Memoirs of the Museum of Victoria 53(2): 267-308 (1992) 

FOUR NEW OCTOPUS SPECIES OF THE OCTOPUS MACROFUS GROUP 
(CEPHALOPODA: OCTOPOD1DAE) FROM THE GREAT BARRIER REEF, AUSTRALIA 

By Mark D. Norman 
Invertebrate Zoology. Museum of Victoria. 285 Russell Street, Melbourne, Victoria 3000, Australia 

Abstract 

Norman, M.D., 1992. Four new octopus species of the Octopus macropus group (Cepha- 
lopoda: Octopodidae) from the Great Barrier Reef, Australia. Memoirs of the Museum of 
Victoria 53: 267-308. 

Four new species of shallow-water octopuses are described from tropical waters of the 
Great Barrier Reef, Australia. All four are members of the "Octopus macropus group" (Rob- 
son. 1 929). characterised by arms of unequal length with the dorsal pair longest (AF 1 .2.3.4), 
moderate to high gill lamellae counts (10-14 per demibranch) and a robust conical copu- 
latorv organ. All are nocturnally active. Two species. O. alpheus and O. aspilosomatis spp. 
nov occur in clear waters foraging predominantly on intcrtidal coral reefs and offshore 
islands. O. dierylhraeus sp. nov. forages intertidally and subtidally on muddy substrates in 
coastal waters. Octopus graptUS occurs in more open waters, on sandy and mud substrata in 
the channels and flat bottoms between islands. Full morphological descriptions arc pro- 
vided, along with details of known distributions, life history and commercial exploitation. 
Delineation of each species from related taxa is discussed. 

Introduction 

A number of workers have recognised the dis- 
tinctive group of octopus species often referred 
to as the "Octopus macropus group" (Robson, 
1929; Adam, 1941; Taki, 1944, 1964; Voss. 
1981). Members of this species group are found 
in most tropical and temperate waters of the 
world and are characterised primarily by 
elongate arms with the dorsal pair longest (AF 
1 .2.3.4), moderate to high gill counts (10- 1 4 per 
demibranch), a moderately large cylindrical 
copulatory organ with deep grooved ligula. and 
nocturnal activity patterns. 

The species from which this group derives its 
name. Octopus macropus Risso, 1826, was 
described from the Mediterranean Sea. This 
species appears limited to the Mediterranean 
Sea and temperate eastern Atlantic (Hochberg, 
Mangold and Norman, in prep.). A number of 
species from Indo-West Pacific waters show 
close morphological and behavioural similar- 
ities with O. macropus. Asa consequence, many 
of these species have regularly, and inappropri- 
ately, been assigned the name O. macropus. both 
within Australian waters [Girard, 1 890; Brazier. 
1892;Odhncr, 1917;Nesis, 1982 (plate in 1987 
abridged translation); Lu and Phillips, 1 985], as 
well as elsewhere in the Indo-West Pacific region 
(Joubin, 1894, 1898: Goodrich, 1896; Hoyle, 
1904: Berry, 1912. 1914; Wiilker, 1913. 1920; 
Massv, 1 91 6; Sasaki. 1920; Robson, 1926. 1929, 
1932: Boone, 1938; Adam, 1939, 1942, 1946, 

1954, 1959, 1960, 1973; Rces and Stuckey, 
1954- Voss, 1963; Roper et al., 1984). 

Sasaki ( 1 920) was the first to question the sup- 
posed wide distribution of O. macropus, when 
tentatively assigning the name to a Japanese 
species: "(there is) much doubt whether the 
species extends as far as the Japanese waters 
from its home; that is, the Mediterranean Sea". 
Significant physical and temperature bound- 
aries separate the distribution of the Indo-West 
Pacific species from that of O. macropus and 
there is no doubt that the Indo-West Pacific 
species are distinct taxa. A major review of the 
Indo-West Pacific members of the O. macropus 
group is required, including re-examination of 
species names from these waters previously syn- 
onymised under O. macropus (Robson, 1929; 
Roper et al., 1984). 

Recent research into the shallow-water octo- 
puses of the Great Barrier Reef and northern 
Australia has uncovered a surprisingly rich octo- 
podan fauna (Norman, 1991, 1992a, 1992b, in 
prep.). At least 25 species have been recognised 
from these waters, of which only five can be 
assigned to previously described taxa. Amongst 
this rich fauna are five members of the O. macro- 
pus group. Octopus ornatus Gould, 1852 is a 
large species widely distributed throughout the 
tropical Indo-West Pacific. Norman (in prep.) 
describes the morphology of this species and its 
occurrence in Australian waters. The remaining 
four taxa are described here as new species: O. 

267 

268 

MARK D. NORMAN 

alpheus, O. aspilosomatis, O. dierythraeus and 
O. graptus spp. nov. 

A diagnostic key to the O. macropus group in 
Great Barrier Reef waters is provided along with 
full species accounts for the four new species. A 
diagnosis and distribution for O. ornatus is pre- 
sented for comparison. 

Material and methods 

Field collection on the Great Barrier Reef was 
carried out between May and November in 1 989 
and 1990. Totals of 82 individuals of O. alpheus, 
26 of O. aspilosomatis and 24 of O. dierythraeus 
were encountered live in the field, primarily at 
night on intertidal reef walks. Active lairs also 
were found intertidally on daylight reef walks, 
and subtidally through day and night snorkel 
and SCUBA dives. The occupants of these lairs 
were flushed using a weak solution of copper sul- 
phate in sea water (<0.5 M) and captured in 
hand nets. Individuals were observed and 
photographed in situ and, where facilities were 
available, within aquaria. 

Representative specimens were retained and 
narcotised in fresh water, and fixed and pre- 
served according to the techniques of Roper and 
Sweeney (1983). These specimens are now 
housed in the Museum of Victoria (NMV). 
Additional specimens of all four new species 
were found in the collections of the Australian 
Museum, Sydney (AM), Queensland Museum, 
Brisbane (QM), National Museum of Natural 
History. Washington, (USNM) and Museum 
National d'Histoire Naturelle, Paris (MNHN). 

In the description and tables, measurements 
and indices follow Roper and Voss (1983). The 
terms "terminal organ" and "copulatory organ" 
are used to replace "penis" and "hectocotylus" 
respectively. These changes follow Hochberg 
and Mangold (in prep.). The structure histori- 
cally known as the "penis" is not an intromittent 
organ and hence should not be referred to by this 
term. Terminal organ is a more appropriate term 
for this element of the male reproductive tract. 
The term "hectocotylus" refers to the entire 
modified arm and not the modified distal tip. 
This tip functions as the intromittent organ dur- 

ing copulation and is more appropriately named 
the copulatory organ. 

The following additional or modified indices 
are also employed: 

Stage of Maturity (StM): Immature (Imm: sex 
indeterminate or reproductive organs minute), 
Submature (S: reproductive organs distinct but 
poorly developed) and Mature (M: developed 
spermatophores or eggs distinct); Head Mantle 
Width Index (HMWI): head width as per cent of 
mantle width; Arm Mantle Index (AMI): arm 
length as per cent of ML; Arm Width Index 
(AWI): arm width at widest point on stoutest 
arm. as per cent of ML; Sucker Count (SC): total 
sucker count for intact arm with the highest 
sucker count; Gill Count (GC): number of gill 
lamellae per demibranch not including the 
medial terminal lamella; Hectocotylized Arm 
Mantle Index (HAMI): length of hectocotylized 
arm as per cent of ML; Hectocotylized Arm 
Sucker Count (HASC): number of suckers on 
hectocotylized arm; Terminal Organ Limb 
Index (TOLI): length of terminal organ as per 
cent of ML: Diverticulum Length Index (DLI): 
diverticulum length as per cent of length of ter- 
minal organ; Spermatophore number (SpN): 
number of spermatophores in Needham's Sac; 
Sperm Cord Whorls (SpCW): number of whorls 
in which sperm cord is coiled: Funnel Length 
Index (FLI): funnel length as per cent of ML; 
Free Funnel Index (FFI): length of free funnel 
portion as per cent of funnel length: Funnel 
Organ Index (FOI): length of outer limb of fun- 
nel organ as per cent of median limb length; 
Funnel Organ Length Index (FOLI): length of 
medial limb as per cent of funnel length. 

In the descriptions, indices are presented for 
both sexes combined, except where significant 
differences were found between the sexes (one 
way ANOVA, p = 0.05). In these indices, range 
and mean for each sex are presented. Where 
ranges significantly overlap, standard devi- 
ations around the mean are also presented. 

Table 1 summarises the key differences 
between Great Barrier Reef members of the 
Octopus macropus group. 

Key to species of the "Octopus macropus group" from Great Barrier Reef waters 

1. Distinctly elongate species with almost cylindrical arms and shallow 
webs, deepest web always less than 1 5% of length of longest arm, typically 

1 0% o 

Robust, moderately elongate species with deep webs, deepest web always 
greater than 1 5% of length of longest arm, typically 20% 3 

2. Large species (to 130 mm ML, 1.2 m TL and 1 kg) with high gill counts 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

269 

a 
o 

00 

s 

»3 

3 

t 

o 

4= 

O 

a 

pi 

CQ 
ca 

a 

PS 
c 

ca 
a 

S 
o 
O 

ca 

H 

*3 

s 

I 

3 

3 



oo 
I 

rn 

o o o ■* I 

CN <N 

OO 

m 

I 

<~n< 

I 

u 

I 

NO 

~^ 

I 

r) 


— i in in m — i — < i> 

no 

to 

I (N o 

0\ (N 00 

I 

O O O ^D 
On r*1 tS 

cn 
I 

u 

on 


OO 
OO 

I 

_L I Oo — 

. "* I I s. 

no On ^O CI IT 

_ __ oo — i ?>■ 

00 

.2© 

X> 

oo 

I 
o^. 

in o O in 

in 

I r- O m 
<n <n oo fS 
,< I <N — i 

On <N 

in ~~i 

<~nj ~-< tj" o 

I | <-h <u m 

rr m I p opro 

in no r- m O csl ^ 3 i 

— m — < cn -* — 'i— ' — ( 

I 

f<N-Hinvot^fnON 
oo 

O 

-a 
<u 

On 

m 

no 
I 

^o 

K 

<"n 

in 

I 

O O od 
O ■* <N ro 

^r no 

— i fN 
I I m 

NO in On 

^ <M in 

I °o 

NO I 

O OO 

I 

oo ■* — ' ""* <^ ^ >~~ 

I 

2£ 

O 
O 
O 

1° 

c/) 

a 
ca 

s 

c 
'c3 

? o 

in 
r--' 

I in oo 
_: I <N 

00 

I 

O O 

o f"> ■* 

I r- 

On On 
© I 
<N -^ CN 
I On — 
CN I I 

03 O 
O <u O 

a oo_ 

— no on NO p 5 ) 

U 

IZJ 

o 



E 

I- 
< 

U 


S i— ' -^ S U l, 

E 

& 
ca 

C 

ca 
oo 

3 3 o 


on 



a 3 3 

oo 007; 
00 00 o 

<^IOfcWWU 

270 MARK D. NORMAN 

(13-14 lamellae per demibranch), high sucker counts (SC 324-342-382, 
HASC 1 52-164-112) and distinctive colour pattern of wide longitudinal 
white stripes on dorsal mantle and paired white spots on dorsal arm 
crown and arms, over brown to deep maroon base colour 

Octopus ornatus Gould, 1 852 

Medium sized species (to 80 mm ML, 440 mm TLand 1 20 g) with mod- 
erate gill counts (10-11 lamellae per demibranch) and moderate sucker 
counts (SC 206-235-267, HASC 78-55-95). Colour pattern of plain red 
or white dorsal mantle lacking spots, arm crown and dorsal arms covered 
with regular paired white spots over red base colour 

Octopus aspilosomatis sp. nov. 

3. Colour patterns consisting of red base colour with large white spots or the 
negative pattern of white base colour with red spots; W- or UU-shaped 

funnel organs 4 

Colour pattern simple, consisting of pale cream to pink base colour with 
dark brown "scribbling" (spots and short irregular lines) over dorsal 
mantle and arm crown (fig. 13a), VV-shapcd funnel organ with outer 
limbs much shorter than medial limbs; very large robust species (to 190 
mm ML, 1.3 m TL, 4.2 kg), gill counts high (13-14 per demibranch) 

Octopus graptus sp. nov. 

4. Large robust species (to 120 mm ML, 715 mm TL, 1.2 kg), with high gill 
counts (12-14 lamellae per demibranch) and moderate sucker counts 
(SC 234-259-280, HASC 103-7/2-125), funnel organ always UU- 
shaped, alarm display of large red spots over white base colour on dorsal 
mantle, arm crown and arms (figs 9a, 12c-d) 

Octopus dierylhraeus sp. nov. 

Medium sized species (to 80 mm ML, 430 mm TL, 340 g), with moderate 
gill counts (1 1-12 lamellae per demibranch) and moderate sucker counts 
(SC 192-209-228, HASC 82-97-97), funnel organ typically W-shaped. 
largest specimens with slight separation of medial limbs, alarm display of 
large white spots over red base colour on dorsal mantle, arm crown and 
arms (figs la, 4a-c) Octopus alpheus sp. nov. 

Octopodidae d'Orbigny, 1839 Reef and several individuals from the warm- 

Subfamily Octopodinae d'Orbigny, 1 839 temperate waters of the New South Wales coast. 

1 his species is widely distributed throughout the 

Octopus ornatus Gould, 1 852 tropical Indo-West Pacific region, from Hawaii 

Figure 17a to East Africa. Figure 17a presents the distri- 

Diagnosis. Large muscular and elongate species bution , of j hlS u , S P ed f ba * ed ° n , ex£miir ! ed 

with unequal arms, dorsal arms longest and matenal and published records that clearly refer 

most robust (AF 1 .2.3.4), dorsal arms typically 6 to th,s distmctive spec.es. 

to 7.5 times mantle length. Webs shallow, Remarks. See Norman (in prep.) for full mor- 

approximately 9% of length of longest arms, phological description, annotated synonymy 

Sucker counts, 320-380 per arm in submature and details of life history based on Australian 

and mature animals, 150- 170 suckers on hecto- material. Callistoctopus arakawai Taki, 1964, 

cotylized arm of submature and mature males, described from southern Japan, is a junior syn- 

Gill lamellae, 13-14 per demibranch, typically onvm. 

14 Funnel organ W-shaped. Eggs small (to Octopus a| heus nov 

3.5 mm long) produced in large numbers 

(>30 000). Alarm colouration of white longi- 

Figs 1-4, 17b 

tudinal stripes on mantle and paired spots along Octopus macropus. — Nesis, 1982 (1987 abridged 
aboral arm surfaces over maroon red base English translation): plate on p. 74. (non Risso. 

colour. 

1826) 

„. ., . „, , , „ Material examined. 82 individuals were encountered 

Distribution, Octopus ornatus is recorded from , ive in thc fidd on the islands of thc Capncorn Bunker 

offshore islands ot the southern Great Barrier Group at the southern end of the Great Barrier Reef. 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

271 

18 were retained and are now in the Museum of Vic- 
toria. Six additional preserved specimens were found 
and examined in Australian museums and the 
National Museum of Natural History. Washington. 

Holotype: Qld: lcf: 74.2 mm ML, NMV F57930, 
Tryon L. Capricorn Bunker Group. 23°15'S. 151°47'E, 
0.1-0.2 m, M. Norman. 2 Nov 1989 (active at 0435- 
0515 hr). 

Paratvpes: Qld: lcf: 61.9; I9: 70.1 mm ML. NMV 
F60100! Tryon L. 23°15'S, 15 WE. 0.1-0.2 m, M. 
Norman. 1 Nov 1989 (active at 0415-0530 hr); I9: 
78.2 mm ML, NMV F65660. Tryon I., 23°15'S. 
1 5 1°47'E, 0.1-0.2 m, M. Norman. 2 Nov 1989 (active 
at 0435-0515 hr). 

