Eledonine Octopods from the Southern Ocean

Antarctic Science 6 (2): 205-214 (1994) Eledonine octopods from the Southern Ocean: systematics and distribution TSUNEMI KUBODERA' and TAKASHI OKUTANP 'National Science Museum Tokyo,Deparment of Zoology, 3-23-1Hyahnin-cho, Shinjuh-ka, Tokyo 169, Japan 2Tokyo University of Fzkheries, Laboratory of Invertebrate Zoology, 4-5-7Konan, Minato-ku, Tokyo 108, Japan

Abstract: Fortyfour octopods from bottom trawls off Palmer Archipelago, south-easternArgentina, south-eastern New Zealand, Crozet Islands and Showa Station were examined. Three species of Pareledone, three species of Graneledone and one species of Megaleledone were identified. All were characterizedby having a single row of arm suckers. Mature males of P. harrissoni, P. adelieana and G. macrotyla were recorded for the first time. Hectocotylus and male reproductive organs of these species are described. On the basis of previously reported distributions and the present localities,P. charcoti, P. harrissoni and P. adelieana appear to have circumantarctic distributions. G. macrotyla wasidentifiedbutthe othertwospeciesof Graneledonecould notbe identifiedto species level because of the poor systematic state of this genus.

Received 5 July 1993, accepted 15 November 1993 Key words: Southern Ocean, eledonine octopods, Pareledone, Graneledone, Megaleledone, systematics, distribution

Introduction Australia and the South African Museum. Eledonine octopods in the Southern Ocean are poorly known In this paper, we provide supplemental descriptions and from both systematic as well as zoogeographic points of view. remarks on each species, especially the male reprodluctive Palacio (1978) summarized systematic characteristics of organs that have not been reported previously, and distribution eledonine octopods and reviewed their general geographic and and depth ranges are summarized. vertical distributions. Several species have also been reported from the stomach contents of seals (Lipinski & Woyciechowski Materials and methods 1981, Rodhouse et al. 1992) and they are considered an important component in the Antarcticfood web. Kuehl(l988) Material was collected indiscriminately from bottom trawl recently reported on the reproductive biology and geographical catches aboard RV Sinkai-Maru (JAMARC) off south-eastern distribution of Antarctic octopods collected off Elephant Island. New Zealand and Crozet Islands in 1983 and 1984, by RVs However, there is still little known of the eledonine octopods of Yoshino-Maru and Banshu-Maru (JAMARC) off Palmer the Southern Ocean. Archipelago and south-eastern Argentina in 1982 and 1985. Eledonine octopods in the region are classified into five Allspecimenswerefrozenonboardandbroughttothelaboraitories genera: Pareledone, Megaleledone, Graneledone, of Tokyo University of Fisheries and/or National Science Thaumeledone and Bentheledone. Currently 17 species have Museum, Tokyo. Specimens were put in 10% formalin fresh been reported but the majority are insufficiently described, and water after thawing and transferred into 40% isopropyl alcohol based only on juveniles and/or females. Several are considered later. Someofthem,ingoodcondition, werecolourphotographed to be synonyms of previously reported species (Berry 1918, before fixation. Collection data for the specimens examined are Robson 1932) and/or nomia dubia (Voss 1988). summarized in Table I. Bottom trawling by the Japan Marine Fishery Resources Species identification and measurement were based on Research Center (JAMARC) off Palmer Archipelago, south- preserved specimens. Counts, measurementsand indicesfiollow eastern Argentina, south-eastern New Zealand, and Crozet Roper &Voss (1983) andmodifiedindicesandcountsproposed Islands during 1982-1985 yielded 44 octopods. A systematic by Norman (1992) are also employed. Abbreviatiom and survey of these specimens revealed that there were three definitions are as follows, StM: stage of maturity (m(ature: Pareledone species:P. charcoti,P. harrissoni andP. adelieana, having developed spermatophore or eggs; premature: having one Graneledone species, G. macrotyla, and oneMegaleledone developingreproductive organ but no spermatophorenor eggs; species, M. senoi. Mature maleP. harrissoni, P. adelieana and immature: having little or no developing reproductive organ, G. macrotyla were collected for the first time. Two unidentified sex indeterminate); ML mantle length; TL total length; MWI: species of Graneledone were also recognized. Additional mantle width index (MWIMLx100); HWI: head width index preserved materialsof Pareledone and Graneledone,including (HW/MLx100); MAI: mantle arm index (ML/longest arm type specimens, were examined at the Museum of Victoria, lengthx100); ALI: arm length index (AWMLx3 00); 205

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Table I. Collection data and registration number of National Science Museum, Tokyo for Eledonine octopods examined.

~~~~~~~ ~~ ~ Species n ML Sex Sampling Date Locality Gear Depth Vessel Reg. No.

