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31 March 1990 Memoirs of the Museum of Victoria 50(2): 457-465 (1990) ISSN 0814-1827 https://doi.org/10.24199/j.mmv.1990.50.17 THREE NEW OF (: CEPHALOPODA) FROM SOUTH-EASTERN AUSTRALIA

By T. N. Stranks

Department of Invertebrate Zoologv. Museum of Victoria. 285-321 Russell Street. Melbourne. Victoria 3000 and. Department of Zoology. University of Melbourne, Parkville. Victoria 3052'

Abstract

Stranks. T.N.. 1990. Three new species of Octopus (Mollusca: Cephalopoda) from south- eastern Australia. Memoirs of the Museum of Victoria 50(2): 457-465. Three new species of Octopus are described and illustrated from south-eastern Australian waters: Octopus warringa. Octopus kaurna and Octopus bunurong. Octopus warringa also occurs in New Zealand.

Introduction Robsonella australis. —Benham. 1942: 227. text fig. 3, pis 18, 19 (partim).—Dell, 1952: 32, pi. 4. figs 2-6, There are about 30 species of octopus de- pi. 5. figs 1, 3, 4. pis 7, 8 (partim).—1959: 95 (non scribed from Australian waters (Lu and Phillips, Octopus australis Hoyle, 1885). 1985). As part of an ongoing study of the fauna, a Octopus Species A. —Stranks. 1988: 54. text figs 21- systematic revision of the inshore benthic octo- 25. podids of south-eastern Australia was under- Tasmania. Maria taken (Stranks, 1988), and three undescribed Material examined. Holotvpe: Island, west of Darlington (42°35'S. 148°03'E), 30 m. species of Octopus were identified. This paper R. Wilson, 23 Apr 1985, NMV F57444 (mature male. describes the new species. More details will be 16.5 mm ML). of a larger included with the future publication Paratvpes: Tasmania. Off east coast of Tasmania monographic revision of the of (42°40'S. 148°28'E). 122 m, RV "Discovery", Station south-eastern Australia. 113, BANZARE, 23 Mar 1931. SAM D 152 19 (sub- The counts, measurements and indices listed mature female, 22.0 mm ML); Maria Island (42°44'S. are as defined by Roper and Voss (1983). Other 148°01'E). D. Clayton, 1985, NMV F53219 (sub- mature males. abbreviations used are ML — mantle length and mature female, 25.9 mm ML; 2 16.0 mm ML and 18.0 mm ML). TL — total length. Material is lodged in the Victoria, eastern Bass Strait (38°10'S, 147°49'E), 48 Australian Museum. Sydney (AM); Museum of m, Station 262, East Gippsland Scallop Survey, 28 Feb Victoria, Melbourne (NMV); Otago Museum, 1971, NMV F31259 (mature male, 14.4 mm ML) Dunedin (OM); and South Australian Museum, New Zealand, Portobello (45°51'S. 170°39'E), C. Adelaide (SAM). Hedlev. 10 Dec 1918, AM CI 59292 (mature male. 20.5 mm ML). Octopodidae Other material: Tasmania. Off east coast of Tas- (42°40'S, 148°28'E), from fish stomach con- Octopus Lamarck, 1 798 mania tents, RV "Discovery", stn 1 13, BANZARE, 23 Mar Type species. Octopus vulgaris Lamarck, 1 798. 1931, SAM D15220 (male, 18.9 mm ML); off Maria

Island (42°38'S, 1 48°05'E), 1 1 9 m, RV "Aurora", Aus- Diagnosis. Benthic octopodids. Mantle saccular, tralasian Antarctic Expedition, 12 Dec 1912, AM cirri, arms with without fins. Eight arms lacking C40887 (mature female, 29.6 mm ML). biserial suckers, third right arm of males hecto- Victoria, eastern Bass Strait (37°55'S, 148°21'E), 50 cotylised with end of arm modified into ligula m, stn 412, East Gippsland Scallop Survey, 15 Feb and calamus. Web well developed. Ink sac pre- 1971, NMV F53214 (immature male, 12.4 mm sent. Mantle aperture wide. Internal shell carti- ML). Zealand. Foveaux Strait (46°32'S, 168WE). laginous and vestigial. New OM A.T5.34 (mature female, 26.5 mm ML, with Octopus warringa sp. nov. eggs); same locality, OM A. '29. 1 1 1 (mature male, 34. mm ML; 2 submature females, 12.9 mm ML and 22.5 Figures la-f mm ML); Portobello (45°51'S, 170°39'E), OM

males, 1 9. 1 and Polypus duplex.—Berry, 1917: II, text fig. 5 (non A.'28.24 (2 mature mm ML 20.6 mm submature females, 16.2 20.6 Octopus duplex Hoyle, 1885). ML; 2 mm ML & mm

