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Systematics and Biology of (Pisces, Ophidioidea)

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Systematics and Biology of (Pisces, Ophidioidea) SYSTEMATICS AND BIOLOGY OF THE (PISCES, OPHIDIOIDEA) By JORGEN 6. NIELSEN Zoological Museum, University of Copenhagen CONTENTS I. Introduction ...................... 7 B. bruuni n. sp. ................... 51 Abbreviations.. .................... 8 B.parfaiti (Vaillant, 1888) ............ 53 Acknowledgments. .................. 8 B. specimen .................... 57 Methods and terminology. .............. 8 Meteoria n. gen. ................... 57 Material ......................... 9 M.erythrops n. sp.. ................ 58 Family diagnosis .................... 10 Sciadonus Garman, 1899 .............. 62 Relationships ...................... 10 S.pedicellaris Garman, 1899 ........... 63 ResumC of opinions .................. 11 LeucochIamys Zugmayer, 191 1 ........... 69 Genera.. ........................ 11 L. cryptophthalrnus Zugmayer, 191 1 ....... 70 Key to the genera of Aphyonidae .......... 12 L.jonassoni Nybelin, 1957 ............ 73 11. Systematic account. .................. 13 L.gaIatheae n. sp. ................. 75 Aphyonus Giinther, 1878 .............. 13 111. General remarks .................... 79 A.geIatinosus Giinther, 1878. .......... 15 Description ...................... 79 A. brevidorsalis n. sp.. ............... 20 Neoteny.. ...................... 79 Nybelinin n. gen. ................... 22 Gonads and reproduction.. ............ 81 N.erikssoni (Nybelin, 1957) ........... 23 Distribution.. .................... 82 Barathronus Goode & Bean, 1886 ......... 28 Biology ........................ 83 B. bicolor Goode & Bean, 1886 ......... 31 IV. Summary ........................ 84 B.diaphanus Brauer, 1906 ............ 44 V. References ........................ 86 B.afJinis Brauer, 1906 .............. 48 VI. Dansk resum&...................... 89 I. INTRODUCTION A systematic revision is presented including the de- The first described aphyonid fish, Aphyonzis gelatinosus Giinther, 1878, was caught during the Challenger Expedition in 1874. Since then, 64 ad- tion of the gonads, one of the few ways in which it ditional specimens of aphyonids have been caught. may be possible to learn something about the biol- Of these, 42 belong to one species and the remaining ogy of these rare, deep-living fishes. Consequently, to 12 species. Most aphyonids can thus be referred all specimens were opened, and the gonads and to as rare species, although many of them were other viscera studied. A histological examination found at unfrequently sampled deep pelagic depths. was made of the gonads, preferably of both a male JORDAN& EVERMANN(1898, p. 2499) were ap- and a female of each of the aphyonid species. parently the first to use the name Aphyoninae, as Osteological studies were based on radiographs only, they placed the then existing genera, Aphyonus and except for one species which was also cleared and Bauathvonus, in a separate subfamily under the stained. The holotypes of all described species were Brotulidae. The subfamily designation was appar- re-examined, as many of the old descriptions are ently overlooked by NYBELINwho later (1957, p. short and inadequate. 308) established a new subfamily, Aphyoninae, which included, i. a., the two genera of JORDAN& MENON,ZSIC, HANNIJSSEN, ZMA, ORVARNYBE- EVERMANN'Ssubfamily of the same name. LIN, NHMG, C. RICHARDROBINS, UMML, JONC. The first to propose the family name Aphyonidae STAIGER,UMML, G. TESTA, MOM, ALWYNE was ZUGMAYER(1911 a, p. 131), but not quite in the WHEELER,BM, and LORENP. WOODS,CNHM. OLE same sense as used in this paper. He mentioned that MUNK,Institute for Comparative Anatomy, Copen- if more genera besides Leucochlamys were found, hagen, sectioned the gonads, NIELSBONDE, Institute lacking the ventral fins and with the anus placed on for Paleontology, Copenhagen, gave vaIuabIe advice the posterior half of the body, they should be ar- on the osteology, and H.V.CHRISTENSENtook the ranged in a special family, the Aphyonidae. ZUG- photographs. Many of the drawings were made by MAYER'S proposal of this family-name seems strange JULIE TESCH and some by ELISABETHTBNDER. as on the same page he mentioned that Aphyonus AGNETEVOLSOE corrected the English manuscript. diKers from Leucochlamys by the presence of ventral Special thanks are due to DANIELM.COHEN for fins in the former, whereby Aphyonus per definition placing the large Barathronus bicolor material at my is excluded from the Aphyonidae (sensu ZUGMAYER).disposal and for making suggestions on the manu- script. Finally, I wish to thank ERIK BERTELSEN with whom I have had many fruitful discussions on Abbreviations deep sea fishes and who furthermore gave valuable advice during the preparation of this paper. BM - British Museurn, London. CNHM - Chicago