THE CARLSBERG FOUNDATION'S OCEANOGRAPHICAL EXPEDITION ROUND THE WORLD 1928-30 AND PREVIOUS “DANA’ -EXPEDITIONS UNDER THE LEADERSHIP OF THE LATE PROFESSOR JOHANNES SCHMIDT

DAHA'BBPORT No.

A STUDY OF THE GENUS ENOPLOTEUTHIS (CEPHALOPODA: OEGOPSIDA) IN THE ATLANTIC OCEAN WITH A REDESCRIPTION OF THE TYPE SPECIES, E.LEPTURA (Leach, 1817)

BV CLYDE F. E. ROPER INSTITUTE OF MARINE SCIENCE, UNIVERSITY OF MIAMI

WITH 24 FIGURES IN THE TEXT

PUBLISHED BY THE CARLSBERG FOVSDATIOS

THESE PAPERS MAY BE REFERFED TO AS: "DANA-BEPORT No. 6S, 1966"

COPENHAGEN AN DR. FR. H0ST * S0N

PRINTED BY BIANCO LUNO a/S 1966 CONTENTS Piige Introduction...... 3

Materials and m ethods...... 4

Historical r6suni6 ...... 5

Family and genus diagnoses...... 8

Enoploteuthis leplura (Leach, 1817)...... 9 D escription...... 9 Batliynietric distribution...... 14 Geographic distribution...... 15 Growth...... 15 Discussion...... 16

Enoploteuthis anapsis R o p e r , 1964 ...... 18 D escription...... 18 External an atom y...... 18 Internal a n a to m y ...... 25 General...... 25 Female reproductive system ...... 27 Male reproductive system ...... 28 Digestive tr a c t...... 29 Bathymetric distribution...... 33 Geographic distribution...... 34 Growtli...... 35

Discussion...... 41

Literature c ite d ...... 45

Conlribiilion So. 640 from The Marine Laboratory, Institute oj Marine Science, L'ninersity of Miami. INTRODUCTION

''he vast cephalopod collections of the “Dana”-l-:xpeditions (1920-22 and 1928-30) have resulted lo date Tin live major “ Dana” -Reports: Jo lb in 's study on the “ Dana” octopods (“Dana”-Heport Xo. 11, 1937); B i u l n s classic work on the biology of Spirilla (“ Dana”-Report \o . 24, 1943); Pickford’s two contributions on Vnmpyroteuthis infernnlis (“Dana”-Reports Xos. 29, 1940, and 32, 1949); and T h o k k 's review of four families of pelagic octopods (“Dana”-Report Xo.33,1949). These larger works, with the exception of B r l l n 's , and the short papers bj- Joubin (1928, 1929, 1929a), Jo ub in and R obson (1929), Robson (1930), and Pick- FORD (1949a), are concerned with the orders Octopoda and Vampyromorpha. Preliminary reports on the “ Dana’ octopods and teuthoids were published by Jo ubin (1931, 1933, 1935), but there are still no reports covering the vast majority of specimens collected by the “ Dana”-Expeditions. The Atlantic teuthoids are presently being studied at the Institute of Marine Science, University of Miami, under the direction of Dr. G ilb f:rt L. V oss, and numerous papers on the collection are now in progress. During a study of the Atlantic Enoploteuthidae in the “Dana” collections, a number of specimens were discovered which represented two species of the genus Enoploteiithis d’Orbignv, 1839, including E. leplura (Lf;acii, 1817) and an undescribed species. L e a c h ’s original description as l.oligo leplura and its synonym, L. Smylliii, and the few subsequent discussions of E. leplura proved to be inadequate for specific identifica­ tion, and it was necessary to examine the original material which L each had deposited in the British Museum (XH). I am indebted to Dr. G. L. Voss for making available his notes on the types which he examined during a visit to the Museum. Subsequently I have examined these types and I concur with ^'oss’s observations. With a knowledge of the types, it is possible to utilize the “Dana” specimens, and other material as it has become available, as a basis for the redescription of E. leplura. The original description of the new species, E.nnapsis Ropf:r, which is well represented in the “ Dana” collections, has already been published (R o p e r, 1964). Teuthologists have long recognized the necessity for the delimitation of the genus Enoploleulhis (C h in , 1910; P f e f f e r , 1912; and Adam, 1960), but the lack of adequate descriptions and material has hampered this delimitation. The collections at hand at the Institute of Marine Science now make possible the diagnosis of Enoploleulhis through the redescription of E. leplura and the further, detailed analysis of E. anapsis. I wish to express my gratitude to the institutions and individuals who loaned material or otherwise rendei ed assistance in the course of this study; the Carlsberg Foundation, Copenhagen, and especially Dr. E. Bertef.- SEN and the late Dr. A. Vedee T.\ning, for making the rich “Dana” collections available for study; the British Museum (Xatural History), Dr. Xorman Tebble, Curator of Mollusks, and Mr. H. O. Ricketts for the loan of live Enoploleulhis specimens and for tracking down the type-Iocalities of Loligo leplura and L. Smylhii] the U. S. Fish and Wildlife Service, Bureau of Commercial Fisheries, and Mr. Harvev R. Bu.i.is, Jr., for the loan of M/\' “Oregon" specimens; Mr. G ilbert W. Bane of Cornell University for the specimens of E. leplura he collected in the Gulf of Guinea; Dr. G. E. Maul for a collection of cephalopods from Madeira; the National Institute of Oceanography, Wormley, England, and in particular Dr. Malcom R. Ci.arke for the loan of excellent male si^ecimens of both Atlantic species of Enoploleulhis taken by the R'V "Discovery”. A special expression of gratitude is extended to Dr. G ilb e r t L. V oss of the Institute of Marine Science, University of Miami, not only for his direction and encouragement of my work, but also for the use of his collections and extensive cephalopod library. The Bureau of Commercial Fisheries, U. S. Fish and Wildlife Service, has provided support of my studies through a fellowship grant, for which I am most grateful. MATERIALS AND METHODS

A series of 26 specimens of Eiioploleiillus unapsis was available. Of these, seventeen were collected by the “ Dana”-Expeditions to the Caribbean and the Equatorial Atlantic. Two large specimens were taken by the U. S. Fish and Wildlife Service M/V “Oregon” , operating in the Gulf of Mexico and the western Caribbean. Two specimens were trawled by the University of Miami Institute of Marine Science R/V “ Gerda” from the mid-Caribbean. Two large specimens, borrowed from the British Museum (Natural History) as E. leptiira, were captured in Funchal Harbor, Madeira. One specimen was found in a collection of cephalopods from Madeira. Two fine male specimens were taken by the R/V “ Discovery” olT Madeira Island. Of the eight specimens used in the redescription of Hiwplolenlhis Icpliira, one was a “Dana” specimen from the Cape Verde Islanils, live were collected by Gii.bkht Ba.ne of Cornell L’niversity from stomach con­ tents of yellowfin tuna {Thiinniis albacarcs) in the Gulf of Guinea, one was taken by the R/V “Gerda” in the southern Straits of Florida, and one was captured by the “Discovery” off Madeira Island. Some specimens were damaged to varying degrees by being crushed by heavier animals in the trawls, or, in very small specimens, by the physical strain of water against the animal while the net was being towed. The five specimens of Enoploleuthis lepliira from the stomachs of yellowfin tuna were partially digested but still identifiable. Most larger specimens were in excellent condition.

Measurements, Abbreviations, and Indices

All measurements were made by dividers and milhmeter rule, except for the smallest specimens and small features which were measured with the aid of an ocular micrometer. Dissection and examination of the specimens were done under a binocular microscope. All illustrations were rendered by the author with India ink on Ethulon. A survey of the use of measurements and indices by previous teuthologists is given by Voss (1963) along with definitions of the various counts, measurements, and proportions currently being used. 1 have attempted to adhere to Voss’s scheme as closely as possible, and the following is a list of the counts and measurements taken on each specimen, listed by the abbreviations used in the text and tables. All measurements are ex­ pressed in millimeters.

Measurements and Counts. ML - Mantle length: length of the mantle along the dorsal niidline from the tip of the tail (posterior-most point of the mantle) to the anterior-most extension of the mantle. MW - Mantle width: greatest width of mantle. HW - Head width: greatest width of head measured across the eyes and including the eyeballs. FL - Fin length: greatest length of fins measured from the midpoint of a line drawn across the anterior fin lobes to the posterior-most extreme of the lins fused to the mantle tip. FW - Fin width: greatest width across both fins measured from the lateral angles. Arm I, II, III, IV - Length of dorsal, dorso-lateral, ventro-lateral, and ventral arms measured trom the proximal-most hook (or sucker) to the tip of the arm. Arm formula: Comparative lengths of the arms in decreasing order, as 4.2.3.1. TL - Tentacle length: total length of tentacle measured from base of tentacular stalk to the tip of the dactylus. CL - Club length: length of club measured from the proximal base of the carpal cluster to the tip of the dactylus. CH - Club hooks: the number of hooks on the manus of the club expressed by ventral and dorsal row- on right and left clubs, as 7, 5/7, 5. CS - Club suckers: the number of suckers on the right manus, daclylus/lefl niaiius, daclylus. AH-S, I, II, III, I\^ - Arm hooks and suckers: lolal number of hooks and suckers on the designated arm, as 25-12. ELO - Eye light organs: number of light organs on the ventral surface of the bulbus of the eye.

Indices. MWI - Mantle width index: greatest width of the mantle as a percentage of the mantle length. HWI - Head width index: greatest width of the head across the eyes as a percentage of the mantle length. FLI - Fin length index: greatest length of the fin as a percentage of the mantle length. FWI - Fin width index: greatest width across both fins as a percentage of the mantle length. ALI, 1, II, III, IV - Arm length index: greatest length of the designated arm measured from the proximal- most hook (or sucke.) as a percentage of the mantle length. TLl - Tentacle length intlex: greatest length of the tentacle as a percentage of the mantle length. CLI - Club length index: greatest length of the club as a percentage of the mantle length.

Other abbreviations are used to designate a particular specimen under discussion: 1) 1160 IV’ indicates a “ Dana” specimen from station 1160, net IV; O 1887 indicates a specimen from the “Oregon” station 1887; UMML 31.237 is the specirnen which bears that accession number in the University of Miami Marine Inverte­ brate Museum; \VA 1 and \VA 2 a-d are labels assigned to the West African, Gulf of Guinea specimens by collection number (1, 2) and by decreasing mantle length (a-d); B.\I (XH) 22 is a British Museum (Natural History) specimen which bears the identification number 22; M 9095 is the specimen number from the Maul Madeira collection; Disc. 4743 represents a “Discovery” haul bearing that number, followed by a, b, or c, an arbitrary identification letter for each specimen. These abbreviations are used mainly in tables.

HISTORICAL RESUME

The first mention in the literature of a member of what is now the genus Enoplofeufhis was made in 1817 by William Le.\ch in the Zoological Miscellany. In this work Le.\ch described two new species, Loligo leptura and Loligo Symlhii. The descriptions in Latin were very incomplete and no illustrations appeared with the descriptions. At that time the genus Loligo was a catchall for nearly every'thing that looked like a s(juid, but subsequently it has been broken down to its present proportions. The species were next mentioned and roughly illustrated in 1818 by Leach in the narrative of the Tuckey Expedition to the River Zaire, Appendix IV, based upon the original specimens collected in the Gulf of Guinea during that expedition. Leach’s work also appeared in French in Blainville’s 1818 Journal de Physique. Five years later Blai.nville (1823, 1823 a) again listed tht two species but gave no additional information. Ferussac (1823) also listed both species and illustrated Loligo leptura in his classical Dictionary of Natural History. In 1826 Orbignv placed the two species in the genus Onycholeiilhis. Ferussac and Orbig.ny’s large work, Histoire Naturelle des Cephalopodes Acetabuliferes, appeared at intervals between 1834 and 1848. Unfortunately, none of the sections bears the actual date of publication, so it has become the tedious task of subsequent workers to interpret, as best they could, the actual dates of publication of names. In 1924 R. Wi.nckworth published the results of his research on the Ferussac and Orbigny work and listed the dates of publication of each section. The systematic section of this great monograph is entirely the work of Alcide d’Orbig.ny. In this work O r­

bigny erected the genus Enoploteiithis. The generic description appears on pages 336-337, which according

to W in c k w o r t h (1924) were published in 1848. Also in this work Loligo Smylhii was first synonymized w ith leptura. However, the generic name Enoploteuthis was used in previously published sections, so the date of the name is questionable. OrbiGxNY first mentioned the name Enoploteuthis as a subgenus on page VII

of the Generalites which was published in 1841 (W in c k w o r t h , 1924). The name is mentioned three more G

