NEW ANDNEW

MemoirsoftheMuseumofVictoria 53(2):345-373(1992)

REVIEW OFTHEOCTOPUSAUSTRALISCOMPLEX FROMAUSTRALIA AND NEW ZEALAND. WITH DESCRIPTIONOFANEWSPECIES(MOLLUSCA: CEPHALOPODA)

ByT. N. Stranks'and M.D. Norman²

"Department ofInvertebrateZoology.MuseumofVictoria,285-321 RussellStreet,Melbourne Victoria 3000, Australia

-Departmentof Zoology.Universityof Melbourne.Parkville. Victoria 3052, Australia and.Department of InvertebrateZoology. MuseumolVictoria,285-321 RussellStreet,Melbourne,Victoria 3000, Australia

Abstract

Stranks.T.N. andNorman,M.D., ¹992.ReviewoftheOctopusauslraliscomplex from Australiaand NewZealand,with descriptionsofanewspecies(Mollusca:Cephalopoda).

MemoirsoflheMuseum_ofVictoria53:345-373.

Fourspeciesofshallow water octopusesbelongingtotheOctopusauslraliscomplexare reportedfromAustraliaandNewZealand.O. auslralis Hoyle,1885,isredescribedbasedon thetype materialandotherspecimensfromsubtropical watersofeastern Australia.Three otherspecies are distinguishedandcharacterised: O.berrimasp. nov.,isdiagnosed from temperate waters ofsouth-eastern Australia;andO.wartlngaStranks.1990.from temperate watersofAustraliaand NewZealand,andO, camphclli (Smith. 1902)fromsubantarctic watersoffNewZealand,areredescribed.

Introduction

Theinshoreand deepwater octopodid fauna ofAustralia isslowly becomingbetterknown.

Stranks (1988)revised the inshoreoctopusesof south-eastern Australia, and togetherwe con- tinueresearch into the systeraaticsof octopuses of temperate,subtropicalandtropical watersof Australia.Stranks (1988: 1990; in press) men- tionedproblemsinvolving acomplexofshallow waterspecies previously attributed to Octopus auslralis Hoyle. 1885. The species within the complex are small tomediumsized octopuses with broadlyovoidmantles, longarms,fineand roundedtuberclesontheskin,andsometimesa fold or ridge of skin on the ventrolateral mantle.

Theapplicationof thespecificnameauslralis to AustralianandNewZealandlaxa^isconfused.

AsTail(1982: 19) stated,"the confusion relat- ing to the identityof OctopusauslralisHoyle^is due ^{largely to} the lack of a mature male type specimen."Thebrief original descriptionofthe immature and submature^{typespecimens,}and lackofadetailedcomparativestudy of thetypes,

have resulted in the name O. auslralis being applied to several similar taxa from ^different biogeographicregions.

Threetaxaconfusedwith O.auslralis,bulnow knownto constitute separate^species,are herein listed as: O. berrima sp. nov., O. camphclli (Smith, 1902)and O. wanittgaStranks, 1990.

Recentpublications dealing with the O.^auslralis

complex are by Tait (1982), Stranks (1988,

1990).O'Shea( 1990) andToll(1991).

Thispaperresolves theconfusion concerning the fourspecies. Detailed diagnoses, basedon type materials plus a comprehensive seriesof specimens,areprovidedfortwoof the taxa(O.

auslralis and O. berrima). The few available specimens of 0. campbelli were examined to diagnosethisspecies.Stranks (1988. 1990) pro- vided detailed descriptionsofO. warringa and the readerisreferredtothoseaccountsfor infor- mation.

Wherethereissufficientmaterial,countsand measurementsare included for 10 representa- tive females and 10 males. Localities for all

specimen lots are mapped. Counts, measure- ments andindices weredefined byRoperand Voss (1983). Measurements and indices are listed throughoutasrangeswith themean itali- cized.Otherabbreviations usedare:F-female.

M-male, ML-mantle length and TL-total length. Material is lodged in collections of:

Auckland Institute and Museum, Auckland (AIM);TheAustralianMuseum,Sydney(AM);

The Natural History Museum, London (BMNH); Museum of Victoria, Melbourne (NMV); Queensland Museum, Brisbane(QM);

Queen Victoria Museum and Art Gallery, Launccston(QVM);SouthAustralianMuseum, Adelaide (SAM); andTasmanian Museumand Art Gallery. Hobart(TM).