Natural History Museum. FRSH - Fisheries Research Station, Hongkong. Methods and terminology - Institut fur Seefischerei, Hamburg. ISH Counts were made and measurements taken in MCZ - Museum of Comparative Zoology, Har- accordance with NUBBS& EAGLER(1958) except vard. that the upper jaw symphysis is used as the anterior- MNHN - Muskuin National d7KistoireNaturelle, most point and not the tip of the snout. Owing to Paris. the indistinctness of the eyes, measurements involv- MOM - MusCum Ocianographique, Monaco. ing the eyes are most often omitted. -The accuracy NHMG - Naturhistoriska Museet, Goteborg. of the measurements makes it reasonable to give UMML - University of Miami, Marine Labora- the morphometric characters with an absolute un- tory. certainty of 0.1 when the characters form less than USNM - United States National Museum, Wash- 10 % of the standard length (e.g., length of snout ington D. C. 8.4 % i0.1 of the SL), with an absolute uncertainty WHO1 - Woods Hole Oceanographic Institution. of 0.5 in the interval 10-50 % (e.g., length of post- - Zoologisch Museum, Amsterdam. ZMA orbital 16.5 % & 0.5 of the SL), and with an absolute - Zoologisches Museum, Berlin. ZMB uncertainty of 1in the interval 50-100 % (e.g., length ZMUC - Zoologisk Museum, Universitetet, K0- of preanal 56 % & 1 of the SL). - When references benhavn. to tables, plates and figures are spelled with a ZSIC - Zoological Survey of India, Calcutta. capital T, P and F they refer to the present work. SL - Standard length. other publications. - Quite often the depth indica- TL - Total length. tions seem very accurate, but this is usually due to conversion of fathoms into metres. - Except for Barathronus bicolor all species of the aphyonids are Acknowledgments represented by so few specimens that an osteological I am indebted to the following persons for provid- examination based on clearing and staining is ex- ing material or data: M.L.BAUCHOT,MNHN, cluded. Radiographs from a lateral as well as from WILLIAML. CHAN,FRSH, DANIELM. COHEN,Sys- a dorsal view were taken of all the specimens. Un- tematics Laboratory, U.S. Fish and Wildlife Ser- fortunately, many of the details are difficult to see vice, Washington D. C., KURT DECKERT,ZMB, on radiographs, especially when dealing with so RICHARDL. KAEDRICH,WHOI, BENGTHUBENDICK, poorly ossified fishes as the Aphyonidae. - In order NHMG, GERHARDKREFFT, ISH, ERNESTA. LACH- to examine, e.g., lateral line papillae, fin-rays and NER, USNM, GILESW. MEAD, MCZ, A.G.K. pseudobranchs, an air-jet was used. The air from an ordinary aquarium pump was led through a transversal plane. The m. infracarinales mediales pointed glass tube, making it possible to dry a appear as a chain of rectangular to square-shaped given area of the fish so that finer details could be muscular "fields". In the specific description the more easily distinguished. - The following method ratio between the length and the width of the was used when measuring the diameter of the eggs. middle muscular "fields" is normally given. The The ovary was divided into small portions and length is the cranio-caudal and the width is the placed under the microscope. By means of a draw- dorso-ventral measurement. ing-tube and 60 times magnification, the diameter of the eggs was drawn and subsequently measured Material on the paper. - The gonads were embedded in tissuemat, cut in 8 and 10 p. sections and stained Due to the kindness of many colleagues I have with Alcian blue (AB)-Kernechtrot, periodic acid been able to examine a11 aphyonid specimens that Schiff (PAS)-Ehrlich's hematoxylin and AB-PAS- have come to my knowledge. The collection com- Ehrlich's hematoxylin. prises a total of 65 specimens belonging to 13 species The following oceanographic terms are those and six genera. Below is listed all the species ex- proposed by BRUUN(1956 and 1957): Epipelagic, amined with indication of the number of recorded mesopelagic, bathypelagic, abyssopelagic, bathyal, and additional (old and new) specimens. For those and abyssal. - The term "viviparous" is used for all specimens, of which the gonads have been examined species in which the eggs are fertilized within the 'histologically, the histological serial number (Ser. female even if a minor part of the development of No.) is included : the embryo takes place outside the female. About half of the more than 200 ophidioid species are Aphyonus gelatinosus Gunther, 1878, including A. referred to viviparous groups, owing to the presence mollis Garman, 1899. Three specimens of which of a copulatory apparatus and/or to the presence of one is new. Ser. Nos. 17 (9) and 64 (3). fertilized eggs in the ovaries. Only very few species, Aphyonus brevidorsalis n. sp. Only the holotype is i.a., two aphyonids, are actually known to bring known. Ser. No. 4 (8). forth free larvae. Like most other authors I do not
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