limes in that section (page IX, X, and XXV'II) without the subgeneric notation. On page XXVII it definitely appears to be intended as a generic designation. However, all these citations are nomina nuda as none is accompanied by a refence to a species. In the synonymy of Enoplolheulhis leplura on page 337 O rbig ny gives the following citation: d’Orbignv, 1839, Ceph. acct., Onychotenthis, pi. 2, fig. 3-4, pi. 6, pi. 11, fig. 6­ 14, pi. 12, fig. 10-24. The name on plates 2, 6, and 11 is Onychotenthis leptiira and, although the title of plate 12 is Onychotenthis, figures 10-24 are indicated as Enoploteuthis leptura. If plate 12 were published in 1839, as OnBiG.w indicates in his synonymy, that should be the correct date of the generic name. S h e r b o rn in Index Animalium (1925) and Nk.vve in Nomenclator Zoologicus (1939) both give 1839 as the publication date ot Enoploteuthis, referring to plate 12, and I presume that this decision was based on O rb ig n y ’s synonymy. WiNCKwoRTii (1924) gives (1839)-1848 as the date of publication for the plates but states that the bracketed dates should not be used in systematic work because exact page and plate sequence has not been established. However, since the name was used by several other authors prior to 1848 (Ruppeli., 1844, and V e ra n v , 1846), as well as by O rbign y (1845, 1845 a), it appears that Plate 12 must have been available earlier, and indeed, probably was published in 1839, even though Winckworth was unable to obtain proof of this. There­ fore, 1839 remains the valid date of publication of the generic name, Enoploteuthis. Orbigny in several publications (1845, 1845 a) listed and figured Enoploteuthis leptura, but these, as those in Mollusques Vivants et Fossiles (1855), seem to be verbatim from the earlier Ferussac and Orbigny mono­ graph. Until the turn of the century no additional information appears in the literature concerning Enoploteu­ this leptura, other than simple listings of the species. The genus was listed by Gr.\y in 1847 and the species appeared in Gray’s 1849 Catalogue. Enoploteuthis appeared in H. and A. Adams’s generic list of 1858. T ry o n (1879 and 1883) listed the species and gave brief but insufficient descriptions and in the 1879 work repub­ lished both Orbigny’s and Leach’s illustrations. The genus was again listed in H oyle’s catalogue (1886), P feffer’s synopsis (1900), H oyle’s generic key (1904), and H oyle’s list of generic names (1910). Ciil'.v (1910) gave a general description of Enoploteuthis leptura but deferred a complete description to a future work which never appeared. Chun' figured his specimen in toto along with the club (pi. 11, fig. 5 and 6), Based on our present knowledge, the specimen is not Enoploteuthis leptura but is strikingly like the new species, E. anapsis R o p e r, 1964. Chu.n’s misidentification, however, is understandable because of the lack of previous detailed descriptions and clear-cut illustrations, and because the specimen was “severly dam ag ed ” . In his extensive work on the Plankton Expedition Cephalopods, P f e f f e h (1912) included a fairly length}' description of Enoploteuthis leptura, but in the main, it is not sufficiently detailed for specific determination and no illustration was given. Pfeffeh commented that unfortunately the type genus of the famih' was by far the least known of the several genera in the Enoploteuthidae. The first of several additional species assigned to the genus Enoploteuthis was described by Ishikawa (1914) from the Japan Sea and bears the name Enoploteuthis chunii. Shortly afterward, in 1918, B e r r y described Enoploteuthis galaxias from Australia. Both descriptions, especially B erry’s, are detailed and accompanied by illustrations, and these constitute the first adequate descriptions in the genus Enoploteuthis. T h ie le (1921) briefly described Chln’s “Valdivia” specimens but added little new information. Grimpe (1925) merely listed Enoploteuthis in the classification section of his work. In his large monograph of Japanese cephalopods, Sasaki (1929) gave a good description of Enoploteuthis chunii accompanied by several excellent illustrations. The genus Enoploteuthis was again only listed in Thiele’s Handbuch (1934). Nothing more appeared in the systematic literature until 1956 when Rees and .Maul reported two specimens of E. leptura from Funchal, Madeira. In 1959 D ell described Enoploteuthis neozelanicn from New Zealand. From the description and illustration it appears that D ell’s specimen should be referred to the genus Abrriliopsis. This is Abrnliopsis gilchristi (Robson) (Voss, personal communication). Finally, in 1960, Adavi described Enoploteuthis dubia from the Gulf of Aqaba, Red Sea, but he was unable to establish with certainty the generic position because the tentacles of the unique specimen were missing and the buccal membrane connective attached to the dorsal surface of the ventro-Iateral arm instead of to the ventral surface, as is normal. Adam emphasized the need for a detailed description of the type species of the genus, Enoploteuthis leptura, for proper delineation of the genus. An opportunity for such a detailed description was presented during the summer of 1960 when the author was engaged in routine sorting and identification of the Enoploteuthidae collected by the “ Dana” - Expeditions in the Atlantic Ocean. Several specimens wer? found to belong to Hnoploleulliis leptiirn, based upon new information obtained by examination of the types in the British Museum (NH). Among the Enoploteuthis material were other specimens subsequently discoAcred to represent a new species which recently has become established as Enoploteuthis nnapsis (R o p e r, 1964). A brief comparison of the Atlantic species of Enoploteuthis appeared with the diagnostic original description of E. anapsis. The more dc'tailed present study will help to delineate the genus Enoploteuthis and will aid in the identification of its species. FAMILY AND GENUS DIAGNOSES

Family Enoploteuthidae Pfeffer, 1900

F.noplotcuthiclae P f e f f e r , 1000, p. 151 152, 163.

Diagnosis: Animal small to medium, mantle tapered posteriorly, with large, almost or entirely terminal fins; gladius feather-shaped, with distinctly developed vane which has a ledge-shaped thickening on each side next to edge; rachis grooved to the rear with somewhat raised mid-rib; conus flat, spoon-shaped to slender bag-shaped; arms usually with strong protective membranes, suckers completely or mostly developed into hooks; carpal cluster of club with two rows of suckers and pads; buccal membrane with eight supports, lobes, and connectives; light organs usually present—on ventral eyeball, as numerous small organs under the skin, or inside the mantle-cavity; one or both ventral arms hectocotylized. (After T h ie le , 1934, p. 961).

Type genus: Enoplotciitbis O kbignv, 1839, pi. 12, figs. 10-24.

Genus Enoploteuthis OnnioNY, 1848.

Loligo, L e a c h , 1817, p. 141. (in part, reference to Loligo leptura a n d Loligo Smylhii only; non Loligo L am a r c k , 1789). Onychoteuthis, O r b ig n y , 1826, p. 61. (in part, reference to Onycholeulhis leptura a n d Onycholeulhis Smylhii only; non Onycholeulhis L ichtenstein , 1818). Enoploteuthis O r b ig n v , 1839, pi. 12, figs. 10-24; 1848, p. 336. - G r a y , 1849, p. 46. - Ch u n , 1910, p. 57, 107. - P f e f f e r , 1900, p. 165, 167. - H o y l e . 1904, p. 12. - P ff.f f e r , 1912, p. 126. - A d a m , 1960, p. 12. Enoploteiithys, Vf.rany, 1846, p. 17. non Enoploteuthis, D e l l , 1959, p. 6 (mis-identified; = Abraliopsis).

Diagnosis: Mantle conical, muscular, but saccate at tail. Fins not terminal, broadly sagittate, V3-V 4 the mantle length. Buccal membrane purplish; supports, lobes, and connectives eight each. Arms in adult with biserial hooks; biserial suckers at lips. Tentacles with well-defined carpal cluster; hiserial hooks on manus, usually quadriserial, sometimes biserial suckers on dactjlus. Right ventral arm hectocotylized, provided with hooks, suckers, and membranes. Gladius penniform, conus present but not enlarged. Numerous small to minute uniform photophores form several longitudinal multiserial rows (or a patchy pattern in E. galaxias) on the ventral surface of the mantle and head, six rows on funnel, three rows on the ventral surface of the fourth arms, and a single row on the third arms. Nine (8-10) small photophores arranged in a series along the ventral periphery of the bulbus, with the terminal photophores the largest.

Type Species: Loligo leplura L each , 1817, Zool. Misc., 3; 141; by subsequent designation, by P p e f fe u , 1900, Mitteil. Naturh. Mus., 18: 167. ENOPLOTEUTHIS LEPTUHA (Leach, 1817) Figs. 1-5, 7.

Loligo leptura L e a c h , 1817, p. 141; 1818, p. 411; 1818, p. 14, pi. 18, figs. 3, 4, French Translation, atlas (not verified); (in BLA.INVILLE) 1818, p. 395, pi. of June, figs. 3-5. - B i.m n v i l l e , 1823, p. 137; 1823 a, p. 126. - F e r u s s a c , 1823, p. 67, pi. fig. 3. Loligo Smylhii L e a c h , 1817, p. 141; (in B l a in v il l e ) 1818, p. 395. - B l a in v il l e , 1823, p. 137; 1823 a, p. 126. - F e ­ r u s s a c , 1823, p. 67. Onycholeulhis leptura, O r b i g n y , 1826, p. 61. Onycholeulhis Smythii, O r b i g n y , 1826, p. 61. Enoploleulhis leptura, O r b ig n y , 1839, pi. 12, figs. 10-24. - R u p p e l l , 1844, p. 129 (not seen). - O r big .n y , 1845, pi. 17, figs. 1-9; 1845 a, pi. 14, figs. 1 9 (not seen). - V e r a n y , 1846, p. 17 (as Enoploteuthys = misspelling) (not seen). - O r b ig n y , 184€*^, p. 336. - G r a y , 1849, p. 46 (not seen). - O r b ig n y , 1855, p. 399, pi. 27, figs. 1 9 . - H o y l e, 1886, p. i47. - P f e f f e r , 1900, p. 167. - H o y l e , 1910, p. 409. - Chu.n, 1910, p. 107, pi. 11, figs. 5, 6. - P f e f f e r , 1912, p. 126. - IsHiKAWA, 1914, p. 410. - B e r r y , 1918, p. 220. - T h ie l e , 1921, p. 444, pi. 54, figs. 6- 8. - Sasa k i, 1929, p. 238. - T h ie l e , 1934, p. 962. - R e e s and M a u l , 1956. p. 265. - D e l l , 1958, p. 6. - .\ dam , 1960, p. 16. - R o p e r , 1964, p. 140-148. Enoploteuthis Smythii (leptura), T ryon, 1879, p. 172, pi. 75, figs. 311-315.

Material: 1 adult male, ML 79.0 mm, “Gerda” Sta. 120; 23°32'N, 82°21'\V; 19 June 1963; 1620-0m. 1 adult male, ML 73.0 mm, “ Discovery” 4743; 32°43'N, 16°45'\V; 20 September 1961; 170-0m. 1 adult female, ML 44.5 mm, “ Dana” Sta. 1160 IV; 15°50'N, 26°32'\V; 4 November 1921; 300 mw. 1 adult female, ML 21.0 mm, West Africa 1; 04°53'N, 0°37' W; 4 March 1960; Surface— from tuna stomach. 4 late juvenile females, ML 13.5-18.5 mm. West Africa 2 a-d; 04°55'N, 0°35'W '; 4 February 1960; Surface— from tuna stomach.

Description The Mantle is elongate, cylindrical, and tapers posteriorly to a blunt point (Fig. 1). The greatest width of the mantle is at the free anterior margin. The antero-dorsal margin is slightly produced in the midline. The ventral margin is slightly concave between the triangular ventro-lateral lappets of the mantle component of the locking apparatus. Most of the mantle is thick-walled and muscular, but the posterior tip becomes very thin and membranous —almost sac-like. The saccate tail apparently is not a continuation of the muscular mantle wall which terminates at the conus of the gladious, but rather it is an extension of integumentary layers which is filled with a reticulum of fluid-filled compartments. A long, thin band of muscle connects the tip of the tail with the mantle at the end of the gladius. The Fins occupy nearly half the mantle length and are much broader than long. They unite posteriorly and merge with the mantle tissue slightly anterior to the tip. The free anterior lobe persists as the convex anterior margin of the fin. The lateral angle of the fin is rounded, and the posterior border is nearly straight at about a 45° angle to the body axis. Leading and trailing edges of the fins are very thin and fine, but in preserved specimens are usually curled under against the stouter fm muscle. The Funnel is broad and muscular. The funnel component of the locking apparatus, located at the ventro­ lateral angle ol the funnel, is a raised oblong structure with a median longitudinal groove which is narrow and deep anteriorly but flares broadly and becomes shallow posteriorly. This groove receives the ridge of the mantle component of the locking apparatus. The dorsal member of the funnel organ is horseshoe-shaped

Dana-Reporl No. 66, 1906 2 with long, broad limbs which taper to a point posteriorly and which terminate on the funnel retractor muscles. About halfway forward on each limb a long flap arises which extends to the anterior crosspiece of the struc­ ture. The ventral pads are irregularly oval and slightly pointed posteriorly. A broad semi-circular flap, the funnel valve, is fused laterally and posteriorly to the dorsal wall of the funnel tube but remains free anteriorly. The Head is large and somewhat broader than the mantle width. The three nuchal folds are conspicuous and well-defined. The eye opening is moderately large, and the ventral half of the margin possesses a row of light organs. A conspicuous optic sinus projects anteriorly from the anterior margin of the eye opening.

Fig. 1. Enoploteulhis leplura. Adult, 44.5 nun mantle length; A. Dorsal aspect; B. Ventral aspect.

The eye is very large, and a row of nine round light organs appears on the ventral surface of the bulbus. The anterior and posterior terminal light organs are considerably larger than the seven closely packed middle organs and also are separated from them by a small space. The male specimen taken by the “ Discovery” olT the Cape Verde Islands has only seven eye light organs, but this is the only specimen of the Atlantic species of Enoploteulhis 1 have examined which shows fewer than the usual nine light organs. The Arms are long, robust, and of nearly equal lengths. Arm II is usually shghtly longer than the re­ maining three arms which are subequal. All arms have biserial rows of strong hooks fitted with cutaneous sheaths. The number of hooks on each arm is given in Table 1. Only the arm tips are suppHed with double rows of suckers which decrease greatly in diameter distally and suddenly become minute at the tips. Table 2 gives the number of hooks and suckers on each arm of three adult specimens of E. leptinn. The dorsal arm is not keeled but has a slight aboral ridge along the distal one-third of the arm. The dorsal protective mem­ brane of Arm I is very weakly developed, but the ventral one is very prominent with strong trabeculae. Arm II is similar to 1 with a slight aboral ridge distally, a weak dorsal, and a strong ventral protective mem­ brane. The aboral keel of the third arm is well-developed as the swimming membrane, which originates Table 1. Measurements (in mm), counts, and indices of Enoplotcuthis leplura ( L k a c i i , 1817), including the type of L. Smythii.

Specimen Type UMML Disc. D W. Afr. W. Afr. W. Afr. W. Afr. W. Afr. by Sta. No. BM(NH) 31.483 4743 a 1160 IV 1 2 a 2 b 2 c 2 (1

Sex...... F .M M F F F F F F ML...... 50.0 79.0 73.0 44.5 21.0 18.5 15.5 13.0 13.5 >nvi. . . 34.0 30.0 31.4 36.4 57.7 49.7 51.3 56.6 53.3 HWI . .. 31.9 25.4 40.4 50.0 42.1 41.9 40.0 47.1 1 -L I.... 44.0 59.5 58.4 42.6 42.9 40.5 38.6 33.3 39.3 !• WI . ,. 80.4 67.1 68.9 74.1 81.0 71.0 67.0 46.6 63.7 AI.I, I . . 67.2 57.0 66.3 62.9 109.2 105.2 113.0 93.4 100.0 ALI, II . 74.0 59.5 66.3 67.4 122.3 111.0 120.3 96.6 118.6 A1,I, III 65.0 59.3 68.5 67.4 123.3 108.0 117.4 100.0 107.:i ALI, IV. 69.0 66.4 74.3 67.4 119.0 113.0 108.3 86.7 !U.(I T '. l . . . . 89.7 89.0 96.6 ,M 135.0 161.1 M 146.0 CLI .... 19.6 20.6 17.9 M 29.8 43.9 M 46.7 CH ...... 6,3/6,3 4,2'3,3 7,5 7,4 M M 5,5 5,5 5,5 M -M AH, I. . . 23 23 22 25 .M 30 28 .M 21 AH, I I . . 22 24 22 25 M 30 26 ,\I 20 AH, III. 25 25 23 26 .M 32 30 .M 22 AH, IV . 38 33 32 38 M 32 30 M 28 ELO . . . 9 7 9 9 9 9 9 9

,\! = character missing. at the base of the arm, becomes broad along the middle half, then tapers to the tip of the arm. Again, the ventral protective membrane is more strongly developed than the dorsal. On the ventral arm there is no aboral keel or ridge per se. However, the tcntacular sheath extends along the dorsal aboral angle of the arm. It originates web-like between III and IV, and occupies nearly the entire length of IV. The ventral protective membrane is well-developed while the dorsal one is considerably weaker.

