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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. (mnorman@museum.vic.gov.au)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
Phuket mar. biol. Cent. Res. Bull.
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
The current state of Octopus taxonomy
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
Phuket mar. biol. Cent. Res. Bull.
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
The current state of Octopus taxonomy
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
The current state of Octopus taxonomy
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
The current state of Octopus taxonomy
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
The current state of Octopus taxonomy
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
Phuket mar. biol. Cent. Res. Bull.
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
The current state of Octopus taxonomy
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
The current state of Octopus taxonomy
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
The current state of Octopus taxonomy
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
Phuket mar. biol. Cent. Res. Bull.
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
The current state of Octopus taxonomy
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
Phuket mar. biol. Cent. Res. Bull.
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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from Australia. Bulletin of Marine Science. 49(1–2), 1991: 73–87.Muus, B. 2002. The Bathypolypus-Benthoctopus problem of the North Atlantic (Octopodidae, Cephalopoda).
Malacologia. 44(2): 175–222.Nateewathana, A. 1997. The octopod fauna (Cephalopoda: Octopoda) of the Andaman Sea, Thailand.
Phuket Marine Biological Center Special Publication. 17(2): 407–452.Nateewathana, A. and M.D. Norman. 1999. On three new species of ocellate octopuses (Cephalopoda:
Octopoda) from Thai waters. Phuket Marine Biological Center Special Publication. 19(2): 445–462.Norman, M.D. 1992a. Octopus cyanea Gray, 1849 (Mollusca: Cephalopoda) in Australian waters:
description, distribution and taxonomy. Bulletin of Marine Science. 49(1–2), 1991: 20–38, 5 figs.Norman, M.D. 1992b. Ameloctopus litoralis gen. et sp. nov. (Cephalopoda: Octopodidae), a new shallow-
water octopus from tropical Australian waters. Invertebrate Taxonomy. 6: 567–82.Norman, M.D. 1992c. Systematics and biogeography of the shallow-water octopuses (Cephalopoda:
Octopodinae) of the Great Barrier Reef, Australia. Unpublished Ph.D. thesis. University of Melbourne.Norman, M.D. 1993a. Four new species of the Octopus macropus group (Cephalopoda: Octopodidae)
from the Great Barrier Reef Australia. Memoirs of the Museum of Victoria 53(2), 1992: 267–308.Norman, M.D. 1993b. Ocellate octopuses (Cephalopoda: Octopodidae) of the Great Barrier Reef, Australia:
description of two new species and redescription of Octopus polyzenia Gray, 1849. Memoirs ofMuseum of Victoria. 53(2), 1992: 309–344.
Norman, M.D. 1993c. Octopus ornatus Gould, 1852 (Cephalopoda: Octopodidae) in Australian waters:morphology, distribution and life history. Proceedings of the Biological Society of Washington.106(4): 645–660.
Norman, M.D. 1998. Family Octopodidae, Benthic octopuses. In: K.E. Carpenter and V.H. Niem (eds.).FAO Species Identification Guide for fishery purposes. The living marine resources of the WesternCentral Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. FAO, Rome. pp.800–826.
Norman, M.D. 2000. Cephalopods: a World Guide. IKAN Publishing, Frankfurt, 320 pp.Norman, M.D. 2001. New octopus species from Queensland, Australia. Memoirs of the Queensland
Museum. 46(2): 677–690.Norman, M.D. and J. Finn. 2001. Revision of the Octopus horridus species group with description of
two member species from the Great Barrier Reef, Australia. Invertebrate Taxonomy. 15: 13–35.
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Norman, M.D. and F.G. Hochberg. 1994. Shallow-water octopuses (Cephalopoda: Octopodidae) of HongKong territorial waters. In: B. Morton (ed.). The Malacofauna of Hong Kong and southern ChinaIII, Proceedings of the Third International Workshop on the Malacofauna of Hong Kong andsouthern China. Hong Kong University Press. pp. 141–160.
Norman, M.D., F.G. Hochberg and C.C. Lu. 1997. Mollusca Cephalopoda: Mid-depth octopuses (200–1000 m) of the Banda and Arafura Seas (Octopodidae and Alloposidae). In: A. Crosnier and P.Bouchet (eds.). Résultats des Campagnes MUSORSTOM, Volume 16. Bulletin de Museum Nationald’Histoire Naturelle, Paris, 172: 357–383.
Norman, M.D. and C.C, Lu. 2000. Preliminary checklist of the cephalopods of the South China Sea.Special edition on the Biodiversity of the South China Sea, Bulletin of the Raffles Museum, Singapore.8: 539–567.
Norman, M.D. and M.J. Sweeney. 1997. The shallow-water octopuses (Cephalopoda: Octopodinae) ofthe Philippine Islands. Invertebrate Taxonomy. (1992), 11: 89–140.
