A Generic Revision of the Cranchiidae (Cephalopoda; Oegopsida)

Home , Cranchiidae, Cranchiinae, Leachia, Taoniinae, Taonius, Teuthowenia

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/iteuthis, 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 exhibit 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 cranchiid 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 specimens 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 reference, 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 identification 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. In 1833 Grant gave the first detailed illustrations of a cranchiid with his description of Loligopsis guttata from the Indian Ocean. The char- acteristic mantle fusions of the family were clearly described by Eschscholtz in an unpublished personal journal apparently written on the first round-the-world cruise of the Russian exploratory vessel RURICKon which Eschscholtz served as ship's physician. These observations were published by Rathke in 1835 in the most accurate anatomical description of a cranchiid to that time. Rathke named the squid collected in the Indian Ocean Perothis eschscholtzii. The next year 368 BULLETIN OF MARINE SCIENCE. VOL. 30. NO.2. ]980

Owen (1836) described the fusions in the first detailed description of Cranchia scabra and made the initial attempt to place the earlier-described cranchiids into the systematic order of cephalopods. Using the common feature of the fusions in the then recognized genera Cranchia and Loligopsis, he separated them into an unnamed section in his large conglomerate family Teuthidae. D'Orbigny (1839) placed the two genera in widely separate families on the basis of his mistaken concept that Cranchia had a skin covering the eyes. Cranchia was put in the myopsid family Sepidae and Loligopsis in the oegopsid family Loligopsidae along with Chiroteuthis and Histioteuthis. In 1848 Adams and Reeve added another new species Loligopsis ellipsoptera, collected by HMS SAMARANGin the North Atlantic. Prosch (1849) first named a systematic group distinguished by the basic common character of the mantle fusions. Prosch separated out from Owen's family Teuthidae a new subfamily Cranchidae comprised of the single genus Cranchia with the subgenera Cranchia and his new Owenia. Important discrepancies in Prosch's work on his new species Owenia megalops were pointed out by March (1850). Gray (1849) also recognized the importance of the internal fusions but retained the mistaken character of the covered eyes in Cranchia. He united the two monotypic families Cranchiadae and Loligopsidae in an unnamed section, and restricted the Loligopsidae by creating a new family Chiroteuthidae for Chiroteuthis and Histioteuthis. Finally Steenstrup in 1861 "united under the name of the Cranchia family all cephalopod forms which agree with the genus Cranchia in that they have the mantle firmly connected with the head at three different points .... " He named the new family Cranchiaformes. Steenstrup pointed out d'Orbigny's error in attributing a "myopsid" eye to Cranchia and in his use of the generic name Lo/i- gopsis. Steenstrup resurrected Lesueur's Leachia for the "Loligopsis" forms, and proposed a new genus Taonius to contain his new species hyperboreus from the North Atlantic and Lesueur's pavo. He also expanded Prosch's concept of Cranchia to contain his own earlier-described species from the North Atlantic Leachia reinhardtii Steenstrup, 1856 as well as scabra and megalops. Unfor- tunately he denied the existence of a funnel valve in Cranchia and gave rise to the misconception continued by some workers that a valve was lacking in the family. Though Steenstrup's work was a tremendous stride forward, it seems to have been largely ignored by succeeding early workers. Tryon's Manual of Conchology (1879) included Steenstrup's new species, but used the old collective genus Loligopsis, uniting it with Cranchia (in which he reconfirmed the presence of a valve) in the Cranchiidae, emending the name of the family to that which we use today. Though Verrill (1881) accepted Steenstrup's Taonius for pavo, he referred to d'Orbigny's classification when he united his Desmoteuthidae with the oegopsid group Taonidea along with the allied forms formerly included in Loligopsis. Des- moteuthidae was created for Taonius and his Desmoteuthis which was based on a specimen that he erroneously considered identical with Steenstrup's Taonius hyperboreus. Rochebrune (1884) greatly expanded d'Orbigny's old family Loligopsidae including in it his four new genera, Phasmatopsis, Dyctydiopsis, Zygaenopsis and Pyrgopsis and three new species of cranchiids from the Atlantic and southern Indian Ocean. His confused, inaccurate, and generally poor systematic work was much criticized by Hoyle in his 1885 paper "On Loligopsis and some other genera. " Here Hoyle reassigned to their rightful genera the species formerly voss: GENERIC REVISION OF THE CRANCHIIDAE 369 erroneously referred to Loligopsis and, among other things, recognized Verrill's earlier mentioned error in identification and synonymized Desmoteuthis with Taonius. Hoyle valued Steenstrup's work and in his Challenger Report (1886) utilized Steenstrup's conception of the family Cranchiaeformes. Pfeffer (1884) described several new species of cranchiids based upon smaIl specimens in the collections of the Hamburg Museum and added two new genera, Megalocranchia and Liocranchia. In his "Synopsis der oegopsiden Cephalopo- den," Pfeffer (1900) reviewed the family, defined genera, and gave the first key to their identification. He errected two new genera, Taonidium for Hoyle's (1886) species Taonius suhmi from south of Australia and Hensenioteuthis for his species jOllbini from the North Atlantic. Though Pfeffer, too, placed Verrill's Desmoteuthis hyperborea in the synonymy of Taonius pavo, he retained the generic name Desmoteuthis for Steenstrup's Taonius hyperboreus and added Goodrich's (1896) Taonius abyssicola from the Laccadive Sea and his own earlier Megalocranchia maxima from off the Cape of Good Hope, thereby synonymizing the genus Megalocranchia. In all, he recognized nine genera-Taonius, Des- moteuthis, Taonidium, Owenia, Hensenioteuthis, Zygaenopsis, Leachia, Lio- cranchia and Cranchia. Pfeffer used Tryon's emended name Cranchiidae for the family and it has been used ever since by most cephalopod workers. The first detailed description of the stalked eye of a larval cranchiid was published by Chun (1903a), soon to be followed by the first detailed description of the photophores on the eye of a cranchiid by Joubin (1905) based on several adults of a species of Leachia found dying at the surface in the vicinity of the Azores and Canary Islands. Chun was later (1910) to greatly expand our knowledge of photophores in the family. After studying the rich coIlections of known and new cranchiids made by the VALDIVIA1898-1899, Chun (1906) more sharply defined the family in an expanded key to the genera. He delineated \3 genera, which included all those recognized by Pfeffer in 1900, except the poorly defined Hensenioteuthis, and added five new genera, Corynomma, Crystalloteuthis, Sandalops, Toxellma and Bathothau- ma, all based on very young stages. He described Liocranchia valdiviae from the Indian Ocean and recognized that Joubin's GaLiteuthis represented the adult form of Taonidium. Joubin (1898) had erected a new family, Cranchionychiae, for his Galitellthis armata from the Mediterranean because of the presence of hooks on the clubs. He considered it to be intermediate between Steenstrup's families Onychii and Cranchiaeformes. Chun (1910) gave a comprehensive treatment of the family in his monograph of the oegosid cephalopods coIlected by the German Deep-Sea Expedition in which he compared and evaluated internal and external characters in detail and described ontogenetic morphological changes. He expanded the descriptions of, and illustrated, the new species from his 1906 work, proposed Teuthowenia for the preoccupied Gwenia and added a new species from the Antarctic, antarctica, to the genus. He also proposed Euzygaena for the preoccupied Zygaenopsis and added a new species from the southeastern Atlantic, pellucida, to Desmoteuthis. In addition, he gave full recognition to Galiteuthis, thereby synonymizing Taonid- ium. It is surprising that despite his detailed work, Chun continued to miss the presence of a funnel valve in Cranchia, Liocranchia and Corynomma and considered its absence an important familial character. This feature, together with the unique dorsal and ventral fusions of the mantle, justified to Chun the division of the oegopsids into the suborder Oegopsida consulta, containing the Cranchi- idae, and the suborder Oegopsida libera, containing all of the others. From 1907 to 1912, new genera and species of cranchiids were described by 370 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

several workers. Massy in 1907 well described a new genus and species Helico- cranchia pJeJJeri taken off the coast of Ireland. And in the following year, Issei (1908) described from the collections of the circumnavigation of the Italian ship LIGURIA (1903-1905) three new species and a new genus Liguriella, the latter based on a distorted 9 mm ML larva from the South Atlantic. A year later, Russell (1909) created another new species based on a larva, Taonidium pJeJJeri, from the Shetland Islands. Berry (1909, 1911) published three new species from the Pacific; and a year later created a new subfamily Galiteuthinae to contain Joubin's Galiteuthis (Berry, 1912a). In another attempt to untangle the confusion caused by the erroneous use of Verrill's Desmoteuthis, Berry (1912b) proposed that Pfeffer's Megalocranchia be used to contain Steenstrup's hyperboreus and the other species assigned to Desmoteuthis by Chun (1910), with the exception of Helicocranchia pJeJJeri. Unlike Chun, Berry recognized the uniqueness of Mas- sy's species after having examined an undescribed but similar form taken by the ALBATROSSin Hawaiian waters. Joubin (1912) added a new genus Leucocranchia and three new species from the northeastern Atlantic to the literature, all inadequately characterized. In his monographic review of the oegopsid cephalopods, Pfeffer (1912) stressed the importance in the cranchiids of the form of the fins and the posterior end of the gladius and their relationships to each other. He separated the Cranchiidae into two subfamilies, the Cranchiinae and the Taoniinae, each composed of three tribes. In the subfamily Cranchiinae he distinguished the tribes: Leachia-like Cranchiinae consisting of Leachia and Pyrgopsis, Cranchia-like Cranchiinae consisting of Liocranchia and Cranchia and Liguriella-like Cranchiinae consisting of Liguriella. In the subfamily Taoniinae Pfeffer distinguished the tribes: Taonius- like Taoniinae consisting of Taonius, Galiteuthis, Desmoteuthis, Megalocran- chia, Phasmatopsis, Phasmatoteuthion, Toxeuma, Crystalloteuthis. Corynomma and Taonidium; Teuthowenia-like Taoniinae consisting of Teuthowenia, further divided into four subgenera-Teuthowenia, Hensenioteuthis, Sandalops and He- licocranchia; and Bathothauma-like Taoniinae consisting of Bathothauma. Pfef- fer resurrected Rochebrune's Pyrgopsis for the early stalk-eyed forms of Leachia and it has been commonly so used ever since. He included in his Megalocranchia the same species assigned to it by Berry (1912) with the exception of Steenstrup's hyperboreus for which he persisted in retaining Desmoteuthis. Two new species were described from the North Atlantic, again based on larval types, and the genus Phasmatoteuthion was erected for Joubin's (1895) Taonius richardi inaccurately described from a distorted incomplete juvenile specimen from the North Atlantic. Over the next few years little was added to the' literature of the family except for Berry's (1916) Megalocranchia pardus from the Kermadec Islands. In a foot- note in the same paper Berry proposed a new genus Verrilliteuthis for Desmo- teuthis tenera Verrill which Berry considered improperly grouped with the Meg- alocranchia forms and without a valid generic name since he had shown in his 1912 paper that Desmoteuthis fell into the synonymy of Taonius. Verrilliteuthis was used by a number of succeeding workers for Steenstrup's hyperboreus into whose synonymy Verrill's species was relegated. A number of new genera and species of cranchiids were described in 1920. Joubin created a new genus and species Fusocranchia alpha for a larva from the northeast Atlantic; Sasaki named a species Crystalloteuthis beringiana from a young specimen taken in the Aleutian Islands; and finally, Berry (1920a) added a subgenus of Teuthowenia, Ascoteuthis, and four new species all based on larval material from the western Atlantic. In a separate paper, reviewing the occurrence voss: GENERIC REVISION OF THE CRANCHIIDAE 371

of photophores in the cephalopods, Berry (l920b) divided the oegopsids into five superfamilies, placing the Cranchiidae as the sole member of the superfamily Cranchioidea. He recognized the family to be composed of 18 genera and 41 species. The following year, Liocranchia gardineri from the Indian Ocean was described by Robson (1921). In his systematic survey of European cephalopods, Grimpe (1922) created a new subfamily Teuthoweniinae for Teuthowenia (+Hen- senioteuthis and Helicocranchia) and Sandalops, and placed Bathothauma in a new family Bathothaumatidae. He reshuffled the oegopsids into six new family- groups, uniting his Bathothaumatidae, along with the Cranchiidae, in the family- group Cranchiina, thereby replacing Berry's recently proposed superfamily. Grimpe's concepts of classification were generally disregarded by following workers. In his "Die Cephalopoden" (1923) Naef rejected Chun's proposed isolation of the Cranchiidae in a separate group "Oegopsida consulta" from the other oegopsids. Studying the post-embryological development of several species of cranchiids, he attempted to show the similarity between the earliest young stages of the Cranchiidae and those of other oegopsids. The family was expanded by new records over the next 10 years, and the addition of four genera and nine species including Liocranchia intermedia and Anomalocranchia impennis Robson, 1924 from South African waters, Mesony- choteuthis hamiltoni Robson, 1925 from the stomach of a sperm whale from the South Shetland Islands, and Teuthowenia tagoi and Teuthowenia elongata Sa- saki, 1929 from Japanese waters. During this period the vast cephalopod collections of the cruises of the DANA yielded the new forms Pyrgopsis atlantica and Desmoteuthis thori Degner, 1925 from the North Atlantic, Drechselia danae Joubin, 1931 from the eastern tropical Pacific and Egea inermis Joubin, 1933 from the North Atlantic. Numerous new records greatly broadened our knowledge of the distribution of the cranchiids during the 1940s and 1950s. In addition, the species Taonidium pacificum and Helicocranchia beebei were described by Robson (1948) from collections made by the Arcturus Oceanographic Expeditio'n to the Eastern Pa- cific, and Dell (1959) described Megalocranchia richardsoni from New Zealand waters. Studying a large collection containing a wide range of sizes collected by the DANA, Muus (1956) showed that Teuthowenia megalops (Prosch, 1849) is the early stage in the development of Steenstrup's (1861) species hyperboreus, but unfortunately settled on Desmoteuthis for megalops. His choice of a generic name and his mistaken placement into the synonymy of megalops of several diverse species, particularly Helicocranchia pJeJJeri, was soon criticized (G. Voss, 1960; Adam, 1962) but Muus reaffirmed his decisions in a later paper (Muus, 1962). In 1957 Lane proposed Belonella for the preoccupied Toxeuma. The number of cranchiids available for study increased many fold during the next 20 years due to the surge in marine research, new oceanographic ships, and new and better sampling techniques. G. Voss (1960, 1962, 1963) described two new genera and four new species from the North Atlantic: Megalocranchia papillata, Carynoteuthis oceanica, Ascocranchia joubini, and Phasmatopsis lucifer. Clarke (1962a) in a paper describing in detail a large mature female cranchiid taken dead at the surface near Madeira mistakenly identified it to Rochebrune's (1884) Phasmatopsis cymoctypus. He redefined Phasmatopsis from his misidentified specimen and synonymized Carynoteuthis. Clarke then attempted to expand Taonius to Steenstrup's original scope to include Prosch's megalops (= hyperboreus Steenstrup) with Taonius pavo, disregarding the marked differences of generic level between the two species. In "A Review of the Systematics and 372 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