Pareledone charcoti 1 50.0 F 22 Jan., 1982 61"Sl'S 55"03'W BT 107m Yoshino-Maru NSMT Mo66775 Pareledone charcoti 1 50.0 M 16 Jan., 1985 61"lOS 55"55'W BT 120m Banshu-Maru NSMT Mo67833 Pareledone charcoti 1 70.0 F 16 Jan., 1985 61"lOS 55"55'W BT 120m Banshu-Maru NSMT Mo67831 Pareledone charcoti 2 35 .O-52.0 F2 16 Jan., 1985 61"lO'S 55"55'W BT 120m Banshu-Maru NSMT Mo67848 Pareledone charcoti 11 41 .O-52.0 M10,Fl 21 Jan., 1985 61"13'S 55"51'W BT 133m Banshu-Maru NSMT Mo67847 Pareledone charcoti 1 55.5 M 27 Jan., 1985 63"28'S 62"58'W BT 475m Banshu-Maru NSMT Mo67829 Pareledone charcoti 1 59.5 F 27 Jan., 1985 63'28's 62"58'W BT 475m Banshu -Maru NSMT Mo67830 Pareledone charcoti 6 48.0-65.0 F6 27 Jan., 1985 63"28'S 62'58'W BT 475m Banshu-Maru NSMT Mo67846 Pareledone charcoti 1 71.5 M 4 Feb., 1985 62"42'S 55"48'W BT 222m Banshu-Maru NSMT Mo67832 Pareledone harrissoni 1 80.0 F 22 Jan., 1982 61'15's 55'03'W BT 107m Y oshino-Maru NSMT Mo67834 Pareledone harrissoni 1 96.0 M 22 Jan., 1982 61"15'S 55"03'W BT 107m Yoshino-Maru NSMT Mo67844 Pareledone harrissoni 1 73.0 M 22 Jan., 1982 61"lO'S 55"55'W BT 120m Yoshino-Maru NSMT Mo67845 Pareledone harrissoni 1 96.0 M 30 Jan., 1985 61"lS'S 54'28'W BT 138m Banshu-Maru NSMT Mo66767 Pareledone adelieana 1 66.0 M 29 Jan., 1982 62"14'S 60°43'W BT 496m Yoshino-Maru NSMT Mo67841 Pareledone adelieana 1 33.5 M 27 Jan., 1985 63"28'S 62"58'W BT 475m Banshu-Maru NSMT Mo67842 Megaleledone senoi 1 115.0 M 30 Jan., 1982 62"59'S 62'09'W BT 803m Yoshino-Maru NSMT Mo66840 Megaleledone senoi 1 145.0 M 16 Jan., 1985 61"lO'S 55"55'W BT 120m Banshu-Maru NSMT Mo63960 Megaleledone senoi 1 110.0 M 16 Jan., 1985 61"lO'S 55"55'W BT 120m Banshu-Maru NSMT Mo63961 Megaleledone senoi 2 125.C-130.0 M 16 Jan., 1985 61"lO'S 55"55'W BT 120m Banshu-Maru NSMT Mo67849 Megaleledone senoi 3 105.0-120.0 F 27 Jan., 1985 63"28'S 62"58'W BT 475m Banshu-Maru NSMT Mo67850 Graneledone macrotyla 1 65.0 F 14 Oct., 1984 47"OO'S 59"37'W BT 972m Shinkai-Man! NSMT Mo67839 Graneledone macrotyla 1 67.0 M 16 Oct., 1984 45"44'S 59"46'W BT 858m Shinkai-Maru NSMT Mo67838 Graneledone sp. A 1 59.0 M 16 Oct., 1984 45"44'S 59"46'W BT 858m Shinkai-Maru NSMT Mo67837 Graneledone sp. B 1 49.0 M 14 Nov., 1983 48'40's 179Y1 'E BT 732m Shinkai-Maru NSMT Mo67835 Graneledone sp. B 1 70.0 F 14 Nov., 1983 48'48's 179' 16'E BT 725m Shinkai-Maru NSMT Mo67836

H: hectocotylized arm; AWI: arm width index (arm width at Diagnosis: Small to medium in size, mature about 40-50mm widest point on stoutest arm/MLxiOO); SDI: sucker diameter ML. Body muscular with short subequal arms, typically index (largest sucker diameterMLx100); WDI: web depth 1.5-1.8 times mantle length. Web shallow. Head narrower index (deepest web length/longest arm lengthx100); GLC gill than mantle with moderate eyes. Triangular small papilla on lamellaecount (number ofoutergilllamellae);ASC: armsucker dorsal surface over each eye. Dorsal surface of mantle, head count (highest number of suckers for intact arm); and arms sculptured in numerous well-marked tubercles. HALI: hectocotylized arm length index (HALMLxlOO); Ventral surface of body smooth with peripheral fold on ON:opposite arm length index (HAWLeft Arm I11 length mantle. Papillae on eyes, dorsal sculpture and peripheral fold ~100);HASC: hectocotylizedarmsuckercount; OASC: opposite variable. Funnel organ VV-shaped. Arm suckers uniserial, arm sucker count; LLI: ligula length index (LL/HALxiOO); typically about 35-45. Gills with 7-9 lamellae on outer CLI: calamuslengthindex(CULLx100);TOLI: terminal organ demibranch. Ligula small, about 10%of hectocotylized arm. limb index (length of terminal organ/MLx100); Ligula groove smooth with well-marked lateral margins. SpLI: spermatophore length index (length of Calamus short with wide, shallow groove. About 26-32 spermatophoreMLx100); SpWI: spermatophore width index suckers on hectocotylized arm. Ink sac completely buried in (greatest width of spermatophore/SpLxlOO); ELI egg length mid-gut gland. index (length of mature egg/MLxiOO); EWI: egg width index (greatest width of mature egg/MLx100); FLI: funnel length Remarks: In the material examined, two forms of P. charcoti index (funnel length/MLxlOO); FFI: free funnel index (length were recognized. One form was identical to P. charcoti but the of free funnel portionfFLxi00). other differed from typical P. charcoti, in having: 1) much The radula was removed from the buccal mass for scanning smaller tubercles of which the top was pigmented dark purplish electron microscopy (SEM). The upper part of the rolled inner in dour,2) the larger size at maturity of males and 3) slightly ribbon was cut and cleaned in an ultrasonic cleaner. After different shape of radula teeth (Fig. la-d). The external drying, the ribbon was stretched on a SEM stub with adhesive appearanceof thelatter formwasvery close to that ofP. aurorae tape. This procedure slightly damaged the ribbon and teeth. (Berry 1917, PL XIII) but this species was synonymized with P. charcoti by Berry himself (1918) without comment. Pareledone charcoti (Joubin, 1905) Fortunately, one of us (TK) had a chance to examine both the Material examined: See Table I for collection data and lectotype of P. charcoti and the holotype of P. aurorae at the Table 11 for measurements and indices (NSMT Mo67829, Museum of Victoria and obtained the following results. 67830,67831,67832,67833). LectotypeofP. charcoti (designatedbyRobson, 1932; female,

Table 11. Measurements (in mm) and indices of Pareledone spp.