Other material: Qld: Id: 19.7 mm ML, NMV 
F57928, One Tree I., 23°30'S. 1 52°05'E. 0.2 m. S. Jack- 
son, 1 6 Oct 1 989 (active at 1 830 hr); 29: 20.2, 25.6 mm 
ML, NMV F57926. Tryon I., 23°15'S, 151°47'E. 0.1- 
0.2 m, M. Norman, 2 Nov 1989 (active at 0435-0500 
hr. retreated into lair, flushed with CuS0 4 ); I9: 20.9 
mm ML. AM C3 1668, Masthead I.. 23°32'S. 1 5 1 °45'E. 
D.B. Fry. no date; I9: 30.8 mm ML. AM CI 59263, 
North West I.. 23°18'S. 151°42'E. M. Ward and W. 
Boardman, "Dec 1929-Jan 1931"; I9: 32.3 mm ML. 
QM Mo35762, Wilson I.. 23°I8'S, l5r55'E. 18 Sep 
1972; 3d. 49: 36. 1-77.0 mm ML, NMV F65662, Tryon 
I.. 23°15'S. 151°47'E, 0.1-0.2 m. M. Norman. I Nov 
1989 (active at 04 1 5-0530 hr); Id: 38.6mm ML.NMV 
F57925. One Tree L. 23°30'S. 1 52°05'E, 0.2 m, R. Fen- 
wick and M. Norman, 7 Sep 1990 (active at 1840 hr): 
1 9 : 40.2 mm ML, NMV F57929, Tryon I.. 23°15'S. 
15P47'E, 0.1 m, M. Norman. 31 Oct 1989 (active at 
0520 hr); Id: 44.0 mm ML. NMV F57927, One Tree I.. 
23°30'S, 1 52°05'E, R. Fenwick and M. Norman, 7 Sep 
1990 (active at 1910 hr): Id: 45.0 mm ML. AM 
CI 59262, Heron L. 23°26'S, 151°57'E, Dec 1964; 2d, 
I9: 45.0-63.9 mm ML. NMV F60099. One Tree I., 
23°30'S, 1 52°05'E. <0.3 m, M. Norman. 1 7 Oct 1 989 
(active at 1925-2100 hr); Id: 49.0 mm ML. USNM 
817786. One Tree I., 23°30'S, I52 C 05'E, 0-2 ft (0-0.7 
m) 22 Nov 1 969 (rotcnone station); 19:65.8 mm ML. 
AM CI 59265. North West I., 23"18'S. 151°42'E. A.A. 
Livingstone and W. Boardman, "Dec 1930 to Jan 
1931". 

Diagnosis. Large muscular species with unequal 
arms, dorsal arms longest and most robust (AF 
1.2.3.4), typically 3 to 4.5 times mantle length. 
Webs moderately deep, approximately 20% of 
length of longest arm. Sucker counts, 200-230 
per arm in submature and mature animals. 80- 
100 suckers on hectocotylized arm of submature 

and mature males. Gill lamellae, 10-12 per 
demibranch. typically 1 1 on the outer demi- 
branch and 1 2 on the inner demibranch. Funnel 
organ typically W-shaped. No mature females 
encountered, submature females with large-type 
eggs, produced in low numbers. Alarm colour 
pattern of large white spots over deep red base 
on dorsal surfaces of mantle, arm crown and 
arms. 

Description. The following description is based 
on 2 submature and 3 mature males, and 7 sub- 
mature females. Counts and indices for this 
material are presented in Tables 2 and 3, with 
data on 5 immature males and 3 immature 
females. 

Moderate to large robust species (fig. la): no 
mature females found in this study, ML of sub- 
mature females and mature males to at least 80 
mm, TL to at least 430 mm; weight to at least 
340 g. Mantle variable from round to elongate 
(MW1 39.7-77.6-89.2), mantle walls moder- 
ately muscular. Stylets present. Pallial aperture 
of moderate width, approximately half mantle 
width. Funnel long, muscular and broad based 
(FLI 48.0-55. 7-69.2) with free portion approxi- 
mately half funnel length (FFI 30. \-46. 7-74.2). 
Funnel organ well developed (fig. 1 b), typically 
W-shaped, except in 2 largest specimens (I9: 
78.2 mm ML and Id: 74.2 mm ML, NMV 
F57930 exhibited slight separation of the medial 
limbs approaching a UU-type funnel organ). 
Outer limbs of funnel organ slightly shorter than 
median limbs (FOI 75.0-79.7-83.6). Funnel 
organ approximately 50% of funnel length 
(FOL1 48.2-5 3 .3-6 1.7). 

Head of moderate width (HWI 43.0-.vS.5- 
74.6). typically narrower than mantle (HMW1 
69.0-82.5- 108.3). Neck distinct, slightly 
narrower than head. Eyes large and slightly 
pronounced. 

Arms of moderate length, typically 3 to 5 
times mantle length (AMI 316.1-427.0-507.5). 
Arms robust (AW1 14.7-20.7-25.9), roughly 
square in cross section and tapering evenly along 
length. Arms unequal in length, dorsal pair 
longest, ventral pair shortest (AF typically 
1 .2.3.4). Suckers of moderate to large size (SDI 
I 1.9- 14. 3- 17. 4), deep with distinct radial 
cushions and scalloped outer rim. Scalloping 
exaggerated on small distal suckers to form ring 
of small digits of skin around rim of sucker. 
Suckers largest on dorsal arms, none especially 
enlarged in either sex. Approximately 200 
suckers per intact normal arm, with slightly 
higher counts in females (SCd 1 92-203 ± 9. 98- 
?16 n = 4d; 9 208-27 7± #.32-228, n = 59). Webs 
of moderate depth (WDI 16.3-27.S-25.0). 
deepest dorsally evenly decreasing in depth to 
shortest ventral web (WF typically A.B.C.D.E). 
Web margins extended on ventral edges of arms 
for majority of length. 

Right third arm in males hectocotylized. 
slightly shorter than opposite arm (OAI: 68.5- 
70.3-72.1; HAMI: 197.8-225.9-274.6). Copu- 
latory organ (fig. lc) of moderate size [LLl(mat) 

272 

MARK D. NORMAN 

5mm 

30mm 

5mm 

Figure 1 . Octopus «//j//tm sp nov a dorsal view of 74.2 mm ML male (holotype, NMV F57930). b, funnel organ 
ol 41.0 mm ML male (NMV F65662). c, copulatory organ of 74.2 mm ML male (holotype, NMV F57930). 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

273 

10mm 

Figure 2 Octopus alpheus sp. nov. a, digestive tract of 43.3 mm ML male (NM V F65662). ASG = anterior salivary 
PlS BM buccal I mass- C = caecum CD = crop diverticulum; CR = crop; DC = d.gcst.ve gland; I - intestine; IS 
ftS^-S&EsS^Sffi salivary gland; R - rectum; S - stomach, b-d, beaks of 77 mm ML 
female (NMVF65662". b, upper beak, lateral view, e, lower beak, lateral view. d. lower beak, ventral view. 

5.3_7.2-9. 1], roughly cylindrical with deep 
liguia groove and moderate calamus [CLI(mat) 
20.9-23. 9-28.9]. Liguia groove contains >20 
fine transverse grooves. Spermatophore groove 
well developed and wide with fine transverse 
grooves. Spermatophore guide distinct with 
moderately deep notch and flattened ridge 
topped with square papillae. Approximately 90 
suckers on hectocotylized arm (HASC 82-W- 

97) 

Gills with 10-12 lamellae on each demi- 
branch, typically 1 1 on outer and 12 on inner 
demibranch. Terminal lamella small. 

Digestive tract illustrated in figure 2a. 
Anterior salivary glands approximately one- 
third of buccal mass length. Posterior salivary 
glands slightly longer than buccal mass. Crop 
diverticulum large in the 2 specimens dissected, 
almost as large as digestive gland but possibly 
distended by contents. Stomach bipartite. 
Caecum clearly striated, coiled in 1.5 to 2 
whorls. Intestine thin walled and reflexcd in 
proximal third. Rectum straight and muscular. 
Ink sac well developed, embedded in ventral sur- 
face of digestive gland. Ink red when released by 
live animals. Fleshy anal flaps present. 

274 

MARK D. NORMAN 

a 

10mm 

10mm 

Figure 3. Octopus alpheus sp. nov. a, reproductive tract of 74.2 mm ML male (holotype, NMV F57930). A = 
appendix; D = diverticulum; MG = mucilaginous gland; NS = Needham's Sac; SGI = spermatophoric gland I; 
SG2 = spermatophoric gland II; T = testes; TO = terminal organ; VD = vas deferens, b-c, spermatophore of same 
specimen, b, whole spermatophore. CB = cement body; EA = ejaculatory apparatus; OC = oral cap; SR = sperm 
reservoir, c, detail of oral cap. 

Upper beak (fig. 2b) with small hood and 
weakly hooked rostrum. Lower beak (figs 2c-d) 
with moderately sharp rostrum, narrow hood, 
widely spread wings and flared lateral walls. 
Ventral view of posterior margin of lateral walls 
deeply concave. Radula with 7 transverse rows 
of teeth and marginal plates (figs 4e-f). Rhach- 
idian tooth has 2-3 lateral cusps, typically 3, on 
each side of large curved medial cusp. Lateral 
cusps in asymmetrical seriation, migrating from 
lateral to medial position over 8-9 rows. First 
lateral teeth small and unicuspidate; second lat- 
eral teeth unicuspidate, of moderate length and 
robust; lateral marginal teeth relatively straight, 

short and robust; marginal plates oblong and 
plain. 

Male reproductive tract illustrated in figure 
3a. Terminal organ in mature males of moderate 
length and very robust (TOLI 19.2, 24.8, 32.4) 
with robust diverticulum (DL1 53.8, 57.5, 58. 1). 
Genital aperture subterminal. Mucilaginous 
gland enlarged at point of attachment to vas def- 
erens. Spermatophoric gland I narrow with large 
recurved coil approximately 80% along length. 
Spermatophoric gland II robust and moderately 
short with reflexed tip. Spermatophores (figs 
3b-c) almost as long as mantle length (SpLI 92.3, 
93.7, 95.1), of moderate width (SpWI 2.6, 2.9, 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

275 

Figure 4. Octopus alpheus sp. nov. a-c, active individuals on reef flats at night on Tryon I. d, dorsal view of 
preserved 74.2 mm ML male (holotype, NMV F57930), showing wavy transverse lines on dorsal arm crown, e-f. 
radula of 77.0 mm ML female (NMV F65662). e. dorsal view. f. lateral view. 

276 

MARK D. NORMAN 

> 

03 

E 

. X) 

> a 
o <*> 

a ii 

S3 

-5- S 
Q c 

s " 

2 ■ " 
O c 

u c 

o n 
Ik" 

"Sj 

O 

U 

S 
3 

Q 

y- r- •£► 

" m o 

> OS 
111 ON 

z ° 

Uh 

Z ° 

> 

a 

<L> 
3 

111 >o 

2 ^> 
>7 in 

Uh 

OS 
ON 

o 
o 

NO 
Uh 

NO 

ON 
m 

U 

1< rN 

2 ON 

z ^ 

U- 

> rN 

1, NO 
2 NO 

Uh 

> m 

1. rN 

2 ON 

Z r "- 

Uh 

oo 

ON 

in 

Uh 

O 

z 

ai 

^ d — in— I oo ^f' oo -■^ : odC2l ° r ^ 00 * c ' Ti:a ' r ' <N ''^ <N — I 

Tf <n m 

no in oo no O r~ on 

(N —■ — ' rN 

rN rN in On 

rN O 

in m oo in 

on 5! oo in on ^r 

m 

NO 

oc 
m 

0O NO 0O ON 

A M 


oo^j-oofNmr--pir-r<-)0"^"oo — i -h r- rN oo rN no on 
-h o -: M on , • • 

rN 

tj- r- in r~ m 

m -h fN -h <n — ; 
nm -h 

r~ <N rN ro in m r— 
rN m in on ■**■ 

it *r 

rN m 

-H CNl 

in 

e: 

ininooONm-Hm-nmooO 

^. inNornoNrnNoodinrf— « © 
N /11-«rnnO\OM — i — i rN 

— i on • ■ °^ • ■ ■ l— ' 

oo 
O r- 

fN 

m c 


rf no 

NO O 

00 

~ | 

Q 


ON o 
oo rN 

oo 
ID 

CI 

CN 


Q 

so 

ON. 

m 
m 

d 

a a 

oo 

NO 

oo tj- 

r— 


rN 

rN 

rN 

Hi 

— i 

— 


rs 




r«i 


— H 




Tf 

rri 


ri 

m 




— 


<^i 








O c/3 r-~ m — ■r'-imrNONOorNooON'^-rN 

in £] — iinr-'rnrNoor^rNosrNi'^j-^ 

<-j ''r^inr^r— rNooONNO — i — i rN — i 

m rn rN — . rN — 

oo in tj- on m 

r-' oo °° n-i ^t- 

QQ 

c c 

on no m 

Hi HH 

o 

ci^rNr—oor^rNQrN^-in'^-o-H 

~ 'rnr-'odrNrn oo-h'onnoon ' 

r-inr-r^rf _h no —■ — i ■— i 

mm rN rN 

r^j 

OO ^ TJ- NO C C 

rN 

O crNmm^t-inoinoomNor--mo 

— < a Dj r--' on no on — < on on r-' in cs id C 

ti n in^oOHVOHOO- '— 'rNrt 

rf m m rN mi _ 

OO NO 0O ON O ^3" 
OO 

on in 

O NO 

fN 

o -=t 
■t 

cooNoi^oor^or^Nor-r^in-H 

S2^NONdoNrNin'r---'r---'odmON~' 
r--ini^ONinooooo — i — i — i 
m m rN rN rN 

m r- rN o O r-~ — i 

ON 

2 s 

cm — aMvotCMNi^miNMM 

00 *0(Ni'*'^ : o6rnModri'tt 
t~~NOoooONin— •■^--h — i rN — i 
m rN rN rN rN -h 

r^ rN 
O r- 
rN 

rN Tt r- 

m rN ^ 

— i oo 
rN 

m in in on 
rN in 

c c ^o c 

d I j I £ § § 


on ^j- on m 
in r- r- in 

rNONO^ONrNQNOor~~-HONrNNONOQoomaNrNinr-Nom t — oorNt~~- 

t on ■*' -t t~-' in ■^-od^}--^-— i^rl^t aN in'o6oNt-~'mrNin in rNrN — i 
rn r~-r-~ONO0— ■ t^r^— i — c ri -h N h- rs-ninON in ininooNO 

O O "<t no 

— i O t~ Tf 
no in t~- rh 

O in O no 

rf no in r- 
in tT r~- in 

ON O NO f- 

in n-> r~- in 

oo rN c C 
■* r- — i— i 

in oo m on 

on — < — ■' in 
in ^- oo ■<*■ 

■* p r-_ — i 

in r-~' — < no 
in in r-~ in 

^f On m t^ 
in m oo in 

U H, h M m hh 

HCHJUHQcri^iyNoritiHti^LLitJL, 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

277 

00 

S 

c 
.2 

S3 
> 

o 

S 

a 

>• s 

oS 

c <~ 

3 6 
3 

§•11 

t) * 

O V 

u n 

73 3 

s a 

,<U C 

i- 3 
O w 

8^ 

'•5 tf 

T3 to 

§'■3 

C II 

c 

> 

<u 

H S 

73 

tu 

O0 

s 

173 

T3 

no ca 

> ^ 

~ SO 

2 so 

so 

Ph 

> ° 

4h NO 
Uh 

in 

SO 
CN 

ON 

u 

ON 
ON 

o 
o 

NO 

rN 

SO 

so 

IT) 

NO 

5 

< 

ON 

n 

ON 

r- 

IT) 

u 

> rN 

2 so 

NO 

P- 

> NO 

2 ON 

z r ~ 

M in 

NO 

<N 

ON 

>n 

Uh 

O 

Z 

do 

<n vi o — i m m —i 

oo 
r-- 

1 Q °° -. 

■ rN — i oo # 
■^■mcNiONO r-t^rNiin'ro^r: ^ 
mvo^ioom ^t -h fN — ■ rN <^ so 

Tt HH fN — ' CNl 

O oo r- no rN 
2 so ■* rn oo 

Ot/iON^r-OM^o-io-^-aoNMO * oi^^ono 

N l" - ^ i— H ON ^^ ■ ^"^ — H 

NO *Nf NO ON SO rN OO — i — i — I 

rn m m rN 

<N _; 

-HC/30000SO— 'SOrNOO^l- 

in O m rN oo * 
-h (ni r~- 

ooaNCNiNOmmrNONrocN 

\04t^OtNWWHC-HNH 

f CflflCl -h 

oo(/)oor^or<"> — -^ Mt^- 

in 

NO 

^ONmooNOooO T 3 - oo(Nin^. 