457 458 T. N. STRANKS

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Figure 1. Octopus warringa sp. nov.: a, dorsal view of holotype, NMV F57444, male, 16.5 mm ML; b, hecto- cotylised arm, and c, dorsal, and d, lateral, detail of hectocotylus, of paratype, NMV F53219, 16.0 mm ML; e, tubercles, branched and unbranched papillae head on and mantle dorsum of NMV F532 1 4, male, 1 2.4 mm ML; f, lateral view of arborescent ocular papillae of paratype, NMV F53219, male, 16.0 mm ML. fHREE NEW SPECIES OF OCTOPUS 459

ML); between Nelson and Stephen Islands, Tasman Distribution. South-eastern Australia, from the Bay(4P42'S, 174WE), 18-55 m,Ci. Thomson, 1900, Greal Australian Bight to eastern Victoria,

OM A. "0.97 (submature female, 24.8 .). mm Ml including Bass Strait and Tasmania. Also in the Description. Small (ML to 35 mrtr.TLto temperate waters of New Zealand, including the 125 mm); mantle broadly ovoid (MW1 56.3- North and South Islands, and Stewart Island. An #0.5-107.0); head wide, but narrower than inshore species, living on rocky bottom, and mantle (HWI 40.9-66.0-82.2), demarked from among sponges and polyzoans, at depths from mantle by moderate constriction; eyes large, 0-144 m (Stranks, 1988). projecting above surface of head. Funnel large, Etymology. The specific epithet warringa is de- stout, bluntly tapered (FuLI 32.6--//. 7-56.7); rived from an Australian Aboriginal word mean- funnel organ W-shaped, limbs thick, outer limbs ing "the sea", and is to be treated as indeclin- three-quarters as long as median limbs. Arms able. long (MAI 21.4-36. 4-52.8) (1.9-3.7 times ML in mature animals), stout, tapering to narrow Remarks. There has been discussion about the tips. Arm lengths subequal, arm order usually generic placement of this species. Octopus III. IV. 11. 1. Arm suckers biserial, raised from arm Warringa has been previously incorrectly iden- surface, moderately sized (ASIn 6. 1- 10. .^-14.3), tified and described under the name Rohsonclla 6th to 10th suckers usually largest, enlarged on australis (Hoyle, 1885). all arms of mature males and females. Third Pickford ( 1 955) reviewed the generic charact- right arm of males hectocotylised, shorter than ers of Rohsonclla, and concluded that the itsoppositenumber(OAI 67.2-77.6-85. 8; HcAI was not valid. Pickford (1955) considered that 149.2-205.5-273.3); ligula 6-10% of third right species which had been assigned to Rohsonclla arm length in mature animals (LLI 6.3-7.9- should be returned to the genus Octopus, a view I 10.2); ligula groove long, well marked and deep, agree with. without transverse ridges; calamus short, Tail (1982) revised Octopus australis Hoyle, acutely pointed (CaLI 24.0- .i5. 7-50.0); hecto- 1885 from south-eastern Australia, and dis- cotylised arm with 51-65 suckers. Web shallow cussed the status of similar species described (WDI 18.8-24.7-33.9), web formula usually from New Zealand, including material de-

