A Monograph of the Cephalopoda of the North Atlantic

Home , Lampadioteuthis, Lycoteuthidae, Lycoteuthis, Selenoteuthis

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 megaleia 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 considered, 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 periphery 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 histological 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. A larval specimen 5.0 mm in mantle length was taken by the ARCTURUS expedition and reported on by Robson (1948). He considered the genus and species to be uncertain and listed it in the Lycoteuthinae which for some reason he made a subfamily of the Onychoteuthidae, following Pfeffer's long rejected scheme. With the beginning of exploratory trawling in the Western Atlantic in the 1950s, several new records and genera and species have come to light. In 1956 Voss reported upon the cephalopods of the Gulf of Mexico, recording three new specimens of L. diadema and a new genus and species, Oregoniateuthis springeri, utilizing the family Lycoteuthidae. Later, in 1958 he described another new genus and species, Selenoteuthis scintillans from the Florida Current and showed that Pfeffer's larval form, Asthenoteuthion planctonicum described in 1912 as an enoploteuthid is actually in the developmental line of Lycoteuthis diadema. Over the years the lycoteuthids have been classic examples of marine luminescent organisms and the names have been utilized over and over in the literature on bioluminescence. In nearly all cases Lycoteuthis diadema is the species referred to and the name is firmly established in the popular literature also. Citations in these fields are not considered here. In 1956 Sir Alister Hardy in his book "The Open Sea, the WorId of Plankton," gives several references to Thaumatolampas (Lyco- teuthis) diadema. While this work is not for taxonomic purposes it is perhaps well to note that the use of the name Lycoteuthis in paren- thesis is not intended as subgeneric but is used in the sense of equal to, without the insertion of the sign.

GENERAL MORPHOLOGY The lycoteuthids are upper bathypelagic squids with strong muscu- lar mantles indicating that they are active swimmers which perhaps explains their paucity in trawl and plankton catches. The mantle is in general .cylindrical and blunt posteriorly except in Oregoniateuthis springeri in which the fins and mantle are drawn out into a long slender tail. The fins in all of the species are broader than long (discounting the tail in springeri) and rather sharply pointed laterally. The head, damaged in most specimens known, is large, rounded somewhat laterally, with normal eyes and moderate size eye openings, 1962] Voss: Lycoteuthidae 269 the lids usually with a small anterior sinus. The arms are strong, slightly compressed, usually with a keel which may be deep on the ventro-lateral arms. The arm order is usually 3.2.4.1 but in this group a peculiar modification may occur of a supposed generic nature, in which either the ventro-lateral (Nematolampas) or the dorso-lateral arms (Oregoniateuthis) are greatly lengthened and bear numerous light organs. The arm suckers are in two rows (except on the sharply modified section of the elongate arms in Nematolampas and Oregoniateuthis) and none are modified into hooks. All of the arms bear trabeculate protective membranes of which the ventral membrane on the ventro- lateral arms may be very deep. The tentacles are of moderate length with expanded clubs. The carpal cluster is distinct. The manus or hand bears four rows of large suckers bordered by trabeculate protective membranes. The suckers of the dactylus are also in four rows but are smaller and more crowded but in regular rows. The buccal membrane has eight lappets with eight supports and eight attachment straps. Chun (1910) states that there are six aquiferous pores but Pfeffer (1912) insists that there are only two. My own material was not suitable to settle this question but the problem is of little importance in the considerations before us. In all of the specimens which I have examined the buccal membrane is deeply pigmented, possessing a purplish color. The beaks are somewhat onychoteuthid in appearance with a long thin rostrum but present no particular characteristic features. The radula is not distinctive. The gladius, except in Lampadioteuthis, is distinctive. It is rather narrow and darkly pigmented. It is widest about in the middle of the gladius but then is constricted and strongly laterally compressed. The posterior end widens into a spoon-shaped conus. In Lampadioteuthis the posterior constriction and conus do not appear and the gladius is typically enoploteuthid. However, the young of Lycoteuthis also have an enoploteuthid-like gladius and it does not attain the adult shape until about half grown. Lampadioteuthis, rather than being more closely related to the enoploteuthids as Berry has suggested, may merely show somewhat larval characteristics as far as the gladius is concerned. The internal anatomy of no species has been sufficiently worked out at present, except for Chun's (1910) observations and illustra- 270 Bulletin of Marine Science of the Gulf and Caribbean [12(2) tions, to permit generalizations. The only known males are the two specimens of Oregoniateuthis springeri and the type of Selenoteuthis scintillans. In both of these genera the male genitalia are paired and both are functional. This is an unusual condition in the Oegopsida and at the present must be supposed to be characteristic of the Lycoteuthidae. The female genitalia appear normal and two nida- mental glands are present. The most distinctive morphological feature to be found in the lycoteuthids is the presence and arrangement of light organs. To our knowledge, no other squids present such a spectacular array of these organs with the possible exception of the histioteuthids and certain species of Abralia, but even these do not present the array and diversification of colors that the lycoteuthids possess. Chun (1910, text figs. 18 & 19, plate 4, figs. 1-22) has illustrated these organs in detail as to their finer structure and given a wealth of description (l.c., pp. 68-77). Seven types of light organs are found in the family, but not all of them need be found in anyone species. These may be listed as follows: surface light organs found in the skin on the mantle, the head, and on certain arms (Oregoniateuthis and Nematolampas); on the arm tips (males of Selenoteuthis, and in Nematolampas); at the posterior end of the mantle (Selenoteuthis); a series of elongate light organs deeply buried in the slender tail (Oregoniateuthis); in tentacle stalks in all genera and at the base of the tentacle in Lampa- dioteuthis; on the eyeball in all genera; and a set pattern within the

TABLE 1 NUMBER AND POSITION OF LIGHT ORGANS IN THE LYCOTEUTHIDAE diadema springeri lorigera scintillans regalis megaleia Eyes 5/5 5/5 5/5 ? 5/5 5/5 3-113-1 Tentacles 2/2 212 5/5 2/2 2/2 4/4 Base of tent. 1/1 Anal 2 2 2 2 2 2 Abdominal 3 3 3 3 3 Branchial 2 2 2 2 2 2 Post. Abdom. 3 1-3 1 ? 3 ? 3 ? Terminal 1 2 Tip of arms II, III I, II Integ. arms 50+ 5+ Integument 25 60- Tail 8 Total 24 105+ 33+ 29 92 23 1962] Voss: Lycoteuthidae 271 mantle cavity in all except Lampadioteuthis in which certain ones are omitted. The arrangement and numbers of light organs in the known species are given in Table 1, which is a revision of Berry's (1916) table. The basic pattern of the internal light organs is very characteristic and finds no deviation except in Lampadioteuthis which represents, in my opinion, a divergent offshoot of the family. Some confusion has resulted from the structure of the posterior abdominal photophore. This is discussed in detail under Lycoteuthis diadema. Various names have been used for the internal light organs. In this paper they are defined as follows. ANAL-The two round light organs lying one on each side of the rectum near its opening and just within the funnel. ABDOMINAL-The central three light organs lying on the visceral mass between the gills. The central one is usually round or longitudinally oval, the laterals oval and oblique. BRANCHIAL-The round light organs near the base of each gill and together with the abdominals forming a band of organs across the middle of the viscera. POSTERO-ABDOMINAL-Eitherone or a complex of three light organs posterior to the middle of the viscera and fastened not to the visceral sac but to the inner wall of the mantle. The positions of all of these internal light organs are shown in Figure 1. One of the interesting finds of this study was the sexual demorphism in Selenoteuthis. The male of this species possesses a quite large round black light organ at the tip of both the dorso- and ventra-lateral arms. When the females were examined these light organs were absent. This led to speculation that other details might show sexual dimorphism such as the elongate arms of Oregoniateuthis which might be the male of Lycoteuthis, still unknown. This, however, seemed illogical and with the finding of the new species of this genus represented by a female, was shown not to be the case.