"(ntO^N"!^- ' — ' rN — 

^■mOHOo 
— i o 

CN| 

r~ £| on on r~ r-' — •' oo in in r--' <-<S C! ^ 

in ^OONOr^rNONNONOrN — ^CnI — i 

rN 

P~-C/3rnOONNOOONOTfO<N — inOCnI 

r~ 

o m CT\ rn — r~ t— oo m rN 

r^ no r- 

t^-oor~-CNi-H— ■cn-h 
-d- co m m •— i 

rn/5O^0'Nl'* | n(N^f^t-;*-^<N 

o S » f> *V d d w ci » io'C 

^- ^ r- no oo O no — i — ■ rN — ■ rN — i 

—i g ro no rN — ' — < on 
no <i . ^J* cv On 
en c m it r~ 

tj- oo in o O <N 
•^•r-NorNjoN^r— ■— 'CN^h 

nO 

in 

COr-4ooP^r~-roornOr-_ONrxi 

S~ '<nr-^o6'tf : r-'oodo6 — i -— ; C! 

C"^NOinooOr- | nro— ■— |<N— i 

mM mm — i 

rN f3fNiOM>^Q' r i , tsDrn^;oo-' 

• H OO — I ' ^ i~J r-J y^i r-^' *~si tHt ' ' 

© a 

CNl ^ 

»o 

NO NO O 

r- r- <N 
no in Tf 
cn| r^l es 


|— < — I CnI CO 

^ -h o c c 
v so,o~~ 

O on rN in o 

5 in Tt on <-! 
in tj- r- in 

V 

O O oo hh m 

2 rn in ON ON 

. . >n m r- Tt 

O O r>-> no — i 

2 — ' ON ON — ' 

7; so ^ r^ m 

rN on oo ** 

on rn on rsi 
so in r- >n 

gQQQQ 
2 c a 

? 

od ro C C 

oo 1— i so so 

oo on rn rn 
tj m oo in 

in — i r- t~- 

n- od o rN 
in tj- t— in 

<oo^OooWWWUhUhUhUh 

278 

MARK D. NORMAN 

3.1) and produced in low numbers (2 and 3 in 
Needham Sacs of 2 specimens). Oral cap simple 
bearing long cap thread. Sperm reservoir 
approximately half of spcrmatophore length 
(SpRI 45.1, 47.7, 55.3), sperm cord coiled in 
approximately 40 regular whorls. 

No mature females were encountered in this 
study. Immature ovaries were visible in sub- 
mature females, poorly developed in even the 
largest female. Large type eggs in low numbers 
clearly visible in undeveloped ovaries. Very 
small ovary of I female (78.2 mm ML: NMV 
F57930) contained approximately 1 30 long thin 
immature eggs up to 5.4 mm long. Approxi- 
mately 22 follicular folds visible on immature 
eggs. Mature eggs would be very large and pro- 
duced in low numbers. 

Colour in life orange (to deep red in alarm dis- 
play) covered in circular white spots over dorsal 
body, arm crown, webs and arms (figs la, 4a-c). 
Spots on arms paired and evenly spaced. 
Additional spots on dorsolateral faces of arms 
including 1 on the base of each sucker on at least 
arms I and 2. Pairs of slightly darker markings 
on dorsal mantle. Iridescent tissue layer present 
in skin, giving iridescent green sheen in torch- 
light at night. 

Skin sculpture simple, consisting of scattered 
low papillae located in the centre of white spots, 
especially on dorsal and lateral mantle. Trans- 
verse pair of large papilla in largest white spots 
on dorsal mantle. Small punctae scattered 
between papillae on dorsal surfaces. Single 
slightly larger supraocular papilla directly above 
each eye, surrounded by low papillae. Pigmen- 
tation and sculpture do not extend to oral 
surface of webs. 

Preserved specimens tend to lose dark red 
colour and definition of spots, base colour 
fading to cream. Pale spots containing central 
papilla sometimes visible on mantle, especially 
on lateral faces. Two to three dark wavy trans- 
verse lines across dorsal arm crown often visible. 

There appears to be little sexual dimorphism 
in this species. Sucker counts were slightly lower 
for males in material examined. 

Distribution. Offshore islands of the Capricorn 
Bunker group at the southern end of the Great 
Barrier Reef, Queensland (fig. 17b). Specimens 
have been collected from Tryon (23°15'S, 
151°47'E), Northwest (23°18'S, 15P42'E), Wil- 
son (23°18'S, 151"55'E), Heron (23°26'S, 
151°57'E), One Tree (23°30 , S, 152°05'E) and 
Masthead Islands (23'32'S, 151'45'E). 
Life history. Nocturnally-activc species which 

forages on exposed intertidal coral reef flats on 
offshore islands. Preys on small crabs and other 
crustaceans by probing crevices and burrows 
with arm tips. This species occupies lairs within 
coral bedrock and under living coral, closing 
entrance during the day with pieces of dead 
coral. It is unclear whether lairs are permanent 
or temporary refuges. Animals only emerge after 
dark. 

A total of 82 individuals was encountered 
actively foraging, always in less than 0.3 m on 
exposed reef flats. Individuals were only 
encountered at night between 1 840 and 0600 hr. 
No specimens were encountered subtidally, 
cither during the day or at night. 

This species often emerges from pools and tra- 
verses areas of exposed reef whilst foraging. 
Water was not retained within the mantle cavity 
while animals traversed exposed sections of reef. 
Individuals were often heard making whistling 
or squeaking noises when in very shallow pools 
or traversing dry ground as they drew air in and 
out of the mantle cavity. 

This species also tolerates low salinities, being 
difficult to narcotise in freshwater. This toler- 
ance may relate to the intertidal habits of this 
species where rain can cause locally low sal- 
inities in closed and shallow pools. 

Foraging individuals often were captured 
carrying small crabs on suckers close to the 
mouth. Several were caught carrying much 
larger crabs (up to 100 mm carapace width). 
Examination of stomach contents revealed only 
crustacean remains. 

Foraging individuals of O. alpheus appeared 
tolerant of conspecifics, often crawling over 
each other whilst seeking prey without any 
observed aggression. This tolerance was not 
observed by congeneric octopuses which appear 
to be a predator of at least smaller members of 
this species. One small individual of O. alpheus 
was attacked and partially devoured by a large 
individual of O. ornatus Gould, 1852 when 
released into an aquarium containing the latter 
species. The stomach contents of O. ornatus has 
been found to contain the beaks of smaller octo- 
puses (Norman, in prep.). 

Large eggs indicate young adopt a benthic 
habit on hatching. 

Etymology. From the Latin "alphus'" meaning 
white spots on the skin, referring to the white 
spots generated in the alarm display of this 
species (fig. 4a). 

Remarks. The colour plates in the 1 987 abridged 
English translation ofNesis( 1982) show an indi- 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

279 

vidual of O. alpheus at Heron I., under the head- 
ing O. macropus. The distinctive colour pattern 
and locality of this specimen clearly identify it as 
an individual of O. alpheus. 

Octopus aspilosomatis sp. nov. 

Figs 5-8, 17c 

Octopus ornatus. — Roper and Hochberg, 1 987: 1 6. 
18. — 1988: 161. (non Gould, 1852). 

Material examined. 25 individuals were encountered 
live in the field on Lizard and Russell islands and on 
the mainland coast at Cape Tribulation. 21 were 
retained and are now in the Museum of Victoria. 4 
additional preserved specimens were found and exam- 
ined in the collections of the Australian Museum and 
the National Museum of Natural History. Washing- 
ton. 

Holotvpe: Qld: \d\ 42.1 mm ML. NMV F67001. 
Russell L Frankland Group, 17°14'S. 146WE, <0.1 
m. M. Norman and S. Troy, 19 Oct 1990 (active at 
0255 hr). 

Paratvpes: Qld: ltf: 42.2 mm ML. NMV F60I48, 
Casuarina Beach. Lizard L. I4°41'S. I45°27'E. 0.1 m. 
M. Norman. 14 Nov 1989 (active at 0440-0530 hr); 
ltf: 44.7 mm ML, NMV F60149. Coconut Beach. Liz- 
ard L, Tryon I.. I4°40'S. 145°28'E, <0.l m. M. Nor- 
man. C. Davies and . I. Martin. 1 5 Nov 1989 (active at 
0450 hr): 1 9 : 56.8 mm ML, NMV F60I47. Russell I., 
Frankland Group, I7"14'S, 146°06'E. <0.l m. M. 
Norman. S. Troy. 19 Oct 1990 (active at 0335-0350 
hr); I?: 67.1 mm ML. NMV F60145. Casuarina Beach. 
Lizard 1.. I4°4I'S, 145°27'E, 0.1 m, M. Norman. 13 
Nov 1989 (active at 0330-0515 hr). 

Other material: Qld: 4tf; 25.4-34.4, 1 9: 36.4 mm 
ML, NMV F67002, Coconut Beach, Cape Tribulation 
National Park. I6°05'S. 145°29'E. <0.2 m, M. Nor- 
man. S. Troy. 16 Oct 1990 (active at 0020-0145 hr); 
I9: 27.5 mm ML, USNM 817667, off Coconut Beach. 
Lizard L, I4°40'S, I45°28'E, 3 m. in open water 25-30 
m deep. Larval Fish Survey, 27 Dec 1986 (in floating 
night light trap); 3d 1 : 30.5-51.8, I9: 50.7 mm ML, 
NMV F60144, Coconut Beach. Lizard I., 14°40'S. 
1 45°28'E, <0. 1 m. M. Norman, C. Davies and J. Mar- 
tin. I 5 Nov 1 989 (active at 0450 hr); Irf: 37.3 mm ML. 
NMV F65659. Casuarina Beach. Lizard L. 14°4I'S, 
145°27'E, 0.1 m, M. Norman, 14 Nov 1989 (active at 
0440-0530 hr); 39: 39.0-45.9 mm ML, NMV F65664, 
Casuarina Beach, Lizard I., I4°41'S. I45°27'E, 0.1 m, 
M. Norman. 13 Nov 1989 (active at 0330-0515 hr); 
\<S: 39.3. I9: 39.8 mm ML, NMV F65661, Russell I., 
Frankland Group. 17°I4'S, 146WE, <0.1 m. M. 
Norman, S. Troy. 19 Oct 1990 (active at 0335-0350 
hr); 1&. 49.3. 83.2 mm ML, AM CI 59275, Casuarina 
Beach, Lizard I., I4"41'S. 145°27'E, W.F. Ponder, P.H. 
Colman, I. Loch, 29 Nov 1974 (active at night): Irf: 
67.9 mm ML, NMV F60I46. Casuarina Beach, Lizard 
L. 1 4°40'S. 1 45°28'E. 0.2 m, M. Norman, 1 3 Nov 1 989 
(active at 0325 hr). 

Diagnosis. Moderate sized, elongate species 
with unequal arms, dorsal pairs longest (AF 

1.2.3.4), typically 4.5-6 times mantle length. 
Webs shallow, approximately 10% of length of 
longest arm. Sucker counts, 200-260 per arm in 
submature and mature animals, 78-95 suckers 
on hectocotylized arm of submature and mature 
males. Gill lamellae, 10-11 per demibranch, 
typically 10 on outer demibranch and 11 on 
inner demibranch. Funnel organ W-shaped. No 
mature females encountered, submature 
females with small-type eggs, produced in large 
numbers (about 30 000 in 1 specimen). Alarm 
colour pattern of red dorsal mantle and arms, 
and paired white spots on arm crown and aboral 
surface of arms only. 

Description. The following description is based 
on 4 submature and 6 mature males, and 1 sub- 
mature females. Counts and indices for this 
material are presented in Tables 4 and 5. 

A number of specimens of O. aspilosomatis 
possess greatly elongated mantles where the 
mantle becomes almost cylindrical (fig. 8d). 
Voss (1981) reported such variation in mantle 
shape in O. ornatus, variation also witnessed in 
Australian material of O. ornatus (Norman, in 
prep.). Certain indices are presented separately 
for normal and elongate material in the descrip- 
tion below. 

Moderate sized, elongate species (fig. 5a): no 
mature females found in this study, ML of sub- 
mature females and mature males to at least 80 
mm, TL to at least 440 mm; weight to at least 
120 g. As in Octopus ornatus, mantle is highly 
variable from ovoid to extremely elongate in 
some preserved specimens (fig. 8d: MWI 27.5- 
52.5-72.9), mantle walls moderately muscular. 
Stylets present. Pallial aperture of moderate 
width, slightly greater than half mantle width. 
Funnel long, muscular and broad based (FLI 
30.9-45. 1-6 1.1) with free portion variable in 
length, typically around half funnel length (FFI 
28.6-57.2-86.8). Funnel organ well developed 
(fig. 5b), W-shaped with broad limbs. Outer 
limbs clearly shorter than median limbs (FOI 
58.4_6<5.O-84.0). Funnel organ approximately 
50% of funnel length (FOLI 44.6-52.5-61.7). 

Head of moderate width (HWI 26.8-46.3- 
55.3). generally narrower than mantle (HMWI 
73.4-90-109.8). Neck distinct, slightly nar- 
rower than head. Eyes large and pronounced, 
especially in elongate form. 

Arms long, tvpicallv 4.5 to 6 times mantle 
length (AMI 438.4-5J7.6-639.8, n = 13, 314.9- 
391. 4-41 1.2 in elongate forms, n = 4). Arms 
robust (AWI 9.9-/4-19.9), square in cross sec- 
tion and tapering evenly along length. Arms 

280 

MARK D. NORMAN 

3mm 

5mm 

20mm 

Figure 5. Octopus aspilosomatis sp. nov. a, dorsal view of 42. 1 mm ML male (holotype, NMV F67001). b, funnel 
organ of same specimen, c, copulatory organ of same specimen. 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

281 

— O 

10mm 

Figure 6 Octopus aspilosomatis sp. nov. a, digestive tract of 50.7 mm ML female (NMVF60 1 44) [abbre viat ions 
lateral view, d, lower beak, ventral view. 

unequal in length, dorsal pair longest, ventral 
pair shortest (AF typically 1.2.3.4). Suckers 
biserial, of moderate size and slightly larger in 
males [SDIrf (normal) 11.4-/2.4-13.2, SDltf 
(elongate forms) 6.4-5.2-9.8; SDI9 6.7-5.5- 
9.6]. Suckers deep with fine radial cushions and 
scalloped outer rim. Suckers largest on dorsal 
arms, none especially enlarged in either sex. 
Approximately 230 suckers per intact normal 

arm in both sexes (SC 206-235-267). Webs shal- 
low (WDI 9.1-77.6-14.6), shortest ventrally, 
remaining webs subequal (WF typically 
A=B=C=D.E). Web margins extended on ventral 
edges of arms for majority of length. 

Right third arm in males hectocotylized. 
slightly shorter than opposite arm (OAI: 55.0- 
60.5-65.9; HAMI: 134.6-270.4-279.6). Copu- 
latory organ (fig. 5c) of moderate size [LLI(mat) 

282 

MARK D. NORMAN 

a 

10mm 


NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

283 

6.1-7.5-8.8, n = 6]. roughly cylindrical with 
deep ligular groove and moderate calamus 
[CLI(mat) 1 5.5-2/. 7-29.3]. Ligula groove deep 
and typically closed. Spermatophore groove well 
developed and wide with fine transverse 
grooves. Spermatophore guide distinct with 
moderately deep notch and flattened ridge 
topped with square papillae. Approximately 85 
suckers on hectocotylized arm (HASC 78-55- 
95). 

Gills with 10-11 lamellae on each demi- 
branch. typically 10 on outer and 1 1 on inner 
demibranch. Terminal lamella small. 

Digestive tract illustrated in figure 6a. 
Anterior salivary glands approximately one- 
third of buccal mass length. Posterior salivary 
glands approximately equal in length with 
buccal mass. Crop diverticulum large in the 
specimen dissected, larger than digestive gland 
but distended by contents. Stomach bipartite. 
Caecum clearly striated, coiled in 1.5-2 whorls. 
Intestine thin walled, reflexed in proximal third. 
Rectum muscular and straight. Ink sac well 
developed and elongate, ink duct long. Ink red 
when released by live animals. Fine elongate 
anal flaps present. Membrane on dorsal surface 
of visceral mass pigmented with large dark chro- 
matophores, presumably remnants of larval or 
founder chromatophores. 

