BULLETIN OF MARINE SCIENCE, 66(2): 417–443, 2000 NEW SPECIES PAPER
SQUIDS OF THE GENUS ABRALIA (CEPHALOPODA: ENOPLOTEUTHIDAE) FROM THE WESTERN TROPICAL PACIFIC WITH A DESCRIPTION OF ABRALIA OMIAE, A NEW SPECIES
Kiyotaka Hidaka and Tsunemi Kubodera
ABSTRACT Squids of the genus Abralia collected by a midwater trawl in the western tropical Pa- cific were examined. Five species were identified, including Abralia omiae new species. This species is characterized by having five monotypic subocular photophores, two lon- gitudinal broad bands of minute photophores on the ventral mantle, and a hectocotylized right ventral arm with two crests. The other four species identified were A. similis, A. trigonura, A. siedleckyi and A. heminuchalis. Materials from new localities for these spe- cies are discussed and comparisons to related species in several additional characters are made.
Squids of the genus Abralia Gray 1849, family Enoploteuthidae, are important mem- bers in the micronektonic communities in the tropical and subtropical world oceans (e.g., Reid et al., 1991). The genus is distinguished in the family Enoploteuthidae by manus of club with one row of hooks and two rows of suckers, absence of enlarged photophores with black coverage on tips of arm IV, buccal crown with typical chromatophores on aboral surface without any other pigmentation, and five to 12 photophores on eyeball (Young et al., 1998). Several systematic studies on pelagic cephalopods in the Pacific Ocean (Quoy and Gaimard, 1832; Berry, 1909, 1913, 1914; Sasaki, 1929; Grimpe, 1931; Nesis and Nikitina, 1987; Okutani and Tsuchiya, 1987; Burgess, 1992) have revealed that nine nominal Abralia species are known from the Pacific. Subgeneric divisions of this genus have been dis- cussed (Grimpe, 1931; Nesis, 1982, 1987; Nesis and Nikitina, 1987; Tsuchiya and Okutani, 1988; Tsuchiya, 1991), and the scheme proposed by Tsuchiya and Okutani (1988) and modified by Tsuchiya (1991) is applied in the present study. The zoogeography of pelagic cephalopods in the Pacific Ocean has been investigated by Young (1972) off California, by Okutani (1974) in the eastern tropical Pacific, by Nesis (1977) in the western tropical Pacific, by Burgess (1982, 1992) in the central Pa- cific, by Riddell (1985) in New Zealand waters, and by Jefferts (1988) in the northeastern Pacific and the regional distributions of certain species of the genus Abralia have been reported. Tsuchiya (1993) summarized the zoogeography of enoploteuthid squids, in- cluding the genus Abralia, in the Northwest Pacific. During the course of ecological studies on micronektonic squids collected by midwater trawl in the western tropical Pacific, a large number of squids belonging to the genus Abralia were collected. They consist of five species, one of which is an undescribed species, presented here as Abralia omiae new species. The other species identified were A. similis Okutani and Tsuchiya, 1987; A. trigonura Berry, 1913; A. siedleckyi Lipinski, 1983; and A. heminuchalis Burgess, 1992. Specimens from new localities for these species are discussed and comparisons to related species in several additional characters are made.
417 418 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 1. Locations where the trawls were taken in 1995. The star shows the station at which the holotype of Abralia omiae new species was collected.
MATERIALS AND METHODS
Midwater trawl survey: A large midwater trawl with a mouth opening of 20 × 20 m and 8 × 8 mm mesh cod end was towed at 20 stations between the surface and 200 m depth in the area of 3–20°N, 135–160°E from October to December 1995 (Fig. 1, Table 1). The survey was conducted by the FTS OMI-MARU under a skipjack (Katsuwonus pelamis) research program organized by the Tohoku National Fisheries Research Institute, Fisheries Agency of Japan. In total, 75 sampling operations were conducted and 5393 cephalopods were collected, including more than 1000 Abralia from which 50 individuals in good condition were examined as specimens for the taxonomic study. All of these specimens are housed at the National Science Museum, Tokyo (NSMT). MEASUREMENTS OF SPECIMENS.—Methods of presenting squid measurements, counts and compu- tation of indices conform to those proposed by Roper and Voss (1983). Arm length is measured from the first basal hook or sucker to the tip of the arm after Burgess (1992). Abbreviations and terminology agree with those generally accepted for squids.
Table 1. Sampling locations of Abralia specimen.
Setation Leatitud Lgongitud Sgamplin Samplin deepth (m) tim 1−01 20° 00' 138° 00' 1500−120 2255−235 1−03 12° 00' 136° 30' 850−100 0245−034 1−04 9° 00' 136° 30' 1500−120 0245−034 1−06 3° 00' 138° 00' 1020−140 0120−022 1−07 3° 00' 140° 00' 1500−120 2155−225 1−09 9° 00' 141° 00' 1080−200 0010−011 2−03 7° 00' 150° 00' 1560−180 2115−221 2−05 2° 00' 154° 00' 03−200 2230−234 2−06 5° 00' 155° 00' 600−80 2000−210 HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 419
SYSTEMATICS
Family Enoploteuthidae Pfeffer, 1900 Genus Abralia Gray, 1849 Subgenus Pygmabralia Nesis and Nikitina, 1987 Abralia (Pygmabralia) omiae new species (Figs. 2,3,4; Table 2)
HOLOTYPE.—female: ML 26.8 mm, sta. 2-06, 5°00'N, 155°00'E, depth 60–80 m, 27 Nov. 1995, NSMT Mo-68678. PARATYPES.—6 females: ML 24.8 mm (NSMT Mo-68679), 21.4 mm (NSMT Mo-68680), 23.4 mm (NSMT Mo-68681), 24.1 mm (NSMT Mo-68682), 19.0 mm (NSMT Mo- 68683), and 22.7 mm (NSMT Mo-68684), and 3 males: ML 18.2 mm (NSMT Mo-68685), 17.5 mm (NSMT Mo-68686), and 19.3 mm (NSMT Mo-68687), sta. 2-06, 5°00'N, 155°00'E, depth 60– 80 m, 27 Nov. 1995. Diagnosis.—Small Abralia species that matures at 19.0–26.8mm ML in females and at 17.5–19.3mm ML in males; large photophores on the ventral mantle arranged in six trans- verse rows and small photophores around the mesial rows set densely in two broad bands; each arm with 6–13 hooks and distal suckers; tentacular club with one or two hooks, ventral flap and aboral keel; subocular photophores five in number, different in size, and monotypic in structure; right ventral arm of male hectocotylized with two lappets, one bilobate and the other semilunar. Description.