Species charcoti charcoti charcoti charcoti* charcoti* charcoti* harrissoni harrissoni adelieana 4adelieana Reg.No. NSMT Mo. 67829 67830 67831 67832 67833 66775 66767 67834 67841 67842

Sex male female female male male female male female male male StM mature mature mature mature mature premature mature premature mature premature ML 55.5 59.5 70.0 71.6 50.0 50.0 96.0 80.0 66.0 33.5 TL 150.0 135.0 170.0 205.0 123.0 135.0 310.0 220.0 165.0 106.0 MWI 74.8 73.3 68.4 86.2 70.2 76.6 87.5 72.5 62.1 83.6 HWI 54.1 47.7 39.4 46.1 43.8 40.0 50.0 45.0 43.9 58.2 MAI 63.8 75.3 72.9 52.3 71.4 62.5 43.6 53.3 69.5 51.5 ALI L1 D 112.6 118.6 176.0 124.0 138.0 197.9 175.0 D 179.1 L2 D 124.4 120.0 177.3 D 72.0 182.3 D 136.4 185.1 L3 156.8 132.8 131.4 183.0 136.0 160.0 229.2 168.8 143.9 194.0 L4 156.8 114.3 137.1 176.0 140.0 140.0 224.0 175.0 143.9 194.0 R1 135.1 105.9 D 174.6 122.0 128.0 187.5 187.5 109.1 185.1 R2 138.7 121.0 125.7 D 128.0 148.0 197.9 181.3 140.9 t79.1 R3 136.9(H) 129.4 135.7 160.6(H) 130.0(H) 156.0 192.7(H) 175.0 119.7(H) '194.0(H) R4 151.4 121.0 135.7 191.3 138.0 136.0 208.3 187.5 136.4 l94.0 Arm formula 4=3>2>1 3>2=4>1 3=4>2>1 4=3=2=1 4>3>2>1 3>2>4=1 4=3>2=1 4=1>3=2 4=3=2>1 3=4>1=2 AWI 13.0 13.8 13.0 15.7 13.0 16.0 15.1 13.8 12.1 16.1 SDI 5.4 5.3 5.1 6.3 6.4 8.8 8.3 7.5 6.4 8.1 WDI 25.3 32.9 33.3 26.0 25.7 31.3 20.5 23.3 28.4 19.7 Web formula B,C,D,E,A C,B=D,A,E C,B,D=A,E D=C,B,A,E C=D,B,A,E B,C=D,E,A C,B,D,A,E B=C=D,A,E D,C,B,A,E B=CtD,A=E GLC 8-9 7 7 7 7 7 9-10 9 7 8 ASC 48 38 38 47 37 37 53 50 46 46 HAL1 136.9 - - 160.6 130.0 - 192.7 - 119.7 194.0 OAI 87.3 - - 87.8 95.6 - 84.1 - 83.2 100.0 HASC 31 - - 32 26 - 38 - 26 32 OASC 48 - - 46 37 - 53 - 46 46 LLI 27.6 - - 7.3 10.8 - 5.0 - 13.7 1.7 CLI 38.8 - - 34.5 42.9 - 34.8 - 55.6 - PLI 52.3 - - 37.7 41.0 - 49.0 - 14.3 - SpLI 142.3 - - 73.3 - - 112.3 - - - SpWI 4.5 - - 5.3 - - 4.9 - - - ELI - 15.9 - - - 24.4 - - - - EWI - 6.7 - - - - - 10.0 - - FLI 37.8 30.3 36.4 33.8 29.4 43.4 32.3 32.5 28.0 32.8 FFI 18.7 14.8 19.9 22.3 15.6 21.0 16.7 20.0 12.9 l3.4

*:"aurorae"type; D=damaged; H=hectocotylized arm.

