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A Generic Revision of the Cranchiidae (Cephalopoda; Oegopsida)

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A Generic Revision of the Cranchiidae (Cephalopoda; Oegopsida) BULLETIN OF MARINE SCIENCE, 30(2): 365-412, 1980 A GENERIC REVISION OF THE CRANCHIIDAE (CEPHALOPODA; OEGOPSIDA) Nancy A. Voss ABSTRACT This paper is the first in a monographic study of the cephalopod family Cranchiidae. Thirteen of the 41 nominal genera are recognized as valid for the family: Cranchia, Lio- cmnchia, Leachia, He/icocranchia, Bathothauma, Sanda/ops, Ligllriella, Taonius, Ga/iteu- this, Mesonychoteuthis, Egea, Megalocmllchia and Teuthowenia. A full definition is given for the family and each valid genus; diagnoses are given for the two subfamilies Cranchiinae and Taoniinae. A generic key is accompanied by illustrations of a representative species of each genus. A comprehensive historical resume for the family is given and generic interre- lationships are discussed. The subfamily Taoniinae comprises roughly three groups of genera based primarily on the development and relationships of the posterior end of the gladius (Ianceola and conus) and the fins and on the photophore pattern on the eye. He/icocrallchia, Bathothauma, Sandalops and Liguriella form a loose group of the more primitive genera that possess a short conus (lost in Bathothallma) and small subterminal or terminal fins. Taollills, Galitellthis and Me- sOllychoteuthis form a group of specialized genera in which the conus and associated terminal fins are elongate and the large suckers of the club are modified into hooks or suckers with one or two large, central, hook-like teeth. Of these, Ga/itellthis and Mesollychoteuthi.\· are closely related. In the third group formed of Egea, Megalocranchia and Teuthowenia, the fins elongate and simultaneously extend up the lateral margins of the mantle as the conus elongates. Egea and Megalocranchia are closely allied. The great variety of sexual dimorphism of the arms in the family is compared. All species of the relatively homogeneous Cranchiinae possess a hectocotylus in the male and brachial end-organs in females. In the more diverse Taoniinae, the males lack a distinct hectocotylus but have a high incidence of secondary sexual modifications of some arms and the females of only one group, Egea-Mega/ocranchia-Teuthowenia, possess brachial end-organs. The cranchiids are among the most numerous squids in the world oceans, both in number of species and in total number of catches (Roper, Gibbs and Aron, 1970; Clarke and Lu, 1974, 1975; Lu and Clarke, 1975a, b; Nesis, 1973, 1974b; Okutani, 1974; Yamamoto and Okutani, ]975; and unpublished data). They ex- hibit a high degree of morphological diversity. Most species undergo such great ontogenetic changes in morphology that various growth stages in a single species have regularly been and still are being described as separate genera and species (Nesis, ]972, 1974a; Lu and Clarke, 1974). K. N. Nesis (1972, 1974a) has recently revised several genera of cranchiids, but his work is not based upon examination of type material. Additional causes of confusion with the cranchiids are that few species are known from the adult stage and many have been described from only larval specimens. Another problem arises with the diverse forms of sexual dimorphism existing in the family. The need for a revision of the family Cranchiidae has long been obvious. Its successful accomplishment requires the study on a world-wide basis of large collections containing the various growth stages and the examination of the widely dispersed type material. I undertook this study several years ago. Since then I have acquired on loan the cranchiid collections from the Carlsberg Foundation-University of Copen- hagen, Oregon State University, the Newfoundland Biological Station, the South 365 366 BULLETIN OF MARINE SCIENCE. VOL. 30. NO.2. 1980 African Museum, the Australian Museum and the ANTON BRUUNIndian Ocean Expedition and the ELTANIN Antarctic collections from the Smithsonian Ocean- ographic Sorting Center. In addition, I have examined or have on loan the cran- chiid collections of Scripps Institution of Oceanography and of the National Mu- seum of Natural History, Smithsonian Institution (general collections , WALTHER HERWIG from South Atlantic, National Marine Fisheries Service from Central Pacific, University of Hawaii from Hawaiian waters). These collections together with the extensive collections at Miami cover most of the world oceans and contain a broad range of growth stages and a sizable proportion of near-mature and mature animals. In the spring of 1975, I visited the British Museum, the National Institute of Oceanographic Sciences at Wormley, England, the National Museum of Natural History in Paris, the Museum of Natural History in Nice, and the Institute of Oceanography in Monaco, where all cranchiid types and supplementary speci- mens were examined. I have also received types on loan from Berlin, Copen- hagen, Tokyo, Wellington, N.Z., and the U.S. National Museum of Natural His- tory, as well as critical specimens from Moscow. Thus, I have studied the major cranchiid collections of the world and have examined most of the type material