—The mantle muscular, short conico-cylindrical, and terminates posteri- orly in a blunt short tail. The tail length about 7–10% of ML. The mantle width 36–41% of ML (Fig. 3A,B). The entire outer surface of mantle, funnel, head and arms covered with a gelatinous epidermis (Fig. 2). The fins rhombic in outline. The fin length 44–52% of ML, and their width 78–87% of ML. The maximum width occurs at about 40% of fin length from anterior end (Fig. 3A,B). The head subcubic in shape and tapers slightly anteriorly, with lateral large eyes. The head little narrower than the mantle opening. The head width 30–37% of ML. The eyelid with a shallow sinus in front. Three pairs of crests present on the posterior periphery of the head: a triangular ridge located at the posterior rim of the funnel groove, a fleshy semilunar crest anterior to the funnel cartilage and a low ridge located behind the eye, respectively (Fig. 4A). The funnel moderate in size (Fig. 4A). The funnel groove broad and shallow. The ante- rior border of the funnel groove continuous to the ventral surface of the head in the middle. The posterior ridge of the funnel groove well developed. The dorsal pad of the funnel organ an inverted-V shape with a short pointed papilla at its anterior end (Fig. 4I). The ventral pads oval in shape (Fig. 4J). The funnel cartilage spatulate with a distinct groove in the middle (Fig. 4E). The mantle cartilage with a simple straight ridge (Fig. 4F). The nuchal cartilage elongated spatulate with a robust mesial longitudinal ridge, and little widened at each end, anterior end wider than the posterior end (Fig. 4D). The arms short for the genus (ALI 27.5–49.2). The arm formula appears IV = II, III = I but variable: Arm IV or Arm II the longest, 34 to 49% of ML. The aboral keel well developed in Arms I to III. The aboral keel of Arm IV continuous to the ventral surface of the arm and covers the base of the tentacle. The width of the aboral keel subequal to the 420 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 2. Abralia omiae new species. Three views of the holotype. Scale bar 10 mm. HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 421
Figure 3. Abralia omiae new species. A. Ventral view of the holotype. B. Dorsal view of the same. Scale bar 10mm. C. Right tentacular club of the holotype. D. Hectocotylized arm of the paratype No. 7. Scale bar 2 mm. 422 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 4. Abralia omiae new species. A. Lateral view of the head of the paratype No. 1. B.Ventral view of the funnel of the same. C. Subocular photophores of the same. D. Nuchal cartilage of the same. E. Left funnel cartilage of the same. F. Left mantle cartilage of the same. G. Gladius of the same. H. Anal pulp of the same. I. Dorsal funnel organ of the same. J. Ventral funnel organ of the same. K. Upper jaw plate of the paratype No. 5. L. Lower jaw plate of the same. M. Distal sucker on the Arm II of the holotype. N. Manus sucker of the same. O. Seminal reseptacle of the paratype No. 5. P. Half row of radular tooth of the same. Q. Spermatophore of the paratype No. 9. Scale: single bar 2 mm; double bar 5 mm; broken bar 0.1 mm. HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 423
Table 2. Measurements, indices and counts of Abralia omiae new species.
H1olotype P2aratype P3aratype P4aratype Paratype N8SMT Mo- 69867 60867 61868 62868 6868 Seex feemal feemal feemal feemal femal D8ML 286. 244. 241. 213. 24. M9WI 356. 379. 490. 318. 36. F7LI 500. 434. 446. 500. 49. F2WI 876. 842. 857. 865. 82. H9WI 366. 340. 396. 229. 33. ALI lrlrlrlrlr I7364. 323. 390. 341. 3-36. 3-53. 229. 28. I3I 397. 316. 315. 315. 491. 474. 339. 432. 345. 34. I6II 333. 311. 357. 395. 396. 386. 333. 336. 300. 29. I3V 317. 358. 337. 378. 389. 328. 400. 358. 341. 34. T8LI 845. 778. 1109. 1408. 1208. 1911. 904. 981. 1600. 99. C0LI 220. 137. 292. 280. 197. 176. 199. 117. 255. 20. AHC I010101+71111011198 I1I 1210101212131310111 I0II 101901111111210101 I1V 1111111111121312121 T1111212211HC
P6aratype 5 P7aratype P8aratype P9aratype Paratype N3SMT Mo- 64868 65868 66868 67868 6868 Seex feemal feemal meal meal mal D0ML 179. 222. 158. 137. 19. M9WI 318. 326. 411. 481. 35. F6LI 551. 408. 510. 485. 50. F1WI 841. 718. 814. 891. 81. H2WI 343. 340. 317. 333. 36. ALI lrlrlrlrlr I7313. 321. 342. 350. 217. 279. 249. 351. 483. 35. I3I 305. 440. 307. 387. 300. 268. 316. 327. 439. 38. I3II 326. 394. 343. 360. 394. 331. 296. 340. 490. 39. I8V 306. 410. 386. 374. 345. 377. 313. 357. 431. 37. T4LI 757. 919. 1018. 889. 961. 1412. 769. 1300. 955. 100. C3LI 261. 262. 251. 263. 138. 290. 242. 223. 197. 18. AHC I5 6 9 9 8 8 7 6 6 6 I6I 811 1 19901987 I6II 6110101019877 I7V 60101888767 T0112000000HC 424 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000 arm width in Arms I and II, and wider than that in Arm III. The protective membrane well developed along the ventral margin of Arms I to III, short trabeculae situated sparsely along the dorsal margin. Arm IV lacks a protective membrane. Each arm with 6–13 hooks and distal suckers. Distal suckers biserial and 12–32 in number, but absent on the hectocotylized arm. Some specimens with one proximal arm sucker. The chitinous ring of the distal suckers with five or six flat teeth on the distal half of the inner margin and numerous small knobs on the inner surface (Fig. 4M). The right Arm IV of males hectocotylized (Fig. 3D). The hectocotylus with two un- equal-sized, offset fleshy crests and 6–8 proximal hooks. The proximal crest large, bilo- bate, and extends from the dorso-distalmost hook to about the distal one third of the arm, situated closely to the dorsal edge of the oral surface. The distal crest small semilunar, situated slightly distal of the proximal crest and does not extends to the distal end of the arm. The terminals of the distal crest attached to the dorsal margin while middle part situated in the middle of oral surface. The protective membrane of the ventral margin expands to a semilunar shape at the distal end of the arm. The modified portion of the hectocotylized arm devoid of armature. The tentacle rather weak and the stalk smooth. The club slightly widened (Fig. 3C). The ventral lappet semilunar in shape, extending from the area between the carpus and the manus to the proximal ventral sucker of the dactylus. The aboral keel extends from the middle of the manus to the distal tip. The carpal group consists of 4–6 suckers and 4–5 pads. The manus with 4–6 dorsal suckers in two longitudinal rows and 1–2 ventral hooks. The dactylus with 75–80 suckers arranged in four longitudinal rows. The chitinous ring of the large suckers of the manus with rectangular teeth along the inner margin and small knobs on the inner surface. The small knobs tri- or quadriserial at distal quarter of the inner surface of the ring while monoserial at the other (Fig. 4N). Integumentary photophores numerous: large photophores with white centers and small darker photophores recognizable in preserved specimens. On the ventral side of the mantle, about 400 small and about 64 large photophores present (Fig. 3A). A photophore-free midventral strip exists. The large photophores set in almost bilateral symmetry. Three rows of large photophores recognizable on both sides of the photophore-free midventral strip, and the small photophores around the mesial rows of large photophores set densely in two broad bands. On the dorsal side of mantle, about 30 small photophores arranged sparsely and in almost bilateral symmetry (Fig. 3B). The ventral side of the funnel with two broad bands of photophores in the middle and a longitudinal row of four photophores on each side (Fig. 4B). Each median broad band composed of one large and about 12 small photophores. Each longitudinal row composed of one large and three small photo- phores. On the ventral surface of head, about 20–24 large and about 150 small photo- phores arranged in almost bilateral symmetry (Fig. 3A). A monoserial row of photo- phores composed of two white and about 10 small photophores recognizable in the mid- line. Photophores in the median area form a strip on each side of the monoserial row. A single row of small photophores on the eyelid. A semicircular row of small photophores with a wide space in the middle situated between the median area and the eyelid. Arm IV with three longitudinal monoserial rows of photophores. The ventral row extends from the base to about 70% of arm length and is composed of one large and about 12–13 small photophores. The mesial row extends from the base to the arm tip and composed of 2–4 large and about 25 small photophores proximally. The dorsal row extends from the base to about 65% of the arm length and composed of 2–3 large and about 10 small photo- HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 425
