Ocellate Octopuses (Cephalopoda: Octopodidae) of the Great Barrier Reef, Australia: Description of Two New Species and Redescription of Octopus Polyzenia Gray, 1849

Memoirs of the Museum of Victoria 53(2): 309-344 (1992) 31 December 1992 https://doi.org/10.24199/j.mmv.1992.53.14 OCELLATE OCTOPUSES (CEPHALOPODA: OCTOPODIDAE) OF THE GREAT BARRIER REEF, AUSTRALIA: DESCRIPTION OF TWO NEW SPECIES AND REDESCRIPTION OF OCTOPUS POLYZENIA GRAY, 1849

By Mark D. Norman

Invertebrate Zoology, Museum of Victoria, 285 Russell Street. Melbourne, Vic. 3000, Australia

Abstract

Norman, M.D., 1992. Ocellate octopuses (Cephalopoda: Octopodidae) of the Great Bar- rier Reef, Australia; description of two new species and redescription of Octopus polyzenia Gray, 1849. Memoirs of the Museum of Victoria 53: 309-344. Four species of ocellate octopuses are reported from the waters of the Great Barrier Reef, Australia. Ocellate octopuses exhibit a false eye-spot (or ocellus) on the arm crown below each eye. Octopus polyzenia Gray, 1849 is reported as an ocellate species for the first time and is redescribed based on examination of the type and specimens from tropical waters across northern Australia. Two new species, O. exannulatus and O. mototi spp. nov. are described and details of distribution and life history are presented. The widely distributed O. cyanea Gray, 1849 is the fourth ocellate species found on the Great Barrier Reef. Octopus cyanea shows different affinites to the other three species and may be of separate origin. A key for ocellate octopuses of Great Barrier Reef waters is provided. Delineation of Great Barrier Reef ocellate octopuses from others of the tropical Indo-West Pacific is discussed. Distribution patterns also are examined.

Introduction ern Queensland (Smith, 1884). In examining these specimens in the British Since the early 19th century, teuthologists Museum (Natural History), it was discovered that both possess have been familiar with ocellate octopuses, ocelli with an iridescent ring and both manv authors discussing them in detail (Brock, were mature males despite their small size (holotype: 1887;Berrv, 1912. 1912a; Robson, 1929; Sasaki, 12.3 mm ML, BMNH 1845.5.14.78 and Thurs- 1929; Adam, 1939, 1959, 1973; Pickford and day I. specimen: 14.1 ML, McConnaughey, 1949; Voss and Solis Ramirez, mm BMNH 1882.2.23.568). An additional 14 specimens 1966). Ocellate octopuses are shallow-water were found in Australian museum collections species characterised by the presence of a pair of (NTM, AMS, WAM, NMV). Octopus polyzenia dark false eye-spots (or ocelli) on the arm crown is here redescribed as a small ocellate species over the bases of arms 2 and 3, one below each which lays large eggs and occurs in coastal waters eye. In many of these octopuses the dark ocelli across northern Australia. Species accounts are each contain a fine ring of iridescent tissue, presented for two new ocellate species, O. exan- which is expressed as a brilliant metallic blue or nulatus and O. mototi, along with information gold ring in live animals. on distribution, life history and live animal attri- Recent research into the shallow-water octo- butes. puses of the Great Barrier Reef, Australia has uncovered many octopus species (Norman, Material and methods 1991, 1992a, 1992b), including four ocellate species: Octopus cyanea Gray, 1849, O. poly- Field work on the Great Barrier Reef was carried zenia Gray, 1849 and two new species, O. out between May and November in 1989 and exannulatus and O. mototi spp. nov. 1 990. Live specimens of O. cyanea and O. mototi Octopus cyanea is a widely distributed tropical were encountered in the field during the day and Indo-West Pacific species, found in coral reefs. at night on intertidal reef walks, snorkel and It has previously been reported from Australian SCUBA dives and caught in octopus pots in waters by Roper and Hochberg ( 1 987, 1 988) and deeper water. Individuals were observed in situ, Norman (1991). Octopus polyzenia was pre- collected, photographed, narcotised in fresh viously known only from the type specimen water, and fixed and preserved according to the from Port Essington, Northern Territory (Gray, techniques of Roper and Sweeney (1983). These I., north- specimens are now in the 1 849) and a small male from Thursday Museum of Victoria 309 .

310 MARK D. NORMAN

(NMV). Preserved specimens of all four ocellate per cent of ML; Hectocotylized Arm Sucker Reef species were found and examined in the Count (HASC): number of suckers on hectoco- collections of the Australian Museum, Sydney tylized arm (see Toll, 1988); Terminal Organ (AM), Queensland Museum, Brisbane (QMB), Length Index (TOLI): length of terminal organ Northern Territory Museum of Arts and Sci- as per cent of ML; Diverticulum Length Index ences (NTM), Western Australian Museum (DLI): diverticulum length as per cent of length (WAM), National Museum of Natural History. of terminal organ; Spermatophore number Washington (USNM), Californian Academy of (SpN): number of spermatophores in Need- Science (CAS), British Museum (Natural His- ham's sac; Sperm Cord Whorls (SpCW): number tory) (BMNH) and Museum National d'Histoire of whorls in which sperm cord is coiled; Funnel Naturelle, Paris (MNHN). Length Index (FLI): funnel length as per cent of In the description and tables, measurements ML; Free Funnel Index (FFI): length of free fun- and indices follow Voss (1963) and Roper and nel portion as per cent of funnel length: Funnel Voss (1983). Terminology has been modified by Organ Index (FOI): length of outer limb of fun- Hochberg and Mangold (in prep.) for several nel organ as per cent of median limb length; anatomical structures: "copulatory organ" Funnel Organ Length Index (FOLI): length of replaces the term "hectocotylus" and "terminal medial limb as percent of funnel length; Ocellus organ" replaces the term "penis". The following Diameter Index (OcDI): diameter of ocellus as additional or slightly modified indices also are a per cent of ML (specified as referring to employed: iridescent ring or entire dark web spot). Stage of Maturity (StM): Immature (I: sex Indices are presented for both sexes com- indeterminate or reproductive organs minute). bined, except where significant differences were Submature (S: reproductive organs distinct but found between the sexes (one-way ANOVA, P = poorly developed) and Mature (M: developed 0.05 level). In these indices, range and mean for spermatophores or eggs distinct); Head Width each sex are presented. Where ranges signifi- Index (HWI): head width as a per cent of ML; cantly overlap, standard deviations around the Head Mantle Width Index (HMWI): head width mean are also presented. as per cent of mantle width; Arm Mantle Index Two additional undescribed ocellate species (AMI): arm length as per cent of ML: Arm Width have been recognised from northern Australian Index (AW1): maximum arm width at widest waters west of Cape York. Due to the scarcity of point on stoutest arm, as per cent of ML; Sucker material and the apparent absence of these taxa Count (SC): total sucker count for the intact arm from the Great Barrier Reef region, these forms with the most suckers; Gill Count (GC): number have not been included in this work. of gill lamellae per demibranch not including the Table 8 summarises the key differences terminal lamella; Hectocotylized Arm Mantle between ocellate octopuses of the Great Barrier Index (HAMI): length of hectocotylized arm as Reef.

Key to ocellate octopuses of the Great Barrier Reef

Ocellus lacks iridescent ring; either simple black spot (figs la, lOe-f) or black spot surrounded by a fine dark outer ring (figs lb, 2a) 2 Ocellus consisting of dark spot containing narrow iridescent blue ring in live animal, visible as white or pink superficial ring over dark ocellus in preserved specimens (figs 1c, 6h, 14g-h) 3 9-11 gill lamellae per demibranch, typically 10; over 400 suckers on normal intact arms of submature and mature animals; approximately 200 suckers on hectocotylized arm; ocellus comprised of dark oval spot bound by additional thin dark ring (fig. lb), iridescent tissue absent; dark zebra bars on ventral faces of all arms on submature and mature animals

(fig. 2d); very large species (ML to 160 mm; weight to 6 kg; fig. lh) .

Octopus cyanea Gray, 1 849

7-8 gill lamellae per demibranch. typically 7; 1 20- 1 90 suckers on normal intact arms of submature and mature animals; approximately 70 suckers on hectocotylized arm; ocellus plain black spot lacking an iridescent ring, oval shaped and clearly defined (figs la, 10e-f); 4 dark broad longitudinal stripes on dorsal body, continuing anteriorly on to dorsal arm crown .

OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 311

(figs 7a, lOa-d); moderate size species (ML to 50 mm; weight to 75 g; fig. 10 Octopus exannulatus sp. nov. 6-7 gill lamellae per demibranch, typically 6; 85- 135 suckers on normal intact arms of submature and mature animals; approximately 50 suckers on hectocotylized arm; widely spaced dark transverse bars on all arms

separated by approximately 3-4 suckers (figs 3a, 6d, g); pattern of faint crucifix of light patches on dorsal body (figs 3a); small species (ML to 38

mm; weight to 20 g; fig. la) Octopus polyzenia Gray, 1849 9-11 gill lamellae per demibranch, typically 11; 143-176 suckers on normal intact arms of mature animals; approximately 100 suckers on hectocotylized arm; circular cluster of dark spots above each eye forming

"flower" pattern (figs 1 la, 14a); alarm pattern of dark longitudinal bars on dorsal body and arm crown over white base (figs 14b, e); well developed frontal white spot with prominent elongate papilla (figs 14a, 14d);

moderate to large species (ML to 100 mm; weight to 300 g; fig. lg) . O. mototi sp. nov.

Octopodidae d'Orbigny. 1839 Pacific from Hawaii in the east to the east Afri- can coast in the west (fig. 15a). Subfamily Octopodinae d'Orbigny. 1 839 Aspects of the life history of O. cyanea were

Octopus cyanea Gray. 1 849 treated in Van Heukelem (1983). Morphological description and observations of live Australian Figs lg, 2a-d, 15a specimens were provided in Norman (1991). See Norman (1991) for full annotated synonymy. Commercial exploitation. There is no commer- Octopus marmoratus Hoyle. 1885. O. horsti Joubin, gra- cial fishery for O. cyanea in Australian waters. 1 898, O. herdmani (Hoyle, 1 904). O. cyanea var. The association of this species with coral and cilis (Robson. 1 928) and Callistoclopus magnocellatus Taki. 1964 are synonyms. rocky reefs excludes it from trawl fisheries which work more open substrates. No small-scale local Diagnosis. Large robust species with black oval exploitation has been noted. Elsewhere through- ocellus surrounded by pale ring and thin dark out the tropical Indo-West Pacific, this species is outer ring. Ring of iridescent tissue within commonly taken in local and subsistence fish- ocellus absent. Dark bars on ventral surfaces of eries (Hoyle, 1907; Berry, 1912; Sasaki, 1929; all 4 arms of larger specimens (ML > about 60 Rees, 1950; Van Heukelem, 1983; Roper et al., forming stripes in alternation with base of mm), 1984). suckers. 3-7 rows of cream spots on aboral arm tips, containing surfaces from web margin to Octopus polyzenia Gray, 1 849 small erectile skin ridges. Skin sculptured in Figs Id, 3-6. 15b irregular patch and groove system. 3 pro- patches on dorsal arm nounced large circular 1879: Octopus polv-enia Gray , 1849: 13. — Tryon, gill especially in younger animals. 9-1 1 crown, 122. — Cox, J882: 787. — Smith, 1884: 34, pi. 4. figs typically 10. More lamellae per demibranch, A-A3. — Hoyle, 1886: 8. 80. — Lu and Phillips, 1985: of sub- than 400 suckers on intact normal arms 33. mature and mature individuals, approximately Octopus granulatus (non Lamarck, 1798). — 4. 200 suckers on hectocotylized arm. Terminal Brazier. 1 892: Ortmann, organ small and fine (TOLI about 20). Sperma- Octopus rugosus (non Bosc, 1792). — 1891:669. — Robson, 1 929: 63, 65, 73. — Flecker and tophores short, approximately 40% of ML and Cotton, 1955: 3. produced in large numbers (SpN >300). Eggs small, ovarian eggs to 2.7 mm [ELI(ov) to 1.7], Material examined. 16 preserved specimens including examined in and produced in very large numbers (EN the extant type (BMNH 1845.5.14.78) museum collections and the British > 100 000). Australian Museum (Natural History). from Distribution. Octopus cyanea is recorded Holotype: NT: lcf: 12.3 mm ML, BMNH from tropical waters of northern Australia, 1845.5.14.78, Port Essington, 11°16'S, 1 32WE, pre- Moreton Bay, southern Queensland (27°23'S, sented by the Earl of Derby. north-west Western Other material: NT: 31mm: 7.4-8.8 mm ML. NTM I 53°1 5'E) to Point Cloates, is PI 393. Port Essington, Coral Bay (1 1°16'S, 132WE), Australia (22°42'S, 113°39'E). This species 4.5-6 m, Helen Larson, 18 Oct 1981 (poison station. widclv distributed in the tropical Indo-West 312 MARK D. NORMAN

^t&Mtete*.

100 mm

Figure l . Ocelli and body sizes of Great Barrier Reef ocellate octopuses, a-c. Ocellus types: a, ocellus simple black spot, lacking iridescent ring or outer ring (as in O. exannulatus). b, dark ocellus contained within a light ring, which is bound by an additional thin black ring (as in O. cyanea). c, dark ocellus containing thin iridescent ring on surface of skin (as in O. polyzenia and O. mototi). d-g. Comparative body sizes: d, Octopus polyzenia Gray, 1 849. e, Octopus exannulatus s

patterns showing range in sculpture and „ rr™ i R4Q a-t I ive colour

mature male). specimen (NMV F57904: 100.0 mm ML

25.8- 20°0l'S, 148°15'E, E.H. Rainsford, no date; 39 : bottom, rock rubble with soft corals and gor- siltv sand CI 64 168, Gulf of Carpentaria, ML, NTM P1451, Darwin. 33 3 mm ML, AM gonians)- 1 9 : 32.8 mm 141°42'E,W. H. Foley, 130°52'E). Weipa, Albatross Bay, 12°40'S. Casuarina Beach, Off Lee Point. (12°2I'S. lrf: ML, AM C164179, Gulf of Car- Pinna shell alter 1962- 26.3 mm C. Hood, 22 Jan 1986 (washed up in pentaria, off Karumba, (about I7°29'S, about storms). , „_ _ CSIRO Prawn Survey, 1963-1964 (trawl); ML, WAM 363-88. Dampier 140°50'E), WA- 1?: 13.9 mm ML, NMV F60112, Cleveland Bay, off Norbill Bay, 1 9 : 37.7 mm Archipelago, Rosemary I., outside Townsville, (19°11'S, 147°01'E), <20 m, G. Jackson. 5 m. Barry Wilson, WAM Crown (20°29'S 116°35'E) Kirby"). 2 17 .7 1989 (trawled, RV "James 21 May 1972 (dredged), 9 . of Thorns Survey, ML WAM mm, body missing from second specimen, Dampier Archipelago. Rosemary I., Norbill 329-88. black oval ocellus 1-2 m, Barry Wilson, WAM Diagnosis. Small species with Bav (70°29'S, 1 16°35'E), (dredged on iridescent blue ring. Dark, widely- Crown of Thorn Survey, 21 May 1972 containing an 305-88, Monte Bello Is, transverse bars along aboral and lateral sand)' 1 J: 24.4 mm ML, WAM spaced, F.E. (20°28'S, 1 15'31'E), 3-4 suckers northern end of Hermitc I., faces of all arms, approximately Wells 70 Aug 1986. m between each bar. Light patches surrounding BM 82.2.23.568, Torres dorsal Old' I* 14.1 mm ML, raised skin ridges form crucifix pattern on (10°35'S, 142°13'E), 7.3-9.2 m. Strait Thursday I., 6-7 gill lamellae per demibranch, typi- AM C48267 BoWCn- mantle. 24 - 9 mm ML ' HMS "Alert"; l* ' 314 MARK D. NORMAN

4 mm

2mm

10 mm

Figure 3. Octopus polyzenia Gray, 1849. a, dorsal view of 33.3 mm ML female (AM C164168): DMR = dorsal mantle ridges; FWS = frontal white spot complex, b, funnel organ of ~>6 same specimen, c, copulatorv organ6 ofu, ^u.j3 mm ML male (AM CI 64 1 79). OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 315

5 mm

tract 33.3 ML female (AM C 1 64 1 68): ASG = anterior Figure 4. Octopus polyzenia Gray, 1 849. a, digestive of mm = = diverticulum; = crop; = digestive gland; I salivary glands; BM = buccal mass; C = caecum; CD crop CR DG salivary gland; S = stomach. intestine; IS = ink sac; O = oesophagus; PSG = posterior lower beak, lateral view, d, lower beak, ventral b-d, beaks of same specimen: b, upper beak, lateral view, c, view. 316 MARK D. NORMAN

