A JVIONOGRAPH OF THE CEPHALOPODA OF THE NORTH ATLANTIC 1. THE FAMILY LYCOTEUTHIDAE1 GILBERT L. VOSS lnstitute of Marine Science, University of Miami ABSTRACT The family Lycoteuthidae is revised on a world wide basis. An historical account is followed by a discussion of the anatomy, food, depth range and geographical distribution. The taxonomic status with full synonymy is given for each species including keys for differentiation. The family and subfamilies are redefined and new generic diagnoses are given along with type citations. Oregoniateuthis lorigera (Steenstrup, 1857) is described in full for the first time and Lycoteuthis diadema, Oregoniateuthis springeri, Selenoteuthis scintillans, Nematolampas regalis, and Lampadioteuthis me- galeia are described and illustrated. Leptodontoteuthis inermis Robson, 1926 is placed in the synonymy of Lycoteuthis diadema (Chun, 1900). The phylogenetic relationships of the Lycoteuthidae are discussed as well as the relationship of the genera within the family. The possibility is con- sidered, on the basis of their occurrence within two different genera, that the males of the Iycoteuthids possess paired, functional genitalia. INTRODUCTION The bathypelagic squids of the family Lycoteuthidae have held the interest of biologists since Chun first encountered the living animals in the nets of the VALDIVIA. His account of their spectacular light dis- plays has become a classic in the literature on bioluminescence. Unfor- tunately, adult specimens are rarely encountered. However, since 1900 when the first species was described, sixteen specimens of adult size have been taken and six genera erected for their disposal of which five are monotypic and three are represented solely by unique specimens. The addition in the last four years of two new genera and the acquisition of additional specimens, both larval and adult, of Lycoteuthis diadema have shed some new light upon the family. In the present revision the worldwide distribution of the family is considered and the keys and descriptions include the three Pacific members. The author wishes to express his thanks to the officials of the British Museum (Natural History) and in particular to Dr. W. J. Rees, formerly in charge of the mollusk collections, for permission to study the material of Lycoteuthis and the type of Leptodontoteuthis inermis Robson; to Dr. Jorgen Knudsen of the Zoological Museum, IContribution No. 381 from The Marine Laboratory, University of Miami. This study was supported by National Science Foundation grants G-5853 and G-17940. 1962J Voss: Lycoteuthidae 265 Copenhagen, for courtesies extended during his visit to that institution and the loan of the type of Oregoniateuthis lorigera (Steenstrup) and to the officials of the Zoologisches Museum, Berlin, and especially Dr. R. Kilias, Curator of Mollusks, for permission to study the types of Lycoteuthis diadema Chun at that institution and for the many kindnesses extended during the writer's visit. The writer is indebted to Lilly King Manning for the execution of the illustrations contained in this paper. This work has been supported by two grants from the National Science Foundation, G-5853 and G-17940, including a visit to the above mentioned museums during the summer of 1961. For this support the writer wishes to express his thanks. HISTORICAL RESUME The first mention of a lycoteuthid squid in the literature appeared in 1875 in a paper by the Danish teuthologist Iapetus Steenstrup. It consisted only of the statement that a squid in the museum at Copenhagen, Onychoteuthis (??) lorigera had the second pair of arms very elongate, a feature now known to be possessed by the genus Oregoniateuthis. Examination of this specimen by the writer has shown this species to be a true lycoteuthid, but at the time none of the extraordinary characters of this family were recognized. Twenty-five years later Carl Chun, in his general narrative of the Valdivia Deep Sea Expedition (1900, p. 532) illustrated and gave a brief description of a squid taken off the Bouvet Islands, which he called Enoploteuthis diadema. His account of the colors emitted by this specimen is as follows. "Unter allem, was uns die Tiefseetiere an wundervoller Hirbung darbieten, Hisst sich nichts auch nur annahernd vergleichen mit dem Kolorit dieser Organe. Man glaubte, dass der Karper mit einem Diadem bunter Edelsteine besetzt sei; das mittelste der Augenorgane glanzte ultramarinblau und die seitlichen wiesen Perlmutterglanz auf; von den Organen auf der Bauchseite erstrahlten die vorderen in rubinroten Glanze, wahrend die hinteren schneeweiss oder perlmutter- tarben waren mit Ausnahme des mittelsten, das einen himmelblauen Ton aufwies. Es war eine Pracht." This species was characterized by Chun by having 24 light organs of large size, distributed in the tentacular stalk, on the ventral peri- phery of the eyeball and within the mantle cavity. The presence of these organs in nearly the same numbers and positions throughout 266 Bulletin of Marine Science of the Gulf and Caribbean [12(2) the members of the family is perhaps the most distinctive