phores. Arm III with two longitudinal rows of photophores. Each row composed of 3–4 small and dark photophores and one large and white photophore on the base of the arm. Arm II with one small photophore at its dorsal base. Arm I devoid of photophores. The eyeball with five semispherical subocular photophores on the ventral periphery. The terminal and middle photophores larger than the others, all orange in color and mo- notypic in structure (Fig. 4C). Anal palps situated on both sides of the anus. Each anal palp leaf-shaped and attached to the anus by a stalk (Fig. 4H). The buccal membrane with eight lappets and darkly pigmented on the aboral surface, but no pigmentation on the oral surface. The buccal connectives attached to the dorsal side of Arms I, II and IV, and the ventral side of Arm III (DDVD-type). The upper beak with an acute rostrum. The jaw angle about 80°. The hood is broad and covers about a quarter of the crest. The crest with inconspicuous two ridges beneath the hood. The edge of the wing continuous to the hood and terminates at the ventral third of the anterior edge of the lateral wall. The lateral wall thin, fragile, inconspicuously con- cave in the anterior middle part and with no ridge. The rostrum and hood darkly pig- mented and lighter in color toward the transparent edge. The lateral wall darkly pigmented at the anterior half of the crest, lightly pigmented in the posterior half and with a transpar- ent edge (Fig. 4K). The lower beak with a rostrum of an angle of about 70°. The jaw angle curved and about 110°. The shoulder slightly concave. The moderately curved hood covers about half of the crest. The crest length about 85% of the base length. The wing thin and wide in the middle. The lateral wall thin and with a low ridge which runs from upper-anterior part to lower-posterior part, its height longer than the base length. The rostrum and hood colored darkly becoming lighter toward the transparent edge. The upper edge of the hood colored lightly and becoming lighter toward the transparent edge. The upper edge of the hood colored lightly at the rear of jaw angle. The lateral wall colored darkly at the crest and lightly elsewhere, becoming slightly darkened in the middle and transparent at the edges (Fig. 4L). The radular teeth occur in seven longitudinal rows. All teeth unicuspid. The outer mar- ginal tooth the longest, more than twice in length of others (Fig. 4P). The seminal receptacles situated at both sides of the nuchal cartilage on the inner side of the collar, purse-shaped and darkly pigmented (Fig. 4O). The gladius penniform. The rachis strong and tapers posteriorly. Rachis width about 4% of the GL. The conus small. The vane length about 60% of the GL and width about 18% of the GL. The vane thin, lateral blunt angles at posterior 65% of the vane length (Fig. 4G). The walls of vane set at an angle of about 90°. The spermatophores (Paratype 9) large (6.5 mm total length, SPLI 33.7) and slender (0.2 mm). The spiral filament 2.5 mm long with two coils behind the cap. The cement gland cylindrical and 1.3 mm long. The sperm reservoir 2.3 mm long (Fig. 4Q). Remarks.—The present species is assigned to the subgenus Pygmabralia Nesis and Nikitina, 1987, because it has monotypic subocular photophores, one or two hooks on the manus, and no caudal photophore. A. redfieldi Voss, 1955, A. grimpei Voss, 1959, and A. similis Okutani and Tsuchiya, 1987 are the other members of this subgenus. Abralia grimpei was reported from the western North Atlantic off Florida (Voss, 1959). This species is separable from A. omiae based on its larger number of subocular photo- phores (7–8 versus five). 426 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Abralia similis is known to be distributed in the North Pacific (Okutani and Tsuchiya, 1987; Tsuchiya and Okutani, 1988), off Tonga Islands (Ridell, 1985), and equatorial western Indian Ocean and eastern Pacific (Nesis and Nikitina, 1987). This species is separable from A. omiae by photophore arrangement on the ventral surface and hectocotylus struc- ture. Small photophores on the ventral mantle of A. similis are distributed uniformly while they form two broad bands in A. omiae. Small photophores on the median area of the ventral surface of the head are scattered uniformly in the A. similis while a monoserial row in the midline is recognizable in A. omiae. The proximal bilobate crest of the hectocotylus of A. similis is situated along the ventral margin of the right Arm IV, while that of A. omiae is situated closely to the dorsal edge of the oral surface. The distal crest of A. similis is situated along the dorsal margin of the oral surface while that of A. omiae is situated with its terminals attached to the dorsal margin and middle part in the middle of oral surface. Abralia redfieldi was reported from the western Atlantic off Cuba (Voss, 1955), and from the western Atlantic from New England to Argentina (Nesis, 1975; Cairns, 1976; Lu and Roper, 1979; Stephen, 1982; Lipinski, 1983; Nesis and Nikitina, 1987) and eastern Atlantic off Dakar (Lu and Clarke, 1975). Abralia redfieldi and A. omiae both have five subocular photophores and similar photophore arrangements on the ventral mantle. Sper- matophore morphology and indices are also very close to each other (Hess, 1987). The differences between A. redfieldi and A. omiae are as follows: A. redfieldi has three hooks on manus while A. omiae has one or two; A. redfieldi has larger number of hooks on each arm than A. omiae (Arm I: 11–16 vs 5–10, Arm II: 12–16 vs 6–13, Arm III: 11–16 vs 6– 12, Arm IV: 6–13 vs 10–16 (normal) and 9-–10 vs 6–8 (hectocotylized)); the arms of A. redfieldi are longer than those of A. omiae on average (ALI I: 37.8 vs 32.3, ALI II: 44.3 vs 37.6, ALI III: 42.5 vs 34.0, ALI IV: 47.5 vs 37.2); the semicircular photophore row on the ventral side of the head is continuous in A. redfieldi, but is widely spaced in the middle in A. omiae. Etymology.—The specific name refers to the FTS OMI-MARU, the ship used to collect the material for this study.