Table 2. Number of hooks-suckers on the right/left arms of Enoplotcuthis leptwa.

Specimen: Dana 1160 IV UMML 31.483 Disc. 4743 a ML: 44.5 mm 79.0 mm 73.0 mm Sc.k: F .M M

-\rm: rigtitleft right left right left AH-S, I ...... 25-20/25-17 23-37/23-40 22-26 23-32 I I ...... 25-21 24-19 24-34 24-36 22-20- 21-25^ I l l ...... 27-2127-17 25-33 24-28 23-22 22-28 IV ...... ' 38-20 35-16 35^39,34-33 32-18- 32-4-

+ Indicates a missing arm tip or lost suckers.

The first three pairs of arms in the male possess numerous, minute fleshy papillae evenly distributed over the entire oral surfaces between the bases of the hooks and suckers. The ventral arms do not exhibit this character. Hectocotylization appears on the right ventral arm of the male in the form of a modification in the ventral protective membrane (Fig. 2). This membrane is extremely narrow and weak along the proximal two- thirds of the arm, followed by a sudden pronounced expansion into a long flap which continues distally for a distance occupied by about six pairs of hooks. Distal to the termination of the flap the protective membrane, still somewhat expanded, tapers to the tip of the arm. The dorsal protective merr.brane appears normal save for its expansion into small flaps along an area opposite to the large flap of the ventral protective membrane. The two rows of hooks extend ah)iig the arm, then are replaced by suckers near the arm tip, so that there appears to be no modification in the number, size, or distribution of hooks or suckers. Suckers are biserially arranged over a small area of the distal arm lips and are relatively few in number. The suckers gradually diminish in size distally and suddenly become minute toward the tip. The outer sur­ face of the sucker ring is papillated, the papillae appearing larger distally. The aperture is smooth proxi­ mally, but there are nine teeth distally arranged in an arc. The middle five teeth are long and slender, while the lateral teeth are short and blunt.

Fig. 2. Enoploteuthis leplura. Fig. 3. Enoploteuthis leplura. Fig. 4. Enoploteuthis leptura. Hectocotylus, right ventral arm. Adult tentacular club. Sperm atophorc, 21.5mm.

Tentacles are short — only a little longer than the arms —and thin and are triangular to oval in cross section. The tentacular club is not expanded but is relatively long and slender (Fig. 3). There are two rows of sheathed hooks on the hand with six to seven large hooks in the ventral row and four to five smaller hooks in the dorsal row. The dactylus has two rows of 10-15 small to minute suckers. The sucker rings possess seven small, narrow teeth around their distal half. The dactylus tapers to a narrow tip and is not expanded or rounded. A single, narrow club membrane arises on the dorsal surface of the hand in line between the first and second hooks of the dorsal row and extends to the tip of the dactylus. The carpal cluster is well- defined but long and narrow and consists of five to six each of suckers and buttons. There are no con­ spicuous ridges and grooves. The Spermatophore of Enoploteuthis leptura has an extremely long sperm mass which occupies approx­ imately one-half of the total length of the spermatophore (Fig. 4). The sperm mass extends to the aboral lip of the spermatophore. The cemcnt body is short, only about oue-fourlh of the length of the sperm mass, and has one collar at its oral end. The spiral filament appears relatively simple with a few short longitudinal ridges in the aboral end of the apparatus. Nearly the entire ventral surface of the animal is covered with Luminescent Organs which appear in orderly arrangements (Fig. 1 B). The ventral and ventro-lateral surfaces of the mantle possess a total of seven complex rows of photophores of two types. The larger type has a dark circumference with a light center, while the smaller type is white with no such dark ring. Least clearly defined is the outer lateral row with its scattered photophores. This row is short, extending from the anterior mantle margin to a point about in line with the anterior edge of the fin. The second row originates just median to the lappet of the locking apparatus and extends posteriorly, then curves laterally slightly and ends at a point in line with the middle

Fig. 5. Enoploleulhis leplura. A.Ventral mandible; B. Dorsal mandible; C. Radular teeth. of the fin. The next row (inner lateral) arises at the mantle margin just lateral to the midline and passes to the tip of the tail. But from a point in line with the posterior union of the fins with the tail tip the row loses its complex arrangement and appears as a single line of Ught organs. The seventh row of light organs has its anterior origin in either the right or left row just lateral to the midline somewhat posterior to the mantle margin. This row arches to the midline then runs straight posteriorly where the light organs become scattered before the row terminates at a point in line with the posterior union of the fins. The entire tail is free of light organs with the exception of the simple ventro-lateral rows already described. Until the anterior portion of the median row reaches the Miidiine, all the light organs are of the small, whitish type only. 1'here are a few scattered photophores on the dorso-lateral surface of the mantle. The mantle margin is ringed with a row of light organs which diminishes dorsally. No photophores appear on either surface of the fins. Light organs on the funnel are also arranged in rows. On the ventral surface are four rows, two of which lie just lateral to the clear midline area, while the other two are lateral to the first pair and median to the anterior part of the funnel component of the locking apparatus. These two lateral rows are quite short. Another row e.xtends along the diagonal dorso-lateral angle on each side of the funnel. Photophores on the ventral and lateral surfaces of the head are very numerous and are arranged in an orderly manner (Fig. 1 B). The ventral midline from the funnel groove to the V-notch formed between the bases of the ventral arms is completely devoid of light organs. All rows are paired. The medianmost row lies just lateral to the clear midline and originates with a scattered few white photophores in the anterior terminus of the funnel groove, then passes anteriorly over the head and runs along the aboral median angle of arm IV nearly to the tip. The next row has its origin on the prominent ventral rounded area between the nuchal folds and the funnel groove lateral to the funnel tube. Light organs here are very thickly clustered, Ihen they thin somewhat and pass anteriorly over the head and along the aboral lateral angle of arm IV. This row runs along the longitudinal base of the tentacular sheath to the tip of the arm. The next row arises just lateral to the previously described row and curves anteriorly to end abruptly at a clear patch of skin which acts as a window to let out the light from the nine light organs on the ventral surface of the eyeball. The row then continues out along the lateral edge of the tentacular sheath of arm IV^ terminating nearly at the tip. A simple row of thinly distributed light organs originates at the ventro-lateral nuchal fold and passes lateral to the eye window to merge anteriorly with the row extending along the protective tentacular sheath of IV. The most lateral row originates at the nuchal folds and runs to the middle of the posterior border of the eye opening. Opposite this on the anterior border and dorsal to the conspicuous optic sinus the row con­ tinues and passes along arm III between the longitudinal base of the swimming membrane and the aboral ventral angle. This row terminates just prior to the tip of the arm. The final row of photophores starts dorsal to the middle of the posterior eye opening margin and runs around the ventral half of the eye opening termi­ nating at the optic sinus. The Mandibles are illustrated in Fig. 5 A, B. The radular teeth are long and slender (Fig. 5 C), and the rachidian possesses a small cusp on either side.

Holotype: British Museum (Natural History).

Type Locality: 01°08' N, 07°26'30” E; collected by J. C r a .n c h , Congo Expedition to West Africa.

Bathymetric Distribution Since there are so few specimens of E. leplura on record, an accurate evaluation of its bathymetric dis­ tribution based on captures is not possible at this time. However, it may be assumed that this species is a mesopelagic animal which lives at moderate depths during the day and ascends toward the surface at night. Table 3 shows that the single “ Dana” specimen was taken at night at 100 m over very deep water off the Cape \’erde Islands, while the Gulf of Guinea specimens appear to be from the surface of moderately shallow water. These West African specimens were taken from the stomachs of two yellowiin tuna, Thunnus albacares.

Table 3. Bathymetric distribution of Enoploleuthis leplura ( L e a c h , 1817).

Specimen by Depth of water Depth of capture Time of capture Station No, (meters) (meters) (2400 hours)

Dana 1160 I V ...... 4400 100 0300 West Africa 1 ...... 200 0 0930 West Africa 2 ...... 95 0 1030 UMML 31.483 ...... 1620 1620-0 0330 Disc. 4 743 a ...... 2200? 170-0 2200

Judging from the partially digested condition of the squid and from the other material found in the stomachs, the tuna either had been feeding at intermediate depths or at the surface during the predawn hours on animals ascending from intermediate depths. Either situation would satisfy the indicated mesopelagic habitat of this species. The UMML specimen was captured in a small otter trawl which had fished the bottom at 1620 m, but in all probability, the capture was made closer to the surface while the net was being lowered or hauled. The “ Discovery” specimen was taken over deep water at night within 170 m of the surface. Geographic Distribution Until the R/V “ Gerda” captured a single male specimen in the southern Straits of Florida in the summer of 1963, there was no record of E. leptura from the western Atlantic or the Caribbean. With this capture, E. leptura displays a distribution which might be expected of a tropical Atlantic mesopelagic cephalopod. The known geographic range of E. leptura extends from Madeira, the Cape Verde Islands, and the Gulf of Guinea to the southern Straits of Florida (Fig. 6).

Fig. 6. Geographic distribution of Enoploteulhis leptura. • E. leptura, present specimens; o E. leptura, type locality.

Growth Unfortunately, no small individual was available for this study, so the early stages remain unknown. The four smallest specimens (mantle length 13.5 to 18.5 mm) represent about the same late juvenile stage of development and the largest of these, \V. Africa 2 a, mantle length 18.5 mm, is illustrated (Fig. 7). Five of

Table 4. Enoploteulhis leptura, raw data in inm.

I MMI. Disc. Type Dana W. .\fr. W. .\fr. W. Afr. \V. Afr. W. Afr. Specimen: 31.183 4743 a BM(NH) 1160 IV 1 2 a 2 b 2 c 2 d

M .M F F FF FF F ML...... 79.0 73.0 50.0 44.5 21.0 18.5 15.5 15.0 13.5 MW ...... 23.7 22.9 17.0 16.2 12.1 9.2 8.0 8.5 7.2 H\V ...... 23.8 18.5 — 18.0 10.5 8.0 6.5 6.0 6.4 r-1...... 47.0 42.6 22.0 19.0 9.0 7.5 6.0 5.0 5.3 FW ...... 53.0 50.6 40.2 33.0 17.0 10.0 10.4 7.0 8.6 Arm I ...... 45.0 48.4 33.6 28.0 23.0 19.5 17.5 14.0 13.5 II ...... 47.0 48.4 37.0 30.0 25.7 20.5 18.7 14.5 16.0 I l l ...... 46.8 50.0 32.5 30.0 25.9 20.0 18.2 15.0 14.5 IV ...... 52.4 54.2 34.5 30.0 25.0 20.9 16.8 13.0 12.7 T L ...... 70.9 65.0 43.0 — 25.0 25.0 — 19.7 (.1...... 15.5 15.0 — 8.0 — 5.5 6.8 — 6.3 the eight available specimens were taken as partial stomach con­ tents of yellowfin tuna, so while still specifically identifiable, most of the more delicate tissue has been digested. The three largest specimens are in excellent condition, but the redescription was originally based upon the female (“ Dana” 1160 IV), because the males did not become available until the manuscript of this work was nearly ready for press. Table 1 gives the measurements and indices of Enoploteuthis leplura. Table 4 lists the raw data. The mantle in the late juvenile stage (Fig. 7) is about one-haJf as wide as the mantle length and curves posteriorly to end in a short, narrow tail. The fins are laterally pointed, united posteriorly, and show slight extension posteriorly along the tail. The fins are short and the fin width is between two-thirds and three-fourths the mantle length. The head is large and the arms are very long. The tentacles are moderately long and very thin. The clubs are slender with the narrow carpal cluster and most of the hooks well-developed. Light organ arrangement on the ventral sur­ faces of the mantle, funnel, and head is already well-defined at this stage. However, the light organs are not nearly so numerous as in larger specimens, and they are rather patchy in distribution along the photophore rows of the mantle. That is, a row may have two or three clusters of eight to ten light organs connected by three or four single organs. The patchy grouping is entirely regular so that the patches appear in the .same position on the paired rows and these alternate with the patches of adjacent rows. Fig. 7. Enoploteuthis leptura. Young, 18.5 mm The adult mantle is long, narrow, and tapers into the tip of the mantle length, ventral aspect. saccate tail (Fig. 1). The mantle is about one-third as wide as the mantle length. The fins are broadly pointed laterally', are united, and extend posteriorly to fuse with the tail. The fins are about one-half as long and three-fourths as wide as the mantle length. The adult arms are proportionately very much shorter in relation to the mantle length than in younger specimens. Perhaps (his is of some biological significance which allows the younger members of the species to capture and hold their prey more efficiently. The adult pattern of light organs shows no patchiness whatsoever, and the light organs are extremely numerous.

Discussion The identity of Enoploteuthis leptura has been a stumbling block in the study of the genus ever since L ea ch first described it. The full original description is given below (L e a c h , 1817, p. 141):

Brachia omnia antliis hamatus; supplementaria antliis inferioribus simpUcibus pedunculatis. Loligo leptura: L. branchiis supplementariis hamis liberis, cauda abrupta tenui. Mus. Brit. Corpus et brachia externe laevia tuberculis nonnullis in lineis longitudinalibus interruptis digestis. Loligo Smythii: L. branchiis supplementariis tiamis infra membrana instructis, cauda gradatim attenuata. Mus. Brit. Corpus et brachia externe tuberculata: tuberculis purpureis apice albidis.