CB-DA=E, Radula with B, 4 serration of the scribed as Rohsonclla australis. Octopus rhachidian. Ink sac present. Gill lamellae 6-8. warringa may be readily distinguished from O. Mature female with small eggs (2-3 mm long, australis by the characteristic skin patterning 1.0-1.5 mm wide), very long stalks, forming fes- (particularly the absence of a ventro-latcral all toons. Male with very long penis (PLI 15.2- integumentary ridge). I he enlarged suckers on 29.3-51.9), with a coiled diverticulum marked arms of mature males and females, the bulbous- by three lobes; spermatophores very long (SpLI shaped ligula, and the small eggs arranged in fes- 7 j. 7_/4ri.0-184.4), slender (SpWI 2.5-3, /- toons. 3.6), with large, coiled sperm reservoir (SpRI Previous accounts, under various names, now 29. 2-.?2. 2-36.4). referable in part or entirely to O. warringa Integumental sculpture consists of a pattern of include: I'olvpus duplex (Hoyle, 1885) (as used fine, rounded and closely set epidermal tu- by Berry, 1917); and Rohsonclla australis bercles; tubercles cover both dorsal and ventral (Hoyle, 1885) (as used by Benham, 1942; Dell, surfaces; branched and unbranched papillae pre- 1952; 1959; Brough, 1965). Two detailed de- sent on dorsum; pattern of papillae on mantle scriptions exist for Octopus warringa (Benham, dorsum includes approximately seven sub- 1942; Dell, 1952). parallel rows of simple, usually unbranched Berry (1917) described a specimen collected papillae along the mantle length; each row has by the "Aurora" off Maria Island, Tasmania, 3-4 papillae, a single larger papilla forms a pos- during the Australasian Antarctic Expedition

tentat i vely idenl i lied terior point on the mantle; a larger arborescent (1911-1914). Berry (1917) specimen as Polypus duplex, and included papilla is obvious in the supraocular region, sur- the rounded by 3-4 smaller, usually unbranched measurements and a figure. The specimen was a C40887), not an immature papillae; lateral integumentary ridge or fold mature female (AM around mantle circumference absent. No infor- female as Berry (1917) reported. Based upon the examination of 30 new speci- mation is available on colouring of live animals. 1 gave a Preserved specimens in ethyl alcohol uniformly mens from New Zealand, Benham ( 942) comprehensive description of Rohsonclla aus- light brown to purple dorsally, cream to light measurements and figures. brown ventrally. Ocelli absent. tralis, with detailed 460 T. N. STRANKS