Ecology of Oceanic Squids" Clarke (1966) tentatively regarded the Cranchiidae to be composed of 24 genera and 51 species, recognizing the fact that some genera would probably prove to be developmental stages of others. His error in including Crystalloteuthis, Ascocranchia and Egea in the subfamily Cranchiinae was cor- rected by Roper, R. Young and G. Voss (1969). There followed an overall increased stress on the morphological changes accompanying growth. McSweeny (1970) described the juvenile form of the giant Mesonychoteuthis hami/toni. Growth stages of several species were described in various works, including Ga/iteuthis phyllura, G. pacifica and a new species Leachia dis/ocata from off southern California by R. Young (1972), Bathot/wuma /yromma by Aldred (1974) and Okutani (1975), Egea inermis by N. Voss (1974) in which Phasmatopsis lucifer was synonymized as the adult stage, Sanda/ops me/ancho/icus and Taonius pavo by R. Young (1975a), TeutholVenia mega/ops by Dilly and Nixon (1976a) and Galiteuthis g/acia/is by McSweeney (1978). Changes in eye shapes with growth were related to the vertical distribution of the growth stages by R. Young (1975a). The different patterns of vertical distribution in the family were presented by Roper and R. Young (1975). The increased availability of cranchiids sparked added interest in behavior, flotation physiology, luminescence, morphology and function, and other aspects of biology exemplified in the works of Clarke (1962b), Denton, Gilpin-Brown and Shaw (1969), J. Young (1970), Dilly and Herring (1974), R. Young (1975b) and Dilly and Nixon (1976b). Working with collections from recent wide-ranging cruises of Russian research vessels, Nesis (1972, 1974a) revised severa] genera of cranchiids: Taonius, Be/- onella, Corynomma, Mega/ocranchia. Sanda/ops and Liguriella. He discussed evolutionary lines and conceived the subfamily Taoniinae to include 14 genera: Taonius, Phasmatopsis, Re/onel/a, Ga/iteuthis, Mesonychoteuthis, Corynomma, Verri/liteuthis, Sanda/ops, Liguriella, He/icocranchia, Ascocranchia, Egea, Bathothauma and his new genus Vossoteuthis. Unfortunately Nesis was ham- pered by the lack of examination of types and crucial previously published upon specimens, as well as incomplete developmental series and the lack of large collections of comparative world-wide material. Lack of knowledge of the ontogenic changes in morphology with growth continued to result in the naming of new taxa for already known species: Ga/iteuthis aspera Filippova, 1972 (=G. g/acia/is), Uranoteuthis bi/ucijera Lu and Clarke, ]974 (=Sanda/ops me/ancho/icus) and Ga/iteuthis tri/uminosa Lu and Clarke, 1974 (= Taonius pavo). Numerous reports from every ocean except the Arctic widened the known distribution of the family. Nesis (1973, 1974b), Clarke and Lu (1974, 1975), Lu and Clarke (1975a, b), Okutani (1974) and Yamamoto and Okutani (1975) showed cranchiids to be the first or second most speciose family and the first or second most numerous in total catch of oceanic squid taken in the Atlantic, Pacific and Antarctic. The obvious part that cranchiids play in the food web in the ocean is of growing interest. That they form a very significant part of the diet of many oceanic animals is revealed in recent work with sperm whales (Clarke and MacLeod, 1976), blue sharks (Clarke and Stevens, ]974) and certain seabirds (Imber, 1973).

SYSTEMATIC SECTION Family Cranchiidae Prosch, 1849

Cranchidae Prosch, 1849: 7\. Cranchiadae Gray, 1849: 36. voss: GENERIC REVISION OF THE CRANCHIIOAE 373

Loligopsidae Gray, 1849: 36. Cranchiaeformes Steenstrup, 1861: 70. Cranchiidae Tryon, 1879: 102. Desmoteuthidae Verrill, 1881: 300. Cranchionychiae Joubin, 1898: 291. Bathothaumatidae Grimpe. 1922: 51.

Definition.-Small to very large sized oegopsids; mantle fused to head at nuchal region and to funnel at its posterolateral corners; collar muscle fused to mantle dorsally along two obtusely diverging lines from nuchal point of fusion and ventrally along a dorsally acutely diverging line from each point of fusion at posterolateral corners of funnel; ventrally diverging lines from posterolateral corners of funnel formed by fusion of ventrolateral margins of funnel to mantle; funnel depressor muscles expanded into broad membrane that inserts laterally onto mantle, thereby dividing mantle cavity into one large, ventral chamber and two dor- solateral chambers with a connecting spiracle medial to each gill; mantle stout to elongate, spindle-, conical- or cylindrical-shaped with posterior end varying from rounded (Bathothauma) to filiform (Taonius); mantle wall thin but usually muscular, outside surface sometimes covered with tubercles, from larval stage to adults (Cranchia) or only in subadults and adults (some species of Galiteuthis); the lanceola and midline of rachis and conus usually visible dorsally through skin; entire body flaccid in gravid females; fins thin (exceptions-very muscular in Mesonychoteuthis), vary greatly from a) widely separate, small, paddle-shaped and subterminal, to b) medium to large, rounded terminal, to c) ovate or lanceolate terminal or terminal-lateral; fins insert laterally on lanceola and conus (and rostrum in Helicocranchia), and sometimes extend onto lateral margins of mantle (Megalocranchia, Egea and Teuthowenia);funnel moderately large to very large, with or without valve; dorsal pad offunnel organ inverted V- or V-shaped, usually with two or three papillae or flaps or both (in Leachia, up to seven papillae present); two small to very large ventral pads of varying shapes (oval, kidney, crescent, L-shaped, etc.); head short; eyes large (sometimes small to medium), sessile, prominent or protruding, usually anteriorly or anterolaterally oriented, with photophores on ventral surface (occasionally extending to dorsal surface); olfactory organ short-stalked or semi-lunar papilla on posteroventral surface of head; buccal membrane with seven lappets and eight connectives, attachment formula DDVV; arms short to medium length, muscular, with trabeculate protective membrane developed on both margins of all arms (trabeculae very weakly developed to absent in Helicocranchia); wide-aperture suckers in two rows, often with varying degrees of enlargement with growth and sometimes associated with sexual dimorphism (enlarged midarm suckers modified into hooded hooks in Me- sonychoteuthis); arms III usually longest, sometimes markedly so; either right or left IV in males modified into hectocotylus or hectocotylus absent; secondary male modifications (usually elongate, attenuate ends of arms with abruptly reduced-sized suckers in two or more rows) often present on arm pairs (some or all arms in adult males sometimes more robust and elongate); near-mature and mature females sometimes with ends of one to four pairs of arms modified into brachial end-organ (possibly a photophore); tentacles short to long, stout to slender, with two to four rows of carpal suckers alternating with pads on major portion of stalk; carpal cluster or lesser-defined carpal group sometimes present at base of short to long, slightly expanded or unexpanded club with short dactylus; suckers on manus and dactylus in four rows display varying degrees of enlargement, with proximal or all suckers in median two rows of manus modified with 374 BULLETIN OF MARINE SCIENCE, VOL. 30. NO.2, 1980

growth into hooded hooks in Galiteuthis and Mesonychoteuthis; gills short; digestive gland stout to slender, spindle-shaped, oriented almost perpendicular to longitudinal axis of body in young, becomes more longitudinal in adults; ink sac usually small, partially embedded in median posteroventral surface of digestive gland (in Megalocranchia digestive gland with growth becomes rounded with large, bilobed, compound photophore overlying ink sac and ventral surface of digestive gland); paired digestive duct appendages situated on posterodorsal surface of digestive gland, on digestive duct or ducts or on caecum; two digestive ducts fused to form medium to long common duct (not fused in Megalocranchia); male genital organs single, situated on left side; female genital organs paired; with maturity, nidamental glands enlarge coequally with oviducal glands; gladius with long, narrow rachis with dorsal keel absent to weakly developed anteriorly (strongly developed in Leachia), bordered posteriorly by short to long vanes that extend to form a long, spindle-shaped to broad, diamond-shaped, weakly to strongly keeled lanceola with short, broad to long, filiform, usually hollow, conus (exceptions-vanes are very short and expand to form short conus with dorsal rostrum in Helicocranchia; vanes modified into transverse bar with expanded, shovel-shaped ends with no conus in Bathothauma). Type genus.-Cranchia Leach, 1817. Early larval form.-Larva with mantle-head and mantle-funnel fusions; mantle elongate with thin walls (occasionally outer mucous coat surrounds mantle and head); fins separate, paddle-shaped (wider than long with narrow base); head with short to long arm pillar (no arm pillar in Cranchia and Liocranchia); eyes oval, usually with ventral rostrum, stalked (unstalked in Cranchia and Liocranchia); arms short; tentacles relatively long with suckers on entire length; gladial conus short. Distribution.-Epipelagic to upper bathypelagic zones with most genera restricted to circumglobal tropical and subtropical waters; Teuthowenia, Galiteuthis, Taoni- us, Liguriella and Bathothauma extend to subarctic and/or subantarctic waters (Galiteuthis extends farthest into Antarctic waters); Mesonychoteuthis is restricted to Antarctic waters.

KEY TO THE GENERA OF THE CRANCHIIDAE (ADULTS) la. Ventral surface of mantle with one or two cartilaginous strips extending posteriorly from anterior apex of funnel-mantle fusions. Funnel fused to head laterally. Eyes with four or more small, round to oval photophores (Subfamily Cranchiinae) 2 lb. Ventral surface of mantle without cartilaginous strips extending posteriorly from anterior apex of funnel-mantle fusions. Funnel free from head lat~rally. Eyes with one usually large photophore, or two to three markedly dissimilar-sized photophores with the largest usually crescent-shaped (Subfami Iy Taoniinae) .. _ 4 2a. Ventral surface of mantle with two cartilaginous strips in inverted V-shaped pattern extending posteriorly from anterior apex of funnel-mantle fusions. Funnel valve present. Dorsal pad of funnel organ with three longitudinal, triangular flaps. Gladius with short conus . ..._ 3 2b. Ventral surface of mantle with one cartilaginous strip extending posteriorly from anterior apex of funnel-mantle fusions. Funnel valve absent. Dorsal pad of funnel organ with three to seven narrow papillae. Gladius with long slender conus Leachia (p. 380, fig. 3) 3a. Mantle covered with cartilaginous tubercles. Eyes with fourteen photophores. Brachial end- organ present on all arms in near-mature and mature females. Suckers in four rows on midportion of hectocotylized right IV in males Cranchia (p, 376, fig. I) 3b. Mantle without cartilaginous tubercles except on ventral cartilaginous strips and sometimes on dorsal median line. Eyes with four or fourteen photophores. Brachial end-organ present only on arms III in near-mature and mature females. Suckers in two rows on mid portion of hectocotylized right or left IV in males Liocranchia (p. 378, fig. 2) voss: GENERIC REVISION OF THE CRANCHIlDAE 375

4a. Fins small, paddle-shaped, subterminal. Eyes with one, usually large, photophore______5 4b. Fins not paddle-shaped, may be short to long, round to lanceola-shaped, terminal or terminal lateral. Eyes with one large and one or two small photophores .______6 5a. Fins fused distally, insert on short rostrum of gladius which projects dorsally free of end of mantle. Eyes small to medium He/icocranchia (p. 382, fig. 4) 5b. Fins widely separated, insert on"lateral expanded ends of transverse extensions of posterior end of gladius. Eyes large to huge _. Bathothauma (p. 384, fig. 5) 6a. Gladial conus short, broad to narrow. Fins short «25% ML), oval to round. Digestive gland long, narrow, spindle-shaped 7 6b. Gladial conus medium to long, narrow, or needle-like to filiform. Fins medium to long (30- 60% ML), narrow, lanceolate to ovate. Digestive gland stout, spindle-shaped, or rounded 8 7a. Posterior fin insertions do not extend to tip of gladius. No tubercles present on funnel-mantle fusion cartilages. Dorsal pad of funnel organ with large, triangular lobe on each lateral arm. Eyes with small, round, anterior photophore indented into median anterior margin of large, round, posterior photophore Sandalops (p. 386, fig. 6) 7b. Posterior fin insertions extend to tip of gladius. Two small tubercles present at anterior end of funnel-mantle fusion cartilages. Dorsal pad of funnel organ with large, spatulate papilla on each lateral arm. Eyes with small, crescent-shaped, anterior photophore lying closely within concavity of large, crescent-shaped, posterior photophore Liguriel/a (p. 388, fig. 7) 8a. Anterior fin insertions on lateral margins of lanceola of gladius .______9 8b. Anterior fin insertions on lateral margins of mantle 11 9a. Tentacular club with hooded hooks, marginal suckers of manus greatly reduced in size or lost. Arms IV the longest in juveniles, approximately coequal to III in adults. Gladial conus narrow or needle-I ike __ _ __ 10 9b. Tentacular club without hood hooks (enlarged suckers with one or two large, central, hook- like teeth on distal margin), marginal suckers of manus not greatly reduced in size or lost. Arms III the longest in juveniles, the longest or approximately coequal to II in adults. Gladial conus filiform ~ Taonius (p. 390, fig. 8) lOa. Arms without hooded hooks. Fins lanceolate becoming attenuate posteriorly ______Ga/iteuthis (p. 392, fig. 9) lOb. Arms with hooded hooks on mid portion. Fins stout, ovate (nearly round in combined outline in juvenile), do not become attenuate posteriorly Mesonychoteuthis (p. 394, fig. 10) Ila. Funnel valve present. Dorsal pad of funnel organ with triangular flap on each lateral arm. Eyes with two photophores (large, roughly crescent-shaped posterior photophore, and within its concavity, smaller, roughly elongate S-shaped anterior photophore). Carpal suckers in two rows on tentacular stalk '12 lib. Funnel valve absent. Dorsal pad of funnel organ with long, spatulate papilla on each lateral arm. Eyes with three photophores (large, crescent-shaped posterior photophore, and within its concavity, smaller, crescent-shaped anterior photophore and third small, oval photophore). Carpal suckers in four rows set in zigzag pattern on tentacular stalk ______.______Teut howenia (p. 400, fig. 13) 12a. Photophores absent on ventral surface of stout, spindle-shaped, digestive gland. Long, single, united digestive duct Egea (p. 396, fig. II) 12b. Large compound photophore present on ventral surface of rounded digestive gland. Two short digestive ducts remain separate Megalocranchia (p. 398, fig. 12)

