13. norman...title 13. norman author nipol subject 13. norman created date 9/12/2005 8:56:37 am

Phuket mar. biol. Cent. Res. Bull. 66: 127–154 (2005)

THE CURRENT STATE OF OCTOPUS TAXONOMY

M.D. Norman1+2 and F.G. Hochberg3

1Sciences, Museum Victoria, GPO Box 666E, Melbourne, Vic 3001, Australia. ([email protected])2Zoology, University of Melbourne, Vic 3010, Australia.

3Department of Invertebrate Zoology, Santa Barbara Museum of Natural History, CA 93105, USA.

ABSTRACT: Much has changed since the 1988 CIAC sponsored workshop on octopus systematicsin Washington DC and the decisions reported in the resulting workshop publications. This paperprovides an update on the status of octopus taxonomy 17 years on and reports revisions, omissions,corrections and additions (new species). Information on 374 nominal species is presented, of whichwe consider around half (186) to represent valid names. This tally is considered a significantunderestimate of the total number of extant species. More than 150 undescribed species have beenrecognised in the past 17 years through examination of museum collections, primary field surveys anddiscovery of many cryptic species. The rate at which new species are being discovered is significantlyhigher than the rate at which new species are being formally described. This is largely due to a small,aging and diminishing cohort of octopod taxonomists. As these animals receive a higher profile infisheries, biodiversity and behavioural studies, the need for thorough and detailed taxonomy is higherthan ever before. The authors call for higher moral and financial support for this critical alpha taxonomy.

INTRODUCTION

In July 1988, the “International Workshop onSystematics and Biogeography of Cephalopods”was held in Washington DC. The workshop wassponsored by the Cephalopod InternationalAdvisory Council (CIAC). The primary objectivesof this workshop were to address the numeroustaxonomic problems associated with manycephalopod groups, particularly the cuttlefishes andoctopuses, and several speciose, commerciallyimportant families of squids. The results of thisworkshop were published in two volumes (Vosset al. 1998), with these works making aconsiderable contribution to the field.

For the benthic octopuses (FamilyOctopodidae), much has changed in the intervening15 years. Considerable research activity has beenfocused on octopus systematics, particularly in theIndo-West Pacific and Australian regions (Stranks,1988a–b, 1990; Gleadall, 1991a–b, 1993a–b, 1997,Gleadall and Naggs, 1991; Lu and Stranks, 1992;Norman, 1992a–c, 1993a–c, 1998, 2000, 2001;Stranks and Norman, 1993; Norman andHochberg, 1994; Allcock, 1997; Norman andSweeney, 1997; Norman et al. 1997; Lu, 1998;

Gonzalez et al., 1998; Hochberg, 1998;Nateewathana, 1997; Nateewathana and Norman,1999; O’Shea, 1999; Guerra et al., 2000; GuerreroKommritz, 2000; Norman and Lu, 2000; Normanand Finn, 2001; Allcock and Piertney, 2002; Muus,2002; Allcock et al., 2003).

The aim of this paper is to revise the species-level taxonomy of the family Octopodidae, updatingthe earlier works that resulted from the Washingtonworkshop. Through both primary field surveys andstudies of museum collections, many taxonomicproblems have been addressed and clarified. Anumber of nominal species of octopuses have beenrevised and numerous new species have beenrecognised. Revisions, omissions, corrections anda list of new taxa are presented. A generic levelrevision and updates on phylogenetic studies onthis group are currently in progress.

The past 17 years have been a productiveperiod for discovery of new species. More than150 new species are currently recognised fromthe Indo-West Pacific region alone (see citedreferences above, and Norman and Hochberg,unpubl. data). This more than doubles theestimated total number of octopus species in theworld. These new discoveries have resulted from

128

Phuket mar. biol. Cent. Res. Bull.

increased fieldwork, improved sampling techniquesand critical review of previously unexaminedpreserved specimens in museum collections. Majorsurveys have been undertaken in tropical,temperate and polar regions, in all habitats fromintertidal areas to the deep sea.

As the value and profile of octopusesincreases as fisheries targets, the need for rigoroustaxonomic knowledge is greater than ever before.Current impediments to progress on octopustaxonomy are discussed along with recognition offuture priorities.

Revision of taxonomic decisionsAs a result of this recent research activity,

many of the decisions made by the octopus workinggroup in 1988 now require updating. Table 1 presentsrevised designations for a number of species treatedin Toll and Voss (1998) and Toll (1998).

Two key species groups previously treatedwithin the genus Octopus have been sources ofconsiderable taxonomic confusion. Norman andFinn (2001) revised the “horridus species-group”sensu Norman (1992c), coining the subgenericname Abdopus to define these small long-armedoctopuses. Recent molecular studies of thephylogeny of octopuses of the family Octopodidae(Hudelot, 2000) have demonstrated that the genusOctopus is polyphyletic and that a major genericrevision of the family is required. We consider thatthe characters defining Abdopus are sufficient towarrant generic status and, as such, it is hereraised to generic level.

In Norman and Finn (2001) the status of manyhistorical names were reviewed. The maintaxonomic decisions were:

· Abdopus aculeatus d’Orbigny, 1834 isresurrected as a valid species (removed fromsynonymy of filamentosus Blainville, 1826 on thebasis of gill lamellae counts).

· The type series of Octopus aranead’Orbigny, 1834 consists of two species, thepoorest condition specimen also being the holotypeof filamentosus. At present, aranea is consideredunrecognisable and unresolved.

· Octopus argus Krauss, 1848 is resurrectedas a valid species (removed from the synonymy

of A. horridus d’Orbigny, 1826 on the basis ofspermatophore characters and the presence ofenlarged suckers).

· Octopus filamentosus Blainville, 1826 isconsidered unrecognisable (and invalid) as thesingle type specimen is in such poor condition.Until well-preserved material from the type locality(Mauritius) is obtained, the status of this speciesremains unresolved.

· Abdopus harmandi Rochebrune, 1882 isplaced in the synonymy of aculeatus (removedfrom synonymy of the unrecognisablefilamentosus, differing in gill lamellae counts).

· Toll and Voss (1998) placed Octopusniveus Lesson, 1830 in the synonymy of O.filamentosus at the same time as designating thetype of A. aculeatus as the neotype for O. niveus.In the absence of a type specimen, any additionalmaterial from the type locality and any validdiagnostic characters, niveus is considered asunrecognisable and unresolved. We do not considerToll and Voss’s designated neotype as valid.

Similar confusion surrounds members of the“aegina species-group” sensu Robson (1929) andNorman (1992c) (= the genus AmphioctopusFischer, 1882). A detailed discussion of key non-ocellate member taxa is provided in the Appendix.The primary taxonomic decisions are:

· Amphioctopus aegina Gray, 1849, A.kagoshimensis Ortmann, 1888 and A. marginatusTaki, 1964 are considered distinct and valid species.

· A. dollfusi Robson, 1928 and A.hardwickei Gray, 1849 are junior synonyms of A.aegina.

· A. striolatus Dong, 1976 is a juniorsynonym of A. marginatus.

In addition, it should be reinforced that A.areolatus de Haan in d’Orbigny, 1839–41, A.ocellatus Gray, 1849 and A. brocki Ortmann, 1888are all junior synonyms of A. fangsiao d’Orbigny,1839–41 (see Gleadall and Naggs, 1991).

Several taxa treated in O’Shea’s (1999)revision of New Zealand octopods requirecomment. Following examination of referencematerial from eastern Australia and New Zealand,we propose that Octopus gibbsi O’Shea, 1999 is asynonym of O. tetricus Gould, 1852, a common

129

The current state of Octopus taxonomy

Table 1. Revised designations to nominal species of the family Octopodidae.

Species 1988 Workshop decision Revised status Comments

INDIAN OCEANOctopus aranea junior synonym of Octopus unresolved see Norman and Finn, 2001Orbigny, 1834 filamentosus Blainville, 1826

Octopus argus junior synonym of Octopus valid species see Norman and Finn, 2001Krauss, 1848 horridus Orbigny, 1826

Octopus filamentosus valid unresolved see Norman and Finn, 2001Blainville, 1826

WEST PACIFICOctopus aculeatus junior synonym of Octopus valid species see Norman and Finn, 2001Orbigny, 1834 filamentosus Blainville, 1826 = Abdopus aculeatus

as well as being designatedneotype of another proposedsynonym, O. niveus Lesson, 1830

Octopus aegina nomen dubium valid species = Amphioctopus aegina seeGray, 1849 discussion on Amphioctopus

in Appendix

Callistoctopus arakawai uncertain junior synonym of junior synonym of see Norman 1993cTaki, 1964 Callistoctopus ornatus Gould, 1848 Callistoctopus ornatus

Gould, 1848

Octopus areolatus valid junior synonym of see Gleadall, 1991de Haan in Orbigny, Amphioctopus fangsiao1839–41 d’Orbigny, 1839–41

Octopus berenice nomen dubium valid species, based on Hochberg, unpubl. dataGray, 1849 additional material

linked to the type inthe British Museum

Octopus brocki valid junior synonym of see Gleadall, 1991Ortmann, 1888 Amphioctopus fangsiao

d’Orbigny, 1839–41

Octopus dollfusi junior synonym of junior synonym of see discussion onRobson, 1928 Octopus hardwickei Amphioctopus aegina Amphioctopus in

Gray, 1849 Gray, 1849 Appendix

Octopus hardwickei valid junior synonym of see discussion onGray, 1849 Amphioctopus aegina Amphioctopus in

Gray, 1849 Appendix

Octopus harmandi junior synonym of junior synonym of see Norman and Finn, 2001Rochebrune, 1882 Octopus filamentosus Abdopus aculeatusBlainville, 1826 d’Orbigny, 1834

Octopus hawiiensis nomen dubium valid species, missing Hochberg, unpubl. dataEydoux and Souleyet, type found in

1852 MNHN, Paris

Octopus marginatus nomen dubium valid species =Amphioctopus marginatusTaki, 1964 see discussion on

Amphioctopus in Appendix

Octopus niveus junior synonym of Octopus unresolved see Norman and Finn, 2001Lesson, 1830 filamentosus Blainville, 1826

Octopus striolatus valid junior synonym of see discussion onDong, 1976 Amphioctopus marginatus Amphioctopus in Appendix

Taki, 1964

130


species in New South Wales. In the absence oftype material, O’Shea resurrects Pinnoctopuscordiformis (Quoy and Gaimard, 1832) as thesenior synonym of Macroctopus maorum Hutton,1880. Quoy and Gaimard’s original description ofcordiformis refers to a large animal with arms ofalmost equal length, the lateral pairs being slightlyshorter. This arm formula does not match withthat of the species previously treated under thename maorum, which has an arm formula of1>2>3>4, the dorsal arms being obviously longerand more robust than the subsequent arm pairs.Quoy and Gaimard’s description more closelymatches Enteroctopus zealandicus, another largeoctopus found in the same general region. Thegenus Pinnoctopus was coined by Orbigny (1845)with the diagnostic character of a fin-like flaparound the margin of the mantle. We believe thatthis structure is an artefact of poor preservation,where loose skin settles in a flange around the

lateral mantle border. We have observed such astructure while preserving specimens ofEnteroctopus dofleini (Wülker, 1910) from thenorthern Pacific Ocean. This has not beenobserved while preserving specimens ofmaorum. This preservation artefact furthersupports the suggestion that Quoy andGaimard’s species is in fact a member of thegenus Enteroctopus (namely zealandicus). Inthe light of this confusion, the limited originaldescription and the absence of a type specimenfor cordiformis, we choose at this stage to leavethe name Pinnoctopus cordiformis as unresolved,making Macroctopus maorum the available namefor the distinctive species found in New Zealandand temperate Australian waters.

