A NOTE ON THREE SPECIMENS OF THE SQUID LAMPADIOTEUTHIS MEGALEIA BERRY, 1916 (CEPHALOPODA: OEGOPSIDA) FROM THE ATLANTIC OCEAN, WITH A DESCRIPTION OF THE MALEl

RICHARD E. YOUNG Institute of Marine Science, University of Miami

ABSTRACT The male and larvae of Lampadioteuthis megaleia Berry are described for the first time. The presence of a single penis and a hectocotylus, and the discovery of a rostrum on the pen prompt a re-examination of the characters that have placed this animal as the sole representative of the subfamily Lampadioteuthinae.

INTRODUCTION In 1916 S. Stillman Berry described a cephalopod, Lampadioteuthis megaleia, from a lone specimen collected on the beach in the Kcrmadcc Islands in the South Pacific Ocean. For this small animal having an unusual assortment of light organs, he erected a new family, Lampadioteuthidae, primarily on the basis of its rather peculiar gladius, which showed affinities to both the Enoploteuthidae and the Lycoteuthidae. No further information on this new group has appeared in the ensuing half century. In 1962, Voss, in his monograph of the Lycoteuthidae, indicated that the similarities between Lampadioteuthis and the lycoteuthid genera were sufficient to include this animal in the latter group at the subfamily level, thus following Thiele's classification (1934). However he was unable to definitely establish its position within the family without the study of additional material. An examination of the collections of cephalopods from the DANA Expeditions, which are presently on loan to The Marine Laboratory of the University of Miami, has revealed three additional specimens of Lampadio- teuthis megaleia, two of which are larvae. All three specimens were taken from the Atlantic Ocean, thus greatly extending the known range of this species. Berry's original specimen was a female. Therefore, it is fortunate that the largest of the animals described here is a male (Fig. 3). This is particularly important since the lycoteuthid genera are characterized by rather unusual features of the male reproductive system. Since this squid is in an excellent state of preservation (which, unfortunately, was not the

lContribution No. 559 from The Marine Laboratory, Institute of Marine Science, University of Miami. Papers from the "Dana" Oceanographical Collections No. 54. This study was supported by grant GB-I090 from the Nalional Science Foundation. 1964] Young: Notes on Lampadioteuthis Megaleia 445 case with the Kermadec specimen), it is now possible to amplify the original description both in the general characters of the species and in the details of the male genitalia.

ACKNOWLEDGMENTS The writer thanks the Carlsberg Foundation for the opportunity to examine the DANAmaterial and Dr. Gilbert L. Voss of The Marine Lab- oratory of the University of Miami for his assistance with the manuscript.

Lampadioteuthis megaleia Berry, 1916 Figures 1-4 Lampadioteuthis megaleia Berry, 1916: 52.-Voss, 1962: 300. Description.-The mantle is short and muscular, its length being slightly greater than twice its width. The mantle margin projects slightly in the nuchal region but more distinctly so in the areas of the mantle locking apparatus. The outer integument of the mantle extends to the posterior termination of the fins, but the mantle muscle ends a short distance anterior to the apex of the fins. The fins are broad, their width approximately equaling the mantle length, and join in the midline posteriorly where they extend past the apex of the mantle. The gladius (Fig. 4b) is distinct from that of any other known cephalo- pod. Its rachis is long and straight anteriorly, and its thin vane extends laterally, the outer margin of which makes a rather sharp angle just posterior to the midpoint of the pen. At its posterior end, the vane flares slightly and its edges turn under to form a narrow ledge which then follows around the blunt end of the pen. Throughout its length, in the midline of the rachis, a slight thickening occurs which emerges a short distance from the posterior end of the pen as a slender, pointed rostrum. The fact that this rostrum is not a continuation of the posteriormost point of the gladius makes this pen unique among teuthoids.