Subfamily Cranchiinae Pfeffer, 1912

Cranchiinae Pfeffer, 1912: 644. Diagnosis.- Ventral surface of mantle with one or two cartilaginous strips bearing tubercles for entire length (except in Cranchia) extending from anterior apex of funnel-mantle fusions along one or both lines of fusion; fins small to large, terminal, elliptical, to nearly circular in combined outline, with anterior insertions confined to posterolateral margins of lanceola; funnel fused to head laterally; eyes with four or more small round to oval photophores; in males, right or left IV modified into hectocotylus, secondary sexual modification of ends of arms may be present on III, or absent; in near-mature and mature females, modification of 376 BULLETIN OF MARINE SCIENCE. VOL. 30, NO.2. 1980

arm tips into brachial end-organ present on III or on all arm pairs; caecum larger than stomach.

Type genus.-Cranchia Leach, ]8]7.

Cranchia Leach, ]8]7 (Figure I)

Cranchia Leach, 1817: 140. Type species, by subsequent designation, Steenstrup, 186]: 72 (27, Eng. trans.): Cranchia scabra Leach, ]8]7: 140, Type specimen deposited, British Museum (Nat. Hist.). Examined; fair condition. Definition.-Medium-sized cranchiids; stout, spindle-shaped mantle tapers posteriorly to broad point (in larva, mantle often contracted to near-round), entire surface covered with closely set, variable-sized cartilaginous tubercles usually 3 to 5 sided with sharp multipoints at each angle; mantle walls thin, leathery; two short, smooth, cartilaginous strips of coequal length diverge from apex of funnel- mantle fusions in inverted V-shaped pattern with large, multipoint tubercle complex at apex; long, narrow, spindle-shaped nuchal fusion cartilage without tubercles; small, nearly circular fins project and unite with median cleft posterior to end of gladius, dorsal surface except for outer margins covered with cartilaginous tubercles, ventral surface papillated; funnel large, broad, extending nearly to base of arms, surface papillated; large, inverted V-shaped dorsal pad of funnel organ with three longitudinal, broad, triangular flaps, a large one extending medially full length of each lateral arm and continuous with anterior margin of pad, and a smaller one on median line of posterior part of anterior midportion; very large, stout, kidney- or crescent-shaped ventral pads; large funnel valve present; head with large, round, prominent, sessile eyes occupying entire lateral surface; small knob-shaped olfactory organ present on posteroventral surface; overall surface papillated; eyes with fourteen oval photophores (seven in outer half circle on ventral surface; one midway on posteroventral surface; six in inner full circle, four ventral and two dorsal) (in larva, head often withdrawn into mantle; eyes oval with short ventral rostrum, sessile; arm pillar lacking); arms short to medium length, narrow, muscular, with length formula III > II = IV> I; moderately developed web connects arms I-III (inner web between arms I and I-II, outer web between II and III);aboral keel well developed on III, poorly developed on II and I; lateral keel well developed on IV, trabeculate protective membrane well developed on both margins of all arms; medium to large suckers largest on midportions of all arms (except the hectocotylus); sucker rings smooth to toothed with small flat, round, square or pointed teeth on distal margin; in males, right IV modified into hectocotylus (arm enlarged, curved laterally with two, then four, rows of small suckers followed terminally by two ventral rows of enlarged, elongate suckers), ends of III secondarily modified (four rows of minute suckers); in near-mature and mature females, brachial end-organ present on all arms; short, stout tentacles with slightly expanded 'c1ubs; diagonally set pairs of carpal suckers alternate with pads on distal two-thirds of stalk; carpal group present, but not distinct carpal cluster; suckers of club largest on midmanus, with central suckers only slightly larger than marginals; suckers abruptly reduced in size on dactylus; rings of manus suckers with numerous sharp conical teeth around entire margin; club with well developed trabeculate protective membrane on both margins and well developed dorsal keel on distal two-thirds (in early larva, only short, stout tentacles and shorter arms II present, buds of remaining arms develop in order voss: GENERIC REVISION OF THE CRANCHIIDAE 377

Figure I. Cranchia: Crallchia scabra. a, subadult male, ventral view, A BH45-215, ML 120 mm; b, hectocotylized right arm IV, of same; c, larva, ventral view, CI 245, ML 8 mm.

I, III, IV); digestive gland stout, spindle-shaped, with digestive duct appendages in two irregular-shaped masses on posterodorsal surface, left mass considerably larger than right; digestive duct long; gladius with long, narrow, delicate rachis, not expanded anteriorly; vanes slightly longer than distal third and moderately expanded to form spindle-shaped lanceola with strong dorsal keel; posterolateral margins of lanceola roll inward, do not overlap, distally fuse to form very short, broad, blunt-pointed, hollow conus. Distribution.-Circumglobal tropical and subtropical waters. Concentrated in upper 200 m. 378 BULLETIN OF MARINE SCIENCE. VOL. 30, NO.2. 1980

Liocranchia Pfeffer, 1884 (Figure 2)

Liocranchia Pfeffer, 1884: 25. Type species, by subsequent designation, Berry, 1914: 346: Leachia reinhardtii Steenstrup, 1856: 200. Type specimen deposited, University Zoological Museum, Copenhagen. Examined; fair condition. Fusocranchia Joubin, 1920: 73. Type species, by monotypy: Fusocranchia alpha Joubin, 1920: 73. Type specimen deposited, lnstitue of Oceanography, Monaco. Examined; fair condition. Dejinition.-Medium-sized cranchiids; elongate, spindle-shaped mantle (larva often inflated to stout, elliptical shape) tapers posteriorly to narrow point; walls thin, leathery; two moderately long cartilaginous strips diverge from apex of funnel-mantle fusions in inverted V-shaped pattern, strips of coequal or unequal length (ventral most strips the longest) bear simple, single point tubercles for entire length with 2-4 point tubercle complex at apex; nuchal fusion cartilage oval to narrow, spindle-shaped with median line of single point tubercles which continue posteriorly along median line overlying gladius for two-thirds to three-quarters of gladial length, or, with median line of tubercles absent; moderately large to large fins nearly circular in combined outline unite posterior to end of gladius and project for one-third to almost one-half their length, with small median cleft; funnel large with broad base, may extend anteriorly to base of arms; large dorsal pad of funnel organ inverted V-shaped with three longitudinal, broad or narrow, triangular flaps, a large one extending medially full length of each lateral arm and a smaller one on median line of posterior part of anterior midportion; very large, stout, kidney- or crescent-shaped ventral pads, separate or overlapping and appearing to unite at midventral line; large to medium-sized funnel valve present; head with medium to large, round, prominent, sessile eyes; olfactory organ small, narrow, or semilunar-shaped papilla on posteroventral surface; eyes with four (in half circle on ventral surface) or fourteen (seven in outer half circle on ventral surface; one midway on posteroventral surface; six in inner full circle, four ventral and two dorsal) round to oval photophores (in larva, head often withdrawn into mantle; eyes oval with short ventral rostrum, sessile; arm pillar absent); arms short, narrow, muscular, with arms I-III connected basally by low to medium depth web (inner web between arms I and I-II, outer between II-III), arm formula III > IV > II > I; moderate to well developed trabeculate protective membrane present on both margins of all arms; aboral keel present on III, absent on I and II, lateral keel present on IV; medium to large suckers largest on midarm on 1- III, may be markedly so on third quarter of III and II, on IV suckers may be largest on midarm or on base; sucker rings smooth or with flat, blunt to pointed teeth on distal margin; in males, right or left IV modified into hectocotylus (arm enlarged and elongate, with two rows of suckers basally, followed by single ventral row of enlarged fleshy suckers on laterally curved end), secondary modification on ends of III (multiple rows of small suckers) present or absent; in near- mature and mature females, brachial end-organ present on III; short, stout tentacles with slightly expanded clubs; carpal suckers in two rows (sometimes appearing as one) alternating in pairs or singly with pads extending one-half to two-thirds length of stalk; carpal group present; on club, suckers largest on midmanus, rings with sharp, conical teeth around entire margin; well developed trabeculate protective membrane on both margins, more highly developed ventrally; dorsal keel present on distal one-half to two-thirds of club (in early larva, only short, stout tentacles and shorter arms II present, buds of remaining arms develop in order I, III, IV); digestive gland stout, oval or spindle-shaped, with digestive duct appendages in two irregular-shaped masses on posterodorsal surface, left mass usually larger than right; digestive duct long; gladius with long, narrow, voss: GENERIC REVISION OF THE CRANCHIIDAE 379

Figure 2. Liocranchia: Liocranchia reinhardtii. a, subadult female, ventral view, 04008 V, ML 157 mm; b, larva, ventral view, D 1157 XIII, ML 6 mm; c, hectocotylized left arm IV, mature adult male, WH 471-1-71, ML 180 mm. delicate rachis, not expanded anteriorly; slender vanes bordering slightly less than distal half of rachis form elongate, moderately expanded lanceola with strong dorsal keel; posterolateral margins of lanceola turn inward, do not overlap, distally fuse to form short, narrow-pointed, laterally compressed, hollow conus. Distribution.-Circumglobal tropical and subtropical waters. Surface to about 1,200 m. 380 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

Leachia Lesueur, 1821 (Figure 3)

Leachia Lesueur, 1821: 89. Type species, by monotypy: Leachia eye/ura Lesueur, 1821: 90. Type specimen not extant. Neotype to be selected, Perothis Rathke, 1835: 149. Type species, by subsequent designation Hoyle, 1910: 411: Pero/his eschscho/tzii Rathke, 1835: ]49, Deposition of type specimen unknown, probably not extant. Peroti.\', Gray, 1849: 41. Erroneous spelling of Perothis Rathke, 1835. Dycrydiopsis Rochebrune, 1884: 10, Type species, by monotypy: Loligopsis ellipsoptera Adams and Reeve, 1848: 2, Type specimen not extant. Dictydiopsis Hoyle, 1885: 324, Erroneous spelling of Dyctydiopsis Rochebrune, 1884, Pyrgopsi.\' Rochebrune, 1884: 17, Type species, by monotypy: Pyr[?opsis ryne/lOphorus Roche- brune, 1884: ]7. Type specimen deposited, National Museum of Natural History, Paris. Exam- ined; poor condition. Drech.\'elia Joubin, 1931: 197. Type species, by monotypy: Drechse/ia danae Joubin, 1931: 197. Type specimen deposited, University Zoological Museum, Copenhagen. Examined; fair condition. Definition.-Small to medium-sized cranchiids; elongate, spindle-shaped mantle tapers posteriorly to slender point, walls thin, leathery; single short to long cartilaginous strip extends from anterior apex of funnel-mantle fusions along dorsal most of two lines of fusion, strip bears single to multipoint tubercles for entire length, independent free standing tubercles mayor may not continue in a line posteriorly; nuchal fusion cartilage with short anterior protrusion that laterally mayor may not bear two to four flattened or rounded tubercles; moderately large to large fins transversely elliptical in combined outline, posterior margins extend and fuse just beyond end of gladius with small median cleft (in larva, stout gladial spine projects beyond posterior margin of fins); funnel moderately large to large, extending anteriorly from mideye level to base of arms; dorsal pad of funnel organ broad, stout, inverted V-shaped with three to seven papillae (one to three short to long flattened papillae on each lateral arm and one short to long papilla on anteromedian line of anterior midportion); moderately large to large ventral pads elongate; funnel valve absent; head with small to medium-sized, round, protruding, anteriorly oriented, or large, round, prominent, sessile eyes; short, stout olfactory papilla present on ventrolateral surface; short arm pillar mayor may not persist in adult (in larva, head with long arm pillar; eyes oval with short ventral rostrum, on long stalks); eyes with five to twenty-one, round to oval photophores in roughly two rows on ventral surface, inner row may extend around dorsal surface; arms short to medium length, stout with arms III markedly more robust and elongate, arm formula III > II > IV > I (in larva, arms III elongate early); trabeculate protective membrane moderately developed on both margins of all arms; aboral keel on I-III and lateral keel on IV low to weakly developed; adult females with small to medium-sized suckers, largest on midarm; sucker rings with flat, pointed teeth on distal and lateral margins (in L. danae, sucker rings on midportion of III have central tooth enlarged and hook-like); adult males display marked sexual dimorphism with increased robustness and elongation of all arms particularly III, with suckers on midportion of all arms except right IV enlarged but with reduced apertures (except in L. danae where basal suckers are enlarged); right IV modified into hectocotylus (elongate, curved laterally with distal one-third to one-half modified by enlarged, reduced-aperture suckers in ventral row on flared ventral margin, suckers in dorsal row reduced and may be lost distally); secondary modification of ends of other arms absent; in near-mature and mature females, ends of III modified into brachial end-organ, except in L. danae where ends of all arms are modified; short, slender tentacles with short, slightly expanded clubs; two rows of carpal suckers alternating with pads extend about half length of stalk; on manus, suckers of two median rows voss: GENERIC REVISION OF THE CRANCHIIDAE 381

Figure 3. Leachia: Leachia atlalltica. a, mature adult female, ventral view, 10M 1880, ML 105 mm; b, hectocotylized right arm IV, mature adult male, P 6904-1, ML 85 mm; c, juvenile, ventral view, MBI 552, ML 32 mm; d, larva, ventral view, MBI 67, ML 8 mm. Leachia lemur. e, brachial end- organ on right arm 111,mature adult female, UMML 31.1648, ML 60 mm. greatly enlarged; sucker rings with small, short, narrow, blunt to pointed teeth on entire margin; well developed trabeculate protective membrane present on both margins; well developed dorsal keel present on distal one-half to two-thirds of club; digestive gland stout, spindle-shaped; two large lobes of digestive duct appendages merge around posterior end of long digestive duct prior to entry into caecum; gladius with strongly keeled rachis, slightly expanded anteriorly, bordered on distal half by long, narrow vanes that form an elongate, spindle-shaped 382 BULLETIN OF MARINE SCIENCE, VOL. 30. NO.2, 1980

lanceola whose posterolateral margins turn inward and fuse to form a long, narrow, hollow, ventrally flattened conus that distally becomes solid and extends to a slender point. Distribution.-Circumg]oba] tropical and subtropical waters. Surface waters to about 2,000 m.