feature of the Lycoteuthidae. Chun placed his species in the family Enoplo- teuthidae. In the same year Pfeffer in his synopsis of the oegopsid squids erected the genus Lycoteuthis (1900, p. 156) which he included in the family Onychoteuthidae. He diagnosed it as follows. "Endspitze des Gladius ganz kurz und dick, komprimiert, Loffel sehr gross, Kiel nicht durch die Rilckenhaut hindurch sichtbar; Schiltzsaume der Arme mit stark ausgebildeten Querbrilcken; Tentakel mit vier Reihen von Haken; Buccalhaut dunkel gefarbt, mit acht Zipfeln und Heftung- en und nur zwei Poren." An examination of the characters listed shows, surprisingly, that none of them are valid criteria for the genus as it is recognized today. In addition, on page 161 under the genus Lycoteuthis a single species is given, Lycoteuthis Jattai, characterized by the presumed presence of hooks on the tentacular clubs, although they were lacking in the specimens before the author. In 1903 Chun (p. 569) mentioned diadema again and placed it in Pfeffer's genus Lycoteuthis along with jattai but later in 1903 he considered on the basis of the light organs that diadema was generic- ally distinct and placed it in a new genus Thaumatolampas, for which he erected a new family, the Thaumatolampadidae (p. 68). Pfeffer (1908) decided that his L. jattai was conspecific with Chun's diadema but considered that Lycoteuthis had priority over Thaumatolampas and retained the genus with its sole species diadema in the Onychoteuthidae but in a new subfamily, Lycoteuthinae. In 1910 appeared ehun's great monograph of the oegopsid squids and the lengthy descriptions of diadema with its accompanying histo- logical and anatomical plates. In this work he lowered the family Thaumatolampadidae to subfamily rank and placed it in the family Enoploteuthidae. He included a synopsis of Pfeffer's 1900 characters, pointed out the many inaccuracies and included jattai as an uncertain specIes. It was inevitable that these two workers should disagree on this question. Two years later (1912) Pfeffer brought out his own monograph of the oegopsids based on the Plankton Expedition's collections. In this he retained the subfamily Lycoteuthinae in the Onychoteuthidae and gave a more detailed description of the original specimens of L. jattai from the Hamburg and Strasburg museums. This showed that in all essential details jattai and diadema were conspecific 1962] Voss: Lycoteuthidae 267 and the Hamburg specimen was illustrated in detail. This specimen, shown in detail in plate 14, figs. 1-9, must be considered as the type of L. jattai. Unfortunately, the specimen was destroyed in the World War II bombing of Hamburg and the illustration, plate 14 figure 4 is here selected as the type illustration of this species. However, Pfeffer himself placed his species jattai in the synonymy of diadema. It thus appears that the genus Lycoteuthis is the valid genus since despite the erroneous diagnosis it was protected by a type species and a type, the latter of which, although no longer extant, was well described and figured in 1912. It would be difficult to prove at this date whether Chun's or Pfeffer's work appeared first in 1900, but the present writer feels that Pfeffer's submerging of his own species should stand and the matter thus settled for the sake of stability. The next mention of a lycoteuthid in the literature was in a paper by Berry (1913) containing the description of a remarkable new cephalopod from the Kermadec Islands, Nematolampas regalis. In a fuller description published in 1914 he removed the lycoteuthids from the family Enoploteuthidae and erected the family Lycoteuthidae for them. He considered that while diadem a had priority over jattai, Lycoteuthis definitely had priority over Thaumatolampas. From 1914 to date, the names Thaumatolampadidae and Thaumatolampadinae have not reappeared in the literature. For nearly 50 years common usage has maintained the names Lycoteuthidae and Lycoteuthinae and the name Thaumatolampadidae should not be retained in the literature, whatever its claims for priority may be. In 1916 Berry again published on Kermadec Island cephalopods, illustrating the gladius and sucker of N. regalis and describing a new genus of lycoteuthid, Lampadioteuthis megaleia. For this genus he formed a new family, the Lampadioteuthidae which he considered fell between the Lycoteuthidae and the Enoploteuthidae. Naef (1923) included the Ly.coteuthinae as a subfamily of the Enoploteuthidae in his monographic work on the Mediterranean cephalopods. The next find of a lycoteuthid was reported by Robson (1924, 1924a) who listed a badly mutilated specimen from off South Africa and later (1926) described a new genus and species Leptodonto- teuthis inermis from the same locality. In 1922 Grimpe published his review of the European cephalopods in which he retained the lycoteuthids in the Enoploteuthinae and maintained the subfamily Lampadioteuthinae. Thiele (1934) retained 268 Bulletin of Marine Science of the Gulf and Caribbean [J2(2) the subfamilies but separated them from the Enoploteuthidae using Berry's Lycoteuthidae.