THE OTHER WESTERN TROPICAL PACIFIC ABRALIA COLLECTED
Of the 19 hitherto known species of Abralia (Young et al., 1998; Sweeney and Roper, 1998), A. trigonura Berry, 1913, A. siedleckyi Lipinski, 1983, A. similis Okutani and Tsuchiya, 1987, and A. heminuchalis Burgess, 1991 were identified from the present material. A brief description was given and comparison with previous description was made for each species.
Abralia (Pygmabralia) Nesis and Nikitina, 1987 Abralia (Pygmabralia) similis Okutani and Tsuchiya, 1987 (Fig. 5, Table 3)
Material Examined.—5 females: ML 19.3 mm (NSMT Mo-68689), 18.5 mm (NSMT Mo-68690), 19.9 mm (NSMT Mo-68692), 18.1 mm (NSMT Mo-68693), and 20.2 mm (NSMT Mo-68694), and 3 males: ML 18.8 mm (NSMT Mo-68688), 20.3 mm (NSMT Mo-68691), and 20.6 mm (NSMT Mo-68695), sta. 1-01, 20°00'N, 138°00'E, depth 100–120 m, 29 Oct. 1995. HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 427
Figure 5. Abralia similis. A. Ventral view. B. Right tentacular club. C. Hectocotylized arm. D. Spermatophore. Scale: single bar 2 mm; double bar 5 mm.
Description.—The mantle width about 42–47% of ML. The tail length about 7–13% of ML. The fins moderate, length 44–54% of ML, width 84–100% of ML. The dorsal pad of the funnel organ an inverted V, carrying a ridge on about 1/3 of anterior edge and a papilla at the anterior end. Ventral pads oval. The arms short for the genus (ALI 24.5–45.2). The 428 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
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Se HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 429
arm formula 2.4.3.1 or 4.2.3.1. Each arm with 6–11 hooks and distal suckers. Distal suckers biserial. Some specimens bear one proximal sucker. The right Arm IV of males hectocotylized (Fig. 5C). The hectocotylus with two un- equal-sized, offset fleshy crests and 6–8 proximal hooks. The proximal crest large, bilo- bate, and extends from the dorso-distalmost hook to about the distal one third of the arm on the dorsal side. The distal crest semilunar, situated on the dorsal margin of oral sur- face, slightly distal of the proximal crest and does not extends to the distal end of the arm. The protective membrane of the ventral margin expands to a semilunar shape at the distal end of the arm. The modified portion of the hectocotylized arm devoid of armature. Tentacles short for the genus (TLI 91.3–126.3), with ventral lappet and aboral keel. The manus with 1–2 ventral hooks in one row and eight suckers in two rows. The dactylus with four longitudinal rows of suckers (Fig. 5B). Integumentary photophores numerous: large photophores with white centers and small darker photophores recognizable in preserved specimens. On the ventral side of mantle, about 450 small and 74 large photophores present. An inconspicuous photophore-free midventral strip exists. Large photophores set almost symmetrical and small photophores scattered around and between them (Fig. 5A). Ventral side of the funnel with two broad bands of photophores in the middle and a short strip of photophores on each side. Each median band with two large and about 15 small photophores. Each lateral strip with one large and three small photophores. On the ventral surface of head, about 26 large photophores arranged in almost bilateral symmetry. Small photophores scattered uniformly in the median area, set in a row on the eyelid, and form a semicircular row between the median area and the eyelid (Fig. 5A). Arm IV with three longitudinal rows of photophores. Each row monoserial. The ventral row composed of four large and about 20 small photophores. The mesial row composed of eight white and about 30 small photophores. The dorsal row composed of 2–3 white and about 15 small photophores. The eyeball with five semispherical subocular photophores on the ventral periphery. The terminal and middle photophores larger than the others, all orange in color and mo- notypic in structure. Seminal receptacle not observed because of lack of female specimen in mature. Spermatophores large (7.7 mm total length, SPLI 37.3), slender (0.3 mm). Spiral fila- ment 3.6 mm long, three coils behind cap. Cement gland cylindrical, 1.1 mm long. Sperm reservoir 2.5 mm long (Fig. 5D). Remarks.—Abralia (Pygmabralia) similis Okutani and Tsuchiya, 1987 was first de- scribed from subtropical North Pacific. The present materials agree well with the original description in photophore arrangements and hectocotylization of right arm IV of the male. Numerical characters are also comparable to those in Okutani and Tsuchiya (1987). Com- parison of this species with the other species of the subgenus Pygmabralia in the present study area, A. omiae, is given in the remarks of A. omiae. 430 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Abralia Gray, 1849 Abralia (Heterabralia) Tsuchiya and Okutani, 1988 Abralia (Heterabralia) trigonura Berry, 1913 (Fig. 6, Table 4)