34mm ML; premature; locality 65"05'S, 63"55'W, 3 Sept., Eyes slightly larger in proportion to mantle and head than in 1904;MNHN 5-7-1095): preserved in good condition. Mantle P. charcoti. Ventral surface of mantle, head and arms IV and funnel cut at mid-ventralline but no internal dissection has smooth with no obvious peripheral fold on mantle. Funnel been done. Skin absent on most of dorsal mantle. Dorsal surface organs VV-shaped with thin limbs and well separated of head, eyes and arms 1-111 havenumerouswell-markedround, components. flat-topped tubercles, measuring about 0.5-2 mm diameter. Observationbased on skin texture and funnel organs oEmore Ventral surface of mantle and arms III-IV smooth with weakly than 20 specimens of P. charcoti deposited in the Museum of definedperipheralfoldon mantle. Onedistinctpapillaondorsal Victoria showed that they varied in form between the type of surface over each eye. Funnel organ W-shaped with thick P. charcoti to the type of P. aurorae. This suggests that skin limbs and very little space between the two components. texture and the papillae over the eyes and funnel organs are Holotype of P. aurorae (Moschites aurorae Berry, 1917; affected by artificial factors such as variation in preservation. male, 24mmML; premature; locality 66"08'S, 94"17'E,27 Jan., Specimensfixed while alive have distincttubercles, aperipheral 1914; AM C.40891): preserved in good condition except for fold and thick limbs on the funnel organ, identical to the form hardening of the body. Beaks and radula removed from buccal seen in the type of P. charcoti. Specimens fixed when dead or mass. Dorsalsurfacesofmantle, head, eyes and arms I-IV have thawed after freezing tend to have low, indistinct tubercles and numerous small round tubercles of which the top is pigmented papillae over the eyes, characteristics found in the type of dark purplish in colour. Tubercles much smaller and elevation P. aurorae. Since no distinct specific difference was found, in lower than in the lectotype of P. charcoti. Two relatively large agreementwith Berry (1918),we conclude that the two species tubercles on central dorsal mantle. No papillae over the eyes. are synonymous.

between 107- 475 m. Pareledone harrissoni (Berry, 1917) Material examined: See Table I for collection data and Table I1 for measurements and indices (NSMT Mo66767, 67834). Diagnosis: Medium in size, mature about 100 mm ML. Body soft, epidermis loose. Arms subequal, typically 1.8-2.3 times mantle length. Web shallow. Head narrower than mantle with large eyes. Surface of body almost smoothwith thick peripheral foldonmantle. Asmallpapillae ondorsalsurfaceovereacheye. Funnel organ W-shaped with thick limbs. Arm suckers uniserial, typically about 44-53. Gills with 9-10 lamellae on outer demibranch. Ligula small, about 5% of hectocotylized arm. Ligula groove smooth with well-marked lateral margins. Calamus short with wide, shallow groove. Thirtyeight suckers on hectocotylized arm. Ink sac completely buried in mid-gut gland. Supplementaldescription:Male(96mmML, NSMTMo66767) right third arm hectocotylized, slightly shorter than opposite arm(OAI=84.1). Ligulasmall(LLI=5,0), alittleflattenedbut not expanded, with round tip. Ligula surface smooth with well developed lateral margins. Calamus well developed (CLI = 34.8) but bluntly pointed with wide, shallow groove that continues to spermatophoric groove on ventral margin of arm (Fig. 2a). Number of suckers on hectocotylized arm 38 and opposite arm 53. Male reproductive organs consist of well developed testis, spermatophoric gland, accessory gland, spermatophore storage sac and terminal organ connected with short weakly coiled vas deferens, very short proximal spermatophoric duct, ill defined medial spermatophoric duct and thick distal spermatophoric duct, respectively (Fig. 2c). Proximal portion of spermatophoric gland strongly coiled. Diverticulum single coiled. One spermatophore in the terminal organ diverticulum and the other in the spermatophore storage sac. Spermatophore long (SpLI = 112.3), slender (SpWI = 4.9), with relatively small sperm reservoir (SpRI = 22) (Fig. 2b). Female (80 mm ML, NSMT Mo67834) reproductive organs with small immature ovary; proximal oviduct short, oviducal gland; distal oviduct long, thick (Fig. 2e). Digestive tract- anterior salivary glands small; posterior salivary glands large, well developed; esophagus short, connecting with large crop; Fig. 1. Pareledone charcoti: a. dorsal view of typical “charcoti” stomach large, muscular, with well defined spiral caecum; (male, ML = 55.5 mm, NSMT Mo67829), b. dorsal view of intestine thin-walled (Fig. 2d). Ink sac completely buried in “aurorae” form (male, ML = 71.6 mm, NSMT Mo67832), mid-gut gland; anal flaps present. Gills with 9-10 lamellae on c. radula teeth of typical “charcoti))(male, ML = 55.5 rnm, outer demibranch. Radula with seven transverse rows of teeth NSMT Mo67829), d. radula teeth of “aurorae”form (female, and marginal plates (Fig. 3a). Rachidian tooth has 1-2 lateral ML = 50 mm, NSMT M066775). cusps on each side. First lateral teeth small and unicuspidate; secondlateral teethunicuspidate with widebase; marginal teeth Distribution: On the basis of published reports (Joubin 1905, long, pointed. Berry 1917,Robson 1932,Okutani 1986)andmaterialexamined here, this species appears to have acircumantarctic distribution. Remarks: Berry (1917) described this species based on only It has been reported from a depth range of 0 m (Robson 1932) three immaturefemales and onejuvenile. The specificcharacters to 383 m (Massy 1916). The present material was collected herecognized are identical in our specimens. Male reproductive

Fig. 2. Pareledone harrissoni:a. tip of hectocotylized right third arm (male, h4L = 96 mm, NSMT Mo66767), b. Spermatophorein spermatophore storage sac (male,ML = 96 mm, NSMT Mo66767), c. male reproductive organs (h4L = 96 mm, NSMT Mo66767), d. digestive tract (female, ML = 80 mm, NSMT Mo67834), e. female reproductive organs (ML = 80 mm, NSMT Mo67834). L = ligula; C = calarnus; TO = terminal organ; GP = genital pore; D = diverticulum; SSS = spermatophore storage sac; AG = accessory gland; SG = spermatophoric gland; MG = Mucilaginous gland; VD = vas deferens; T = testis; BM = buccal mass; ASG = anterior salivary glands; E = oesophagus; PSG = posterior salivary glands; CR = crop; S = stomach; C = caecum; I = intestine; MGG = mid-gut gland; 0 = ovary; OD = oviducal gland; PO = proximal oviduct; DO = distal oviduct.

