A New Species of Cranchiid Squid, <I>Phasmatopsis Lucifer</I>, From

A NEvV SPECIES OF CRANCHIID SQUID, PHASMATOPSIS LUCIFER, FROM THE GULF OF MEXICO!

GILBERT L. VOSS Institute of Marine Science, University of Miami

ABSTRACT A new species of cranchiid squid, Phasmatopsis lucifer is described from the Gulf of Mexico. Its relationship to the genera Phasmatopsis and Taonius is discussed and despite certain characters not common to Phasma- topsis, it is placed in that genus until more information is available. Among the cephalopods collected by the U.S. Fish and Wildlife Service exploratory vessel OREGON during the year 1961 was a single specimen of an unusual cranchiid squid. Examination of the specimen and comparison with representatives of other species in the family revealed that it belongs to a new species, differing in several important features from all others so far described. I am indebted to Harvey R. Bullis, Jr., Base Director, Gulf and South Atlantic Fisheries Explorations, Pascagoula, Mississippi for permission to examine and report upon the specimen. The illustrations are from the pen of Mrs. Lilly King Mannin)!. This paper is part of the results of a study of the cephalopods of the North Atlantic supported by the National Science Foundation under grant G-5853. Phasmatopsis lucifer, new species Figures 1-2 Holotype.-Gravid female, mantle length 95.0 mOl, from Oregon Sta. 3219, 29D06'N, 88°02'W, 60' midwater trawl at a depth of 275 meters, February 9, 1961. UMML 31.298. Description.- The mantle is long and slender, flaring at the anterior margin, widest at the anterior third, and tapering gradually to a slender posterior point. The anterior margin is slightly produced in thc dorso- median line and is shallowly excavated beneath the funnel with lappets on either side at the point of attachment. The mantle is attached to the head in the nuchal region by a narrow union indicated by an oval cartilage which is the anterior end of the gladius. This portion is distinctly papillated. On each side of the funnel there is a slender diamond-shaped cartilage at the point of fusion of the mantle with the funnel base. The fins are badly torn but are slender and marginal, bordering the gladius in the posterior part but not extending past its base. The funnel is moderately stout and extends anteriorly almost to the level of the center of the eyes. The dorsal funnel bridle is paired, each

IConlribulion No. 454 from The Marine Laboratory, Institute of Marine Science, Univers'ity of Miami. 78 Bulletin of Marine Science of the Gulf and Caribbean 113(1 ) unit consisting of closely associated muscle bundles. The funnel adductor muscle is continuous with the outer wall of the funnel. It is very prominent and the two continue up the ventral surface of the head as two prominent

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FIGURE 1. Phasmatopsis lucifer, n. sp.: holotype, mantle length 95.0 mm. a.-Dorsal view. b.-Ventral view. C.-Left tentacle and club. d.-Ventral view of gladius. 1963] Voss: New Species of Squid 79 white ridges which terminate near the base of the ventral arms. There is a small, distinct, semicircular or slightly triangular, funnel valve. The dorsal funnel organ is trilobed with a large triangular flap on each posterior lobe. The ventral pads are large, somewhat oval, with sharply pointed, elongate anterior projections. The head is large, joined to the mantle by a small constricted neck, and bears large, prominent, sessile eyes with widely open lids. There is an olfactory organ below and posterior to each eye, consisting of a long, slender, stalk terminating in a goblet-shaped orifice. The eyeballs each bear on their outer ventral surface two large semilunar photophores, the outer much larger than the inner one and enclosing the latter. The arms are short, in the order 3.4.2.1. The arms are all damaged; only arms IV, left III and right I and II are intact to the tip and on all of these the skin is largely rubbed off. It cannot now be ascertained whether there ever were keels on I and II, but III has a prominent keel along its entire length. The arm suckers are biserial and are bordered on each side by large, well-developed, trabeculate protective membranes. All of the arms terminate in normal though slender tips except the intact left III which bears a large oval purplish black light organ at its extremity. The arm suckers vary considerably in their dentition even on the same arm. On III