Upper beak (fig. 6b) with small hood and 
weakly hooked rostrum. Lower beak (figs 6c-d) 
with moderately sharp rostrum, narrow hood, 
widely spread wings and flared lateral walls. 
Ventral view of posterior margin of lateral walls 
deeply concave. Radula with 7 transverse rows 
of teeth and marginal plates (figs 8g-h). Rha- 
chidian tooth has 2-3 lateral cusps on each side 
of large curved medial cusp. Lateral cusps in 
asymmetrical seriation, migrating from lateral 
to medial position over 8-9 rows. First lateral 
teeth small and unicuspidate; second lateral 
teeth unicuspidate, of moderate length and 
robust; lateral marginal teeth curved and rela- 
tively fine; marginal plates oblong and plain. 

Male reproductive tract illustrated in figure 
7a. Terminal organ in mature males of moderate 
length and robust [TOLl(mat) 21.4-26.2-33.6] 
with robust diverticulum (DLI 48.5-55.4-61.3). 
Genital aperture subterminal. Mucilaginous 
gland enlarged at point of attachment to short 
and robust vas deferens. Spermatophoric gland I 
narrow with large recurved coil approximately 
80% along length. Spermatophoric gland II nar- 
row and moderately long with reflexed tip. Sper- 
matophorcs (fig. 7b) approximately 80% of 
mantle length, proportionally snorter in 

elongate forms [SpLI (normal) 75.1, 82.9, SpLI 
(elongate) 47.8], of moderate width (SpWI 2.5- 
2.6) and produced in low numbers (1-3 in 
Necdham Sacs of 3 specimens). Oral cap simple 
bearing long cap thread. Sperm reservoir 
approximately half of spermatophore length 
(SpRI 46.2, 54.3), sperm cord coiled in approxi- 
mately 55 regular whorls. 

No mature females encountered. Immature 
ovaries were visible in submature females. Small 
type eggs in high numbers were clearly visible in 
undeveloped ovaries. Largest female (67.1 mm 
ML paratype, NMV F60I45) possessed an 
immature ovary 14 mm in diameter with 
elongate narrow oviducts and oviducal glands 
containing approximately 15 braiding chambers 
(fig. 7c). This ovary contained about 30 000 
immature eggs up to 1.1 mm long. 6 follicular 
folds were visible in these undeveloped eggs. 
Mature eggs would be small and produced in 
large numbers. 

Colour in life orange to deep red on the 
mantle, arm crown and aboral surfaces of all 
arms. White spots on arm crown and arms in 
most colour patterns (figs 8a, e-f). Spots on arms 
paired and evenly spaced. Transverse pair of 
small dorsal white spots visible on dorsal mantle 
in some colour patterns (fig. 8a). White spots as 
found on arm crown never present on dorsal 
mantle. Distinct, but faint, patch and groove 
trellis present (fig. 8,b c) on dorsal and lateral 
mantle. Dorsal mantle often mottled with 
irregular darker markings. Alarm colour pattern 
of white mantle, dark eye bars and red arm 
crown and arms with prominent white spots 
observed in several specimens. 

Skin sculpture simple, consisting of scattered 
low papillae evenly distributed over dorsal sur- 
faces. Small punctae scattered between papillae 
on dorsal surfaces. Single slightly larger supra- 
ocular papilla directly above each eye, sur- 
rounded by low papillae. Pigmentation and 
sculpture do not extend to oral surface of 
webs. 

Preserved specimens tend to lose dark red 
colour and definition of spots, base colour 
fading to cream. Pale spots containing central 
papilla sometimes visible on arm crown and dor- 
sal arms. Two to three dark wavy transverse 
lines across dorsal arm crown often visible. 

Sexual dimorphism was not marked in the 
material examined. 

Distribution. Moderately clear waters of north- 
ern Queensland (fig. 1 7c). Specimens have been 
collected from intertidal reef flats at Lizard I. 

284 

MARK D. NORMAN 

Figure 8. Octopus aspilosomatis sp. nov. a, active individual on reef flats at night on Lizard I. (photograph 
courtesy of C.F.E. Roper), b-c, live aquarium photographs of Lizard I. specimens, d, dorsal view of preserved 
67.9 mm ML male (NMV F60 1 46), showing greatly elongated mantle, c, active individual on reef flats at night on 
Lizard I. f, live photograph of 42.1 mm ML male (holotype, NMV F6700I). g-h, radula of 46.7 mm ML male 
(NMV F60I44). g, dorsal view, h, lateral view. 

(14°41'S, 145°27'E), Russell I. (17°I4'S, 
146WE) and Cape Tribulation (16°05'S, 
145°29'E). One specimen was collected swim- 
ming on the surface in open water at night oft" 
Lizard I. in 25-30 m. 

The distribution of this species may prove to 
be considerably wider, since eggs are small and 
hatehlings are presumably planktonic. 

Life history. Nocturnally active species which 
forages on exposed intertidal coral reef flats in 
moderately clear water of the offshore islands 
and mainland coast of northern Queensland 
where coral reefs are well developed. This 
species occupies lairs within coral bedrock and 
under living coral, blocking entrance during the 
day with pieces of dead coral. It is unclear 
whether lairs are permanent or temporary ref- 
uges. No prey remains were found surrounding 
lairs. Animals only emerge after dark. 

A total of 25 actively foraging individuals was 
encountered, always in less than 0.3 m on 
exposed reef flats. Individuals were only 
encountered at night between 0020 and 0535 hr, 
which corresponded with low tides at time of 
collection. No specimens were encountered sub- 
tidally, either during the day or at night. 

This species preys on small crabs and other 
crustaceans by probing crevices and burrows 
with arm tips. Foraging individuals were cap- 
tured carrying small crabs on suckers close to the 
mouth. Cannibalism is also likely to occur in the 
wild as several larger individuals readily 
attacked and commenced devouring smaller 
conspccifics when placed in the same con- 
tainer. 

This species tolerates low salinities, being dif- 
ficult to narcotise in freshwater. As was noted 
with O. alpheus, this tolerance may relate to the 
intertidal habits of this species where rain can 
result in temporary low salinities in closed and 
shallow pools at low tide. 

Small eggs indicate young adopt a planktonic 
habit on hatching. 

Several specimens contained large numbers of 
nematodes in the connective tissue membranes 
surrounding the visceral mass. 

Etymology. From the Greek "aspihs" meaning 
without spots and "soma" meaning body, refer- 
ring to the absence of red or white spots on the 
dorsal mantle in all colour patterns. 

Remarks. Roper and Hochberg (1987, 1988) 
reported O. ornatus Gould. 1852 from Lizard I. 
Examination of photographs of live material 
forming the basis of this record, Roper and 
Hochberg's reference to large numbers on the 
reef flats adjacent to the research station, and the 
low hectocotylized arm sucker count (about 90. 
R. Toll. pers. coram.), all clearly indicate that 
these reports refer to O. aspilosomatis and not O. 
ornatus. 

Octopus dierythraeus sp. nov. 

Figs 9-12. 17d 

Material examined. 24 individuals were encountered 
live at 6 sites along the Queensland coast from Lizard 
I. (I4°40'S. 145°28'E) in the north, to Sarina Beach 
(2L24'S, I49°I9'E) in the south. 21 were retained and 
are now in the Museum of Victoria. 10 additional pre- 
served specimens were found and examined in the 
collections of the Museum of Victoria, Australian 
Museum. National Museum of Natural History, 
Washington, and Museum National d'Histoire Natii- 
rellc, Paris. 

Holotype: Qld: Id 1 : 135.0 mm ML, NMV F67007, 
oil" W coast of Cape Yorke, Gulf of Carpentaria. 
I0°5I'S. I40°27'E, 57 m.CC.LU, "Susan Wright", 14 
Sep 1982 (trawl, 1940-2040 hr). 

Paratypes: Qld: \<S\ 59.8 mm ML, NMV F60129, 
Orpheus I., Cattle Bay. 18°34'S, 146°29'E. 1.5 m. M. 
Norman, 1 1 Oct 1 990 ( 1 630 hr, flushed from lair with 
CuS0 4 ); lef: 89.1 mm ML, NMV F60127. Lizard I.. 
Watsons Bay, Coby Hole (14°40'S, 145°28'E), 16 m, 
M. Norman, C. Davies, 23 Nov 1 989 ( 1 100 hr, flushed 
from lair with CuS0 4 ); l 9 : 118.8 mm ML, NMV 
F60128, Sarina Beach, 2L24'S. 149°19'E„ 0.3 m, M. 
Norman, 16 Sep 1990 (active at 0330 hr. handnet). 

Other material: Qld: 2?: 13.8, 19.4 mm ML, AM 
C50580, Hayman I., 20°03'S, 148°53'E. E.H. Rainford 
(on reef): I9: 20.3 mm ML, AM CI 59268, "near 
Townsville", near 19°I6'S, I46°50'E, E. Worrell, Mar 
1956 (in estuary); 5tf, 3?: 25.7-80.7 mm ML, NMV 
F60I33, Alexandra Reef (near I6°35'S. I45"30'E). 
0.1-0.2 m, M. Norman, S. Troy, 17 Oct 1 990 (active 
on reef flats between 0100-0200 hr, handnet); 2<S: 
31.1, 61.1 mm ML, AM CI59267. Torres Strait, 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

285 

286 

MARK D. NORMAN 

3 

X> 
3 

O 

a 

u 

"1 

o 

LV-S 

63 

CO 


^ c 

4> 1— 1 

S « 

« .2 
.a 11 

cd 

M 

ea 00 

<" 
U ^ 

■* cd 

*j <3 

.0 -a 

r ^ 'en 

h c 
o 
o 

u 

a 

cd 

S 

w 

r- 
cn 

ON 

in 

u 

> NO 
7 ° 

Z ° 

Ph 

m 
cn 

ON 

in 

U 

Ph 

> 00 

4n NO 

>5 

2 o 

Ph 

ON 

in 
in 

NO 
P_ 

> CN 

2 o 
Z r - 

Ph 

o 

z 

00 
u 

y^r~<noo^ONCNcnNO^ON^o^^NOONOoininincNooincNcnr~-_ONQQ 

00 m(N(N-rf — 1 

— rx ■* "* 

li| , ?0\OOMON't"t(MOO»M(NOOONOOOOOM5> 

on «^ rf _•„■_■ .„■ : ^ „• _j *-: ^ - ^; — rn „• ,-; 00 

._ O cn on m — 00m \t or^fM 
m cncnOoocnr~mm — 1 
— 1 cn m cn — 1 cn 

_. <^ 

OO CN NO ^T 
CN CN m 

[jjr/2>oOrn— 'r-voOOfNooONfN— iioOt~-vOfNoo<nc3N 
00 ?^ONNdrn<^>nrNiin^'oc»o^S'noo 00 TfrnaN^t : 

cN "*d" 

r 1 

in 


— o 

<N 

On in 

(j.) c/D on cn — r^rNrN-^-NDinooONOOONNDP-rNicx^inoin 

O 
in 

^d- 00 — ooONOOin— 1 
Tt cn cn — 1 cn 

_h ^ 

t o — -<d- 

:s 

•<d- 

r- in 00 no cn cn 

JT; cn 00 cn cN r~ 

' ' no in ON On O 

]Q— ;OOncn— <0-h— ;Q 

00 cn on cn cn 
OO — — — 

CN CN. 

CN 

^ r--' on O 00 

cn cn >— c m 

On in no cn 
o on o no 

O Tf OO NO 

-too 

cn r- a C 
tj- in 1— 1 1— 1 

m no t~- m 
■d- rcit-^ 't 
^d- in r- in 

cn 00 cn O no 

00 r~ ■* -*d- 

Tf ^f 00 in 

Tt 00 00 — NOoocN^d-in— cNONDin — iONTj-NOcno\NOcncNi — vom^ 
-h cn _• ^ 00 

cn *J — 1 00 r- 

cNC^inincnONincNONO^- — — "U.'OnO 

no in V cn ^t — 1 cn 

t— o cn — 1 cn cn -<d- 

cn cn no 00 in 

r- o "<d- no 
in no r- in 

-^•50cNcncncnr—oooNinoONNO — OON-^-^d-oomcNoo 

CN 

<d- 

S2 

r—NOooOONOcnr- — 1 — 1 — 1 
no rf ■* cn cn 

NO O Tf O 
CN CN NO 

cn cn —1 -h cn 
O no 00 tt 

NO ^f NO NO 

cn w cn on rf r~- 

. • ■< 00 
CN 

Ooooor--Or---H — 1 

cn 

cNr~~ooocN^f^tr-r--cn-- 

t--int--cncNO>nm— 1 — 1 <-< 

no in tj- cn ■* 

— ■ On r-- On 00 — 1 cn -^- in — ■ in 
CN 

■* CN 

in no 

CN 

NO On — 1 r- in CN 

cn cn in r- 

tMO^^M^OO- '— 'r--incNOr--— 1 

cn 

'n 
CN 

cn On cn in o r^ r^ 

cn r- cn — 1 

r-moooNOO-d-r-— .— ._ 
in in ^- cn cn 

• — 1 CN 
CN 

r» o 

■* NO 

cn 

c~- "<d" cn -rt in cn 

N CO 

in -rt cn — 1 
cn — 1 in 

— ( — < — < CN ON 

— < O cn 00 

NO NO t~~ NO 

—_ O ■* O 

OO NO On ON 

in in no "d" 

g — 1 cn cn K -3- 

u 

S — , C/3 „ J „ " i> 
l> O W5 

<<JJOJ a^ a a J E 5 O 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

287 

o 

c 

Q 
c 

i, ȣ 

O 00 
C 00 

■a 'c 

5 o 

a 1 

s — ' 
B. II 

£ a 

E I 

« «3 

y oo 

So 

g U 

IS 

TD OS 

o ft 
rt -g 

in .-h 

C C 
3 O 

o o 

U u 

in c 
U g 

X) 

H 

UJ 

3 

&0 

7 ° 

z ° 

<^ NO 

>3 

4h no 
Ph 

no 

Uh 

> "* 

U. no 

Zm 
NO 
Uh 

Zm 
SO 

U- 

m no 
2 no 

Zin 
no 

Uh 

> 

z 

© 
o 
r- 

no 

u- 

o 

Z 

00 

u 

a! 

— i C/3 On 00 NO NO NO 
CO 

NO 

On On On r~- 

tI- m r- 

in 

r-r--NOTi-L/->or--* 

<N| U-) — I — I tN — ' 

O t-- — ' IT) o 

"* oo oo od r- 

" * l#~v -^- /-** *f-v *-»- ^-1 V/~> i i *■ ^ 

NO 
in 

Or--<Nooinr<->r--rnoo^r 
Tf rn <N (N 

£T r^ • ■••_••• • • • -h -rf- 

n^-^ooo^ ^ ' — i on ■ — i 

o 
in 

Tf -st 

os no — i m m — i 

>t * rnrn 

0Niy30000CN|ON0ininininr--O — no 
in ""io-<doo-i(N(N-ioddCS 

NMmTt-.0\'JQQOOr 

rn 9°, oo on o <n 
-3- n rt moom 

in 

-H (N NO ON 

m — i 
m 

— © 

OO GO On <N in © VO 00 © t~- vq © "* © 00 

On 
en 

inr-in^— i^oomoocNj 
— i ^r m m cni 

O&ONO-HOOmoo-^-fN— _rnr-~;— ;-h 
on ^Jr^— '©©r-'NO^i — r^o"" 

m "viNf Mm^NtO-- - 
m m m r»-> 

© 
rn 

Ttc^moNON©r-NO-.<?NrNiNor«-i-H^ 
^A ^r^'^oin'oo'oONONOON— i ^ 
rn ^inmCKNyON-^ — < 
_ in rr rn rn 

°°^ Q Q 
NO — c c 

PlNt" 1 - 1 

m no oo cni 

On — "' tf — >' 

tt in no >n 

in oo ^r Cn) 

tJ- no oo in 
in oo in -t 

O Cn) — < CN] 

O — < no rn 
in rf no in 

© no fN "=t 
rf od tN -h 
rt cn) r~ in 

-NQQ 
in ro C C 

Tj" ^1- M h-4 

no rn on no 

— ! -(T — — < 

in in no in 

I— i — i <n <~n '* 

288 

MARK D. NORMAN 

5ffwn 

►————« 

50mm 

5mm 

Figure 9. Octopus dierythraeus sp. nov. a, dorsal view of 1 35.0 mm ML male (holotype, NM V F67007). b, funnel 
organ of 37.5 mm ML male (NMV F60134). c, copulatory organ of 135.0 mm ML male (holotype, NMV 
F67007). 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

289 

10mm 

Figure 10. Octopus dierythraeus sp. nov. a, digestive tract of 60.7 mm ML male (paratype, NMV F60133) 
[abbreviations as in fig. 2a]. b-d, beaks of same specimen, b, upper beak, lateral view, c, lower beak, lateral view, 
d. lower beak, ventral view. 