FOOD The stomachs of all of the specimens available in the collections at Miami were opened for an examination of the contents in order to learn something of their food habits. The three specimens of L. 272 Bulletin of Marine Science at the Gulf and Caribbean [12(2)

ana.l photo

bra.I'lc.h iQ./ photo fabdomina.1 photo

pO$tero -a.bdom. photo

FIGURE 1. Ventral view of Lycoteuthis diadema with mantle cut away to show location of internal light organs. diadema contained fine remains of small fish, in one case a myctophid, and Crustacea possessing luminescent organs. O. springeri had crustacean remains among which were numerous photophores possibly those of euphausiids or sergestids. The large specimen of O. lorigera contained fish scales, two large fish eyes and fragments of the gladius and beaks of a squid. The beaks, however, were fragmentary and not identifiable. The three specimens of Selenoteuthis had only unidenti- fiable remains. It seems from the above scanty evidence that like most oceanic squids, the diet of the lycoteuthids consists of other small bathypelagic organisms, primarily fish and shrimp, although they do not hesitate to eat other squids and may even be cannibalistic. 1962] Voss: Lycoteuthidae 273

DEPTH DISTRIBUTION Despite the depth records given in the literature, there is little doubt that the members of the family are confined to the upper bathypelagic region. Unfortunately most of the records are from open nets and all too often the depth of capture is assumed to be the greatest depth of the net or the trawling depth, rather than the possibility that capture was achieved on the way up or down. There is much literature on this and the writer is well versed on the subject. However, in the present case the evidence is on the side of the shallower depths. As shown above the morphology indicates a habitat in the shallower depths. The actual depths as given in the literature and from the trawl data are given below according to species. Lycoteuthis diadema-Adults: 366 m, 366 m, 403 m, 589 m, 0-1500 m, 0-3000 m, 2195 m. Larvae: 46 m, 57 m. Oregoniateuthis springeri-367 m. O. lorigera-no depth data. Nematolampas regalis-on beach. Selenoteuthis scintillans-46 m, 3290 m. Lampadioteuthis megaleia-on beach. In the above data the depth given is the depth fished for each record. In those listed as 0-, the hauls were vertical tows from the greatest depth indicated to the surface. Thus the shallow tows are the only ones of significance and these indicate a habitat from about 300-600 m or that area described by Bruun as the mesopelagic characterized by a large number of animals with a great variety of luminous organs (Bruun, 1957, p. 643). Although the data is insignificant, there is a suggestion from the depth of capture of the larvae (46-57 m) that in common with so many mesopelagic animals the larvae are found near the surface, moving downward with age.

KEY TO THE SUBFAMILIES, GENERA AND SPECIES OF THE LYCOTEUTHIDAE 1. Five light organs in a single line on the ventral periphery of the eyeball ...... " Subfamily Lycoteuthinae 2 Four light organs on the eyeball, 3 ventral and 1 lateral ...... Subfamily Lampadioteuthinae Lampadioteuthis megaleia 2. Arms subequal in length; no integumentary light organs 3 Either second or third arms greatly elongate; integumentary light organs present 4 274 Bulletin of Marine Science of the Gulf and Caribbean [12(2) 3. A terminal posterior light organ present; terminal light organs at tip of second and third arms in males, absent in females .. Selenoteuthis scintillans No terminal light organs on mantle; no terminal light organs on tips of arms Lycoteuthis diadema 4. Third arms greatly elongate with numerous light organs; tip of body not elongate into tail Nematolampas regalis Second arms greatly elongate , 5 5. Posterior end of body drawn out into a tail; two large light organs in tentacular stalks; tips of third arms normal Oregoniateuthis springeri Posterior end of body not forming a tail; two large and three small light organs in tentacular stalks; tips of third arms drawn out, whip-like, smooth, without suckers Oregoniateuthis 'origera

Order TEUTHOIDEA Suborder OEGOPSIDA Family LYCOTEUTHIDAE Berry, 1914 Thaumatolampadidae Chun, 1903, p. 68. Lycoteuthidae Berry, 1914, p. 140. Animals small to large; buccal membrane with eight lappets and supports; suckers unmodified, in two rows on arms, in four rows on tentacular club; light organs in tentacular stalks, on the eyeball and within the mantle cavity, occasionally dermally; fins short, wide and pointed laterally, with a tail in Oregoniateuthis springeri; male genitalia paired (Oregoniateuthis, Selenoteuthis). Type genus.-Lycoteuthis Pfeffer, 1900.

Subfamily Lycoteuthinae Pfeffer, 1908 Lycoteuthinae Pfeffer, 1908, p. 294. Gladius slender, distinctly concave, constricted behind midpoint, spoonshaped at posterior end. Five light organs on ventral border of each eye, two or more in tentacular stalk and eight or ten in the mantle cavity. Genus Lycoteuthis Pfeffer, 1900 Enoploteuthis, Chun, 1900, p. 532. Lycoteuthis Pfeffer, 1900, p. 161. Thaumatolampas Chun, 1903, p. 67. Arms subequal in length, no external light organs except on tentacular stalk. Type species.-By monotypy, Lycoteuthis Jattai Pfeffer, 1900 (=Eno- ploteuthis diadema ehun, 1900). 1962] Voss: Lycoteuthidae 275

FIGURE 2. Lycoteuthis diadema (Chun, 1900) ML 65.0 mm. a. ventral view; b. dorsal view. Lycoteuthis diadema (Chun, 1900) FIGURES 2-5 Enoploteuthis diadem a Chun, 1900, p. 532. Lycoteuthis Jattai Pfeffer, 1900, p. 161. Thaumatolampas diadema, Chun, 1903, p. 67. Lycoteuthis diadema, Chun, 1903, p. 569. - Pfeffer, 1908, p. 294; 1912, p. 114. - Thiele, 1934, p. 961. - Voss, 1956, p. 118; 1958, p. 374. Asthenoteuthion planctonicum Pfeffer, 1912, p. 172. - Voss, 1958, p. 374. ?Lycoteuthis sp. A. Robson, 1924, p. 2. ?Lycoteuthis sp. Robson, 1924a, p. 599. Leptodontoteuthis inermis Robson, 1926, p. 2. Material examined.-Types of Enoploteuthis diadema, 1 ~ , ML 30.0 mm, VALDIVIA Sta. 89, 31°21'S, 15°58'E, 0-3000 m-I ~,ML 21.5 276 Bulletin of Marine Science of the Gulf and Caribbean (12(2~

FIGURE 3. Lycoteuthis diadema (Chun, 1900) ML 65.0 mm. a. funnel organ; b. left tentacular club; c. gladius, dorsal view; d. gladius, side view of posterior end; e. upper mandible; f. lower mandible. mm, VALDIVIASta. 118, 40031'S, 15°06'E, 0-1500 m all in the Zoo- logisches Museum, Berlin.-Holotype of Leptodontoteuthis inermis BMI925.9.5.10, Sta. 522, Cape, (Cape of Good Hope), 2195 m, otter trawl, Globigerina ooze, Survey No. V4639.-1 <;?, ML 65.0 mm, OREGON Sta. 481, 28°57'N, 88°40.5'W, Sept. 7, 1951, UMML 31.174. -1 <;?, ML 83.0 mm, OREGON Sta. 321, 29°27'N, 87°19'W, 403 m, bottom temp. 10°C, April 28, 1951.-1 <;? ML 51.0 mm, OREGON Sta. 382, 29°11.5'N, 88°07.5'W, 366 m, June 21, 1951. 1962] Voss: Lycoteuthidae 277 -2 ~ ~, ML 50.0-59.0 mm, SILVER BAY Sta. 1198, 24°11'N, 83°31'W, 366 m, June 9, 1959, UMML 31.228.-1 spec., Lyco- teuthis sp., Natal, Gilchrist Sta. 156, BM 1924.9.9.26a, 589 m. Larvae: 2 sex indet., ML 8.3 and 7.2 mm from 26°35'N, 79°28'W in 57 m, Feb. 7, 1954, UMML 31.194.-1 sex indet., ML 8.7 mm, from 20012'N, 65°19'W in 46 m, Feb. 25, 1954, UMML 31.194.