Subfamily Taoniinae Pfeffer, 1912

Taoniinae Pfeffer, 1912: 644. Galiteuthinae Berry, 1912a: 315. Teuthoweniinae Grimpe, 1922: 51. Diagnosis .-Mant]e without cartilaginous strips along lines of funne]-mantle fusion; fins vary from small, separate, subterminal to rounded, terminal, to long, lanceo]ate, terminal or terminal-lateral, with anterior insertions on lateral margins of ]anceola or of mantle; funnel free from head laterally; eyes with one to three markedly dissimilar-sized photophores, with the largest one usually crescent- shaped (exceptions-rough]y triangular in Helicocranchia, half round in Batho- thauma, round in Sandalops); in males, hectocoty]us absent, secondary sexual modification of ends of arms may be present on I, II, III, or on all arm pairs, or absent; in near-mature and mature females, modification of arm tips into brachia] end-organ may be present on I, II, III or on all arm pairs, or absent; caecum smaller than stomach. Type genus.-Taonius Steenstrup, 1861.

Helicocranchia Massy, ]907

(Figure 4)

He/icocranchia Massy, 1907: 382. Type species, by monotypy: Helicocranchia pfefferi Massy, 1907: 382. Type specimen deposited, British Museum (Nat. Hist.), BM 1909.11.27 .10. Examined; fair condition. Ascocranchia G. Voss, 1962: I. Type species, by original designation: Ascocranchia JOLlbini G. Voss, 1962: I. Type specimen deposited, Institute of Oceanography, Monaco. Examined: fair condition. Definition.-Small cranchiids; mantle cylindrical, stoutest in midportion, tapering posteriorly to a blunt point that is free from support of g]adius; mantle walls thin but muscular with gelatinous or membranous outer layer that mayor may not be covered with closely set papillae (in larva, mucous outer coat present on mantle and head); funnel-mantle fusion area broad with, barely discernable narrow cartilage, tubercles absent; mid-dorsal anterior mantle margin free, with median cleft; nuchal fusion cartilage irregular oval-shaped, tubercles absent; small, subterminal fins paddle-shaped, fused distal to tip of gladius, with free posterior lobes, all but very anterior margin insert on lateral margins of posterior tip of g]adius that projects dorsally free of end of mantle; funnel very large, broad, extending beyond base of arms; dorsal pad of funnel organ on posterior wall of funnel, small, stout, inverted V-shaped with three slender papillae (one on mid-anterior portion of each lateral arm and one centra] in midapical region); ventral pads on anterior wall of funnel, curved c1ub- to roughly L-shaped with posterior portion shorter and stouter than anterior portion; funnel valve absent; head with small to medium-sized, protruding, anteriorly oriented, ova] to round eyes (in larva and juvenile, head with short arm pillar; eyes ova] with pronounced ventral rostrum, on short, stout stalks); eye with single small to large, often poorly defined, oval voss: GENERIC REVISION OF THE CRANCHIIDAE 383

d ,~ (1

~ 'til t ... ~f~ ~ >' . fr·~~;.·::' "j:.!

,.... ':::.!•.:~.:.!.'

~:':: w

~~':;':: ... ' '.:.):~'~~~ ;:,".~~;.;::'". ..,' :.~f.~~"';':"~ ...; .... -- ~:.~::. ",,: ::.:;.~ ::-:...... ):

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Figure 4. Helicocranchia: Helicocranchia papillata. a, immature adult female (outermost skin layer missing), ventral view, D 1185, ML 67 mm; b, partial ventral view of mature adult male showing papillation of intact skin, D 1189 II, ML 63 mm; c, juvenile, ventral view, OA 9-14-N, ML 33 mm; d, larva, ventral view, OA 9-18-N, ML 8 mm. Helicocranchia pfefferi. e, left eye, posterior and lateral views showing photophore, subadult male, Eit 26-3, ML 60 mm.

l to triangular photophore on posteroventral surface ; olfactory organ a short, stout papilla; short to medium length muscular arms joined by low web; arm formula III > II > I = IV or III > II > I > IV; trabeculae weakly developed to absent in low to medium protective membrane on both margins of all arms; suckers

I Reexamination of the ho1otype of AsrocfOnchia joubini revc~led that the two or three small round spots on the anterior sunace of the eye do not appear to be photophores as originally described. 384 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

medium to large with fine teeth on only distal or on entire margin (in juvenile, arms short, slender with small suckers); marked sexual dimorphism present with adult males showing enlargement of all suckers and increased robustness of all arms particularly III, with ends of I-IV additionally secondarily modified (multiple rows of suckers with or without expanded protective membranes), to greatest extent on I-II; adult females with no suckers disproportionately enlarged or with three or four pairs greatly enlarged on third quarter of III and to decreasing 2 degree on third quarter of II, I and IV; brachial end-organs absent ; long, low aboral keel present on I-III, lateral keel on IV; complete tentacles not known in adult; in juvenile, tentacles long, muscular with bases stouter than arms (in adult, bases of arms usually much the stouter); two rows of carpal suckers set in pairs alternating with pads extend nearly to base of stalk with carpal cluster absent; small, slightly expanded club with suckers on long pedestals; on manus, suckers of median two rows slightly enlarged and on dactylus, suckers show gradual enlargement from dorsal to ventral margin, or, suckers on both manus and dactylus show enlargement from dorsal to ventral margin with largest suckers in ventral row on distal manus and proximal dactylus; digestive gland stout, spindle- shaped with digestive duct appendages in two large masses on posterodorsal surface; digestive duct of medium length; gladius with long, delicate rachis, expanded anteriorly, very narrow medially; vanes very short, slightly expanded or flared, turn ventrally and fuse to form very short and broad, or short and narrow, hollow conus with terminal, very short, broad, or short, narrow, solid dorsal rostrum that is ventrally flattened and dorsally keeled, Distribution.-Circumglobal tropical and subtropical waters. Subsurface waters to greater than 1,000 m.

Bathothauma Chun, 1906 (Figure 5)

Batholhauma Chun, 1906: 86. Type species, by monotypy: Balholhauma lyromma Chun, 1906: 86. Type specimen deposited, Zoological Museum, Humboldt University, Berlin. Examined; poor condition. Lellcocranchia Joubin, 1912: 397. Type species, by monotypy: Lellcocranchia pfefferi Joubin, 1912: 397. Type specimens deposited, Institute of Oceanography, Monaco. Examined; good to fair condition. Definition.-Medium-sized cranchiids; mantle elongate, cylindrical, sac-shaped, rounded posteriorly; mantIe wall thin, membranous; funnel-mantle fusion broad with barely discernable narrow cartilage, tubercles absent; broad nuchal fusion without defined cartilage, tubercles absent; fins small, rounded, paddle-shaped, widely separated, subterminal, inserted on lateral expanded ends of transverse extensions of posterior end of gladius; funnel large, extending approximately to mideye level; dorsal pad of funnel organ inverted V-shaped with swollen mid- anterior portion and with posterior portion of lateral arms swollen into bluntly pointed, stout cones; ventral pads elliptical to kidney-shaped; funnel valve absent; head dominated by large to huge, protruding, anteriorly oriented, nearly spherical eyes with majority of posteroventral surface covered by single, large, half round- shaped photophore indented in mid-anterior margin (in larva and juvenile, head with long arm pillar; eyes oval with ventral rostrum, on long stalks); olfactory organ small, trumpet-shaped, narrow-stalked; well developed buccal membrane

2 On reexamination of holotype of Ascocranchiu joubin;, a mature female, the originally described three rows of suckers on ends of arms appear to be an artifact of contraction. voss: GENERIC REVISION OF THE CRANCHIJDAE 385

Figure 5. BlI1hothallma: Bathothallma Iyromma. a, subadult female, ventral view, 0 4713, ML 165 mm; b, early juvenile, dorsal view, GS 102, ML 70 mm; c, larva, dorsal view, 0 1140 I, ML 6 mm. forms low inner web between arms; arms of juveniles and adult females short and slender with arm formula in females III > IV = II > I, all arms of adult males stouter and longer with arm formula I > III > IV = II; weak trabeculate protective membrane present on both margins of all arms; aboral and lateral keels absent; medium-sized suckers gradually enlarge to midportion of all arms then decrease distally; sucker rings incised with low, flat, round to pointed teeth on entire or distal margin with rings of distal suckers more distinctly incised with fewer teeth confined to distal margin; rings in large adults often nearly smooth, in adult males with reduced apertures; in adult males, distal three-quarters of 386 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

arms I additionally secondarily modified (elongate with abruptly reduced-sized, laterally compressed suckers in two or four rows); in near-mature and mature females, brachial end-organs absent; tentacles long, muscular, with carpal suckers in two rows from near base of stalk, then in roughly four rows proximal to club forming long carpal group (in larva, distal portion of tentacles markedly stouter than proximal portion); club moderately long with suckers of manus on long pedestals, medium-sized suckers gradually enlarge to midmanus then decrease distally; sucker rings half round-shaped with conical teeth confined to distal and lateral margins; trabeculate protective membrane present on both margins diminishes proximally and extends onto stalk for most of length of carpal group; short dorsal keel present; long, thin, spindle-shaped digestive gland with long, thin, spindle-shaped ink sac embedded in median posteroventral surface and projecting beyond end, digestive duct appendanges in two large masses on posterodorsal surface; gladius with long, delicate rachis, slightly expanded anteriorly, very delicate and barely discernable medially, and very slightly expanded posteriorly prior to insertion of modified posterior end (vanes transformed into transverse bar that gradually expands laterally to shovel-shaped ends on which fins insert); conus absent. Distribution.-Circumglobal tropical and subtropical waters, extending to subantarctic. Depth range from approximately 100 to greater than 2,000 m.

Sandalops Chun, 1906 (Figure 6)

Sanda/ops Chun, 1906: 86. Type species, by monotypy: Sanda/ops me/aneho/iells Chun, 1906: 86, Type specimen deposited, Zoological Museum, Humboldt University, Berlin. Examined single remaining syntype; fair condition. Vrano/ell/his Lu and Clarke, 1974: 985. Type species, by monotypy: Vrano/ell/his bi/llcifera Lu and Clarke, 1974: 985. Type specimen deposited, British Museum (Nat. Hist.). Not examined. Definition.-Medium-sized cranchiids; mantle stout, cylindrical, broadest at approximately one-third mantle length from anterior margin, tapering gradually to broad posterior point; mantle wall thin, leathery; no tubercles present on narrow funnel-mantle fusion cartilage or on small, roughly oval nuchal cartilage; small, separate, terminal fins oval with anterior and posterior lobes, confined to posterolateral margins of lanceola with broad-pointed conus of gladius projecting slightly beyond posterior insertions; funnel very large, extending to or beyond base of arms; large, stout dorsal pad of funnel organ inverted V-shaped with large, triangular lobe obliquely set on each lateral arm and small, lanceolate papilla centered on mid-anterior portion; ventral, pads large, stout ovals; funnel valve absent; head slightly dorsally oriented, dominated by large, protruding, slightly compressed to nearly spherical, anteriorly oriented eyes with large, two- photophore complex (small, round to elliptical anterior photophore indented into, and contiguous with, mid-anterior margin of large, round posterior photophore) covering posteroventral surface (in larva, head with short, stout arm pillar; eyes on moderately long stalks, laterally compressed with long, pointed, cone-shaped ventral rostrum; in juvenile, eyes tubular); olfactory organ small, moderately long-stalked; arms short; arm formula in large females IV > I =0 II =0 III with low to medium trabeculate protective membrane on both margins of all arms; arm formula in large males III = II = I > IV or I > II = III > IV with all arms longer and stouter basally than in females, with trabeculate protective membranes expanded on all arms on basal half and absent on terminal, additionally modified half of I-III (arms I attenuated, with abruptly reduced, small suckers in two rows; voss: GENERIC REVISION OF THE CRANCHllDAE 387

Figure 6. Sandalops: Sandalops sp. a, mature adult male, dorsal view, CI 71-6-26, ML 102 mm; b, juvenile, lateral view, FIll, ML 42 mm; C, larva, ventral view, D 3782 I, ML 24 mm: d, right eye, posteroventral view (slightly diagrammatic) showing photophores, malure adult male, CI 7]-6-26, ML 102 mm; e, funnel organ, of same.