Of this series in the Otago Museum, nine speci- specimen in the Portobello Aquarium laid ap- mens have been re-examined. proximately 1000 eggs during January 1963, Dell (1952) also described new material of R. which she brooded for about 80 days, until australis from New Zealand, and in 1959 listed hatching occurred (Brough, 1965). O. warringa two specimens collected by the "Discovery" off has mature eggs with a small egg length index (7- Tasmania during the British, Australian and 13% of mantle length), and the newly hatched New Zealand Antarctic Research Expedition juveniles are small, with a total length of ap- (BANZARE) (1929-1931). One specimen, a proximately 4 mm (Brough, 1965; and this submature female (SAM D 152 19), has been re- study). Based upon the relative sizes of eggs and examined; a second specimen, a male (SAM juveniles, it may be assumed that the juveniles of D15220) collected from fish stomach contents, O. warringa adopt a planktonic existence ini- has been examined but was unsuitable for tially. The duration of the planktonic phase is measurement. unknown. The systematic status of several lots of ma- terial described from New Zealand waters can- Octopus kaurna sp. nov. not be identified with certainty. These include Figures 2a-f Octopus campbelli Smith, 1 902 and Polypus aus- 1966: 241, text- tralis Massy, 1916. These accounts differ from Octopus jlindersi.— Macpherson, fig. pi. figs 1-3 (partim) (non Octopus flindersi descriptions of both Octopus australis from 1, 2, Cotton, 1932). south-eastern Australia, and O. warringa from Octopus Species B. —Stranks. 1988: 6 1 . text figs 26- south-eastern Australia and New Zealand. O. 30. campbelli Smith. 1902, from Campbell Island, New Zealand, which was subsequently redes- Material examined. Holotvpe: Victoria. Hobsons Bay (37°52'S. 144°56'E), NMVF24494 (mature male, 34.0 cribed by Robson (1929), possesses enlarged mm ML). suckers, an unusual ligula with a long calamus, Paratvpes: Victoria. Port Phillip Bav, Rosebud and 10 gill lamellae. Polypus australis, described (38°22'S, 144°54'E). shallows, J.H. Black, 5 Feb 1969, by Massy (1916) from Spirits Bay, New Zealand, NMV F53228 (mature female. 42.3 mm ML): Port and subsequently redescribed by Robson ( 1 929), Phillip Bav, Carrum (38°05'S, 145°07'E), beached. possesses a stout ligula, 6-7 gill lamellae, and a Port Phillip Authority, Feb 1981, NMV F52317 characteristic colour pattern of spots and bars on (mature female, 51.0 mm ML). the arms. Definite conclusions regarding the South Australia. Brighton (35°01'S, 138°31'E), Sept 1937, SAM 13283 (immature male, 31.3 species' identities and affinities await examin- D mm ML); Glenelg (34°58'S. 138°32'E). A. Robb, 29 Mar 1949. ation of the respective materials. SAM D16195 (mature female, 60.9 mm ML); Great Specimens of Octopus warringa from south- Australian Bight (32°24'S, 133°30'E), 49 m, FRV eastern Australia and New Zealand are almost "Explorer", P. Symonds, 23 Aug 1973, NMV F53226 exactly the same; no geographical variation was (mature male, 38.6 mm ML). detected. O. warringa is a distinctive species Other material: Victoria. Port Phillip Bay. Mordial- endemic to temperate waters of south-eastern loc(38°01'S. 145°05'E), W.Kershaw, Nov 1888,NMV Australia and New Zealand. It can be distin- F24488 (immature female, 22.7 mm ML); Port Phillip guished easily from other species of Octopus on Bay, Mentone (38WS, 145WE), L. Kershaw, 28 May 1928. the basis of a combination of characters: a NMV F24505 (mature male. 57.2 mm ML). broadly ovoid mantle; skin with a characteristic pattern of rounded tubercles and both branched Description. Medium-sized animals (ML to 85 and unbranched papillae on the dorsum, and mm; TL to 420 mm); mantle elongate ovoid enlarged papillae over each eye; large and promi- (MWI 40.0-69.9-106.8); head wide, slightly nent eyes; long, subequal arms (1.9-3.7 times narrower than mantle (HWI 36.7-5S. 7-88.8), ML in mature animals); moderately sized suck- demarked from mantle by minor constriction; ers, enlarged on all arms of mature males and eyes small, not projecting far above surface of females; a medium sized ligula (6-10% of third head. Funnel large, slender, bluntly tapered right arm length in mature animals); small eggs (FuLI 43.9-67.4-88.8); funnel organ VV- (2-3 mm long), arranged in festoons; and 6-8 gill shaped, limbs thick, outer limbs three-quarters lamellae. as long as median limbs. Arms very long (MAI Brough ( 1 965) described the morphology and 14.5-2J. 7-33.3) (3.1-6.2 times ML in mature brooding of eggs, and the hatching and behav- animals), slender, tapering to narrow tips. Arm iour of juveniles of Robsonella australis, here lengths unequal, arm order IJI.III.IV. Arm re-identified as Octopus warringa. A female suckers biserial, deeply set in arms, small sized THREE NEW SPECIES OF OCTOPUS 461

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Figure 2. Octopus kaurna sp. nov.: a, dorsal view, and b, hectocotyliscd arm, of holotype, NMV F24494, 34.0 mm e, tubercles mantle ML; c, dorsal, and d, lateral, detail of hectocotylus of NMV F24505, 57.2 mm ML; rounded on dorsum of paratype, NMV F52317, female, 51.0 mm ML; f, lateral view of ocular region of NMV F24488, female, 22.7 mm ML.

(ASIn 3.0-5. 3-8.7), all suckers similarly sized, length in mature animals (LLI 4.7-6.0-8.0); without sucker enlargement. Third right arm of ligula groove long, well marked and moderately males hectocotylised, shorter than its opposite deep, with incomplete transverse ridges; cala- number (OAI 53.5-67.2-69.3; HcAI 143.5- mus short, pointed (CaLI 32.1-41. 7-48.0); hec- 248.9-325.5); ligula 4-8% of third right arm tocotylised arm with 66-129 suckers. Web very 462 T. N. STRANKS

animals); shallow (WDI 10.5-/4. /-18. 7), web formula arms (3.1-6.2 times ML in mature usually scriation small suckers, without enlargement; a small ABCDE. Radula with B 4 _ s of the rhachidian. Ink sac present. Gill lamellae 9- sized ligula (4-8% of third right arm length in long), I animals); large eggs (9-11 mm 1. Mature female with large eggs (9-1 1 mm mature long; 2-3 mm wide); method of egg attachment with unknown method of attachment to sub-

to substrate unknown. Male with long penis (PL1 strate; and 9-1 1 gill lamellae. L7.7-2J.9-35.5), with a large, single coiled di- The biology of the species is unknown. verticulum; spermatophores relatively short (SpLI 51.7-

