chilensis has a color pattern of seven or eight distinct and often dark gray or black saddle blotches with a few nearly round dark gray or black spots between the saddles, these spots somewhat larger in diameter than the spiracles. Some specimens have few such spots on the ventral surfaces. Some individuals have a few small white spots. The ground color of some specimens is light gray dorsally and white below but more or less general melanism may occur. Some of the pigment occurs in the skin, but in the darker individuals it is also incorporated within denticles. Discussion. Schroederichthys chilensis frequents shallow water, at least in the southern part of its range, as might be inferred by its small eye (smaller than eyes of Halaelurus canescens) and large denticles, somewhat smooth surfaced in the head region. It is known only from the west coast of South America. Its range overlaps that of S. bivius in southern Chile and extends northward to Peru. I have not verified a record of its occurrence south of lat. 39°42'S. I have not seen an egg capsule of S. chilensis or any reference to one. Schroederichthys maculatus Springer Figures 75, 76 Figure 75.— Schroederichthys maculatus, adult male holotype, 330 mm, USNM 185556. ,Dorsal saddle blotches in this drawing^are somewhat more prominent than is usual for the species. Drawing by Mildred Carrington. Schroederichthys maculatus Springer 1966:605, fig. 4A (Caribbean Sea, NNW Cape Gracias a Dios, Honduras). Material examined. USNM 185556, ad. S, 328 mm, holotype, off Honduras, 410 m, 21 August 1957; USNM 185557, ad. ?, 335 mm, paratype, from same haul as holotype; USNM 187684, about 35 specimens, off Rosalind Bank, western Caribbean Sea, 274 m; USNM 187687, 4 specimens, off Rosalind Bank; USNM 187690, 7 specimens, off Quita Suena Bank, western Caribbean; USNM, 2 specimens, off western Colombia, 274 m. Diagnosis. Schroederichthys maculatus is a small scyliorhinid, the maximum length about 342 mm. It has a short trunk and a very long caudal section, the caudal section in adults about two-thirds total length, proportionally longer than in adults of any other scyliorhinid except possibly S. tenuis. Caudal fin is only about one-fifth total length. 120 Figure 76. — Schroederichthys maculatus. A. Ventral side of head of 328-mm adult male holotype, USNM 185556. B. Egg capsule taken from oviduct of 342-mm female, midline length of capsule 39 mm, anterior end of capsule (with- out horns and tendrils) probably incomplete. Schroederichthys maculatus is not sexually dimorphic in tooth shape. It has broadly triangular nasal flaps unlike the narrow lobelike flaps in S. tenuis and S. bivius. It has no enlarged dorsal denticles such as are to be found on S. chilensis. Schroederichthys maculatus is tan or light brown dorsally with randomly distributed, approximately round, yellowish or white spots, the spots about the size of the pupil. Young or half-grown S. maculatus have seven to nine sad- dle blotches which may faintly persist on adults or may be absent. In color and pattern Schroederichthys maculatus and Scyliorhinus torrei are identical and they may be expected to oc- cur in some of the same localities although available material does not show overlapping ranges. Of course, maculatus may be recognized by its much longer tail section and also by the characters that distinguish Schroederichthys from Scyliorhinus. Schroederichthys has labial furrows continuous around the mouth corners. Scyliorhinus does not and lacks furrows along the upper jaw. Schroederichthys also lacks the inconspicuous tabs of the upper lip that overlap the lower lip at the mouth corners in all species of Scyliorhinus. The pelvic fins of Schroederichthys males are not united along their inner margins as they are in Scyliorhinus. Description. In the series of Schroederichthys maculatus examined, 40 males ranged from 145 to 330 mm long and all of those longer than 280 mm were sexually mature. Nine females ranged from 220 to 342 mm. The 342-mm specimen was gravid with one fully formed, opaque, greenish, egg capsule. The egg capsule was striated longitudinally and was 44 by 14 mm with threadlike tendrils 225 mm long. Head broad and somewhat flattened; trunk short postpelvic section very long, about two-thirds total length, but caudal fin short, about one-fifth length; trunk rather slender; snout with a rounded point. Nasal flaps triangular, their points crossing or covering posterior part of nasal apertures; mouth with a moderately high arch, somewhat higher and more V-shaped in the adult male; labial furrows extending around the mouth corners, not long, the upper and lower furrows about equal; eye moderately large, its length more than one-half length of snout, subocular gutter extends under full length of eye but does not include spiracle, subocular gutter lined with denticles; spiracle somewhat larger than in most scyliorhinids; gill slits not long, the last two above the pectoral base. Origin of first dorsal fin over posterior end of pelvic fin base, origin of second dorsal fin over or slightly posterior to posterior end of anal fin base; second dorsal fin slightly greater in area than first dorsal, the distance between their bases greater than the head length; anal fin low, its