Table 2 presents the details and proposed status ofan additional eight historical species names omittedfrom the 1988 workshop publications.

Old species omitted Type locality Status Comments

Octopus appendiculatus (Indian Ocean) nomen nudum unsourced reference inDe. de Montford (date?) Blainville, 1826

Octopus cuvieri Pondicherry, India junior synonym of -Orbigny, 1826 Callistoctopus lechenaulti

Orbigny, 1840

Octopus cyanea var. Madras, India junior synonym of see Norman, 1992agracilis Robson, 1929 Octopus cyanea

Gray, 1849

Octopus dubius Bourbon Is. nomen dubium by Thomas, 1977Souleyet, 1852 (= Reunion Is.),

Indian Ocean

Eledonenta filholiana Fiji unresolved member of CallistoctopusRochebrune, 1884

Octopus hoylei var. annae Persian Gulf junior synonym of potential mixed lot:Robson, 1929 Pteroctopus hoylei paratypes from shallow

(Hoyle, 1909) water

Octopus lechenaulti Pondicherry, India unresolved member of CallistoctopusOrbigny, 1826

Callistoctopus magnocellatus Iyeno-Kushi, Japan junior synonym of see Norman, 1992aTaki, 1964 Octopus cyanea (not Callistoctopus)

Gray, 1849

Table 2. Nominal species omitted from Toll and Voss (1998) and Toll (1998).

131


Revisions to the “Classification of RecentCephalopoda” (Sweeney and Roper, 1998)

The following octopus taxa have beendescribed since the 1988 workshop and should beadded to Sweeney and Roper’s (1998)Classification of Recent Cephalopoda:· Abdopus abaculus Norman and Sweeney, 1997· Abdopus capricornicus Norman and Finn, 2001· Adelieledone piatkowski Allcock et al., (2003)· Amphioctopus neglectus Nateewathana and

Norman, 1999· A. rex Nateewathana and Norman, 1999· A. siamensis Nateewathana and Norman, 1999· Callistoctopus nocturnus Norman and Sweeney,

1997· Bathypolypus pugniger Muus, 2002· Benthoctopus clyderoperi O’Shea, 1999· B. karubar Norman, Hochberg and Lu, 1997· B. tangaroa O’Shea, 1999· B. tegginmathae O’Shea, 1999· Galeoctopus lateralis Norman, Boucher and

Hochberg, 2004· Graneledone gonzalezi Guerra, Gonzales and

Cherel, 2000· G. taniwha taniwha O’Shea, 1999· G. taniwha kubodera O’Shea, 1999· G. yamana Guerrero Kommritz, 2000· Microeledone mangoldi Norman, Hochberg and

Boucher-Rodoni, 2004· Octopus bulbus Norman, 2001· O. harpedon Norman, 2001· O. kaharoa O’Shea, 1999· O. mernoo O’Shea, 1999· O. micros Norman, 2001· O. pumilus Norman and Sweeney, 1997· O. pyrum Norman, Hochberg and Lu, 1997· Thaumeledone marshalli O’Shea, 1999· T. zeiss O’Shea, 1999· Vulcanoctopus hydrothermalis Gonzalez et al., 1998

Table 3 is an alphabetical list of nominal species inthe family Octopodidae. Generic placements andtaxonomic status for each name are presented. Anumber of species previously placed in the genusOctopus are treated here under recently resurrectedgeneric names. These include: AmphioctopusFischer, 1882; Callistoctopus Taki, 1964;Enteroctopus Rochebrune and Mabille, 1889;

Macroctopus Robson, 1928; Macrotritopus Grimpe,1922 and Paroctopus Naef, 1923. Taxa designatedas “unplaced” are currently treated under the genusOctopus until a major generic-level revision of thefamily Octopodidae is undertakenjects of a majorgeneric revision in progress by the authors.

Potential species names are also identified fora number of regional populations of octopusescurrently treated under the catchall names“vulgaris”, “macropus” and “defillipi”. Thesenames are available should current studiesdemonstrate that any of these populations warrantdistinct species status.

A total of 374 species names are listed forthis family (Table 3) of which 186 we consider tobe valid. A total of 102 names are consideredsynonyms of other species. Five names areconsidered to lack any form of description ortype material (nomina nuda), while 23 namesare considered dubious (nomina dubia) due toproblems with type material (poor, lost or nonedesignated), type locality (il l-defined,indecipherable or none designated) and/orinsufficient original descriptions to allow speciesrecognition. Four names were pre-occupied ofwhich three were assigned replacement names.The remaining 55 names are consideredunresolved and await further investigation.

We consider the tally of valid species to be aconsiderable underestimate of the total number ofextant species in this family. We have amassedobservations, museum records, data anddescriptions in progress for more than 150additional unnamed species within this family. It isclear that there is a considerably higher diversityof octopuses than previously suspected. Currentlythe rate at which new species are being discoveredgreatly exceeds the rate at which species are beingformally described.

Hidden diversityRegional surveys

The bulk of new species discoveries arecoming from recent regional reviews of octopodfaunas. Nateewathana (1997) reported 11 speciesof benthic octopuses from the waters surroundingThailand of which five were new to science. Lu(1998) reported 22 species from Taiwanese waters

132


Tabl

e 3.

Alp

habe

tical

list

ing

of n

omin

al s

peci

es o

f the

Fam

ily O

ctop

odid

ae

Val

id s

peci

es in

bol

d. *

= ty

pe s

peci

es o

f re

cogn

ised

gen

era.

Aut

hori

ties

for

gene

ra: A

bdop

us N

orm

an a

nd F

inn,

200

1; A

deli

eled

one

Allc

ock

et a

l. (2

003)

; Am

eloc

topu

s N

orm

an, 1

992;

Am

phio

ctop

us F

isch

er, 1

882;

Bat

hypo

lypu

sG

rim

pe, 1

921a

; Ben

thel

edon

e R

obso

n, 1

932;

Ben

thoc

topu

s Gri

mpe

, 192

1; C

allis

toct

opus

Tak

i, 19

64; C

isto

pus

Gra

y, 1

849;

Ele

done

Lea

ch, 1

817;

Ent

eroc

topu

s R

oche

brun

ean

d M

abill

e, 1

889;

Eua

xoct

opus

Vos

s, 1

971;

Gal

eoct

opus

Nor

man

, Bou

cher

and

Hoc

hber

g., 2

004;

Gra

nele

done

Jou

bin,

191

8; G

rim

pell

a R

obso

n, 1

928;

Hap

aloc

hlae

naR

obso

n, 1

929;

Mac

roch

laen

a R

obso

n, 1

929;

Mac

roct

opus

Rob

son,

192

8; M

acro

trit

opus

Gri

mpe

, 192

2; M

egal

eled

one

Taki

, 196

1; O

ctop

us C

uvie

r, 17

97; M

icro

eled

one

Nor

man

, H

ochb

erg

and

Bou

cher

-Rod

oni,

2004

; P

arel

edon

e R

obso

n, 1

932;

Par

octo

pus

Nae

f, 1

923;

Pte

roct

opus

Fis

cher

, 18

82;

Rob

sone

lla

Ada

m,

1938

; Sc

aeur

gus

Tro

sche

l, 18

57; T

eret

octo

pus

Rob

son,

192

9; T

etra

chel

edon

e V

oss,

195

5; T

haum

eled

one

Rob

son,

193

0; V

elod

ona

Chu

n, 1

915;

Vos

sele

done

Pal

acio

, 197

8; V

ulca

noct

opus

Gon

zale

z et

al.,

199

8. Taxa

Gen

eric

pla

cem

ent

Stat

us

abac

ulus

(N

orm

an a

nd S

wee

ney,

199

7: 1

09)

Abd

opus

valid

abru

ptus

(Sas

aki,

1920

: 173

)B

enth

octo

pus

valid

acul

eatu

s (d

’Orb

igny

, 183

4: p

ls 7

, 8)

Abd

opus

valid

syno

nym

: har

man

di R

oche

brun

e, 1

882:

73

adam

si B

enha

m, 1

944:

259

unpl

aced

syno

nym

of

hutt

oni

*ade

liean

a (B

erry

, 191

7: 1

7)A

deli

eled

one

valid

aegi

na (

Gra

y, 1

849:

7)

Am

phio

ctop

usva

lidsy

nony

ms:

dol

lfus

i Rob

son,

192

8: 4

3ha

rdw

icke

i Gra

y, 1

849:

8sm

edle

yi R

obso

n, 1

932:

24

alat

us S

asak

i, 19

20: 1

80un

plac

edun

reso

lved

albe

rti J

oubi

n, 1

895:

18

unpl

aced

juve

nile

: nom

en d

ubiu

mal

bida

(B

erry

, 191

7: 1

5)B

enth

eled

one

valid

albi

dus

(Tak

i, 19

62: 3

97)

Pte

roct

opus

syno

nym

of

hoyl

eial

bus

Raf

ines

que,

181

4: 2

9O

ctop

ussy

nony

m o

f vu

lgar

isal

cock

i Rob

son,

193

2: 2

51Te

reto

ctop

usva

lidal

deri

(Ver

any,

185

1: 3

2)C

allis

toct

opus

syno

nym

of

mac

ropu

sal

drov

andi

(M

ontf

ort,

1802

: 55

)E

ledo

nesy

nony

m o

f ci

rrho

saal

ecto

Ber

ry, 1

953:

56

unpl

aced

valid

alph

eus

(Nor

man

, 199

3: 2

70)

Cal

listo

ctop

usva

lidam

boin

ensi

s B

rock

, 188

7: 5

98un

plac

edno

men

dub

ium

amer

ican

us B

aker

in D

enys

de

Mon

tfor

t, 18

02: 3

8O

ctop

usun

reso

lved

pote

ntia

l nam

e fo

r Wes

t Atla

ntic

“vu

lgar

is”

syno

nym

s: b

aker

ii d

’Orb

igny

, 182

6: 1

44eu

dora

Gra

y, 1

849:

9ge

ryon

ea G

ray,

184

9: 7

anna

e (R

obso

n, 1

929:

220

)P

tero

ctop

ussy

nony

m o

f ho

ylei

anta

rctic

a (T

hiel

e, 1

920:

434

)P

arel

edon

eva

lidan

tarc

tica

Vos

s, 1

976:

448

Gra

nele

done

valid

133


anti

llar

um “

d’O

rbig

ny ?