Figure 5. Octopus polyzenia Gray, 1849. a, reproductive tract of 26.3 mm ML male (AM C164179): MG = mucilaginous gland; NS = Ncedham's Sac; = SG 1 spermatophoric gland I; SG2 = spermatophoric gland II; T = testes; TO = terminal organ; VD = vas deferens, b, spermatophore of same specimen: CB = cement body SR = sperm reservoir, c, ovary of 33.3 female mm ML (AM C 1 64 1 68): DO = distal oviduct; O = ovary; OG = oviducal gland; PO = proximal oviduct. OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF. AUSTRALIA 317

cally 6. Approximately 115 suckers per arm in Third right arm in males hectocotylized and mature females (SC$ 103-775-135, n = 9<&), slightly shorter than opposite arm (OAI: 76.9, slightly fewer in mature males (SCrf 85, 86, 108), 77.4; HAMI: 152.1, 155.7, 195.1). Spermato- approximately 50 suckers on hectocotylized phore groove well developed and wide with fine arm. Terminal organ short and robust (TOL1 transverse ridges. Spermatophore guide distinct about 15). Spermatophores short, approxi- with small papillae. Copulatory organ small and mately one third of ML and produced in low robust (fig. 3c: LLI 4.7, 5.4, 6.8), ligula roughly numbers (SpN about 10). Eggs large, spawned conical with blunt tip; groove shallow with capsules to 7.5 mm [ELI(sp) to 22.9] and pro- raised longitudinal midrib. Calamus well devel- duced in moderately low numbers (EN about oped and large (CLI 40.7, 44.4, 69.2). Approxi- 250). mately 50 suckers on hectocotylized arm (HASC: 45, 49, 52). Description. Based on 3 mature males, and 3 Gills with 6-7 lamellae, typically 6, on both submature and 4 mature females. Counts and inner and outer demibranchs, plus a terminal indices are in Tables 1 and 2. lamella. Small robust species (figs Id, 3a), males to at Digestive tract illustrated in figure 4a. least 26 mm ML. females to 37.7 mm ML, TL to Anterior salivary glands moderately large, length. Pos- at least 130 mm; weight to at least 19 g. Mantle approximately half of buccal mass ovoid (MWI 50.4-69.0-85.4) with muscular terior salivary glands well developed, slightly walls. Stylets not found. Pallial aperture of mod- larger than buccal mass. Crop diverticulum dis- erate width, approximately half of mantle width. tinct but not greatly developed. Stomach bipar- Funnel muscular and broad-based (FLI 34.1- tite. Caecum striated, coiled in single whorl. intestine reflexed approximately one- 39.6-5 1 .2) with free portion usually greater than Muscular half funnel length (FFLI 44.0-65.5-78.1). Fun- third along length from proximal end. Ink sac in ventral surface of nel organ W-shaped with moderate limbs (fig. well developed, embedded iridescent tissue layer. 3b). Outer limbs slightly shorter than median digestive gland below limb (FOI 90.2-95.7-100.0). Funnel organ Anal flaps present. moderate hood and slightly large, approximately 60% of funnel length Upper beak with (FOLI 48.0-67.7-73.5). hooked rostrum, concave on cutting edge (fig. Head of moderate width (HWI 41.1-49.5- 4b). Lower beak (figs 4c-d) with blunt rostrum, hood, widely spread wings and slightly 63.3), always narrower than mantle (HMWI narrow view of posterior 60.0-72.5-89.5). Neck distinct, slightly flared lateral walls. Ventral concave. Radula with seven narrower than head. Eyes large and slightly margin moderately in each transverse pronounced. teeth and two marginal plates Rhachidian tooth typically has 1 Arms of moderate length, 2-3 times mantle row (figs 6i-j). lateral cusp on each side of medial cone (2 in length; from limited material arms appear fourth row). Lateral cusps in symmetrical slightly longer in females (AMIrf 200.8, 209.1, every seriation. migrating from lateral to medial pos- 252.0; 9 236.4-267. 7-309.4). Arms robust (AWI four transverse rows. First lateral 13.0-75.2-25.9), roughly circular in cross sec- ition over teeth unicuspidate with cusp towards lateral tion and taper evenly to fine tips. Dorsal arms edge; second lateral teeth unicuspidate and shortest, lateral and ventral arms longer and marginal teeth robust, straight more robust (AF generally 4=3.2.1 or 4.3.2.1). robust; lateral moderately short; marginal plates oblong Suckers of moderate size, slightly elevated with and and plain. distinct radial cushions and scalloped rim. Male reproductive tract illustrated in figure Mature males can possess 1-3 enlarged suckers 5a. Terminal organ in mature males short and on all arms, beyond margin of webs at level of robust (TOLI 14.4, 16.3. 16.8) with robust seventh sucker pair(SDLn 0.3, 11.7, 13.5;97.7- diverticulum (DLI 34.2, 56.1, 70.0), genital 9.4-12.9). Arm 4 possesses most suckers, deferens short and in females aperture subterminal. Vas approximately 1 1 5 suckers per arm robust. Mucilaginous gland enlarged at point of (SC9 103-/75-135, n = 999). slightly fewer in attachment to vas deferens. Spermatophoric males (SO? 85, 86, 108). Webs moderate to deep gland I short and robust with recurved coil (WD1 26.1-57.4-39.1), deepest laterally, dorsal approximately three-quarters along length. web distinctly shorter (WF typically D.OE.B.A Spermatophoric gland II relatively short and or D=E.C.B.A). Web margins poorly developed, robust with reflexed tip. No distinct appendix at extending less than halfway along ventral edge junction of spermatophoric glands and Need- of arms. 318 MARK D. NORMAN

„-..•''.-'...... •. W^'-.

a b c OCELLAIE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 319

Figure 6. Photographs ol' preserved specimens of Octopus polyoma Gray, 1849. a. dorsal view ol" 25.8 mm ML female (AM CT 64 168). b. dorsal view of 26.3 mm ML male (AM CI 641 79). c, dorsal view of 24.4 mm ML male bars.e, enlarged suckers (WAM 305-88). d, posterior view of 1 3.9 mm ML female (WAM 363-88), indicating arm

1 arms and of 24.4 mm ML male (WAM 305-88). f, lateral view of 8.8 mm ML immature specimen (NTM 393). g, ocellus of 32.8 mm ML female (NTM F1451), indicating position of arm bars, h, ocellus of same specimen, containing iridescent ring, i-j, radula of 32.5 mm ML female (AM C164168).

papilla. Posterior ham's sac. Needham's sac relatively short and coloured containing primary clearly distinct robust. Spermatophores (fig. 5b) short (SpLI spot pink to peach coloured, pigmentation. 34.2) and moderately robust (SpWl 5.6), pro- from surrounding crown covered in small duced in low numbers (10 in Needham's sac of Body, arms and arm of AM C164179). Sperm reservoir approximately round papillae, extending on to oral surface half spermatophorc length (SpRl 55.6). Sperm male Octopus cord iridescent gold in colour and coiled in Table 1. Counts and indices of approximately 45 whorls. Oral cap with con- polyzenia Gray. 1 849. = = * not stricted collar bearing long cap thread. (M mature; D damaged; — * = as diameter of Mature ovary oval shaped (fig. 5c). Oviducts recorded; OcDl measured ring). short, robust and straight. Oviducal glands large iridescent with approximately 14 radiating (braiding) AM chambers. Eggs large, capsule of mature ovarian Museum BMNH WAM 1845.5.14.78 305-88 C164179 eggs to 4.9 mm long [ELl(ovarian) to 19.7], Reg. No. spawned eggs with capsule to 7.5 mm (Holotype) [ELKspawned) to 22.9]. Eggs moderately wide to 9. and ML 12.3 24.4 26.3 [EWKovarian) to 6.8: EWI(spawned) 1] M M produced in low numbers (226 spawned eggs StM M 46 75 84 accompanying 32.8 mm ML female: NTM TL 85.4 50.4 70.3 P1451). Approximately twelve follicular folds MWI 60.2 45.1 45.2 with cross striations on ovarian eggs. Ovarian HW1 64.3 stages of develop- HMWI 70.5 89.5 eggs in I female at 2 distinct 195.1 151.6 174.9 ment, suggesting spawning over an extended AMI: 1 243.9 168.0 205.3 period. Eggs laid in short festoons attached by- 2 3 252.0 D 197.7 short stalks to a fine central thread. 152.1 pre- II 195.1 155.7 Colour in life unknown. Base colour ol 4 252.0 200.8 209.1 served specimens generally cream to pink brown 13.0 14.3 20.2 with dark mottlings and light patches on dorsal AW1 coloured SD1 11.4 13.5 10.3 body. Dorsal mantle pattern of light cen- WDI 32.2 34.9 32.7 oval patches containing raised skin ridges, forming GC 6 6 6 tral four arranged tn diamond shape 195.1 155.7 152.1 rec- HAM I faint crucifix pattern (fig. 3a). Dark brown 77.4 D 76.9 block between and slightly behind eyes OA1 tangular 49 52 to black oval HASC 45 in some specimens. Dark brown LL1 5.4 4.7 6.8 either side of arm crown between ocellus on 44.4 40.7 small iri- CLI 69.2 bases of arms 2 and 3. containing iridescent TOLI 16.3 16.8 14.4 ring (fig. 6h: OcDl for descent blue 34.2 spot ol the DLI 70.0 56.1 ring6. 1-9. 7-14.6). Dark background — leaving SpLI — 34.2 ocellus may fade in preserved specimens bars SpWl — — 5.6 ring visible. Dark transverse only iridescent — — 55.6 surfaces of all arms, widely SpRl on aboral and lateral — 10 each bar (tigs SpN — spaced with 3-4 suckers between FLI 40.7 32.8 36.5 dark longitudinal line along dor- 3a 6d f-g). No 78.1 motolt and O. FFLI 44.0 51.3 face of arms as found in O. sal' FOI 91.7 97.9 95.7 exannulaius, ,, spots FOL1 48.0 58.8 71.9 Frontal white spot complex ol 2 light midpoint ot OcDl* 14.6 6.1 8.0 present on dorsal arm crown below Anterior spot cream eyes (FWS in fig. 3a). 320 MARK D. NORMAN

Table 2. Counts and indices for female Octopus polyzenia Gray, 1849. (S = SLibmature; M = mature; D = damaged; — = not recorded; * = OcDl measured as diameter of iridescent ring).