Thursday I., 10°35'S, 142°13'E, G.P. Whitley, 1928; 
lcf; 35.5 mm ML, NMV F60132, Orpheus I., Pioneer 
Bay, 1 8°36'S, 146°29'E, 0. 1 m, M. Norman, 6 Oct 1 990 
(active at 0410 hr, handnet); lcf: 36.5 mm ML, AM 
C164181, Lindeman I., 20°27'S, 149°02'E, G.P. Whit- 
ley, 1935; 2d: 37.5, 48.6 mm ML, NMV F60134, Sar- 
ina Beach, 21°24'S, 149°19'E, 0.1-0.2 m, M. Norman. 
1 6 Sep 1 990 (active between 0305-03 1 5 hr, handnet); 
lcf: 43.2 mm ML, NMV F601 31, Orpheus I., Pioneer 
Bay, 18°36'S, 146°29'E, 1 m, M. Norman, 8 Oct 1990 
(active at 2005 hr, retreated to lair, flushed with 
CuS0 4 ); 1$: 43.2 mm ML, NMV F60138, Cape Hills- 
borough, 20°55'S, 149°05'E, 0.1 m, M. Norman, 18 
Sep 1990 (in mouth of lair at 0430 hr, flushed with 
CuS0 4 ); I9: 45.3 mm ML, NMV F60136, Bowen, 
Cape Edgecombe, 20°01'S, 148°15'E, 0.1 m, M. Nor- 

man, 20 Sep 1990 (active at 0400 hr, handnet); 29: 
50.5, 52.5 mm ML, NMV F60137, Magnetic I., Geof- 
frey Bay, 19°09'S, 1 46°52'E,0. 1-0.3 m, M.Norman, S. 
Troy, 4 Oct 1 990 (active between 0230-0330 hr, hand- 
net); lrf: 53.1, 19:77.1 mmML, USNM 817785, Gulf 
of Carpentaria, offWeipa, 12°35.4'S, 141°36.2'E, 2 m, 
G. Hendler, "Alpha Helix" stn M-18, 4 Jun 1979; I9: 
54.6 mm ML, NMV F60135, Magnetic I., Horseshoe 
Bay, 19°07'S, 146°51'E, <0.1 m, M. Norman, J. Mar- 
tin, 9 Nov 1989 (within lair at 1200 hr, flushed with 
CuS0 4 ); lcf: 66.6 mm ML, NMV F60130, Orpheus I., 
Cattle Bay, 18°34'S, 146°29'E, 1.5 m, M. Norman, 9 
Oct 1 990 (within lair at 1 600 hr, flushed with CuS0 4 ); 
lcf; 90.5 mm ML, MNHN 4.2.853, Torres Strait, 
Thursday I., 10°35'S, 142°13'E, M. Lix, No. 27, 
1891. 

290 

MARK D. NORMAN 

a 

10mm 

20mm 

Figure ll. Octopus diciythnicus sp. nov. a, reproductive tract of I35.0 mm ML male (holotypc, F67007) 
[abbreviations as in fig. 3a], b-c, spermatophore from same specimen, b. whole spermatophore. CB = cement 
body; EA = cjaculatory apparatus; OC = oral cap; SR = sperm reservoir, e. detail of oral cap. 

Figure 12. Octopus dierythraeus sp. nov. a, active individual at night on reef flat on Magnetic I., showing rest- 
ing/foraging colour pattern, b. intermediate colour pattern of 54.6 mm ML female (NMV F60 1 35), showing first 
signs of red spots within white spots, c. alarm colour pattern of same specimen, d. alarm colour pattern of 66.6 
mm ML male (NMV F60 1 30). e, dorsal view of preserved 59.8 mm ML male (NMV F60I29). f, lair of 66.6 mm 
ML male (NMV F60 1 30). g. lateral view of preserved 60.7 mm ML male (NMV F60 1 33). showing pale spots each 
containing a central papilla, h-i, radula of 80.7 mm ML female (NMV F60133). h, dorsal view, i, lateral 
view. 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

291 

292 

MARK D. NORMAN 

WA: lc?:71.6mmML,NMVF67013,offKimberlev 
coast, 13°12'S, 125°03'E, 77-78 m, C.C. Lu, FV "Hai 
Kung", 30 March 1981 (on sand). 

Diagnosis. Large muscular species with unequal 
arms, dorsal arms longest and most robust (AF 
1.2.3.4), typically 4 to 5 times mantle length. 
Webs moderately deep, approximately 20% of 
length of longest arms. Sucker counts, 200-280 
per arm in submature and mature animals, 1 00- 
125 suckers on intact hectocotylized arm of sub- 
mature and mature males. Gill lamellae, 12-14 
per demibranch, typically 13. Funnel organ UU- 
shaped. No mature females encountered, sub- 
mature females with large-type eggs, produced 
in low numbers. Alarm colouration of white base 
colour and red circular spots on dorsal body, 
arm crown and arms. 

Description. The following description is based 
on 1 mature and 5 submature males, and 1 
submature/maturc female and 4 submature 
females. Counts and indices for material exam- 
ined presented in Tables 6 and 7, with data for 
immature specimens (4cf, 3$). 

Moderate to large robust species (fig. 9a): ML 
to at least 135 mm for males and 120 mm for 
females, TL to at least 8 1 mm; weight to at least 
1.5 kg. Mantle from round to ovoid, never 
greatly elongated (MWI 62.2-69.7-80.3), 
mantle walls moderately muscular. Stylets pres- 
ent, poorly developed. Pallial aperture of mod- 
erate width, approximately half mantle width. 
Funnel long, muscular and broad based (FLI 
43.6-60.9-72.8) with free portion usually 
greater than half funnel length (FFI 36.9-56.S- 
64.8). Funnel organ well developed (fig. 9b), 
always UU-shaped with broad limbs, outer 
limbs slightly shorter than median limbs (FOI 
16.2-83. 6-92J). Funnel organ approximately 
50% of funnel length (FOLI 42.2-51.8-62.9). 

Head of moderate width (HWI 46.8-5S.6- 
71.1), typically narrower than mantle (HMWI 
69.3-84.2-107.1). Neck distinct, slightly nar- 
rower than head. Eyes large and slightly 
pronounced. 

Arms moderate to long, dorsal pair longest 
and typically 4 to 5 times mantle length (AMI 
365.7-477.5-574.0) and robust (AWI 11.3- 
79.3-25.3), roughly square in cross section and 
tapering evenly along length. Arms unequal in 
length, dorsal pair longest, ventral pair shortest 
(AF typically 1.2.3.4). Suckers of moderate to 
large size (SDI 13.0-75.5-20.2), deep with dis- 
tinct radial cushions and scalloped outer rim, 
scalloping pronounced on small distal suckers. 
Suckers largest on dorsal arms, none especially 

enlarged in either sex. 200-280 suckers on intact 
normal arms of larger specimens (SC 234-259- 
280, n = 5cf, 39). Webs of moderate depth (WDI 
1 7.6-20. J-27. 8), deepest dorsally receding ven- 
trally to shortest ventral web (WF typically 
A.B.C.D.E). Web margins extended on ventral 
edges of arms for most of length. 

Only 1 mature male ( 1 35.0 mm ML holotype) 
encountered. Hectocotylized arm of holotype 
almost fully regenerated, opposite arm dam- 
aged. Right third arm hectocotylized and 
slightly shorter than opposite arm in submature 
specimens [OAl(submature) 69.2- 73. 9-77.0, n 
= 4; HAMI(submature) 234.6-264.9-298.2]. 
Copulatory organ of mature male large [fig. 9c: 
LLI(holot\ pe) 6.4 on regenerating arm], roughly 
cylindrical with deep ligula groove and small 
sharp calamus [CLI(holotype) 16.6]. Spermato- 
phorc groove well developed with fine trans- 
verse ridges. Regenerating hectocotylized arm of 
holotype with 93 suckers, approximately 112 
suckers in submature males [HASC(submature) 
103-772-125]. 

Gills with 12-14 lamellae, typically 13 on 
each demibranch. plus small terminal lamella. 
Digestive tract illustrated in figure 10a. 
Anterior salivary glands approximately 40% of 
buccal mass length. Posterior salivary glands 
approximately equal in length with buccal mass. 
Crop diverticulum well developed, approxi- 
mately 70% of length of digestive gland. 
Stomach bipartite. Caecum clearly striated, 
coiled in 1.5-2 whorls. Intestine thin walled and 
reflexed approximately one-third from proximal 
end. Rectum straight and muscular. Ink sac well 
developed, embedded in ventral surface of 
digestive gland. Ink red when released by live 
individuals. Anal flaps present. 

Upper beak (fig. 10b) with small hood and 
slightly hooked rostrum. Lower beak (figs 10c- 
d) with moderately blunt rostrum, narrow hood, 
widely spread wings and flared lateral walls. 
Ventral view of posterior margin of lateral walls 
deeply concave. Radula with 7 transverse rows 
of teeth and marginal plates (figs 12h-i). Rha- 
chidian tooth has 2-3 lateral cusps, typically 3, 
on each side of large curved medial cusp. Lateral 
cusps typically in symmetrical seriation, 
although seriation asymmetrical in 1 specimen 
(52.5 mm ML ? , NMV F601 37). This specimen 
differs from other material in no other character. 
The symmetry' of seriation does not appear 
fixed. In all specimens, lateral cusps migrate 
from lateral to medial position over 9-10 rows. 
First lateral teeth small and unicuspidate; sec- 
ond lateral teeth unicuspidate, of moderate 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

293 

u. 

3 

B 

o S 

c .5 

2 <D 

3 y 

?5 3 

M . - 

E o 

h,a 

•2.2 

•3 ii 

■So 

13 C 

C " 

C/} I— 

O E 

° a 

SO "" 
O » 

S E 

h a 

■o 

<u 

00 
03 

C3 

■o 

Z ^ & 

<*-> so o 

^ CM 

Z 5 

> 2 
z s 

<- sO 

> 2 

z 5 

*-H SO 

z ° 

Ch so 

> St 

^ m 

Z 5 

^H SO 

> <2 
^~ so 

> 3: 
z ° 

« OO 

so 

U 

z 5 

o 
Z 

ob 

PC 

O5Orvjr~ i ^oo^Q^r^voTffN'i-Op^Q0^o^ , >ot~- ; oosqw-ifN'^- 

•n" Hcid-'t^OO t^ N f) * OS C 1 ?! t~- Clt^o<tOSiO-<'-< 

<N o —I r— oo 
so m oo r- -*j- 

r- fN m tj- os 

„ OO (N -h -, m 

gro - 

m 

o 

so 

■^ — i (N in t-~ , '* 

-m ^ 

oo _; so 

-st ON 

r- ro 

CO — i t~~ OS 

V) o rsi rsi 

rf r- os so 

c/^oooooomosot— ■osinmot--rorsiosrsit--osoor- 
so ,,-, t^. /— », ,^1 , ^i _j. _j_ , -— r- — *—■< r- 

sornotN — i rsi "st ■<♦ — i m t-~ 

in ^t m (N ro 

r~) ** as 

04 so so 
rsi 

rsi m os oo 

*W00ifl!N^ , i'10>fl00Cl/lO(NC1* 

so rrmTtinm— 'Tto^ostnos 

SO 'sDi/IMinaiNinON-H-i-' 

— SO 

M/l-i-nOnmQQN*OvmQm^(NQini~;qQQ 

o 

SO 

r- (N m os 

SO sO Os OO 

oo m ■* >n oo 
os r- rsi — i 
<N m 


2! r4 in c 

rsi « 

A 

oowoM Y iONoooNOMJvosair*inooQ(NO;-|oo-;i r iitt-- 

5 in rsi 2 ^ fi ■* r "-' 

O oo in — OMNinMinOO 
oosooososooot~-r--r-ifNrsi 
in "* m <N m 

so C/3 O m 
od "~> 

•^-r— — t--insosor~rimrn 

>, l-^ O ■* r~ r— ' ■* ■* - ' '•O fi rsi i C^ 

"'soinr^os^-or^r-— ■— 'r-irsi 

m m t<i m m — i 

O 

c m O ro os Tl- 

Oinr*^ — sosO'J'ro 

£:; os (N od — < — < ^t "st O Os in O ^ 
™ r-sor^osOininm — ■ — fN cn 

1—1 ^^ClNn —i 

^do'in'^tsO^fdvDoo'iNriC 
""■oosor-oom — mos — -^ Mm 

cmoorsimsoo^rp— ; — pinm 
g 1 ^-— isdod— ittodd- iin— ' ui C 
C~'ininOsor--r^iO"* — ' — ' — '^l 
i— i -. r»i rl M ts (N — ' 

CO — . Tl" 

so o in — i ci oo os 

"* t-- 2 r4 c*S r-' t> 

ro t~~ "" rn in 
rsi 

t — C: — c c c 

in r~ mmm 

rsi 

O OS 2 — ' c c c 
in r— — <-* i— i 

rs 

Om" C Cm 
O oo i— i i— i 
rsi 

m g rsi tT r<-i o ^T r-; ; rf — < O rs] I — ; ; rn rn 

ir; 5 — d*d<>oor-ivdodo>*t^C 

r*-, C rs| ^mxmoomr^oo-H-'-'N 

i— i Tt m m <N m — ' 

- - oo oo oo Q Q 
so rn 2 — f~~ C C 
r~ oo rsi " — 

— < f~ G G 

s5m«M 

oo ro rn r- oo so rsi 

"*«QQ 

O r-^ ^j rsi r-^ oo rn 

O rf G G 

mh" m so 

so ^r — < Hi 

<N 


os — os rsi 

os — ^ os rsi 
so in oo ^r 

so r-~ n oo 
r- r-- so in 

lOini^- \t 

— i os oo in 

os o rsi o 
in so oo in 

— rsi o as 

rn so in -^ 
so so r- ■* 

in in ro m 

so — < oo ^}- 
in in oo Tf 

QQQQ 

n rsi — os 

os in od — ■ 
in so oo in 

i— i — i rsi rn 


H-UOJaaaa-lU-OO 

J U H D M 73 M V5 n t t Ph 

294 

MARK D. NORMAN 


> 


11) 


s 

a 


z 

nO 

si 

C/3 

5n 


Ph 

eg 

0- 

<u 



w 

c 




ca 








Vh 




ca 




> 




O 

> 

m 


l-H 

s 

C 1 


3 

z 

O 

NO 


3 


fc 


> 

<A 

> £ 

O 

3 

S'^ 

r 

<u 


s 

7 ° 

^ NO 
Ph 

t>5 

3 


r^ 

c/5 


Rl 

3 


3 

s 


-. 

II 

> £ 

tu 

* 

g- 

5s 


7 ° 

<-i NO 

3 

3 

P. 

to, 

ks 


a 

s 





O 



— 
S3 

3 

3 

s 

2^ 

<S 

3 

z ° 

^ *o 

— 

w 

Ph 

■H go 


"O . * 


.2 o 

-a .5 

§2 

§3 

7 ° 

^ no 

«> "2 

P. 