?” in

Dal

l, 18

96: 2

7un

plac

edno

men

nud

umap

olly

oni (

Ber

ry, 1

912:

284

)E

nter

octo

pus

syno

nym

of

dofl

eini

appe

ndic

ulat

us B

lain

ville

, 182

6: 1

85un

plac

edno

men

nud

um*a

raka

wai

Tak

i, 19

64: 2

92C

allis

toct

opus

syno

nym

of

orna

tus

aran

ea d

’Orb

igny

, 183

4: p

l. 5

unpl

aced

unre

solv

edar

aneo

ides

Tak

i, 19

64: 3

07un

plac

edun

reso

lved

arbo

resc

ens

Hoy

le, 1

904:

189

unpl

aced

unre

solv

ed*a

rctic

us (P

rosc

h, 1

847:

59)

Bat

hypo

lypu

sva

lidsy

nony

ms:

faer

oens

is R

usse

ll, 1

909:

446

grön

land

icus

Dew

hurs

t in

Stee

nstr

up, 1

856:

17

sasa

kii R

obso

n, 1

927:

257

areo

latu

s (d

e H

aan,

183

9–18

41: 6

5)A

mph

ioct

opus

syno

nym

of

fang

siao

argu

s K

raus

s, 1

848:

132

unpl

aced

valid

aspe

r (A

kim

ushk

in, 1

963:

138

)E

nter

octo

pus

syno

nym

of

dofl

eini

aspi

loso

mat

is (N

orm

an, 1

993a

: 279

)C

allis

toct

opus

valid

aust

ralis

Hoy

le, 1

885:

224

unpl

aced

valid

bair

dii (

Ver

rill,

187

3: 5

)B

athy

poly

pus

valid

syno

nym

s: le

ntus

Ver

rill,

188

0: 1

38ob

esus

Ver

rill,

188

0: 1

37pr

osch

i M

uus,

196

2: 1

3ba

keri

i d’O

rbig

ny, 1

826:

144

Oct

opus

syno

nym

of

amer

ican

us(i

f lat

ter i

s va

lid n

ame

for W

. Atla

ntic

“vu

lgar

is”)

balb

oai V

oss,

197

1: 1

6un

plac

edva

lidba

nden

sis

Hoy

le, 1

885:

227

unpl

aced

juve

nile

: nom

en d

ubiu

mbe

reni

ce G

ray,

184

9: 1

1un

plac

edva

lidbe

rmud

ensi

s (H

oyle

, 188

5: 2

28)

Cal

listo

ctop

ussy

nony

m o

f fu

rvus

(if

latte

r is

val

id n

ame

for W

. Atla

ntic

“m

acro

pus”

)be

rrim

a St

rank

s an

d N

orm

an, 1

993:

355

unpl

aced

valid

berr

yi R

obso

n, 1

924:

658

Ben

thoc

topu

sva

lidbi

mac

ulat

us V

erri

ll, 1

883b

: 121

Oct

opus

valid

bim

acul

oide

s Pi

ckfo

rd a

nd M

cCon

naug

hey,

194

9: 1

4O

ctop

usva

lidbi

tent

acul

atus

Ris

so, 1

854:

61

Oct

opus

syno

nym

of

vulg

aris

bock

i Ada

m, 1

941:

1un

plac

edva

lidbo

reop

acif

ica

Nes

is, 1

982:

322

Gra

nele

done

valid

syno

nym

: pac

ific

a V

oss

and

Pear

cy, 1

990

bosc

ii L

esue

ur, 1

821:

110

unpl

aced

nom

en n

udum

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

134

Phuket mar. biol. Cent. Res. Bull.br

evip

es d

’Orb

igny

, 183

5: 2

2un

plac

edju

veni

le: n

omen

dub

ium

*bre

vis

(Hoy

le, 1

885:

230

)T

haum

eled

one

valid

brev

itent

acul

atus

Bla

invi

lle, 1

826:

187

Oct

opus

syno

nym

of

vulg

aris

bria

reus

Rob

son,

192

9: 6

12un

plac

edva

lidbr

ocki

(O

rtm

ann,

188

8: 6

45)

Am

phio

ctop

ussy

nony

m o

f fa

ngsi

aobu

lbus

Nor

man

, 200

1: 6

80un

plac

edva

lidbu

nuro

ng (

Stra

nks,

199

0: 4

62)

Cal

listo

ctop

usva

lidbu

rryi

(Vos

s, 1

950:

76)

Am

phio

ctop

usva

lidsy

nony

ms:

vin

cent

i Pic

kfor

d, 1

955:

159

burs

ariu

s St

eens

trup

in

Hoy

le, 1

886:

14

Cis

topu

ssy

nony

m o

f in

dicu

sca

erul

esce

ns B

lain

ville

, 182

6: 1

89un

plac

edno

men

dub

ium

calif

orni

cus

Ber

ry, 1

911:

590

unpl

aced

valid

cam

pbel

li Sm

ith, 1

902:

201

unpl

aced

valid

cant

hylu

s V

oss

and

Pear

cy, 1

990:

69

Ben

thoc

topu

sva

lidca

part

i Ada

m, 1

950:

7E

ledo

neva

lidca

pens

is E

ydou

x an

d So

uley

et, 1

852:

11

unpl

aced

juve

nile

: nom

en d

ubiu

mca

pric

orni

cus

Nor

man

and

Fin

n, 2

001:

24

Abd

opus

valid

carl

gren

i Tho

re 1

945:

52

Ele

done

syno

nym

of

schu

ltze

ica

roli

nens

is (V

erri

ll, 1

884:

235

)A

mph

ioct

opus

unre

solv

edca

ssio

peia

Gra

y, 1

849:

9O

ctop

ussy

nony

m o

f vu

lgar

isce

phea

Gra

y, 1

849:

15

unpl

aced

unre

solv

edch

alle

nger

i (B

erry

, 191

6: 4

9)G

rane

ledo

neva

lid*c

harc

oti (

Joub

in, 1

905:

22)

Par

eled

one

valid

*cha

rrua

Pal

acio

, 197

8: 2

82Vo

ssel

edon

eva

lidch

ierc

hiae

Jat

ta, 1

889:

64

unpl

aced

valid

chro

mat

us (

Hei

lpri

n, 1

888:

324

)C

allis

toct

opus

syno

nym

of

furv

us(i

f la

tter

is v

alid

nam

e fo

r W. A

tlant

ic “

mac

ropu

s”)

chun

i Gri

mpe

, 192

2un

plac

edun

reso

lved

cilia

tus

Ran

g, 1

837:

65

unpl

aced

unre

solv

edci

rrho

sa (L

amar

ck, 1

798:

130

)E

ledo

neva

lidsy

nony

ms:

ald

rova

ndi

Mon

tfor

t, 18

02:

55oc

topo

dia

Lin

ne, 1

758:

658

(as

Sepi

a)pe

nnan

ti a

utho

r un

know

n, in

Hoy

le, 1

909:

296

sepi

oide

a R

oche

brun

e, 1

884:

156

clyd

erop

eri O

’She

a, 1

999:

209

Ben

thoc

topu

sva

lidco

cco

(Ver

any,

184

6: 1

09)

Pte

roct

opus

syno

nym

of

tetr

acir

rhus

coer

ules

cent

es F

ra P

iero

, 189

5: 2

67O

ctop

ussy

nony

m o

f vu

lgar

is

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

135


colo

ssus

Gra

y, 1

849:

19

unpl

aced

apoc

ryph

al:

nom

en n

udum

com

mun

is (

Park

, 188

5: 1

98)

Mac

roct

opus

syno

nym

of

mao

rum

coni

spad

iceu

s Sa

saki

, 191

7: 3

67un

plac

edva

lidco

rdif

orm

is Q

uoy

and

Gai

mar

d, 1

832:

87

unpl

aced

unre

solv

edse

e di

scus

sion

corn

utus

Ow

en, 1

881:

131

unpl

aced

nom

en d

ubiu

mcu

vier

i (d’

Orb

igny

, 182

6: 1

8)C

allis

toct

opus

syno

nym

of l

eche

naul

tii

cyan

ea G

ray,

184

9: 1

5O

ctop

usva

lidsy

nony

ms:

gla

ber

Rüp

pell

in W

ülke

r, 1

920:

51

grac

ilis

Rob

son,

192

9: 9

8he

rdm

ani H

oyle

, 190

4: 1

87ho

rsti

Jou

bin,

189

8: 2

3m

agno

cell

atus

Tak

i, 19

64: 2

98m

arm

orat

us H

oyle

188

5: 2

27da

na R

obso

n, 1

929:

140

Mac

rotr

itop

ussy

nony

m o

f de

fill

ipi

defi

lippi

(V

eran

y, 1

851:

30)

Mac

rotr

itop

usva

lidsy

nony

ms:

dan

a R

obso

n, 1

929:

140

kem

pi R

obso

n, 1

929:

312

Unr

esol

ved

Wes

t Atla

ntic

for

m:

Ava

ilabl

e na

me:

sco

rpio

, Ber

ry, 1

920:

299

didy

nam

us (R

afin

esqu

e, 1

814:

28)

Cal

listo

ctop

ussy

nony

m o

f m

acro

pus

dier

ythr

aeus

(Nor

man

, 199

3: 2

84)

Cal

listo

ctop

usva

lid*d

igue

ti Pe

rrie

r an

d R

oche

brun

e, 1

894:

770

Par

octo

pus

valid

dofl

eini

(W

ulke

r, 1

910:

7)

Ent

eroc

topu

sva

lidsy

nony

ms:

apo

llyo

ni B

erry

, 191

2: 2

84as

per

Aki

mus

hkin

, 196

3: 1

38gi

lber

tian

us B

erry

, 191

2: 2

84pu

ncta

tus

Gab

b, 1

862:

170

doll

fusi

(Rob

son,

192

8: 4

3)A

mph

ioct

opus

syno

nym

of

aegi

nadu

plex

Hoy

le, 1

885:

226

unpl

aced

syno

nym

of

supe

rcil

iosu

sel

egan

s B

rock

, 188

7: 5

97un

plac

edno

men

dub

ium

*equ

ivoc

us (

Rob

son,

192

9: 1

69)

Mac

rotr

itop

usju

veni

le: u

nres

olve

dsy

nony

m: g

raci

lis

Ver

rill,

188

4: 2

36er

gast

icus

(Fis

cher

and

Fis

cher

, 189

2: 2

98)

Bat

hypo

lypu

sva

lidsy

nony

ms:

loth

ei (

Chu

n, 1

913:

18)

prof

undi

cola

(M

assy

, 190

7: 3

77)

eudo

ra G

ray,

184

9: 9

Oct

opus

syno

nym

of

amer

ican

us(i

f lat

ter i

s va

lid n

ame

for W

. Atla

ntic

“vu

lgar

is”)