II-III attenuated, with numerous rows of minute suckers); aboral keel on I-III and lateral keel on IV present; small to medium-sized suckers largest on basal third or midportion of arms; in adult male, suckers on basal portion of all arms enlarged, to greatest extent on III, apertures reduced; sucker rings smooth to irregularly incised into low, blunt teeth; in near-mature and mature females, brachial end-organs absent; tentacles long, with two rows of carpal suckers on distal half; carpal cluster absent; short, slightly expanded club with suckers on long 388 BULLETIN OF MARINE SCIENCE. VOL. 30. NO.2. 1980

pedestals; suckers slightly enlarge to midmanus then decrease distally; rings of manus suckers with pointed teeth on entire margin with longest teeth on distal portion; protective membrane present on both margins; dorsal keel present on distal portion; long, thin, spindle-shaped digestive gland with long, spindle- shaped ink sac embedded in midposteroventral surface and often projecting beyond end, digestive duct appendages in two large masses on posterodorsal surface; digestive duct of moderate length; gladius with long, narrow, delicate rachis; vanes that border slightly more than distal quarter expand to form broad, diamond-shaped, weakly keeled lanceola; lanceola with gradual anterior and steep posterior taper; posterolateral margins of lanceola sharply turn inward and fuse distally to form short, broad-pointed, hollow conus with flat ventral surface. Distribution.-Circumglobal tropical and subtropical waters. Subsurface waters to greater than 2,000 m.

Liguriella Issei, 1908 (Figure 7)

Lif?lIriel/a Issei, 1908: 228. Type species, by monotypy: Liguriella podophtalma Issei, 1908: 228. Deposition of type specimen unknown, probably not extant (extensive search was carried out). Neotype to be selected. Vossotellthis Nesis, ]974a: 12. Erroneus identification to Desmotellthis pel/lIdda Chun, ]910. Ne- sis' specimen deposited, Institute of Oceanology, Academy of Sciences of USSR, Moscow. Examined; good condition.

Definition.-Moderately large cranchiids3; elongate, moderately stout mantle widest in midportion, tapering posteriorly to narrow, blunt point (to broad, blunt point in larva and juvenile), anterodorsal mantle margin free except at midline fusion; mantle wall thin, leathery; stout, slightly curved funnel-mantle fusion cartilage with two small tubercles at anterior end; kite-shaped nuchal fusion cartilage without tubercles; short, oval, terminal fins with small anterior lobes and small posterior lobes that appear to fuse distal to end of gladius with median cleft, all but anteriormost margin of fins insert on posterolateral margins of lanceola; funnel stout, with broad base, extending to approximately mideye level; large, stout dorsal pad of funnel organ inverted V-shaped with very broad anterior margin and with three papillae (a large, spatulate papilla on midportion of each lateral arm and a small conical papilla with spatulate tip centered on mid-anterior portion); ventral pads large, kidney-shaped; funnel valve absent; short neck tapers to head dominated by large, protruding, nearly spherical, laterally oriented eyes; ventral surface of eyes nearly covered by two photophores (large, wide, crescent-shaped posterior photophore, and smaller, oblong-shaped anterior photophore contiguous within its concavity) (in larva, head with medium to long arm pillar; eyes on medium to long stalks, oval with short ventral rostrum; in juvenile, eyes elliptical); olfactory organ small, cup-shaped, delicate-stalked; arms short, muscular, with formula III > II > IV > I (III > IV > II > I in juveniles); medium-sized suckers markedly enlarged on third quarter of III, progressively less so on II, I and IV; sucker rings on basal three-quarters of arms with numerous low, flat, square or blunt teeth on distal and lateral margins; medium width trabeculate protective membrane present on both margins of all arms; aboral keel appears to be lacking on I-III, lateral keel present on IV; secondary sexual modifications in males not known; in near-mature and mature females, brachial end-organs absent; medium length, slender but muscular tentacles with two rows

;, Adult size estimated from immature stage of development of genitalia in largest specimens available and on which definition is primarily based. 243 mm ML ~, 138 and approx. \00 mm ML 00. voss: GENERIC REVISION OF THE CRANCHIIDAE 389

d e

>:':':.::"

Figure 7. LiRl/riel/a: LiRl/riel/a podophtalma. a, subadult female, dorsal view, WH 417-1-71, ML 243 mm: b,juvenile, ventral view, WH 363-11-71, ML68 mm: c,larva, dorsal view, SAM SM34, ML 12 mm: d, right funnel-mantle fusion cartilage, WH 417-1-71: e, left eye, ventral view showing photophores, of same. of carpal suckers set in pairs; carpal cluster absent; club long, unexpanded; manus with medium, roughly coequal-sized suckers on long pedestals; sucker rings with about five to seven long, slender, pointed teeth on distal and lateral margins; weakly trabeculate protective membrane well developed on both margins; dorsal keel present on distal quarter to third of club (larva with short, slender arms; tentacles usually stout with distal portion heavier than proximal, appearance of two then four rows of carpal suckers gradually change in juvenile to two rows of carpal suckers and long, poorly defined club on long, stout tentacles); elongate, 390 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

spindle-shaped digestive gland (markedly long and slender in young) with partially embedded ink sac projecting from ventral end, digestive duct appendages in two large masses on posterodorsal surface; digestive duct of medium length; long, slender, delicate rachis of gladius very slightly expanded in anterior portion and slightly expanded just anterior to vane insertions with some thickening of margins; vanes that border approximate distal third expand to form keeled, diamond-shaped lanceola with narrow, blunt point; ventrally turned posterolateral margins of lanceola do not meet except at very tip to form short, narrow conus (lanceola of larva, very delicate with broad, blunt point). Distribution.-Subtropical and northern subantarctic waters of South Atlantic; tropical, subtropical and northern subantarctic waters of Indian and western Pa- cific Oceans, Subsurface to greater than] ,000 m.

Taonius Steenstrup, ]86] (Figure 8) Taollius Steenstrup, 1861: 84, Type species, by original designation: Loligo palla Lesueur, ]821: 96. Type specimen deposited, National Museum of Natural History, Paris. Examined, fair condition. Desmoteuthis Verrill, ]881: 300. Erroneus identification to TeutholVellia megalops (Prosch, 1849). Verrill's specimen deposited, U.S. National Museum of Natural History, USNM 883. Examined; fair condition. Toxeuma Chun, 1906: 86. Type species, by monotypy: Toxel/ma be/aile Chun, 1906: 86. Deposition of type specimen unknown (not in Leipzig or Berlin). Invalid junior homonym of Toxeuma Walker, 1833 (Insecta). Be/ollella Lane, 1957: 220. Substitute name for Toxel/ma Chun, ]906. Definition.-Moderately large cranchiids; mantle very long and slender, widest near anterior margin, tapering posteriorly to filiform tip; mantle wall thin, leathery; small, narrow to stout, roughly oval funnel-mantle fusion cartilages without tubercles (single tubercle may be present in young); oblong nuchal fusion cartilage extending onto head, tubercles absent; terminal fins long, narrow, lanceolate, with small anterior lobes, very attenuate posteriorly, extending as barely detect- able fringe to tip of gladius; except for anteriormost margin, anterior fin insertions on posterolateral margins of lanceola (young assume general adult appearance of mantle and fins very early in development);funnellarge, extending to or slightly beyond mideye level; dorsal pad of funnel organ small, stout, inverted V-shaped, with three papillae (a short to medium height, fat, round papilla on each short lateral arm and a fat, round papilla with thin terminal flap centered on large mid- anterior portion); small to large, oval to kidney-shaped ventral pads; funnel valve absent; short tapering neck joins mantle to head 'dominated by huge protruding, anteriorly oriented; nearly spherical eyes; ventral surface of eyes with two photophores (large, crescent-shaped posterior photophore with small, crescent- shaped anterior photophore within its concavity) (in larva, head with long arm pillar; eyes on long stalks, oval, with short ventral rostrum; in juvenile, eyes tubular, anteriorly oriented, with large double photophore complex usually having compact tripartite appearance on posteroventral surface); olfactory organ small, cup-shaped, stout-stalked; short, muscular arms connected by low inner web, with arm formula in adult and juvenile III > II > IV > I or III == II > IV == I; aboral keel present on I-III, lateral keel on IV; trabeculate protective membrane moderately developed on both margins of all arms; suckers of approximately coequal size on IV, become large on midportion of I-III; rings of basal suckers usually smooth, on midarm become incised with broad, blunt teeth on distal or entire margin; in large males, ends of I-III secondarily modified (two rows of voss: GENERIC REVISION OF THE CRANCHIIDAE 391

:, 'rl:. ;i' 11 Ii !l

Figure 80 Taonius: Taonius pavoo a, immature adult female, dorsal view, 0 3664, ML 539 mm (tentacle reconstructured from mature adult male, 0-4812, ML 540 mm); b, juvenile, ventral view, D 1139 III, ML 80 mm; c, larva, ventral view, D 1323 II, ML 20 mm; d, right eye, ventral view showing photophores, immature adult female, 0 3664, ML 539 mm; e, left tentacular club, large juvenile male, OA 12-62, ML 198 mmo abruptly reduced, small suckers); in near-mature and mature females, brachial end-organs absent; short, muscular tentacles with two rows of carpal suckers on approximate distal two-thirds of stalk; poorly defined carpal cluster present; strong, moderately long club slightly expanded, with well developed trabeculate protective membrane on both margins and low dorsal keel on distal third to half; suckers on manus enlarge, become increasingly unequal with growth with those of ventral median row becoming very large; rings of marginal suckers with long, 392 BULLETINOF MARINESCIENCE,VOL. 30, NO.2, 1980

sharp teeth on distal and lateral margins; rings of median suckers become compressed laterally with distal margin drawn out into one or two large, central, hook-like teeth, with large suckers of ventral median row most strongly modified (modification begins at about 50-60 mm ML); digestive gland stout, spindle- shaped; digestive duct appendages extend from posterodorsal surface of digestive gland down long digestive duct to anterior surface of caecum; gladius with very long, slender rachis and long (may exceed half gladius length) moderately expanded vanes that posteriorly turn ventrally, overlap, and extend into hollow, filiform conus and appear to fuse in posteriormost portion. Distribution.-Circumglobal subantarctic to subarctic waters. Subsurface to greater than 2,000 m, concentrated below 200 m.

Galiteuthis Joubin, 1898 (Figure 9)

Galiteuthis Joubin, 1898: 279. Type species, by monotypy: Galiteuthis armata Joubin, 1898: 279. Type specimen deposited, Museum of Natural History, Nice. Examined; poor, very fragile condition. Taonidium Pfeffer, 1900: 192. Type species, by monotypy: Taonius suhmi Hoyle, 1886: 192. Type specimen deposited, British Museum (Nat. Hist.), BM 90,1.24.19. Single remaining syntype examined (CHALLENGER Sta. 159); poor condition. Crystalloteuthis Chun, 1906: 85. Type species, by monotypy: Crystal/oteuthis glacia/is Chun, 1906: 85. Type specimen deposited, Zoological Museum, Humboldt University, Berlin. Not examined. Phasmatoteuthioll Pfeffer, 1912: 728. Type species, by monotypy: Taonius richardi Joubin, 1895: 46. Type specimen deposited, Institute of Oceanography Museum, Monaco. Examined; poor condition. Cristal/oteuthis Naef, 1921: 49, Erroneus spelling of Crystalloteuthis Chun, 1906. Definition.-Moderately large cranchiids; mantle long, slender, broadest in anterior half, tapering posteriorly to slender point; mantle wall thin, muscular, and, in subadults and adults mayor may not bear numerous cartilaginous tubercles over outer surface; tubercles mayor may not be present at anterior end of roughly comma-shaped funnel-mantle fusion cartilages and of broad, triangular nuchal fusion cartilage; fins terminal, medium to long, lanceolate, becoming attenuate to varying degrees posteriorly and extending as low delicate fringe to end of gladius, anterior lobes lacking (in larva, except for very early stage, fins with intervening stout gladial spine); except for anteriormost margin, anterior fin insertions on posterolateral margins of lanceola; medium-sized funnel extends approximately to mideye level; dorsal pad of funnel organ roughly inverted V-shaped with three large, slightly flattened papillae (one on each lateral arm and one centered on mid-anterior portion); small to medium-sized, oval ventral pads; funnel valve absent; head dominated by large, protruding, anterolaterally oriented eyes; ventral surface of eyes with two photophores (large, cresent-shaped posterior photophore with small, crescent-shaped or nearly round anterior photophore within its concavity) (in larva, head with short to medium arm pillar; arms very short of approximate coequal length; eyes oval without ventral rostrum, on short to medium stalks); olfactory organ small, cup-shaped, short- stalked; arms short, muscular with formula IV == III > II > I in adult females, IV = III > II > I or I > II > IV == III in adult males (IV > III > II > I in juveniles); well developed trabeculate protective membranes more expanded on ventral margins of all arms; low aboral keel present on I-III, lateral keel on IV; moderately large suckers largest on midportions of all arms; rings smooth or with flat, square, rounded or pointed teeth on distal and lateral margins; in large males, terminal portions of I and II may be secondarily modified (two rows of abruptly voss: GENERIC REVISION OF THE CRANCHIIDAE 393

c ~~ IJ,:~!J I.;i ;~;ta 'f\l~""~i'~ ,·tt Ii ~\;~,i~;~, 1~4J \ d I) , e

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Figure 9. Galitellthis: Galiteuthis g/acia/is. a, immature adult female, dorsal view (redrawn from McSweeny, 1978), Elt 1323, ML 333 mm; b, left tentacular club (redrawn from McSweeny, 1978), immature adult male, Elt H-371, ML 297 mm; c, left eye, lateral and ventral views showing photophores (redrawn from McSweeny, 1978), immature adult female, Elt 1323, ML 333 mm; d, juvenile, ventral view, Elt 21-272, ML 54 mm; e, larva, ventral view, Elt 46-295, ML II mm.