Material Examined.—5 males: ML 28.4 mm (NSMT Mo-68699), 29.9 mm (NSMT Mo-68701), 30.3 mm (NSMT Mo-68703), 29.8 mm (NSMT Mo-68704), and 28.2 mm (NSMT Mo-68705), and 4 females: ML 22.4 mm (NSMT Mo-68698), 33.9 mm (NSMT Mo-68702), 30.9 mm (NSMT Mo- 68706), and 35.1 mm (NSMT Mo-68707), sta. 1-01, 20°00'N, 138°00'E, depth 100–120 m, 29 Oct. 1995. 1 female: ML 32.9 mm, sta. 1-03, 12°00'N, 136°30'E, depth 80–100 m, 2 Nov. 1995, NSMT Mo-69696. 1 female: ML 32.5 mm, sta. 1-04, 9°00'N, 136°30'E, depth 100–120 m, 3 Nov. 1995, NSMT Mo-68697. 1 male: ML 31.3 mm, sta. 2-03, 7°00'N, 150°00'E, depth 160–180 m, 21 Nov. 1995, NSMT Mo-68700. Description.—The mantle width about 33–41% of ML. The tail length about 13–17% of ML. The fins large, length 60–66% of ML, width 94–104% of ML. The dorsal pad of the funnel organ an inverted V with weak ridges. Ventral pads oval. The arms moderate in length (ALI 31.7–50.3). The arm formula variable: Arm IV or Arm II the longest and Arm I the shortest in most specimen. Each arm with 8–16 hooks and distal suckers. Distal suckers biserial. The left arm IV of males hectocotylized. The hectocotylus with two offset crests and 11–14 proximal hooks. Ventral crest larger and positioned posterior to the other. Distal suckers absent (Fig. 6C). Tentacles long (TLI 86.2–161.0), with ventral lappet and aboral keel. The manus with 2–4 ventral hooks in one row and 6-8 manus suckers. The dactylus with suckers arranged in four longitudinal rows (Fig. 6B). Integumentary photophores numerous: large photophores with white centers and small darker photophores recognizable in preserved specimens. On the ventral side of mantle, 650–700 small and about 70 large photophores present. A photophore-free midventral strip exists occasionally, especially for small specimen. The large photophores set in al- most bilateral symmetry and the small photophores scattered around and between them (Fig. 6A). Ventral side of the funnel with two broad bands of photophores in the middle and a short strip of photophores on each side. Each median band with one large and 22 small photophores. Each lateral strip with one large and six small photophores (Fig. 6D). On the ventral surface of head, about 20 large photophores arranged in almost bilateral symmetry. Small photophores scattered in the median area, set in a row on the eyelid, and form a semicircular row between the median area and the eyelid (Fig. 6A). Arm IV with three longitudinal rows of photophores. The ventral row monoserial and composed of 4–5 large and about 40 small photophores. The mesial row either mono- or biserial and composed of five white and about 40 small photophores. The dorsal row monoserial and composed of 1–2 white and 15–20 small photophores (Fig. 6A–F). The eyeball with five subocular photophores present on the ventral periphery of the eyeball. Terminal photophores oval, creamy white, and larger than mesial three, posterior terminal one extra large. Mesial three photophores small, round and orange, of which the middle one larger than the others. Seminal receptacle sac-like, situated posterior to nuchal cartilage (Fig. 6E). HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 431
Figure 6. Abralia trigonura. A. Ventral view. B. Right tentacular club. C. Hectocotylized arm. D. Ventral view of funnel. E. Seminal reseptacle. F. Ventral views of the right Arm IV. G. Spermatophore. Scale: single bar 2 mm; double bar 5 mm. 432 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Table 4. Measurements, indices and counts of Abralia trigonura Goodrich, 1896.
NSMT M6o- 67869 68869 69869 60869 61870 6870 Seex feemal feemal feemal meal meal mal D9ML 352. 342. 242. 238. 391. 29. M1WI 363. 384. 355. 399. 386. 39. F9LI 672. 603. 685. 664. 695. 60. F9WI 983. 958. 909. 1500. 925. 101. H1WI 302. 332. 360. 462. 346. 30. ALI lrlrlrlrlrlr I8315. 387. 413. 451. 315. 307. 327. 337. 490. 388. 353. 37. I7I 348. 421. 439. 570. 464. 462. 420. 454. 402. 475. 416. 39. I5II 318. 460. 434. 425. 379. 348. 430. 451. 480. 470. 331. 38. I9V 433. 421. 497. 438. 424. 417. 424. 463. 463. 432. 410. 41. T0LI 1-437. 1561. 1348. 1121. 1821. 1412. 1914. 981. 907. 947. 114. C7LI 2-7. 207. 9296. 284. 291. 223. 220. 128. 200. 148. 20. AHC I911 1 1211111212111211101 I8I 13 2 1313131214131313131 I1II 11111110101411111111121 I1V 12111412131112131213121 THC 4-4334334433 4- 4334 334433
NSMT M2o- 63870 64870 65870 66870 67870 6870 Seex feemal meal meal meal feemal femal D9ML 333. 380. 229. 298. 310. 35. M1WI 399. 368. 366. 410. 450. 39. F8LI 559. 664. 675. 602. 633. 60. F3WI 1504. 1500. 997. 1302. 957. 99. H2WI 354. 314. 336. 307. 395. 32. ALI lrlrlrlrlrlr I6463. 337. 336. 353. 382. 313. 357. 308. 369. 336. 316. 36. I5I 442. 425. 490. 460. 384. 349. 422. 482. 481. 432. 402. 39. I3II 490. 306. 359. 397. 366. 377. 326. 388. 491. 401. 319. 39. I6V 458. 402. 431. 358. 317. 398. 443. 481. 442. 461. 493. 40. T3LI 1911. 1519. 1625. 1238. 836. 829. 1512. 805. 1636. 1611. 1231. 110. C4LI 265. 202. 230. 169. 138. 179. 188. 240. 119. 252. 271. 21. AHC I913 1 1214131312141514141 I2I 12141415141615151515131 I1II 1119512121415161614121 I2V 1118311141014131515151 THC 343333343442 3 43333343442
Spermatophores small (4.7 mm total length, SPLI 16.7), slender (0.2 mm). Spiral fila- ment 1.7 mm long, three coils behind cap. Cement gland cylindrical, 0.7 mm long. Sperm reservoir 2.3 mm long (Fig. 6G). Remarks.—Abralia (Heterabralia) trigonura Berry, 1913 was first described from Ha- waiian water and was recently redescribed by Burgess (1992). The present materials agree well with the original and redescription in photophore arrangements and hectocotylization HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 433
of left arm IV of the male. Spermatophore morphology agrees with Hess’s (1987) de- scription and illustration. The maximum values of FLI and FWI of the present A. trigonura specimens are smaller than that reported by Burgess (1992) based on Hawaiian material (FLI 66 vs 71; FWI 104 vs 125), however, the ranges are comparable (FLI 59.8–65.6 vs 61.5–71.4, FWI 93.9–104.0 vs 88.9–125.1). The other numerical characters are compa- rable to those in original description and Burgess (1992).