known to be extant. The study is of such a scope that if the results were published as a single monographic work, it would be a number of years in preparation. In order to make the results more readily available, I will publish first this redefinition of the valid genera with a key for their determination and a historical resume for the family; a series of generic revisions will follow. A preliminary paper, "A Rede- scription of Egea inermis Joubin, 1933," has already appeared (N. Voss, 1974). There are 41 nominal genera in the family Cranchiidae: Cranchia Leach, 1817 He/icocranchia Massy, 1907 Leachia Lesueur, 1821 Liguriella IsseI, 1908 Perothis Rathke, 1835 Zygocranchia Hoyle, 1909 Owenia Prosch, 1849 Teuthawenia Chun, 1910 Taonius Steenstrup, 1861 Euzygaena Chun, 1910 Desmateuthis Verrill, 1881 Lellcocranchia Joubin, 1912 Procalistes Lankester, 1884 Phasmatoteuthian Pfeffer, 1912 Phasmatopsis Rochebrune, 1884 Verrilliteuthis Berry, 1916 Dyctydiopsis Rochebrune, 1884 Fusocranchia Joubin, 1920 Zygaenopsis Rochebrune, 1884 Ascoteuthis Berry, 1920 Pyrgopsis Rochebrune, 1884 Parateuthis Thiele, 1921 Megalocranchia Pfeffer, 1884 Anamalocranchia Robson, 1924 Liocranchia Pfeffer, 1884 Mesonychotellthis Robson, 1925 Cali/ell/his Joubin, 1898 Drechse/ia Joubin, 1931 Taonidium Pfeffer, 1900 Egea Joubin, 1933 Henseniotellthis Pfeffer, 1900 Belonella Lane, 1957 Corynomma Chun, 1906 Carynoteuthis G. Voss, 1960 Crystal/otellthis Chun, 1906 Ascocranchia G. Voss, 1962 Sandalops Chun, 1906 Vranatellthis Lu and Clarke, 1974 Toxeuma Chun, 1906 Vossotell/his Nesis, 1974 Ba/ho/hallma Chun, 1906 Each nominal genus is either stated to be valid, placed into synonymy, or considered dubious. The type species for each genus is given with original ref- erence, as well as place of deposit of the type and its present state of preservation if extant. A full definition is given for the family and each valid genus and a diagnosis is given for the subfamilies. The genera in the Taoniinae are arranged in the order of their apparent increased modificati;)H from the most primitive form. The key based on adult characters, together with the figures of a species of voss: GENERIC REVISION OF THE CRANCHIIDAE 367 each genus, will facilitate the determination of the genera. Though the identifi- cation of young stages is of secondary importance in the paper, larval and juvenile characters are included in the generic diagnoses, and figures of a larva and usually a juvenile of a species of each genus are given. The term "hectocotylus" is restricted in this paper to the extensive male sexual modification of a single arm presumably used in handling spermatophores. The lesser degree of male sexual modification found on arm pairs or on all the arms, which suggests a holding or caressing function in courtship and copulation, is referred to as "secondary sexual modification." The organ found on the tips of arm pairs of near-mature and mature female cranchiids is here called "brachial end-organ." The new names proposed by Bidder (1976) for digestive organs in coleoid cephalopods are used in this paper: "digestive gland" for the "liver," "digestive duct appendages" for the "pancreas," and "digestive duct" for the "hepatopan- creatic duct." The following abbreviations are used in figure captions for source or deposition of material illustrated: A-R/V ACONA, Department of Oceanography, Oregon State University; B-M/V BRANDAL,Newfoundland Biological Station; C-R/V A. T. CAMERON,Newfoundland Biological Station; CI-R/V COLUMBUSISELIN, University of Miami; CI-"Clarke" cruise, University of Hawaii deepsea biology project; D-R/V DANA, Denmark; Elt-USNS ELTANIN; F-"Fido" cruise, University of Hawaii deepsea biology project; GS-R/V JAMESM. GILLIS, Uni- versity of Miami; 10M-Institute of Oceanography, Monaco; MBI-Marine Biomedical Institute, University of Texas; O-M/V OREGON, National Marine Fisheries Service (formerly USFWS); OA-Ocean Acre Program, Smithsonian Institution; P-R/V PILLSBURY,University of Miami; SAM-South African Mu- seum; TM-R/V TONAN MARO, Japanese Exploratory Fisheries; UMML-Uni- versity of Miami, Rosenstiel School of Marine and Atmospheric Science; WH- FRS WALTHERHERWIG, Institut fur Seefischeri, Hamburg. HISTORICALRESUME The first cranchiid described was Cranchia scabra named by Leach in 1817 in honor of Mr. John Cranch, Collector of Objects of Natural History, on the Tuckey expedition to the river Zaire during which the unique specimen was captured off West Africa. The initial accompanying diagnosis "C. sacco tuberculato scabro; tuberculis duris scabriusculis" was expanded the next year when Leach (1818) stated, when diagnosing the genus, "neck with a frenum behind, connecting it with the sack, and with two other frena connecting it with the sack before." A second species from off West Africa, Cranchia maculata, based only on a mantle was originally placed in the genus by Leach but was soon disregarded by workers as being a doubtful form. Lesueur in 1821 described the next members of the family, Leachia cyclura from the Indian Ocean and Loligo pavo from the North Atlantic. The former species proved troublesome from the beginning because the description was based solely on a drawing.
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