■ii 3 


c — < 


=> II 


o 


UQ 


. 3 
r-~ i— i 

> oo 

1 & 

H 2 

z ° 

^ no 

Ph 

S 


6 


• OH 


II 


S 

«5 oo 

fl 

< <N 


ON 


IT) 

"0 

— H 

00 

u 

03 


s 


3 


"O 


II 


Q 

e 4 

^z 


4> 



oo *J m in oo m ro 

oo $ IT* r~' no On (N 

Q 

in -- ^h r-. no m 

m O on in r— 

in Tt -h — i h 

* * o no on on in 
°^ °^ £> rn no od r»' 

—i — Y/ ■* ^ f"- ""> 

vOWin^oOMOO | nf f i' , liNT^ 

JS n t Nt ci d --<-'■*' ni rt 
tC h~> m ro 

it 

NO 

2 r-° (N © oo en cn r-' t - r-° oo C 

'r^NooooN-NTNOin-H-H— m 

*t *t m m h 

in 

in 

CNOTtl^l^ — — m»lOVOh 

M 

m 

cJJiodoNrNJr^oOTror-'rNiod 
-3- m m m 

r-"-ONinr^t^-^-ooONTtin-Hr<-iNO 
^~ONHH*coincN|CNiooodc'"ioQ''"' 

VD in 1^ N \D M O — i — i — ' 

it m m fri 

rs 

fN 

rn c/3 "* ON -h t~~ 
9 ^ 

OO^ooinTf-Hmo 

in — ' rn Tt ON On — < -h [-.' 0O ~* Ifi 

r^r~ONt^r 1 o — ■ -h — <N 

m tt Tt tt 

(NWON^OM-'^vOCTiOr'imoOrONt 

-h ni 

i N in ^n rN \t~i i — s ^n i/-\ /vi . /^i i " 


^ONt^ininroO'^tinr- 

NONOONOONOincNi-HrNi 
-immMN 

rO C-^-^(Nl^iOOiOi-MXlOm^ 
O c^2in'r^— ^rNirnin'in— .'in^^TJTi 
N B osooONONOmOiN — rvicNi 
* t m m _h 

no oo tt in 
r-' rt ^ —.' 

no no oo in 

OO On 3 3 

NO in HH HH 

in ^- m on 
(N«dd 

NO t ON rt 

NO in On Cn| 

On On —• On 
m\|-ON \t 

00 O — f*N 

(N-r<ld 
r~ no oo in 

(n|M)0 \t 
it © O rn 
in no oo in 

O — < ON Tt 

no ro r^ in 

^ -h <n m Tt 

2 c/3 H 2 K X < 

^5>^U J^Z jEoo 

<c^>OcOPJPJPJPhPhP-P- 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

295 

length and robust; lateral marginal teeth rela- 
tively straight, short and robust; marginal plates 
oblong and plain. 

Male reproductive tract illustrated in figure 
1 1 a. Terminal organ in mature male of moderate 
length and robust [TOLI(holotype) 24.7] with 
robust diverticulum (DLI(holotype) 59.8). 
Genital aperture subterminal. Mucilaginous 
gland enlarged at point of attachment to vas def- 
erens. Spermatophoric gland I narrow with large 
recurved coil approximately 80% along length. 
Spermatophoric gland II elongate and narrow 
with coiled tip. Spermatophores (fig. llb-c) 
approximately three-quarters of mantle length 
(SpLI 74.8. 75.6). narrow (SpWI 1.3, 1.5) and 
produced in low numbers (4 in Needham's Sac 
of holotype). Oral cap simple bearing long cap 
thread. Sperm reservoir approximately 40% of 
spermatophore length (SpRI 39.6, 41.2), sperm 
cord coiled in approximately 30 whorls in best 
condition spermatophore. 

No mature females encountered. Immature 
ovaries were visible in submature females, pos- 
itioned on the right hand side of the septum 
midway along visceral mass. The ovary of the 
largest female (118.8 mm ML, NMV F60128) 
occupied approximately 20% of visceral mass 
and contained approximately 350 long thin 
immature eggs up to 14.1 mm long[ELI(Imm)to 
1 1.9; EWI(Imm) to 1.9]. Approximately 20 fol- 
licular folds visible on immature eggs. This 
species clearly produces large type eggs in 
moderately low numbers. 

Colour in life variable, 3 distinct colour pat- 
terns displayed most frequently. Foraging ani- 
mals have orange base colour with mottled 
cream and red/brown patches (fig. 12a). Dis- 
turbed animals have red brown base colour with 
white circles each containing a central papilla 
(fig. 12b). This pattern is similar to that of O. 
alpheus (fig. 4a-c) displaying the same large 
transverse pair of white spots on dorsal body. 
This pattern is an intermediate stage in attaining 
the full alarm display. Alarm display of the nega- 
tive pattern: white base colour and circular red 
spots over dorsal body, arm crown, webs and 
arms (figs 9a. 12c-d). Iridescent tissue layer in 
the skin produces an iridescent green sheen in 
torchlight at night. 

Skin sculpture simple, consisting of scattered 
low papillae with smooth tips, located in the 
centre of white spots in intermediate colour pat- 
tern and centre of red spots in alarm pattern (figs 
9a, 12c-d). Smaller punctae occur over dorsal 
surfaces between papillae. Single slightly larger 
supraocular papilla directly above each eye, sur- 

rounded by low punctae. Oral surfaces of all 
webs smooth and unpigmented. 

Preserved specimens lose darker red colours 
and definition of spots, base colour fading to 
pink or cream. Ventral surfaces smooth and 
scattered with small fine chromatophores. 2-3 
dark wavy transverse lines often visible across 
dorsal arm crown. Pale spots containing central 
papilla sometimes visible on mantle, especially 
on lateral faces (fig. 1 2g). 

Sexual dimorphism was not marked in the 
material examined. 

Distribution. Coastal waters and inshore islands 
of the Great Barrier Reef and northern Aus- 
tralia, from Sarina Beach (21°24'S. 149°19'E) in 
the south, north to Thursday I. (10°35'S, 
142°13'E) and west to the Kimberley coast, 
northern Western Australia (13°12'S, 125°03'E) 
(fig. 17d). Specimens were collected from 
exposed intertidal reefs and subtidally to 78 m. 

Life history. Octopus dierythraeus is a noctur- 
nally active species which forages on intertidal 
rock and mud flats, and shallow subtidal habi- 
tats in coastal muddy waters and round inshore 
islands. 

This species is an active predator, observed 
probing amongst crevices and rubble mainly 
with arms 1 and 2 on exposed reef flats at night. 
Active animals were encountered carrying fresh 
bivalves, crabs, a large polychaete worm and in 
one case a decapitated fresh octopus of indeter- 
minate identity. Active lairs were surrounded by 
clean bivalve shells and carapaces of small crabs. 
Large strombs were also found at two lairs. One 
specimen flushed from a lair at Orpheus I. (66.6 
mm ML male: NMV F60130) had accumulated 
the remains of over 100 bivalves. Only one small 
crab carapace was found amongst this midden. 
Octopus dierythraeus has had a significant influ- 
ence on James Cook University's Giant Clam 
(Tridacna spp.) mariculture project on Orpheus 
1.(1 8°36'S, 1 46°29'E). The empty shells of young 
clams were regularly found around lairs of this 
species (P. Lee, pers. comm.). Captive individ- 
uals readily take frozen fish (J. Hoey, Reef 
Wonderland Aquarium, Townsville. pers. 
comm.). 

Octopus dierythraeus occupies lairs amongst 
rocks on muddy substrates and within coral bed- 
rock, closing the entrance during the day with 
pieces of dead coral. Lairs are permanent or at 
least long term, judging by the large number of 
prey remains (primarily bivalve shells) sur- 
rounding lair entrances. Animals only emerge 
after dark. 

296 

MARK D. NORMAN 

The large eggs of this species indicate benthic 
hatchlings. 

Etymology. From the Greek " dierythros" mean- 
ing spotted with red, referring to the red spots 
generated in the alarm display of this species 
(fig. 9a, 12c-d). 

Octopus graptus sp. nov. 
Figs 13-16, 17c 

Material examined. None encountered live in the 
field. 17 specimens obtained from commercial prawn 
trawl operators, now in the Museum of Victoria. 10 
preserved specimens were found and examined in the 
collections of the Australian Museum, Northern Ter- 
ritory Museum of Arts and Sciences, and Museum of 
Victoria. 

Holotype: Qld: 1<?: 88.8 mm ML, NMV F67006, 
Cleveland Bay, Townsville (19°1I'S, I47°01'E), A. 
Cabanban, 1 2. Ian 1990 (trawl). 

Paratypes: Qld: 1?: 107.0 mm ML. NMV F67008, 
Gulf of Carpentaria, 16°09'S, I38°51'E, 32 m. QDPL 
grid 6188, port stn. 8, shot 3, 1 2 Apr 1 983 (at 02 1 hr); 
Irf: 1 1 1.6 mm ML, NMV F67009, 0.3 mile (0.6 km) 
south of Lucinda wharf (~18°32'S, I46°22'E), 15-17 
m, D. Sutton, 12-18 Nov 1989 (prawn trawl on 
sand). 

NT: I9: 116.1 mm ML, NTM P1478. W of Orontcs 
Reef, olTCobourg Peninsula, 1 1°06.0'S, 132°04.3'E.C. 
Johnson, 10 Aug 1990 (trawl). 

Other material: Qld: 2rf: 68.2, 76.9 mm ML, 89: 
57.3-86.9 mm ML, NMV F67009, 0.3 mile (0.6 km) 
south of Lucinda wharf (near 18°32'S, 146°22'E), 15- 
17 m, D. Sutton, 12-18 Nov 1989 (prawn trawl on 
sand); I9: 75.6 mm ML, lrf: 92.0 mm ML, AM 
CI 70702, W of Fitzmaurice Point, SE Gulf of Carpen- 
taria, 17 o 10.5'S, !40°30.7'E, 13 m, 1. Loch, Dec 1976 
(trawl); lrf: 85.4 mm ML, NMV F67012. Gulf of Car- 
pentaria, I0°59'S, 140°29'E, stn 058, SS05/91, 29 Nov 
1 99 1 (trawl, 0400 hr); I9: 87.8 mm ML, NMV F67003, 
50 km west of Port Musgrave, Gulf of Carpentaria. 
1 2°07'S, 1 4 P27'E, 36 m, C.C. Lu, FV "Susan Wright", 
16 Jun 1982 (prawn trawl, 1930-2030 hr); 2rf: 23.6, 
157.9 mm ML, 4?: 104.8-190.8 mm ML, NMV 
F6701 1, East side of Cape Yorke, somewhere N of 
Cairns (no exact locality data), commercial prawn 
trawler, 1989 (purchased from Rosslyn Bay Fisher- 
man's Co-operative); lrf: 111.3 mm ML, AM 
C 1 7070 1 , W of Fitzmaurice Point, SE Gulf of Carpen- 
taria, 17°12'S, 140°37.8'E, 11 m, 1. Loch, Dec 1976 
(trawl). 

NT: lrf: 76.3 mm ML, NMV F67004, Off Groote 
Evlandt, (about 14WS, 137WE), 29 Nov 1980 
(prawn shot, 2400 hr). 

WA: I9: 40.6 mm ML, NTM P015795, Joseph 
Bonaparte Gulf, UWWS, 128°08'50"E, 27 Jun 
1990. 

Diagnosis. Large muscular species with unequal 
arms, dorsal pairs longest (AF 1.2.3.4), typically 

4.5-7 times mantle length. Webs moderately 
deep, approximately 20% of length of longest 
arm. Sucker counts, 200-280 per arm in sub- 
mature and mature animals, 86-88 on hecto- 
cotylized arm of submature and mature males. 
Gill lamellae, 13-14 per demibranch, typically 
13 on outer demibranch and 14 on inner demi- 
branch. Funnel organ VV-shaped. Eggs large (to 
28 mm long), produced in moderately low num- 
bers (680 in mature female). Colour pattern of 
pale cream to pink base colour with dark pink 
brown irregular scribbling over dorsal mantle 
and arm crown. 

Description. The following description based on 
2 submature, 1 submature/mature and 1 mature 
males, and 1 mature and 4 submature females. 
Counts and indices presented in Tables 8 and 
9. 

Large robust species (fig. 1 3a): ML to at least 
1 90 mm, TL to at least 1 300 mm; weight to at 
least 4.2 kg. Mantle round to ovoid (MWI 47.4- 
59.5-76.8), mantle walls thick and muscular. 
Stylets well developed. Pallial aperture of mod- 
erate width, slightly greater than half mantle 
width. Funnel long, muscular and broad based 
(FL1 40.9-5tt 7-66.6) with free portion variable 
in length, typically around half funnel length 
(FFI 20.9-5 /.2-60. 1). Funnel organ moderately 
developed (fig. 1 3b), VV-shaped with narrow to 
medium width limbs. Outer limbs considerably 
shorter than median limbs (FOI 56.3-62.1- 
70.4). Funnel organ approximately 40% of fun- 
nel length (FOLI 39. 2-40.8-44 J). 

Head of moderate width (HWI 33.1-42.9- 
60.2), typically narrower than mantle (HMWI 
55.5-72.2-92.6). Neck distinct, slightly 
narrower than head. Eyes large and pro- 
nounced. 

Arms long, typically 4.5-7 times mantle 
length (AMI 446.9-5^7.9-706.8). Arms robust 
(AWI 15.1-/9.6-26.2), roughly circular in cross 
section and tapering evenly along length. Arms 
unequal in length, dorsal pair longest, ventral 
pair shortest (AF typically 1 .2.3.4). Suckers large 
[SDI(femalesand submature males): 1 3. 1- 14.5- 
16.3], larger in mature male [SDI(mature male): 
21.5], but no individual suckers specially 
enlarged. Suckers deep with moderate flared 
rims, fine radial cushions and scalloped outer 
rim. Suckers largest on dorsal arms, none 
especially enlarged in either sex. Approximately 
240 suckers per intact normal arm in both sexes 
(SC 194-240-280). Webs moderately deep 
(WDI 16.4-20.0-22.0). deepest dorsally 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

297 

10mm 

50mm 

5mm 

Figure 1 3. Octopus graptus sp. nov. a, dorsal view of 88.8 mm ML female (holotype, NMV F67006). b, funnel 
organ of 87.8 mm ML female (NMV F67003). c, copulatory organ of 157.9 mm ML male (NMV F67011). 

298 

MARK D. NORMAN 

— o 

PSG 

20mm 

Figure 1 4. Octopus graptus sp. nov. a, digestive tract of 87.8 mm ML female (NMV F67003) [abbreviations as in 
fig. 2a]. b-d, beaks of 1 57.9 mm ML male (NMV F670 1 1 ). b, upper beak, lateral view, c, lower beak, lateral view, 
d, lower beak, ventral view. 

decreasing evenly in depth to ventral web (WF 
typically A.B.C.D.E). Web margins extended on 
ventral edges of arms for majority of length. 

Right third arm in males hectocotylized, 
slightly shorter than opposite arm (OAI: 59.2- 
68.4-13.0; HAMI: 187.4-265.7-365.4). Copu- 
latory organ (fig. 13c) of moderate size 
[LLI(mat) 6.5], roughly cylindrical with ligular 
groove deep and closed. Calamus of moderate 
size [CLI(mat) 19.9]. Spermatophore groove 
well developed and wide with fine transverse 
grooves. Spermatophore guide not distinct in 
frozen mature male. Approximately 87 suckers 
on hectocotylized arm (HASC 86-57-88). 

Gills with 13-14 lamellae on each demi- 
branch, typically 13 on outer and 14 on inner 
demibranch. Terminal lamella small. 

Digestive tract illustrated in figure 14a. 
Anterior salivary glands approximately one- 
third of buccal mass length. Posterior salivary 
glands approximately equal in length with 
buccal mass. Crop diverticulum well developed 
in the specimen dissected. Stomach bipartite. 
Caecum clearly striated, coiled in 1.5-2 whorls. 
Intestine thin walled, reflexed in proximal third. 
Rectum muscular and straight. Ink sac poorly 
developed and elongate, ink duct long. Fine 
elongate anal flaps present. 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

299 

30mm 

20mm 

Fie.ire 1 5 Octoous xraotus sp nov. a, reproductive tract of 1 57.9 mm ML male (NMV F6701 1 ) [abbreviations as 
fn £ 3a 1 » soTrm Shorefrorn same specimen. CB - cement body; EA = ejaculatory apparatus; OC = oral cap; 
SR f 'sperm SS^Twrf 1 90.8 mm ML female (NM V F670 1 1 ). DO - distal ov.duct; O = ovary; OE = 
mature ovarian egg; OG = oviducal gland. 