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Figure 3, Octopus bunurong sp. nov.: a, dorsal view ofholotype, NMV F'53223, male, 55.0 mm ML; h, hecto- cotyliscd arm of NMV F5 101 , 25.0 mm ML; c, dorsal, and d, lateral, detail of heetoeotylus of NMV F53220, 39.3 mm ML; e, rounded and elongate tubercles on mantle dorsum of NMV F53222, male, 37.5 mm ML; f, lateral view of ocular region of NMV F53220, male, 39.3 mm ML.

largest, without sucker enlargement. Third right with incomplete transverse ridges; calamus very arm of males hectocotylised, shorter than its short, acutely pointed (CaLI 12.8-/7.5-22.1); opposite number (OAI 45.9-6/.6-81.8; HcAI hectocotylised arm with 70-96 suckers. Web 166.9-20^.6-278.9); ligula 9-12% of third right very shallow ( WDI 9.4-/2 4- 14.8), web formula arm length in mature animals (LLI 9.0-9.5- usually ABCDE. Radula with A.3.4 sedation 11.8); ligula groove long, well marked and deep, of the rhachidian. Ink sac present. Gill lamellae 464 T. N. STRANKS

9-10. Mature female with large eggs (8-10 mm acters: an elongate ovoid mantle; skin with a long; 2-3 mm wide); method of egg attachment characteristic pattern of rounded and elongate to substrate unknown. Male with long penis (PLI tubercles on the dorsum, without large papillae 4.5-20.7-39.2), with a single coiled diverticu- over the eyes; large and prominent eyes; very lum; spermatophores relatively short (SpLI long, unequal arms (4. 1-7.3 times ML in mature 41.2-65.7-102.9), slender (SpWI 3.1-4.4-5.1), animals): moderately large suckers, without with large, coiled sperm reservoir (SpRI 40.6- enlargement; a medium sized ligula (9-12% of 49.7-55.9). third right arm length in mature animals); large Integumental sculpture consists of a pattern of eggs (8-10 mm long), with unknown method of fine, rounded and closely set epidermal tu- attachment to substrate; and 9-10 gill la- bercles; some irregularly spaced tubercles are mellae. larger and more elongate than the former type; The biology of the species is unknown. tubercles reach the largest size on the dorsum, Acknowledgements and those on the ventral surface are smaller and less prominent; no larger papillae in ocular I am grateful to Dr C.C. Lu, Department of region; lateral integumentary ridge or fold Invertebrate Zoology, Museum of Victoria, for around mantle circumference absent. No infor- his assistance, and comments on the manu-