reduced, small suckers) or, I may become very elongate and II less so; in near- mature and mature females, brachial end-organs absent; short to medium length tentacles muscular with two rows of carpal suckers set in pairs on distal two- thirds to three-quarters of stalk; carpal cluster present; short, slightly expanded club with four to eight pairs of median suckers on manus modified with growth into large, hooded hooks (in juvenile, forming hooks appear at about 35-60 mm ML); suckers of marginal rows greatly reduced or lost; moderately developed, 394 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, ]980 weakly trabeculate protective membrane on both margins; small dorsal keel present or absent; digestive gland stout, spindle-shaped; digestive duct appendages confined to posterodorsal surface of digestive gland or may extend entire length of long digestive duct; gladius with long, slender rachis bordered on approximate distal half by long, narrow vanes that roll inward posteriorly, overlap and, with or without fusing, extend into hollow, needle-like conus. Distribution.-Circumglobal Antarctic to subarctic waters. Subsurface waters to approximately 1,500 m. Mesonychoteuthis Robson, 1925 (Figure 10)

Mesonychoteuthis Robson, 1925: 272. Type species, by original designation: Mesonychoteuthis hamiltoni Robson, 1925: 272. Type specimens (two brachial crowns) deposited, British Museum (Nal. Hist.). Not examined. Definition.4-Very large cranchiids; mantle elongate, moderately stout in anterior quarter, tapering sharply then gradually to form long, narrow posterior half (in juvenile, mantle stout, displaying with growth gradual elongation of narrow posterior portion); mantle wall thin, muscular; short, stout, curved funnel-mantle fusion cartilages without tubercles (in larva and juvenile, 4-6 pointed tubercle complex present at anterior end); triangular nuchal fusion cartilage without tubercles (in larva and small juvenile, single tubercle present at anterior end); large (>50% ML), terminalfins stout ovate, without anterior lobes, tapering posteriorly to broad point at end of gladius; except for anteriormost margin, anterior fin insertions on posterolateral margins of lanceola; fins very muscular medially, tapering to thin outer margins (in larva, fins with intervening very short, broad gladial spine: in small juveniles, fins nearly round in combined outline with posterior margins extending to end of gladius); funnel moderately large, extending to slightly less than mideye level, funnel opening small; dorsal pad offunnel organ not known in adult (in juvenile, inverted V-shaped with three papillae, a short, stout papilla on each lateral arm and long, narrow, conical papilla centered on mid-anterior portion); small ventral pads oval; funnel valve absent; head dominated by large, nearly spherical, protruding, anterolaterally oriented eyes; ventral surface of eyes with two photophores (large, wide, crescent-shaped posterior photophore and small, oblong anterior photophore within its concavity) (in larva, head with very short, stout arm pillar; eyes oval without ventral rostrum, on short, stout stalks; in juvenile, arm pillar absent; large, protruding eyes nearly spherical); olfactory papilla short, stout; arms robust, of medium length, with approximate formula in adult female III == IV == II > I (not known for adult male; IV > III > II > I in juvenile); trabeculate protective membrane well developed on ventral margins of all arms, lesser developed on dorsal margins; low aboral keel present On distal one-half to two-thirds of I-III; lateral keel present On IV; suckers On all arms gradually enlarge to approximately midarm, are then replaced by three to approximately nine pairs of large hooded hooks (arms I through IV have increasing number of hooks) (in juvenile, forming hooks appear at about 45 mm ML); rings of suckers basal to hooks smooth to irregular on distal and lateral margins (more distinctly incised in juveniles); rings distal to hooks with broad, flat, triangular teeth on distal margin; in large males, all arms elongate with terminal portions of II and III appearing to be secondarily modified (attenuated with

• Definition of adull based primarily on a nearly complete 125 em ML 9 (inner organs missing) specimen, judged to be immature, and several large panial specimens, one a mature <5, in the collections of the National Museum of Natural History, Smithsonian Institution. voss: GENERIC REVISION OF THE CRANCHIIDAE 395

Figure 10. Memnychoteuthis: Mesonychoteuthis hamiltoni. a, subadult female, ventral view, TM 2, ML 1,250 mm; b, juvenile, ventral view (redrawn from McSweeny, 1970), Elt 142, ML 86 mm; c, larva, dorsal view, Elt 33-5, ML 23 mm; d, left arms I-IV (redrawn from McSweeny, 1970),juvenile, Elt ]421, ML 86 mm. abruptly reduced-sized suckers in two rows); suckers immediately distal to hooks on I-III of large males larger than those immediately preceding hooks rather than smaller as on IV and on all arms of large females; in near-mature and mature females, brachial end-organs probably absent; moderately long tentacles robust; alternating, diagonally set pairs of suckers and pads extend approximately two- thirds length of stalk; carpal cluster present; short, strong, only slightly expanded club with suckers of two median rows of entire manus modified into approximately twelve pairs of hooded hooks that coequally enlarge to midclub, then 396 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

decrease distally (hooks well formed on proximal manus at 59 mm ML); suckers of marginal rows minute; protective membranes absent except for low portion on both margins of greatly reduced dactylus; dorsal keel absent on rounded aboral surface; digestive gland stout, spindle-shaped with digestive duct appendages confined to posterodorsal surface; digestive duct long; long, medium width vanes of gladius border approximate posterior three-quarters of narrow rachis; posterolateral margins of vanes roll ventrally, and distally overlap and fuse to form long, narrow, strong, hollow conus. Distribution.-Circumglobal Antarctic waters. Depth range little known. Shal- lowest taken in haul of 0-494 m with most larvae and juveniles taken in hauls of 0-1,000 to greater than 2,000 m. Available adults and subadults all taken from stomachs of sperm whales.

Egea Joubin, 1933 (Figure 11)

Euea Joubin, 1933: 48. Type species, by monotypy: Egea inermis Joubin, 1933: 48, Type specimens deposited, University Zoological Museum, Copenhagen. Examined; excellent to good condition, Lectotype selected, N. Voss, 1974: 947, Definition.5-Medium-sized to moderately large cranchiids; long, narrow, spindle-shaped mantle tapers posteriorly to attenuated tip; mantle walls thin, leathery; no tubercles present on conspicuous, spindle-shaped nuchal fusion cartilage or on elongate, triangular funnel-mantle fusion cartilages; pair of noncartilagi- nous, nontubercular diverging lines extending posteriorly from funnel-mantle cartilages sometimes apparent (external evidence of internal lines of funnel-mantle fusions); fins lanceolate, terminal-lateral without anterior lobes, tapering distally to end of gladius; fins extend anteriorly onto lateral margins of mantle for approximately one-third mantle length with anteriormost insertions anterior to level of origin of gladial vanes; moderately large funnel extends to approximate mideye level; large dorsal pad of funnel organ inverted V-shaped with broad anterior margin and triangular flap on each lateral arm, mid-anterior portion without papilla; large ventral pads irregular oval to kidney-shaped with wide cleft in anterior margin; well developed funnel valve present; short, tapering neck joins mantle to small head dominated by large, nearly spherical, protruding, anterolaterally oriented eyes; ventral surface of eyes with two long, narrow photophores (long, crescent-shaped posterior photophore and, within its concavity but close to and encircling approximately two-thirds circumference of lens, a shorter anterior photophore, crescent-shaped except for short turn at posterior end) (in larva, head with short arm pillar; eyes oval with short ventral rostrum, on short to medium stalks; in juvenile, eyes elliptical); small olfactory organ goblet-shaped; arms short, muscular with formula III > IV == II > I; strong dorsal keel on I-III and lateral keel on IV; trabeculate protective membrane well developed on both margins of all arms; on all arms, moderately large suckers gradually enlarge to midarm then decrease distally with enlargement greatest on III and progressively less on II, I and IV; sucker rings with numerous, low, flat, blunt to rounded teeth on distal, distal and lateral or entire margins; in large males, ends of all arms secondarily modified (minute suckers in four rows), to greatest extent on I; in subadult females, brachial end-organ present on III, not known in mature females; medium length tentacles muscular, with slightly expanded club; carpal suckers

, Mature animals not available. Definition based On subadults, largest available were 207 and 210 mm ML 00. voss: GENERIC REVISION OF THE CRANCHIIDAE 397

i;~. /1 \\ d II 1\ iI nIi !I !J II

Figure II. Egea: Egea if/ermis. a, subadult male, dorsal view (redrawn from N. Voss, 1974),0 2945, ML 198 mm; b, juvenile, ventral view (redrawn from N. Voss, 1974). UMML 31.1737, ML 48 mm; C, larva, dorsal view (redrawn from N. Voss, 1974),0 1214 IV, ML approximately 7 mm; d, funnel organ and valve, subadult male, WH 471-11-71, ML 207 mm; e, left eye, lateral view showing photophores, of same; f, left eye, ventrolateral view showing photophores, of same. and pads in two rows arranged in alternating, diagonally set pairs on distal three- quarters of stalk; carpal cluster absent; manus with medium-sized suckers on long pedestals; suckers gradually enlarge to midmanus, then decrease distally; suckers also show progressive enlargement from dorsal most to ventral most rows with largest suckers in ventral marginal rows; sucker rings with approximately 7-12 sharp, conical teeth confined to distal and lateral margins; well developed trabeculate protective membrane present on both margins; long, well developed dorsal keel extends for most of length of club; digestive gland stout, spindle- 398 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

shaped with long, narrow ink sac partial1y embedded in posteroventral surface; digestive duct appendages extend in clusters from digestive gland along entire length of long digestive duct; gladius with long, delicate, very narrow rachis with no anterior expansion; lateral margins of rachis gradual1y thicken resulting in formation of median ventral groove on distal third to half of rachis; delicate vanes bordering approximate posterior third of rachis form narrow, elongate lanceola; distal posterolateral margins of vanes overlap without fusing and extend into hol1ow, needle-like conus. Distribution.-Tropical waters occasional1y to subtropical waters of Atlantic, Indian and western Pacific Oceans. Subsurface waters to possibly 2,000 m.

Megalocranchia pfeffer, 1884 (Figure 12) Megalocranchia Pfeffer, 1884: 24. Type species, by monotypy: Megalocranchia maxima Pfeffer, 1884: 24. Type specimen deposited, Hamburg Natural History Museum. Not extant, destroyed in World War II. Neotype to be designated. Corynomma Chun, 1906: 85. Type species, by monotypy: Corynomma speculator Chun, 1906: 85. Type specimens deposited, Zoological Museum, Humboldt University, Berlin. Examined; good condition. Lectotype selected, Nesis, 1974a: 6. Xenoteuthis Berry, 1909: 419. Error. In Berry's paper, his species He/icocranchia fisheri appears in a faunal list as Xenoteuthis fisheri Berry. Ascoteuthis Berry, 1920a: 295 (subgen.). Type species, by original designation: Teuthowenia (As- coteuthis) corona Berry, 1920a: 296. Type specimen deposited, U.S. National Museum of Natural History, USNM 338695. Examined; fair condition. Carynotellthis G. Voss, 1960: 434. Type species, by original designation: Carynoteuthis oceanica Voss, 1960: 434. Type specimen deposited, Chicago Natural History Museum, CNHM 78326. Examined; good condition.

Definition.6-Moderately large to large cranchiids; slender, elongate mantle tapers posteriorly to slender point; mantle walls thin, leathery (in larva, mucous outer coat present); no tubercles present on elongate, usually roughly triangular funnel-mantle fusion cartilages or on conspicuous spindle-shaped nuchal fusion cartilage; long, lanceolate, terminal-lateral fins extend from tip of gladius onto lateral margins of mantle for approximately one-half mantle length, with anterior fin insertions approximately at level of origin of gladial vanes, anterior lobes absent; stout, well developed funnel extends to approximately mideye level; large, stout dorsal pad of funnel organ with triangular flap on each lateral arm and with broad anterior margin, mid-anterior portion without papilla; small to medium-sized ventral pads roughly oval to kidney-shaped; well developed valve present; short neck tapers to head dominated by very large, nearly spherical, protruding, anterolaterally oriented eyes; major portion of ventral surface of eye covered by two photophores (large, crescent-shaped posterior photophore with irregular-shaped inner margin with central bulge, and within its concavity, smaller, narrow, roughly elongate S-shaped anterior photophore (in larva, head with slender, moderately long arm pillar; eyes oval with short ventral rostrum, on long stalks); olfactory organ cup-shaped, heavy-stalked; arms short, strong, with approximate formula in large males III > II == I > IV, and in large females III > II > IV == I (in juveniles and early subadults, usual formula III > IV > II > I); strong, well developed dorsal keel on I-III and lateral keel on IV; wide, trabeculate protective membrane present on both margins of all arms; on all arms, suckers small basally, enlarge to midarm then decrease distally with marked

• Only one mature animal available (in poor condition, 810 mm ML 9, M. Clarke, 1962); most of definition based on subadulls of about 450-520 mm ML. voss: GENERIC REVISION OF THE CRANCHIIDAE 399

Figure 12. Megalocranchia: Megalocranchia sp. a, subadult female, dorsal view, P 6407, ML 254 mm; b, right eye, lateral and ventrolateral views (slightly diagrammatic) showing photophores, juvenile, 0 1110 VI, ML 82 mm; c, compound photophore on ventral surface of digestive gland, of same; d, juvenile, ventral view, D 1108 I, ML 54 mm; e, larva, dorsal view, MBJ 48-52, ML 17 mm. enlargement on III, less on II, and much less on I and IV; rings of basal suckers usually with numerous, small, flat, blunt to pointed teeth on distal and lateral margins, enlarged midarm suckers with similar teeth sometimes on entire margin, distal suckers with approximately four to eight relatively large, pointed teeth confined to distal margin; in large males, ends of I-III secondarily modified (attenuated with abruptly reduced-sized, minute suckers in two rows); in subadult and adult females, brachial end-organ present on I-III, only on III or occasionally only on II; tentacles short, strong, with two rows of carpal suckers on mid-third 400 BULLETIN OF MARINE SCIENCE, VOL. 30. NO.2. 1980

of stalk, changing to four rows prior to forming strong carpal group in large animals at base of moderately long, strong, slightly expanded club (long, stout tentacles of larva with two rows of carpal suckers on basal half, four rows on distal halt); club suckers set on short to medium height pedestals; suckers of manus only moderately enlarged, with those of two median rows the largest; sucker rings with large, pointed, conical teeth on distal and lateral margins, proximal margin irregular to finely incised; trabeculate protective membrane developed on both margins; short to long dorsal keel present; large, rounded digestive gland with large, bilobed, multilens photophore covering entire ventral surface and overlying partially embedded ink sac (in larva, digestive gland stout, spindle- shaped; forming photophore appears at about 23-30 mm ML); digestive duct appendages in clusters on separate, short digestive ducts prior to their union at entrance to caecum; gladius with long, very slender rachis without anterior expansion; lateral margins of rachis gradually thicken and roll inward resulting in formation of deep ventral groove on distal two-thirds of rachis; long, moderately expanded vanes border posterior half of rachis; distolateral margins of vanes roll inward and fuse to form long, hollow, needle-like conus. Distribution.-Circumglobal tropical and subtropical waters. Subsurface waters to possibly greater than 1,000 m.