Museum WAM AM AM AM NTM AM NMV Reg. No. 363-88 C48267 C164168 C164168 P1451 C164168 F60112

ML 13.9 24.9 25.8 32.5 32.8 33.3 37.7 StM S M S S M M M TL 59 93 90 119 121 126 133 MWI 74.1 71.1 69.4 72.9 57.6 77.5 61.5 HW1 63.3 53.0 49.2 48.3 43.0 46.5 41.1 HMW1 85.4 74.5 70.9 66.3 74.7 60.0 66.8 216.2 183.0 AMI: 1 230.2 188.8 186.0 206.2 210.4 2 251.8 224.9 220.9 224.6 207.3 228.2 209.5 3 309.4 236.9 236.4 258.5 247.0 270.3 212.2

4 302.2 26 1 .0 232.6 261.5 247.0 261.3 246J AWI 25.9 17.7 17.1 18.2 18.6 17.1 19.9 SDI 12.9 9.2 9.3 8.9 9.1 8.4 7.7 WD1 39.1 29.2 32.8 27.1 30.9 27.8 26.9 GC 6 6 6 6 7 7 7 ELI 19.7 22.9 12.6 — EWI 6.8 9.1 3.9 — EN 100s 226 100s 100s FLI 48.9 34.1 41.5 40.0 38.7 46.8 36.1 FFLI 76.5 71.8 61.7 70.8 63.8 62.2 54.4 FOl 100.0 94.0 91.4 97.0 93.8 90.2 99.0 FOLI 60.3 58.8 65.4 51.5 63.8 59.0 73.5 OcDl* 12.9 9.2 9.7 9.5 8.5 9.0 9.5

dorsal and dorsolateral webs. Sculpturing pres- (20'01'S, 148°15'E) west to Rosemary L, Damp- ent on ventral surface of body, but absent from ier Archipelago, Western Australia (20°29'S. ventral arm crown. Longitudinal raised skin 1I6°35'E). ridges visible on dorsal body (fig. 3a), largest 4 Life history. This taxon appears restricted to forming diamond arrangement. Additional shallow, coastal waters on open substrata of sand shorter raised ridges on dorsal and lateral or mud. Most specimens were collected by trawl mantle. 1 large and 2 small branching papilla or dredge between 1 and 20 m. The large eggs above each eye. and well developed arms in very small juveniles Sexual dimorphism not marked in the few from the type locality (31mm: 7.4-8.8 mm ML, specimens available. Mature males can possess NTM PI 393) indicate hatchlings adopt a bent- 1-3 enlarged suckers on all arms at level of sev- hie habit. enth sucker pair. Two other trends are visible in Two specimens were accompanied by eggs the limited material: i) mature males appear to laid in bivalve shells. have longer arms than females (AMlcf mean 220.6; a mean 261. 7); yet, ii) females appear to Commercial exploitation. There is no known have higher sucker counts than males (SC9 mean exploitation of this species although it is likely to 115\ d mean 93). When converted to a ratio of occur in low numbers in prawn trawl by-catch, sucker count/arm length, the trend is clearer particularly in the Gulf of Carpentaria. (SC/AL? 130.0-754.4-193.3, n = 699,

Robson, 1929: Flecker and Cotton, 1955). a of Carnarvon, (about 24'53'S. I13"40'E), L. Marsh species also lacking type material. The type and M. Sinclair. 3-4 Jul 1975 (trawl). rugosus. refer- locality of O. and the absence of Diagnosis. Small to moderate sized species with ence to ocelli in both O. granulatus and O. rugo- simple black ocellus lacking an iridescent ring. 4 sus justify removing O. polyzenia from the dark broad longitudinal stripes present on dorsal of synonymies of these species. The status these body extending on to arm crown. Dark leading two nominal taxa requires review. edges present along length of dorsal edges of all arms. 7-8 gill lamellae per demibranch. Octopus exannulatus sp. nov. Approximately 150 suckers per arm in females Figs lc, 7-10, 15c (SC9 136-/54-188, n = 8), slightly fewer in

males (SCrf 1 23- 13 7- 1 62, n = 8); approximately Octopus inemhranacetis (non Quov and Gaimard. 70 suckers on hectocotylizcd arm. Terminal 1832). — Lu and Phillips, 1985: 33. organ large and recurved (TOL1 about 40). Sper- Material examined. 137 preserved specimens exam- matophores very long, up to 1.5 times mantle ined in Australian museum collections and the length and produced in low numbers (SpN National Museum of Natural History, Washington < 10). Eggs moderately small, ovarian eggs to 3.9 (USNM). The material listed was used to generate the mm [ELl(ov) to 7.3] and produced in large num- description. bers (>5000) Holotypc: Qld: lef: 46.7 mm ML. NMV F60I43. Moreton Bav. 3 mi (5 km) N of E corner of Mud l„ Description. The following description based on (about 27'23'S, about I53°15'E). W. Stephenson. 9 9 mature males, and 6 submature and 4 mature Dee 1966. females. Counts and indices in Tables 3 and 4, Paratypes: Qld: I* 39.8 mm ML. QMB Mo29473, with data from an immature male (1 7.1 mm ML, off Cairns, about 15 km SE of Fitzroy L. (about QMB Mo29469). 17WS.about 1 46°05'E). 30m. C.Jones, 25 Apr 1982 Small lo medium-sized robust species (figs le, (Seibenhausen net): lo: 41.5 mm ML. USNM 817673, for both sexes, TL 7a, I Oa): ML to at least 50 mm Moreton Bay. 3 mi N of E corner of Mud I., (about to at leasl 200 mm; weight to at least 75 g. Mantle 27°23'S, about 1 53°15'E), W. Stephenson. 9 Dec 1966; round to ovoid (MW1 56.2-67.4-79.3), mantle I9: 41.8 mm ML, NMV F60105. Gulf of Carpentaria. not found. Pal- 130 km WofPnnceofWalesL. I0°56'S, 14(r55'E,44 walls thick and muscular. Stylets m. C.C. Lu. 10 Sep 1982 (trawl. 1900-2000 hr): I?: lial aperture of moderate width, approximately 48.0 mm ML, QMB Mo29329. Moreton Bay, Peel 1.. half mantle width. Funnel muscular and broad 53*1 J.M. Horseshoe Bay. (about 27'23'S. about 1 5'E. based (FLI 3 1.8- JS. 0-45.0) with free portion F60I07, Raven. 9 Oct 1975'. I9: 53.3 mm ML, NMV usually greater than half funnel length (FFLI Musgravc. Gulf of Carpentaria, 50 km W of Port 40.4-57,5-78.6). Funnel organ W-shaped with 11'57.5'S. 141'22'E. 38 m, C.C. Lu, 7 Sept 1982 broad limbs (fig. 7b). Outer limbs slightly- (prawn trawl, 2052-2137 hr). shorter than median limb (FOl 79.4-S9.6- Other material: Qld: ttf: 17.7 mm ML. QMB Funnel organ large, approximately two- Mo29469, Moreton Bay. Middle Banks, (about 97 2). funnel length (FOL1 53.3-67.0-85.9). 27'23'S, about I53"I5'E), W. Stephenson. Mar, 1974; thirds of Moreton moderate width (HW1 41.7-50.7- 24. 3e 27.5-41.5 mm ML. NMV F60104, Head of smallest Bay (about 27'23'S, about 153'15'E). M. Potter. 22 60 7) narrower than mantle except in E ol dis- Jul 1981; 1* 38.9 mm ML. QMB Mo29468, specimen (HMWI 53.3-74.9-91.7). Neck m. Queensland Cairns. 17'03.0'S. I46°07.8'E. 30.6 tinct, slightly narrower than head. Eyes large and "Southern Ocean". 28 Jan Fisheries Service. FRV slightly pronounced. with tickler chains); lc': 1981 (single rigged |rv nets Arms of moderate length, approximately 2-3 of Carpentaria. 175 39 3 ML, NMV E60 1 06. Gulf mm times ML (AMI 214.5-245.2-285.4). robust l„ IO"27'S. 140'45'E. 47 m. C.C. km W of Thursday in cross 1 1), roughly circular 2tf; 41.7 ( AWI 3.3-/6.S-20. Lu, 13 Sep 1982 (trawl. 1900-2000 hr); 41.3, tapering rapidly to line tips in distal mm ML. NMV F60I03, Moreton Bay, (about 27'23'S. section and arms shortest, lateral and ventral about 153°I5'E). Jan or Feb 1982. third. Dorsal 352-88. Dampier more robust (AF generally WA; lv: 24.4 mm ML, WAM arms longer and Wil- Archipelago, 5-6 mi orfBrazom I.", 43 in. BR. 4.3.2.1 or 3.4.2.1). Suckers of moderate size, dredge on son FV "Daveno", 5 Jun 1960. (honolulu slightly elevated with distinct radial cushions, F60I09. North sand)' V 23.6, 31.1 mm ML, NMV rim scalloped and often incurved. Mature males 17-55.5 E, 42 m. CSIRO, 22 West Shelf 19'55.5'S, 1 possess 2-4 considerably enlarged suckers at trawl. 0845hrs); I5: 44.5 mm ML. Apr 1983 (beam level of third sucker pair on arms 2 and 3 (SDIrf F60I08. 50 km off Wickham, 20*15 S, NMV 10.2-y