organs, hectocotylized arm and digestive tract are described short arms with fewer arm suckers than P. turqueti. here for the first time. Voss (1988, table 2) considered P. harrissoni a nomen dubium without giving any reasons for Distribution: Previously recorded from three stations off his decision. One of us (TK) examined the holotype of Queen Mary Land, Antarctica, Indian Ocean Sector, at P. harrissoni at the Museum of Victoria and confirmed that the 500-712 m (Berry 1917). The presentlocalities are theopposite species, although in very poor condition, is valid. This species side of Antarctica. Depth range of the present collection was resembles P. turqueti but is distinguished by having relatively 107-138 m.

not expanded, with round tip. Ligula groove with 7-8 well marked transverse ridges; lateral margins low. Calamus well developed (CL1=55.6) but pointed triangular small with long narrow groove that continues to spermatophoric groove on ventralmarginof arm (Fig. 4a). Sucker number on hectocotylized arm26; onopposite arm46. Malereproductiveorganswithwell developed testis; spermatophoric gland, accessory gland, spermatophore storage sac and terminal organ connected with very short vas deferens, ill defined proximal spermatophoric duct, ill defined medial spermatophoric duct and thick distal spermatophoric duct, respectively (Fig. 4b). Proximal portion of spermatophoric gland strongly coiled. Diverticulum large, sac-like, not coiled. Digestive tract-anterior salivary glands small; posterior salivary glands extremely large, we11 developed. Esophagus relatively long, connecting with well defined crop. Stomach small but muscular; spiral caecum also small; intestine short, wide thin-walled; anal flaps present (Fig. 4c). Ink sac completely buried in mid-gut gland. Gills with 7 - 8 lamellae on outer demibranch. Radula with seven transverse rows of teeth and marginal plates (Fig. 3b). Rachidian tooth has no lateral cusps on each side. First lateral teeth small, triangular in shape; secondlateralteeth unicuspidatewithwidebase;marginal teeth relatively short, pointed. Remarks: Voss(1988, table2)consideredP. adelieana anomen dubium without justifying his decision. Berry (1917) described P. adelieana based on one female and two juvenile specimens. This species shares a number of morphological characterswith two other Pareledone species: P. umifakae Taki, 1961 and P. polymorpha (Robson, 1930), including a W-shaped funnel organ and a concave ligula with 7-8 transverse ridges. No Fig. 3. SEM of radula teeth: a. Pareledone harrissoni (female, justification was found to recognize this species as a nomen ML = 80 mm, NSMT Mo67834), b. P. adelieana (male, ML = dubium. P. adelieana possesses an almost straight or upward 66 mm, NSMT Mo67841), c. Craneledone rnacrotyla (male, curling rostra1 edge of the lower beaks Perry 1917, fig. 12). ML = 67 mm, NSMT M067838). This character was also found in our specimens and may aid identification of beaks from stomach contents of predators. The Pareledone adelieana (Berry, 1917) rostrum of the lower beak of P. umitakae (Taki 1961, text- Material examined:See Table Ifor collection data andTable I1 fig. 11) appears similar to that of P. adelieana. Detailed for measurements (NSMT Mo67841,67842). examination on the type specimens is required to resolve the status of these species. Diagnosis:Small to medium insize, male mature about 60 mm ML. Body soft but muscular with short subequal arms, typically Distribution: This specieswas previously known only from the 1.4-2.0 times mantle length. Web very shallow. Head narrower type locality, off the Mert Glacier Tongue, AdClie Land, than mantle with large eyes. Surface of body almost smooth Antarctica, 533-555 m (Berry 1917). The material described without peripheral fold on mantle. No distinct papillae over here was collected from the opposite side of Antarctica, from eyes. Funnel organ W-shaped. Arm suckersuniserial,typically 215496 m. Lu & Stranks (Unpublished data) collected this about 44-46. Gills with 7-8 lamellae on outer demibranch. speciesfromPrydz Bay, suggesting distribution inIndo-Atlantic Ligula size moderate, about 14% of hectocotylized arm. Ligula sector in the Antarctic Ocean. groove with well-marked 7-8 transverse ridges. Calamus short Megaleledone senoi (Taki, 1961) with narrow groove. 26-32 suckers on hectocotylized arm. Posterior salivary glands extremely large. Ink sac completely Material examined: See Table I for collection data and buried in mid-gut gland. Kubodera & Okutani (1986) for measurements of specimens (NSMT Mo63960,63961). Suppiementaldescription:Male (66 mML, NSMTMo67841) right third arm hectocotylized, shorter than opposite arm (OAI Diagnosis: Large species, male mature about 150 mm ML. = 83.2). Ligulasizemoderate (LLI= 13.7), slightlyflattenedbut Body soft but muscular with relatively short subequal arms,