Teuthowenia Chun, 1910 (Figure 13)

Owenia Prosch, 1849: 71 (subgen.). Type species, by monotypy: Cranchia (Owenia) megalops Prosch, 1849: 71. Type specimen deposited, University Zoological Museum, Copenhagen. Ex- amined; poor condition. Invalid junior homonym of Owenia Chiaje, 1844 (Polychaeta). Teuthowenia Chun, 1910: 376. Substitute name for Owenia Prosch, 1849. Verrilliteuthis Berry, 1916: 64. Type species, by monotypy: Desmoteuthis tenera Verrill, 1881: 412. Type specimen deposited, U.S. National Museum of Natural History, USNM 574850. Examined; fair condition. Anomalocranchia Robson, 1924: 9. Type species, by original designation: Anomalocranchia impennis Robson, 1924: 9. Type specimen deposited, British Museum (Nat. Hist.), BM 1924.9.9.38. Examined; poor condition. Definition.-Medium-sized to moderately large cranchiids; conical mantle elongate, tapering from widest point near anterior margin to narrow posterior point; mantle wall thin, leathery; small, spindle-shaped funnel-mantle fusion cartilages with one to four tubercles at mantle margin; small nuchal fusion cartilage elongate, diamond-shaped, without tubercles; long, narrow, terminal-lateral fins with growth extend progressively up lateral margins of mantle to approximately 40-60 percent mantle length, with anterior fin insertions opposite point of origin of gladial vanes, anterior lobes absent; fins taper very gradually posteriorly termi- nating in small lobes whose posterior margins extend beyond tip of gladius for very short distance and fuse with median cleft; large, broad-based funnel extends to mideye level or beyond; dorsal pad of funnel organ with three long papillae (spatulate one on each narrow lateral arm and lanceolate one on mid-posterior margin of anterior midportion); medium-sized ventral pads roughly triangular; funnel valve absent; head dominated by huge, protruding, anteriorly oriented eyes with three photophores (large, crescent-shaped posterior photophore with smaller, narrow, crescent-shaped anterior photophore within its concavity on ventral surface, and third (very occasionally an aberrant fourth) small, oval photophore on inner anterodorsal margin) (in larva, head with short arm pillar; eyes oval, without ventral rostrum, on stout stalks, with two photophores; in juvenile, third small photophore forms at about 45-50 mm ML); olfactory organ small, voss: GENERIC REVISION OF THE CRANCHIIDAE 401

Figure 13. Teuthowenia: Teuthowenia megalops. a, immature adult female, dorsal view, B 17, ML 352 mm; b, juvenile, dorsal view, C 149-2, ML 92 mm; c, larva, dorsal view, D 4370 III, ML II mm; d, funnel organ, subadult male, D 1387 III, ML 136 mm; e, left eye, ventral and anterolateral views showing photophores, subadult male, C 62-431, ML 147 mm; f, left tentacular club and distal portion of stalk, of same. narrow-stalked; short to medium length arms muscular with formula III > II > I > IV in adult females, III = I > II > IV in adult males (in juveniles, III > II = IV > I; in subadults, III > II > IV > I); medium-sized suckers greatly enlarge on midportions of III and II exceeding width of arms, similar but progressively less enlargement on I and IV; rings of basal and midarm suckers usually smooth, rings of terminal suckers with flat, blunt teeth on distal margin; well developed trabeculate protective membrane on both margins of all arms basally fuse to form 402 BULLETIN OF MARINE SCIENCE, VOL. 30. NO.2. 1980 low inner web between all arms; aboral keel present on III, not known for I and II; lateral keel present on IV; arms I of mature males strongly modified, become elongate, sometimes swollen in midportion, and with increased and more pointed dentition of sucker rings, ends with suckers in two or four rows; ends of II exhibit lesser degree of modification with marked palisading appearance of enlarged pedestals; in near-mature and mature females, brachial end-organ present on all arms; glandular processes present or absent near base of midarm suckers on I-III of mature females; tentacles short, muscular, with four rows of carpal suckers set in zigzag fashion on distal one-half to two-thirds of stalk (long, slender tentacles of early larva with two rows of carpal suckers on basal third, four rows on distal two-thirds; tentacles more robust and relatively shorter in late larva; in juveniles, tentacles with four rows of carpal suckers throughout); carpal cluster absent; club slightly expanded with suckers on long pedestals; suckers gradually enlarge to midmanus with slight increased enlargement from dorsal to ventral margins; sucker rings incised on entire margin with numerous short, sharp teeth, longest on distal portion; trabeculate protective membrane well developed on both margins; dorsal keel present on distal half; digestive gland stout, spindle-shaped; digestive duct appendages extend from posterodorsal surface of digestive gland down entire length of long digestive duct; gladius with long, narrow rachis; moderate width vanes bordering posterior one-third to one-half, turn ventrally, overlap without fusing and extend into narrow, but not needle-like, hollow conus. Distribution.-Circumglobal southern subtropical to subantarctic waters, extending through tropical to subarctic waters in Atlantic. Upper 50 to 2,000 m or more.

INVALIDGENERA

Invalid genera of the family Cranchiidae that have appeared in the literature but do not belong to the synonymies of the preceding described valid genera are presented below.

Loligopsis Lamarck, 1812: 123. Type species, by monotypy: Loligopsis peronii Lamarck, 1812: 123. Type specimen not extant. The type species is unidentifiable even to family from the original description. Loligopsis is a nomen dubium. It is treated here because so many of the early described cranchiids were mistakenly identified to it.

Phasmatopsis Rochebrune, 1884: 9. Type species, by monotypy: Phasmatopsis cymoctypus Roche- brune, 1884: 9. Type specimen deposited, National Museum of Natural History, Paris. Appears to have been lost, could not be located by author or museum staff in 1975, or by museum staff in the 1976 reorganization of the cephalopod collection. The type was inadequately and inaccurately described and illustrated by Roche- brune as pointed out by Hoyle (1885) on reexamination of the large (1 m ML) unique specimen. Even with Hoyle's corrections, the type remains unidentifiable from the literature. The type was subsequently reexamined by Anna Bidder, and a short time later in 1961 by my husband, Gilbert Voss. Clarke (1962) using the notes on Bidder's external examination of the type, resurrected the genus by identifying and attributing to it the characters of a large taoniin, the specimen of which I have since reexamined and definitely identified to the genus Megalo- cranchia. A number of recent authors following Clarke's work have used Phas- matopsis as the generic name for specimens belonging to Megalocranchia. In the reexamination of the type by G. Voss, permission was granted to open the mantle. His notes describe the digestive gland as being long, "elongate like voss: GENERIC REVISION OF THE CRANCHIIDAE 403 that of Desmoteuthis hyperborea" (=Teuthowenia megalops), with no trace of photophores, quite different from the large, rounded digestive gland with the large, compound photophore covering the ventral surface present in Megalo- cranchia. In my paper on Egea inermis (1974), I discussed this Phasmatopsis problem in more detail. Since the two exhaustive attempts to relocate the type in Paris have been unsuccessful, I have no misgivings in the absence of the type in considering Phasmatopsis a nomen dubium.

Zygaenopsis Rochebrune, 1884: 13. Type species, by monotypy: Lo/igopsis zygaena Verany, 1851: 125. Deposition of type unknown. Invalid junior homonym of Zygaenopsis Felder, 1874 (Lepi- doptera). The larval type, inadequately illustrated and described by Verany, appears to be a cranchiid and has been historically regarded as such. If the character of the arms of approximately coequal length can be relied upon, then the specimen most clearly resembles a larval Galiteuthis as recognized by Chun (1910). Pfeffer (1900) introduced the more commonly held concept that Verany's specimen was a Leachia-like form, when he placed Pyrgopsis Rochebrune into its synonymy. Without the type specimen, identification is impossible. This genus is now considered a nomen dubium.

Proca/istes Lankester, 1884: 311. Type species, by monotypy: Proca/istes suhmii Lankester, 1884: 311. Type specimens deposited, British Museum (Nat. Hist.), BM 90.1.24.20.21.22. Examined; poor condition. All three types are old slide mounts; the largest measures approximately 9 mm ML. Absolutely no specific identification is possible. I have no hesitation in considering Procalistes a nomen dubium.

Henseniotellthis Pfeffer, 1900: 193. Types species, by monolypy: Hensenioteuthis joubini Pfeffer, 1900: 193. Type not extant, destroyed in World War II. Pfeffer illustrated and described the distorted larval type in 1912 but they are inadequate for identification. With the type lacking, nothing remains but to con- sider Hensenioteuthis a nomen dubium.

Zygocranchia Hoyle, 1909: 276. Substitute name for Zygaenopsis Rochebrune, 1884. Since Zygaenopsis is a nomen dubium, then Hoyle's substitute name has no validity.

Euzygaena Chun, 1910: 354. Substitute name for Zygaenopsis Rochebrune, 1884. Chun's work was in press at the time Hoyle's substitute name Zygocranchia appeared in print. Euzygaena therefore is invalid by reason of being a synonym of Zygocranchia Hoyle, as well as being based on a nomen dubium.

Paratellthis Thiele, 1921: 463. Type species, by monotypy: Paratellthis tunicata Thiele, 1921: 463. Deposition of type unknown. Thiele included the genus in the Cranchiidae, but the original text and illustrations indicate it is definitely not a cranchiid. It has been considered possibly to be a larval ALLuroteuthis by Odhner (1923) and Roper, Young and G. Voss (1969).

OTHER NAMES Cranchiidarum Chun, 1903. This name appears in the synonymies of Corynomma speculator and Toxeuma be/one in Clarke (1966). "Cranchiidarum Chun, 1903" was first used by Chun 404 BULLETIN OF MARINE SCIENCE, VOL. 30, NO.2, 1980

(1910) coupled with "nov. gen." or "n. gen," in the synonymies of the above two species and Crystalloteuthis glacialis, all new species of his. Chun obviously did not mean Cranchiidarum to be the name of a new genus but rather a concise latinized word meaning "of cranchiid." It does not appear in the 1903(b) work referred to by Chun.

DISCUSSION It is difficult at first to see much unity in such a diversified group as the Cran- chiidae. Study however shows that all of the diverse morphological forms in the family develop from a common early larval type (p. 374). The different early larval characters are lost or modified at varying stages during growth in each genus. The larval stalked eyes, for example, are completely lost in the development of Cranchia and Liocranchia and persist for varying periods in the development of the other genera. They appear only in the early larva of Mesonycho- teuthis, and are retained longest, up to the near-mature stage, in some species of Leachia. The paddle-shaped fins are retained in near-larval form in the adult stage of Helicocranchia and Bathothauma, become longer and rounded with growth in Cranchia, Liocranchia, Leachia, Sandalops and Liguriella, elongate further in Taonius, Galiteuthis and Mesonychoteuthis and extend anteriorly along the lateral margins of the mantle in Egea, Megalocranchia and Teuthowenia. The short conus of the larva is retained in Cranchia, Liocranchia, Helicocran- chia, Sandalops and Liguriella, is elongated to varying degrees in the other genera, and is lost completely in Bathothauma. The short arms generally are retained by all genera, but a dimorphic sexual elongation sometimes occurs in adult males. The tentacles shorten proportionately with growth except in Helicocranchia, Bathothauma, Sandalops and Mesonychoteuthis. A comparison of some basic morphological characters for all genera of the Cranchiidae (Table I) shows that the division into two subfamilies is a natural one. The relatively homogeneous Cranchiinae is characterized by the cartilaginous strengthening along one or both lines of funnel-mantle fusions, the lateral fusion of the funnel to the head, four or more small photophores on the eye, a hectocotylus, and a caecum larger than the stomach. Cranchia and Liocranchia are closely allied genera with few species; Leachia is more speciose. The more speciose and morphologically diverse Taoniinae may be divided into three groups of genera. Helicocranchia. Bathothauma, Sandalops and Liguriella form a loose, perhaps unnatural, group of the more primitive genera which retain a short conus (except for Bathothauma in which it is lost) and small fins. Of these, Helicocranchia and Bathothauma share paddle-shaped, subterminal fins, a single large photophore on each eye and broad funnel-mantle fusion areas with barely discernable cartilages. A second and smaller photophore occurs on the eye in Sandalops and Liguriella, but it remains contiguous with the large photophore rather than being removed, as in the remainder of the taoniins. Taonius, Galiteuthis and Mesonychoteuthis form a group in which the conus and associated terminal fins are elongated and the large suckers of the club are modified into hooks or laterally compressed suckers with one or two large, central, hook-like teeth. The second or anterior photophore on the eye is separate, short and small. Galiteuthis and the more specialized Mesonychoteuthis are closely related, sharing a similar larval type, clubs with fully developed hooks, an arm formula of IV > III > II > I in juveniles and more highly developed ventral trabeculate protective membranes on all arms. Egea, Megalocranchia and Teuthowenia form a third group in which the paddle-shaped fins of the larva are retained for a longer period during development voss: GENERIC REVISION OF THE CRANCHIIDAE 405