No illustration accompanied the original descriptions, but later L ea ch (1818) in the Narrative of the Expedition to Ihe Kiver Zaire figured the types of J.oligo leptura and Smythii in lateral view which give little indication of the specific characters. O kbigny’s illustrations (1848, 18r)5) could easily be either this species or another, since again the characters shown an> inconclusive. Indeed, from the evidence given of the earlier works there was no reason to beUeve that L. leplura and L. Smythii actuallv were conspecific, as O kbignv (1848) and subsequent authors contended. For these reasons, later workers have expressed doubt concerning the relationship of the newer species of Enoploleiithis, both with each other and with the type of the genus. Even the exact nature of the genus was in doubt as late as 1960 when Ad.\m described the most recent possible addition, Eiwploleiilhiii dubia. During the course of this study. Dr. G i l b e r t L. V oss, Curator of the Marine Museum, Institute of .Marine Science, visited several European museums and while in Great Britain re-examined Lkac.h’s original material of Loligo leptura and L. Siuylhii in the type collection of the British Museum (Natural History). The results of Ihe examinalion were given to me upon Dr. Voss’s return to Miami. Several interesting facts were revealed. First, the type of Loligo leptura is a small specimen of about 25.0 mm mantle length and represents the very late juvenile stage described in the present work. It is damaged but has the typical plump body with the small attenuation at the tip of the mantle. It has the characteristic rows of light organs on the body of which Ihe middle row converges with an inner lateral row on one side. Secondly, the jar containing the type of Loligo Smythii is marked “syntypes” and contains a large adult Eiioploteiilhis leptura of 50.0 mm mantle length. Obviously this is the type of L. Smythii and is the specimen figured in Leacii, 1818. It is conspecific w ith L. leplura and represents the adult stage at which the entire body becomes somewhat attenuate, losing the plumpness of the juvenile stage. The middle row of light organs coalesces anteriorly with one of the inner lateral rows as in L. leplura. Finally, the second syntype is not a specimen of Eiwploteutliis leptura, but consists of the head and arms of a specimen of Abraliopsis gilchristi (R o bso n). Dr. Voss examined the specimens of E. leptura reported upon in the present paper and declared them conspecific with the specimens he examined at the British Museum (XH). His notes upon the adult specimen of E. Smythii (= leptura) at the British Museum are appended, along with the counts and measurements (Table 5):

Table 5. Counts and measurements (in millimeters) of Enoploteuthis Smythii (L each, 1817).

.ML 50.0 Arms I ,33.6 Hooks I 23 .MW 17.0 II ,37.0 II 22 r i. 22.0 III 82..T III 25 I-'W 40.2 IV 34..') IV 38

Loligo Smythii L e a c h , 1817. Syntype. The arms are long for an enoploteuthid, and as far as can be told from the largest specimen of Loligo Smythii, only the third arms are keeled for about the distal ^ j. The arms bear hooks over most of the length, but these are followed by a small group of pairs of small suckers and on I. II, and III terminate in two rows of minute suckers at the tip. The tentacles are very slender but have their clubs intact. The hand in the largest specimen bears eight liooks which appear to be in a single row, but this may be due to drying out of the club. The light organs on the mantle are of two kinds: large black ones with clear centers and small white ones. These are arranged in seven distinct rows, diverging somewhat laterally posteriorly, and the center one does not reach the anterior l)order of the mantle. Apparently only those of the lirst two (ventrally) lateral rows reach to the tip of the body or nearly so. There are six rows of light organs on the funnel. The ventral rows are the longest and extend over most of the length of the funnel. The lateral rows are short and are ventro-anterior to the anterior edge of the funnel locking groove. The linal row on each side is almost within the funnei-head groove and slightly dorsal. The head bears four rows of light organs marked by dark photophores. but within the ventral paired rows lie two more rows of white organs making six rows in all. There are circles of light organs around the eyes. The ventral arms bear two main rows of light organs, ventral and dorsal, and a thin, scattered row of organs along the narrow membrane on the dorsal side. Only the median row reaches the end of the arm. The III arm bears a row of light organs along the base of the keel for nearly ® 5 of the arm length (?). There are no light organs visible on the re­ maining arms. This is a gravid female. No. 63 a in G r a y ’s manuscript of the Catalogue. This is Enoploteuthis leptura presented by J. Crancii, I-:sq. Congo Expedition. The locality data reads; West .Africa, Lat. 01 08' N, Long. 07°26'30" K.

Uana-Reporl No. 66, 1966. ^ ENOPLOTEUTHIS ANAPSIS Roper, 1964 Figs. 8-24.

1 Enoploteuthis leptiira, Chun, 1910, p. 107, pi. 11, figs. 5, 6. Enoploteuthis anapsis R o p e r , 1964, p. 140. Holotype: 1 adult female, ML 68.2 mm, “Oregon” Sta. 1887; 16°55'N, 81°10'W ; 23 August 1957; 300 fms; USNM 575605. Paralypes: 1 adult male, ML 79.0 mm, Disc. 4743 b; 32“34' N, 16°45' W; 20 September 1961; 170-0m. 1 adult female, ML 76.9 mm, G. E. M a u l coll. no. 2963; Madeira; date unknown; BM(NH) 1958. 12.30.22. 1 adult female, .ML 69.0 mni, G. li. M a l l coll. no. 3487; Funchal Harbor, Madeira; 1 June 1946; BM(NH) 1958. 12.30.23. 1 adult male, ML 54.0 mm. Disc. 4743 c; 32^34' N, 16°45' W; 20 September 1961; 170-0ni. 1 adult female, ML 27.13 mm, “Dana” Sta. 1292 V; 17°43' N, 64°56' W ; 16 April 1922; 100 mw (meters of wire). 1 adult female, ML 73.2 mm, "Oregon” Sta. unknown; Gulf of Mexico; date unknown. 1 adult female, .ML 29.4 mm, UMML 31.237; 16=55' N, 74°16' %V; 10 -May 1959; 100 m. 1 adult female, ML 16.9 mm, “ Dana” Sta. 1168 V; 09°30' N, 42°41' W ; 12 November 1921; 100 mw. 1 late juvenile female, ML 14.2 mm. “ D ana” Sta. 1238 111; 26°13' N, 78°48' W; 11 February 1922; 300 raw. 1 late juvenile female, .ML 13.5 mm, “D ana” Sta. 1287 III; 16^04' N, 61°52' \V; 8 April 1922; 300 mw. 1 late juvenile female, ML 13.4 mm, “ D ana” Sta. 1269 II; 17“13' N, 64°58' %V; 15 .March 1922; 4,000 mw. 1 late juvenile female, .ML 13.0 mm, “ D ana” Sta. 1247 IV; 17°57' N, 72°51' W; 20 F'ebruary 1922; 100 mw. 1 late juvenile female, ML 11.1 mm, “ Dana” Sta. 1168 V; 09°30' N, 42°41' \V; 12 November 1921; 100 mw. 1 juvenile female, ML 10.0 mm, “ Dana” Sta. 1180 II; 12°11'N, 57°12'W ; 20 November 1921; 300 mw. 1 juvenile female, ML 9.8 mm, “ Dana” Sta. 1202 IV; 09°40' N, 79"’56' W; 10 January 1922; 100 mw.

Additional .Material:

1 adult female, M L 64.2 mm. G. E. .Maul coll. no. 9095; Funchal Fish .Market, Madeira; from Aphanopus carbo stomach; 13 August 1956. 1 late juvenile female, ML 11.0 mm, “ D ana” Sta. 1247 IV; 17°57' N, 72°51' W; 20 February, 1922; 100 mw. 1 juvenile female, ML 9.1 mm, “ Dana” sta. 1223 III; 22°06' N, 84°58' W ; 1 February 1922; 100 mw. 1 juvenile female, .ML 9.0 mm, “ Dana” Sta. 3981 IV; 19°16'S, 01°48'W ; 12 February 1930; 100 mw. 1 juvenile female, ML 8.8 mm, “ Dana” Sta. 1289 IV; 17°43' N, 64°56' W ; 15 April 1922; 300 mw. 1 juvenile female, ML 8.7 mm, “ Dana” Sta. 1243 IV; 21=04' N, 73°48' W ; 16 February 1922; 100 mw. 1 juvenile female, ML 8.5 mm, “ Dana” Sta. (?) Caribbean; date unknown. 1 juvenile female, ML 4.5 mm, “ Dana” Sta. 1238; 26=13' N, 78 48' W, 11 February 1922. 1 juvenile female, ML 4.5 mm, “ Dana” Sta. 1289 IV; 17°43' .N, 64°56' W; 14 April 1922; 200 mw. 1 late juvenile female, ML 10.8 mm, UM.ML 31.236; 17°08' N, 75°02' W ; 10 May 1959; 200 mw.

Description External Anatomy

The elongate, cylindrical Mantle (Fig. 8) tapers gradually posteriorly to a blunt point. The mantle wall is moderately thick and muscular e.xcept at the posterior tip, which is thin, membranous, and saccate. This saccate tail is formed by the extension of the integumentary layers into a cone-shaped bag. The mantle wall muscle terminates around the conus of the gladius. The internal structure of the saccate tail is made up by a reticulum of numerous small compartments arranged in honey comb fashion. The anterior margin of Ihe mantle is free at the neck and on both sides of the funnel, and is held securely in place hv the nuchal and funnel-mantle locking apparatuses. Dorsally the anterior margin is produced in the midline, forming a broadly rounded lobe (Fig. 8 A). The ventral margin is slightly concave between the two small ventro-lateral lappets of the mantle locking apparatus (Fig. 8 B). The Fins extend anteriorly a little more than half the mantle length and are broader than long (Hg. 8 A). The anterior lobes are free and the anterior margins of the fins are convex. The lateral margin forms a sharply

Fig. 8. Enuploteuthis anapsis. .Mantle length 68.6 nini; Dorsal aspect; B. Ventral aspect, (after H o i’e r , 196-1). rounded point, riie posterior margins are straight or slightly concavc and diagonal to the body axis. The fins are imited posteriorly and merge into the posterior tip of the mantle, terminating immediately anterior of the saccate tip. The dorsal and ventral surfaces of the fins are devoid of light organs, but the dorsal surface bears scattered chromatophores, while the ventral surface is clear. The Funnel is well-developed with thick, muscular walls. The paired mantle-funnel locking comple.xes are essentially simple longitudinal grooves and ridges located on each side of the funnel. The funnel com­ ponent (Fig. 9 A) is an oblong, cartilaginous structure with a deep longitudinal sulcus which receives the straight cartilaginous ridge on the inner surface of the mantle (Fig. 9 B). Posteriorly the sulcus becomes broad and shallow. The locking ridge on the mantle is longer than the furrow into which it fits, allowing some 3* independent mantle and/or funnel niovenient along the body axis, while restricting lateral movements when the eoniponents are locked. The dorsal member of the Tunnel Organ forms a broad, inverted V-shaped patch with convex antero­ lateral margins (Fig. 10 C). There are two movable lappets on the ventral surface of the dorsal member which are nearly united at the anterior end. The ventral members are simple, oblong structures aligned with the hmbs of the dorsal member. The Funnel Valve is a semicircular flap attached to the antero-dorsal wall of the funnel tube. In cross-section the Head is nearly round and is slightly narrower than the mantle; the dorsal surface is convex, the ventral surface nearly tlat, and the sides well-rounded. The dorsal surface is continuous with

Fig. 9. Enoploleulhis anapsis. A. Manlle locking apparatus, funnel componenl; B. Manlle locking apparatus, mantle component; C. Nuchal locking apparatus, mantle component; D. Nuchal locking apparatus, nuchal component. the neck, which contains the elongate nuchal-mantle locking apparatus along the mid-line (Fig. 9 C, 1)). The posterior dorso lateral angles of the head contain on each side three nuchal folds and the olfactory crest. The postero-ventral portion of the head is excavated to receive the funnel.

The Eye Opening is nearly oblong in shape. Its margins are slightly rounded, with the optic sinus in the anterior margin just below the level of the pupil of the eye (Fig. 10 B). The Eye is large and bulbous. There are nine nearly circular light organs arranged in a longitudinal row on the ventral surface of the bulbus (Fig. 10 A). The terminal organs are larger than the seven intermediates, which are usually closely spaced or even crowded and sometimes irregular in outline. Three Nuchal Folds occupy the posterior portion of the lateral surface of the head and are divided by a thin, diagonal, membranous ridge into a dorsal triangular and a ventral oblong section. The dorsal border of the triangle consists of an epithelial lappet. The anterior border of the oblong section is a prominent reti­ cular, granular ridge, w hich curves posteriorly to form the dorsal border and the large, rounded olfactory crest.

The Buccal Membrane has eight strong supports with the connectives attached to the dorsal oral angles of arms I, II, and IV and to the ventral oral angle of arm III. The inner surface of the membrane is entirely rugose and papillated and is a brownish purple color in preserved specimens. The lappets have delicate Fig. 10. Enoploleuthis anapsis. A. Bulbus of eye, ventral aspect showing photophores; B. Lateral aspect of eye; C. l unnel organ.

u

i .

Fig. II. Enoploteuthis anapsis. .\. Right ventro-lateral arm, B. Arm sucker; C. Proximal arm hook: D. Lateral aspect of C. D.-H. Series of arm hooks, proximal through distal, lateral view. The chitinoiis Gladius is well developed (Fig. 14 A, B). Us rachis is strong, rounded anteriorly, and forms the heavy midline ridge along the full length of the gladius. The vane is thin and broad and the lateral borders are slightly concave. The broadest point of the vane occurs shortly posterior to the point of origin of the vane from the rachis. A slight thickening of the chitin begins at this broadest angle and passes diagonally to the posterior tip of the gladius. The vane, especially in the anterior portion, is marked with very line striations parallel to the antero-lateral angle. The vane is nearly flat anteriorly in the horizontal plane, but posteriorly it becomes strongly arched to lit the taper of the body. The gladius terminates posteriorly in a bluntly rounded cone. One of the most striking characteristics of Enoploleuthis nnapsis is the presence of Luminescent Organs, or photophores, on the ventral surfaces of the mantle, funnel, head, and arms. These circular organs are of graded sizes from small to minute. The centers are clear white sur­ rounded by a ring of black pigment. The ventral surface of the mantle is profusely supplied with these luminescent organs, some of which are arranged in two miiltiserial longitudinal row s on each side of the clear ventral midline (Fig. 8 B). Lateral to the distinct rows, the photophores are more thinly scattered and form a broad band fr. Ef­ ferent branchial vessel; 16. Branchial heart; 17. Branchial heart appendage; 18. Vena cava; 19. Posterior aorta; 20. Posterior mantle vein; 21. I’osterior mantle artery; 22. Oviduct; 23. Ovum; 24. Ovary; 25. Ventricle of systemic heart; 2G. Fin vein; 27. Median mantle arterv.