FIGURE 1. Lampadioteuthis megaleia Berry. a, Large sucker from left ten- tacular club.-b, Ventral sucker from base of second right arm.-c, Sucker from middle of second right arm. 446 Bulletin of Marine Science of the Gulf and Caribbean [14(3) The funnel is short, compact and muscular. The funnel groove is well defined laterally but not anteriorly where it merges with the ventral surface of the head; therefore, a foveola is lacking. The funnel organ is of the typical teuthoid form, consisting of a large, inverted V-shaped dorsal pad and two oval ventral pads. The funnel locking apparatus is pointed anteriorly, blunt posteriorly, and contains a simple, straight groove which opposes a straight ridge on the mantle. The head is approximately the same width as the mantle but narrows slightly anteriorly and ventrally. Two pairs of poorly defined nuchal folds are present; the dorsalmost one arises in the mid-lateral point on the back of the head and travels nearly to the nuchal region, while the ventral most is shorter and arises just lateral to the funnel. Near the dorsal border of the ventral nuchal fold lies a small lobe upon which the olfactory papilla is located. The eyes are extremely large, extending almost the entire length and depth of the head. The eye opening is oval and without a marginal sinus, although there is a slight swelling of the tissue at its anterior border. The arms are roughly subequal in length with the exception of the first pair which is much the smallest. The arm formula is 2> 3 > 4 > 1. The suckers are small and biserial; those of the ventral arms are about half the diameter of the rest and arc restricted to a narrower portion of the arm width. A sucker from the middle of the second arm (Fig. Ie) contains eight bluntly pointed teeth on its distal border. The ventral margin of the

FIGURE 2. Lampadioteuthis megaleia Berry. Ventral view of larva. 1964] Young: Notes on Lampadioteuthis Megaleia 447 first three pairs of arms contains extremely well developed trabeculate protective membranes, while the membranes on the ventral pair are reduced. The protective membranes on the dorsal borders of the arms are all greatly reduced, being represented primarily by stubby trabeculae. The swimming membrane is well developed on all arms except for the dorsal pair where it forms only a narrOw ridge. The buccal membrane consists of two highly papillated lobes, one extending between the dorsal pair of arms and the other up the third left arm. There is no evidence to indicate that this area was damaged during capture. However, in light of the

FIGURE 3. Lampadioteuthis megaleia Berry. Male, ML 31 mm. Ventral view. 448 Bulletin of Marine Science of the Gulf and Caribbean [14(3) fact that Berry's specimen had an eight-lobed membrane, as do the two larval specimens, the unusual nature and asymmetrical position of these lobes must indicate an abnormal condition. The tentacles are robust and taper slightly, being about one and a half times the length of the arms. The tentacular stalks are smooth except for the presence of a few carpal suckers and light organs. The club (Fig. 4c) is small, being between one-fourth to one-fifth the length of the stalk and slightly expanded by the presence of narrow protective membranes. The suckers are in four rather irregular rows on the manus and are set on somewhat elongate stalks. They become progressively smaller on the dactylus and are there arranged into four very regular rows. The distal edge of the largest sucker ring contains eleven small pointed teeth (Fig. la). The carpus contains six scattered suckers on the left stalk and five on the right which are easily distinguished by their narrow and smooth chitinous rings. Knobs on the carpus which would be expected to oppose the suckers could not be found. A narrow swimming membrane runs along the aboral edge of the dactylus. The light organs are large and distinctive (Fig. 3). Each tentacle contains five pearly white organs, the first four being nearly equally spaced while the fifth, lying near the carpus, is somewhat more separated from its fellow. They all protrude slightly from the surface of the stalk except for the peculiar basal organ, which is raised completely off the tentacle by a short pedicel. The light organs of the eyes are perhaps the most easily recognized characteristic feature of this species. Berry (1916, p. 55) has described them as follows: "Subocular photophores large, circular in outline, whitish; four in number on each eye; three, of which the median is somewhat the smaIlest, occupy the usual situs on the ventral periphery; the fourth is larger than any of these, and situated just within the boundary of the pupil, at a point almost exactly behind the centre of the lens." The present specimen differs from this description in that the photophores are brownish in color; the anterior member of the left eye is missing (apparently due to injury), and all the organs are approximately the same size. On either side of the rectum lie two large circular anal photophores with a rather granular appearance and a striking magenta color. At the base of each gill lies a transversely elongate abdominal light organ with a pearly white appearance. Near the end of the mantle, apparently within the median septum and attached to both the inner surface of the mantle and the visceral mass, is located a single oval postero-abdominal photophore with a pearly white sheen. As indicated above, the larger specimen at hand is a male. A small penis is present on the left side of the visceral mass, protruding from the medial side of the gill base. The right ventral arm is hectocotylized by the presence of a smaIl flap on its ventral border nearly midway down the 1964] Young: Notes on Lampadioteuthis Megaleia 449 length of the arm (Fig. 4a) . No sperm atophores were present. The color in preservation is provided by scattered small brown chromato- phores which lie in the portion of the torn integument that still remains with the animal. Both larvae have a mantle length of approximately eight millimeters. They differ from the larger specimen primarily in the following ways: the fin is proportionally smaller; the abdominal photophores are only slightly elongate; the light organs of the eye decrease slightly in size posteriorly; the tentacles are more robust with only the basal light organ visible; the body proportions are slightly different (Fig. 2). In spite of these differ- ences, the larvae are strikingly similar to the larger specimen; thus therc is almost no doubt of their specific identification. Type.-S. Stillman Berry Collection 416. Type locality.-Washed ashore at Sunday Island, Kermadec Islands. Discussion.-Other than the points already mentioned, the specimens here discussed disagree with Berry's description primarily in the structure of the pen. The pen illustrated by Berry had definitely undergone some distortion, since no indication of the presence of a rostrum is given. However this is not surprising since the slender rostrum, which apparently

