OCTOPUS MAGNIFICUS (CEPHALOPODA - CORE

BULLETINOFMARINESCIENCE49(1-2) 39-561991

OCTOPUS MAGNIFICUS (CEPHALOPODA OCTOPODIDAE)A NEW SPECIES OF LARGE OCTOPOD FROM

THE SOUTHEASTERN ATLANTIC

R Villanueva P Sanchez and M A Compagno Roeleveld

ABSTRACTA new species of octopod Octopus magnificus is described based on a total of 131 specimens

from the southeastern Atlantic The species has been collected from 2603S 13deg43pound to34deg29S 25deg36pound between 2-560 m of depth mainly on fine sandy bottom The species ischaracterized by its large overall size long ligula the anatomy of the male reproductive tractthe shape of the funnel organ skin loose and elongate folds present in live animals Omagnificus is compared to three large octopuses namely O dojleini O maorum and Enteroc-topus megalocyathus

Large octopod specimens collected for the South African Museum since 1971were first considered to be atributable to the giant octopus of the North PacificOctopus dojleini Wiilker 1910 However satisfactory assessment of the relation-ships between the South African and North Pacific forms was hampered by thepoor state of knowledge regarding the systematics of the latter The only reviewby Pickford (1964) was based largely on immature animals Recently 14 speci-mens of this large octopod were caught in the southeast Atlantic off Namibia fromJanuary to July 1988 Upon examination the specimens were considered to bedistinct from all previously described taxa and to represent a new species in thegenus Octopus The new species is described based on 14 type specimens and 117additional specimens Information also is provided on the biology and ecologyof the species in southeastern Atlantic Records of O dojleini from Crozet IslandsPrince Edward Island and Marion Island (Lu and Mangold 1978 Roeleveld1986) are attributed with some reservation to the new species Definitions ofmeasurements and indices after Robson (1929) Pickford (1964) Roper and Voss(1983) and Toll (1988) The relationship between ML and TW was calculated asfollows TW = amiddotMLh

Abbreviations -ICM-Instituto de Ciencias del Mar Barcelona Spain R S AF-RICANA-Research Vessel Africana SAM-South African Museum Cape TownSouth Africa USNM- United States National Museum Washington DC USA

Octopus magnificus new speciesFigures 1-8 Tables 1-5

Octopus dojleini (non Wiilker 1910) Augustyn and Smale 198997-104Octopus dojleini (non Wiilker 1910) Lu and Mangold 1978567-573 Roeleveld 19864

Material Examined-TypE MATERIALHolotype and paratypes measured fresh (Tables 12 Fig1) two males (ICM91 and ICM92) and one female (ICM90) from Namibian waters collected byFreezer Trawler SUEVEJanuary-March 1988 plus five males (ICM95 ICM94 USNM817139 SAM-S2363 ICM96) and six females (SAM-S2364 ICM99 ICM102 ICMI01 USNM817140 SAM-S2365) from the same area collected by the Freezer Trawler CHICHA-TOUZAJuly 1988 The specimenswere frozen at - 20degC immediately after capture They were measured weighed dissected and pho-tographed in the laboratory after thawing and then fixed in 10 formalin and preserved in 80 ethylalcohol

ADDITIONALMATERIALSPECIMENSMEASUREDFRESHForty one males and 57 females (Tables 12) The following data were collected ML TL and in males length of hectocotylized arm ligulacalamus penis penis-diverticulum and spermatophore as well as hectocotylized arm sucker count

39

40 BULLETINOFMARINESCIENCEVOL49 NO1-2 1991

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Figure I O magnificus new species Geographical distribution

Total weight and gonad weight were obtained After having been examined the specimens werediscarded Beaks were extracted from 93 specimens and preserved in 70 ethanol

SPECIMENSMEASUREDAFTERPRESERVATIONNine males and 10 females (Tables 15)

ADDITIONALDISTRIBUTIONRECORDSOne male 305 mm ML and 126 Kg TW 2 males 2 juveniles3 un sexed specimens 30003S to 36deg26S 15deg21E to 21deg33E 80-296 m depth November 1980 toSeptember 1987 crayfish trap and bottom trawl 6 males 2 females 29deg12S to 34007S 14deg45E to17deg36E 224-406 m depth 6-22 February 1988 bottom trawl R S AFRICANACruise 059 3 females33deg27S to 34deg31 S 17deg32E to 18deg22E 240-427 m bottom trawl 8-12 March 1988 R S AFRICANACruise 060 7 males I female 6 unsexed specimens 352-6675 g TW 34deg56S to 35deg42S 21deg40Eto 25deg36E 86-450 m depth bottom trawl 17-29 May 1988 R S AFRICANACruise 063 2 juveniles30006S to 30014S I 5deg54E to 16deg25E 182-207 m bottom trawl 21-25 January 1990 R S AFRICANACruise 079