FIGURE 4. Lycoteuthis diadema (Chun, 1900) ML 65.0 mm. a-b. largest suckers of second right arm; cod. largest suckers of third right arm. 278 Bulletin of Marine Science of the Gulf and Caribbean [12(2)

FIGURE 5. Lycoteuthis diadema (Chun, ]900) Asthenoteuthion stages. a. larva of 8.0 mm mantle length; b. larva of 7.0 mm mantle length. 1962] Voss: Lycoteuthidae 279 Description.-Mantle moderately stout, cylindrical, pointed posteriorly; anterior margin slightly produced in dorso-median line; mantle width about 1/3 of length. Fins moderately long, very broad, triangular, terminal in position. Head small, compact; eyes large with five round light organs on ventral periphery. Funnel short and stout, free for about 1/3 of length; funnel organ broad, inverted V-shaped with rounded lateral arms; ventral pads oval, thick and prominent. Arms in order 2.3.4= 1 with dorsal and ventral pairs about equal in length, only slightly shorter than lateral. Third pair bordered by broad swimming membrane which extends full length of arm. Suckers in two rows throughout, ventral suckers bordered by slender protective membrane with strong supports. Suckers small with chitinous rings which bear numerous teeth, those on the distal half long, slender, sharp, curving, those of the proximal half short, truncate. Tentacles moderately long, round in cross-section. Clubs with four rows of suckers which are large on hand, gradually decreasing distally. Suckers with about eight or nine long, slender, blunt teeth on distal margin, proximal portion smooth.

TABLE 2 MEASUREMENTS(IN MM) OF SIXFEMALESOF Lycoteuthis diadema (CHUN) Station Lycoteuthis OREGON SILVERBAY VALDIVIA (Robson, 1924a) 481 1195 89 118 ----- Mantle length 89.0 65.0 59.0 50.0 30.0 21.5 Mantle width 28.0 20.0 20.0 15.0 12.0 10.5 Head width 13.0 11.5 Fin length 42.0 35.0 29.0 23.0 14.0 11.0 Fin width 78.0 57.0 41.0 40.0 28.0 22.0 Arm length I 34.3 26.0 24.0 19.0 12.0 10.0 II 34.0 33.5 26.0 23.0 14.5 12.5 III 31.0 24.0 23.5 12.5 10.5 IV 36 28.0 28.0 21.0 13.0 10.0 Tentacle length 64.0 64.0 20.0 Club length 17.5 14.0

Two light organs embedded in each tentacular stalk, one near base, one about midway of stalk. Five large light organs on ventral periphery of eyeball; central one dark purplish blue, slightly granular, other four smooth, reddish. Ten large light organs within mantle 280 Bulletin of Marine Science of the Gulf and Caribbean [12(2) TABLE 3 INDICES OF BODILY PROPORTIONS OF Lycoteuthis diadema (CHUN) ------Station Lycoteuthis OREGON SILVER BAY VALDIVIA (Robson, 1924a) 481 1195 89 118 ------Mantle length 89.0 65.0 59.0 50.0 30.0 21.5 MWI 31.5 30.8 33.9 30.0 40.0 48.9 HWI 43.3 53.3 FLI 47.1 53.9 49.2 46.0 46.6 51.2 FWI 87.6 87.7 69.5 80.0 93.4 102.0 ALI I 38.5 40.0 40.6 38.0 40.0 46.5 II 38.2+ 51.5 44.1 46.0 48.3 58.2 III 47.7 40.7 47.0 4].7 48.4 IV 40.4 43.2 47.5 42.0 43.4 46.5 cavity: single reddish photophore on each side of anus, five across middle of mantle cavity consisting of one near base of each gill and three, two large, one small one transversely across the midline; three large light organs in line across viscera near posterior end, closely fused together appearing as one elongate light organ. Gladius strongly formed, rachis narrow anteriorly, broad in mid-section and tapering to posterior quarter where it again broadens into spoon-shaped area at end. Type.-Zoologisches Museum, Berlin. Type locality.-West wind drift, 40031'S, l5°6'W, vertical net to 1500 meters. Discussion.-Lycoteuthis diadema was described by Chun (1910) in his popular narrative of the VALDIVIAexpedition as Enoploteuthis diadema. Pfeffer, in the same year described Lycoteuthis jattai based upon disfigured specimens from dolphin stomachs into which he read some characters which were non-existent. These specimens were destroyed at Hamburg during World War II. While his generic diagnosis is inaccurate, his specimens are clearly diadema, and his genus is accepted as the valid one though Chun in 1903 erected the genus Thaumatolampas for his own species, pointing out that Pfeffer's diagnosis was erroneous. Asthenoteuthion planctonicum was described by Pfeffer (1912) who considered that these young larval forms were early stages of A bralia, basing his opinion upon the enoploteuthid-like gladius which they possess. Voss (1958) has demonstrated through the development 1962] Voss: Lycoteuthidae 281 of the light organs and their arrangement, and the changes in the gladius that they belong to the developmental cycle of L. diadema and the name must be placed in the synonymy of the latter. In 1924 Robson reported briefly upon a mutilated lycoteuthid from off South Africa which he designated Lycoteuthis sp.A. Later, in 1926 he described Leptodontoteuthis inermis also from off the Cape, based upon a detached head and arms. He apparently considered his earlier sp. A to be the same species. He separated Leptodontoteuthis from Lycoteuthis on the basis of a somewhat different radula, a different distribution of the light organs on the eyeball, strongly compressed arms and the different shape of the mandibles. The present writer examined the type at the British Museum and found a number of discrepancies between Robson's description and the actual specimen. Firstly, the arms do not appear to be more compressed than in diadema, secondly a close examination of the rather mangled eyes revealed only five light organs on the ventral aspect of the right eyeball instead of seven as shown by Robson and eight as given in his description, while the left eyeball had only fragments, and thirdly, the mandibles do disagree with those figured by Chun (1910, pI. 3, figs. 3-5) and are much closer to those from the Gulf of Mexico specimens. The latter also differ somewhat strongly from Chun's in shape and pigmentation, but other characters found in the specimens are the same. As a result of the detailed examination, no characters could be found that warranted separation of the species and hence Robson's species must be placed in synonymy. I also consider his Lycoteuthis sp. A to be L. diadema. Some discrepancies occur in the various descriptions, concerning the number of light organs within the mantle cavity. Chun (1910) states that there are eight light organs within the mantle cavity whereas Pfeffer (1912) counted ten. Thiele (1934) gives eight in his diagnosis of the Lycoteuthinae. The cause of these discrepancies is the structure of the posterior abdominal light organ or organs. Chun has shown this as one long transverse organ. Pfeffer indicates it as three organs closely appressed and Voss (1956) also found this condition. In Nematolampas, Berry (1913) shows an elongate transverse organ but both the figure and the description give no hint as to whether it was partly divided or not. In Selenoteuthis Voss (1958), there is a single large luminous patch. This is distinctly three parted with three separate lenses. In Oregoniateuthis springeri Voss (1956), these organs are in three parts but in O. lorigera Steenstrup they are fused into one long 282 Bulletin of Marine Science of the Gulf and Caribbean [12(2) oval band with no trace of three elements. It seems likely that this set of organs has a tendency toward fusion both within the species and the subfamily. Much of this confusion concerning the light organs probably originated from Chun's figures, so beautifully and apparently accur- atelyexecuted (1910, pI. 2). Actually one wonders on close examination how Chun could have been so confused and inaccurate. In figures 2 and 5 the anal organs are shown as being attached to the dorsal wall of the funnel. Actually they are embedded in the ventral surface of the fleshy wall containing the rectum, oesophagus and associated organs. The posterior abdominal organs are illustrated in figure 3 and are shown attached to the viscera when actually they are firmly attached to the inner surface of the mantle wall and form a depression only in the viscera. This point has been missed by nearly all who have worked with the family. In life the mantle is transparent and the orientation of the light organ is inconsequential. If the posterior abdominal light organs became separated from the mantle along with the oval ridge or pad on which they lay, they could be illustrated in the above fashion and considered as one organ. If they are fused together as in Selenoteuthis, the three major elements are probably still present. The types were examined in Berlin. Both are badly damaged by dissection and parts are missing for histological study so that some characters could not be checked. The anal organs are nearly perfectly round, not oval. The abdominal light organs are fastened to the mantle wall, not the viscera and are slightly three parted. The funnel organ illustrated by Chun (figure 4) is semidiagrammatic; the organ is not so strongly shouldered, the limbs are thinner, and the posterior cleft is deeper. In view of these discrepancies, I think there is no question of the identicalness of the other specimens with Chun's species. For a detailed critique of Robson's Leptodontoteuthis see the present writer's South African Cephalopods (Voss, in press). Remarks.-Apparently only eight adult specimens of this species are recorded in the literature of which all but two were taken from the stomachs of marine vertebrates. The vertical distribution is apparently restricted to the upper bathypelagic from near the surface to about 500 meters (see earlier discussion of depth). Distribution.-Gulf of Mexico and Florida Current (Voss); West 1962] Voss: Lycoteuthidae 283 coast of South America, Indian Ocean in 46°S, 1200E, Atlantic Ocean (all Pfeffer); Benguela Current in 31°21'S, 15°58'E, West Wind Drift in 40031'S, 15°6'E (all Chun); South Africa, Cape Marine Province (Robson). Genus Oregoniateuthis Voss, 1956 Oregoniateuthis Voss, 1956, p. 120. Lycoteuthids with greatly elongate dorso-Iateral arms; 8-10 light organs within mantle, 5 on each eye or more (unknown in lorigera), 2 or more on each tentacular stalk and few to many on mantle, head and arms; posterior end of mantle drawn out into long tail in springeri, not so much in lorigera, male testis and penis paired and both functional in springeri. Type species.-By original designation, Oregoniateuthis springeri Voss, 1956. Oregoniateuthis springeri Voss, 1956 FIGURES 6 - 8 Oregoniateuthis springeri Voss, 1956, p. 120, figs. 7b-f. Material examined.- ~ , holotype, mantle length 80.0 mm, OREGON Sta. 382, 29°11.5'N, 88°07.5'W, in 367 meters, June 21, 1951. USNM 575090.- ~, mantle length 97.0 mm, OREGON Sta. 3296, 28°36'N, 89°48'W, 244-520 fms, August 21, 1961, UMML 31.376. Description.-Mantle slim, cylindrical, tapering posteriorly to long, slender tail; anterior median dorsal margin slightly produced; ventral margin straight. Fins wide, pointed laterally, terminating indistinctly posteriorly in well defined ridge which extends to distal end of tail. Funnel small, compact; mantle locking apparatus with simple straight funnel member and corresponding slightly raised ridge on inner surface of mantle. Dorsal funnel organ member broadly and sharply shouldered with short anterior ridge on each side of apex. ventral pads small. Head moderate size with large eyes; eyelids widely open, semicir- cular, posterior margin straight, anterior margin rounded with clearly defined sinus. Head connected with mantle dorsally by simple straight nuchal locking cartilage. Funnel groove deep and clearly defined; three nuchal folds of which second lateral is olfactory organ. Arms, with exception of second pair, moderately long, order 2.3.4.1. First pair shortest with slight ridge on dorsal surface. Second pair longest, longer than mantle, rounded for most of length, distally 284 Bulletin of Marine Science of the Gulf and Caribbean [12(2)