300 

MARK D. NORMAN 

c 

$ 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

301 

Figure 16. Octopus graptus sp. nov. a, dorsal view of preserved 88.8 mm ML female (holotype, NMV F67006) 
b, dorsal mantle of same specimen, c, lateral view of preserved 1 16.1 mm ML female (NTM PI 478) d, dorsal 
view of preserved 40.6 mm ML female (NTM P015795). e-f, radula of 57.3 mm ML female (NMV F67009). 
e, dorsal view, f, lateral view. 

Upper beak (fig. 1 4b) with weakly hooked ros- 
trum, small hood and large lateral walls. Lower 
beak (figs 14c-d) with sharp rostrum, narrow 
hood, widely spread wings and flared lateral 
walls. Ventral view of posterior margin of lateral 
walls deeply concave. Radula with 7 transverse 
rows of teeth and marginal plates (figs 16e-f). 
Rhachidian tooth has 2-3 lateral cusps, typically 
3, on each side of large curved medial cusp. Lat- 

eral cusps in symmetrical sedation, migrating 
from lateral to medial position over 9-10 rows. 
First lateral teeth small and unicuspidate; sec- 
ond lateral teeth unicuspidate, of moderate 
length and robust; lateral marginal teeth curved 
and relatively fine; marginal plates oblong and 
plain. 

Male reproductive tract illustrated in figure 
1 5a. Terminal organ in mature male large and 

Table 8. Counts and indices for male Octopus graptus sp. nov. 
(D = damaged; InD = indistinct; M = mature; S = submature). 

Museum 

NMV 

NMV 

NMV 

NMV 

Reg. No. 

F67004 

F67009 
(Paratype) 

F67011 

F67005 

ML 

76.3 

111.6 

123.6 

157.9 

StM 

S 

S 

S/M 

M 

TL 

420 

778 

749 

1286 

MWI 

47.7 

65.9 

61.0 

76.8 

HWI 

35.9 

36.6 

35.0 

60.2 

HMWI 

75.3 

55.5 

57.3 

78.4 

AMI: 1 

446.9 

562.7 

D 

706.8 

2 

327.7 

465.9 

484.6 

589.6 

3 

256.9 

337.8 

449.8 

526.9 

H 

187.4 

243.7 

266.2 

365.4 

4 

249.0 

330.6 

368.9 

499.7 

AWI 

15.1 

17.6 

20.2 

26.2 

SDI 

13.1 

14.3 

16.3 

21.5 

WDI 

22.0 

19.1 

D 

D 

GC 

13/14 

13/14 

13 

14 

SC 

194 

220 

230 

280 

HAMI 

187.4 

243.7 

266.2 

365.4 

OAI 

73.0 

72.1 

59.2 

69.4 

HASC 

86 

86 

86 

88 

LLI 

1.5 

6.1 

10.3 

6.5 

CLI 

InD 

29.4 

26.0 

19.9 

TOLI 

6.0 

10.7 

12.1 

31.2 

DLI 

37.0 

67.2 

56.4 

82.4 

SpLI 




85.5 

SpWI 




1.9 

SpRI 




38.5 

SpN 




5 

FLI 

41.7 

55.2 

52.0 

66.6 

FFI 

44.3 

60.1 

34.8 

32.8 

FOI 

70.4 

InD 

InD 

InD 

FOLI 

39.3 

InD 

InD 

InD 

302 

MARK D. NORMAN 

Table 9. Counts and indices for female Octopus graptus sp. nov. 
(D = damaged; InD = indistinct; M = mature; S = submature). 

Museum 

NMV 

NMV 

NMV 

NTM 

NMV 

Reg. No. 

F67003 

F67006 

F67008 

PI 478 

F67011 



(Holotype) 

(Paratype) 

(Paratype) 


ML 

87.8 

88.8 

107.0 

116.1 

190.8 

StM 

S 

S 

S 

S 

M 

TL 

510 

587 

764 

679 

1229 

MW1 

59.3 

58.2 

56.9 

60.0 

49.7 

HWI 

47.4 

53.9 

41.5 

42.6 

33.1 

HMWI 

79.8 

92.6 

72.9 

71.0 

66.6 

AMI: 1 

479.5 

542.8 

574.8 

455.6 

534.1 

2 

381.5 

412.2 

474.8 

386.7 

410.4 

3 

325.7 

346.8 

396.3 

288.5 

370.0 

4 

289.3 

316.4 

356.1 

282.5 

D 

AWI 

15.5 

18.9 

21.6 

21.8 

19.5 

SD1 

13.2 

14.4 

15.0 

14.4 

15.1 

WDI 

21.4 

19.4 

16.4 

21.6 

D 

GC 

13/14 

13 

13/14 

13/14 

13/14 

SC 

237 

225 

265 

258 

249 

ELI 





14.7 

EWI 





17.8 

EN 





680 

FLI 

45.9 

49.5 

46.8 

52.0 

40.9 

FFI 

25.1 

24.8 

20.9 

25.4 

33.5 

FOI 

62.7 

56.3 

63.5 

57.4 

InD 

FOLI 

39.2 

41.6 

39.3 

44.7 

InD 

robust [TOLI(mat) 31.2] with robust diverticu- 
lum (DLI 82.4). Genital aperture subterminal. 
Mucilaginous gland enlarged at point of attach- 
ment to short and robust vas deferens. Sperma- 
tophoric gland I narrow and elongate with large 
recurved coil approximately 80% along length. 
Spermatophoric gland II narrow and elongate 
with reflexed tip. Spermatophores (fig. 15b) 
approximately 80% of mantle length in mature 
male (SpLI 85.5), of moderate width (SpWI 1.9) 
and produced in low numbers (5 in Needham's 
Sac of mature male). Oral cap simple bearing 
long cap thread. Sperm reservoir approximately 
40% of spermatophore length (SpRI 38.5). 
Sperm cord deteriorated in spermatophores 

examined, appears regularly coiled in approxi- 
mately 30 whorls in best condition spermato- 
phore. 

1 mature female specimen encountered 
(NMV F67011, 190.8 mm ML). Ovary con- 
tained approximately 680 almost mature 
ovarian eggs up to 28.0 mm long (ELI 14.7) (fig. 
1 5c). Eggs exhibited approximately 16 follicular 
folds. Mature ovary bore 2 large oviducal glands 
with around 26 braiding chambers. Oviducts 
robust and relatively short. 

Colour in life unknown. Preserved specimens 
exhibited a fixed colour pattern of cream to pink 
base colour with irregular fine "scribbling" over 
dorsal mantle and arm crown (fig. 13a, 16a, c). 

Figure 17. Distributions of Great Barrier Reef members of the Octopus maewpus group, a, Octopus omatus 
Gould, 1852: ▼ = type locality; ♦ = type locality of synonym Callistoctopus arakawai Taki, 1964; # = other 
material examined; A = additional published records, b. Octopus alphcus sp. nov. c, Octopus aspilosomatis sp. 
nov. d, Octopus dierythraeus sp. nov. e, Octopus graptus sp. nov. 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

303 

a 
























*«V 

? 

i 

j 






•< 

-v 

*• 

/ 









\ 

\* 



\ 

v 



-A 

r 

\ 

e 

^ 

[» 






•• 






^ 

N 


s 

W 

s 


\ 

y 

\ 

J 


% 


• 


i 

k 



T 





X 



2 




) 

\ 

* 

K) 

o 

• 
















• 


• 



^J 


Sf-.'. 

^>V 












.• 

Y, 








^ 

"X 

/\ 



• 
• 

■ . 



% 






/ 

u 

•" 






c 



s t 

¥ 




% 





• 










v. 

^— 

X 














40"N 

20°N 

20°S 

40°S 

140°E 

180° 

140°W 

10°S 

20°S 

30°S 

40° S 

h^ fS°'°° 

10°S 

20°S 

30°S 

40°S 

120°E 

140°E 

160°E 

120°E 

140°E 

160°E 

10°S 

20°S 

30°S 

40°S 

10°S 

20°S 

30-S 

40°S 

120°E 

140°E 

160°E 

120°E 

140°E 

160°E 

304 

MARK D. NORMAN 

In several specimens base colour had become 
pink brown, darkest in distal portions of arms 
(fig. 16a). 

Skin sculpture simple, consisting of scattered 
low papillae evenly distributed over dorsal sur- 
faces. Small punctae scattered between papillae 
on dorsal surfaces. Single slightly larger supra- 
ocular papilla directly above each eye, sur- 
rounded by low papillae. Pigmentation and 
sculpture do not extend to oral surface of 
webs. 

Sexual dimorphism was not marked in the 
limited material examined. 

Distribution. Open substrata in the coastal 
waters of northern Australia (fig. 1 7c), from 1 1 
to 36 m. Collected primarily in prawn trawls 
from Joseph Bonaparte Gulf, Western Australia 
(13°09 , 70 ,, S. 128°08 , 50"E) east and south to 
Cleveland Bay. Townsville (19°11'S. 
147°01'E). 

Life history. Little is known of the life history of 
this species. Where time of capture information 
was available, all specimens were captured at 
night indicating that animals wereemerged from 
lairs. This species probably has nocturnal 
activity patterns. 

Stomach contents were examined in several 
specimens and were found to contain soft tissue 
remains, suggesting a diet of soft bodied animals 
such as shellfish. No crustacean or polychaete 
remains were found. 

The large eggs produced by this species indi- 
cate that young adopt a benthic habit on hatch- 
ing. 

Etymology. From the Greek "gtaptos" meaning 
inscribed or marked, referring to the irregular 
"scribbled" dark markings visible on the dorsal 
mantle and arm crown in preserved material 
(fig. 13a). 

Remarks. Commercial prawn trawl operators 
report catches of this species when trawling at 
night over sandy substrata. This species is fig- 
ured in Queensland Department of Primary 
Industries posters promoting bycatch species for 
human consumption. Annual catch figures are 
not available, however it is likely that this large 
species makes up the bulk of the annual catch of 
octopuses in Queensland waters. 

Discussion 

These new species with other members of the 
Octopus macropus group form a distinct 
assembly of shallow-water octopus species ex- 
hibiting many similarities in morphology and 

behaviour. Shared characters include elongate 
arms (typically 4-7 times mantle length), dorsal 
arms considerably longer than ventral pair (AF 
1.2.3.4), moderate to high gill lamellae counts 
(10-14 per demibranch), a multicuspid radula 
(rhachidian tooth typically bears three lateral 
cusps on either side of a medial cone), a moder- 
ately large cylindrical copulatory organ with 
deep ligula groove, forked lateral walls on the 
lower beak (deeply concave in ventral view), 
absence of enlarged suckers in both sexes, and 
nocturnal activity patterns. 

In describing four new members of the "Octo- 
pus macropus group" from the tropical Indo- 
Wcst Pacific, it is necessary to compare these 
taxa with related species already described from 
these waters. Table 10 lists ten nominal taxa 
from the tropical Indo-West Pacific which share 
these characters and hence are considered mem- 
bers of the O. macropus group. Except for O. 
ornatus, little has been published on these taxa, 
most of which are known only from their orig- 
inal descriptions. Based on examination of type 
material for seven of the nominal taxa, and 
counts and measurements provided in original 
descriptions, the new species described here can 
be clearly delineated. 

Three of the new species (O. alpheus, O. die- 
rythraeus and O. graptus) lay large eggs (to 
28 mm long) in low numbers. This egg size and 
number indicate that hatchlings adopt benthic 
habits on hatching. This feature clearly dis- 
tinguishes these three species from O. lechenaul- 
tii d'Orbigny, 1826 (and its synonym O. cuvieri 
d'Orbigny, 1826) and O. luteus Sasaki, 1929 
which both lay small eggs in large numbers. The 
limited dispersal of the large-egg Australian 
species also reduces the probability of these 
species being synonymous with other Indo-West 
Pacific members of the O. macropus group, 
especially species such as O. rapanui from Easter 
I. 

Octopus ornatus and O. aspilosomatis are 
small-egg species whose planktonic hatchlings 
would be capable of wider dispersal. O. ornatus 
is delineated from the other taxa listed in Table 
10, primarily on the grounds of high sucker 
counts, high gill count (13-14 per demibranch) 
and its distinctive colour pattern (see Norman, 
in prep.). 

Octopus lechenaullii (and its synonym O. 
cuvieri) is known only from the original type 
material. This species shows some similarities 
with O. aspilosomatis including small eggs and 
short webs. O. aspilosomatis appears distinct, 
however, in possessing shorter arms (AMI to 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

305 

Table 10. Nominal taxa described from tropical Indo-West Pacific waters belonging to the Octopus 

macropus group. 

Species 

Type locality 

Valid taxa: 

* O. lechenaullii d'Orbigny. 1826 Pondicherry, Southern India 
Synonym: * O. cuvieri d'Orbignv. Pondicherry. Southern India 
1826 

* O. omatus Gould, 1852 Hawaii 
Synonym: Callistoclopus arakawai Japan 
Taki. 1964 

* O. rapanui Voss. 1979 Easter Island 

Nomen dubium: 

O. machikii Brock, 1887 

Uncertain: 

O. linens Sasaki. 1 929 

O. nanhaiensis Dong, 1976 

* O. taprohtirwnsis Robson. 1926 

* O. teuthoides Robson, 1929 

* Eledonenta filholiana 
Rochebrune, 1884 

Amboina. Indonesia 

Taiwan 

China 

Sri Lanka 

New Hebrides (Vanuatu) 

Fiji 

Nature of type material 

Reasonable condition (MNHN). 
Reasonable condition (MNHN). 

Good condition neotype (USNM). 
Taki's personal collection. 

Good condition (USNM). 

Very poor condition (ZMUG). 

Hokkaido Imperial Museum. 
Inst. Oceanol. Acad. Sin. 
Juvenile specimen (BMNH). 
Juvenile specimen (BMNH). 
Poorly preserved (MNHN). 

type material examined by author 

639.8 vs 669.0 for O. lechenaullii), slightly lower 
gill lamellae counts (10 vs 12 on outer demi- 
branch) and slightly more suckers on intact 
normal arms (206-255-267 vs 168, 210). 
Additional material from southern India would 
be required to confirm this separation. 

Octopus luteus has a similar morphology to O. 
aspilosomatis sharing production of small eggs. 
Octopus aspilosomatis appears distinct in having 
longer arms (AMI 438.3-557.6-639.8 vs 400, 
410forO. luteus), fewer gill lamellae(10 vs 12on 
outer demibranch) and slightly shallower webs 
(WDI 9.1-//.6-14.6 vs 14.0, 17.7). 

Octopus rapanui Voss, 1 979 is described from 
Easter I. The type and additional specimens of 
O. rapanui were examined in the collections of 
NMNH and MNHN. Octopus rapanui is charac- 
terised by deep webs (WDI 18-23 (Voss, 1979)); 
about 105 suckers on hectocotylized arm; 
medium sized spcrmatophores [SpLI 66-75 
(Voss. 1979)]; 11-12 gill lamellae; and distinct 
colour pattern of white base colour with fine 
purple brown chromatophores on dorsal sur- 
faces. These characters clearly delineate this 
species from species occurring in Great Barrier 
Reef waters. 

The status of some species of the Octopus 
macropus group cannot be resolved until 
additional material becomes available. Octopus 
nanhaiensis Dong, 1 976 is known only from the 

brief original description based on a single male 
from Quangdong Province, China. The type has 
not been examined. 

Two species are only known from juvenile 
specimens. Octopus taprobanensis Robson, 1926 
was described from the pearl banks off Sri 
Lanka. It is known only from the type, a 14.4 
mm ML specimen of indeterminate sex, which 
shows the characteristic arm formula (1.2.3.4) 
and high gill count ( 1 3). Octopus teuthoides Rob- 
son, 1929 was described from Vanuatu. It is 
known only from the type, a 1 5.8 mm ML speci- 
men also of indeterminate sex, which shows the 
same arm formula (1.2.3.4) and slightly fewer 
gill lamellae (11). Until the juvenile stages of all 
Indo-West Pacific members of the O. macropus 
group are determined, a process commenced by 
Young et al. ( 1 989) and Hochberg et al. ( 1 992), it 
will not be possible to determine the status of 
these species. 