SUBANTARCTICDISTRIBUTIONRECORDSTwo females 155 and 190 mm ML Transvaal Cove MarionIsland 15m depth gill net and fish trap 19 April and 25 August 1985 1 female Marion Island46deg4053S 37deg5098E 460-488 m depth dredge 24 August 1987

COMPARATIVEMATERIALOctopus dojleini dojleini 1 male 242 mm ML and 1 female of 189 mmML Wakkanai Hokkaido Japan January-March 1978 Octopus dojleini martini I male 265 mmML and I female 198 mm ML Howe Sound British Columbia Canada December 1989

Diagnosis-Animals large not exceeding 362 mm ML and 126 kg total weightfully mature female unknown length of hectocotylized arm 81-95 of opposite

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Figure 2 O magnijicus new species a Dorsal view of the holotype male 275 mm ML SAM-S2363 b Ligula of holotype (ligula length = 627 mm)

arm ligula 14-22 ofhectocotylized arm in mature males spermatophores verylarge 160-283 ML beaks without grooves on rostrum gills with 12-15 lamellaein outer demibranch funnel organ wide W-shape with lateral limbs about 50as long as medial limbs skin loose elongate folds present in live animals withoutpapillae one large supraorbital papilla over each eye

Description-Octopods attaining a large size maximum total length 1673 mmand total weight 126 kg Mantle saccular slightly ovoid mantle wall muscularwell developed Mantle opening broad reaching to posterior limit of eyes Headnarrow set off by narrow well defined neck and brachial crown (Fig 2a) Eyesmoderate in size not prominent somewhat protruding in preserved specimensA single large conical primary supraorbital papilla (10-15 mm) located mediallyabove each eye more readily apparent in fresh or thawed specimens than inpreserved specimens A second small papilla occasionally present over each eyelocated anterior to large papilla (Fig 3) Funnel tubular free for half or more itslength Funnel organ broadly W-shaped lateral limbs considerably shorter thanmedial limbs Brachial crown well developed Arms long with attenuated tips

VILLANUEVA ET AL OCTOPUS MAGNlFICUS NEW SPECIES 45

Figure 3 O magnificus new species Female 139 mm ML SAM-S2347 in aquarium

muscular and robust proximally Arm pairs I and II subequal in length longerthan arm pairs III and IV which also are subequal Arm formula typically I = IIgt III = IV with considerable variation Suckers biserial proximal seven pairsof suckers small to medium followed distally by eight larger pairs remainingsuckers decreasing in size to the tip Specially enlarged suckers absent in malesand females Average arm sucker count 206-294 in type specimens Averagesucker count in each pair of arms in type specimens was 246 243 264 280 inarms I-IV respectively (excluding hectocotylus) Web well developed always shal-lowest in sectors A and E remaining sectors variable general formula not dis-cernible Web extending 84 along ventral side of arms I and II and 80 alongarms III and IV (excluding hectocotylus) Right third arm of males hectocotylizedshorter than opposite arm (OAI = 81-95 in mature males) Hectocotylized armsucker count ranging from 92-126 in 46 males examined (Fig 4) Spermatophoricgroove well defined ending at base of calamus Ligula large (LLI = 14-222 inmature males) inner surface ofligula sparsely pigmented flat not subcylindricalwith sides slightly curved inwards in the holotype (Fig 2b) Ligula tip slightlypointed Several rows of copulatory papillae present in medial portion of ligulalongitudinal or transverse striations absent Calamus small (CaLI = 46-187)unpigmented with median longitudinal groove tip pointed and slightly erectGills with 12-15 lamellae in outer demibranch (usually 13-14) including terminallamella Digestive tract typical of genus (Fig 5a) A pair of flattened medium-sized anterior salivary glands attached to posterior portion of the buccal massPosterior salivary glands almond-shaped yellowish after fixation joined by sal-ivary duct to posterior portion of buccal mass Esophagus narrow followed by alarge expanded longitudinally striated crop without crop diverticulum Stomach