5 mm

20mm 2 mm

Figure 7. Octopus c.xaninilalus sp. nov. a, dorsal view of 41.5 mm ML female (NMV F60104): DMR = dorsal mantle ridges; DWS = dorsal white spols; FWS- frontal white spot, b, funnel organ of 4 1.7 mm ML male (NMV F60I03). c, copulatory organ of same specimen. OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 323

Figure 8. Octopus exanmtlatus sp. nov. a. digestive tract of 37.2 mm ML male (NMV F60I04): hatched line indicates extent of'ink sac embedded within digestive gland (for key to symbols see fig. 4). b. upper beak, lateral view of 34.9 mm ML female (NMV F60104). c, lower beak, lateral view of same specimen, d, lower beak, ventral view of same specimen.

in females (SCo 1 36-/54-1 88, n - 899), slightly Third right arm in males hectocotylized and (not significantly) fewer in males (SCo" 123- slightlv shorter than opposite arm (OA1: 74.6- 137-162, n = idd). Webs moderate to deep 83.0-97.6; HAMI: I58.6-/,V7.5-218.5). Sper- (WDI 24.3-2P.S-37.1), deepest laterally, dorsal malophore groove well developed and wide with web distinctly shorter (WF typically D.C=E.B.A fine transverse ridges. Spermatophore guide or D=C.E.B.A). Web margins well developed on deep with elevated square papillae. Mature ventral edges of arms, extending along approxi- copulatory organ small (fig. 7c: LLI 3.5-4.3- mately 80% of arm length. 5.8). ligula roughly conical with blunt tip; groove 324 MARK D. NORMAN

5mm

I 10mm 1

10mm

Figure 9. Octopus exummlatus sp. nov. a, male reproductive tract of 37.2 mm ML specimen (NMV F60104)- for key to symbols fig. see 5.; A = appendix, b, spcrmatophorc of 39.8 mm ML specimen (QMB Mo29473)- SR = sperm reservoir, c. ovary of 44.5 mm ML specimen (NMV F60108): = : distal DO oviduct; = ovary; OG • oviducal gland; PO = proximal oviduct. OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 325

wide and shallow with a raised longitudinal mid- from tip of oral end in all preserved material rib. Calamus medium size, slightly raised (CLI examined. Sperm cord very fine and irregularly 19.4-26.2-34.2). Approximately 70 suckers on packed within the sperm reservoir, not forming hectocotylized arm (HASC: 62-69-77). regular whorls in material examined. Cement Gills with 7-8 lamellae on both inner and body not distinct in material examined. outer demibranchs. plus a terminal lamella. Mature ovary oval shaped (fig. 9c). Oviducts Digestive tract illustrated in figure 8a. robust and of moderate length. Oviducal glands Anterior salivary glands large, approximately large with approximately 1 5 radiating (braiding) 60% of buccal mass length. Posterior salivary chambers. Eggs small, ovarian eggs to 3.9 mm glands well developed, slightly larger than buccal long [ELI(ovarian) to 7.3], moderately wide mass. Crop diverticulum distinct but not greatly [EWI(ovarian) to 1.9] and produced in large developed. Stomach bipartite. Caecum coiled in numbers (at least 5000 ovarian eggs estimated single whorl. Muscular intestine rellexed from 1 specimen). 4-5 follicular folds on approximately one-third of intestine length ovarian eggs. from proximal end. Ink sac well developed, Live specimens not witnessed. Preserved embedded in ventral surface of digestive gland specimens exhibit 4 broad dark brown to black below iridescent tissue layer. Anal flaps present. longitudinal stripes on dorsal body, over cream Upper beak (fig. 8b) with moderate hood and to red-brown base colour (fig. 7a). Inner pair of slightly hooked rostrum, concave on cutting stripes extend along length of body joining at edge. Ventral edge of hood with regular vertical midpoint above eyes, continuing as single ridges. Lower beak (figs 8e-d) with blunt ros- medial stripe to margin of dorsal web. Lateral trum, narrow hood, widely spread wings and body stripes commence from midway along lat- slightly flared lateral walls. Ventral view of pos- eral mantle, pass through the eye and extend on terior margin moderately concave. Radula with arm crown to margin of dorsolateral web. Thin seven teeth and two marginal plates in each dark brown to black lines extend along bases of transverse row (figs lOe-f). Rhachidian tooth suckers on dorsal faces of all arms. Plain dark with 1-2 lateral cusps, typically I. on either side brown to black ocellus present on each side of of a short robust medial cone. Lateral cusps are arm crown between bases of arms 2 and 3 (figs in symmetrical serialion. migrating from lateral lOc-f: OcDI of black spot 14.8-20.6-28.8). Ring to medial position over 2-3 rows. First lateral of iridescent tissue absent. teeth unicuspidate with cusp towards lateral Several small circular pink spots arc visible edge: second lateral teeth unicuspidate and long over base colour. I pair on dorsal body approxi- with curved base; lateral marginal teeth long, mately one-third along body from level of eyes straight and robust; marginal plates oblong and (fig. 7a). Circular frontal white spot on dorsal plain. arm crown present just below eyes, in centre of Male reproductive tract illustrated in figure medial dark stripe. Pigmentation reduced on 9a. Terminal organ in mature males very long, ventral body and arm crown surfaces, but well thin and recurved (TOLL measured from lip to developed on oral surface of dorsal web. apex of curve: 34.1-4/. J-48. 1) with robust Body, arms and arm crown covered in small diverticulum (DLI 27.3-i4.4-38, 1), genital aper- oval to round papillae, extending to arm faces ture subtcrminal. Vas deferens relatively robust and oral surface of dorsal and dorsolateral webs. and short. Mucilaginous gland enlarged at point Sculpturing present on ventral surface of body, of attachment to vas deferens. Spermatophoric but papillae almost twice diameter of those on dorsal surfaces. Sculpturing absent from ventral gland I elongate and narrow with large recurved coil approximately 80% along length. Spermato- arm crown. 4 longitudinal raised skin ridges vis- phoric gland II long and narrow with reflexcd ible on dorsal body, in diamond arrangement tip. Appendix present at junctions of spermato- (fig. 7a). Single large branching papilla above phoric glands and Needham's sac. Needham's and slightly behind each eye, surrounded by sac greatly elongated and narrow curved to fol- additional smaller papillae (fig. 10a). Raised low curve of spermatophoric glands. Spcrmato- papilla present in centre of frontal white spot on crown. phores (fig. 9b) very' long (SpLI 136.8, 152.3, dorsal arm

in low- Photographs of a live juvenile (1 5. 1 mm ML. 1 62.4) and fine (SpWI 0.8- I.I), produced numbers (

326 MAKRD. NORMAN

—1 -^ vo ** rq ' ^ rn 06 NO rj

fr r f 000,,0,0 ^^' ~ o:i I ON tS^! ! 1^ 1^"! * hMfN toc '-Nt NO Q 00 "- - Q *£ 12 „Mddjn-im'd«6^ fi t-.' Tf on' no o< vo — r- I no ^r ^MSr;,^~ '0»iooominCM r-i on r~ — m ^ m tn _. rN rNj rN rN) — _ ,