Internal Anatomy General To study the organs contained within the mantle, a longitudinal inci.sion in the mantle along the ventral midline was made from the anterior border to the posterior lip. The mantle was then laid open exposing the ventral organs in situ (Fig. 15). In the same way the funnel was opened to expose the funnel organ, the funnel valve in the funnel opening, and the anal opening. The anterior border of the funnel valve lies just inside the funnel opening (Fig. 15: 1). The funnel organ with its ventral and dorsal components lies in the inner wall of the funnel as described in the previous section (Fig. 15: 2). The retractor muscles (Fig. 15: 6) of the funnel lie dorsal to the funnel organ and run posteriorly from the postero-lateral lobes of the dorsal member to their points of origin on the interitu’ dorsal mantle

Dinia-Keport No. 66, 1066. 4 wall along the gladius. A longitudinal band of muscle lies in the dorsal midline of the funnel and sends off small lateral bands to the median borders of the retractors. These, with the retractors, form the dorsal wall of the funnel. Protruding well into the funnel chamber along the midline lies the terminal intestine with rectum, anus, anal flaps, and ink sac

Fig. 16. EnojAoteuihis anai)sis. Female reproducUve system. .A. Nidamental gland and oviduct, dorsal aspect; B. Nidamenlal gland and oviduct, ventral aspect; C, lintjre female reproductive system, ventral aspect. 1. Internal oriface of oviduct: 2. Oviduct: 3. Ovum; I. Oviducal gland; 5. Oviducal gland oriface; (i. Nidamental gland: 7. Membrane: 8. Blood vessels: 9. Nidamental tube; 10. Nida­ mental oriface; 11. Ovary.

The internal orifice of the oviduct (I'ig. 16: 1) lies on the ventral surface of the ovary just beneath the ovarian membrane. The dorsal wall of the oviduct extends further posteriorly bound with the ovarian mem­ brane and thus forms a half-tube leading to the orifice. The oviduct (Fig. 16: 2), bound by its own mesentery to the ventral surface of the ovary, inside the ovarian sac, passes anteriorly in a series of convolutions. These convolulions and the oviduct itself increase in diameter as they approach the oviducal gland anti tlie nida­ mental gland. The anterior loop of the oviduct is filled with ripe ova (Fig. 16:3) waiting to pass through the oviducal gland. A ring of fine lamellae surrounding the antenor portion of the oviduct comprises the oviducal gland (Fig. 16: 4). The base of the bilobed nidamental gland (Fig. 16: 6) surrounds the oviducal gland, and the nidamental gland passes well forward between the gills. The lobes are united at the postero-dorsal base by a continuous band of the laminated, glandular tissue. 'I'he terminus of the oviduct, the oviducal gland, and the base of the nidamental gland cannot be seen in situ because they pass dorsal to the gill bases and the branchial hearts. After passing through the oviducal gland, the oviduct emerges dorsally between the lobes of the nidamental gland and terminates shortly as the oviducal gland orifice (Fig. 16: 5). The entire nidamental gland is enclosed in membrane (Fig. 16: 7) and is well-supplied with blood vessels (Fig. 16: 8), dorso-medi- ally. The space between the nidamental lobes, being covered by the membrane, serves as a tube into which the oviduct opens (Fig. 16: 9). The nidamental gland, like the oviducal gland, consists of many fine lamellae. The anterior tips of the nidamental lobes are also bound in membrane, and a short tube is formed from the ventro-lateral lobe (a continuation of a tube between the lobes). This nidamental orifice (Fig. 16: 10) empties into the anterior pallial cavity just posterior to the funnel border. The two nidamental glands are united by a membranous cavity through which the renal papillae protrude into the pallial cavity.

Male Reproductive System (Fig. 17). The male reproductive system occupies much of the area en­ closed in the posterior portion of the mantle. The conical testis, enveloped in a membranous sac, extends from the posterior ter­ mination of the viscera to the end of the mantle cavity' where it is incased dorsally by the vane of the gladius. The posterior portion

of the N e e d h a m ’s sac (Fig. 17: 1) rests in a deep groove which

extends along the ventral midline of the testis. N e e d h a m ’s sac then curves to the left, passes anteriorly dorsal to the posterior mantle vein, the vena cava and the left branchial heart, and emerges into the mantle cavity dorsal to the basal section of the left efferent

branchial vein. The free anterior portion of N e e d h a m ’s sac, the “penis” (Fig. 17:2), extends a short distance into the mantle cavity between the left gill and the intestine and ink sac before it terminates. The tip of the finishing gland also protrudes into the mantle cavity alongside of the penis. The ampulla of the vas deferens (Fig. 17: 3), bound to the membrane which enfolds the I'ig. 17. Htwploleiilhis anapsis. Male reproduc­ testis, lies to the right of N e e d h a m ’s sac at the anterior ventral end tive system, ventral aspect. 1. Needham’s sac; 2. “Penis ,'i. .Ampulla of vas deferens; 4. Vas of the testis. The remainder of the male reprotluctive system, i.e., deferens; Spermatophore gland I; 6. Spernia- the vas deferens and the spermatophore glands (in addition to tophore gland II; 7. Spermatophore gland III; portions of N e e d h a m ’s sac) lies, bound in strong membranes, 8. Ciliated canal; 9, Spermatophore gland duct; 10. Accessory gland; 11. Finishing gland (ap­ anterior to the testis, dorsal to the branchial hearts, vena cava, pendix); 12. Vas efterens. posterior aorta, and posterior mantle veins, and ventral to the stomach.

D r e w (1919) examined the male reproductive system of live specimens of Loligo pecilei and assigned names to the various segments of the spermatophore glands based upon their observed specific function. However, in the present study, since only two preserved male specimens were available for examination, both of which have been designated as paratypes, it was not possible to undertake a detailed analysis of the spermatophoric organ and associated glands. Thus the nomenclature employed here will follow more closely

that of M a r c h a n d (1907), who simply numbered the spermatophore glands in the order in which the sperm

packets passed through them in route from the vas deferens to the N e e d h a m ’s sac. The illustration of the male reproductive system (Fig. 17) is schematic in that the various structures are shown separated and out of their natural positions. Ordinarily the spermatophore glands and accessory gland are bound tightly together in a membranous envelope. From its point of origin at the anterior end of the testis, the vas deferens (Fig. 17: 4) extends anteriorly in a series of convolutions along the right side of N e e d h a m ’s sac and dorsal to spermatophore gland I, then il makes a loop before enlering the anterior end of the granular spennalophore gland I (Fig. 17:5). Gland II (Kig. 17:6) is a continuation of spermatophore gland I and is in the form of a large hook-shaped lobe which appears to have several distinct sections. A tube arises in the posterior end of gland II near the point of insertion of the vas deferens into the anterior end of gland I. The continuation of this tube forms spermatophore gland III (Fig. 17; 7), first by a slight bulbous swelling of the tube, then by its broad expansion into the long sigmoidal gland. Spermatophore glands II and III are tightly coiled together with the tube which connects them lying between the glands. Spermatophore gland 111, in fact, encloses gland II in its coiling, then swings anteriorly to its bluntly rounded termination. Along thin tube arises at the anterior end of gland 111, and, bound to the gland, it extends posteriorly to terminate w ithout connection. The function of this tube is inknown and is termed the ciliated canal by M a r c iia n d (Fig. 1 7 : 8 ) .

A duct, called the spermatophore gland duct by M a r c h a n d (Fig. 17:9), connects the anterior end of gland I I I with the accessory gland (Fig. 17: 10), a long, posteriorly flattened, boomerang-shaped gland which forms a blind sac with only one duct. Continuous with the accessory gland duct is the appendix (Fig. 17: 11), a hook-shaped, tubular structure with a slightly bulbous termination. This is called the blind sac of the distal vas deferens by VIa r c h a n d , but because there are two openings this term is not applicable. Because of its position at the end of a series of glands which make up the spermatophoric organ, this appendix may be best referred to as the finishing gland (Fig. 17: 11), a term applied by

D r e w . The vas efferens (Fig. 17: 12) (called spermatophoric duct by D r e w , 'A distal vas deferens by M a r c h a n d ) originates in the anterior end of the finishing gland near the terminal swelling and then passes posteriorly to empty into the

N e e d h a m ’s sac. N e e d h a m ’s sac, or the spermatophoric sac, the largest component of the spermatophoric complex, serves as a storage place for the completed spermatophores. The anterior end of the muscular N e e d h a m ’s sac passes into the mantle cavity and terminates as the “penis” (Fig. 17: 2). The spermatophore of Enoploteuthis anupsis is characterized by having the cement body component of the ejaculatory apparatus approximately one-half the length of the sperm mass (Fig. 18). The sperm mass occupies about one- Fig. 18. lutui>l(jteiithis ana/jsis. third of the total length of the spermatophore but does not extend to the aboral Spermatophore, 22.) mm. tip. The oral end of the cement body has two distinct collars, the aboral one being the larger. The aboral end of the spiral filament component has a complex of interconnected ridges which forms a prominent sculpturing for about one-third of the length of the spiral filament. A description of the hectocotylus (Fig. 12), which presumably is used to transfer spermatophores to the female, is given under the description of the arms on page 22.

Digestiuc Tract (Fig. 19). The esophagus (Fig. 19: 6) emerges from the postero-dorsal angle of the head along with the anterior or cephalic aorta and passes posteriorly in a distinct longitudinal groove along the dorsal midline of the long, large liver (Fig. 19: 7). At the posterior tip of the liver two small protrusions extend over the esophageal groove from the dorsal surface at each side of the groove and form a tube around the esophagus and aorta just before they leave the posterior region of the liver. The esophagus enters the antero-dorsal angle of the stomach (Fig. 19: 10). The stomach is an irregularly shaped, bulbous organ which Hes to the right of the midline. To the left of the stomach lies the spiral caecum (Fig. 19: 11), a coiled organ united at its anterior end with the stomach. The extension of this union forms the intestine (Fig. 19: 13). Between the left posterior angle of the Uver and the spiral caecum lies the pancreas (Fig. 19: 9) enclosing the hepatic duct which passes Pig. 19. Enoploteulhis anajisis. A. Digestive Iracl, dorsal aspect: H. Ventral rnandilde and pharynx, dorsal aspcct; C. Buccal mass, ventral aspect. 1. Buccal lip; 2. Ventral mandible rostrum; .3. Buccal mass; 1, Buccal artery; 5. Brachial ganglion; 6. Ksophagus; 7. Liver; 8. .Mantle nerve groove; 9. Pancreatic tissue; 10. Stomach; 11. Spiral caecum; 12. Ink sac; 1.3. Intestine; II. Anus; lo. Anal papilla; 16. .^nal flap; 17. Rostral lamella; 18. Gular lamella; 19. Tongue; 20. Radula; 21. Pharynx; 22. Pharyngeal teeth; 23. Pala­ tine lobe; 24. Palatine groove; 2.5, Esopageal ring; 26. Buccal membrane; 27. Buccal lappet; 28. Buccal support; 29. Anterior buccal artery; 30. Posterior buccal artery; 31. Buccal nerve; 32. Buccal ganglion. 33. Brachial nerves. 34. Buccal connective. into the anterior spiral caecum. The liver is long and cylindrical with bluntly rounded ends and is a single organ, not bilobed. It fills the anterior visceral cavity. A prominent groove lies along the dorsal midline which holds the esophagus and the anterior aorta. Grooves (Fig. 19: 8) from the antero-lateral angles contain the large mantle or pallial nerves which run from the head to the stellate ganglia on the posterior part of the head retractor muscles. These muscles form a tube fused ventrally and postero-dorsally around the liver. The anterior end of the liver is nearly flat where it lies adjacent to the posterior wall of the cranial cartilage. A groove nms along the midline for a short distance on the antero-ventral surface of the liver. This groove contains the visceral nerve. The intestine emerges from the stomach and spiral caecum, turns ventrally, then passes anteriorlv along the midline terminating at the anus (Fig. 19: 14) in the funnel chamber. The long tubular ink sac (Fig. 19: 12) is bound by mesentery to the dorsal surface of the intestine. Its posterior terminus lies between the spiral caecum whorl and the postero-dorsal angle of the pancreas. It then joins the intestine and passes anteriorlv with it. The intestine and the ink sac are bound fast to the fused ventral e.vtensions of the head retractor muscles. The ink sac empties into the intestine or rectum adjacent to the anal opening. (In the illustration the intestine and the ink sac have been displaced to the side to make them visible.) The anal oyiening is formed of two double components: the anal papillae and the anal flaps. The anal papillae (Fig. 19: l.'>) form the actual anal opening when they are held apart. The anal flaps (P'ig. 19: 16) are asymmetrically Y-shaped and arise at the base of the anal slit perpendicular to the papillae bases. The flaps extend well beyond the ends of the papillae and into the funnel chamber, thus insuring the direct extrusion of waste material and ink through the funnel tube and into the water. The esophagus (Fig. 19: 6) passes anteriorly to the head through the esophageal foramen in the cranial wall. Before entering the cranium, the esophagus is slightly swollen for a short distance. The anterior aorta accompanies the esophagus to the cranial wall but forks symmetrically before penetrating the wall and en­ tering the head region. Once inside the cranial chamber the esophagus turns ventrally and passes anteriorly through the center of the brain. Between the brain and the buccal mass the esophagus is bound with nerves, ganglia, and vessels. Branches of the aorta follow the route of the esophagus through the brain and into the buccal mass (Fig. 19: 3). One branch divides just ventral to the buccal ganglion and serves the posterior buccal mass (Fig. 19: 30), while the other branch passes anteriorly through the buccal muscle to serve the anterior buccal mass (Fig. 19: 29). The large, bilobed brachial ganglion (Fig. 19: 5) lies just dorsal to the esophagus posterior to the buccal mass and radiates nerve branches to each of the arms. Also a large branch from each lobe of the brachial ganglion passes antero-ventrally to form the lobes of the buccal ganglion (Fig. 19: 32) located on the posterior border of the buccal mass ventral to the point of origin of the esophagus. Nerves from the buccal ganglion serve the several components of the buccal mass. The buccal mass (Fig. 19: 3) itself is a large, globular structure encased in strong muscles. The posterior hemisphere is internal and receives blood vessels and nerves and gives rise to the esophagus. The anterior hemisphere is external, covertd with skin and surrounded by the buccal membrane (Fig. 19: 26) which is strengthened by the eight buccal supports (Fig. 19: 28). The buccal connectives (Fig. 19: 34) connect the free edge of the membrane to the dorsal side of arms. I, II, and IV and to the ventral side of 111. Inside and concentric with the buccal membrane is the muscular papillated lip (Fig. 19:1) which nearly covers the jaws and which forms the an­ terior boundary of the anterior hemisphere of the buccal mass. The mandibles (Fig. 20 A-D) are chitinous, black to brown in color, and superficially resemble parrot beaks. The beaks or rostra are very strong and pointed (Fig. 20: 1). Both mandibles are held in place and motivated by powerfid muscles which insert on all surfaces except the rostra themselves. The dorsal mandible (Fig. 20 C, D) is the more simply constructed, consisting of the sharp rostrum and the broad, hood-like, palatine lamella (Fig. 20: 2), which serves as dorsal and lateral walls for the pharynx and for the attachment of the masticatory niusclcs. The ventral mandible (Fig. 20 A, B) consists of the blunt, overlapping rostrum, the broad, dorso-lateral rostral lamellae (Fig. 20: 4), and the triangular, ventral median gular lamella (Fig. 20: 3). The upper mandibular rostrum and lamella fit inside of the lower rostrum, rostral lamella, and gular lamella. The lower rostrum overlaps the upper, forming an efficient cutting edge. A longitudinal cut through the muscles of the dorsal surface of the buccal mass exposes the palatine lamella of the upper mandible which lies quite close to the surface. Further separation of the muscles reveals the flat dorso-lateral rostral lamellae of the lower mandible. The muscular tongue (Fig. 19: 19) fits into the hollow of the rostrum of the lower mandible and between the two vertical throat lamellae (Fig. 19). The tongue surface is covered with papillae, and the anterior salivary gland is located between the anterior lobes of the tongue. The posterior border of the tongue is crescent-shaped duo to the presence of the curved chitinous radula with its protective sheath.