Abralia (Heterabralia) siedleckyi Lipinski, 1983 (Fig. 7, Table 5)
Material Examined.—3 females: ML 30.2 mm (NSMT Mo-68708), 37.6 mm (NSMT Mo-68709), and 34.3 mm (NSMT Mo-68710), sta. 1-04, 9°00'N, 136°30'E, depth 100–120 m, 3 Nov. 1995. 3 females: ML 27.2 mm (NSMT Mo-68711), 26.3 mm (NSMT Mo-68712), and 29.8 mm (NSMT Mo-68713), and 3 males: ML 25.8 mm (NSMT Mo-68714), 26.2 mm (NSMT Mo-68715), and 26.3 mm (NSMT Mo-68716), sta. 1-06, 3°00'N, 138°00'E, depth 120–140m, 9 Nov. 1995. 1 female ML 33.3 mm, sta. 1-09, 9°00'N, 141°00'E, depth 180–200 m, 12 Nov. 1995, NSMT Mo-68717. Description.—The mantle width about 37–43% of ML. Tail about 14–20% of ML. The fins large, length 64–71% of ML, width 97–110% of ML. The dorsal pad of funnel organ inverted V with a distinct fleshy ridge on top and a papilla at the anterior end. Ventral pads oval. The arms moderate in length (ALI 39.0–58.9). The arm formula 4.2.3.1 or 4.2.1.3. Each arm with 11–19 hooks and distal suckers. Distal suckers biserial. The left arm IV of males hectocotylized. The hectocotylus with two offset crests and 16 proximal hooks. Ventral crest larger and positioned posterior to the other. Distal suckers absent (Fig. 7C). Tentacles long (TLI 106.0–220.4), with ventral lappet and aboral keel. The manus with 2-3 ventral hooks in one row and 7–8 manus suckers. The dactylus has suckers arranged in four longitudinal rows (Fig. 7B). Integumentary photophores numerous: large photophores with white centers and small darker photophores recognizable in preserved specimens. On the ventral side of mantle, about 750 small and about 160 large photophores present. Both types of photophores scattered and especially dense in the posterior 2/3 of the midventral surface of the mantle. There is no midventral strip, except for small specimen (Fig. 7A). Ventral side of the funnel with two broad bands of photophores in the middle and a short strip of photophores on each side. Each median band with one large and about 30 small photophores. Each lateral strip with one large and 5–7 small photophores (Fig. 7D). On the ventral surface of head, about 20 large photophores arranged in almost bilateral symmetry. Small photophores scattered in the median area, set in a row on the eyelid, and form a semicircular row between the median area and the eyelid (Fig. 7A). Arm IV with three longitudinal rows of photophores. The ventral row monoserial and composed of about 10 white and about 45 dark photophores. The mesial row tri- or quadriserial and composed of about 10 white and about 80 dark photophores. The dorsal row monoserial and composed of about 30 dark photophores (Fig. 7A,G). The eyeball with five subocular photophores on the ventral periphery. Terminal photo- phores oval, creamy white, and larger than mesial three, posterior terminal one extra large. Mesial three photophores small, round and orange, the middle one larger than the others. Seminal receptacle not sac-like, but rugose pad (Fig. 7E). 434 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 7. Abralia siedleckyi. A. Ventral view. B. Right tentacular club. C. Hectocotylized arm. D. Ventral view of funnel. E. Seminal reseptacle. F. Spermatophore. G. Ventral views of the right Arm IV. Scale: single bar 2 mm; double bar 5 mm. HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 435
Table 5. Measurements, indices and counts of Abralia siedleckyi Lipinski, 1983.
NSMT M8o- 69870 60870 61871 62871 6871 Seex feemal feemal feemal feemal femal D2ML 360. 337. 324. 237. 26. M8WI 386. 388. 358. 307. 43. F6LI 626. 645. 791. 667. 69. F7WI 967. 1905. 1109. 1209. 112. H7WI 461. 318. 359. 475. 43. ALI lrlrlrlrlr I7415. 492. 464. 483. 447. 496. 401. 493. 407. 46. I7I 438. 418. 571. 485. 552. 552. 498. 475. 551. 52. I7II 446. 403. 461. 462. 442. 406. 460. 432. 423. 45. I7V 500. 570. 479. 419. 567. 506. 541. 591. 502. 54. T4LI 1941. 1961. 1930. 1451. 2020. 2816. 1170. 1574. 1747. 135. C9LI 127. 210. 188. 156. 204. 253. 159. 159. 280. 19. AHC I31214121313131512141 I5I 1414151515151515131 + I3II 13141412111313121 + 10+ I6V 16171715151514131 + 15 T2HC + 6+ 54323232
NSMT M3o- 64871 65871 66871 67871 6871 Seex feemal meal meal meal femal D8ML 289. 225. 236. 236. 33. M2WI 387. 328. 411. 481. 37. F4LI 697. 720. 637. 644. 69. F7WI 1507. 1208. 1604. 966. 103. H6WI 480. 463. 421. 319. 38. ALI lrlrlrlrlr I6491. 339. 485. 474. 437. 437. 478. 408. 379. 41. I3I 531. 498. 531. 522. 459. 488. 594. 528. 476. 44. I9II 401. 455. 471. 455. 433. 454. 502. 501. 339. 42. I3V 571. 520. 531. 534. 545. 593. 565. 578. 467. 48. T3LI 1845. 1963. 1160. 1365. 1744. 1742. 1240. 1061. 1506. 125. C5LI 283. 221. 250. 220. 210. 282. 202. 234. 261. 21. AHC lrlrlrlrlr I51416141412161512131 I5I 141 + 157 1615181815151 I2II 1315141313161613131 I6V 1613161310221917161 THC 2 322223333 436 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Spermatophores small (3.3 mm total length, SPLI 12.5), slender (0.1 mm). Spiral fila- ment 1.1 mm long, two coils behind cap. Cement gland cylindrical, 0.4 mm long. Sperm reservoir 1.3 mm long (Fig. 7F). Remarks.—Abralia siedkeckyi Lipinski, 1983 was described from South Atlantic water off Capetown. The original description based on three specimens is the only record of this species. The present materials agree well with the original description in photophore ar- rangements, hectocotylization of left arm IV of the male, and spermatophore morphol- ogy. Specimens of A. siedleckyi from the study area have wider heads than reported in the originally described for South Atlantic material (HWI 38.1–45.5 vs 21.2–25.8), such in- traspecific variations are often seen in other species of the genus (e.g. A. similis in Bur- gess, 1992). The other numerical characters are comparable to those in Lipinski (1983).