~ r — c '. "^ ~ "KlOMK- l^rNi— — 1 >o "^ 2 8 ^ '. i ? . iomnoo»U(NnhO- lr , ' "!f' i--' u-i ^ sr ob no ooVi o no ~ o\ vi _" 25 5 ASS:IC!!G2ttli^fNO «* t*s ON ^t OO <~) — H Q ^f o rH (N 00 r- rs Tf cnJ trt t Nt 00 ^D(N O 2£ g ^^^^^^^^^NOôt^QOrôoNOfNlOIÔtNO I inm« vcr-. i/-i Z^ s "T __ 00 — o st'no r-' 06 10 06 o r»~ t> ^ tr) r4 I I I •T o M omhcoONOvn- — tn onoo r«-i t+ m -~ r-l H — c*l F-t

- r- rf r* } rn >sO crs —' r" ^o i> fn iv-i ' 2 * M 3 so rH ri ri rn en —< \o ^d Co b, 2t

a eft >o ^fr (N w-i vo io 2n — — I O^ ^ od n-i ' r-' 06 ^ 25 3 ^ ^ O — m" t-~' rn Q\ Tt nn II -* in a Zl?

u DO c 2 no>nto\ i ^° ^^oon^o^m'odad^'^ co ro rn n 06 -! ' 1 r-- t vo n o-, n _ ,^,-si^,-^ —i a\ Mrt m Co

50. «"> r^ in i> ov r* ^d^r^da^a^d^r-^ cK 00 ^° -) -- fc< !—

Co — —1 —<

OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 327

~ r^s fvi r»~i l/"~i —« ** i/^i i/~i r-~. t+ r-- ^— 3? ON C] O OO — ~ <"*"> **! Ov «n CN

o c/3 — ooh^Oh^wnrnmoo \o -< ^ (^"O

Co Sa — — (N rj 5?-.

-^0000^|-r>-;^O'^t-'s000r^00<^ lOOr>100 — Tfr^i

: —^ 2 00 —' > > o iri r*i lo -rf r- Tf ^ vo 00 —1 r

. CD Dh p B5 O O r- —< ^o r- 9 i> r~-' rn 2S "^in^t^-ONrs»n>n-^-MfNtN rs ^n fS o so

r 'OMN^^rr)O vC |A^ ,t'Nooirih-OoooOfN( iON ^oo^ta^oodo^^tcocKoN in — £5 — r-^ r--' o »n ' — \£> >n r~- t^- o O —1 —'

>o OM/l'to^-^MfncocM^rnoo rfr r^ TJ" 06 MD Ci ^oincO'OONO^- — (N rn in c-> r- -h z £

00 o on O m

' — fi r*~i \o m oo so c> O ' — SO - - o >— •— sd 00 < GO m so — -HfNrsn

Q "* *K "0 in CTsri 06

'— Tf r- so

tONQQM •— > O |— *

3 25

r/i 3* S* 328 MARK D. NORMAN OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 329

Figure 10. ('holographs of preserved specimens of Octopus exannulatus sp. nov. a, dorsal view of 4 1.5 mm ML malelUSNM h. 81 7673). lateral view of 4 1.8 mm ML female paratypc(NMV F60l05).c. lateral view of 46.7 mm MLmaleparatype(USNM8l7673).d.dorsalviewof4l.8mmMLfemale(NMVF60I05).e,ocellusof4l.8mm

ML female paratype (NMV F60I05). f. ocellus ol 46.7 mm ML male paralype (USNM 81 7673). g, i, radula of 34.9 mm ML female (NMV F60I04). h, enlarged suckers of 41.5 mm ML mature male (USNM 817673).

ation consisted of red-brown longitudinal bars Material examined. 2 live O. mototi were encountered over white base colour on offshore islands at the southern end of the Great Barrier Sexual dimorphism is not marked in this Reef. 8 preserved specimens were examined in species. Both sexes attain approximately equal Australian museum collections, the National Museum of Natural History. Washington and the Museum maximum size. Mature males possess 2-4 National d'Histoiie Nalurelle. Paris. enlarged suckers on arms 2 and 3 at level of third Holotype: Qld: h* 66.2 mm ML, NMV F6010I. sucker pair (SDlcf mean 14.3; 9 mean 8.4). Capricorn BunkerGroup, Heron L, 23°50'S, 1 52°25'E, Females appear to possess more suckers than 31 m. M. Norman and R. Fenwick, 30 Aug 1990 males (SCo 136-/54-188; rf mean 123-7J7-162. (raised in pot at 0945 hr. on sandy bottom in channel difference not significant at P = 0.05 level). on N side of island). Paratypes: Qld: l»: 58.6 mm ML. NMV F60I02. Distribution. Octopus exannulatus is recorded Capricorn Bunker Group, One Tree I., 23°30'S. from tropical Australian waters (fig. 15c). from I52°05'E, I m, M. Norman and R. Fenwick, 7 Sep Shark Bay, Western Australia (24°49'S, 1990 (deep in lair, 1415 hr. in dead coral within

I I3°33'E) to Gulf of Carpentaria, Torres Strait lagoon, on sand. Hushed with CuS0 4 ): I9: 73.3 mm and south to Moreton Bay. southern Queens- ML, AM C 1 54277, Coral Sea, Wreck Reef, NE of West land (27°15'S, 153°15'E). Islet, 22"12'S, I55°10'E, 12 m. P. Cook and B. Bately, 30 Oct 1983 (in lair during day, under dead staghorn Life history. It has been collected from exposed clump on sand, caught by hand on SCUBA); I9: 76.7 intertidal mudflats to 84 m deep, on muddy, mm ML. QMB Mo29325, Swains Reef, 2I°46.9'S, sandy and shelly sand substrata. This species I52°50.0'E. 54 m. C Jones, Swains RecT Survey Station 12. 27 July 1987 (trawled with appears to inhabit open bottom substrata and Seibenhausen net). seagrass beds. Austral Is: IS: 70.5 mm ML, USNM 817681, Rapa Prawn trawler operators fishing within Great I., about 27"36'S, I44°20'W, G. Paulay, 6 Jun 1980 Barrier Reef waters report catches of this dis- (captured by local islanders). tinctive species in trawls on sand or muddy sub- Other material: Qld: \&. about 55 mm ML (in two strates. pieces, arm crown with intact arms and bulk of body; head missing), QMB Mo29466, no locality data; lrf; Commercial exploitation. Octopus exannulatus 60.6 mm ML, NMV E60I42, off Caloundra, 26'40'S. occurs in fairly low numbers in the by-catch of 153°16'E, 22 fm (40.3 m), Adam Butcher, QDPI, "San Moreton Bay. Great Barrier Reef and Gulf of Antone", 22 Jul 1991 (prawn trawl, just before Carpentaria trawl fisheries, particularly prawn dusk). fisheries. This species is often retained for use as New Caledonia: I9: 100.0 mm ML, MNHN 2010, off I66°40'E), bait. No information is available on scale of Noumea, (about 22°30'S, Alan Ger-

bault.nodate: 1 9: 100.5 catch. mm ML, MNHN 201 1, North Lagoon, (about 22\30'S. 166°40'E), 1990. Etymology. From the Latin, ex (without) and Diagnosis. Moderate sized species with black annuius (ring), referring to the simple black oval ocellus containing iridescent blue ring. Cir- ocellus which lacks an iridescent ring. cular cluster of dark spots above each eye forms Remarks. Lu and Phillips (1985) reported this "flower" pattern. Alarm colour pattern of white species under the name O. membranaceus Quoy base colour and 6 dark maroon longitudinal and Gaimard. 1 832. Despite the poor condition stripes on dorsal body and arm crown. 9-1 1 gill of the type of O. membranaceus, it clearly pos- lamellae per demibranch, typically 1 1. Approxi- sesses an iridescent ring within its ocellus, easily mately 160 suckers per arm, roughly equal in distinguishing it from O. exannulatus. number in both sexes in the material available (SCV 149. 176; 9 143-/59-172, n = 499), Octopus mototi sp. nov. approximately 100 suckers on hectocotylized Figs If, 11-14, 15d arm. Terminal organ short and robust (TOLI about 20). Spcrmatophores of moderate length Octopus membranaceus. — Loch, 1987: 8, tcxlfigs. (SpLI about 70) and produced in low numbers (non Ouoy and Gaimard. 1832) 330 MARK D. NORMAN

5mm

20mm

Figure 1 1 . Octopus mololisp. nov. a, dorsal view of 73.3 female = mm ML paratype (AM C 1 54277): DMR dorsal mantle ridges; DWS = dorsal white spots; FWS = frontal white spot, b, funnel organ of 58.6 mm ML female paratype (NMVF60 102). c. copulalory organ of 66.2 mm ML holotype(NMV F60101) d copulatorv organ of 70.5 mm ML paratype (LISNM 817681). OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 331