Octopus machikii Brock, 1887 is only known 
from a single female type specimen. This speci- 
men is still extant in the University Museum, 
Gottingen. It is in very poor condition and was 
considered by Pickford (unpublished MS) as 
being a nomen dubium. Robson (1929) 
described this specimen as having dorsal arms 
longer than ventral ones, and dorsal webs deeper 
than ventral ones, suggesting affinities of this 
species with O. macropus. 

;,l(, MAKk I) NOUMAN 

Korlirl.iuiK* I, SKI) proposed ;i new r.eiius and AcltiiowlcdKi-mi'iils 

species, Eledonenta filhollana, <>n the basis of a | )lls reseaf ch forma pan of ;i I'lii) research 

single poorly-preserved Bpecimen from Fiji. ,„.,,,,,..„„ into octopods of the Great Barriei -Reef 

This genus was proposed as a new member of the and northern Australia, carried oul in associ- 

Eledonidae, reported as possessing the single a tion with the Museum of Victoria and the Uni- 

iows oi suckers typical oi the family. the type versity of Melbourne. Sincere thanks to C.C. Lu, 

specimen oi this species was examined in the ,.- (; Hochberg, G.F. Watson and T. Stranks for 

mini in. md is a poorly fixed elongate specimen, :lsslsl;nUT with the manuscripl and project; R. 

The elongation ofthe arms has probably resulted Tu |, l()l n aspilosomatls information; C, 

from poor fixation oi deterioration prioi to fix Davies, R. Fenwick, G. Greenwood, J. Martin, 

ation The suckers are arranged in two rows bul j gwartz, S. Troy, M. Turner, J. and S. Whyte 

elongation ol the arms has spaced suckers suf« gn< j ( | u . Breidahl family for field assistance; M. 

Sciently loi Rochebrune to interprel them as Dunning (QDPl), D. Sutton and A. Cabanban 

formings zig sag single row, Eledonenta filhoi (jcu) foi provision of specimens; I. Loch, P. 

tana belongs in the Octopodinae and is clearly a coiman and B. Rudman (AM), F. Naggs, 

poorly preserved member of the Octopus macro (BMNH), T, Gosliner (CAS), R. and G. 

pit* group It has the characteristic arm formula Boucher-Rodoni (MNHN), D. White and L. 

ol i 2 'i. 10 c.iii lamellae and a copulatory v . li( ( ntm). .i. Stanisic (QM), S, Slack-Smith 

organ typical ol the group. Additional, well- (WAM)andC. Roper and M.Sweeney (USNM) 

preserved material of this species from the type | ol assistance and access to collections; D. Paul 

locality will be required to resolve the status ol ;MK | ( > Rowley for assistance with photographic 

this species, plates, This research was made possible through 

Octopus macropus has-been reported a num- support grants From the Museum of Victoria, 

bei of limes from Australian waters Girard Victorian Institute of Marine Sciences, Ainv 

(1890) recorded il based on a single specimen tmlian Museum Postgraduate and Keith Suther- 

donated by Baron de Mueller, providing the |. 1IU | Awards, Great Barrier Reef Marine Park 

locality simply as "Australia" Inadequate Authority, M.A. Bartletl travel Grant and 

locality information and the absence of the orig- Write-up Award (University of Melbourne), 

inal specimen leave this record unresolved. j ovce w. Vickery Research Fund (Linnean 

Odhnej (1917) reported Octopui cuvieri from Society of NSW), Ethel Mary Read Fund (Royal 

the tropical waters of north-wesl Western Aus- zoological Society of New South Wales). Ian 

tralla,as"OrtopM.yct<v^r/d*Orbigny macropus p tter Foundation, Hawaiian Malacological 

Kisso, 45 miles WSW of Pearl Banks off < ape society and Royal Society of Victoria, Western 

Jaubert, Western Australia". At this stage, insuf* Society ofMalacologists, Southwesl Shell club. 

Relent material has emerged from the tropical s . ml; , Barbara Shell Club and San Diego Shell 

waters ofWestern Australia to identify the octo- ( ;j U n 
poti.m fauna of this region, t lencc, this record is 

also unresolved, Temperate Australian waters References 

also contain members ofthe Octopus macropus , , , 

, , , ii ,, i uuii , ... i ., . \ilam W p)V) l he ( cphalopoda in the Indian 

group Octopus maorum Hutton, 1880 (and its ^ww, . • ' . ' , , , ,, 

' ,, i ,. .. ,,,,,, , ,, Museum. Cakutln. Records of the Indian Museum 

synonym, 0, fitnaersi Cotton, 1932) trora ": (| M() 

southern Australia and New Zealand waters Wam%w .,~i94l Notessurlesc4phtl<jpo<les.PartXV 

belongs in the macropus group (for full treat- § ul u valeur diagnostique de la radulechei les 

menl of 0. nnii'iuni. see Stranks. 1 4 > S S > An Cgphalopodes Octopodcs, Bulletin du Mwtee 

additional, undescribed species, reported as <> royal d'Histoire naturefle de Belgian? XVII (38) 

macropus by I u and Phillips (1985), occurs in I 19,3 figs, 

the warm temperate waters of southern Queens- Adam.V/ . 1942 LesCephalopodeadelaMei Rouge, 

land and New South Wales, rhis species is prob- Bulletin of Institute oj Oceanography 822(39); I 

ably the same as that reported by Braaier( 1892) .. i0 - ... _ . , . ,. _ *. , .. ... 

„ , , , N , .. ,. .. , , ,. \dam.W., 1946. Ccphaopodo from D SixtcnBocks 

from Port Jackson, New South Wales under the ,, , ,, £ ,i, i>,.r, ■ i -i,„ i 1,1 ,,. r„ 

expedition n> the south racinc islands, irKfv joi 

name <> macropus, Ttoobttl, Stockholm 57A (5): I 25, 2 pis 

lhe phylogcnettc affinities and taxonomte Vlim NN |l)v) cephalopoda. Part I. IV Cephalo- 

status of the Octopus macropus group is the sub ,, lH ies ['Exclusion des genres Sepia, Sepielh et 

ieet of ongoing research (Hochberg, Mangold Sepioteutfm Siboga Expeditiel (Vc): 123 193 

and Noun. m, in piep .). Ad.mi. \\ . L959, Les Cephalopodes dc la Mer Rouge 

NEW SPECIES OF OCTOPUS FROM THE GREAT BARRIER REEF 

307 

Result scientific de Mission de Robert Ph. Doll- 
fus en Egvpte 1927-1929 3eme partie (28): 125- 
193. 

Adam. W\, 1960. Cephalopoda from the Gulf of 
Aqaba. Contributions to the knowledge of the Red 
Sea, No. 16. Bulletin of the Sea Fisheries Research 
Station. Haifa 26: 1-26. 1 pi. 

Adam. W„ 1973. Cephalopoda from the Red Sea. 
Contributions to the knowledge of the Red Sea. 
No.47. Bulletin of the Sea Fisheries Research 
Station. Haifa 60: 9-47. 

Berry. S.S.. 1912. A catalogue of Japanese Cephalo- 
poda. Proceedings of the Academy cf Natural Sci- 
ences of Philadelphia': 380-444. textfigs. 1-4. pis. 
V-IX. 

Berry. S.S.. 1914. The Cephalopoda of the Hawaiian 
Islands. Bulletin of the U.S. Bureau of Fisheries 32: 
255-362. 1 1 pis. 

Boone. L., 1938. Cephalopoda, In: Scientific Results 
of the World Cruises of the yachts "Ara" 1928- 
1929 and "Alva" 1931-1932. "Alva" Mediter- 
ranean Cruise 1933. and "Alva" South America 
Cruise 1935. Bulletin of the Vaiulerhilt Marine 
Museum 7: 328-361. 

Brazier. J.. 1892. Catalogue of the marine shells of 
Australia and Tasmania. Part 1. Cephalopoda. 
Australian Museum Catalogue 15: 1-19. 

Dong. Z.. 1976. On three new species of the genus 
Octopus (Octopoda: Cephalopoda) from Chinese 
waters. Studio Marina Siniea II: 211-215 (In 
Chinese). 

Girard. A. A.. 1890. Revision des Cephalopodcs du 
Museum dc Lisbon ne. Journal tie Sciencias Math- 
ematieas Physicas e Naturaes (2) 1: 233-245. 

Goodrich. E.S., 1896. Report on the collection of 
Cephalopoda from the Calcutta Museum. Trans- 
actions of the Linnean Society of London Scr. 2 
(Zoology) 7: 1-24. 5 pis. 

Gould, A.A., 1852. Mollusca and shells. United States 
Exploring Expedition . . . under the command of 
Charles Wilkes. 12: i-xv. 1-510. atlas of 52 
plates. 

Hochberg, F.G., Nixon. M. and Toll. R.B., 1992. 
Order Octopoda. Pp. 213-280 in: Sweeney, M.J.. 
Roper. C.F.E.. Mangold. K.M., Clarke, M.R. and 
Bolctzky. S.v. (eds), "Larval" and juvenile cepha- 
lopods: a manual for their identification. Smith- 
sonian Contributions to Zoology 513: 1-282. 

Hoyle, W.E.. 1904. Report on the Cephalopoda col- 
lected by Professor Herdman at Ceylon in 1902. 
Report Ceylon Pearl Oyster Fisheries. Supplemen- 
tary Reports 14: 185-200. 3 pis. 

Joubin. L., 1894. Cephalopodcs d'Amboinc. Rente 
Suisse Zoology 2: 23-64. 4 pis. 

Joubin, L, 1898. Note II: Surquelques cephalopodcs 
du Musee Royal de Leyde el description de trois 
especes nouvellcs. Notes Leyden Museum 20: 
21-28. 

Lu. C.C. and Phillips, J.U., 1 985. An annotated check- 
list of the Cephalopoda from Australian waters. 
Occasional Papers from the Museum of Victoria 2: 
21-36. 1 fig. 

Massy, A.L., 1916. XVI. The Cephalopoda in the 
Indian Museum. Records of the Indian Museum 
12(5): 185-247, pis 23-24.' 

Nesis, K.N., 1982. Cephalopods of the World: squid, 
cuttlefish, octopuses and their allies. [1987 
English abridged translation by B.S. Levitov]. 
T.F.H. Publication: Neptune City. 351 pp. 

Norman. M.D.. 1991. Octopus cyanea Gray, 1849 
(Mollusca: Cephalopoda) in Australian waters: 
description, distribution and taxonomy. Bulletin 
of Marine Science 49(1-2): 20-38, 5 figs. 

Norman, M.D.. 1992a. Ameloclopus liloralis. gen. et 
sp. nov. (Cephalopoda: Oclopodidae), a new shal- 
low water octopus from tropical Australian 
waters. Invertebrate laxonomy 6(5): 567-582. 

Norman, M.D., 1992b. Ocellate octopuses (Cephalo- 
poda: Octopodidae) of the Great Barrier Reef, 
Australia: description of two new species and 
redescription of Octopus polyzenia Gray, 1849. 
Memoirs ol the Museum of Victoria 53: 309- 
344. 

Norman, M.D., (in prep.). Octopus omatus Gould, 
1852 (Cephalopoda: Octopodidae) in Australian 
waters and a review of its description, life history 
and distribution. Submitted to Proceedings of the 
Biological Society of Washington. 

Odhncr. N.H.J.. 1917. Results of Dr. E. Mjoberg's 
Swedish scientific expeditions to Australia, 1910- 
1913. Part XVII. Mollusca. Kunliga Svenska 
Vetenskapsakademiens. Handlingar (4th ser.), 
52(16): 1-115, 3 pis, 51 figs. 

d'Orbigny. M.A.. 1826. Plates released in advance of 
text of Ferussac and d'Orbigny, 1835-1848. 

Pickford. G.E.. (unpublished manuscript). A revision 
of the Indo-Malayan Octopodinae based on the col- 
lections of the British Museum. Bingham Oceano- 
graphic Laboratory, Yale University. 

Recs, W.J. and Stuckcy, A., 1954. The "Manahine" 
expedition to the Gulf of Aqaba, 1948-1949: VI: 
Mollusca. Bulletin of the British Museum (Natural 
History) 1: 183-201, pis 28-30. 

Robson. G.C., 1926. Notes on the Cephalopoda, 1. 
Descriptions of two new species of Octopus from 
southern India and Ceylon. Annals and Magazine 
of Natural History (ser. 9) 17: 159-167, 7 figs. 

Robson, G.C.. 1929. .1 monograph of the Recent 
Cephalopoda. I. Octopodinae. British Museum 
(Natural History): London. 236 pp. 

Robson, G.C., 1932. Report on the Cephalopoda in 
the Raffles Museum. Bulletin of the Rallies 
Museum. Singapore. Straits Settlements 7: 21-33, 
4 ligs. 

Rochebrune, A.T, de, 1884. Etude monographiqucde 
la famiile des Eledonidae. Bulletin de la Societe 
Philomathique de Paris 7(8): 152-163, 1 pi. 

Roper, C.F.E. and Hochberg, F.G., 1987. Cephalo- 
pods of Lizard Island, Great Barrier Reef, Aus- 
tralia. Occasional Papers from the Museum of 
Victoria 3: 1 5-20. 

Roper. C.F.E. and Hochberg. F.G., 1988. Behaviour 
and systematics of cephalopods from Lizard 
Island. Australia based on colour and bodv 

308 

MARK D. NORMAN 

patterns. Makwologia 29(1): 153-193. 

Roper, C.F.E. and Sweeney, M.J., 1983. Techniques 
for fixation, preservation, and curation of cepha- 
lopods. Memoirs of the Museum of Victoria 44: 
29-47. 

Roper, C.F.E., Sweeney, M.J. and Nauen, C.E., 1984. 
FAO Species Catalogue, Volume 3. Cephalopods 
of the World. FAO Fisheries Synopses, (125) 3: 
196 pp, textfigs. 

Roper, C.F.E. and Voss, G.L., 1983. Guidelines for 
taxonomic descriptions of cephalopod species. 
Memoirs of the Museum of Victoria 44: 49-63. 

Sasaki, M., 1 920. Report of cephalopods collected dur- 
ing 1906 by the United States Bureau of Fisheries 
Steamer "Albatross" in the northwestern Pacific. 
Proceedings of the United States National 
Museum 57: 163-203, 4 pis. 

Sasaki, M., 1929. A monograph of the dibranchiate 
cephalopods of the Japanese and adjacent waters. 
Journal of the Faculty of Agriculture, Hokkaido 
Imperial University 20 (suppl.), 357 pp., 30 pis. 

Stranks, T.N., 1988. Systematics of the family Octo- 
podidae (Mollusca: Cephalopoda) of south- 
eastern Australia. Unpublished MSc Thesis, 
University of Melbourne, Victoria. 114 pp. 

Taki, I., 1944. Studies on the octopus. (2) The sexes 
and the reproductive organs. Japanese Journal of 
Malacology (The Venus) 13 (5-8): 267-310. 

Taki, I., 1964. On eleven new species of the Cephalo- 
poda from Japan, including two new genera of 
Octopodinae. Journal of the Faculty of Fisheries 
and Animal Husbandry, Hiroshima University 
5(2): 277-343. 

Voss, G.L., 1963. Cephalopods of the Philippines. 
Smithsonian Institution Bulletin 234: 1-180, 
36 figs. 

Voss, G.L., 1979. Octopus rapanui, new species from 
Easter Island (Cephalopoda: Octopoda). Proceed- 
ings of the Biological Society of Washington 92(2): 
360-367. 

Voss, G.L., 1981. A redescription of Octopus ornatus 
Gould, 1852 (Octopoda: Cephalopoda) and the 
status of Cullistoctopus Taki, 1964. Proceedings of 
the Biological Society of Washington 94(2): 525- 
534. figs 1-3. 

Wiilker, G., 1913. Cephalopoden der Aru und Kei 
Inseln. Abhandlungen Senckenbergischen Natur- 
forschenden Gesellschaft 34: 451-488, pi. XXII. 

Wiilker, G., 1920. Uber Cephalopoden der Roten 
Meeres. Senckenbergiana 2(1): 48-58. 

Young, R.E., Harman, R.F. and Hochberg, F.G.. 1 989. 
Octopodid paralarvae from Hawaiian waters. The 
Veliger 32(2): 152-165, 5 figs.