a b c

FIGURE 2. Phasmalopsis lucifer, n. sp.: holotype, mantle length 95.0 mm. a.-Left eye. showing shape and position of light organs. b.-Oral view of terminal light organ of third arm. c.-Lateral view of same. d.-Funnel organ. e.-Large sucker from left tentacular club. f.-Basal sucker from right third arm. g.-Median sucker from right third arm. 80 Bulletin of Marine Science of the Gulf and Caribbean [/3(1) basally and scattered throughout the ventral row the ring may bear only five round irregular teeth distally with an irregular entire margin laterally and basally, or it may be more or less covered with teeth as is found in the dorsal row. There is a very narrow papillated area mostly basally. The teeth on the margin are broad, semicircular or flattened, but noteworthy for their irregularity both in size and shape. The tentacles are moderately long, flattened on the oral surface, and bear small expanded clubs. The clubs are not differentiated into manus and dactylus but bear four rows of small suckers on long pedicles. The suckers have a wide papillated ring surrounding the aperture. There arc about two to four irregular large teeth on the distal portion; the remainder of the ring is smooth. The club is bordered on each side by a well-developed, trabeculate, protective membrane but there is no aboral swimming keel. The stalk bears two rows of minute suckers along about four-fifths of its length. These suckers are in pairs alternating with pairs of round pads. The gladius was not dissected out but was clearly visible from within the mantle cavity. It consists of an oval papillated anterior expansion after which it abruptly narrows and passes down the mantIe as a narrow band to about the midpoint of the fins. Here it widens in a broad V and turns around the ventral side but does not form a distinct conus, since it appears to remain open ventrally to the tip where it is slender and pointed but rather short. The liver is cigar-shaped, and free for its anterior third. It is traversed on its ventral side by the rectum. The ink sac is long and slender, deeply set into the liver. The liver in life probably was aligned at an angle of about 45° to the long axis of the body. There is no trace of any luminescent organ on the ventral surface of the liver. The specimen is a female and apparently ripe eggs are distributed in small, compact clusters along the posterior one-third of the mantle between the base of the gills and the stomach. The eggs all appear to be uniform in size but this is not certain. The buccal membrane has six indistinct, rounded lobes and seven supports. The dorsal support is stout, and is single up to the immediate point of attachment to the dorsal arms, where it bifurcates abruptly. The supports are dorsally attached to all of the arms except III, where they are ventral. The supports attached to IV are closely appressed, parallel, and appear at first glance to be single, but when the membrane is stretched they are seen to be paired. Although the skin is rubbed off of almost the entire mantle, it is present on parts of the arms, their bases, the head and the neck. In these places the skin bears large, reddish to reddish-purple, crowded chromatophores, so a complete specimen undoubtedly would be deeply colored. 1963] Voss: New Species of Squid 81 TABLE 1

MEASUREMENTS OF THE HOLOTYPE OF Phasmatopsis lucifer. NEW SPECIES

Mantle length 95.0 Arm length I 15.0 Mantle width 20.0 Arm length II 17.0 Head width 23.0 Arm length III 20.0 Fin length 33.0 Arm length IV 18.0 Fin width 14.0± Tentacle length 52.0 Club length 13.0