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FIGURE 6. Oregoniateuthis springeri Voss, 1956. ML 97.0 mm, male. a. dorsal view; b. ventral view showing internal light organs; c. funnel organ; d. left eyeball with photophores; e. right tentacular club. flattened and broadened; suckers moderate size, stalked, in 2 rows proximally, about 1/4 length of arm upper row disappears followed at about 1/3 of total length by disappearance of ventral row, replaced by small pointed papillae which disappear toward end of arm. Suckers bordered dorsally by narrow protective membrane with supports which disappear at distal fourth of arm, ventrally by broad membrane 1962] Voss: Lycoteuthidae 285 with prominent large supports which in distal third becomes very broad with slender supports which in preservation appear as rugose ridges. Sixteen widely spaced round light organs embedded in dermal layer, becoming somewhat crowded distally, terminating in small light organ near tip. Arms of third pair slightly longer than first, broad and bordered laterally by strong swimming membrane. Suckers in two rows, rings bearing 22-25 sharp teeth longest on distal border, with small protective membrane dorsally and greatly expanded one ventrally, both with prominent supports. Nine large light organs present in dorsal side of arm, decreasing in size distally with a bare area near distal third of arm. Fourth arms normal with slight skin fold ventrally, suckers protected by 2 low membranes with supports. Tentacles missing in holotype. In specimen from OREGON Sta. 3296 tentacles nearly as long as mantle, moderately stout and nearly round in cross section, with two large light organs deeply buried in stalk,

FIGURE 7. Oregoniateuthis springeri Voss, 1956. a-b. large suckers of right tentacular club; c. large sucker from basal portion of second right arm; d-e. large suckers from near basal se<::tionof third right arm. 286 Bulletin of Marine Science of the Gulf and Caribbean [12(2) TABLE 4 MEASUREMENTS (IN MM) OF TWO MALES OF Oregoniateuthis springe>ri VOSS, 1956 Type Mantle length 80.0 97.0 Mantle width 20.5 20.0 Head width 18.5 18.5 Fin length 48.0 60.0 Fin width 53.0 54.0 Arm length I 28.0 31.0 II 90.0 75.0 III 32.0 31.5 IV 29.0 34.5 Tentacle length 87.0 Club length 21.2

1 near base, 1 near club. Club compact, slightly expanded, bordered on both margins by trabeculate protective membrane; aboral surface with large well developed swimming keel extending over two thirds length of club. Carpal cluster consists of four to five smooth ringed suckers and four to five large white pads. Manus with four rows of large suckers about equal in size with horny rings equipped with about 18 teeth, those of the distal half long, slender and sharp, proximal teeth small, triangular. Dactylus with small suckers in four orderly rows. Eyes large, prominent, with 5 large light organs of equal size on ventral periphery; middle organ much darker than others, with purple sheen. Buccal membrane unpigmented with exception of few scattered chromatophores; 8 lappets and 8 supports attached dorsally to arms I, II and IV, ventrally to III; oral surface thickly covered with fleshy overlapping papillae. Outer intergument missing over much of mantle and fins. Head with 8 light organs; 1 large one in front of each near base of third arms; 1 small light organ at upper corner of eyelid; I large one on lower corner of eyelid, and a pair of light organs, one on either side of posterior part of head. Mantle light organs few, arranged as follows: 1 on each side of anterior edl!e of mantle, 1 on each side of midline half way between anterior edge of mantle and fin insertion, 1 on each side of mantle just anterior to and below lobes of fin, 3 light organs in a diagonal row beneath fins of which 1 at midpoint of fins is round and large, 1 on each side of central one slender, oval, pointed; 1 pair dorsally 1962J Voss: Lycoteuthidae 287 just posterior to insertion of fin lobes, 1 large median photophore near posterior end of fin in midline, and 1 pair of closely set organs near midpoint of tail. In addition, 8 long slender rod-like organs deeply buried in dorsal midline of tail terminating in a tear drop shaped organ at distal extremity. A total of 105 light organs on body, eyes, in mantle, on head, arms, tentacles and in tail. The mandibles and radula were not dissected out. The gladius is surprisingly short, extending posteriorly only to point of convergence of posterior edge of fins. Gladius typically lycoteuthid, broadened slightly in anterior half, narrowing in posterior