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subspherical muscular with single connection to pinkish caecum and two con-nections to digestive gland Ink sac embedded in digestive gland visible at surfaceIntestine long ending in rectum at level of posterior salivary glands Anal flapsminute Mandibles lack distinctive features without grooves on rostrum (Fig5b) Radula exhibit great individual variability (Fig 5c d e) According to radulaformula defined by Adam (1941) seriation of ectocones on rachidian teeth typeA3222 in two specimens (ICM92 150 mm ML and ICM99 268 mm ML) andseration of type B221 in another specimen (ICM102 270 mm ML) First lateraltooth in specimens ICM99 and ICMI02 with acute mesocone but devoid ofectocones first lateral tooth in specimen ICM92 bears an ectocone but no me-socone Second lateral tooth with a broad endocone and curved base Third lateraltooth long and sabre-shaped Marginals rectangular plate-like Female reproduc-tive system (Fig 6a) comprised an ovary and two yellowish cream colored ovi-ducts The oviduct is common when it leaves the ovary Distal oviducts longerthan proximal oviducts Distal hemisphere of oviducal glands dark brown inmaturing females Maturing ovarian eggs elongate tear-shaped with 10-12 follic-ular folds attached in ovary in yellowish white clusters by a long thin stalkLength of chorion chamber in ovarian eggs 6-8 mm Ovary of maturing female(268 mm ML ICM99) weighed 413 g and contained about 20000 eggs Spawnedeggs unknown Fresh gonad of maturing females 62-87 of total weight (gonadweight 450-700 g in 53 females of5300-1O350 g TW) Male reproductive system(Fig 6b) with single and extremely long penis diverticulum up to 918 of penislength Fresh gonad of mature males 46-95 (mean 62) of total weight (15mature males of 4500--11400 g TW had gonad weights of 280-795 g) Sper-matophores very long 532-870 mm in fresh specimens (SpLI = 161-283) andfew in number always one spermatophore found in penis-diverticulum and 2-8in Needhams sac Oral cap of spermatophore simple rounded Flagellum emerges


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Figure 7 O magnificus new species Statoliths A Female 270 mm ML ICM102 anterior view ofright statolith B Female 268 mm ML ICM99 ventral view of right statolith C Female 270 mmML ICM102 posterior view ofleft statolith

from oral cap Sperm reservoir coiled occupying approximately half of totalspermatophore length (Fig 6c) Skin soft smooth and loose in recently dead andfrozen specimens In life skin erected into large longitudinal folds and smallgranular papillae of varying size and density Large supraorbital papilla presentabove posterior margin of each eye Color oflive animals varying from yellowishochre to reddish brown or chocolate brown on dorsal surfaces of mantle headand arms paler ventrally Edges of longitudinal folds frequently white Recentlydead animals with 25-35 faint longitudinal rows of dark yellow chromatophoreson dorsal mantle surface also extending to head Ventral surface of mantle yel-lowish white After fixation dorsal surface of mantle and arms reddish brownwith more distinct rows of chromatophores taking on a purplish blue or darkreddish blue Stellate ganglia (ICM91 240 mm ML) with 16-17 large nerves and6-8 very thin lateral nerves radiating outwards Optic complex dissected in twoimmature animals (ICM9l 240 mm ML and ICM99 268 mm ML) Optic glandsmall rounded located at the base of olfactory lobe yellow in color in bothindividuals White body well developed spongy enveloping optic nerves andattached to inner surface of the eye ball Statoliths (Fig 7a-c) with flat ovalattachment area oval border unserrated and groove extending halfway into thearea Posterior surface acutely pointed projecting beyond border of anterior sur-face and with narrow median furrow along entire length of posterior surfacestarting at the point and ending at opposite end Smaller diagonal grooves radiateover posterior surface from the median furrow Shell vestige (ICM99 268 mmML) consist oftwo symmetrical rods located above gills V-shaped with widelyseparated limbs

Holotype -Maturing male 275 mm ML preserved in 80 ethyl alcohol SouthAfrican Museum SAM-S2363

Type locality-Southeastem Atlantic off Namibia 29deg4lS l4deg4lE in 415 m

Etymology - The specific name magnificus refers to its large size and impressiveappearance