Discussion.- The discovery of the new species described above again raises the question of the biological validity of certain genera within the family Cranchiidae. Even the subfamily criteria are now considered unsatisfactory in the light of the problematical position of the genus A scocranchia (Voss, 1962). However, the Cranchiinae may be considered satisfactorily stable in comparison with the Taoniinae, where at present almost utter chaos confronts the student. This confusion is due, in major part, to the prolonged larval and juvenile stages of the species involved, and to the great morpho- logical changes which occur during development from the larva to the adult. It is also complicated by the lack, in many species, of any conception of the appearance of the adult. Further confusion has resulted from a lack of thorough study and evaluation of cranchiid taxonomic characters, poor descriptions, and lost or badly preserved types. The problems within the Taonius-Desmoteuthis-Teuthowenia-Helico- cranchia-Megalocranchia complex have been discussed but not solved by Muus (1956, 1962) and Voss (1960), while the Taonius-Phasmatopsis problem has been discussed by Clarke (1962). The situation in the Taoniinae has long since passed the stage mentioned for the Histioteuthidae by Dell (1952), who stated "What does become readily apparent from the above is that the genera of the Histioteuthids is at present separated and diagnosed are a thoroughly artificial assemblage. Practically every new species obtained appears to require a new generic name." This was somewhat similarly stated by me (Voss, 1962) for the cranchiids with the following observation, "What would be useful in the cranchiids is a catch-all genus which would be used for the reception of new species until their generic affinities can be worked out." The usefulness of such a category can be seen from the following discussion. Clarke (1962) has given a lengthy and detailed history of the status of the genera Taonius and Phasmatopsis based upon new material and reexamination of the type of Phasmatopsis cymoctypus. He has preferred to group Phasmatopsis de Rochebrune (1884) and Carynoteuthis Voss, 1960 together under the first name because they share in common photophores on the liver, a tripartite dorsal funnel organ with two large 82 Bulletin of Marine Science of the Gulf and Caribbean IJ 3(1) flaps, and a funnel valve. These certainly seem to be important similarities. However, they differ in the possession by Phasmatopsis of light organs on the arm tips, pointed teeth on the arm suckers, and a buccal membrane with eight supports. The first two are lacking in Carynoteuthis, which in addition has but seven distinct buccal supports. These are probably all specific differences with the possible exception of the number of buccal supports which may have been caused by the great disparity in size of the animals examined. For the present I consider, with Clarke, that Carynoteuthis is a probable synonym of Phasmatopsis. Clarke incorporated two species under the name Taonius: T. pavo (Leseuer) and T. megalops (Prosch). He characterized the genus by the lack of a funnel valve, lack of flaps on the dorsal funnel organ, and the statement that the light organ on the eye "appears to consist of two or three elements according to the stage of development (MullS, 1956)." This last statement is partly erroneous since pavo as an adult has only two photophores while megalops has three. Perhaps it would also be well to clarify the description of the eyeball photophores in the two genera. In both there are basically two strip photophores, but in pavo and megalops these have the thickened ridge medially with the thin expansion toward the eyeball. In cymoctypus and oceanica the inner, inclosed photophore is reversed, with the raised ridge nearest the eyeball and the thin expansion extending toward the inner area of the ventral organ. This feature among the cranchiids seems to be unique to these two species. It may now readily be seen that lucifer does not fit well into either of the presently defined genera. While it shares with Clarke's Phasmatopsis a light organ on the tip of one pair of arms, a funnel valve and a tripartite dorsal funnel organ with flaps, it has no photophores on the liver, again the buccal membrane has seven supports, and the inner photophore of the eyeball is not reversed. Similarly, it shares with Clarke's Taonius the shape and nature of the photophores on the eyeball, the shape and position of the liver and lack of a photophore on it, and the seven buccal supports. It differs, however, in that lucifer has a light organ at the end of the third arms, there is a funnel valve, and the dorsal funnel organ has two triangular flaps instead of the three long slender papillae. From the above considerations I believe that it is still premature to diagnose these genera as Clarke has done, and in my opinion both the generic and the specific affinities in these groups are still indefinite. Since lucifer shows the strongest affinities with Phasmatopsis, however, and there is no catch-all genus available to us, I am placing lucifer tenta- tively in the genus Phasmatopsis until its true relationships can be worked out. The specific name lucifer is from the Greek meaning "light bearer" and refers to the large light organ borne on the tip of the third arms. 1963] Voss: New Species of Squid 83 LITERATURE CITED

CLARKE, MALCOLM R. 1962. A large member of the squid family Cranchiidae, Phasmatopsis cymocty pus de Rochebrune 1884. Proc. Malac. Soc. London, 35 (1): 27-42, 3 pIs. DELL, R. K. 1952. The recent Cephalopoda of New Zealand. Bull. Dominion Mus. New Zealand, No. 16: i-157. Muus, BENT J. 1956. Development and distribution of a North Atlantic pelagic squid, family Cranchiidae. Medd. Danmarks Fisk. Havunders., N.S., 1 (15): 1-15. 1962. Cephalopoda. The Godthaab Expedition 1928. Medd. om Gr~nland, 81 (5): 1-23. DE ROCHEBRUNE, A. T. 1889. Etude monographique de la famille des Loligopsidae. Bull. Soc. Philom. Paris, ser. 7, 8 (1): 7-28. Voss, GILBERT L. 1960. Bermudan Cephalopods. Fieldiana: Zoology, 39 (40): 419-446. 1962. Ascocranchia joubini, a new genus and species of cranchiid squid from the North Atlantic. Bull. Inst. Oceanogr. Monaco, No. 1242: 1-6.