LS

-----2

Fig. 20. Enoploteulhis anaf)sis. A. Ventral mandible, lateral aspect; B. Ventral mandible, ventral aspect; C. Dorsal mandible, lateral aspect; D. Dorsal mandible, dorsal aspect; E. Radular teeth. 1. Rostrum; 2. Palatine lamella; 3. Gular lamella; 4. Rostral lamella.

The radular ribbon (Fig. 19; 20) originates ventrally in the radular sac, passes dorsally to just above the tongue surface, and curves slightly posteriorly into the pharynx. The posterior or dental surface of the radular sheath has indentations corresponding to the rows of teeth on the radula. This close-fitting sheath protects the radula from damage. The radular ribbon (Fig. 20 E) possesses seven rows of sharp, closely overlapping, curved teeth which are directed posteriorly; the rows converge slightly at the terminus. The teeth of the outer two rows on each side are curved toward the three center rows, which point straight posteriorly and are curved only in the horizontal plane and not laterally. The old, working teeth toward the tip of the radula are considerably more blunt than are the young, unused teeth which continuously grow from the radula sac. None of the radular teeth possesses cusps (Fig. 20 E). The wall of the pharynx (Fig. 19: 21) is formed by two large vertical papillaled flaps, the palatine lobes (Fig. 19: 13), which arise ventrally and are contained dorsally by the sides of the curved palatine lamella of the upper mandible. The cuticular lining of the pharynx, esophagus, and stomach originates along the dorse- median surfaces of the palatine lobes. In the pharvtigeal area the cuticle gives off several rows of sharp posteriorly directed teeth (Fig. 19: 22), which insure the direction of the passage of food. The esophagus (Fig. 19: 23) arises as a tube at the posterior border of the buccal mass. The dorsal wall of the pharvnx is formed by the palatine lamella of the upper mandible, the posterior tip of which is inserted under and united with the antero-most dorsal wall of the esophagus. Table 6 gives the counts and measurements of the holotype of E. anapsis.

Table 6. Counts and measurements (in mm) of the holotype of Enoploleulhis nnnpsis Ropkr, 1964.

Sex - F Al., I - 39.0 Hight U-fl ML - 6 8 .6 I I - 43.0 CH 7,5 6,0 MW - 22.:J III - 45.1 .^H-S, 1 - 24-29- 24-27; HW - 21.0 I V - 47.0 II - 25-2ti- 25-20 I-'L -44.1 TL - 1.34.5 III - 20-24 25-10 - FW - 52.0 CL - 21.0 IV - 29-2.i- 28-27

Holotype: U. S. National Museum (USNM 575605). Paratypes are deposited in the U. S. National Museum, the Zoological Museum, Copenhagen, the British Museum (Natural History), and the University of Miami Institute of Marine Science Marine Invertebrate Museum.

Type Locality: 295 statute miles WSW of Kingston, Jamaica and 165 statute miles .S of Grand Cayman Island at 16°55' N, 81°10'W. “Oregon” Sta. 1887; 23 August 1957; 540-0ni.

Bathymetric Distribution Table 7 compiles available bathymetric data for the 26 specimens of Enoploleulhis nnapsis. The two speci­ mens taken by the “ Oregon” and the two taken by the “ Gerda” (these bear UM.ML accession numbers) have the actual depth of capture, calculated from the wire angle, recorded in the original data. The data recorded from the “Dana”-Expeditions, however, do not include actual depth of capture but indicate the number of meters of wire out. The wire angle was not recorded. Therefore, the depth of capture is here calculated by the method used by Brlln (1943). For lengths of wire up to 1000 meters the depth of capture is calculated at one-third the length, and for lengths greater than 1000 meters of wire (mw) the depth of capture is recorded as one-half the length. Only one specimen comes under the second category. All specimens except one were captured from 1800 hours to 0450 hours when these animals might be expected to be in the upper levels of water. At least one of the three Madeira specimens was caught at the sur­ face at night BM(NH) 22. One other (M 9095) was taken from the stomach of Aphanolus carbo, a mesopelagic feeder. No data is available for the third specimen (BM(NH) 23). The “ Discovery” specimens, taken south of .Madeira, also were captured at night over deep water. The catch was made with an open net which was lished in the upper 170 meters of water. Most of the captures were at quite shallow depths of less than 200 meters. I'ufortunately, it was not feasible for the “ Dana” to make deep (greater than 1000 meters) tows at night and all such tows were made during the day; shallow hauls at less than 1000 meters were made at night. For this reason it is not possible to show vertical migration by direct comparison because day and night hauls from the same depths were not taken. However, it is reasonably safe to assume that the animals had migrated chjse to the surface or were in the process of migrating toward the surface when they were caught at night. This assumption may be justified because this species, although possessing light organs characteristic of mesopelagic organisms, in all other features is adapted for life near the surface and does not exhibit the reddish coloration, soft or gelatinous mantle and other characteristics of deep dwelling squids. That no specimen was taken in the numerous deep, daytime hauls (1000 m and deeper) indicates that the species probably is limited to the mesopelagic and upper bathypelagic zones as defined by Bruu.n (1957).

Dana-Report No. 66, 1966 ML Depth of water Depth of capture Time of capture station number (mm) (meters) (meters) (2400 hrs)

D 1289 IV ...... 4.5 100 0040 D 1238...... 4.5 620 16-266 0400 DC?) ...... 8.5 D 1243 IV ...... 8.7 2,220 33 1930 D 1289 IV ...... 8.8 100 0040 D 3981 IV ...... 9.0 5,070 33 2000 D 1223 III .... 9.1 33 1940 D 1202 IV ...... 9.8 1,,500 • 33 0450 D 1180 II ...... 10.0 100 2130 IM.ML 31.2.36.. 10.8 1,800-2,400 200 1050 D 1247 IV ...... 11.0 2.500 33 0400 D 1168 V ...... 11.1 33 0400 D 1247 IV ...... 13.0 2.500 33 0400 D 1269 II ...... 13.4 4,250 2000 1800 D 1287 III .... 13.5 100 2010 D 1238 III ___ 14.2 f)20 100 0400 D 1168 V ...... 16.9 33 0400 D 1292 V ...... 27.3 33 1930 U.M.ML 31.237 . . 29.4 3,000 100 2130 Disc. 4743 c . . . . 54.0 2,200? 170-0 2200 -M 9095 ...... 64.2 O 1887 ...... 68.G 600 B.M(NH) 2 3 ___ 69.0 O (?)...... 73.2 B.M(NH) 2 2 ...... 76.9 0 Night Disc. 4743 b ...... 79.0 2 ,2 0 0 ? 170-0 2200

As all the nets used by the four vessels involved in collecting the present specimens were open type nets, some captures could have been made at lesser depths than indicated while the nets were being set or hauled. Brlu.n (1943), however, considers that the time spent in lowering and raising is insignificant in comparison with the time spent fishing at the desired depth. This would be especially true if a few or, as in most cases, only one specimen were taken. With deep hauls, of course, time spent lowering and raising becomes an im­ portant factor, and nets set to fish at 2000 meters (as in “ Dana” Sta. 1269 II) would require about 40 minutes to reach the surface with 4000 mw and an average drum speed of 100 meters per minute. The drum speed of the "Dana" trawl winch was 150 meters per minute full and 60 meters per minute empty (Schmidt, 1929). Apparently adults and juveniles share the same bathymetric distribution, neither preferring a particular layer. Geographic Distribution Enoploteuthis nnapsis has been captured throughout the tropical Atlantic and Caribbean and at Madeira (Fig. 21). The majority of specimens was collected from the northern Caribbean, but this is presumably a factor of sample collection pressure rather than of actual biological distribution. The specimens from Madeira and the one taken southeast of St. Helena at 19^6' S, 01“48'W (F'ig. 21), provide good evidence that the species is widely distributed throughout the tropical Atlantic. However, no conclusions can be drawn as to relative abundance throughout the known range of the species because so few stations have been made in the mid-Atlantic between 10° X and 20° S. Actually, this species with its mesopelagic habitat should be expected to be widely distributed. The specimens used in this study come from the following localities: the Tongue of the Ocean, Bahamas; northern central Gulf of Mexico; just north of Haiti; the Caribbean Sea, especially along the Antilles; just north of the Panama Canal Zone; 220 statute miles east of Barbados; 1400 statute miles east of Trinidad a j ill the mid-Allanlic; Madeira; and jusl southeast of St. Helena Island. If Cm \ s specimen is referable to Enoploleulhis anapsis, as it appears to be, its location closes the gap between the St. Helena specimen and those of Madeira and the mid-Atlantic, since it was taken in the South liquatorial Current at 0^29' N, IS^oT'W.

Growth The growth of Enoploteuthis anapsis, as in most other cephalopods, shows a smooth, continuous develop­ ment to the adult with little indication of separate larval or juvenile stages as are present when marked metamorphosis occurs. Larval and juvenile stages are therefore recognizable only through the state of devel­ opment of such characters as the hooks, the development of the manus, the complexity of the carpal ckister, light organ patterns, and others. The smallest specimens available (4.5 mm mantle length), are in unsatis­ factory condition for description or illustration. The smallest specimen which can be utilized (1) 1243 IV) is 8.7 mm in mantle length and is considered an early juvenile (Fig. 22). Figs. 23, 24, and 8 show the mid­ juvenile, late juvenile, and adult forms respectively. Table 8 gives the measurements and indices of all 26 specimens available to this study. Table 9 lists the raw data from which the indices were calculated. The 8.7 mm specimen is short and stout; the mantle width about two-thirds to three-fourths of the mantle length. 'I'he mantle is bullet-shaped and plump, curving sharply posteriorly to a bluntly rounded tij) w hich projects only slightly beyond the fins. The conus of the gladius is clearly visible through the thin skin of the ventral surface. The fins are round, about twice as wide as long, and are united posteriorly but do not extend to the end of the mantle. The fins are less than one-half the mantle length, and broad, their width about 80 to 90 per cent of the mantle length. The head width is equal to or slightly greater than the mantle width. The arm formula is 3.4.2.1 or sometimes 3.2.4.1 in other specimens of about the same size. There are fewer than ten hooks on each arm with Arms II and ill possessing the greater number. A number of suckers arc in the process of developing into hooks. The tentacles are robust and even at this size are considerably longer than the mantle. The club bears no hooks but the four large and four to six small suckers show indications of transformation into the future hooks of the adult. The carpal cluster is not distinct, and the club membranes appear only as indistinct ridges. Table 8. Measurements and indices of Enoploteuthis anapsis. Abbreviations are explained in materials and methods section.