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Biological Data - The Namibian and South African waters are characterized bythe cold Benguela Current upwelling system (Shannon 1985) O magnificus hasbeen collected at depths from 2-560 m on soft bottom sediments consisting ofvery fine clay and calcareous sand in waters with 62-13deg bottom temperaturesThe south coast of South Africa is influenced both by the cold Benguela Systemin the west and the wann Agulhas Current from the east in this region O mag-nificus was collected in temperatures of83deg-135degC at 86-450 m The occurrenceof O magnificus on rocky area in Benguela and Agulhas currents has not beenassessed (Fig 1) In a total of 131 specimens examined the sex ratio approachedparity with 435 males Length-weight relationships showed no significant dif-ferences between males and females (t-Student P gt 005) (Table 3 Fig 8) Nodifferences in depth distribution were found for either sex or size Mature maleswere found in summer autumn and winter but only one immature male has beencollected in spring The smallest mature male examined weighed 425 kg (ML184 mm) Mature spaWJOingfemales are unknown

DISCUSSION

O magnificus displays several morphological characters similar to several oc-topod species O dojleini Willker 1910 O maorum Hutton 1880 and Enter-octopus megalocyathus (Gould 1852) Despite the high commercial value of Odojleini(Kanamaru 1964 Kanamaruand Yamashita 1967 Mottet 1975 Nesis1987) extensive work on its physiology (Mann et al 1970 Martin et al 1970Packard and Hochberg 1977) and ecology (Gabe 1975 Hartwick 1983 Hartwicket al 1984) its systematic status is unclear Current opinion is that it probablyconstitutes a group of closely related species (K Mangold and F G Hochbergpels comm) The most recent systematic account by Pickford (1964) concludedthat the giant Pacific octopus should be treated as three separate subspecies of O


Table 3 O magnificus new species Parameters of the power curve TW = amiddotDMLb (n number ofspecimens Vb standard error of b coefficient)

MalesFemalesAll specimens

a

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b

322304311

r

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n

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Vb

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dojeini in adjacent geographic regions O d dojeini in the temperate westernPacific (Japan and Korea) O d apollyon in the subarctic northern Pacific (SiberiaAleutians and Alaska) and O d martini in the temperate eastern Pacific (Wash-ington to California USA) Redescriptions of all three subspecies based on anadequate number of specimens are urgently required for the resolution of thesystematic tangle surrounding the O dojeini species complex

Comparison with O dofleini - The relationship of O magniicus with O dojeini(Tables 4 5) is indicated by the similar sucker counts on the hectocotylus (HASC= 98 and 109 in O dojleini HASC = 92-126 in O magniicus large ligula LLI= 16-25 in O dojeini (Pickford 1964) LLI = 14-22 in mature O magnificusextremely large spermatophores SpL = 600-1150 mm SpLi = 342-346 in Odojleini (Kanamaru 1964 Mann et al 1970 Sasaki 1929) and SpL = 532-870mm SpLi = 1375-3056 in O magnificus and long penis diverticulum PdLlapproximately 86 in O dojeini (Akimushkin 1965) and PdLl = 65-918 in Omagnificus However the two species also possess clearly distinct characters Omagnificus differs from O d martini in the relative length of the hectocotylizedarm OAI rarely less than 90 (range 81-109 mean 94) in preserved O d martiniand OAI = 76-88 in preserved O magnificus (Table 5) Two small O d dojleinihad opposite arm indices very similar to those of O magnificus (all maturitystages) but fresh mature males had non-overlapping ranges for this characterOAI = 669-783 in O d doj1eini (Sasaki 1929) and OAI = 786-946 in Omagnificus Preserved specimens of O d apollyon had OAI very similar to rangesfor preserved O d dojleini and O magnificus O magnificus differs from O dapollyon in sucker size In O d apollyon the suckers are relatively larger especiallyin females (SI = 155-205 for males SI = 144-185 for females) than in Omagnificus (SI = 97-227 for males SI = 89-135 for females) O d doj1eini isalso distinct in having a funnel organ with lateral limbs almost as long as themedial limbs whereas in O d apollyon O d martini and O magnificus thelateral limbs of the funnel organ are considerably shorter than the medial limbsO d apollyon differs from O magnificus and the other two subspecies O ddoj1eini and O d martini in having a conspicuous groove on the rostrum of bothupper and lower beaks in large specimens Transverse striations in the ligula havebeen reported for O d dojeini by Sasaki (1929) and Akimushkin (1965) In Omagnificus copulatory papillae may be present in the ligula but distinct transversestriations are not present Skin color and texture of octopods (Packard and Hoch-berg 1977 Hanlon 1988) have not received much systematic attention but maywell prove to be of systematic value once intraspecific variation has been estab-lished and character states codified The skin type observed in O dojleini sparselyornamented with small papillae (of which) four larger papillae may form a dorsalrhomb in young animals (Pickford 1964 p 48) was not found in either freshor preserved O mangificus Sasaki (1929 p 73) also reported warts of variousshape but usually pointed at apex and more or less stellate at base often connected