Table I. Morphological characters of the genera of Cranchiidae

Cranchiinae Taoniinae ~ .~ "" .~ "" "" ~ ~ u .~ ~u E <; ~ .S "" ~ ~ ~ u ."-1 "" <:: E ~ ..s "" u :<; ~ u "" ~ ~ ~ ~ u <:: :; <:: :<; u <; .S ~ ~ ~ u ~ "" ~ "" ~ ~ ~ .~ Sl <:: .~ ~ ;; ~ .~ c c; ~ ~ :; Chamcter 8 --l --l ~ ., '" --l ~ <.:l ~ ~ ::;; ~ Mantle Ventral cartilaginous strips present X X X Fins Small, paddle-shaped, subterminal X X Elliptical, oval or circular, terminal X X X X X Lanceolate or stout ovate, terminal X X X Lanceolate or long, narrow, terminal- lateral X X X Funnel Fused to head laterally X X X Valve present X X X X Eyes Four or more small photophores X X X One usually large photophore X X One large + one small contiguous photophore X X One large + one small, short non-contiguous photophore X X X One large + one long, narrow non-contiguous photophore· X X X Armst Hectocotylus present in males X X X Brachial end-organs present in females X X X X X X Hooks present X Clubs Hooks present :j: X X Digestive gland Stout, spindle-shaped X X X X X X X X X Elongate, spindle-shaped X X X Rounded with large photophore X Digestive duct appendages On digestive gland X X X X X X X§ X On digestive gland and ducts X X§ X X On posterior end of ducts or on caecum X X Caecum Larger than stomach X X X Smaller than stomach X X X X X X X X X X Gladius Short conus X X X X X Medium to long conus X X X X X X X Conus lacking X

• In Te",holVe"ia a third and occasionally an aberrant fourth small photophore forms with growth. t Additional dimorphic sexual features of the arms are compared in Table 2. * Rings of large suckers laterally compressed, with one or two large, central, hook-like teeth on distal margin. § Nol presenl in all species. 406 BULLETIN OF MARINE SCIENCE. VOL. 30. NO.2. 1980

than in the preceding group. The conus and fins elongate with growth and simultaneously the fins extend anteriorly along the lateral margins of the mantle. In this group the second photophore on the eye is separate, long and narrow and in one genus, Teuthowenia, a third and occasionally fourth small photophore forms with growth. The three are the only genera in the Taoniinae known to have the brachial end-organs in the females. Egea and Megalocranchia are closely allied sharing a similar larval type, a funnel valve, and a similar form of dorsal funnel pad and funnel-mantle fusion cartilage. The above grouping of genera is based more strongly on morphological characters than that proposed by Nesis (1974a) in which the Taoniinae was divided into four groups based solely on the extent of the development of the "tail," or the terminal end of the gladius. Nesis' groups were as follows: "Long-tailed" squids-Taonius, Phasmatopsis (=Megalocranchia), BelonelLa (=Taonius), Gal- iteuthis, Mesonychoteuthis; "Medium-sized tailed" squids-Corynomma (= Meg- alocranchia), Verrilliteuthis (=Teuthowenia); "Short-tailed" squids-Vossoteu- this (=LigurielLa), Sandalops, H elicocranchia, Ascocranchia (=Helicocranchia); "Tailless" squids-Bathothauma; with Egea as a transitional form from "long- tailed" to "short-tailed" and LigurielLa as a transitional form from "short-tailed" to "tailless." Unfortunately, Nesis lacked the large comparative world-wide collections and the developmental series that would have permitted him to distinguish between specific, generic and ontogenetic characters. Furthermore, the development of the "tail" alone is not as systematically valuable as the relationship of the fins and the posterior end of the gladius (lanceola and conus). The importance of this relationship was first recognized by Pfeffer (1912), but his systematic conclusions were premature; he confused adult and larval characters, because most of the genera at that time were only known from larval or juvenile forms. Table 2 shows the high degree of sexual dimorphism of the arms that exists in the family. Despite the generic diversity, there is a common basic form for each major sexual modification. The structure of the hectocotylus, the secondary modifications in the male and the brachial end-organ in the female is similar between genera, varying chiefly in extent of development. The secondary sexual modifications in the male are more variable, as would be expected, in the Taoniinae. The first and third pairs of arms are frequently the most strongly affected and the fourth pair the least. These diverse sexual modifications of the arms suggest that equally diverse behavior patterns occur in courtship and copulation. The sexual modification of the tips of the arms in females into an organ is unique in the teuthoids. The brachial end-organ is more common and proportionately larger in the Cranchiinae. It develops at a later growth stage than does the hectocotylus or secondary male modifications. This is expected, since the male matures at a smaller size, and then it continues to grow while the female reaches sexual maturity at near-maximum size. R. Young (1975b) suggests that the brachial end-organ is a photophore used possibly for sexual attraction. The function of the other sexual modifications in the females of some species of Helicocranchia, where midarm suckers are greatly enlarged, and of one species of Teuthowenia, where glandular processes develop at the base of midarm suckers, is not known.

ACKNOWLEDGMENTS

I am most grateful to the numerous individuals and institutions who have loaned or made available collections, types and supplementary material for this study: S. S. Berry, Redlands, Calif.; P. Bouchet voss: GENERIC REVISIONOF THE CRANCHIIDAE 407

Table 2. Dimorphic sexual features of the arms for the genera of Cranchiidae

.; 2 Secondary ModificationsPresent Other BrachialEnd- Modi- Hect<>- Ends of Arms' En- Organ Present fica- cotylus tions Subfamily Pres- I- I- I- ll- ~f~ri I- I- Pres- Genera ent II III IV III III Arms2 III IV III ent

Cranchiinae Cranchia X X X Liocranchia X X· X Leachia X X X· X Taoniinae Helicocranchia X X X· Bathothauma X X Sandalops X X Liguriella3 Taonius X Galiteuthis X Mesonychoteuthis ?X X Egea X X7 Megalocranchia X X X8 Teuthowenia XS X XIO

I To varying degrees, ends of arms elongate, usually become attenuate. with two or more rows of abruptly reduced-sized suckers. a All arms elongate, usually become more robust, generally with enlarged suckers often with reduced apertures. In Mesonychoteulhi .•• all arms elongate. but only the suckers distal 10 the hooks on I-III show enlargement,with no reduction of apenures. 3 Secondary sexual modifications in males not known for Uguriella . .•Not present in all species. 5 In addition, arms 1 sometimes become very swollen in mid-portions . • One species. L. danae. has the brachialend-organon all arms. T In subadulls: no mature females available. 8 Some species have the brachial end.organ only on III. occasionally only on f! . • In at least one species. H. papilla/a, several pairs of suckers greatly enlarge on third quaner of III and to decreasingdegree on II. I and IV. 10 In mature females of one species. T. mega!ops, glandular processes are present near base of midarm suckers on I-If!.

and F. Danrigal, Laboratory of Malacology, National Museum of Natural History, Paris; F. M. Climo, Mollusc Department, National Museum of New Zealand, Wellington; P. M. David and R. G. Aldred, Institute of Oceanographic Sciences, Wormley, England; A. Fleminger and H. G. Snyder. Inverte- brate Collection, Scripps Institution of Oceanography, La Jolla; R. Hanlon, Marine Biomedical In- stitute, University of Texas, Galveston; R. Kilias, Zoological Museum, Humboldt University, Berlin; J. Knudsen, University Zoological Museum, Copenhagen; G. Krefft and associates, WALTHER HER- WIG, Institut fUr Seelischeri, Hamburg; M. C. Mercer, Newfoundland Biological Station, St. John's; F. C. Naggs, Mollusca Section, British Museum of Natural History, London; K. M. Nesis, Institute of Oceanology, Moscow; W. G. Pearcy, Department of Oceanography, Oregon State University, Corvallis; W. F. Ponder, Department of Malacology, The Australian Museum, Sydney; M. A. Roe- leveld, South African Museum, Cape Town; C. F. E. Roper and M. J. Sweeney, Division of Mollusks, National Museum of Natural History, Smithsonian Institution, Washington; H. Terayama and T. Sato, University of Tokyo, Tokyo; G. Testa, Oceanographic Museum, Institute of Oceanography, Monaco; R. E. Young, Department of Oceanography, University of Hawaii, Honolulu; and the of- licials of the Museum of Natural History. Nice. I would like to thank my husband, Gilbert, for the numerous discussions during this study and for critically reading the manuscript. I would like to thank C. S. McSweeny for the accompanying illustrations. The support for this study by National Science Foundation grants GB-24994 and DEB77-13945 is gratefully acknowledged. This is a scientific contribution from the Rosenstiel School of Marine and Atmospheric Science of the University of Miami.

Note Added in Proof The recent revision of the cranchiids by M. J. Imber [The squid families Cran- chiidae and Gonatidae (Cephalopoda:Teuthoidea) in the New Zealand region. 408 BULLETIN OF MARINE SCIENCE, VOL. 30. NO.2. 1980

N .Z. J. Zool. 5: 445-484, 1978] is a sad piece of systematic work that reflects a total lack of knowledge of the family. Imber unwisely undertook the task when, studying the food of procellariiform seabirds in the New Zealand region, he found himself unable to specifically identify the numerous beaks of cranchiids that oc- curred, indicating the importance of cranchiids in the diet, particularly of Ptero- droma petrels. Working with inadequate material-larvae and juveniles from the New Zealand region and beaks primarily from his food studies, augmented by only 20 squid from other areas, which included but eight of the myriad of essential types-and the confusing literature, Imber distinguished 21 species, including 2 subspecies, belonging to 14 genera: Cranchiinae-Cranchia, Liocranchia and Leachia; Taoniinae- Taonius, Megalocranchia, Galiteuthis, H ensenioteuthis, Sandalops, Bathothauma, Helicocranchia, Teuthowenia, Fusocranchia, Meso- nychoteuthis and Egea. He considered all the genera to be monotypic except for Taonius, containing three species, Liocranchia, Leachia, Ga/iteuthis and He/i- cocranchia, containing two species each, and Teuthowenia, containing two species, including two subspecies. My study has revealed that the cranchiids are not a family of predominately circumglobal species as portrayed by Imber, and so considered by many early workers. This misconception is understandable for the latter who did not have access to large, world-wide collections. Imber has no such excuse. With no apparent hesitation, Imber synonymized such distinctly different species as Pyrgopsis atlantica Degner, 1925 from the North Atlantic, Leachia dislocata Young, 1972 and Drechselia danae Joubin, 1931, from the North and Equatorial Pacific respectively, with an unidentifiable species from off Madagas- car, Leachia eschscholtzii (Rathke, 1835). His decisions were based on the literature and six juveniles from a 4° x 9° area near New Zealand. His Galiteuthis is another example of indiscriminate lumping. Ignoring differences, Imber synonymized the clearly separate G. phyllura Berry, 1911 and G. pacifica (Robson, 1948) with G. armata Joubin, 1898. Again, without examining the types or knowing the animals, he synonymized Hoyle's (1886) distinct subantarctic "suhmi" form with the antarctic G. glacialis (Chun, 1906). Imber's Taonius is a confusion of misidentifications. Specimens from the literature identifiable to T. pavo (Lesueur, 1821) are assigned to three different species: (1) pavo is rightly recognized as valid from Lesueur's type; (2) T. pavo, Joubin, 1900 and 1920 (correct identifications if one examined the specimens) are placed into synonymy of his T. cymoctypus (de Rochebrune, 1884) (another mis- identification; de Rochebrune's poorly described, lost type is definitely not of this genus) and (3) Galiteuthis triluminosa Lu and Clark, 1974 (small juveniles of pavo) is synonymized with T. belone (Chun, 1906) (an unidentifiable species; Chun's small juvenile type is not extant). Other misidentifications abound. Pfeffer's Hensenioteuthis, which Imber resurrected, is based on an unidentifiable, distorted 7 mm ML larva whose deposition is unknown. Into the synonymy of his monotypic species, H. joubini Pfeffer, 1900, Imber relegated two distinctly different species, Megalocranchia papil/ata G. Voss, 1960 and Ascocranchia joubini G. Voss, 1962 together with Joubin's (1933) misidentified Teuthowenia megalops. All three rightly belong to Helicocranchia. The reinstatement of Fusocranchia Joubin, 1920 is another error arising from not examining the type. Joubin's unique larval specimen, which I examined at Monaco, unquestionably belongs to Liocranchia. From the conglomeration of species in the synonymy of his monotypic F. pellucida (Chun, 1910), it is obvious that Imber was misled by Nesis' (1974) erroneous identification of his type specimen of Vossoteuthis to Desmoteuthis pellucida Chun, 1910 and by Nesis' incIu- voss: GENERIC REVISION OF THE CRANCHIIDAE 409

sion of Megalocranchia pardus Berry, 1916 in Vossoteuthis. With a developmental series of larvae through subadults, I can show that the juvenile type of Vossoteuthis, which I have examined, belongs to Issei's (1908) long vanquished Liguriella, as also does Liguriella sp., Nesis, 1974. Both pellucida and pardus correctly belong to Teuthowenia. These are but the most glaring of the errors in this unfortunate work. How little Imber knows the cranchiids. In his closing paragraph, he states that it is "not likely that many, if any, new forms remain to be discovered." How wrong he is-to date, I can distinguish 42 species. I expect the number to increase.

LITERATURE CITED

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DATE ACCEPTED: February 22, 1979.

ADDRESS: Rosenstiel School of Marine and Atmospheric Science, University of Miami. 4600 Rick- enbacker Causeway, Miami, Florida 33149.