Specimen Disc. BM(NH) BM(NH) O 1887 M Disc. UMML D D D D D by sta. 4743 b 22 23 Type 9095 4743 c 31.237 1292 V 1168 V 1238 III 1287 III 1269 II

Sex...... M F F F F F M F F F F F F MI...... 79.0 76.9 73.2 69.0 68.6 64.2 54.0 29.4 27.3 16.9 14.2 13.5 13.4 MW I ___ 34.4 35.1 34.2 32.4 32.2 38.9 33.5 35.7 36.6 41.6 76.0 50.3 66.4 H W l ___ 29.0 33.9 29.8 30.4 30.8 36.4 28.7 30.6 32.2 37.2 59.8 44.4 44.7 I'Ll...... 68.4 70.5 73.6 73.0 64.4 59.0 69.7 58.2 50.9 49.1 50.0 51.8 50.7 I'-fl . . . . 73.7 88.5 73.8 82.3 76.0 95.9 74.1 79.0 72.1 79.8 99.2 74.0 108.2 .\LI, I... 54.8 70.5 68.7 65.6 56.8 82.5 58.4 47.6 44.3 51.4 57.0 51.8 60.4 ALI, II . . 59.2 74.2 71.2 69.5 63.4 93.4 61.1 54.5 51.2 60.3 56.3 59.2 67.1 ALI, III . 58.5 74.2 69.8 74.6 65.7 97.1 61.1 53.4 49.4 64.4 66.1 59.2 75.3 ALI, IV . 68.4 72.9 76.5 87.5 68.5 110.7 67.8 63.3 54.9 59.1 78.8 59.2 74.6 TLI ...... 163.0 225.9 M 172.5 196.0 275.7 209.5 125.2 159.3 171.0 140.8 139.2 205.2 CLI ...... 29.1 35.7 M 35.1 30.8 47.1 37.0 40.5 30.4 35.5 35.2 36.2 36.5 CH ...... 7,6 6,6 7,6 6,6 M 7,6,6,6 7,5 6,6 7,5 6,6 7,6 6,6 6,6,7,6 7,7/7,6 7,6/7,5 5,4/5,3 5,2,5,3 5,3/5,3 AH-S, I . 23-14^ 25-38 25-34 26-34T 24-29 + 28-M 24-37 16-22 16-18 + 13-22 11-13­ 10-22 14-21 AH-S, II. 23-33 28-33 + 24-32 27-32 25-26 + 26-M 21-38 19-22 18-25 16-24 15-14 12-24 17-20 AH-S, III 25-29, 27-37 25-24 + 26-35 26-24 30-M 24-35 19-25 18-28 17-23 15-17 12-20 17-19 AH-S, IV 24-M 29-22 + 30-32 31-11-t 29-23 ^ 27+-M 25-22^ 20-22 20-26 19-21 18-15, 12-22 20-22 ELO .... 9 9 9 9 9 9 9 9 9 9 9

Specimen D D D UMML D D D D D D D D M D by sta. 1247 IV 1168 V 1247 IV 31.236 1180 II 1202 IV 1223 III 3981 IV 1289 IV 1243 IV ? 1238 1289 IV

Sex FF F FF FFFFFF FF ML 13.0 11.1 11.0 10.8 10.0 9.8 9.1 9.0 8.8 8.7 8.5 4.5 4.5 .MW I ___ 46.9 54.0 54.5 56.5 63.0 59.1 F 65.9 66.6 56.8 71.2 58.8 68.8 60.0 HWI ___ 40.0 51.3 58.1 40.7 61.0 50.0 64.8 58.0 56.8 71.2 70.5 80.0 66.6 FLI 53.0 54.0 48.1 50.0 47.0 54.0 45.0 44.4 40.9 45.9 50,5 40.0 35.5 FW I .... 88.4 81.0 87.2 74.9 90.0 90.8 89.0 88.8 78.4 91.9 82.3 93.3 66.6 ALI I. . . . 46.1 52.2 64.5 49.0 40.0 61.2 56.0 62.2 55.6 49.4 69.4 42.2 48.8 ALI, 11 . . 55.3 64.8 65.4 69.5 60.0 72.4 71.4 80.0 68.1 57.4 70.5 66.6 60.0 ALI, III . 61.5 81.0 72.7 71.3 61.0 71.4 76.9 77.7 70.4 72.4 71.7 75.5 55.5 ALI, IV. . 60.0 55.8 70.0 66.6 59.0 60.2 65.9 70.0 61.3 68.9 81.1 62.2 14.4 TLI 150.7 173.8 151.8 143.4 119.0 173.4 141.7 145.5 1.53.4 172.4 175.2 160.0 113.3 CLI ...... 39.2 M 43.6 46.3 30.2 40.8 43.9 33.3 0 4 .o 44.8 37.6 42.2 44.4 C H ...... 5,4,6,4 1,0 5,0 5,2 5,3 3,4,3,3 3,0,'4,0 5,0/M :{,0/2,0 0,0 0,0 4,0 3,0 1.2/.M ■M/3,0 O/'O 2,0/2,0 AH-S, I . 7-22 6-19 6-21 8-9 + 8-18 7-21 7-18 5-22 0-26 7-14 3-16 0-19 0-19 AH-S, II. 12-21 7-15 + 9-24 8-12 + 12-17 11-24 11-20 8-22 4-25 9-19 7-20 0-24 0-21 AH-S, III 14-20 10-15^ 11-22 12-14 + 12-20 9-23 11-19 8-21 2-21 9-20 5-12 + 0-20 0-25 AH-S, IV 11-20 0-35 3-34 4-20 + 8-22 4-35 4-33 0-28 1-28 7-34 0-25 0-25 0-22 ELO .... 9 9 9 M 9 9 9 9 9 9 9 M M

M = character missing.

The light organs of the ventral surface of the mantle, funnel, and head, although few, are arranged in the regular adult pattern. It is mainly this feature which permits the identification of the smallest specimens available. At this size the light organs are mainly of one size and constitute the large organs of the adult which has many minute organs as \vell. The nine ocular light organs are small and closely packed in a single row with the terminal organs, even at this stage, considerably larger than the others. These two organs are a rust red in color while the seven intermediates are bronze to cream colored. At around 10 mm mantle length the mantle is proportionately longer and slightly more tapered posteriorly (“ Dana” 1180 II, Fig. 23). The head and mantle widths are two-thirds to one-half the mantle length and are nearly equal to each other. The posterior end of the mantle is slightly longer than in the previously de- Fig. 22. Enoploleulhis anapsis. Juvenile, 8.7 mni mantle length; A. Dorsal aspect; B. Ventral aspect.

Fig. 23. Enoploleulhis anapsis. Juvenile, 10.0mm mantle length; A. Dorsal aspect; B. Ventral aspect. scribed specimen and extends somewhat beyond the posterior border of the fins. The conns of the gladius is broad and slill visible through the skin of the ventral mantle. The fins are more pointed laterally than in smaller specimens, and they are united posteriorly but do not yet extend posteriorly along the tail. The fin width and the fin length are in the same proportion to the mantle length as in smaller specimens and the fin length-width ratio is 1:2. The arm formula is 3.4.2.1 og 3.2.4.1. The arms bear up to a dozen well- developed hooks on II and 111 with fewer on I and IV. The club is better developed than previously and three or four hooks appear in the ventral row. The carpal cluster is just distinguishable, and the distal aboral

Fig. 24. Enoplotealhis anapsis. Late juvenile, 14.2 mm mantle length; A. Dorsal aspect; B. Ventral aspect.

keel is distinct, while the proximal ventral membrane is barely visible. The integument light organs of the ventral mantle, funnel, and head are more numerous and a few minute organs arc present. The ocidar light organs appear larger in relation to the eyeball than in smaller specimens. In specimens between 10 and 27.3 mm the mantle becomes more elongate and tapered and the mantle width decreases from greater than to around Vs of the mantle length. Although the 14.2 mm ML specimen (D 1238 III) shows some deviation from this trend (it appears to have been damaged in preservation), it has been illustrated (Fig. 24) because it is nearly adult in most characters. The posterior lip of the mantle is better developed and extends considerably beyond the posterior margin of the fins. The conus of the gladius is less conspicuous than in younger specimens. The fins are broad and about one-half as long as the mantle length. The fin length-width ratio remains at 1:2 as in younger specimens, although other specimens in the size range between 10 mm and 27 mm show a 2:3 ratio, which I believe is more nearly correct. The anterior fin border forms a lobe anterior to its point of attachment with the dorsal mantle. The lateral border is broad, Disc. BM(NH) O B.M(NH) 1887 M Disc. UMML Dana Dana Dana Dana Dana Specimen: 4743 b 22 ? 23 Type 9095 4743 c 31.237 1292 V 1168 V 1238 III 1287 III 1269 11

Sex...... M F F F F F M F F F I- F I- MI...... 79.0 76.9 73.2 69.0 68.6 64.2 54.0 29.4 27.3 16.9 14.2 1.3.5 l.'ll MW.... 27.2 27.0 25.0 22.3 22.3 25.0 18.2 10.5 10.0 7.8 10.8 6.8 8.9 H\V . . . . 22.9 26.0 21.8 21.0 21.0 23.4 15.5 9.0 8.8 6.3 8.5 6.0 6.0 ir...... 54.1 54.2 53.8 50.4 44.1 37.9 37.6 17.1 13.9 8.3 7.1 7.0 6,8 1- \V .... 58.2 68.0 54.0 56.8 52.0 61.8 40.0 23.2 19.7 13.5 14.1 10.0 14.5 Arm I.. 43.3 54.2 50.4 45.3 39.0 53.0 31.5 14.0 12.1 8.7 8.1 7.0 8.1 II. 46.7 57.0 52.1 48.0 43,5 60.0 33.0 16.0 14.0 10.2 8.0 8.0 9.0 III 46.2 57.0 51.0 51.5 45.1 62.4 33.0 15.7 13.5 10.9 9.4 8.0 10,1 IV 54.0 56.0 56.0 60.3 47.0 71.1 36.6 18.6 15.0 10.0 11.2 8.0 10.0 T L ...... 128.8 178.0 M 119.0 134.5 177.0 113.1 36.8 43.5 28.9 20.0 18.8 27.5 Cl...... 23.0 27.5 M 24.2 21.0 30.3 20.0 11.9 8.3 6.0 5.0 4.9 4.9

Dana Dana Dana UMML Dana Dana Dana Dana Dana Dana Dana Dana Dana Specimen: 1247 IV 1108 V 1247 IV 31.236 1180 II 1202 IV 1223 III 3981 IV 1289 IV 1243 IV 12.38 1289 IV

Sex...... F F F F F F F F F F F F F Ml...... 13.0 11.1 11.0 10.8 10.0 9.8 9.1 9.0 8.8 8.7 8.5 4.5 4.5 MW .. .. 6.1 6.0 6.0 6.1 6.3 5.8 6.0 6.0 5.0 6.2 5.0 3.1 2.7 IIW . . . . 5.2 5.7 6.4 4.4 6.1 4.9 5.9 5.3 5.0 6.2 6.0 3.6 3.0 I-L...... 6.9 6.0 5.3 5.4 4.7 5.3 4.1 4.0 3.6 4.0 4.3 1.8 1.6 1\V__ 11.5 9.0 9.6 8.2 9.0 8.9 8.1 8.0 6.9 8.0 7.0 4.2 3.0 Ann I. . 6.0 5.8 7.1 5.3 4.0 6.0 5.1 5.6 4.9 4.3 5.9 1.9 2.2 II . 7.2 7.2 7.2 7.5 6.0 7.1 6.5 7.2 6.0 5.0 6.0 3.0 2.7 III 8.0 9.0 8.0 7.7 6.1 7.0 7.0 7.0 6.2 6.3 6.1 3.4 2.r> IV 7.8 16.2 7.7 7.2 5.9 5.9 6.0 6.3 5.4 6.0 6.9 2.8 2.0 T[...... 19.6 19.3 16.7 15.5 11.9 17.0 12.9 13.1 13.5 15.0 14.9 7.2 5.1 i;i...... 5.1 •M 4.8 5.0 3.2 4.0 4.0 3.0 4.8 3.9 3.2 1.9 2.0

slightly pointed, and curves into the posterior border, which curves somewhat posteriorly at its junction with the tail. The arms bear between eleven and eighteen hooks with Arm 1 possessing the fewest, .\rnis II and HI an equal number, and Arm IV the greatest number. The tentacles are long and the club has at least five hooks in the ventral row and three or four smaller hooks in the dorsal row. The third and fourth hooks in the ventral row are very large. The carpal cluster distinctly shows the pads, suckers, ridges, aiul grooves. The club membranes are well-developed, especially the distal aboral keel. The integumentary light organs are very numerous with both sizes now present. A single photophore row extends along each lateral margin to the tip of the tail. The ocular light organs are completely formed. The 16.9 mm mantle length specimen ( ‘Dana” 1168 V) has most adult characters well-developed and little change takes place between it and the larger specimens (27.3 to 76.9 mm) except for the increase in the number of arm hooks, integument light organs, etc. The mantle is slender and tapers into a long tail which e.xtends well posterior to the posterior lin border. The mantle and head widths are about one-third of the mantle length. The fm length is greater than one-half the mantle length, and the fm width is three-fourths or more of the mantle length. The fins are proportionately longer in adults because they are measured from the tip of the long tail to the anterior lobe. Laterally, each (in is bluntly pointed and the posterior border is directed diagonally to fuse with the other fin and the tail. Apparently, as the tail grows posteriorly, it “car­ ries” the united fins with it, thus changing the posterior fin border from nearly perpendicular to diagonal to the body axis. With continued growth the number of arm hooks increases so that the largest female speci­ men (76.9 mm mantle length) has 29 hooks on IV, 28 on 11, 27 on 111 and 2.') on 1. Table 10 shows the iiuiiiber of hooks and suckers on the arms of nine adult specimens of E. anapsis. The tentacles are long and robust. 0 1887 Disc. BM(NH) O?BM(NH) Specimen Type 4743 b 22 23

Sex...... F M F F F ML...... 68.6 mm 79.0 mm 76.9 mm 73.2 mm 69.0 mm A rm ...... right left right/left right/left right/left right/left AH S, I . 24-29+ ,2-1-27. 23-14+,24-37 25-38/27-37 25-34/25-28 + 26-34+ 27-254 II. 25-26+ 25-26 23-33 22 35 28-33 + /24-39 24-34,24-14 + 27-32 27-35 r III 26-24/26-16^ 25-29t 23 Ifi I 27-37/25-39 2.5-24/24-22 20-35/25-25 + IV 29-23t 28-27 24-+ .32-20 29-22+/29-22- 30-32 30-31 31-11+30-21 +

Disc. U.M.ML Dana Dana Specimen 4743 c 31.237 1292 V 1168 V

M F FF Ml...... 54.0 mm 29.4 mm 27.3 mm 16.9 mm A rm ...... right/left right/left right left right,left AH-S, I ... . 24-37 20-35 16-22 17-21 16-18+ 16-20 13-22,13-20 11. . . . 21-.38;24-35 19-22 19-22 18-25 18-22 16-24 16-21 III.. . 24-35,23-34 19-23 20-21 18-28 17-24 17-23 17-22 IV . . . 25-22- 29-30 20-20 : 2\-\6^ 20-26/20- • 19-21 18-22

indicates a missing arm tip or lust suckers.

with fully developed clubs, membranes, and carpal clusters. The Ught organs of the ventral mantle, funnel, and head are extremely numerous and of all sizes. Those on the mantle extend well dorso-laterally and also in a single row laterally along the tail. DISCUSSION

Comparison of Enoploteuthis leptura (L e a c h , 1817) and Enoploteuthis anapsis R o p e r , 1964