10 mm

Figure 12. Octopus moteti sp. nov. a, digestive tract of 58.6 mm ML female paratypc (NMV F60102): hatched line indicates volume of ink sac embedded within digestive gland (for key to symbols see fig. 4). b, upper beak, lateral view of same specimen, c. lower beak, lateral view of same specimen, d, lower beak, ventral view of same specimen.

numbers (SpN <10). Eggs moderately small, females, TL to at least 320 mm; weight to at least ovarian eggs to 6 mm, spawned egg capsules to 300 g. Mantle ovoid (MWI 52.8-64.3-10.4), 3.2 mm [ELI(ov) to 7.8, ELl(sp) to 4.2], pro- mantle walls thick and muscular. Stylets not duced in large numbers (EN > 10 000). found. Pallial aperture of moderate length, approximately half mantle width. Funnel mus-

Description. Based on 3 mature males, and I cular and broad based (FLI 34.5-J7./-40.0) submature/mature and 4 mature females. with free portion usually greater than half funnel Counts and indices in Table 5 and 6. length (FFLI 43.5-64.0-82,6). Funnel organ Moderate sized, robust species (figs If, 11a): W-shaped with broad limbs (fig. lib). Outer slightly shorter than median ML to at least 70 mm for males and 100 mm for limbs limb (FOl 332 MARK D. NORMAN

3 "°V - ' SP a ' male re - ducti ve tract P™ of 66.2 mm ML holotype )• fjmhnll °f*T T°"' (NMV F60 1 1 for kev to° Spermat hore from I VcX CT-caJtE'sR ^ °P sa ™ ~en. b, entile spermatopnor°s CB OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF. AUSTRALIA 333

Table 5. Counts and indices for male Octopus mototi sp. nov. (M = mature; D = damaged; — = not recorded; ID = indistinct; # = specimen frozen, hectocotylized arm distorted at tip; * = OcDI measured as diameter of iridescent ring).

Museum NMV USNM NMV QMB Reg. No. F60101 817681 F60142 Mo29466 (Holotype) (Paratype)

ML 66.2 70.5 60.6 -55 StM M M M M TL 255 260 D -192 MWI 70.4 65.7 52.8 D HWI 52.7 46.4 42.7 D HMW1 74.8 70.6 80.9 D -201.8 AMI: 1 226.6 228.4 D 2 271.9 235.5 D — 232.7 3 268.9 249.6 D D H 250.8 217.0 236.0 -221.8 4 249.2 252.5 D -258.2 -12.4 AW1 18.6 14.9 13.4 SDI 13.0 9.4 10.1 -6.9 WDI 25.6 29.8 D D 10 GC 11 11 10 -221.8 HAMI 250.8 217.0 236.0 OAI 93.3 86.9 D D HASC 100 106 95 103 -2.0 LLI 5.4 3.1 6.7# CLI 36.7 48.9 54.2# D -15.5 TOL1 25.7 20.0 24.9 49.4 DLI 37.1 30.5 46.4 — SpLI 70.2 59.4 D SpWI 1.3 1.2 D SpRI 34.8 35.3 D SpN 5 6 35.9 34.5 D FLI 40.0 D FFLI 82.3 43.5 49.3 86.6 100 D FOl ID D FOLI ID 56.1 63.6 — 15.6 OcDI* 14.4 15.9 18.2

pairs longer and more robust 85.5-9/.0-100.0). Funnel organ large, approxi- eral and ventral typically 4=3.2. ). Suckers of moderate size, mately 60% of funnel length (FOLI 56.1-6/./- (AF 1 slightly elevated with distinct radial cushions rim. Suckers approximately equal Hcad of moderate width (HWI 46.4-49.3- and scalloped avail- mantle (HMWI in size for both sexes in limited specimens 56 6) always narrower than about 6.9, 9.4, 10.1, 13.0; 9 10.5- 70.6-76.7-81.1). Neck distinct, slightly able (SDW The holotype exhibits slight enlarge- narrower than head. Eyes large and slightly ll.3-\ 3.6). suckers at level of fifth to sixth pronounced. ment of several sucker pair on arms 2 and 3. Arms 3 or 4 possess Arms of moderate length, 2.5-3 times ML, examined most suckers, approximately 160 per arm, slightly longer in females of material roughly equal in number in both sexes in avail- (AMIcf about 258, 268.9; 9 261 .7-292.2- 252.5, 143-/59-172, n = (AWI 10.5-/5.5-24.6), able material (SCrf 149, 176; 9 314.0). Arms robust 25.6-30.4- rapidly to 4). Webs moderate to deep (WDI in cross section and tapering square dorsal web distinctly shortest, lat- 38.8), deepest laterally, fine tips in distal third. Dorsal arms 334 MARK D. NORMAN OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 335

Figure 14. Pholographs of Octopus mototi sp. nov. a-e, colour patterns of live animals: a. resting colouration of

66.2 mm ML male holotype(NMV F60 1 1 ); arrows indicate dorsal white spots, b, alarm coloration of 58.6 mm ML female paratype (NMV F60I02). c, wild animal in Coral Sea. emerging from conch shell displaying eye bar and "flower" pattern above each eye (Photo courtesy of Neville Coleman), d, posture of arms drawn back along body exposing suckers, beak and displaying frontal white spot and spotted oral surface of dorsal web (58.6 mm

ML female paratype: NMV F60I02). e, alarm colouration of same specimen, f, colour pattern in preserved 73.3 mm ML female paratype (AM C 1 54277). g, preserved ocellus of same specimen, h, preserved ocellus of 76.7 mm

ML female paratype (QMB Mo29325). i, festoon of spawned eggs from same specimen, j, k, radula of 58.6 mm ML female paratype (NMV F60102).

shorter (WF typically D.C=E.B. A). Web margins conical; groove wide and shallow bordered lat- extend along approximately 60% of ventral erally by fine skin ridges. Calamus medium size edges of arms. and slightlv raised (CLI 36.7, 48.9). Copulatory Third right arms in males hectocotylized and organ on male from Rapa I. (USNM 817681) slightly shorter than opposite arm (OAI: 86.9, slightly shorter and broader (fig. lid) than that 93.3; HAM1: 217.0, about 222, 236.0, 250.8). of holotype, but still bears distinctive skin ridges Spermatophore groove well developed and wide bordering ligular groove. Approximately 100 with fine transverse ridges. Spermatophore suckers on hectocotylized arm (HASC: 95-101- guide distinct with cross ridges but no obvious 106).

papillae. Copulatory organ small (fig. 1 lc: LLI Gills with 9-1 1 lamellae, typically 11, on both 3.1. 5.4 for 2 mature males, 6.7 in distorted inner and outer demibranches, plus a terminal frozen specimen, NMV F60I42), ligula roughly lamella.

Table 6. Counts and indices for female Octopus mototi sp. nov. (S = submature; M = mature; D = damaged; — = not recorded; ID = indistinct; # = from ovarian eggs, capsule of spawned eggs much shorter; ## = counted from festoons of spawned eggs; * = OcDI measured as diameter of iridescent ring).

Museum NMV AMS QMB MNHN MNHN Reg. No. F60102 C154271 Mo29325 2010 2011 (Paratype) (Paratype)

ML 58.6 73.3 76.7 100.0 100.5 StM M S/M M S/M M TL 228 290 324 425 366 MW1 69.6 65.3 69.8 D 56.5 41.9 HWI 51.9 52.7 56.6 D HMW1 74.6 80.7 81.1 D 74.1 252.4 D 271.0 210.0 AMI: 1 240.6 294.0 D 2 261.1 274.2 286.8 305.1 293.0 261.7 3 D 307.0 314.0 243.8 4 273.0 286.5 D 12.8 AWI 16.2 10.5 24.6 16.1 10.5 10.5 SDI 11.6 10.5 13.6 38.8 D WDI 26.9 32.0 29.5 9 10 10 GC 11 10 — — ELI — 7.8# — 1.7# EW1 — z EN ~10 700## >>1000 36.7 D FLI 39.2 38.5 34.6 58.3 D FFLI 73.5 58.2 82.6 ID ID ID FOI 91.8 85.5 ID ID ID FOL1 63.9 61.0 14.3 14.0 OcDI* 13.7 19.1 16.6 336 MARK IX NORMAN

j < f' \ *.v ;V > j « •, i\ \ • K s / • • \u \V .* • 5 «» J • • • %P$.. f. fc a.* • \ ss • • • • «« • /. / 7 20'S U •* • } f • L