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FIGURE 8. Oregoniateuthis springeri Voss, 1956. a. upper mandible; b. lower mandible; c. gladius; d. side view of posterior tip of gladius; e. Si'ermatophore, length 7.3 mm. 288 Bulletin of Marine Science of the Gulf and Caribbean [12(2) third, forming spoon-shaped end posteriorly. Internal anatomy not dissected out but both specimens have well developed penis, Needham's sac and testis on each side of similar size and shape and filled with ripe spermatophores. Holotype.-U.S. National Museum. Type locality.-Gulf of Mexico at 29°11.5'N, 88°07.5'W in 356 m. Discussion.-This species was originally described from a single male taken from the stomach of a shark in the Gulf of Mexico. The second specimen, a quite well preserved male captured alive in the trawl was in such good condition except for the partial loss of skin that it was utilized to amplify the description given above. Of particular importance was the discovery of the paired male genitalia not often found in the Oegopsida. On reexamination of the type of Selenoteuthis scintillans it was also found to have paired genitalia and the possibility must now be considered that this condition is found throughout the family. Another remarkable discovery was the presence of the 8 long rod-like light organs, if such they are, buried deeply in the midline of the tail. After removal of the skin and overlying tissue they were easily removed intact, each organ having a silvery sheen and enclosed in a tough outer cover. With the discovery of sexual dimorphism in Selenoteuthis, the possibility arose that springeri was the male of some already known species. However, with the discovery of O. lorigera, represented by a female, this possibility is now disposed of. Oregoniateuthis springeri must rank as one of the most striking of all of the Atlantic luminescent squids, surpassing even the remarkable Nematolampas described by Berry. Distribution.-Gulf of Mexico (Voss).

Oregoniateuthis lorigera (Steenstrup, 1875) FIGURES 9 - 10 ?Onychoteuthis (?) longimanus Steenstrup, 1857, p. 120. Onychoteuthis (??) lorigera Steenstrup, 1875, p. 473.

Material examined.-Holotype, ('j? ML 180.0 mm, fraen Kaskelot- mave, Sydhavet, Mus. Zool. Skibslaege Mj1l11er.Universitetets Zoolo- gisk Museum, Copenhagen. Description.-Mantle stout, thick-walled, cylindrical, tapering to point 1962] Voss: Lycoteuthidae 289

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FIGURE 9. Oregoniateuthis Iorigera (Steenstrup, 1875). Holotype ML 180.0 mm. a. dorsal view; b. ventral view. posteriorly from about ongm of fins; margin produced slightly in dorso-median line, not emarginated under funnel. Fins large, muscu- lar, considerably wider than long, forming short point posteriorly; margins convex anteriorly, concave posteriorly. Funnel large, stout, little projecting, with nearly straight but weakly developed locking apparatus. Funnel valve present, funnel organ with slightly shouldered limbs, ventral pads roundly triangular. Head with parallel sides, nearly as wide as mantle, but badly damaged. Eyes partially torn from sockets. Buccal membrane eight pointed and with eight supports. Arms squarish in cross-section, rounded aborally equipped with two protective membranes and biserial suckers and in order 2.3.4.1. 290 Bulletin of Marine Science of the Gulf and Caribbean [12(2) I appears slightly keeled on the distal 1/3; stout at base but tapering to narrow, attenuate tip; II is extraordinarily developed: first 65.0 mm normal, stout, with deep membranes; beyond this point it is flattened, very thin, slender and attenuate, approximately 2-1/2 times length of mantle. III is very wide, flattened, with deep keel bordering proximal 2/3; proximal half of arm normal with deep membranes, distal half smooth, suckerless, flat and attenuate. IV is squarish with dorsal keel; suckers extended to within 10.0 mm of tip after which arm is smooth, sharply tapered. Tentacles damaged; long, flattened, with stout lower 1/3, becoming a slender strap. Clubs moderately expanded with swimming keel

FIGURE 10. Oregoniateuthis [origera (Steenstrup, 1875). a. funnel organ; h. right tentacular club; c. gladius; d. side view of posterior end of gladius; e. dorsal view, posterior end of glad ius. 1962] Voss: Lycoteuthidae 291 distally; small carpal cluster of 4 suckers and 2 pads arranged in nearly a straight line; manus with four rows of small suckers followed by four rows of minute suckers on the dactyl. Sucker rings mostly missing, deformed or· broken. Stalk with 5 photophores deeply imbedded: one 1/3 from base, large, three small ones widely separated from each other in middle of stalk; large distal organ near base of carpus. Gladius typically lycoteuthid as illustrated. Light organs present. Eyes badly damaged but remains of two photophores of possible (?) five remaining on left eye. Five photo- ph ores on each tentacular stalk as described above. Skin mostly missing but there are no photophores on mantle or head; left arm II has four small photophores widely spaced on strip of skin extending along sucker portion, right II has only two near base; left III has single organ on surface part way out on dorsal side of swimming keel, none on right III. Eight photophores within mantle cavity: 2 round anal organs, one on each side of rectum just below funnel; 5 across base of gills, central one longitudinally oriented, sharp oval, median laterals oval, oblique, laterals round; one on abdomen, transverse, oval, pointed at ea.ch end, strap-like.

TABLE 5 MEASUREMENTS (IN MM) OF HOLOTYPE OF Oregoniateuthis [origera (STEENSTRUP) Total length 660.0 Arm length I 64.0 Mantle length 180.0 II 426.0 Mantle width 43.0 III 94.0 Fin length 95.0 IV 82.0 Fin width 132.0 Tentacle length 282.0 Club length 49.0

Type.-Zoological Museum, Copenhagen. Type locality.-South Sea. Discussion.-This specimen is badly damaged as would be expected from its coming from a sperm whale stomach. As a result the skin is missing over almost the entire animal and nothing can be deter- mined concerning skin photophores. The extensive eye damage also prevents a description of the ocular light organs. Hoyle (1886, p. 39) cites this species in the synonymy of Ony- choteuthis (?) longimanus Steenstrup, 1857. Steenstrup's paper has 292 Bulletin of Marine Science of the Gulf and Caribbean [12(2) not been available to me but a search of the type material at Copen- hagen revealed only a few suckers of the original O. longimanus, so that it is impossible to tell whether this species is the same as lorigera or not. Because of the lack of a true des.cription and loss of the type I prefer to treat Onychoteuthis (?) longimanus as a species dubia. Steenstrup's description of lorigera (1875, p. 473) consists of little more than a recording of the specimen. "Det er naesten, som om denne smukke Sepia derved gjorde et begyndende Till~b til de saa aldeles abnormt og pidskeformigt forlaengede Armpar, som findes hos en oceanisk Blaeksprutte fra det samme store Verdenshav, hvilken jeg i Museet har forel~big opstillet under Navnet Onycho- teU/his (??) lorigera; men sandsynligvis er den en ny Slaegt. Dennes andet Armpar ere dobbelt saa lange som Kroppen og de andre Arme* *. **Individet er redskaaret af en Kaskelot, derfor kan det ikke fuldstaendigen beskrives, da Armene ere blevne staerkt angrebne af Mavesaftan og derved have mistet med faa Undtagelser sine Suge- kopper." From this only one good character is present: the second arms are twice as long as the mantle. At first appearance, this species might be confused with Oregoni- ateuthis springeri but in springeri the ends of arms III and IV are not modified as in lorigera, the photophores are much more numerous, and the shape of arm II is quite different. It is indeed unfortunate that our knowledge of these two interesting species is based upon so few specimens.