These hvo closely related species are very similar in general body shape and construction, but there are several points by which they may be readily distinguished. Shape and structure of the mantles are similar, even to the saccate tips of the tails, although E. leplura may be slightly more tapered. It is impossible to assign a specific function to the peculiar saccate structure of the posterior mantle until a detailed examina­ tion of its anatomy has been made. However, gross examination reveals numerous, small, thin-walled cham­ bers fdled with fluid and arranged in a honey-combed pattern, with a thin muscle strand running from the mantle termination to the tip of the tail. These features suggest that the saccate tail may function as a bouv- ancy mechanism. The fins are similar in shape but those of Enoploteuthis anapsis are slightly longer than the fms of Eno­ ploteuthis leptura. Funnel structure is nearly the same, as are the locking apparatus components. The funnel organs differ somewhat between the two species, but there is also considerable variation within each species. The funnel organ of Enoploteuthis leptura appears to have slightly longer, nearly parallel limbs with well- developed flaps on the dorsal component. In Enoploteuthis anapsis these limbs form more of an inverted-V and are somewhat shorter and more stubby than in Enoploteuthis leptura. Flaps are present in Enoploteuthis anapsis as well as a lappet from the broader anterior point of the dorsal member. Apparently, from the material at hand, there is some variation in the structure of the funnel organ within each species. This may reflect differences in preservation and disproportionate growth. The funnel valves are similar in both species. The heads with their conspicuous three nuchal folds are similar. The optic sinus of Enoploteuthis leptura is more pronounced than that of Enoploteuthis anapsis, being narrower and more produced anteriorly. Both species have nine similarly arranged light organs on the ventral surface of the eye-balls. The terminal organs on Enoploteuthis leptura tend to be more circular and more separated from the remaining, smaller seven than in Enoploteuthis anapsis. Both species possess long, nearly subequal arms. The arm tips of Enoploteuthis leptura are not fine and attenuate as they are in Enoploteuthis anapsis, and the sucker bearing area is proportionately longer on E. anapsis. E. anapsis has proportionately a greater number of suckers which gradually decrease to a minute size. The arm suckers of E. leplura are fewer in number, and they become minute very abruptly just proximal to the tip. The sucker rings of anapsis possess 5 to 7 teeth, while E. leplura has 9 longer, more slender teeth. The arm keels are generally similar in construction, but those of Enoploteuthis anapsis are slightly stronger or broader. 'I’he protective trabeculate membranes on the first two arms of both species are very similar; the ventral membranes are very strong, while the dorsal membranes are much weaker. Arm III of both species po.ssesses a strong ventral membrane, but the dorsal membrane of Enoploteuthis leplura, while con­ siderably weaker than the ventral membrane, is somewhat stronger than the corresponding membrane of Enoploteuthis anapsis. Arm IV of Enoploteuthis anapsis possesses very weak dorsal and ventral protective membranes which do not have supporting trabeculae. The protective membranes on Arm IV of Enoplo­ teuthis leptura, however, are well enough developed so that trabeculae are present. The hectocotyli of the two species under discussion may be distinguished by the absence of several pairs of hooks (or suckers) at the distal ends of the rows of hooks in £. anapsis. E. leptura, on the other hand, has all of the hooks and suckers in the normal number and position. The large llap of the ventral protective

Dana-Report No. 66, 1960 membrane does not appear to difTer markedly in the two species, hut the small flap on the dorsal protective membrane is smaller in E. lephirn than in E. onapsis. A further distinct dill'erenrc exists between the males of the two species. The first three pairs of arms in the male of E. leptura possess numerous, minute, fleshy papillae which arc evenly distributed over the entire oral surfaces of the arms between the bases of the hooks and suckers. The first three pairs of arms in E. annpsis, and the foin th pair in both species, are completely devoid of papillation. Although there is no previous mention of papillation in any species of Enoploteuthis, some other mem­ bers of the Knoploteuthidae exhibit this interesting, though unevaluated, character, H o y l k (1904 b) reported ihal Abraliopnis hoylci (= Abrnliopsis affiiiiis (PFuri-EK, 1912)), possesses “ minute conical papillae” on portions of all of the arms of the males. These papillae appear along the edge of each “ lappet” (tiabecula) and at the base of each arm where they increase in number from three to four on the dorsal arm to “about a dozen” on the right ventral arm. The left ventral arm of the male is hectocotylized and also bears papillae. Cni N (1910) recorded similar papillae in a species he called Abraliopsis morisii (^'I.RA^v, 1837), which in all probability should be referred to A. a ffin is or A. lineata (Goodhich, 1896) (Voss, in press). This interesting male-associated character, which appears in some species of at least two genera of the Knoploteuthidae, should be evaluated when more adequate series of specimens become available. A striking dillerence exists between the tentacles and clubs of the two species. The tentacles of Enoplo- teutliis leptura are thin and short, not much longer than the arms, but those of Enoploteuthis anapsis are robust and very long, 2 to 3 limes longer than the arms. The tentacular club of leptura is small and narrow, and bears only a poorly developed dorsal keel. The club of anapsi.i is large and expanded and has broad, well-developeil dorsal and ventral keels. The ilactylus of leptura bears two rows of approximately 10-15 small to minute suckers which decrease in diameter toward the pointed tip. The dactylus of annpsis bears four rows of 40-50 small suckers which decrease slightly in dianieler toward the tip, but at the expanded tip there are fewer than ten enlarged suckers. Both species possess two rows of hooks on the hand with live to seven hooks (occasionally eight) in each row, those in the ventral row being larger and usually one to two more in number. The three to four hooks in the middle of the ventral row are extremely long and large in iinapsis, but those of leptura, w hile larger than the others, are not so out of proportion as those of E. anapsis. The carpal cluster of leptura is long and narrow and has five to six suckers with corresponding knobs, but there are no interlocking ridges and grooves. Enoploteuthis anapsis has a broad, rectangular carpal cluster with three to four suckers and corresponding knobs plus three to four interlocking ridges and grooves. Although the male reproductive systems of the two species do not exhibit notable dilTerences, the s])erma- tophori's of E. leplura and E. anapsis dill'er significantly (I'igs. 4 and 18). The relative lengths of the sperm masses and the cement bodies are the most easily recognizable characters by which the spermatophores of the two species may be distinguished. The sperm mass in E. leptura occupies approximately one-half the total length of the spermatophore, but the sperm mass in E. anapsis occupies only one-third of the total spermatophore length. 'I'he sperm mass extends to the aboral tip of the spermatophore in E. leptura, while in E. anapsis the sperm mass terminates short of the aboral end of the spermatophore. The cement body apparatus in the spermatophores of E. leptura is only about one-fourth the length of the sperm mass, while the cement body of E. anapsis is nearly one-half the length of its sperm mass. The oral end of the cement body of E. leptura has one collar, but there are two distinct collars on the cement body of E. anapsis. Tlie aboral section of the spiral filament in E. leptura appears relatively simply constructed, but the corresponding segment in E. anapsis shows a complex reticulation for about one-third of the length of the spiral filament. Light organ row patterns on the ventral surfaces of the body are very distinct in the two species. On the ventral mantle of Enoploteuthis leplura are seven rows of light organs, the most conspicuous feature being the single row along the posterior midline (excluding the tail) which curves away from the midline anteriorly and joins the first lateral row on either the left or the right side. This first lateral pair of rows is the only pair extending from mantle margin to tail tip. 'I’he lateral-most row on each side is short and sometimes inconspicuous. Enoploteuthis anapsis has only four distinct and two indistinct photophore rows on the ventral mantle, and the entire ventral midline is devoid of light organs. Only the second lateral rows extend to the lip of the tail. Both species have scattered light organs ove r the lateral and dorso lateral siirfaees and a row nearly around the anterior mantle margin. Roth species possess three pairs of rows of photophores ou the funnel. The light organ configuration of the head differs notably in the two species. In the anterior funnel groove of Enoploteuthis leptura there are two small patches of photophores, one on each side of the clear midlinc, which continue anteriorly along the head as the median rows. They extend anteriorly aliuig the ventral aboral border of the ventral arms nearly to the arm tips. The midlinc of the head is free of light organs for its entire length. The anterior funnel groove of Enoploteuthis ftmtp.sis has one median, isolated, triangular patch of light organs. The ventral midline of the head is free from ])hotophores for about three-fourths of the head length anterior to the funnel groove patch. .lust posterior to the junction of arms IV, a row of light organs arises which passes anteriorly on the head, then forks at the base of the ventral arms and sends a row along the ventral aboral border nearly to the tip of each arm 1\'. The second lateral row on the ventral surface of the head of Enoploleiitliis lepiiirn arises at the posterior ventro-laleral angle of the head, passes anteriorly close to the first row, then extends along the fusion of the dorsal aboral border of the ventral arm and base of the tentacular sheath to the tip of IV. The corresponding row in Enoploteulliis diuijisis arises in the same area and passes along the head and to the arm tips in much the same manner as in Enoplotenthifi leptura. There is a short row which branches from this main row lateral to the funnel groove, passes an­ teriorly, then rejoins the main row posterior to the base of IV. This short row is absent from Enoploleulhis leptura. The third lateral row in Enoploteuthis leptura originates lateral to the origin of the second row and extends anteriorly to run along the edge of the tentacular sheath, nearly to the tip of IV. This row is inter­ rupted by the clear window below the ventral eyeball light organs. The corresponding row in Enoploteuthis anapsis is quite similar. The fourth lateral row of Enoploteuthis leptura arises from the mid-nuchal-fold region, passes lateral to the window and merges with the third lateral photophore row posterior to the base of the tentacular sheath on IV. This row has few light organs and may be inconspicuous. Enoploteuthis anapsis has no such row. The lateral-most series of light organs in Enoploteuthis leptura originates as a scat­ tered patch between the eye and the nuchal folds. A row of closely packed light organs passes around the ventral half of the eye opening and terminates at the optic sinus. Dorsal to the optic sinus a row extends from the margin of the eye opening and along the ventral longitudinal base of the swimming nunibrane nearly to the tip of arm 111. The pattern is much the same in Enoploteuthis anapsis except that the row along the base of the swimming membrane does not extend more than two-thirds the length of arm III. The radulae of the two species are similar with the exception that the rachidian tooth in E. leptura bears two small cusps at the base. In summary, Enoploteuthis anapsis may be readily distinguished from Enoploteuthis leptura by (he pos­ session of the following characters: four distinct light organ rows on ventral mantle; entire ventral mantle midline devoid of light organs; ventral aboral light organ rows from ventral arms merge into one short median row on anterior one-fourth of head; only four pairs light organ rows on head (lateral-most row along 111 and eye margin row counted as one); very long, robust tentacles; large, expanded clubs with tlorsal and ventral membranes, very Ivgc hooks in ventral row, dactylus with four rows of numerous small suckers and expanded lip, carpal cluster broad with three to four suckers and knobs and three to four ridges and grooves. Discussion of the Species of Enoploteuthis The following discussion is based on the original descriptions and illustrations of the species in the liter­ ature. The only species which were available for examination were specimens of Enoploteuthis anapsis and Enoploteuthis leptura. Therefore, a preliminary key has been constructed using mainly the arrangement of the hght organs on the ventral surfaces of the mantle, funnel, head, and arms. Of the six nominal species of Enoploteuthis, two are uncertain Ad.\m (1960) questions the generic position of his species because the tentacles are missing from the single specimen known and because of the dorsal attachment of the ventro-lateral buccal connectives. Other characters, especially the number and arrange- 6* nient of ocular light organs and the arrangement of light organs on ventral mantle, funnel, head, and arms, indicate that the species is correctly placed in the genus. Dki.l (1959) described Enoploteuthis neozelanica from New Zealand waters. From the description and the accompanying illustrations it appears that the species is incorrectly referred to Enoploteuthis because it possesses bulbous terminal light organs on the tips of arms IV, it has only “ four” light organs on the ventral eyeball, and the fins are proportionately very large. This would place it in Abraliopsis rather than in Enoplo­

teuthis. According to Voss (personal communication) the species is ^4. gilchristi (R o b s o n ).

A Provisional Key to the Species of Enoploteuthis 1. Light organs on ventral surface of mantle and head arranged in four to eight distinct rows ...... 2. Light organs on ventral surface of mantle not arranged in distinct rows but checkered with circular clear patches. Seven rows of light organs on ventral surface of h e a d ...... Enoploteuthis galaxias B e r r y , 1918. 2. Four distinct light organ rows on ventral mantle; ventral mantle midline clear along its entire length; clear tail... '.i. Seven or eight distinct light organ rows on ventral mantle; midline with light organ row or cluster in posterior one- half of ventral m antle...... 4. 3. Second lateral pair of light organ rows on ventral mantle extends to end of tail. Eight complete light organ rows on ventral surface of head; one short row in anterior head midline posterior to bases of IV which forks onto ventral aboral edge of each arm IV. Light organ row on III extends two-thirds arm length. Buccal connective to ventral oral surface of I I I ...... Enoploteuthis anapsis R o p e r , 1964. Second lateral pair of light organ rows on ventral mantle extends only along anterior one-half of mantle. Six light organ rows on ventral surface of head; no midline row, but light organs scattered between all rows. Row on ventral aboral edge of IV originates on arm and not on head. Row on III extends nearly entire arm length. Buccal connec­ tive attached to dorsal oral surface of III ...... Enoploteuthis dubia A d a m , 1960. 4. Seven distinct light organ rows on ventral mantle. Single midline row merges with first lateral row on either side well short of anterior mantle margin. First lateral rows extend to tail tip; second lateral pair does not extend posterior to midpoint of fin: third lateral pair extends very shortly posterior to mantle margin. Ten more or less distinct light organ rows on ventral head surface; ventral midline of head entirely devoid of light organs. Six light organ rows on fu n n e l...... Enoploteuthis leptura (L e a c h , 1817). Eight distinct light organ rows on anterior one-third of ventral mantle, merging together in posterior two-thirds. Anterior midline clear. Seven light organ rows on ventral surface of head, one row in the midline extending from the funnel groove over the head and forking onto arms IV. Four light organ rows on funnel Enoploteuthis chunii Is h ik a w a , 1914.

Although the key compares character similarities and dilTerences of the species, a few other points deserve

mention. I sh ik a w a (1914) compares Enoploteuthis leptura and Enoploteuthis chunii based mainly on body proportions, with Enoploteuthis chunii having relatively smaller proportions and five longitudinal rows of light organs on the ventral surface of the head instead of the six in Enoploteuthis leptura. The circumorbital rows were excluded from the count. Berry (1918) separates his Enoploteuthus gala-rias from Enoploteuthis leptura and Enoploteuthis chunii by the greater number of light organs and the absence of longitudinal rows of light organs on the ventral surface of the mantle. Other dilTerences of E.gala.vias from Enoploteuthis chunii are the more reduced sucker bearing area on the arms, far fewer suckers on the club dactylns, and the absence of small suckers between the carpal cluster and the rows of hooks. Enoploteuthis dubia A d a m , 1960, likewise has its distinctive light organ arrangement on mantle, head, and arms and has buccal supports to the dorsal oral surface of MI instead of to the ventral oral surface as in the other species. LITERATURE CITED

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(eontlnaed on back of coverj^