Abralia (Heterabralia) heminuchalis Burgess, 1992 (Fig. 8, Table 6)
Material Examined.—1 female: ML 22.0 mm, sta. 1-03, 12°00'N, 136°30'E, depth 80–100 m, 2 Nov. 1995, NSMT Mo-68718. 1 female: ML 25.1 mm, sta. 1-04, 9°00'N, 136°30'E, depth 100–120 m, 3 Nov. 1995, NSMT Mo-68727. 1 male: ML 25.5 mm (NSMT Mo-68719) and 1 female: ML 21.2 mm (NSMT Mo-68720) sta. 1-07, 3°00'N, 140°00'E, depth 100–120 m, 9 Nov. 1995. 2 females: ML 26.2 mm (NSMT Mo-68721) and 27.7 mm (NSMT Mo-68722), sta. 1-06, 3°00'N, 138°00'E, depth 120–140 m, 9 Nov. 1995. 3 females: ML 24.3 mm (NSMT Mo-68724), 30.3 mm (NSMT Mo-68725), and 35.3 mm (NSMT Mo-68726), and 1 male: ML 21.9 mm (NSMT Mo-68723), sta. 2-05, 2°00'N, 154°00'E, depth 0–200 m, 25 Nov. 1995. Description.—The mantle width about 36–52% of ML. The tail length about 17–25% of ML. The fins large, length 63–72% of ML, width 99–117% of ML. The dorsal pad of the funnel organ inverted V with a papilla at the anterior end. Ventral pads oval. The arms moderate in length (ALI 40.0–63.3). The arm formula 4.2.3.1. Each arm with 11–21 hooks and distal suckers. Distal suckers biserial. Arm I-III bear one proximal sucker occasionally. The left arm IV of males hectocotylized. The hectocotylus with two offset crests and 17–18 proximal hooks. Ventral lappet larger and positioned posterior to the other. Distal suckers absent (Fig. 8B). Tentacles long (TLI 118.1–179.5), with ventral flap and aboral keel. The manus with 2–3 ventral hooks in one row and 7–8 manus suckers. The dactylus with suckers arranged in four longitudinal rows (Fig. 8C). Integumentary photophores numerous: large photophores with white center and small darker photophores are recognizable in preserved specimens. On the ventral side of mantle, about 750 small and about 160 large photophores present. There is distinct circular space around large white photophores which located on anterior 1/2 of the mantle. There is no midventral strip (Fig. 8A). Ventral side of the funnel with two broad bands of photophores in the middle and a short strip of photophores on each side. Each median band with one large and about 30 small photophores. Each lateral strip with one large and 5–7 small photophores (Fig. 8D). On the ventral surface of head, about 20 large photophores arranged in almost bilateral symmetry. Small photophores scattered in the median area, set in a row on the eyelid, and form a semicircular row between the median area and the eyelid (Fig. 8A). HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 437
Figure 8. Abralia heminuchalis. A. Ventral view. B. Hectocotylized arm. C. Right tentacular club. D. Ventral view of funnel. E. Seminal reseptacle. F. Ventral views of the right Arm IV. G. Spermatophore. Scale: single bar 2 mm; double bar 5 mm. 438 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Table 6. Measurements, indices and counts of Abrlia heminuchalis Burgess, 1991.
NSMT M8o- 69871 60871 61872 62872 6872 Seex feemal meal feemal feemal femal D0ML 2m2. m52m5. m22m1. m22m6. m72m7. m M9WI 348. 414. 497. 450. 35. F2LI 686. 667. 682. 656. 69. F2WI 1407. 1312. 938. 1706. 100. H1WI 491. 386. 356. 334. 31. ALI l rlrlrlrlr I4430. 473. 416. 503. 440. 450. 431. 472. 474. 47. I7I 468. 498. 553. 574. 498. 435. 520. 444. 533. 50. I5II 485. 406. 497. 552. 446. 434. 404. 423. 484. 43. I9V 499. 439. 653. 554. 489. 506. 542. 510. 551. 51. T9LI 1845. 1949. 1451. 1574. 1379. 1560. 1469. 1771. 1837. 142. C1LI 210. 139. 244. 244. 273. 285. 274. 204. 264. 22. AHC I31315141111121113131 I5I 1516161314151414151 I3II 1314141211131113141 I6V 1618102313121 + 13+ 175 1 THC 3312 01 2233
NSMT M3o- 64872 65872 66872 67872 6872 Seex meal feemal feemal feemal femal D9ML 2m1. m32m4. m33m0. m33m5. m12m5. m M9WI 581. 483. 410. 441. 46. F0LI 693. 682. 730. 695. 71. F5WI 1716. 1609. 1802. 1406. 103. H7WI 336. 305. 440. 366. 36. ALI lrlrlrlrlr I6477. 407. 463. 474. 473. 464. 456. 466. 476. 49. I3I 581. 556. 426. 434. 438. 520. 590. 531. 501. 56. I3II 466. 407. 466. 426. 465. 404. 407. 416. 571. 48. I7V 527. 557. 521. 570. 543. 512. 588. 595. 660. 57. T--6LI 1120. 1518. 1545. 1245. 146. --- C--8LI 117. 107. 204. 255. 22. --- AHC I41615131515151413151 I5I 141 + 167 1615151515141 I5II 1214131314131413121 I7V 1129171616191026161 THC --43333 ---
Arm IV with three longitudinal rows of photophores. The ventral row monoserial and composed of about 5–6 white and about 50 dark photophores. The mesial row either tri- or quadriserial and composed of about 10 white and about 75 dark photophores. The dorsal row monoserial and composed of about 25 dark photophores (Fig. 8F). HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 439
The eyeball with five subocular photophores on the ventral periphery. Terminal photo- phores oval, creamy white, and larger than mesial three, posterior terminal one extra large. Mesial three photophores small, round and orange. Among mesial photophores, the middle one is larger than the others. Seminal receptacle not sac-like, but rugose pad (Fig. 8E). Spermatophores small (3.6 mm total length, SPLI 14.1), slender (0.2 mm). Spiral fila- ment 1.3 mm long, two coils behind cap. Cement gland cylindrical, 0.4 mm long. Sperm reservoir 1.3 mm long (Fig. 8G). Remarks.—Abralia heminuchalis Burgess, 1992 was described from the Central Equa- torial Pacific. The present materials agree well with the original description in photo- phore arrangements and hectocotylization of left arm IV of the male. Numerical charac- ters are also comparable to those in Burgess (1992). Morphological characters of these four species and Abralia omiae are summarized in Table 7.