Genus Nematolampas Berry, 1913 Nematolampas Berry, 1913, p. 208. Lycoteuthid with greatly elongate ventro-Iateral arms bearing suckers on basal half, distally devoid of suckers, and thread-like. Light organs 8 (possibly 10) within mantle, but numerous on arms, two near end of mantle. Type species.-By monotypy, Nematolampas regalis Berry, ]913.

N ematolampas regalis Berry, 1913 FIGURE l1a Nematolampas regalis Berry, 1913, p. 208; 1914, p. 140; 1916, p. 50. - Thiele, 1934, p. 961. Description.-Mantle small, cylindro-conical; ending in a blunt point. Fins twice as wide as long, sharply pointed laterally. Head large, short, round with large eyes. 1962] Voss: Lycoteuthidae 293 Arms unequal in length, in order 3.2.4.1. Third arms very long, sucker bearing portion about equal to normal length of other arms, distal portion very slender, thread-like and devoid of suckers, knobby in appearance due to presence of numerous deeply imbedded light organs. Tentacles about as long as mantle, with slightly expanded clubs armed with four rows of suckers. Each tentacle with deeply imbedded light organ near base and another just below club. Gladius very narrow, consisting of rachis and narrow vanes, termi-

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a FIGURE 11. a. Nematolampas regalis Berry, 1913 (redrawn after Berry); b. Lampadioteuthis megaleia Berry, 1916 (redrawn after Berry). 294 Bulletin of Marine Science of the Gulf and Caribbean [12(2) nating in small delicate spoon-shaped .cone supported by small, solid, bulbous swelling at end of rachis. Light organs 90 in number. Five light organs on each eye on ventral periphery, of which central one is largest. Small photophore near tip of dorsal and dorso-lateral arms and 31 on each ventro-Iateral arm. Two light organs imbedded in stalk of each tentacle. Pair of large light organs on external surface of mantle ventrally near tip and between fins. Within mantle cavity are eight photophores: one on each side of anus, a transverse row of five photophores on level of bases of gills, posteriorly transversely oval abdominal light organ which may be three parted. Type.-Dominion Museum, Wellington, New Zealand. Type locality.-Kermadec Islands. Discussion.-This species is so distinct that it presents no problem for identification. It was described from only two specimens washed ashore at the Kermadec Islands and no further specimens are known. Distribution.-Kermadec Islands (Berry).

Genus Selenoteuthis Voss, 1958 Selenoteuthis Voss, 1958, p. 370. Lycoteuthids with large round light organ at end of mantle; one large round light organ on tip of dorso-Iateral and ventro-lateral arms m males, absent in females; genitalia paired in males. Type species.-By original designation, Selenoteuthis scintillans Voss, 1958.

Selenoteuthis scintillans Voss, 1958 FIGURES 12 - 15 Selenoteuthis scintillans Voss, 1958, p. 370. Material examined.-Holotype, t in alcohol, ML 32.5 mm from ATLANTIS Sta. RHB 587, 26°22'N, 76°1O'W, Isaacs Kidd mid-water trawl from 46 m, 2003-2111 hours. March 21, 1954. MCZ.-3

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FIGURE 12. Selenoteuthis scintillans Voss, 1958, female, ML 31.0 mm. a. dorsal view; b. ventral view; c. funnel organ; J. left eyeball with photophores. due to fixation and break in dorsal surface of mantle. A large round light organ at end of mantle entirely projecting beyond fins. Fins are short, very broad, terminal except for black photophore projecting beyond them; anterior margins convex, posterior margins concave; lateral corners sharply pointed. Funnel large and stout, free for distal 1/4 of its length, provided with small but strong dorsal support. Funnel organ uncomplicated, consisting of inverted V -shaped dorsal member with short, shouldered limbs, and fat, thick ventral pads. Funnel valve large, sub-terminal, slightly bilobed. Mantle locking apparatus simple, slightly curved funnel groove on each side, with corresponding ridge on either side of mantle. Nuchal cartilage elongate, swollen, rounded posteriorly, 296 Bulletin of Marine Science of the Gulf and Caribbean [J 2(2) widely pointed anteriorly. Gladius typically lycoteuthid, broad in midsection, constricted in posterior third, spoon-shaped distally. Head short, broad, nearly as wide as mantle, with large eyes. Eyes equipped with five light organs around ventral periphery; terminal ones small, round, central three large, round, all with distinct reddish hue. Beaks not dissected out, normal in appearance. Buccal membrane reddish purple, strongly papillose on outer surface, 8 lobed with 8 prominent supports. Arm order 2= 3.1= 4 or 2= 3= 4.1. No keels seen, tips very

FIGURE l3. Selenoteuthis scintillans Voss, 1958. a. dorsal view of male holotype, ML 32.5 mm; b. right tentacular club of holotype; c. left eyeball of holotype; d. apical light organ of left dorso-lateral arm; e. posterior mantle light organ; f. spermatophore, length 3.5 mm. 1962] Voss: Lycoteuthidae 297 attenuate. Single small brownish black light organ proximal to tip on 3rd arm of right side, 2nd arm of left side; tips of corresponding arms of these pairs missing. Arm suckers biserial, bordered dorsally by low indistinct protective membrane, ventrally by broad membrane with stout supports, often fringed in appearance due to torn membranes. Suckers equipped with about 7-9 long teeth on distal margin of horny ring, proximal edge smooth. No apparent hectocotylization of any of arms. Left tentacle alone present; long, stout, bearing short club, broad basally, attenuate for distal 2/3. Suckers in four rows on manus with entire margin finely toothed. Carpal cluster compact, consisting of five suckers with three discernible buttons. Protective membrane with prominent supports ventrally; swimming membrane occupying about 1/3 of the distal border on dorsal side. Two large silvery light organs embedded on aboral surface of tentacular stalks, one basally, other near carpal section. Ten light organs internally; two elongate, oval organs within funnel chamber, one on each side of rectum. Five light organs in midsection of mantle cavity, small longitudinally oval central organ, large transversely oval light organ on each side, small round light organ attached to base of each gill. Three posterior abdominal photophores arranged in transverse row, organs badly damaged, exact shape and number not ascertainable. Spermatophores short, stout, with only few turns of sperm reservoir. Structure of spermatophores unique, illustrated in Figure 1. Holotype.-U. S. National Museum. Type locality.-Off Florida, 26°22'N, 76°10'W in 46 m. Discussion.-Selenoteuthis is very closely related to Lycoteuthis. With

TABLE 6 MEASUREMENTS (IN MM) OF 4 SPECIMENS OF Selenoteuthis scintillans VOSS, 1958

Sex 3 ~ ~ ~ () ~ ~ ~ ML 32.5 31.0 30.0 19.0 Arms, I 12.0 11.5 MW 12.0 11.0 12.5 II 12.9 13.0 HW 10.0 III 12.9 13.0 FL 11.5 13.5 13.5 8.0 IV 12.0 13.0 FW 30.0 30.0 28.5 18.0 TL 27.5 49.0 Diam. terminal light organ 3.0 298 Bulletin of Marine Science of the Gulf and Caribbean [12(2)

FIGURE 14. Selenoteuthis scintillans Voss, 1958. a. upper mandible; b. lower mandible; c. gladius; d. lateral view of posterior end of gladius.