L C

Fig. 4. Pareledone adelieanu (male, ML = 66 mm, NSMT Mo67841): a. tip of hectocotylized right third arm, b. male reproductive organs, c. digestive tract without mid-gut gland. [abbreviations as in Fig. 21

typically 2.0-2.3 times mantle length. Web deep, about half of 38-39 suckers on hectocotylized arm. Ink sac partially buried armlength. Headnarrowerthanmantlewithmoderate eyes. No in mid-gut gland. distinct papillae over eyes. Surface of body smooth without peripheral fold on mantle. Epidermis thick and flabby, loose Remarks: Taki (1961) proposed a new subfamily and easily dislocated. Funnel organ W-shaped with thicklimbs. Megaleledoninae, new genus Megaleledone and new species Arm suckers uniserial, typically about 55-65. Gills with 11-12 M. senoibasedontwofemalespecimenscollectedat630-680 m. lamellae on outer demibranch. Ligula very small, about 4% of A supplemental description of male reproductive organs has hectocotylized arm. Ligula groove smooth with round tip. been reported elsewhere (Kubodera & Okutani, 1986). Voss Calamus rounded, short with wide spermatophore groove. (1988) failed to include this species. Although the propriety of

Table 111. Measurements (in mm) and indices of Graneledone macrotyla. outer demibranch. Ligula small, about 8% of hectocotylized arm. Ligula groove with shallow 4-5 transverse weak ridges. Reg.No. NSMT Mo. 67838 67839 Calamus short with shallow groove. Thirtyone suckers on Sex male female hectocotylized arm. No ink sac. StM mature immature ML 67.0 65.0 Supplemental description: Male (67mm ML, NSMTMo67838) TL 210.0 230.0 right third arm hectocotylized, shorter than opposite arm. MWI 87.3 115.4 Ligula small (LLI = 8.2), not expandedwith bluntly pointed tip. HWI 68.4 84.6 Ligula groove with 4-5 shallow transverse ridges; lateral 38.2 MAI 49.6 marginslow. Calamuswell developed (CLI = 49.1), raised well ALI L1 194.0 215.0 L2 186.6 246.2 above ligula surface. Posterior groove short, shallow, wide, L3 D 238.5 continuing to spermatophoric groove on ventral margin of arm L4 171.6 233.8 (Fig. 5a). Sucker number on hectocotylized arm 31; opposite R1 201.5 261.5 arm lacks tip but other arms have 60-65 suckers. Male R2 198.5 238.5 R3 149.3(H) 238.5 reproductive organs with well developed testis, long R4 179.1 223.1 spermatophoric gland, long thick accessory gland, thin Arm formula 2=1>3>4 1>2=3>4 spermatophore storage sac and terminal organ, each connected AWI 17.9 24.6 with long, almost straight, vas deferens, ill defined proximal SDI 7.2 9.1 spermatophoric duct, ill defined medial spermatophoric duct WDI 33.3 26.9 Web formula A=B=C,D,E D=C=B,A,E andshortthick distal spermatophoricduct,respectively (Fig. 5b). ASC 65 70 Proximal portion of spermatophoricglandwith two strong coils. GC 7 7 Diverticulum large, sac-like, weakly coiled. Spermatophores HAL1 149.3 - absent. Immature female (65 mm ML, NSMT Mo67839) with - OAI D small undeveloped ovary and oviduct. Radula, as shown in HASC 31 - OASC D - Fig. 3c,and all theothercharactersagreewellwithVoss(1976). - LLl 8.2 Remarks: Judging from the condition of the reproductive CLI 49.1 - PLI 29.9 - organs, this species may attain a larger size at maturity than the FLI 39.6 49.2 present specimens. FFI 20.6 23.8 Distribution: This specieswas previously only known from the D=damaged; H=hectowfylized. typelocality, 54"43'S, 55'30'W. The present collection extends the known distribution to the north of 45"s along south-eastem Argentina. The known depth range is 858-2044 m. a sub-familial rank is questionable, we consider that M. senoi has sufficient distinct diagnostic characters to be a valid genus Discussion and species. Although little isknown of theoctopodsof theSouthem Ocean, Distribution: This species is known to occur off Showa station it isstillapparent thatPareledonespeciesarethemostdiversified and thePalmer Archipelago from 120-803 mdeep (Taki 1961, and dominant octopods in the Antarctic (Roper et al. 1985, Kubodera & Okutani 1986). Lu & Stranks (unpublished data) Kuehl 1988). Among the nine Pareledone species hitherto collected this speciesfromPrydzBay, suggesting distribution in reported from the Southern Ocean, we recognize three valid Indo-Atlantic sector in the Antarctic Ocean. species: P. churcoti, P. adelieanu and P. hurrissoni, in our Graneledone macrotyla (Voss, 1976) collection. A detailed systematic study ofPareledonecollected from Prydz Bay revealed three valid species: P. churcoti, Material examined: SeeTableI for collection data and Table I11 P. adelieana and P. turqueti and two new Pareledone species for measurements and indices (NSMT Mo67838,67839). (Lu & Stranks, in press). Combined with our findings, we have Diagnosis: Medium size species, male mature about 70 mm a total of six distinct Pareledone species, all of which are ML. Body muscularwith relatively short subequalarms,typically considered to be Antarctic endemics. 