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together into longitudinal lines and some larger tubercles in addition to thesewarts ( ) clearly perceptible in regular order on the dorsal surface Thesecharacters were also present in our comparative specimens of O d dojleini andO d martini but were not found in O magnificus

Comparison with other Large Octopodsfrom the Southern Hemisphere -0 mao-rum Hutton 1880 is distributed in the temperate waters of southeastern Australiaand also in the temperate and subantarctic waters of New Zealand O maorumattain 255 mm ML (Table 5) O maorum differs from O magnificus in havingspermatophoral groove poorly developed LLI = 3-6 in mature animals peniswith a single coiled diverticulum marked with three lobes (PLI = 76-141-214)(middle number separated by - indicate the mean) spermatophores relativelyshort (SpLI = 348-685-1013) mature eggs without follicular folds males ma-turing at approximately 90 mm ML and females attaining ovarian maturity atabout 120 mm ML (Stranks 1988)

Enteroctopus megalocyathus (Gould 1852) is distributed in the southwesternAtlantic from Argentinian waters (Gulf of San Jose 42degS) to the Straits of Ma-gellan Beagle Channel and Tierra de Fuego (Re 1980) E megalocyathus attain240 mm ML (Re 1984) Although the male genital system ligula and calamusare similar in both species (Table 5) E megalocyathus differs from O magnificusin having large ovarian eggs (9-17 mm) 11-13 gill lamellae in outer demibranchfunnel free for 13 its length well developed anal flaps relatively small head (HWI= 429-499-563 in males HWI = 328-41-465 in females) spermatophoresranging from 95-390 mm (SpLI = 70-221) in males attaining sexual maturity at115 mm ML and females at 120 mm ML (Re 1984)

Similarities within the group of large octopods (0 dojleini O maorum Omagnificus and Enteroctopus megalocyathus) suggest probable generic relation-ships that may distinguish them from the genus Octopus but this requires seriousrevision (K Mangold and F G Hochberg pers comm)

Comparison with other Octopods from the Genus Octopus in the southeasternAtlantic and southwestern Indian Oceans - Three species of the genus Octopusare present in the southeastern Atlantic O vulgaris Cuvier 1797 O schultzei(Hoyle 1910) O burryi Voss 1950 and three more species occur in the watersof the southwestern Indian Ocean off southern Africa O aegina Gray 1849 OfontanianusOrbigny 1834 and O horridus Orbigny 1826 (Robson 1929 Adam1962 Voss 1962 Roeleveld 1974)

O vulgaris Cuvier 1797 is differentiated from 0 magnificus by its small ligula(LLI = 12-21) relatively long calamus (CaLI = 47-52) and medium sized sper-matoprdres (SpLI = 31-61) (K M Mangold and F G Hochberg pers comm)

O schultzei (Hoyle 1910) is differentiated by arm tips with numerous filamentsinstead of suckers except hectocotylus and hectocotylus with bluntly roundedligula devoid of longitudinal depression calamus absent (Roper and Mangold1991)

O burryi Voss 1950 is differentiated by its small size maximum 45 mm MLin African specimens (Adam 1960a) longitudinal dark purplish brown stripealong the dorsolateral surface of each arm and LLI = 2-6

O horridus Orbigny 1826 is differentiated by reddish-purple body surfaceornamented with white circular patches each with a central papilla ligula withof a conspicuous transverse groove below the calamus and LLI = 4 (Adam 1959)

O fontanianus Orbigny 1834 is differentiated by small circular papillae andgranules over the body surface ligula with two conspicuous cheeks and prom-inent calamus (Robson 1929)

O aegina Gray 1849 is differentiated by the dorsal arm pair shorter than the


rest hectocotylus with an indistinct spermatophore groove LLI 5 (Adam1960b 1962 Okutani et at 1987)

Records of Octopus dofteini in Subantarctic waters - O dojleini Wiilker 1910has been reported in sub-Antarctic waters Prince Edward Marion and CrozetIslands (Lu and Mangold 1978) Transvaal Cove Marion Island (Roeleveld1986) These records are attributed with some doubt to O magnificus The spec-imens cited by Roeleveld (1986) are all immature females and have relativelylower gill counts the Marion Island specimens have 10-11 filaments in the outerdemibranch compared with 12-15 in the southern African specimens