the exception of the terminal light organs of the mantle and arms, few differences are discernible except for its small size. These light organs, however, are of such a size and nature, that it does not seem logical to unite this genus with Lycoteuthis, especially as there is no knowledge of the male genitalia in the latter genus. The structure of 1962] Voss: Lycoteuthidae 299

FIGURE 15. Se/enoteuthis scintillans Voss, 1958. a-h. large right tentacular suckers of 'jl; c. large tentacular sucker of holotype; d-e. largest suckers of left third arm of 'jl; j. largest sucker of right second arm of 'jl. the spermatophore may give us some clue to the type found within the family. Apparently no hectocotylization occurs. Remarks.-The type was taken at a depth of only 46 m. The females were taken in a haul at 1800 fms to the surface but from the depth of the type and the structure and general appearance of the specimens it is most likely that this species belongs in the upper bathypelagic zone. The stomach of the type was opened but only a few fine, un- identifiable crustacean remains were found. Distribution.-Off Florida at 26°22'N, 76°10'W. Subfamily Lampadioteuthinae (Berry, 1916) Grimpe, 1922 Lampadioteuthidae Berry, 1916, p. 51. Gladius enoploteuthid-like, with straight rachis, vane widest at middle, tapering to end where it surrounds rachis but does not form 300 Bulletin of Marine Science of the Gulf and Caribbean [12(2) cone. Five light organs in mantle cavity, 4 on each eyeball and 5 in each tentacle. Type genus.-Lampadioteuthis Berry, 1916.

Genus Lampadioteuthis Berry, 1916 Lampadioteuthis Berry, 1916, p. 52. - Thiele, 1934, p. 961. Body loliginiform with broad, saggittate terminal fins; three light organs on ventral periphery of eye, one lateral near pupil. Type species.-By original designation, Lampadioteuthis megaleia Berry, 1916.

Lampadioteuthis megaleia Berry, 1916 FIGURE lIb Lampadioteuthis megaleia Berry, 1916, p. 52. Description.-Mantle cylindro-conical, firm, fleshy, flaring anteriorly; fins large, fairly thick, slightly more than half mantle length, projecting beyond posterior end of the body. Head large, somewhat flattened, with large eyes. Funnel broad with subterminal valve, straight, simple locking grooves. Arms slender, slightly attenuate, long in order 2=2.4.1, keeled, latero-ventral and ventrals conspicuously so. Suckers in two rows, minute, toothed on distal margin. Tentacles with slender clubs equipped with four rows of minute suckers toothed only on distal margins. Buccal membrane eight pointed, brownish to purple, lappets white. Light organs distributed in orderly fashion. Four organs on eyeball, three on ventral periphery of which middle is smallest, one on postero-ventral lateral surface of eye. Tentacular stalks with four normal photophores embedded in each, with peculiar somewhat stalked organ on base of each. Five photophores within mantle cavity are: one brownish organ on each side of arms, large silvery elongate papi11i- form transverse organ at base of each gill, bright silvery organ posteriorly. Type.-S. Stillman Berry Collection 416. Type locality.-Washed ashore at Sunday Island, Kermadec Islands. Discussion.-Since no further specimens of this unique form have been reported since its original finding, nothing can be added to the description given in detail by Berry, 1916, pp. 52-58. I am not convinced that this species represents a group midway between the Enoploteuthidae and the Lycoteuthidae as Berry has suggested. It 1962] Voss: Lycoteuthidae 301 seems more probable that it represents a branch from the main line along which we find the Lycoteuthidae and the Enoploteuthidae. Actually, our knowledge of the relationships of the two groups is too poor to permit any generalizations at this time. The gladius shape is enoploteuthid, as is the presence of a lateral photophore on the eyeball, but the other light organs are more lycoteuthid. Distribution.-Kermadec Islands (Berry).

GENERAL DISCUSSION The posItIon of the student of phylogeny of the bathypelagic cephalopods is similar to that of a swimmer in deep water; there is very little firm ground upon which to stand. This is partly due to the paucity of specimens and partly to the nearly complete lack of detailed anatomical studies. Thus, probable relationships can only be drawn through certain differences and similarities of obvious struc- tures. Three opinions concerning the position of the Lycoteuthidae are found in the literature. One, proposed first by Chun (1903a) but then rejected by him (1910) and later reinstated by Berry (1914) is that the lycoteuthids represent a separate family and are independent of any others. The second, proposed by Chun ( 1910) places the lycoteuthids in the family Enoploteuthidae, while the third, proposed by Pfeffer (1900), places them in the family Onychoteuthidae. Berry (1916) placed them in their own family, the Lycoteuthidae but separated the genus Lampadioteuthis from the main stock, erecting for it the family Lampadioteuthidae, placing this as an intermediate between the Lycoteuthidae and the Enoploteuthidae. Let us first examine their relationship to the Onychoteuthidae. According to Pfeffer (loc. cit.) the main reason for this position was the similarity of the gladius of Lycoteuthis to the general onychoteuthid gladius. This supposed similarity is, however, more apparent than real. With the exception of the somewhat problematical Chaunoteuthis the onychoteuthid gladius terminates posteriorly in a long straight or slightly curved distinct conus. This type of conus is completely lacking in the lycoteuthids and in fact the only similarity between the two is in the general outline of the vane and the narrowing of this part in the posterior third. Even this, upon closer study, is found to be attained quite differently, since in young lycoteuthids there is not even a simple pocket-like conus and the sides of the vane do not narrow in the 302 Bulletin of Marine Science of the Gulf and Caribbean [12(2) posterior third. It was, in fact, this juvenile condition that caused Pfeffer to describe the genus Asthenoteuthion for the then unrecog- nized young of Lycoteuthis and which he related to the Enoploteu- thidae upon the basis of the structure of the gladius. Other than this superficial resemblance of the lycoteuthid gladius to that of the Onychoteuthidae there is no true relationship. The Onychoteuthidae have 7 buccal lappets and supports, the Lycoteuthi- dae possess eight. The Onychoteuthidae generally have some of the tentacular manus suckers modified into hooks, the Lycoteuthidae do not possess hooks. The Onychoteuthidae possess only the simplest of light organs in one species, Onychoteuthis banksi; the light organs of the Lycoteuthidae are numerous and complex. It is therefore incom- patible with our present knowledge to associate the Lycoteuthidae with the Onychoteuthidae as Pfeffer suggested. The relationship with the Enoploteuthidae is somewhat closer, as Chun has indicated. These two groups share in common, and almost exclusively, an eight parted buccal membrane with eight supports and eight attachments. From our knowledge of this organ considerable emphasis must be placed upon this arrangement. Indeed, the eight parted plan should, on theoretical grounds, appear to be the most complete plan of attachment considering the eight arms as contrasted with both seven and six parted plans as is most often seen (in Histio- teuthis bonellii the young have a seven lappet buccal membrane which in the adult becomes six parted). Another point of strong resemblance is found in the nature of the light organs which are both numerous and composed of several different types with wide distribution over the animal in the Enoplo- teuthidae. Berry (1920, 1920a) has reviewed in detail the distribution of photophores in the cephalopods and has shown that most of these are distributed very widely throughout the most diverse types. He also considers the photophores to have strongly polyphyletic origins. In the arrangement and numbers of light organs the Lycoteuthidae strongly resemble the Enoploteuthidae but it must be stressed, they most closely resemble the family, rather than any genus or species within it. Thus, embedded tentacular photophores are shared exclusively with Pyroteuthis and Pterygioteuthis and are not found apparently in any other families. However, a true posterior abdominal series is not found in the Enoploteuthidae, but is likewise missing in Lampa- dioteuthis. Branchial and abdominal organs are found both in the Lycoteuthidae and some members of the Enoploteuthidae. 1962] Voss: Lycoteuthidae 303 However, the differences of characters between the two families are striking. As mentioned above, the young of Lycoteuthis have an enoploteuthid gladius (Asthenoteuthion) but in the adult lycoteuthids, with the exception of Lampadioteuthis, the gladius is strikingly different. Most teuthologists place strong reliance upon the form of the gladius at the generic and familial level. Contrasting strongly with the enoploteuthids is the absence of hooks. The Enoploteuthidae possess strongly developed hooks on either the manus or the arms in all species, whereas none are known in the Lycoteuthidae. Of perhaps even greater significance is the structure of the hectocotylized arm. In the Lycoteuthidae no hectocotylization is now known in the males of either Oregoniateuthis or Selenoteuthis. In the Enoploteuthidae the hectocotylus may be either a simple double flap as in Abralia or a complex large structure as in Pyroteuthis and Pterygioteuthis. There are many other striking differences, too numerous to list. One thing that strikes the phylogenist when dealing with the Enoploteuthidae in general is the great heterogeneity of this remarkable family and it is certainly possible that when more is known of the detailed anatomy of the various genera future students will find it necessary to split them into several distinct families. Several important features serve to distinguish the Lycoteuthidae from all of the other families of the Oegopsida. The most significant is the form of the gladius, which as we have seen, is distinctive. The only questionable genus in the family is Lampadioteuthis which in this respect lies in a somewhat intermediate position. The possession of paired male genitalia is not found in either the Onychoteuthidae or the Enoploteuthidae and finds its ,counterpart for instance in the widely removed Calliteuthis dofleini of the Histioteuthidae. The posterior abdominal photophores are unique, not being homologous with those found in Pyroteuthis, and only superficially resembling that found in Octopodoteuthopsis megaptera. Also the lack of any modified suckers forming hooks sets the family apart from both of its possible near relatives. The position of the family Lycoteuthidae in the teuthoid classification seems to be close to the Enoploteuthidae. Although lack of sufficient anatomical studies in the latter family makes basic assump- tions shaky, it seems best to derive the Lycoteuthidae from a basic lycoteuthid-enoploteuthid stock sharing in common a simple gladius, polymorphic light organs, and a buccal membrane with eight lappets, 304 Bulletin of Marine Science of the Gulf and Caribbean [12(2) supports and attachments. From this basic stock the Enoploteuthidae and the Lycoteuthidae have arisen, diverging in the characters discussed above. In this scheme Lycoteuthis and Selenoteuthis are considered to be less specialized, closely related genera arising from a separate line from that of the long armed genera Oregoniateuthis and Nematolampas. Lampadioteuthis is considered as a slightly aberrant and more primitive genus and is therefore derived from near the base of the lycoteuthid line, somewhat more closely related to the Eno- ploteuthidae. While this classification has certain inherent difficulties which cannot be solved at this stage of our knowledge, it seems to offer the best explanation of the facts as discussed above, and presents a working hypothesis for the student of this highly spe.cialized group of teuthoids.