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

136

Phuket mar. biol. Cent. Res. Bull.eu

reka

(R

obso

n, 1

929:

179

)B

enth

octo

pus

syno

nym

of

hyad

esi

eury

ceph

ala

(Tak

i, 19

64: 3

13)

Pte

roct

opus

unre

solv

edex

annu

latu

s (N

orm

an, 1

993:

321

)A

mph

ioct

opus

valid

faer

oens

is (R

usse

ll, 1

909:

446

)B

athy

poly

pus

syno

nym

of

arct

icus

fang

siao

(d’

Orb

igny

, 183

9–18

41: 7

0)A

mph

ioct

opus

valid

syno

nym

s: a

reol

atus

de

Haa

n, 1

839–

1841

: 65

broc

ki O

rtm

ann,

188

8: 6

45fa

ngsi

ao e

tchu

anus

Sas

aki,

1929

: 53

fang

siao

typi

cus

d’O

rbig

ny, 1

839–

1841

: 70

ocel

latu

s G

ray,

184

9: 1

5fa

ngsi

ao e

tchu

anus

(Sas

aki,

1929

: 53)

Am

phio

ctop

ussy

nony

m o

f fa

ngsi

aofa

ngsi

ao ty

picu

s (d

’Orb

igny

, 183

9–18

41: 7

0)A

mph

ioct

opus

syno

nym

of

fang

siao

fasc

iata

(H

oyle

, 188

6: 9

4)H

apal

ochl

aena

valid

syno

nym

: pic

tus

Bro

ck, 1

882:

603

robu

stus

Bro

ck, 1

887:

317

favo

nius

Gra

y, 1

849:

9un

plac

edva

lidfi

lam

ento

sus

Bla

invi

lle, 1

826:

188

unpl

aced

unre

solv

edfi

lhol

iana

Roc

hebr

une,

188

4: 1

57un

plac

edun

reso

lved

filo

sus

How

ell,

1867

: 240

unpl

aced

valid

syno

nym

: hum

mel

inck

i Ada

m, 1

936:

1fi

mbr

iatu

s (R

üppe

ll in

d’O

rbig

ny, 1

841:

64)

Abd

opus

syno

nym

of

horr

idus

fitc

hi B

erry

, 195

3: 5

4un

plac

edva

lidfl

inde

rsi (

Cot

ton,

193

2: 5

43)

Mac

roct

opus

syno

nym

of

mao

rum

font

ania

na a

fric

ana

Rob

son,

192

9: 1

89un

plac

edun

reso

lved

*fon

tani

anus

(d’

Orb

igny

, 183

4: 2

8)R

obso

nella

valid

fram

ensi

s L

u an

d St

rank

s, 1

994:

227

Par

eled

one

valid

fray

edus

(Raf

ines

que,

181

4: 2

8)C

allis

toct

opus

syno

nym

of

mac

ropu

sfu

jita

i Sas

aki,

1929

: 70

unpl

aced

unre

solv

edfu

rvus

(G

ould

, 185

2: 4

75)

Cal

listo

ctop

usun

reso

lved

pote

ntia

l nam

e fo

r W

est A

tlant

ic “

mac

ropu

s”sy

nony

ms:

ber

mud

ensi

s H

oyle

, 188

5: 2

28ch

rom

atus

Hei

lpri

n, 1

888:

324

fusc

us T

aki,

1964

: 316

Ben

thoc

topu

sva

lidfu

sifo

rmis

Bro

ck, 1

887:

601

unpl

aced

nom

en d

ubiu

mga

rdin

eri H

oyle

, 190

5: 9

76un

plac

edva

lidga

ucha

Hai

mov

ici,

1988

: 82

Ele

done

valid

gery

onea

Gra

y, 1

849:

7O

ctop

ussy

nony

m o

f am

eric

anus

(if l

atte

r is

valid

nam

e fo

r W. A

tlant

ic “

vulg

aris

”)

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

137


gibb

si O

’She

a, 1

999:

120

Oct

opus

syno

nym

of

tetr

icus

giga

s O

ken,

183

5: 3

45un

plac

edap

ocry

phal

: no

men

nud

umgi

lber

tian

us (B

erry

, 191

2: 2

84)

Ent

eroc

topu

ssy

nony

m o

f do

flei

nigl

aber

(Sas

aki,

1920

: 172

)B

enth

octo

pus

syno

nym

of

hokk

aide

nsis

glab

er R

üppe

ll in

Wül

ker,

192

0: 5

1O

ctop

ussy

nony

m o

f cy

anea

glob

osus

App

ellö

f, 1

886:

7un

plac

edun

reso

lved

gonz

alez

i Gue

rra,

Gon

zale

s an

d C

here

l, 20

00G

rane

ledo

neva

lidgr

acil

is (V

erri

ll, 1

884:

236

)M

acro

trit

opus

pre-

occu

pied

nam

ere

plac

ed b

y eq

uivo

cus

grac

ilis

Rob

son,

192

9: 9

8O

ctop

ussy

nony

m o

f cy

anea

gran

ulat

us (L

amar

ck, 1

799:

130

)A

mph

ioct

opus

unre

solv

edgr

aptu

s (N

orm

an, 1

993:

296

)C

allis

toct

opus

valid

grim

pei R

obso

n, 1

924:

208

Bat

hypo

lypu

ssy

nony

m o

f va

ldiv

iae

grön

land

icus

(D

ewhu

rst i

n St

eens

trup

, 185

6: 1

7)B

athy

poly

pus

syno

nym

of

arct

icus

gron

osus

(Bla

invi

lle, 1

826:

186

)C

allis

toct

opus

syno

nym

of

mac

ropu

sgu

angd

onge

nsis

(Don

g, 1

976:

213

)A

bdop

usun

reso

lved

gunt

eri R

obso

n, 1

930:

392

Tha

umel

edon

eva

lidha

rdw

icke

i (G

ray,

184

9: 8

)A

mph

ioct

opus

syno

nym

of

aegi

naha

rman

di (R

oche

brun

e, 1

882:

73)

Abd

opus

syno

nym

of

acul

eatu

sha

rped

on N

orm

an, 2

001:

677

unpl

aced

valid

harr

isso

ni (

Ber

ry, 1

917:

24)

Par

eled

one

valid

hatta

i Sas

aki,

1929

: 87

unpl

aced

valid

haw

iiens

is E

ydou

x an

d So

uley

et, 1

852:

9un

plac

edva

lidhe

rdm

ani H

oyle

, 190

4: 1

87O

ctop

ussy

nony

m o

f cy

anea

hete

ropu

s R

afin

esqu

e, 1

814:

28

Oct

opus

syno

nym

of

vulg

aris

hoek

i Jou

bin,

189

8un

plac

edun

reso

lved

hokk

aide

nsis

(B

erry

, 192

1: 3

52)

Ben

thoc

topu

sva

lidsy

nony

m: g

labe

r Sa

saki

, 192

0: 1

72ho

ngko

ngen

sis

Hoy

le, 1

885:

224

unpl

aced

unre

solv

edsy

nony

m: t

enui

cirr

us S

asak

i, 19

29: 7

8*h

orri

dus

(d’O

rbig

ny,1

826:

144

)A

bdop

usva

lidsy

nony

ms:

fim

bria

tus

Rüp

pell

in d

’Orb

igny

, 184

1: 6

4ho

rsti

Jou

bin,

189

8: 2

3O

ctop

ussy

nony

m o

f cy

anea

hoyl

ei (

Ber

ry, 1

909:

407

)P

tero

ctop

usva

lidsy

nony

ms:

alb

idus

Tak

i, 19

62: 3

97an

nae

Rob

son,

192

9: 2

20hu

bbso

rum

Ber

ry, 1

953:

53

Oct

opus

valid

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

138

Phuket mar. biol. Cent. Res. Bull.hu

mm

elin

cki A

dam

, 193

6: 1

unpl

aced

syno

nym

of

filo

sus

hutto

ni B

enha

m, 1

943:

53

unpl

aced

valid

syno

nym

: ada

msi

Ben

ham

, 194

4: 2

59hy

ades

i Roc

hebr

une

and

Mab

ille,

188

9: H

6un

plac

edva

lidsy

nony

ms:

eur

eka

Rob

son,

192

9: 1

79m

agel

lani

cus

Rob

son,

193

0a: 3

33hy

alin

us R

ang,

183

7: 6

6un

plac

edju

veni

le: u

nres

olve

d*h

ydro

ther

mal

is G

onza

lez

et a

l., 1

998:

172

Vulc

anoc

topu

sva

lidin

cert

us T

argi

oni-

Tozz

etti,

186

9: 5

89un

plac

edno

men

dub

ium

inco

nspi

cuou

s B

rock

, 188

7: 6

03un

plac

edun

reso

lved

*ind

icus

(R

app,

183

5: 2

4)C

isto

pus

valid

syno

nym

: bu

rsar

ius

Stee

nstr

up i

n H

oyle

, 188

6: 1

4*i

ndic

us R

obso

n, 1

929:

608

Tere

toct

opus

valid

janu

arii

(Hoy

le, 1

885:

229

)B

enth

octo

pus

valid

joub

ini

Rob

son,

192

9: 1

61un

plac

edva

lidka

gosh

imen

sis

(Ort

man

n, 1

888:

664

)A

mph

ioct

opus

valid

kaha

roa

O’S

hea,

199

9: 1

74un

plac

edva

lidka

ruba

r N

orm

an, H

ochb

erg

and

Lu,

199

7: 3

60B

enth

octo

pus

valid

kaur

na S

tran

ks, 1

990:

460

unpl

aced

valid

kem

pi R

obso

n, 1

929:

312

Mac

rotr

itop

ussy

nony

m o

f de

fill

ipi

kera

lens

is (

Oom

men

, 196

6: 5

6)P

tero

ctop

usun

reso

lved

*lat

eral

is N

orm

an, B

ouch

er a

nd H

ochb

erg,

200

4G

aleo

ctop

usva

lidke

rmad

ecen

sis

(Ber

ry, 1

914:

138

)C

allis

toct

opus

valid

lech

enau

ltii

(d’O

rbig

ny, 1

826:

18)

Cal

listo

ctop

usva

lidsy

nony

m: c

uvie

ri d

’Orb

igny

, 182

6: 1

8le

iode

rma

(Ber

ry, 1

911:

590

)B

enth

octo

pus

valid

lent

us (V

erri

ll, 1

880:

138

)B

athy

poly

pus

syno

nym

of

bair

dii

leuc

oder

ma

Sang

iova

nni,

1829

: 317

Ele

done

unre

solv

edle

vis

(Hoy

le, 1

885:

229

)B

enth

octo

pus

valid

*lito

ralis

Nor

man

, 199

2: 5

69A

mel

octo

pus

valid

lobe

nsis

Cas

tella

nos

and

Men

ni, 1

969:

92

Oct

opus

syno

nym

of

tehu

elch

uslo

ngim

anus

(d’

Orb

igny

, 184

0: 1

8)C

allis

toct

opus

syno

nym

of

mac

ropu

slo

ngip

es L

each

, 181

7: 1

39un

plac

edno

men

dub

ium

long

ispa

dice

us S

asak

i, 19

17: 3

66un

plac

edun

reso

lved

loth

ei (C

hun,

191

3: 1

8)B

athy

poly

pus

syno

nym

of

erga

stic

us*l

unul

ata

(Quo

y an

d G

aim

ard,

183

2: 8

6)H

apal

ochl

aena

valid

lute

us (S

asak

i, 19

29: 4

5)C

allis

toct

opus

valid

mac

hiki

i Bro

ck, 1

887:

599

unpl

aced

nom

en d

ubiu

m

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

139


mac

roph

allu

s V

oss

and

Pear

cy, 1

990:

82

Ben

thoc

topu

sva

lidm

acro

podu

s (S

an G

iova

nni,

1829

: 319

)C

allis

toct

opus

syno

nym

of

mac

ropu

sm

acro

pus

(Ris

so, 1

826:

3)

Cal

listo

ctop

usva

lidsy

nony

ms:

ald

eri V

eran

y, 1

851:

32

didy

nam

us R

afin

esqu

e, 1

814:

28

fray

edus

Raf

ines

que,

181

4: 2

8gr

onos

us B

lain

ville

, 182

6: 1

86lo

ngim

anus

d’O

rbig

ny, 1

840:

18

mac

ropo

dus

San

Gio

vann

i, 18

29: 3

19ru

ber

Can

trai

ne, 1

841:

18

Unr

esol

ved

Wes

t Atla

ntic

for

m:

Ava

ilabl

e na

me:

furv

us G

ould

, 185

2: 4

75sy

nony

ms:

ber

mud

ensi

s H

oyle

, 188

5: 2

28ch

rom

atus

Hei

lpri

n, 1

888:

324

mac

roty

la V

oss,

197

6: 4

54G

rane

ledo

neva

lidm

acul

atus

Raf

ines

que,

181

4: 2

9O

ctop

ussy

nony

m o

f vu

lgar

ism

acul

osa

(Hoy

le, 1

883:

319

)H

apal

ochl

aena

valid

mad

okai

Ber

ry, 1

921:

352

unpl

aced

unre

solv

edre

plac

emen

t nam

e fo

r pre

occu

pied

pus

tulo

sus

Sasa

ki, 1

920:

176

mag

ella

nicu

s R

obso

n, 1

930:

333

Ben

thoc

topu

ssy

nony

m o

f hy

ades

im

agni

ficu

s (V

illan

ueva

, San

chez

and

Com

pagn

o, 1

992:

39)

Ent

eroc

topu

sva

lidm

agno

cell

atus

Tak

i, 19

64: 2

98O

ctop

ussy

nony

m o

f cy

anea

*man

gold

i Nor

man

, Hoc

hber

g an

d B

ouch

er-R

odon

i, 20

04*m

aoru

m (

Hut

ton,

188

0: 1

)M

acro

ctop

usva

lidsy

nony

ms:

com

mun

is P

ark,

188

5: 1

98fl

inde

rsi C

otto

n, 1

932:

543

mar

gina

tus

(Tak

i, 19

64: 3

04)

Am

phio

ctop

usva

lidsy

nony

m: s

trio

latu

s D

ong,

197

6: 2

12m

arin

um K

oelr

eut (

date

unk

now

n)O

ctop

ussy

nony

m o

f vu

lgar

ism

arm

orat

us H

oyle

188

5: 2

27O

ctop

ussy

nony

m o

f cy

anea

mar

shal

li O

’She

a, 1

999:

249

Tha

umel

edon

eva

lidm

assy

ae V

oss,

196

4: 5

11E

ledo

neva

lidm

aya

Vos

s an

d So

lis, 1

966:

617

Oct

opus

valid

med

oria

Gra

y, 1

849:

14

unpl

aced

unre

solv

edm

egal

ocya

thus

(Gou

ld, 1

852:

471

)E

nter

octo

pus

valid

syno

nym

s: b

ruce

i Hoy

le, 1

912:

276

jutt

ingi

Rob

son,

192

9b: 6

16m

embr

anac

eus

Roc

hebr

une

and

Mab

ille,

1889

:H7

Taxa

Gen

eric

pla

cem

ent

Stat

usTa

ble

3. (c

ontin

ued)

140


pa

tago

nicu

s L

önnb

erg,

190

7: 5

0

pe

nthe

rinu

s R

oche

brun

e an

d M

abill

e, 1

889:

H7

meg

alop

s Ta

ki, 1

964:

310

unpl

aced

unre

solv

ed*m

embr

anac

eus

(Quo

y an

d G

aim

ard,

183

2: 8

9)A

mph

ioct

opus

nom

en d

ubiu

m*m

embr

anac

eus

Roc

hebr

une

and

Mab

ille,

1889

:H7

Ent

eroc

topu

ssy

nony

m o

f m

egal

ocya

thus

mer

cato

ris A

dam

, 193

7: 7

6un

plac

edva

lidm

erno

o O

’She

a, 1

999:

165

unpl

aced

valid

mic

roph

thal

mus

Goo

dric

h, 1

896:

20

unpl

aced

valid

mic

ropy

rsus

Ber

ry, 1

953:

52

unpl

aced

valid

mic

ros

Nor

man

, 200

1: 6

83un

plac

edva

lidm

icro

sicy

a (R

oche

brun

e, 1

884:

158

)E

ledo

nesy

nony

m o

f m

osch

ata

mim

us G

ould

, 185

2: 4

73O

ctop

usva

lidm

inor

min

or S

asak

i, 19

20: 1

81un

plac

edun

reso

lved

min

or p

arda

lis

Sasa

ki, 1

929:

94

unpl

aced

unre

solv

edm

inor

typi

cus

Sasa

ki, 1

929:

90

unpl

aced

unre

solv

edm

olli

s G

ould

, 185

2un

plac

edno

men

dub

ium

mon

tero

sato

i Fra

Pie

ro, 1

895:

268

unpl

aced

unre

solv

edm

onte

vide

o R

ang,

183

7: 6

2un

plac

edno

men

dub

ium

mos

chat

a (L

amar

ck, 1

798:

130

)E

ledo

neva

lidsy

nony

ms:

mic

rosi

cya

Roc

hebr

une,

188

4c: 1

58m

osch

atus

Raf

ines

que,

181

4: 2

8O

ctop

ussy

nony

m o

f vu

lgar

ism

osch

ites

Car

us, 1

824:

319

Ele

done

unre

solv

edm

otot

i (N

orm

an, 1

993:

329

)A

mph

ioct

opus

valid

mut

ilans

Tak

i, 19

42: 7

1un

plac

edva

lidna

nhai

ensi

s D

ong,

197

6: 2

11un

plac

edun

reso

lved

nanu

s Ada

m, 1

973:

42

unpl

aced

valid

negl

ectu

s (N

atee

wat

hana

and

Nor

man

, 199

9: 4

52)

Am

phio

ctop

usva

lidni

erst

rasz

i (A

dam

, 193

8: 1

4)H

apal

ochl

aena

valid

nige

r R

afin

esqu

e, 1

814:

29

Oct

opus

syno

nym

of

vulg

aris

nigr

a (H

oyle

, 191

0: 2

62)

Ele

done

syno

nym

of

schu

ltze

ini

veus

Les

son,

183

0: 2

39un

plac

edun

reso

lved

noct

urnu

s (N

orm

an a

nd S

wee

ney,

199

7: 1

17)

Cal

listo

ctop

usva

lidno

rman

i (M

assy

, 19

07:

379)

Ben

thoc

topu

sva

lidob

esus

(Ver

rill,

188

0: 1

37)

Bat

hypo

lypu

ssy

nony

m o

f ba

irdi

ioc

cide

ntal

is S

teen

stru

p in

Hoy

le, 1

885:

77

Oct

opus

valid

ocel

latu

s (G

ray,

184

9: 1

5)A

mph

ioct

opus

syno

nym

of

fang

siao

ocho

tens

is S

asak

i, 19

20: 1

74un

plac

edun

reso

lved

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

141


octo

podi

a L

inne

, 175

8: 6

58 (a

s Se

pia

octo

podi

a)E

ledo

nesy

nony

m o

f ci

rrho

saoc

topo

dia

Try

on, 1

879:

113

Oct

opus

syno

nym

of

vulg

aris

ocul

ifer

Hoy

le, 1

904:

14

Oct

opus

valid

syno

nym

: roo

seve

lti S

tuar

t, 19

41: 1

oliv

eri B

erry

, 191

4: 1

36un

plac

edva

lidor

egon

ae T

oll,

1981

: 88

Ben

thoc

topu

sva

lidor

egon

ensi

s V

oss

and

Pear

cy, 1

990:

73

Ben

thoc

topu

sva

lidor

natu

s (G

ould

, 185

2: 4

76)

Cal

listo

ctop

usva

lidsy

nony

m:

arak

awai

Tak

i, 19

64:

292

oshi

mai

Sas

aki,

1929

: 44

unpl

aced

unre

solv

edov

ulum

(Sas

aki,

1917

: 364

)A

mph

ioct

opus

valid

paci

fica

Vos

s an

d Pe

arcy

, 199

0: 8

5G

rane

ledo

nesy

nony

m o

f bo

reop

acif

ica

pala

ri L

u an

d St

rank

s, 1

992:

73

unpl

aced

valid

palli

dus

Hoy

le, 1

885:

223

unpl

aced

valid

*pan

amen

sis

Vos

s, 1

971:

25

Eua

xoct

opus

valid

parv

us S

asak

i, 19

17: 3

65un

plac

edva

lidpa

tagi

atus

Ber

ry, 1

913:

564

Scae

urgu

sva

lidpe

nici

llife

r B

erry

, 19

54:

66un

plac

edva

lidpe

nnan

ti a

utho

r un

know

n, in

Hoy

le, 1

909:

296

Ele

done

syno

nym

of

cirr

hosa

pero

nii L

esue

ur, 1

821:

101

unpl

aced

unre

solv

edpi

atko

wsk

i Allc

ock,

Hoc

hber

g, R

odho

use

and

Tho

rpe

(200

3)A

delie

ledo

neva

lid

pict

us (B

rock

, 188

2: 6

03)

Hap

aloc

hlae

nana

me

pre-

occu

pied

syno

nym

of

fasc

iata

pict

us B

lain

ville

, 182

6: 8

unpl

aced

nom

en d

ubiu

mpi

ctus

Ver

rill,

188

3: 1

12un

plac

edna

me

pre-

occu

pied

repl

aced

by

verr

illi

ver

rill

ipi

llsbu

ryae

Vos

s, 1

975:

346

Eua

xoct

opus

valid

pilo

sus

Ris

so, 1

826:

4O

ctop

ussy

nony

m o

f vu

lgar

is*p

isca

toru

m (V

erri

ll, 1

879:

470

)B

enth

octo

pus

syno

nym

of

Bat

hypo

lypu

s ba

irdi

i (s

ee M

uus,

200

2)A

s a

cons

eque

nce,

gen

us B

enth

octo

pus

requ

ires

revi

sion

poly

mor

pha

(Rob

son,

193

0: 3

90)

Ade

liel

edon

eva

lidpo

lyze

nia

(Gra

y, 1

849:

13)

Am

phio

ctop

usva

lidpr

asha

di A

dam

, 193

9: 1

03un

plac

edun

reso

lved

pric

ei B

erry

, 191

3: 7

3un

plac

edun

reso

lved

prof

undi

cola

(M

assy

, 190

7: 3

77)

Bat

hypo

lypu

ssy

nony

m o

f er

gast

icus

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

142

Phuket mar. biol. Cent. Res. Bull.pr

ofun

doru

m R

obso

n, 1

932:

237

Ben

thoc

topu

sva

lidpr

osch

i M

uus,

196

2: 1

3B

athy

poly

pus

syno

nym

of

bair

dii

pryz

dens

is L

u an

d St

rank

s, 1

994:

232

Par

eled

one

valid

pseu

dony

mus

(Gri

mpe

, 192

2: 4

1)B

enth

octo

pus

valid

pugn

iger

Muu

s, 2

002:

195

Bat

hypo

lypu

sva

lidpu

lche

r (B

rock

, 188

7: 6

07)

Am

phio

ctop

usju

veni

le: n

omen

dub

ium

pum

ilus

Nor

man

and

Sw

eene

y, 1

997:

127

unpl

aced

valid

punc

tatu

s (G

abb,

186

2: 1

70)

Ent

eroc

topu

ssy

nony

m o

f do

flei

nipu

sill

us G

ould

, 185

2: 4

78un

plac

edno

men

dub

ium

pust

ulos

us B

lain

ville

, 182

6: 1

86un

plac

edno

men

dub

ium

pust

ulos

us S

asak

i, 19

20: 1

76un

plac

edpr

e-oc

cupi

ed n

ame,

repl

aced

by

mad

okai

pyru

m N

orm

an e

t al.,

199

7: 3

65un

plac

edva

lidra

bass

in R

isso

, 185

4: 6

7O

ctop

ussy

nony

m o

f vu

lgar

isra

panu

i (V

oss,

197

9: 3

60)

Cal

listo

ctop

usva

lidre

x (N

atee

wat

hana

and

Nor

man

, 199

9: 4

47)

Am

phio

ctop

usva

lidro

bson

i (A

dam

, 194

1: 1

)A

mph

ioct

opus

valid

robu

stus

(B

rock

, 188

7: 3

17)

Hap

aloc

hlae

nasy

nony

m o

f fa

scia

taro

bust

us V

oss

and

Pear

cy, 1

990:

67

Ben

thoc

topu

sva

lidro

osev

elti

Stu

art,

1941

: 1O

ctop

ussy

nony

m o

f oc

ulif

er*r

otun

da (

Hoy

le, 1

885:

230

)B

enth

eled

one

valid

rube

r (C

antr

aine

, 184

1: 1

8)C

allis

toct

opus

syno

nym

of

mac

ropu

sru

ber

Raf

ines

que,

181

4: 2

8O

ctop

ussy

nony

m o

f vu

lgar

isru

besc

ens

Ber

ry, 1

953:

51

unpl

aced

valid

rugo

sus

Bos

c 17

92: 2

4un

plac

edun

reso

lved

sale

bros

us S

asak

i, 19

20: 1

82un

plac

edva

lidsa

lutii

Ver

any,

183

9: 9

3un

plac

edva

lidsy

nony

m: s

aluz

zii N

aef,

192

3: 6

99sa

luzz

ii N

aef,

192

3: 6

99un

plac

edsy

nony

m o

f sa

luti

isa

ncta

ehel

enae

Rob

son,

192

9: 7

4O

ctop

usva

lidsa

phen

ia G

ray,

184

9: 1

1un

plac

edun

reso

lved

sasa

kii

Rob

son,

192

7: 2

57B

athy

poly

pus

syno

nym

of

arct

icus

sasa

kii T

aki,

1942

: 75

unpl

aced

unre

solv

edsc

alen

us (

Hoy

le, 1

904:

13)

Eua

xoct

opus

valid

schm

idti

(Jou

bin,

193

3: 4

)P

tero

ctop

usva

lidsc

hultz

ei (

Hoy

le, 1

910:

261

)E

ledo

neva

lid

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

143


syno

nym

: car

lgre

ni T

hore

194

5: 5

2ni

gra

Hoy

le, 1

910:

262

thys

anop

hora

Vos

s, 1

962d

: 265

scor

pio

(Ber

ry, 1

920:

299

)M

acro

trit

opus

unre

solv

edpo

tent

ial n

ame

for

Wes

t Atla

ntic

“de

filli

pi”

sele

ne V

oss,

197

1: 1

1un

plac

edva

lid*s

enoi

Tak

i, 19

61: 2

97M

egal

eled

one

syno

nym

of

sete

bos

sepi

oide

a R

oche

brun

e, 1

884:

156

Ele

done

syno

nym

of

cirr

hosa

sete

bos

Rob

son,

193

2: 3

13M

egal

eled

one

valid

syno

nym

: sen

oi T

aki,

1961

: 297

siam

ensi

s (N

atee

wat

hana

and

Nor

man

, 199

9: 4

56)

Am

phio

ctop

usva

lidsi

biri

cus

Loy

ning

, 193

0: 1

Ben

thoc

topu

sva

lidsi

nens

is d

’Orb

igny

, 183

4: p

l. 9

Oct

opus

unre

solv

edpo

tent

ial n

ame

for

Japa

nese

“vu

lgar

is”

smed

leyi

(Rob

son,

193

2: 2

4)A

mph

ioct

opus

syno

nym

of

aegi

na*s

pini

cirr

us V

oss,

195

5: 1

07Te

trac

hele

done

valid

spin

osus

Sas

aki,

1920

: 177

unpl

aced

unre

solv

edsp

onsa

lis (F

isch

er a

nd F

isch

er, 1

892:

297

)B

athy

poly

pus

valid

stri

olat

us (D

ong,

197

6: 2

12)

Am

phio

ctop

ussy

nony

m o

f m

argi

natu

ssu

perc

ilios

us Q

uoy

and

Gai

mar

d, 1

832:

88

unpl

aced

valid

syno

nym

s: d

uple

x H

oyle

, 188

5: 2

26w

este

rnie

nsis

d’O

rbig

ny, 1

834:

pl.

10ta

ngar

oa O

’She

a, 1

999:

202

Ben

thoc

topu

sva

lidta

niw

ha k

ubod

era

O’S

hea,

199

9: 2

33G

rane

ledo

neva

lidta

niw

ha ta

niw

ha O

’She

a, 1

999:

222

Gra

nele

done

valid

tapr

oban

ensi

s (R

obso

n, 1

926:

165

)C

allis

toct

opus

nom

en d

ubiu

mte

ggin

mat

hae

O’S

hea,

199

9: 2

00B

enth

octo

pus

valid

tehu

elch

us d

’Orb

igny

, 183

4: 2

7 [i

n 18

34–1

847]

Oct

opus

valid

syno

nym

: lob

ensi

s C

aste

llano

s an

d M

enni

, 196

9: 9

2te

nebr

icus

Sm

ith, 1

884:

35

unpl

aced

valid

tenu

icir

rus

Sasa

ki, 1

929:

78

unpl

aced

syno

nym

of

hong

kong

ensi

ste

nuip

ulvi

nus

Sasa

ki, 1

920:

182

unpl

aced

unre

solv

ed*t

etra

cirr

hus

(Chi

aie,

183

0: p

l. 72

)P

tero

ctop

usva

lidsy

nony

m: c

occo

Ver

any,

184

6: 1

09ti

tano

tus

Tro

sche

l, 18

57: 5

1te

trad

ynam

us R

afin

esqu

e, 1

814:

28

Oct

opus

syno

nym

of

vulg

aris

tetr

icus

Gou

ld, 1

852:

474

Oct

opus

valid

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

144

Phuket mar. biol. Cent. Res. Bull.sy

nony

m: g

ibbs

i O’S

hea,

199

9: 1

20te

utho

ides

Rob

son,

192

9: 1

33un

plac

edju

veni

le: n

omen

dub

ium

*tha

umas

toch

eir

Rob

son,

192

8: 1

10G

rim

pella

valid

thie

lei R

obso

n, 1

932:

233

Ben

thoc

topu

sva

lidth

ysan

opho

ra V

oss,

196

2: 2

65E

ledo

nesy

nony

m o

f sc

hult

zei

tita

notu

s (T

rosc

hel,

1857

: 51)

Pte

roct

opus

syno

nym

of

tetr

acir

rhus

*tog

ata

Chu

n, 1

915:

479

Velo

dona

valid

syno

nym

: tog

ata

cape

nsis

Rob

son,

192

4: 2

06to

gata

cap

ensi

s R

obso

n, 1

924:

206

Velo

dona

syno

nym

of

toga

tato

ngan

us (

Hoy

le, 1

885:

225

)A

bdop

usva

lidtr

iten

tacu

latu

s R

isso

, 185

4: 6

3O

ctop

ussy

nony

m o

f vu

lgar

istr

osch

eli T

argi

oni-

Tozz

etti,

186

9: 1

13O

ctop

ussy

nony

m o

f vu

lgar

ists

ugar

ensi

s Sa

saki

, 192

0: 1

75un

plac

edun

reso

lved

tube

rcul

atus

Tar

gion

i-To

zzet

ti, 1

869:

156

Oct

opus

syno

nym

of

vulg

aris

turq

ueti

(Jou

bin,

190

5: 2

9)P

arel

edon

eva

lid*u

nici

rrhu

s (C

hiai

e, 1

839–

1841

: 70)

Scae

urgu

sva

lidva

ldiv

iae

(Thi

ele,

in C

hun,

191

5: 4

85)

Bat

hypo

lypu

sva

lidsy

nony

m: g

rim

pei

Rob

son,

192

4: 2

08va

lidu

s Sa

saki

, 192

0: 1

83un

plac

edun

reso

lved

vari

abil

is S

asak

i, 19

29un

plac

edun

reso

lved

vari

olat

us B

lain

ville

, 182

6: 1

86un

plac

edno

men

dub

ium

varu

nae

(Oom

men

, 197

1: 6

9)A

mph

ioct

opus

valid

velig

ero

Ber

ry, 1

953:

57

unpl

aced

valid

venu

stus

Ran

g, 1

837:

66

unpl

aced

juve

nile

: unr

esol

ved

vent

rico

sus

Gra

nt, 1

827:

309

unpl

aced

unre

solv

edve

rril

li p

alli

ata

Rob

son,

192

9: 1

63un

plac

edju

veni

le: n

omen

dub

ium

verr

illi

ver

rill

i Hoy

le, 1

886:

93

unpl

aced

juve

nile

: nom

en d

ubiu

m*v

erru

cosa

(Ver

rill,

188

1: 1

05)

Gra

nele

done

valid

syno

nym

s: v

erru

cosa

med

ia J

oubi

n, 1

918a

: 5

verr

ucos

a m

edia

Jou

bin,

191

8: 5

Gra

nele

done

syno

nym

of

verr

ucos

ave

rruc

osus

Hoy

le, 1

885:

222

Oct

opus

valid

vinc

enti

(Pic

kfor

d, 1

955:

159

)A

mph

ioct

opus

syno

nym

of

burr

yivi

oles

cens

Tak

i, 19

64: 3

20B

enth

octo

pus

valid

vitie

nsis

Hoy

le, 1

885:

226

unpl

aced

valid

*vul

gari

s C

uvie

r, 1

797:

380

Oct

opus

valid

syno

nym

s: a

lbus

Raf

ines

que,

181

4: 2

9bi

tent

acul

atus

Ris

so, 1

854:

61

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

145


brev

itent

acul

atus

Bla

invi

lle, 1

826:

187

cass

iope

ia G

ray,

184

9: 9

coer

ules

cent

es F

ra P

iero

, 189

5: 2

67he

tero

pus

Raf

ines

que,

181

4: 2

8m

acul

atus

Raf

ines

que,

181

4: 2

9m

arin

um K

oelr

eut ?

?m

osch

atus

Raf

ines

que,

181

4: 2

8ni

ger

Raf

ines

que,

181

4: 2

9oc

topo

dia

Try

on, 1

879:

113

pilo

sus

Ris

so, 1

826:

4ra

bass

in R

isso

, 185

4: 6

7ru

ber

Raf

ines

que,

181

4: 2

8te

trad

ynam

us R

afin

esqu

e, 1

814:

28

trit

enta

cula

tus

Ris

so, 1

854:

63

tros

chel

i Tar

gion

i-To

zzet

ti, 1

869:

113

tube

rcul

atus

Tar

gion

i-To

zzet

ti, 1

869:

156

Unr

esol

ved

Wes

t Atla

ntic

for

m:

Ava

ilabl

e na

me:

amer

ican

us B

aker

in D

enys

de

Mon

tfor

t, 18

02: 3

8sy

nony

ms:

bak

erii

d’O

rbig

ny, 1

826:

144

eudo

ra G

ray,

184

9: 9

gery

onea

Gra

y, 1

849:

7U

nres

olve

d Ja

pane

se fo

rm:

Ava

ilabl

e na

me:

sine

nsis

d’O

rbig

ny, 1

834:

pl.

9w

arri

nga

Stra

nks,

199

0: 4

57un

plac

edva

lidw

este

rnie

nsis

d’O

rbig

ny, 1

834:

pl.

10un

plac

edsy

nony

m o

f su

perc

ilio

sus

*win

ckw

orth

i (R

obso

n, 1

926:

161

)M

acro

chla

ena

valid

wit

jazi

Aki

mus

hkin

, 196

3: 1

45P

tero

ctop

usun

reso

lved

wol

fi W

ülke

r, 1

913:

458

unpl

aced

valid

yam

ana

Gue

rrer

o K

omm

ritz

, 200

0G

rane

ledo

neva

lidya

quin

ae V

oss

and

Pear

cy, 1

990:

76

Ben

thoc

topu

sva

lidye

ndoi

Sas

aki,

1920

: 179

unpl

aced

unre

solv

edze

alan

dicu

s (B

enha

m, 1

944:

256

)E

nter

octo

pus

valid

zeis

s O

’She

a, 1

999:

246

Tha

umel

edon

eva

lidzo

natu

s V

oss,

196

8: 6

47un

plac

edva

lid

Taxa

Gen

eric

pla

cem

ent

Stat

us

Tabl

e 3.

(con

tinue

d)

146


of which 13 were new to science. O’Shea (1999)reported 30 shallow and deep-water species fromNew Zealand waters of which 11 were new toscience. Australia has received more attention thanmost regions. In 1985, Lu and Phillips were thefirst to generate a list of octopus species fromAustralian waters. A total of 31 species wererecorded of which 12 proved to be valid speciesrecords or names. Stranks (1998) listed 26shallow-water octopus species names of whichhe proposed 15 to be valid. Through Australia-widefield collection over the past six years, the tallyhas now reached 76 valid species of which 55have been recognised in the past decade (worksof Norman, Stranks, Lu, Finn and Hochberg, seeliterature cited above).

Works in progress in other regions of theworld are uncovering similar numbers of newspecies. New Caledonian waters (Norman,Hochberg and Boucher, unpublished data) containat least 20 shallow-water species (of which 14 arenew to science) and 13 deep-water species (all ofwhich are new to science, including three newgenera) Norman et al., 2004a, 2004b. A preliminarycensus of the benthic octopuses of South Africaby Norman, Hochberg, Allcock and colleagues atthe CIAC conference in South Africa in 1997recognised 27 species of which 16 are new toscience. Hochberg (unpublished data, see also Younget. al., 1989) found 16 species of octopodidparalarvae in Hawaiian waters while only 8 specieshad been recognised in the adult fauna. The totaltally for this region is likely to be even higher asthe planktonic paralarvae only represent small-eggspecies. Additional large-egg species with benthicyoung also occur around Hawaii. Elsewhere in thePacific Ocean, many new species are coming tolight including in Tonga (Huffard, unpubl. data),Indonesia (Norman, unpubl. data; Huffard, unpubl.data), Marquesas (Hochberg, Boucher andNorman, unpubl. data), Fiji (Herb and Roper,unpubl. data), Peru (Hochberg and Cardoso,unpubl. data), and Mexico to Panama (Hochbergand Sweeney, unpubl. data).

Recent studies on Antarctic benthic octopuseshave found a high diversity of new species in (andrelated to) the genus Pareledone (Allcock et al.,2003). Allcock (pers. comm.) suggests more than

20 new species from the genus Pareledone alonemay be present in southern polar waters.

Cryptic speciesCryptic species are also coming to light

amongst the better-known faunas. Pygmy speciesare being discovered in all regions of the worldfrom tropical shallow waters (Norman, 2000, 2001;Hochberg, unpubl. data) to the deep sea (Norman,Hochberg and Boucher, 2004 and unpubl. data,Vecchione, this volume). Molecular studies areuncovering sympatric cryptic species, as foundrecently for Amphioctopus marginatus in Taiwan(Ho et al., 2000). Purported anti-tropicaldistributions are also being questioned for siblingsubtropical taxa split on the periphery of thetropical Pacific Ocean (Norman, 2000), namelyAmphioctopus kagoshimensis Ortmann, 1888(Japan) and A. cf kagoshimensis Norman, 2000(eastern Australia); Hapalochlaena fasciata Hoyle,1886 (eastern Australia) and Hapalochlaena sp. 2Norman, 2000 (Japan/China). An Octopus“vulgaris”-like species in Japan and O. tetricus andO. cf tetricus in Australia appear to be similar siblingspecies.

It is clear that there is much higher diversityof octopuses than previously considered. Voss(1977) calculated the rate at which new speciesof cephalopods were being described each decadesince 1835, in order to assess the rate of discoveryover the previous 140 years. He suggested anaverage rate of 6.7 species being describedannually. As there was no obvious decline of thisrate with time, he suggested that “one is forced tothe belief that we have nowhere near exhaustedthe number of undescribed species either on ourmuseum shelves or in the sea”. Voss went on tostate that “it seems clear that we are indeed still inthe descriptive or alpha stage of systematics andthat we are far from knowing the total number ofspecies of cephalopods”. These statements are stillvalid 28years later, particularly for benthic octopuses.The total tally for octopus species and genera islikely to rise dramatically should an in-depth, globalcensus be undertaken. There are many species yetto be discovered, studied and described. Effortsto date may just be skimming the surface of thereal diversity that is present in the world’s oceans.

147


Impediments to taxonomic progressVoss (1977) also discussed the problems

impeding better taxonomic knowledge, listing themas:

· lack of detailed morphometric studies· lack of adequate study material· failure to analyse character variations· need to examine type material· prevalence of uncritical reviews of literature,

perpetuating historical errors.Today many of these taxonomic problems are

still evident. Some factors have shownimprovement. Historically there has been littleconsistency in approach to species recognition anddescription. This is slowly changing. In the pastfive years in particular, more detailed and consistentdiagnoses and descriptions have been produced indescribing new species of octopuses (see list ofrecent taxonomic works in Introduction). Otherimprovements include better international links,faster electronic communication and easier travelthan ever before. There is also better technologyfor detection, sampling, photography and videodocumentation of taxa and the visual attributes oflive animals.

Other factors have deteriorated. Therecontinues to be problems with poor retention ofmaterial, few replicates from a paucity of localities,poor curation and a lack of defined characters.There is disturbingly little support for alphataxonomy research, with the few extant cephalopodtaxonomists being an aging and diminishing cohort.The changing nature of research institutions andavailable funding means few of these researchershave younger understudies on to which the mantle(!) can be passed. Diminishing support for thetraditional curation roles of museums has also takenits toll. As museum curators retire, they are notbeing replaced. Many of the major researchcollections of cephalopods around the world areneither actively worked nor growing. At best theyare maintained by collection managers responsiblefor multiple and diverse collections of invertebratesand other biological resources. As a consequenceof reduced science funding (and priority)worldwide, there are fewer primary field surveysbeing undertaken, compared with the many peaksof activity over the past century.

Despite the decline of support for this primarytaxonomic research, the demands for informationare greater than ever. As many finfish harvestscollapse around the world, commercial fisheriesattention is shifting to exploitation of cephalopodresources. Octopuses, squids and cuttlefishes arefast growing, fecund marine animals which makethem favoured and profitable fishery targets. Thecapacity of fisheries authorities to effectivelymanage octopus fisheries is severely hampered bymany problems, most of which stem from poortaxonomy. These include species lumping, lack ofdetailed diagnostic tools (regional keys), inadequatefisheries catch statistics (including lack of catchper unit effort data), dearth of biological informationfor target species, lack of detailed regional censuses(or even comprehensive fish market surveys) anda lack of well-preserved and well-managedreference and voucher collections.

Sound taxonomy underpins all other fields ofbiological research. At this stage, octopod taxonomyis still in need of major revision and stabilization.An immediate priority is production of detailed andaccurate descriptions for all species (includingredescriptions of named species supplemented byfresh reference material collected from typelocalities). Support should be sought for alphataxonomy on a scale sufficient to clarify the extantfauna, and should include encouragement ofstudents to enter and become active in this field.Regional revisions should also be instigated andsupported, particularly in regions where ourknowledge is poorest or where fisheries aredeveloping fastest.