mation is available on colouring of live animals. script. I thank Ms S.E. Boyd and Dr G.C.B. Preserved specimens in ethyl alcohol light Poore of the same institution, for their com- brown to red brown dorsally, creamy red to light ments on the manuscript. I wish to thank indiv- brown ventrally. Some regions on the dorsum iduals and museums for their help in making have a mottled appearance. Surface of the raised collections available for study and providing tubercles usually darker than the background, information: The Australian Museum (Dr W.B. coloured brick red to dark brown, giving a speck- Rudman. Mr I. Loch, Mr P.H. Colman); led appearance. Ocelli absent. Museum of Victoria (Dr C.C. Lu, Ms S.E. Boyd, Ms R.J. Plant); Otago Museum (Dr A. Harris); Distribution. South-eastern Australia, from the and South Australian Museum (Mr W. Zeidler. Great Australian Bight to southern New South Ms K.L. Gowlett-Holmes). This work is from a Wales, including Bass Strait and northern Tas- thesis submitted for the partial fulfilment of the mania. An inshore species, living on reefs, or requirements of the Master of Science degree at rockv areas of sand, and among seagrass, at the University of Melbourne. The research was depths from 1-130 m (Stranks, 1988). done under the affiliation arrangement of the Museum of Victoria and the University of Mel- Etymology. The specific name bunurong is de- bourne, and was partially supported by: a Keith rived from the name of the Australian Aborigi- Sutherland Award, from the Australian nal clan which once inhabited the south-eastern Museum; Supplementary Grants for Student region of Melbourne, Victoria, and is to be Research, from the Victorian Institute of treated as indeclinable. Marine Sciences; and a Postgraduate Writing Award, from the Office for Research, Uni- Remarks. Octopus bunurong is a new species of Up medium sized octopus with an elongate ovoid versity of Melbourne. mantle and very long arms, from south-eastern References Australia. Macpherson (1966) had identified Benham, W.B., 1942. The octopodous Mollusca of two(NMVF5101 andNMV Fl 516; a juvenile New Zealand. 1. The midget octopus of the coas- male and a mature male respectively) from Port tal waters. Transactions and Proceedings of the Phillip Bay, Victoria, as O. flindersi. The speci- Royal Society ofNew Zealand 72(3): 226-236. pis mens are now re-identified as O. bunurong. 18. 19. Octopus bunurong appears closely related to Berry, S.S.. 1917. Cephalopoda. Australasian Antarc- O. kaurna, but may be readily distinguished tic Expedition. 1911-1914. Scientific Reports. C. from the latter by the characteristic skin pattern- Zoology and Botany 4(2): 5-38, pis 10-14. ing, the larger and more prominent eyes, the Brough, E.J.. 1965. Egg care, eggs and larvae in the midget octopus, Robsonella australis (Hoyle). larger suckers on all arms, and the longer Transactions ofthe Royal Society ofNew Zealand ligula. Zoology 6(2): 7-19, pis 1.2. O. bunurong is a distinctive species endemic Cotton, B.C., 1932. Notes on Australian Mollusca, to temperate waters of south-eastern Australia. with descriptions of new genera and new species. It easily species can be distinguished from other Records of the South Australian Museum 4(4): of Octopus on the basis of a combination of char- 537-547. THREE NEW SPECIES OF OCTOPUS 465

Dell. R.K.. 1952. The recent Cephalopoda erf New Pickford. G.E.. 1 955. A revision of the Octopodinae in Zealand. Dominion Museum 16' Bulletin 1- the collections of the British Museum. Bulletin of 157. the British Museum (Natural History). Zoology Dell. R.K.. 1959. Cephalopoda. British, Australian 3(3): 151-167. and New Zealand Antarctic Research Expedition Robson. G.C.. 1929. .-1 Monograph of the Recent Reports B N4r 89-105. Cephalopoda. Part 1. Octopodinae. British Hoyle. W.E.. 1SS5. Diagnoses of new species of Museum (Natural History): London. 236 pp.. 17 Cephalopoda collected during the cruise of pis. H.M.S. "Challenger". Pan 1. The Octopoda. Roper. C.F.E. and Voss. G.L.. 1983. Guidelines for Annals and Magazine of Natural History 5(15V taxonomic descriptions of species. 222-236. Memoirs of the National Museum of Victoria 44: Lamarck. J.B.. 1 798. Extrait d"un memoire sur le genre 49-63. la Seche. de du Calmar et du Poulpe. \ ulgairement Smith. E.A.. 1902. Mollusca. Pp. 201-213. pis 24. 25 nommes. Polypes de mer. Bulletin des Sciences. in: Reports on the collections of natural history par la Societe Plulomathique. Paris 2: 129-131. made in the Antarctic regions during the voyage of Lu. C.C. and Phillips. J.U.. 1 985. An annotated check- the "Southern Cross". British Museum (Natural list of the Cephalopoda from Australian waters. History): London. Occasional Papers from the Museum of Victoria 2: Stranks. T.N., 1988. Systematic of the Family Octo- 21-36. podidae (Mollusca: Cephalopoda) ofsouth-eastern Macpherson. J.H.. 1966. Port Phillip Survey 1957- Australia. Unpublished M.Sc. Thesis. University 1963. Mollusca. Memoirs ofthe Sational Museum of Melbourne. Victoria. 1 14 pp. of Victoria 27: 201-263. Tait. R.W.. 1982. A taxonomic revision of Octopus Massy. A.L.. 1916. Mollusca. Pan 2. Cephalopoda. australis Hoyle, 1885 (Octopodidae: Cephalo- British Antarctic ("Terra Nova") Expedition, poda), with a redescription of the species. 1910. NaturalHistory Reports. Zoology 2(7): 141- Memoirs of the National Museum of Victoria 175. 43(1): 15-23, pi. 1.