345

https://doi.org/10.24199/j.mmv.1992.53.15 31 December 1992

346 T.N.STKANKSAND^M. D,NORMAN

Oiiopodidae Octopus Lamarck, ^I748

Typexpetitai. OctopusvulgarisLamarck, 1798.

Diagnosis.Benthicoctopodids.Mantlesaccular, without lins. 8 arms lacking cirri, arms with liiscrialsuckers, thirdrightarmofmaleshecto- cotylised withend ofarm modified intoligula

andcalamus.Webwelldeveloped.Ink sac pres- ent. Mantleaperturewide. Internalshellcarti- laginousand vestigial.

Octopusaustralis Floyle Figures 1-5, II

OnanitiauwatliHoyie, \Wy,v, 224.— 1885b: 9K.

1886:88.pi. 3, figs4. 5. Brazier, 18915. -

RobSOrt, 1929: 144.texttig.SI(partun).—1nil.1982:

15,texttigs I, 2. pi. I (piirlnii).

Materialexamined, See Table I.

Types. ^Iectotype (here designated): female, 23.3

mmML:paralectotvpc: male. 12.1)mmML: BMNtl

1889.4.24,28-29; preservedinethylalcohol.

/V/v locality. Australia, New South Wales. Port Jackson I.Lr.sb'S. IJWEj, ^(>-15 fathoms [11-28 m].

Description.Counts,measurements andindices listedinTables2-4.Mediumsizedanimalswilh firm consistency (Fig- la). Mantle saccular, broadly ovoid (MWI 56.3-80. 7-98.5); mantle wall moderately thin, muscular. Head narrow (HWI41.0-5/.7-64.6):demarked from mantle bymoderateconstriction.Eyessmall, projecting above surface of head. Funnel large, slender.

bluntly tapered(Fig. lb:I uLl40.4-45.⁷-54.6):

free forabout halfitslength (FFul 18.0-2 7V 36.4). Funnel organ consisting of 2 closely

opposed V-shaped units; outerlimbs approxi- matelyas long asmedianlintbs (Fig.Ic).Mantle aperturewide (PA1 71.0-97.7-125.5},

Brachialcrownstrong, welldeveloped. Arms long (MAI 23,4-.</.4-44.3) (2,7-4.5 times mantlelengthinmatureanimals);slender(AWI

6.4-<>.6-13.6):taperingto linetips.Armlengths SUbequal,arm orderusuallyII1.II.IV.I.Suckers bisenal.without obviousradial grooves',moder-

ately sized (ASI females 6.3-^.7-13.7, males 7.7-//..'-15.3): ¹6th lo 20th suckers enlarged onarms^IIand IIIofmaturemalesonly.

Web formula variable. CDBAF lo OCBE.V.

dorsal and ventral sectors alwavs shallower.

Webshallow(WD1 15.3-22 7-31.6);webrem- nants extend up ventral sides of arms lor approximatelythree-quartersoftheirlength.

Thirdrightarmof maleshectocotylised (Figs Id-f); shorterthan its opposite number(OA1

72.0-7,0-89.0; McAl 170.5-2.f2. 7-258.8).

Heclocotylised arm with 62-77 suckers;

oppositearmwith 124-217suckers.Spermato- phoralgroovewelldeveloped, with conspicuous thickeningofweb membrane. Ligula8-17%of hectocotylised arm length in mature animals (LLI7.5-/2. 7-16.7). Ligulastout,bulbous,with well marked and deep groove, and 2 rows ot minute papillae present along the groove.

Calamus very short, acutely pointed (CaLl 15.0-/9.5-29.4).

Gillswith 7-9lamellaeonouterdemibranch, plus the terminal lamella.

Digestivetract typicallyoctopodan(Fig. 2a).

Cpperbeak has ^{short, blunt,} curved rostrum;

curvedcrest: largewings: largelateral walls,with posterior margin deeply indented (Fig. 2b).

Lower beak has short, blunt rostrum; long, curvedcrest;largelateralwalls;largewings(Fig.