ACKNOWLEDGMENTS

The authors with to thank Dr A Guerra Instituto Investigaciones Marinas Vigo Spain for criticalcomments of the first draft and bibliography provided Dr K Mangold and Dr S v BoletzkyLaboratoire Arago Banyuls-sur-Mer France were helpful with systematic discussions Special thanksto Dr F G Hochberg Santa Barbara Museum of Natural History California USA for criticalreading ofthe last draft and helpful comments For providing invaluable specimens of O dojleini wegratefully acknowledge Dr T Okutani Tokyo University of Fisheries Japan and Dr B HartwickSimon Fraser University Vancouver Canada We are also grateful to J M Fortuno for technicalassistance with scanning electron micrography to 1 Biosca D Gerneke M van der Merwe and LHoenson for photographic and technical assistance M A Compagno Roeleveld would like to thankthe Director and staff of the Sea Fisheries Research Institute Cape Town and the captain and crewof the RS Africana for permission to participate in the Hake Biomass Surveys and for additionalspecimens and data She would also like to thank other donors of specimens Dr P ZoutendykNational Institute for Oceanology Stellenbosch Dr G Ross previously of the Port Elizabeth MuseumDr O Gon J L B Smith Institute ofIchthyology Grahamstown and Dr L 1 V Compagno SouthAfrican Museum for advice and assistance with computer problems This study was supported by theproject Investigaciones en las costas de Namibia Instituto de Ciencias del Mar Barcelona Spain

LITERATURE CITED

Adam W 1941 Notes sur les Cephalopodes 15 Sur la valeur diagnostique de la radule chez lesCephalopodes Octopodes Bull Mus Hist Nat Belg 17(38) 1-19

-- 1959 Les Cephalopodes de la mer Rouge Result Scient Miss R P Dollfus en Egypte1927-192928 125-193 pis 1-9

-- 1960a Les Cephalopodes de Ilnstitut Francais d Afrique Noire 2 Bull Inst Franc AfrNoire A 22(2) 465-511

-- 1960b Cephalopoda from the Gulf of Aqaba Bull Sea Fish Res Stn Israel 26 3-26-- 1962 Cephalopodes de lArchipel de Cap-Vert de IAngola et du Mozambique Trabhs

Cent BioI Pisc Lisb 32 7-64Akimushkin I I 1965 Cephalopods of the Seas of the USSR (Transl of 1963 Russian ed by

A Mercado) Israel Program for Scientific Translations Jerusalem 223 ppAugustyn C J and M Smale 1989 Cephalopods Pages 91-104 in A I L Payne and R J M

Crawford eds Oceans of life off southern Africa VIaeberg Publishers Cape TownGabe S 1975 Reproduction in the giant octopus of the North Pacific Octopus dojleini martini

Veliger 18 146-150Hanlon R T 1988 Behavioral and body patterning characters useful in taxonomy and field iden-

tification of cephalopods Malacologia 29 247-264Hartwick B 1983 Octopus dojleini Pages 277-291 in P R Boyle ed Cephalopod life cycles Vol

I Species accounts Academic Press LondonHartwick E B R F Ambrose and S M C Robinson 1984 Dynamics of shallow-water populations

of Octopus dojleini Mar BioI 82 65-72Kanamaru S 1964 The octopods of the coast of Rumoi and the biology ofMizudako Mon Rep

Hokkaido Fish Exp Stn 21(4amp5) 189-210 (Translated by M Mottet)--and Y Yamashita 1967 The Octopus Mizudako Part I Ch 12 Investigations of the marine

resources of Hokkaido and developments of the fishing industry 1961-1965 In M Mottet Atechnical report on the fishery biology of Octopus dojleini (Translated by M Mottet) WashingtonState Dept of Fisheries Seattle

Lu C C and K Mangold 1978 Cephalopods of the Kerguelenian province of the Indian OceanProc Int Symp Mar Biogeogr Evol Southern Hemisphere 2 567-573


Mann T A W Martin and J B Thiersch 1970 Male reproductive tract spermatophores andspermatophoric reaction in the giant octopus ofthe North Pacific Octopus dojleini martini ProcR Soc Lond B 175 31--61

Martin A W J B Thiersch H M Dott A P Harrison and T Mann 1970 Spermatozoa ofthegiant octopus of the North Pacific Octopus dojleini martini Proc R Soc Lond B 175 63-68