COMPARISON AMONG THE SUBGENUS HETERABRALIA SPECIES
Among the five species in the present study, Abralia trigonura, A. siedleckyi and A. heminuchalis, belong to the subgenus Heterabralia by having two types of subocular photophores, a hectocotylized left ventral arm, biserial arm sucker arrangement, and no caudal photophores. These three species closely resemble each other, but detailed mor- phological comparisons have not yet been made. During the present study, we recognized that the photophore arrangements of the ventral side of the funnel and ventral arms are important specific characters. These characters are described in the section for each spe- cies above. Type of seminal receptacle is also described for each species and used to separate species, as it is known to be specific character of the species of Abralia (Burgess, 1998). A. trigonura is easily separable from the other two species by having fewer photophores on the ventral side of funnel and fewer photophores on the mesial photophore rows of the ventral arms. A. trigonura is also different to the other two species in having sac-like seminal receptacles. A. siedleckyi and A. heminuchalis have the same photophore arrange- ments on the ventral side of funnel and ventral arms and the same type of seminal recep- tacle, however, A. heminuchalis has a distinct circular space around the large white photo- phores on the ventral mantle while A. siedleckyi does not. A. heminuchalis is also sepa- rable from A. siedleckyi by the occasional presence of proximal suckers on all arms ex- cept the hectocotylized arm. In addition to A. heminuchalis, A. siedleckyi and A. trigonura, the subgenus Heterabralia includes one other species, A. andamanica Goodrich, 1896 (We regard A. robsoni Grimpe, 1931 as a subspecies of A. andamanica after Nesis [1982, 1987]), but no specimen were examined in this study. Judging from the description and figures presented by Lipinski (1983) and Burgess (1992), this species seems to have the same photophore arrange- ments on the ventral mantle, funnel and arms as A. trigonura. A. andamanica has 9–10 distinct rows of photophores on the ventral head, while those of the other three species are diffused. A. andamanica may also be separable from the others by having “whip-like” arm tips (Voss, 1963). 440 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
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S55555 HIDAKA AND KUBODERA: SQUIDS OF ABRALIA FROM THE WESTERN TROPICAL PACIFIC 441
GEOGRAPHIC DISTRIBUTION
Abralia similis has a broad distribution in the tropical Indo-Pacific, in which the study area is included (see remarks for the species description of A. omiae for detail). Abralia trigonura has been recorded from Hawaiian waters (e.g., original description; Hess, 1987; Burgess, 1992; Reid et al., 1991) and Northwest Pacific (Tsuchiya, 1993). Our specimens represent the southernmost record for A. trigonura. Abralia siedleckyi has only been recorded from the South Atlantic off Cape Town (original description), thus this is the second record for the species. These two widely separated records suggest that this species may possibly have a broad distribution. Further investi- gation should be necessary to reveal the range and continuity of distribution area of this species. Abralia heminuchalis has been recorded from the central equatorial Pacific (original description). The specimen of Abralia andamanica reported by Okutani (1974) from the eastern equatorial Pacific has a similar photophore arrangement as A. heminuchalis but the species identity of the material has not yet been made because of the lack of the information about seminal receptacle (Burgess, 1992). Therefore this is the second record of the species. These records suggest that this species is distributed in the western to central area of the equatorial Pacific. In total, five species of genus Abralia were found in the study area. The number of species is comparable to or slightly larger than in other localities as suggested by previous studies of pelagic areas of the Pacific: Okutani, 1974 (two species, eastern tropical Pa- cific); Riddell, 1985 (three species, Tonga Island waters); Burgess, 1992 (four species, central tropical Pacific); Tsuchiya, 1993 (three species, northwest Pacific). The present study area shares the occurrence of A. trigonura, A. similis and A. heminuchalis with the central tropical Pacific; the first two species have also been reported from the Northwest Pacific. Among the three species reported from off New Zealand, only A. similis was found in the present material. Thus, the species of Abralia from the present study area mainly consist of north tropical Pacific species sensu Tsuchiya (1993), however, the fauna of the area may be distinguished from the broader region by the added occurrence of A. omiae and A. siedleckyi.
ACKNOWLEDGMENTS
Samples for this study were offered kindly from the Tohoku National Fisheries Research Insti- tute. We are also grateful to K. Tsuchiya of the Tokyo University of Fisheries for his beneficial comments and encouragement. This study would not have been possible without the assistance and co-operation of the captain, officers and crew of the FTS OMI-MARU. We send our gratitude to L. A. Burgess, New Jersey and M. Lipinski, University of Cape Town for their encouragement and ac- commodation in the loan of the type specimen of Abralia heminuchalis and Abralia siedleckyi which were examined preparatorily for our reference. We are also grateful to the National Museum of Natural History, Washington D.C. and the Zoological Museum, Copenhagen for the loan of the type specimen of above species. We thank T. Okutani for kind and meaningful comments on an earlier draft of the manuscript. 442 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
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DATE SUBMITTED: March 20, 1999. DATE ACCEPTED: November 11, 1999.
ADDRESSES: (K.H.) Planktonology Division, Ocean Research Institute, University of Tokyo, 1-15-1 Minamidai, Nakano-ku, Tokyo, 164-8639 Japan. (T.K) Department of Zoology, National Science Museum, 3-23-2 Hyakunincho, Shinjuku-ku, Tokyo, 169-0073 Japan.