2c). Rostrum, hood, crest and lateral walls of both upperandlowerbeaks, heavilypigmented darkbrowntoblackincolour;margins ofwings.

hood,crestandlateralwallsofbothbeakstrans- parent. Radula typically octopodan, with 7 transverse rows of teeth and marginal plates (Figs 3c, d). Rhaehidian tooth has 1-2 lateral cuspsoneachsideoflargemedialcusp. Lateral cuspsinasymmetricalseriation.migratingfrom medialto lateral position over6-7 rows^(B(,_7 type). First lateralteethsmallandunicuspidatc:

secondlateralteethlongwithcurvedbase; third lateral teeth long and curved: marginal plates rectangularand plain.

Anteriorsalivaryglandssmall,borderingpos- terior buccal mass Posterior salivary glands stout anteriorly, tapering posteriorly, with 1

duet from each gland running forward then uniting toform single duct running alongside oesophagustobuccal mass.Cropwith anterior caecum ofabout 30% ofits length. Posterior oesophagus short. Stomach typicallybipartite.

Caecumwithasingleloosecoil.2separate ducts connectdigestivegland withcaecum. Intestine undifferentiated, although I coil occurs mid- way,butitisnotenlarged toformpouch. Inksac- large, lying superficially in groove on ventral faceofdigestivegland.Ashort,stoutductcon- nectsink sacwithdorsal side ofintestinenear anus.Anusbears a pairofanal Haps.

Testisposteriorinposition.Vasdeferenslong, delicate, tightlycoiled,enteringspermatophoral gland at proximal end. Spermatophoral gland swollen proximally. with muscular walls, but becomingthinwalledtowardsitsjunction with thelong accessory gland.Ashorttubeconnects accessoryglandandNecdham'ssac.Needham"s

THE OCTOPUSAUSTR.4USCOMPLEX 347

Figure1 0,v»/,afl„,(rafcHoyle:^a,dorsalviewofAMCI 56204M.67¹mm^ML(scalebap20mm):b.ventral vcwof mantleopemngand^tunnelolAMCI 56203.^F,507mm^^ff-lOMY^^wMM

Ffi55« F 454mmML^{(scalebar =5mm):d.}hectocotylisedarmofAM^C166899,40.5mmML^(scalebar-10

mm" ^{dorsal andf lateral,detailof}hec.ocotylusof'NMVF65533.68.7mmML^(scalebar-5mm).

348 T.N.STRANKSANDM.D.NORMAN

a

Figure_2.OclopmaustralisHoyle:a,digestive tractofNMVF65533,M.63.2mm ML(scalebar=₁₀mm)- b upperbeak,andc.lower beak, ofNMV F65533.M,68.7mm ML(scalebar -5mm) ^{'' '}

sac long, conical, pointed at apex. Penis long (PLI 16.6-/5.5-19.8), with a single coiled diverticulum. Genital aperture subterminal,on

rightsideof penis(Figs 4a-c).

Spermatophores relatively long (SpLI 626- 75.7-84.7), slender (SpWI 1.6-2.5-3.7) (Figs 4d-g). Oral cap simple, expanded, with_a long cap thread. Ejaculatory apparatus is a tightly coiled tube,which narrowsorally,with2-3coils close to the oral end. Small, bulbous cement bodyconnects with bothoralandaboralendsby narrow necks. Sperm reservoir spirally wound witharoundedaboral end; comprises approxi- matelyhalfofthespermatophore_length(SpRI 34.7-42_7-55.6); forms widest region of spermatophore.

Ovarylarge,ovoid, displacing adjacent organs when mature (Fig. 5a). Proximal oviducts initially common, then dividing _{into 2} long, curvedducts.Oviductsattach to spherical ovi- ductal glands,whicharedarkerincolour. Distal oviducts straight, tapering gradually. No

females wereobserving broodingeggs. Mature eggsfromfemales with enlarged ovaries(NMV

F65533: 54mm.62mm, 65mmML)are large (8-12 mm long; 1.5-2.0 mm wide), yellow, translucent,with eggstriation(EgLI 13.7-770- 21.4;EgWI2.6-3.2-4.4).Methodof eggattach- ment unknown.

Integumental sculptureconsistsofapattern of fine, rounded and closely set epidermal tubercles. The tubercles are largest and most denseonthedorsum; thoseonthe ventral sur- face are similarly sized but more scattered.

Unbranched papillae present in ocular region (Fig. 5b), with arowof ¹ largeand 3-4 small supraocularpapillae. Ventrolateral integumen- taryridgepresentonmantle;ridgeobvious and continuousaroundentiremantlecircumference, with theridgeforming ansharply angledpeak on theposteriormantle(Figs5c, d).