Mottet M G 1975 The fishery biology of Octopus dojleiniWUlker Tech Rep Wash Dep ofFish16 1-39

Nesis K N 1987 Cephalopods of the world (Translated by B S Levitov) TFH PublicationsNeptune City 351 pp

Okutani T M Tagawa and H Horikawa 1987 Cephalopods from continental shelf and slopearound Japan Jap Fish Res Consv Assoc 196 pp

Packard A and F G Hochberg 1977 Skin patterning in Octopus and other genera Symp zoolSoc Lond 38 191-231

Pickford G E 1964 Octopus dojleini (WUlker) the giant octopus of the North Pacific Bull BinghamOceanogr Coli 19 1-70

Re M E 1980 Estudio taxon6mico de Enteroctopus megaocyathus (Gould) (Cephalopoda Oc-topoda) con notas sobre su biologia y pesca Centro Nac Pat 53 1-34

-- 1984 Maduraci6n sexual de Enteroctopus megaocyathus (Cephalopoda Octopoda) CentroNac Pat 93 1-28

Robson G C 1929 A monograph of the recent Cephalopoda Part 1 Octopodinae British MuseumLondon 236 pp 7 pIs

Roeleveld M A 1974 Cephalopoda Pages 743-752 in K H Barnard ed Contributions to theknowledge of South African marine Mollusca Part VII Revised fauna list Ann S Afr Mus47(5) 663-781

-- 1986 Octopus dojleini from Marion Island SASCAR Newsletter 22 4Roper C F E and K M Mangold 1991 Octopus schutzei (Hoyle 1910) a redescription with

designation of Aphrodoctopus new genus (Cephalopoda Octopodinae) Bull Mar Sci 49 57-72-- and G L Voss 1983 Guidelines for taxonomic descriptions of cephalopods species Mem

Nat Mus Vict 44 49-63Sasaki M 1929 A monograph of the dibranchiate cephalopods ofthe Japanese and adjacent waters

J Fac Agr Hokkaido Imp Univ 20(1) 1-357Shannon L V 1985 The Benguela ecosystem Part I Evolution of the Benguela physical features

and processes Oceanogr Mar BioI Ann Rev 23 105-182Stranks T N 1988 Systematics of the Family Octopodidae (Mollusca Cephalopoda) of South-

eastern Australia M Sc Thesis Univ of Melbourne Victoria 114 ppToll R B 1988 The use of arm sucker number in octopodid systematics (Cephalopoda Octopoda)

Amer Malac Bull 6(2) 207-211Voss G L 1962 South African cephalopods Trans R Soc S Afr 36 245-272

DATEACCEPTED December 17 1990

ADDRESSES (R V) (PS) Instituto de Ciencias del Mar Paseo Naciona sin 08039-Barcelona Spain(MACR) South African Museum PO Box 61 Cape Town 8000 South Africa


15middot 20middot 25middot 30middot 35middot

15middot

20middot

25middot

30middot

35middot

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Current

15middot

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25middot

30middot

35middot

40middot5

Figure I O magnificus new species Geographical distribution

Total weight and gonad weight were obtained After having been examined the specimens werediscarded Beaks were extracted from 93 specimens and preserved in 70 ethanol

SPECIMENSMEASUREDAFTERPRESERVATIONNine males and 10 females (Tables 15)

ADDITIONALDISTRIBUTIONRECORDSOne male 305 mm ML and 126 Kg TW 2 males 2 juveniles3 un sexed specimens 30003S to 36deg26S 15deg21E to 21deg33E 80-296 m depth November 1980 toSeptember 1987 crayfish trap and bottom trawl 6 males 2 females 29deg12S to 34007S 14deg45E to17deg36E 224-406 m depth 6-22 February 1988 bottom trawl R S AFRICANACruise 059 3 females33deg27S to 34deg31 S 17deg32E to 18deg22E 240-427 m bottom trawl 8-12 March 1988 R S AFRICANACruise 060 7 males I female 6 unsexed specimens 352-6675 g TW 34deg56S to 35deg42S 21deg40Eto 25deg36E 86-450 m depth bottom trawl 17-29 May 1988 R S AFRICANACruise 063 2 juveniles30006S to 30014S I 5deg54E to 16deg25E 182-207 m bottom trawl 21-25 January 1990 R S AFRICANACruise 079

SUBANTARCTICDISTRIBUTIONRECORDSTwo females 155 and 190 mm ML Transvaal Cove MarionIsland 15m depth gill net and fish trap 19 April and 25 August 1985 1 female Marion Island46deg4053S 37deg5098E 460-488 m depth dredge 24 August 1987