Only this approach will enable production ofappropriate identification tools, particularly guidesand keys for field and fisheries identification. Armedwith such information and tools it will be possibleto develop appropriate monitoring and managementsystems for octopuses of current and potentialfisheries value, as well as identifying speciespotentially vulnerable to such exploitation. Additionof molecular studies in such studies will also providevaluable insights into the population genetics andorigins of these creatures. Without even basicinformation on the composition, distribution andbiology of these animals, sound management isunlikely.

148


ACKNOWLEDGEMENTS

The authors wish to thank Louise Allcock andCrissy Huffard for feedback on the manuscriptand data on regional fauna. This manuscript was

completed during research visits to Paris,supported by the Museum National d’HistoireNaturelle. We extend our appreciation to RenataBoucher Rodoni for facilitating and hosting thesevisits.

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APPENDIX

Clarification of the non-ocellate members ofAmphioctopus Fischer, 1882

Considerable confusion surrounds the groupof small to moderate-sized octopuses historicallyknown as the Octopus aegina species-group(Robson, 1929; Norman, 1992c, 1993b).Members of this genus are characterised by armsapproximately 2–3 times mantle length, deep lateralwebs (approximately 20% of arm length) and avery shallow dorsal web. The oral surface of theshallow dorsal web has the same sculpture andcolour pattern as the dorsal surfaces of body andarms. Species included in the genus may be eitherocellate or non-ocellate. Norman (1993b) andNateewathana and Norman (1999) have treated theocellate members of this species-group.

Member species are found throughout theIndo-Pacific region (A. aegina Gray, 1849, A.exannulatus Norman, 1993b, A. fangsiaod’Orbigny, 1839–41, A. kagoshimensis Ortmann,1888, A. marginatus Taki, 1964, A. mototi Norman,1993b, A. neglectus Nateewathana and Norman,1999, A. polyzenia Gray, 1849, A. rex Nateewathanaand Norman, 1999, A. robsoni Adam, 1941, A.siamensis Nateewathana and Norman, 1999, A.varunae Oommen, 1971 and at least sixundescribed species).

Fewer non-ocellate species are representedin the Atlantic Ocean (A. burryi Voss, 1950 and atleast one undescribed species). The status orvalidity of several old names variously applied tomembers of this group in the Atlantic still need tobe resolved (rugosus Bosc 1792; granulatusLamarck 1799; carolinensis Verrill 1884; andvincenti Pickford 1955).

There has been considerable confusion overthe non-ocellate members of the “aegina species-group”, primarily due to inadequate originaldescriptions, type material that is in very poorcondition or not extant, and historical lumping ofdistinct species (see review in Adam, 1960). Afailure to formally describe type and other materialin detail for members of this group has meant thisconfusion continues today.

The species pivotal in creating most of theconfusion is Amphioctopus aegina Gray, 1849.The species was treated as dubious by Toll andVoss (1998). The type of A. aegina exists (BM1928.2.14.1) and has been examined by theauthors. It is a mature female, which clearly showsthe colour pattern diagnostic of this species; namelya pale longitudinal stripe along the dorsal mantle, anarrow transverse head bar and numerousdistinctive circular reticulations over the mantle,arm crown and arms. We believe that thisrecognisable species is valid. Examination of thetype of A. hardwickei Gray, 1849 from the IndianOcean (presumed Singapore) found it to beidentical in form and colour pattern with the typeand other material attributed by us to A. aegina.Toll and Voss (1998) recognised that the taxon theytreated under the name A. hardwickei has “armed”spermatophores. This type of spermatophore ischaracterized by the presence of sharp teeth in theejaculatory apparatus that may aid penetration ofthe sperm bulb as the spermatophore everts withinthe oviduct of the female or may prevent removalby a competing male. All males attributed by us tothe genuine A. aegina possess such armedspermatophores. No other octopuses of similarmorphology throughout the region possess sucharmature. Remains of spermatophores in the typeof A. dollfusi Robson, 1928 from Vietnam clearlyshow this armature. As a consequence, both A.hardwickei and A. dollfusi are placed in thesynonymy of A. aegina, the former on the basisof page priority by A. aegina in Gray’s (1849)original descriptions.

Subsequent confusion has arisen in thisspecies-group through the use of the nameAmphioctopus aegina for two related but distinctoctopus species, namely A. kagoshimensis and A.marginatus. The former species is easilydistinguished from A. aegina based on charactersof the patch and groove skin sculpture which hasregular polygonal patches (versus round patches).The ligula is of moderate length (4–7% of armlength) and the terminal organ (“penis”) is greatlyelongated and recurved to accommodate the verylong, unarmed spermatophores of this species

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(1.5–2 times mantle length versus 0.6–1.2 inaegina). Amphioctopus marginatus Taki, 1964 isrecognised by its dark purple colour pattern insharp contrast to the white to blue suckers alongthe dorsal faces of arms 1–3. The skin is sculpturedin rounded elongate patches that form irregularlongitudinal rows, displayed as branched veins onthe lateral arm crown. In this species, the ligula issmall (1.5–3.5% of arm length) and thespermatophores are unarmed and only moderatelylong (approximately 0.75 to 1.2 times mantle length).

Robson (1929) initiated much of the confusionsurrounding this group by lumping records of allthree distinct species under the name aegina. Inthe same publication, Robson also treated membersof this species group under the name Octopusrugosus (Bosc, 1792), due to the nature of the“rugose” skin found in well-preserved material.

Bosc (1792) coined the name rugosus on thebasis of an animal from Senegal, west Africa thathad granular skin. There is no type material. Thename often has been treated in a long list ofsynonyms for Octopus vulgaris. A non-ocellatemember of Amphioctopus has been reported offthe coast of Senegal under the name O. burryi Voss,1952 (Roper et al., 1984) and a further species,O. vincenti Pickford, 1955 was described from anearby locality.

Re-examination of the original material listedby Robson (1929) under the name rugosus turnedup a number of Indo-Pacific members ofAmphioctopus, including three ocellate species(e.g., A. polyzenia Gray, 1849 from Torres Strait,Australia: Robson, 1929: 65 and an undescribedspecies from Amirante Atoll, Indian Ocean,Robson, 1929: 64) plus two non-ocellate species(A. aegina Gray, 1849 from Gulf of Martaban, IndianOcean: Robson, 1929: 64 and an undescribedspecies from Madras, India: Robson, 1929: 64).

The other name associated with the non-ocellate members of Amphioctopus is granulatusLamarck, 1798. Sasaki (1929) used this name forhis treatment of A. kagoshimensis. AlthoughLamarck (1798) did not provide a type locality forhis species and no type material has been tracedwe have determined that the material he examined

came from Martinique in the Caribbean. Other thanhaving granular skin, there are no charactersenabling recognition of this species from the brieforiginal description. Robson (1929) placed thistaxon in the synonymy of rugosus whereas manyother authors treat it as a synonym of Octopus vulgaris.At this stage, the name granulatus remains unresolved.

In order to help clarify the named Indo-Pacificnon-ocellate members of Amphioctopus, table 4presents the primary diagnostic characters for thenamed member species considered here to be valid:A. aegina, A. kagoshimensis and A. marginatus.Data for A. aegina and A. marginatus are basedon multiple reference specimens. Data for A.kagoshimensis are based on limited availablematerial and treatments of this species in historicalliterature (including Sasaki’s [1929] treatment ofthis octopus under the name Octopus granulatus).

In an attempt to clarify past literature recordsof these three member species, we also presentthe following annotated synonymies:

Amphioctopus aegina Gray, 1849(treated and illustrated in Norman and Sweeney,1997, Norman, 1998, 2000)

Octopus rugosus (non Bosc, 1792). Robson, 1929.O. hardwickei Gray, 1849. Toll and Voss, 1998.O. dollfusi Robson, 1928. Voss and Williamson,

1972, Li, 1983, Roper et al., 1984, Chikuni, 1987, Dong,1988, Chaitiamvong, 1993, Chotiyaputta, 1993.

A. kagoshimensis Ortmann, 1888(treated and illustrated in Norman, 2000; under O.granulatus in Sasaki, 1929)

O. granulatus (non Lamarck, 1798). Sasaki, 1929.O. aegina (non Gray, 1849). Robson, 1929

(in part), Taki, 1965, Dong, 1988, Okutani et al.,1987 (in part).

O. dollfusi (non Robson, 1928). Roper et al.,1984 (in part).

A. marginatus Taki, 1964(treated and illustrated in Norman, 1993c, 1998,2000; Norman and Hochberg, 1994)

O. aegina (non Gray, 1849). Voss andWilliamson, 1972, Chaitiamvong, 1993.

O. dollfusi (non Robson, 1928). Roper et al.,1984 (in part).

O. striolatus Dong, 1976. Dong, 1979, 1988.

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Synonyms: - striolatus dollfusihardwickei

Misidentifications: granulatus aegina rugosusaegina, dollfusi dollfusi

Distribution: NW Pacific Ocean Indian and Indian and(southern Japan to Taiwan) W Pacific oceans W Pacific oceans

Affinity: subtropical tropical tropicalMantle length: to 86 mm to 80 mm to 90 mmTotal length: to 295 mm to 280 mm to 300 mmArm length: 2.4, 2.6 x ML 2.0–2.8 x ML 2–3 x MLWeb depth: 25.6, 21.4% AL 26.3–33.3% AL 20.0–32.8% ALNormal arm sucker count: 157, 164 (m), 169 (f) 127–152 (both) 96–130 (both)Hectocotylised arm sucker count: 68–88 61–84 55–70Ligula length: 5, 6.7% HAL 1.6–3.6% HAL 4–6% HALCalamus length: 17% LL 37.9–65.0% LL 19–25% LLEnlarged suckers: none 4–5 slightly enlarged 2–3 enlargedMale sucker diameter: to 9.0% ML to 14.5% ML to 16.6% MLSpermatophores: no internal teeth (unarmed) no internal teeth (unarmed) internal teeth (armed)Spermatophore length: 117–160 mm 37–68 mm 25–51 mmSpermatophore length versus ML: 160, 186% ML 73–116% ML 58–115% MLColour pattern: Cross around eye, reticulate White triangle under eye, Longitudinal pale stripe

markings produced by dark dark purple leading edges on dorsal mantle. Darkgrooves. of arms in contrast to circular reticulations.

white-blue suckers.Skin sculpture: Polygonal patches over all Oval to elongate patches in Circular raised patches

surfaces with distinct grooves. irregular longitudinal rows. with indistinct grooves.

Table 4. Comparison of Indo-Pacific non-ocellate members of the genus Amphioctopus.[m = male, f = female, AL = arm length, HAL = hectocotylised arm length, LL = ligula length, ML =mantle length (eye to posterior tip)].

kagoshimensis marginatus aegina

13. norman...title 13. norman author nipol subject 13. norman created date 9/12/2005 8:56:37 am

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