1.8-2.5 times mantle length. Web relatively deep, about 30% With regard to Graneledone species in the Southern Ocean, of arm lengths. Head almost same as mantle width with large Hoyle (1886) reported thepresenceofEeledoneverrucosafrom eyes. Dorsal surface of mantle, head and base of arms with the Kermadec Islands (29"45'S, 178"ll'W)because of the close numerous large distinct cartilaginous tubercles. Other surface resemblancebetween his specimen and E. verrucosa described of body smooth. Gigantic cartilaginous papillae over eyes. by Verrill (1881) from Halifax, Cape Sable and New York Funnel organ W-shaped with thick limbs. Arm suckers (north-west Atlantic). Berry (1916) erected a new name, uniserial, typically about 60-70. Gills with seven lamellae on challengeri,for Hoyle's Kermadec specimen on the basis of the

Fig. 5. Graneledone rnacrotyla (male, ML = 67 mm, NSMT Mo67838): a. tip of hectocotylized right third arm, b. male reproductive organs without testis, c. digestive tract. [abbreviations as in Fig. 21

different localities of capture. But neither Hoyle nor Berry dubium due to insufficient specific characters given by Robson provided detailed description, nor figures in their publications. (1932) from the badly decomposed type specimen. Voss (1976) Joubin (1918) proposed a new genus Graneledone based on pointed out that Robson's (1932) description of G. challengeri E. verrucosa Verrill and included challengeri and his new is confused by mixing two species from different localities. subspecies media in this genus. Robson (1932) summarized According to Voss (1976, 1988), only two valid species of previous reports and gave a short description and remarks on Graneledone,namely, G. antarctica and G. macrotyla, and one G. challengeri. He also described a new Graneledone species, poorly defined species, G. challengeri, occur in the Southern G. setebos based on a (? female) specimen from a rockpool in Ocean. McMurdo Sound, that was reported as Moschites sp. by Massy In the present collection, two unidentified Graneledone (1916). In 1976, Voss examined deep sea octopods from off species, G. sp. A (NSMT Mo67837) and G. sp. B (NSMT Antarctica and southern Argentina and reported two new Mo67835,67836), were recognized,inaddition to G. macrotyla. Graneledone species, G. antarctica and G. macrotyla. In the G. sp. Aresembles G. antarctica except for slight differencesin discussion, Voss (1976) thought that G. setebos was a species tubercle distribution on the arms and in the structure of the

radula teeth. However, the localities of the present material and HowW.E. 1886. Report on the CephalopodacollectedbyH.M.S. “Challenger” of G. antarctica are so far distant from each other (present during the years 1873-76. Voyage of the “Challenger”, 16,l-246. JOUSIN,M.L. 1905. Description de deux ElCdones provenant de I’expCdition du material; off Crozet Islands, 45’44’S, 59’46’W at 858 m, Dr Charcot dans 1’Antarctique. Mkmoires de la Sociktr! Zoologique de G. antarctica type locality; Ross Sea, 74’05.6‘S, 175’05.2’W at France, 18,22-31. 3241 m) that we hesitate to conclude that the material is JOUSIN,M.L. 1918. Ckphalopodesrecueillis au cours descroisi6res de S.AS.le conspecific. G. sp. B is easily separable from G. antarctica and Prince de Monaco. 5 Note: Moschites verrucosa (Vemll). Bulletin de G. macrotyla and was collected off Southern New Zealand l‘lnstitut Ockanographique, No. 339,l-10. KUBODERA,T.