Norman, M.D., R. Boucher and F.G. Hochberg. 2004. The Sharkclub Octopus, Galeoctopus lateralis, anew genus and species of deep-water octopus from thewestern Pacific Ocean(Cephalopoda:Octopodidae). Journal of Molluscan Studies. 70: 247–256
Norman, M.D., F.G. Hochberg and R. Boucher-Rodoni. 2004. Microeledone mangoldi gen. and sp. nov.,a deep-water pygmy octopus from the Norfolk Ridge, New Caledonia (Cephalopoda: Octopodidae).Molluscan Research. 24: 193–209
Okutani, T., M. Tagawa and H. Horikawa. 1987. Cephalopods of continental shelf and slope aroundJapan. Japan Fisheries Resource Conservation Association, Tokyo. 194 pp.
D’Orbigny, A. 1845. Molluscques vivants et fossiles, ou description de toutes les espèces de coquilles etde mollusques. Adolphe Delahays, Paris. 605 pp.
O’Shea, S. 1999. The marine fauna of New Zealand: Octopoda (Mollusca: Cephalopoda). NIWA BiodiversityMemoir. 112: 1–280.
Quoy, J.R.C. and J.P. Gaimard. 1832. Voyage de l’Astrolabe, pendant les anees 1826–1829. Zoology, 2.Paris. 320 pp.
Robson, G.C. 1929. A monograph of the Recent Cephalopoda. Part I. Octopodinae. (British Museum:London.) 236 pp.
Roper, C.F.E., M.J. Sweeney, and C.E. Nauen. 1984. FAO species catalogue. Volume 3, Cephalopods ofthe world: an annotated and illustrated catalogue of species of interest to fisheries. FAO FisheriesSynopsis. (125)3: 1–277.
Sasaki, M. 1929. A monograph of the dibranchiate cephalopods of the Japanese and adjacent waters.Journal of the College of Agriculture, Hokkaido Imperial University, 20 (supplement): 1–357.
Stranks, T.N. 1988a. Systematics of the family Octopodidae (Mollusca: Cephalopoda) of south-easternAustralia. M.Sc. thesis. University of Melbourne, Victoria. 114 pp.
Stranks, T.N. 1988b. Redescription of Octopus pallidus (Cephalopoda: Octopodidae) from south-easternAustralia. Malacologia. 29(1): 275–287.
Stranks, T.N. 1990. Three new species of Octopus (Mollusca: Cephalopoda) from south-eastern Australia.Memoirs of the National Museum of Victoria. 50(2): 457–465.
Stranks, T.N. and M.D. Norman. 1993. Review of the Octopus australis complex (Cephalopoda:Octopodidae) and description of a new species. Memoirs of the Museum of Victoria. 53(2), 1992:345–373.
Stranks, T.N. 1998. The systematic and nomenclatural status of the Octopodinae described from Australia(Mollusca: Cephalopoda). Smithsonian Contributions to Zoology. 586(2): 529–547.
Sweeney M.J. and C.F.E. Roper. 1998. Classification, type localities, and type repositories of recentCephalopoda. Smithsonian Contributions to Zoology. 586(2): 561–599.
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Taki, I. 1965. Cephalopoda: 307–326. In: OKADA et al., (eds.). New Illustrated Encyclopedia of theFauna of Japan, vol. 2.
Thomas, R.F. 1977. Systematics, distribution, and biology of cephalopods of the genus Tremoctopus(Octopoda: Tremoctopodidae). Bulletin of Marine Science. 27: 353–392.
Toll, R.B. 1998. The systematic and nomenclatural status of the Octopodinae described from the IndianOcean (excluding Australia) and the Red Sea. In: N.A. Voss, M. Vecchione, R.B. Toll and M.J.Sweeney (eds.). Systematics and Biogeography of Cephalopods. Smithsonian Contributions toZoology. 586(2): 475–487.
Toll, R.B. and G.L. Voss. 1998. The systematic and nomenclatural status of the Octopodinae describedfrom the West Pacific region. In: N.A. Voss, M. Vecchione, R.B. Toll and M.J. Sweeney (eds.).Systematics and Biogeography of Cephalopods. Smithsonian Contributions to Zoology. 586(2):489–520.
Vecchione this volume (do not have citation).Voss G.L. 1977. Present status and new trends in cephalopod systematics. Symposia of the Zoological
Society of London. 38: 49–60.Voss, G.L. and G.R. Williamson. 1972. Cephalopods of Hong Kong. Hong Kong Government Press: Hong
Kong. 138 pp. [Title page dated 1971, actual publication date 1972].Voss, N.A., M. Vecchione, R.B. Toll and M.J. Sweeney. 1998. Systematics and Biogeography of
Cephalopods. Smithsonian Contributions to Zoology. 586 (2 volumes).Young R.E., R.F. Harman and F.G. Hochberg. 1989. Octopodid Paralarvae from Hawaiian Waters. The
Veliger. 32(2): 152–165.
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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
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