COMPARATIVEMATERIALOctopus dojleini dojleini 1 male 242 mm ML and 1 female of 189 mmML Wakkanai Hokkaido Japan January-March 1978 Octopus dojleini martini I male 265 mmML and I female 198 mm ML Howe Sound British Columbia Canada December 1989

Diagnosis-Animals large not exceeding 362 mm ML and 126 kg total weightfully mature female unknown length of hectocotylized arm 81-95 of opposite


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b

322304311

r

092091091

n

435699

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ACKNOWLEDGMENTS


LITERATURE CITED












































+ o

300200-+-1 bullbull

100

o

Qmagnificus

o males+ females

2




DISCUSSION





a

000007620000229100001479

b

322304311

r

092091091

n

435699

Vb

015013010




00

Do00

NN00N

Do


00

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bull

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bullltt- 00

ac

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tvlIVI_jli-ON0--0

VI-r-NJJN-N

1 10- -00000-

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llVI --N

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~l- 00

or00I

VI-

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obullI

00-0V

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1

-0j

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00

VI

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INr


JJotjN bull-

-0 00000 I 10- 0-0010- 0-bull

NV

N00 bull1-0l1 -11N 00-0 N


r----~oocilIlfl~--

0000

IortoIl

v

00N

I00j-J0-

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10-bullcA--

oNN


lo
















ACKNOWLEDGMENTS


LITERATURE CITED







































a

000007620000229100001479

b

322304311

r

092091091

n

435699

Vb

015013010




00

Do00

NN00N

Do


00

JbullI

bull

ltt00

Jt-I

bull

ltt00I

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~ Ifl- rtI QO t--


ooN0 --q

000 - C7I

bullltt- 00

ac

rJ


~~00+-

INj

I-

VI

tvlIVI_jli-ON0--0

VI-r-NJJN-N

1 10- -00000-

_N

jNN

1 1bullNtoojIro

llVI --N

I 1-0-N

~l- 00

or00I

VI-

I00j

obullI

00-0V

N0-0-

1

-0j

I-00I

00

VI

-0j-Ior00

INr


JJotjN bull-

-0 00000 I 10- 0-0010- 0-bull

NV

N00 bull1-0l1 -11N 00-0 N


r----~oocilIlfl~--

0000

IortoIl

v

00N

I00j-J0-

orN

10-bullcA--

oNN


lo
















ACKNOWLEDGMENTS


LITERATURE CITED





































00

Do00

NN00N

Do


00

JbullI

bull

ltt00

Jt-I

bull

ltt00I

00bullet-

o00IbullbullIbull




~ Ifl- rtI QO t--


ooN0 --q

000 - C7I

bullltt- 00

ac

rJ


~~00+-

INj

I-

VI

tvlIVI_jli-ON0--0

VI-r-NJJN-N

1 10- -00000-

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jNN

1 1bullNtoojIro

llVI --N

I 1-0-N

~l- 00

or00I

VI-

I00j

obullI

00-0V

N0-0-

1

-0j

I-00I

00

VI

-0j-Ior00

INr


JJotjN bull-

-0 00000 I 10- 0-0010- 0-bull

NV

N00 bull1-0l1 -11N 00-0 N


r----~oocilIlfl~--

0000

IortoIl

v

00N

I00j-J0-

orN

10-bullcA--

oNN


lo
















ACKNOWLEDGMENTS


LITERATURE CITED





































~~00+-

INj

I-

VI

tvlIVI_jli-ON0--0

VI-r-NJJN-N

1 10- -00000-

_N

jNN

1 1bullNtoojIro

llVI --N

I 1-0-N

~l- 00

or00I

VI-

I00j

obullI

00-0V

N0-0-

1

-0j

I-00I

00

VI

-0j-Ior00

INr


JJotjN bull-

-0 00000 I 10- 0-0010- 0-bull

NV

N00 bull1-0l1 -11N 00-0 N


r----~oocilIlfl~--

0000

IortoIl

v

00N

I00j-J0-

orN

10-bullcA--

oNN


lo
















ACKNOWLEDGMENTS


LITERATURE CITED



















































ACKNOWLEDGMENTS


LITERATURE CITED




































OCTOPUS MAGNIFICUS (CEPHALOPODA - CORE

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Transcript of OCTOPUS MAGNIFICUS (CEPHALOPODA - CORE