&OKUTANI,T. 1986. New andrarecephalopodsfromthehtarctic (43’9’S, 177’49’E at 780 m, 48”48‘S, 179’6’E at 725 m, and waters. Memoirs ofNationa1 Institute of Polar Research Specialhue, 44, 48”40’S,179’1’E at 732 m). These localities areclose to the type 129-141. locality of G. challengeri (29’45’S, 178’11’W, in 1165 m) and KUFHL,S. 1988. A contribution to the reproductive biology and geographical it is possible that these specimens are conspecific. However, the distribution of Antarctic Octopodidae (Cephalopoda). Malacologia, 29, large triangular papilla over the eye, the number of gill lamellae 89-100. and the structure of the radula teeth of our specimens disagree LIPINSKI,M. & WOYCIECHOWSKI,M. 1981. Cephalopods in the food of Weddell seals from the Admiralty Bay (King George Island, South Shetland Islands). with Robson’s (1932) description of G. challengeri. O’Shea Polish Polar Research, 2,163-167. (1990) included a report on a Graneledone species from Lu, C.C. & STRAW, T.N. In press. Synopsis of Pareledone species, with southern New Zealand and considered it to be an undescribed description of two new species from Antarctica (Cephalopoda: Octopodidae). species. G. sp. B may be identical with O’Shea’s species but a Memoirs of the Museum of Victoria. MASSY,A.L. 1916. Mollusca. Part 11. British Antarctic “Terra Nova” detailed comparison with the holotype of G. challengeri is Expedition, 2,141-175. necessary before final conclusions can be drawn. NORMAN,M.D. 1992. Four new octopusspeciesofthe Octopus macropus group A systematic survey of the Graneledone specimens deposited (Cephalopoda: Octopodidae) fromtheGreatBarrierReef,Australia. Memoirs in the Museum of Victoria, Australia, and the South African of the Museum of Victoria, 53,267-308. Museum indicatesthat severalundescribedspecies aredistributed OKUTANI,T. 1986. Anoteon Antarctic benthic mollusks collected with a beam- trawl from Breid Bay by the 25th Japanese Antarctic Research Expedition. allopatrically and/or sympatrically in subantarctic waters from Memoirs OfNationalInstitute ofpolar Research Specialhue,40,277-287. Tasmania, Kerguelen Islands and South Africa. A detailed O’S~A,S.J. 1990. The systematics of the New Zealand Octopodidae systematic study of these specimens is in preparation. (Cephalopoda: Octopoda). MSc thesis, University Auckland,New Zealand, 144 pp. [Unpublished.] PALACIO,F.J. 1978. Vosseledone charrua: a new Patagonian cephalopod Acknowledgements (Octopodidae)with notes on related genera. Bulletin ofMarine Science, 28, 282-296. Heartfelt thanks to the researchers of the Japan Marine Fishery ROBSON,G.C. 1930. Cephalopoda: Octopoda: Discovery Reports 2,371-402. Resources Research Center for their great effort insupplyingthe ROBSON,G.C. 1932. A monograph of the recent Cephalopoda based on the present material. We are indebted to Dr C.C. Lu and Mr T. collections in theBritish Museum (NaturalHistory). Part2. TheOctopoda. Stranks, Museum of Victoria,Australia, andDr M.C. Roeleveld, London: British Museum (Natural History). 359 pp. South African Museum, for their kind hospitality and RODHOUSE,P.G., ARNBOM, T.R., FFDAK, Ymm,J. &MURRAY,A.W.A. 1992. Cephalopodpreyofthesouthemelephant seal,Mirounga leomina L. Canadian considerationduring the examination of specimens deposited in Journal ofZoology, 70,1007-1015. these museums. They also provided useful discussion and ROPER,C.F.E. & Voss, G.L. 1983. Guidelines for taxonomic descriptions of comments on the present paper. We thank Dr F.G. Hochberg, cephalopod species. Memoirs of the National Museum of Victoria, 44, Santa Barbara Museum of Natural History, for his continuous 49-63. support and help with our studies of octopod systematics. Drs RoPER,C.F.E.,SWEEVEY, M.J. &CLARKE,M.R. 1985. Cephalopoda. InFIsaim, W. & Hmu J.C. eds. FA0 species identification sheets for fuhery M. Norman, Museum of Victoria, Australia and K.N. Nesis, purposes. Southern Ocean (Fishing areas 48, 58 and 88) (CCAMLR Institute of Oceanology, Academy of Sciences, Russia kindly Convention Area). Vol. 1. Rome: FAO, 117-205. commentedonthemanuscript. Dr K Tsuchiya, Tokyo University TAKI, I. 1961. On two new eledonid octopods from the Antarctic Sea. Journal of Fisheries, helped with measurements and preparation of of the Faculty of Fisheries and Animal Husbandry, Hiroshima University, tables. 3,297-316. VERRILL.,AE.1881. Thecephalopodsofthenorth eastemcoastofAmerica.Part 11. The smaller cephalopods including the squids and the octopi with other References allied forms. Transactions of the Connecticut Academy of Science, 5, 259-446. BERRYS.S. 1916. Cephalopoda of the Kermadec Islands. Proceedings of the Voss, G.L. 1976. Two new species of octopods of the genus Graneledone Academy of Natural Science of Philadelphia, 68,4546. (Mollusca: Cephalopoda) from the Southern Ocean. Proceedings of the BERRY,S.S. 1917. Cephalopoda.AustralasianAntarcticExpedition, 1911-1914. Biological Society of Washington, 88,447-458. Scientific Reports. Series C. Zoology and Botany, 4,5-38. Voss ,G.L. 1988. The biogeography of the deepseaoctopoda. Malacologia, BERRY,S.S. 1918. Postscript to the Cephalopoda of the Australasian Antarctic 29,295-307. Expedition. A sheet unpaged, inserted in Scientific Reports. Series C. Zoology and Botany, 4.