base only a little longer than the second dorsal fin base; lower caudal fin low, without an anterior lobelike projection; pectoral fin very wide, its distal margin nearly straight; pelvic fins rather long and narrow (not projecting much laterally), not united along their inner margins. The teeth are in about 48 to 53 vertical rows in the upper jaw, 36 to 42 in the lower. Symphyseal teeth are not always well-differentiated from other teeth. About two to four series are functional. Most of the teeth have long central cusps with one or two much smaller cusps on each side of the main cusp. The largest upper jaw tooth in a 330-mm male was 0.9 mm high. Denticles of dorsolateral surfaces of adults are three-pointed but the middle point of each blade is much longer than the lateral points. The long denticle points overlap other denticles to some extent but skin is visible between most den- ticles. The largest on a 330-mm male were a little less than 0.5 mm long. Radiographs show 29 to 32 monospondylous vertebrae in 23 specimens except one count of 36 that apparently was due to the presence of 4 short (diplospondylous) vertebrae in the trunk where monospondylous vertebrae usually are present. 121 1 Total numbers of vertebrae, 132 to 145 in this series, may include incomplete counts due to poor resolution in radiographs of terminal caudal vertebrae. Measurements for the 328-mm holotype are given here expressed as percentages of the total length. Tip of snout to: front of mouth, 4.2; eye, 4.6; spiracle, 7.2; first gill slit, 13.1; fifth gill slit, 16.1; origin pectoral, 15.2; origin first dorsal fin, 38.3; origin pelvic fins, 30.4; origin second dorsal, 61.1; origin anal, 52.3; origin upper caudal fin, 80.5; anterior end cloacal opening, 31.0. Eyes: horizontal diameter orbit, 3.0; vertical distance from eye, 0.9. Spiracles: greatest diameter 0.7; least distance from eye, 0.9. Mouth: width, 5.8; length, 3.3; length upper labial furrow, 1.2; lower labial furrow, 1.2. Gill slits: height of first, 1.5; height of fifth, 0.5. First dorsal fin: length base, 4.3; anterior margin, 7.0. Second dorsal fin: length base, 5.5; length anterior margin, 8.8. Anal fin: length base, 7.3; length anterior margin, 5.5. Pectoral fins: width base, 3.6; length anterior margin, 9.4; width, 7.3. Distance between: posterior nasal apertures, 2.4; bases first and second dorsal fins, 20.2; bases pectoral and pelvic fins, 10.7. Clasper: length inner margin, 8.5; reach past pelvic fin tips, 2.5. Discussion. Schroederichthys maculatus is known only from the western Caribbean Sea at depths from 190 to 410 m, but the series available for study includes a better representation of medium and adult specimens of both sexes than is available for most scyliorhinids. Catch data suggest a habitat preference of sea bottom of fine, white, calcareous material. This is supported by radiographs of specimens showing nasal rosettes packed with mineral material and dis- sections show the valvular intestines contain much of the white, finely granular material. One stomach examined con- tained a 60-mm length of teleost vertebrae, one small squid or octopus beak, four eye lenses from an unidentified source, several pieces of algae, and several fish scales about 10 mm in diameter. Schroederichthys maculatus shows very little difference in body proportions between immature individuals 145 to 220 mm long and 280 to 342 mm adults. Schroederichthys tenuis Springer Figures 77, 78 Figure 77. — Schroederichthys tenuis, 230-mm immature male holotype, coast of Brazil, USNM 188052. Drawing by Mildred Carrington. Schroederichthys tenuis Springer 1966:606, figs. 16B, 18 (off mouth of Amazon River). Material examined. USNM 188052, imm. S, 230 mm, holotype, off mouth of Amazon River, lat. 01°49'N, long. 46°48'W, 410 m; USNM 188053, imm. 8, 180 mm, paratype, from same locality as holotype. Diagnosis. The narrow nasal flaps of S. tenuis (Fig. 78A) separate it from S. maculatus which has broadly triangular nasal flaps (Fig. 76A). The color pattern of S. tenuis (see Fig. 77) in the two immature specimens of the type-series dis- tinguishes it from the three other species of the genus as indicated in the preceding key. Gerhard Krefft has informed me (pers. commun.) of the capture of a 70-cm specimen of this species off the coast of Brazil. Evidently S. tenuis is a much larger species than S. maculatus, equal to or larger than S. bivius. Separation of the small S. tenuis, 180 mm, from small S. bivius, 155 mm, seems adequate on the basis of color pattern. In S. tenuis a pattern of small spots outlines saddle blotches whereas in S. bivius the saddle blotches are made up of solid color without bordering spots. Description. Comparison of measurements of the 220 and 180 mm S. tenuis with measurements of S. maculatus of equal length shows no significant differences. 122 Figure 78. — Schroederichthys tenuis, 230-mm immature male. A. Ven- tral view of head holotype, USNM 188052. B. Upper and lower jaw teeth, upper with cusps slightly constricted at base as in many juvenile scyli- orhinids. C. Dermal denticles of flank below first dorsal fin. The two specimens of S. tenuis had 32 and 34 monospondylous vertebrae and both had counts of 107 diplospondylous vertebrae. The teeth in