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FIGURE 4. Lampadioteuthis megaleia Berry. a, Right ventral arm showing slight hectocotylization.-b, Gladius, side and ventral views.-c, Left tentacular club. 450 Bulletin of Marine Science of the Gulf and Caribbean [14(3) lies embedded in the fin cartilage, will break off with unusual ease. In fact, while removing the pen with great care under a binocular microscope, I was unsuccessful in extracting it intact. Instead of clarifying the affinities of this group, the examination of this additional material at first seems only to have added to the existing confusion. The relationship of this group to the Lycoteuthidae has been somewhat weakened by the discovery of a single penis and a hectocotylus (male lycoteuthids have been found only in Oregoniateuthis springeri Voss, 1956, and Selenoteuthis scintillans Voss, 1958; each of these species has paired penes and no hectocotylus). The supposed relationship to the Enoploteuthidae on the basis of the pen is also greatly weakened by the discovery of a uniquely situated rostrum on the pen of Lampadioteuthis. Thus, a brief review of the problem would appear to be in order. As Voss (1962) pointed out, there appears to be a close relationship between the Enoploteuthidae and the Lycoteuthidae as evidenced by similarities in shape, photophore patterns and the presence of an eight- parted buccal membrane. There appears to be no doubt that Lampadio- feuthis belongs to this general grouping, and within this group it seems to be more closely related to the Lycoteuthidae than to the Enoploteuthi- dae. As Berry pointed out, the complete lack of hooks on the arms or tentacles precludes the inclusion of Lampadioteuthis in the Enoploteuthi- dae. Its resemblance to the Enoploteuthidae is to the family as a whole rather than to any single species. For example, Lampadioteuthis closely resembles Pyroteuthis in the arrangement of visceral and tentacular light organs, but more nearly resembles Abralia in the arrangement of eye photo- phores and body shape. On the other hand, these resemblances are com- bined in the lycoteuthid species, Selenoteuthis scintillans. The tentacular club of the Lycoteuthidae is of a very uniform appearance throughout all of the species and differs drastically from seven of the eight enoploteuthid genera. The fact that the club of Lampadioteuthis is nearly identical to that of the Lycoteuthidae, greatly strengthens the argument for relating these two groups. The basic similarities of the shape and appearance of Lampadioteuthis to other species, although somewhat subjective, should be of considerable importance in indicating relationships. The general body form of this animal (including relative eye size, arm formula and relative arm size, fin shape, appearance of nuchal crest, funnel size, etc.) resembles very closely that of some of the lycoteuthid genera. In fact, it requires careful examination to distinguish Lampadioteufhis megaleia from Selenoteuthis scintillans on any basis other than the photophore pattern. These similarities between Lampadioteuthis and the Lycoteuthidae appear sufficient to conclude that these groups are closely related. Thus Lampadioteuthis could be included in the Lycoteuthidae at the subfamily 1964] Young: Notes all Lampadioteuthis Megaleia 451 level, or its own family could be resurrected while acknowledging that it is closely related to the Lycoteuthidae. Either arrangement would be phylogenetically acceptable. It is, at present, difficult to resolve the two possibilities by relying, as one must, on the precedent set in the establishment of other oegopsid families, since classification there is far from stabilized at the family level and therefore cannot provide a sound basis for family designations. Thus, at our present state of knowledge, it would seem prudent to leave Lampadioteuthis megaleia in its present position as the sale representative of the subfamily Lampadioteuthinae. At that level it can be distinguished easily from other members of the family by the shape of its gladius, the presence of a hectocotylus and a single penis, and the somewhat distinctive nature of its photophore pattern. Distribution.- The larger specimen was captured at DANA station 4173 III (40046'N., 18°35'W.), which lies approximately between Spain and the Azores, at night in an open net with 400 meters of wire out. The two larvae were taken at DANA station 1156 II (25°11'N., 20027'W.), just north of the Cape Verde Islands, at night in an open net with 600 meters of wire out. These locations are approximately on the opposite side of the globe from the type locality in the Kermadec Islands.