LITERATURE CITED

BERRY. S. STILLMAN 1913. Nematolampas, a remarkable new cephalopod from the South Pacific. BioI. Bull., 25 (3): 208-212, 1 fig. 1914. Notes on a collection of cephalopods from the Kermadec Islands. Trans. N. Z. Inst., 46; 134-149. 1916. Cephalopoda from the Kermadec Islands. Proc. Acad. Nat. Sci. Philad., 1916: 45-66, 22 figs., 4 pIs. 1920. Light production in cephalopods, I. An introductory survey. BioI. Bull., 38 (3): 141-169. 1920a. Light production in cephalopods, II. An introductory survey. BioI. Bull., 38 (4): 171-195.

BRUUN, ANTON FR. 1957. Deep sea and abyssal depths. In Treatise on Marine Ecology and Paleoecology. Volume 1. Ecology. The Geological Society of America Memoir 67, pp. 641-672.

CHUN, CARL 1900. Aus den Tiefen des Weltmeeres. Gustav Fischer, Jena, pp. i-viii+ 1-549, 390 figs., 46 pis. ]903. Aus den Tiefen des Weltmeeres. Zweite Auflage, Gustav Fischer, Jena, pp. i-x + 1-592, 482 figs., 46 pIs. 1903a.Ober Leuchtorgane und Augen von Tiefsee-Cephalopoden. Verh. deutsche Zool. Ges. 13: 67-91. 1910. Die Cephalopoden. 1. Tei]: Oegopsida. Wiss. Ergebn. "Valdivia," 18 (1): 1-401, atlas of 61 plates.

GRIMPE, GEORG 1925. Zur Kenntnis der Cepha]opodenfauna der Nordsee. Wiss. Meere- sunter. Komm. Kiel, 16 (3): 1-122, figs. 1-32, 1 pI. 1922. Systematische ilbersicht der europaischen Cephalopoden. Sitz. Naturf. Ges. Leipzig, 45-48: 36-52. 1962] Voss: Lycoteuthidae 305

HARDY, ALISTER 1956. The Open Sea. Its Natural History: The World of Plankton. Collins, London, pp. i-xvi + 1-335, 103 figs., 24 pis. HOYLE, W. E. 1885. Report on the Cephalopoda collected by H. M. S. Challenger during the years 1873-76. Challenger Rep., 16 (44): 1-246, 33 pIs. NAEF, ADOLF 1923. Die Cephalopoden. Fauna u. Flora Golfes v. Neapel, Monogr. 35, Teil I, Band I, Lieferung 2: 149·863, 473 figs. PFEFFER, GEORG 1900. Synopsis der oegopsiden Cephalopoden. Mitteil. naturhist. Mus., 17 (Jahrb. Hamb. Wiss. Anstalt 17): 145-198. 1908. Die Gattungen Abralia, Abraliopsis and Asteroteuthis. Teuthologische Bemerkungen. Mitteil. naturhist. Mus. 25 (Jahrb. Hamburg. Wiss. Anstalt. 25, 1908): 287-295. 1912. Die Cephalopoden des Plankton Expedition. Ergebn. Plankton Exped. Humboldt-Stiftung., 2: 1-815, atlas of 48 plates. ROBSON, G. C. 1924. Preliminary report on the Cephalopoda (Decapoda) procured by the S. S. "Pickle." Rep. Fish. Mar. BioI. Surv. South Africa, No.3 (9): 1-14. 1924a. On the cephalopoda obtained in South African waters by Dr. J. D. F. Gilchrist in 1920-21. Prec. Zool. Soc. London, 1924 (1): 589-686, 51 text-figs., 1 pI. 1926. The Cephalopoda obtained by the S. S. Pickle. Supplementary report. Rep. Fish. Mar. BioI. Surv. South Africa, No.4 (8): 1-6, 1 pI. 1948. The Cephalopoda Decapoda of the Arcturus oceanographic expedition, 1925. Zoologica, 33 (3): 115-132, 18 text-figs. STEENSTRUP,JAPETUS 1857. Vid. Meddel. nat. Foren. Kjl'lbenhavn, Aar 1856, p. 120. 1875. Hemisepius, en ny Slaegt af Sepia-Blaeksprutternes Familie, med Bemaerkninger om Sepia-Formerne i Almindelighed. K. danske Vidensk. Selsk. Skr., sel. 5, 10: 465-482, 2 pIs. THIELE, JOHANNES 1934. Handbuch der Systematischen Weichtierkunde. Dritter Teil. Jena, 1934. Voss, GILBERT L. 1956. A review of the cephalopods of the Gulf of Mexico. Bull. Mar. Sci. Gulf & Carib., 6 (2): 85-178. 1958. The Cephalopods collected by the R/V Atlantis during the West Indian cruise of 1954. Bull. Mar. Sci. Gulf & Carib., 8 (4): 369-389. In Press. South African cephalopods. Trans. Roy. Soc. S. Africa.