Colouroflive animals unknown. Colour of specimenspreservedinethanol uniformly light

browntopurpledorsally,creamtolight brown

ventrally. Ocelli absent.

Males mature at approximately 20-25 mm

ML. Femalesattain ovarian maturityat about 50-60 mm ML. The largest specimen studied

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Figure_3,Octopus Iwrrimasp.nov.:aandb.radulaofNMVF31264.M.71.4mmMI.OctopusaustralisHovlcc andd.radulaofNMVF65533. M,68.7 mmML.

wasafemaleof72mm MLfromoffNewcastle

NSW(AM C36577).

Distribution. Eastern Australia, from Hcrvey Bay, Queensland (about 25°S) to Jervis Bay,

NewSouth Wales(about 35°S)(Fig. 1

1 ).Bathv- metric records range from 3 to 134 m. The species is common in subtropical inshore waters, living on sand and mud bottom, and amongsponges.

Remarks. Hoyle (1885a)described O. australis based on specimens collected in eastern Aus- tralia duringthe cruise ofHMS "Challenger"

(1873-1876). Additional details appeared in a followingpaper(Hoyle,1885b).whichwasthen again expandedwith the inclusionofmeasure- ments and illustrations (Hoyle, 1886). Hoyle

(1886)providedsomedetailsofwhatisnow^des-

ignated to be the submature female lectotype, butdid not giveanydataontheimmaturemale paralectotypc.

Brazier(1892)laterrecordedO. australisfrom Georges Beach,NSW.Tothatdate, O. australis had been recorded fromtwolocalities inNSW.

Subsequently, the name O. australis was only employed for the morphologicallysimilar, but distinct, species from south-eastern Australia, herein named O. berrima(seeseparate entry)'

Fromthe 1890stothe present, the speciesnow

identifiedasoccurringinNSWandQueensland waterswasconsideredunnamed.

Robson ₍₁₉₂₉₎ and Tait (1982) had oppor- tunitiestoexamineindetailthe typespecimens of O. australis. and both authors continued usage ofthe name for the south-eastern Aus- tralian species. It was not until the present authors had beguna studyofthe eastern Aus- tralianoctopodid _fauna,andcomparativework carried outbetween easternandsouthern Aus- tralian octopuses, that the applicability ofthe

name O. australis to the eastern and not the southern_specieswasrealised.

Thepresentdeterminationand redescription

isbased on examination ofthetwo type speci- mens and31 otherspecimenlotsofO.australis, fromthe collectionsoftheAM. BMNH NMV

and QM.

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figure5.OctopusaustralisHoyle:a,female reproductive organs ofNMVF65533,54.3mmML(scalebar=10 mm);b,lateralview of unbranchedocularpapdlae(scalebar =10mm),c,lateralview,andd,posteriorview,of mantleandventrolateralintegumentaryridge(scalebar =20mm),ofNMVF65533,F,45.4mmML.

immaturemaleand submaturefemale syntypes ofO. australisfall largely within the rangesof variationfromthesample of10femalesand 10 malesfromeastern Australia(seeTable4).Any

significant discrepancies in morphometry between the type specimens and the larger sample(such aswiththeMAI,ALI, LLI, EgLI and EgWI indices) may be attributed to allo- metricvariablity.

Octopusaustraliscan be distinguishedfrom other speciesofthegenusonthe basisofthefol- lowingcharacters:abroadlyovoid mantle;skin with a pattern offine, rounded andclosely set tubercles onthedorsum, acontinuousventro- lateral ridge around the mantle, and a large

papillaover eacheye;small butprominenteyes;

long, subequal arms (2.7-4.5 times ML in

matureanimals);moderatelylarge suckers,with 4-5suckersslightlyenlargedonthelateralarms ofmalesonly; a medium sized, stout,bulbous ligula(8-17%ofthird rightarmlength), and a very short calamus;largeeggs(8-12mmlong), with unknown method of attachment to sub- strate;and7-9gilllamellae.

Thisspeciesiscommon in inshorewatersof eastern Australia,whereitisfrequentlytakenas acommercial by-catchbyprawntrawlers oper- ating in northern New South Wales and southern Queensland waters. Examination of catchesbyoneof us(TNS) showsthatthe species