TABLE 1 MEASUREMENTS (IN MM) OF THREE SPECIMENS OF Lampadioteuthis megaleia BERRY, 1916. Sex c; Mantle length 31.0 8.0 8.0 Mantle width 14.0 4.0 4.0 Head width 13.0 4.0- 4.0- Fin length 18.0 2.5 2.5 Fin width 33.0 4.0 4.0+ Arm length I 14.0 3.0- 3.0 11 22.0 4.5 4.0+ III 21.0 4.0 IV 18.0 3.0+ 3.0 Tentacle length 35.0 8.0 8.5 Club length 6.0+ 1.5 1.0+

SUMARIO ANOTACIONES EN TRES EJEMPLARES DEL CALAMAR Lampadioteuthis megaleia BERRY, 1916, (CEPHALOPODA: OEGOPSIDA) DEL OCEANO ATLANTICO CON DESCRTPCI6N DEL MACHO En 1916 S. S. Berry describi6 el cefal6podo Lampadioteuthis megaleia, par un s610 ejemplar de la hem bra de esta especie, que el seiial6 como el "nico representante de la familia Lampadioteuthidae. Thiele, en 1934, 452 Bulletin of Marine Science of the Gulf and Caribbean [14(3) prefiri6 incluir este animal en la familia Lycoteuthidae al nivel de sub- familia, donde ha permanecido hasta e1 presente. Un macho y dos ejemplares larvales de esta especie fueron reciente- mente descubiertos en la colecci6n Dana. Un examen del macho ha re- velado la presencia de hectocotilos y pene simple (lycoteutidos machos han sido hallados s610 en Oregoniateuthis springeri Voss, 1956 y Seleno- teuthis scintillans Voss, 1958; cada uno de los cuales tiene pene par y no hectocotilo). Ademas la pluma se ha visto que posee un rostrum situado Iigeramente anterior al margen posterior del gladius. Estos caracteres han conducido a una revisi6n de la posici6n sistema- tica de Lampadioteuthis megaleia. Se concluye que esta especie esta es- trechamente relacionada con Enoploteuthidae y con Lycoteuthidae. Sin embargo, su relaci6n con la ultima familia mencionada parece ser mas fuerte. Por tanto, se acepta retener Lampadioteuthis dentro de Lycoteu- thidae al 'nivel de sub-familia para ser distinguida del resto de la familia principalmente por la forma de su gladius, la presencia de hectocotilos y e1pene simple y por la naturaleza algo distinta del patron de sus fotoforos.

LITERATURE CITED BERRY, S. STILLMAN 1916. Cephalopoda from the Kermadec Islands. Proc. Acad. nat. Sci. Philad,. 1916: 45-66.

THIELE, JOHANNES 1934. Handbuch der systematischen Weichtierkunde. Dritter Teil. Jena, Gustav Fischer. p. 961.

Voss, GILBERT L. 1962. A monograph of the Cephalopoda of the North Atlantic. I. The fam- ily Lycoteuthidae. Bull. Mar. Sci. Gulf and Carib., 12(2): 264-305.