60°E

*s < *\> ^ ' \ 1 - \ ^ s° ^ \ > y > X y s Vs s hh ^ •0 J ^6 > a-a. *[>V

- /Y, x , 20° S u ^ \ '• } • . ( ^- f Y X / v^ r-i | 2

a. Gray, 1849: • = materia] Figure 1 5. Distributions of Great Barrier Reef ocellate octopuses, Octopus cyanea examined, b, Octopus polyzcniti Gray, 1849: A type locality; • = other material examined, c. Octopus exan- = nulatUS sp. nov.: A - type locality, • = other material examined, d. Octopus motoli sp. nov.: A = type locality, • other material examined: o = photographic records. OCELLATC OCTOPUSES OF THE GREAT BARRIER REEF. AUSTRALIA 337

Digestive tract illustrated in figure 12a, Mature ovary oval shaped (fig. 13d). Distal Anterior salivary glands of moderate size, oviducts robust and reflexed midway along approximately half buccal mass length. Pos- length, extending to level of anus and flattened terior salivary glands well developed, ap- distally. Proximal oviducts originate laterally on proximately equal in length with buccal mass. ovary, not obviously from common origin in Crop diverticulum distinct. Stomach bipartite. specimen examined. Ovidueal glands large with Caecum coiled in single whorl. Intestine poorly 16-20 radiating (braiding) chambers. Eggs

preserved in dissected specimen, reflexed at (fig. 1 3g) medium sized, ovarian eggs to 6 mm proximal third, rectum not distinct. Ink sac well total length [ELl(ovarian) to 7.8], spawned eggs developed, embedded in ventral surface of with smaller capsule to 3.2 nun [ELI(spawned) digestive gland. Anal Haps present and line. to 4.2|. Eggs moderately wide JEWl(ovarian) Membrane on dorsal surface of visceral mass 1.7; FWI(spawncd) 2.0] and produced in large pigmented with large dark chromatophores, numbers (about 10,700 eggs accompanied 76.7 remnants of larval or founder chromato- mm ML female. QMB Mo29325). Eggs laid in phores. long branching festoons (fig. I4i). Approxi-

Upper beak (tig. 1 2b) with moderate hood and mately 6 follicular folds on immature ovarian short hooked rostrum, concave on cutting edge. eggs. Lower beak (figs I2c-d) with blunl rostrum, Colour in lile variable. Ocellus on each sideol narrow hood, widely spread wings and moder- arm crown between bases of second and third ately flared lateral walls producing moderately arms, each containing a large iridescent blue ring concave posterior margin from ventral view. (figs I4e, g-h: OcDl for iridescent ring 13.7- Radula with seven teeth and two marginal plates /5.9-19. 1). Iridescent ring masked in some in each transverse row (figs I4j-k). Rhachidian colour patterns, exaggerated by dark back- tooth with 1-2 lateral cusps, typically I. on ground in alarm colouration (fig. 14e). either side of a robust medial cone. Lateral cusps Resting animals exhibit orange-cream base are in symmetrical seriation. migrating from lat- colour with darker brown spots in six longitudi- eral to medial position over 4-6 rows. First nal series along dorsal body (fig. 1 4a). Cluster of lateral teeth unicuspidate with cusp towards lat- 5 large spols around a central spot above each eral edge: second lateral teeth unicuspidate and eye. form 'flower" pattern (lig. 1 1 a, 14a). long with curved base: lateral marginal teeth '"Flower" pattern often visible in specimens long, straight and robust; marginal plates oblong within, or partially emerged from, lairs (fig. 14c). visible dorsal and plain. Circular frontal white spot on arm Male reproductive tract illustrated in figure crown just below eyes, containing thin erectile dorsal white spots 13a. Terminal organ in mature males moderate papilla(fig. I la). Small pairof dorsal visible over darker to large and robust (TOLI about 15.5. 20.0. 24.9, midway along mantle

I 14a). 25.7). diverticulum moderate sized (DL1 30.5- resting base colour (fig. la, striped colour 411.9-49.4). genital aperture subtcrminal. Vas Alarm display of distinctive

1 solid dark deferens thin and of moderate length. Mucilagi- pattern over while base (lig. 4b. e). 6 longitudinal stripes form on dorsal body nous gland enlarged at point of attachment to maroon extending on to arm crown and down ventral vas deferens. Spermatophoric gland I robust Median pair of dorsal stripes with large recurved coil approximately three- edges of arms. dorsolateral pair continue quarters along length. Spermatophoric gland II extend to near eyes, down ventral edge ol arm 1. and long and robust, tip reflexed. Appendix present through eyes lateral pair cease at pallial aperture. Additional at junction of spermatophoric glands and Need- medial stripe commences between eyes, divides ham's sac. Needham's sac moderately short, each side of frontal white spot and rejoins to tapering to line point at posterior end. Sperma- continue to margin of dorsal web, line stripe tophores (fig. 13b-e) of medium length (SpLI continuing down dorsal edge ofarm I. Dark oval 59.4, 70.2) and line(SpWI 1.2. 1.3). produced in spot can be expressed under iridescent ring of low numbers (< 10 in Needham's sac of holo- Oral ocelli, extending distally lo form wide stripe typc;4 in Rapa I. male. 6 in NMV F60I42). down ventral edge of arm 2 and line stripe down cap simple, showing slight restriction below tip. dorsal edge of arm 3 (fig. 1 4e). "Flower" pattern bearing long cap thread. Sperm reservoir above eyes sometimes pronounced in approximately one third of spermatophore of spots alarm displays, other times suppressed. Alarmed length (SpRI 34.8, 35.3). Sperm cord coiled in individuals often retreat into crevices or corners approximately 75 coils (holotypc: 76; Kapa I. and raise arms over body lo display frontal white male: 75). 338 MARK D. NORMAN

spot, suckers and beak. In this posture, dark exoskeleton fragments. The lair of one specimen spots arc clearly visible against white base on collected from One Tree I. lagoon was sur- oral face of dorsal and dorsolateral webs (fig. rounded by discarded gastropod shells. It is dis- 1 4d). This characteristic pattern is clearly visible likely that O. mount preys on hermit crabs, on individuals deep within lairs. carding empty gastropod shells outside the Skin sculptured in patch and groove system mouths of lairs. Loch (1987) documented the consisting of oval to round patches (fig. 14a), shells surrounding the lair of the Swains Reef extending on to oral surface of dorsal and dorso- specimen (AM C 1 54277) which included 1 5 gas- lateral webs and arm faces. Raised erectile tropod species, predominantly Fusinus undatus papilla in centre of frontal white spot distinct in and Stromhus spp. Loch noted many shells most colour patterns. 4 longitudinal skin ridges exhibited two drill holes, one produced by nati- in diamond arrangement on dorsal mantle (fig. cid gastropods, the other by an octopus. Loch

1 la). Single small supraocular papilla above and suggested that this was consistent with this slightly behind each eye in centre of "flower" octopus species drilling these shells to prey on pattern of spots. secondary occupants, namely hermit crabs. Skin sculpturing and some colour patterns are Eggs are laid in large numbers in festoons (fig. visible in preserved specimens (figs I4f-h). 14i). It is not known whether eggs arc attached to Sexual dimorphism is not marked in the few the substrate or carried in the web of the female. specimens examined. Females attained largest This species lays small eggs indicating hatchlings size in available material, 100 mm ML versus 70 are planktonic. mm ML in males, females appear to have Octopus mototi may prove to be venomous. slightly longer arms than males (AMlcf 252.5. This species has a prominent warning colour- about 258.2, 268.9; 9 261.7-292.2-314.0). ation and is known locally on Rapa I. as the "poison octopus" (G. Paulay, pers. comm.). The Distribution. Octopus mototi is reported from two specimens encountered in the field showed a eastern Australian waters: Capricorn-Bunker willingness to bite objects such as aquarium Group, southern end of the Great Barrier Reef; nets, a behaviour not observed in other Great Swains Reefs and Wreck Reef, Coral Sea. Barrier Reef octopuses. Neville Coleman (pers. comm.) provided an additional record of Ihis species from Saumarez Etymology. On Rapa I., South Pacific (27°36'S.

Reef, Coral Sea (21°45'S, I 53°40'E), photo- 1 44°20'W) the local name attributed to this octo- '<*" graphing an individual at 10 m on sand occupy- pus is "fe 'c mototi". "fi meaning octopus and ing a large gastropod shell (fig. 14c). This species "mototi" meaning poison, referring to the is also recorded from the South Pacific (fig. 1 5d): reported poisonous nature of this distinctive

New Caledonia (22°33'S, 1 66°40'E) and Rapa I. species (G. Paulay, USNM, pers. comm.) (noun (27°36'S. I44°20'W). Dr Okutani (Tokyo Uni- in apposition). versity of Fisheries) has a 77 mm ML female specimen of this species from Okinawa, south of D Japan (26 30'N, I28°00'E). Photographs of this Discussion specimen kindly provided by Dr Okutani show Systemcttics, Occllatc octopuses are found in two the characteristic "flower" pattern above the separate regions of the world: i) the Indo-West eyes, longitudinal rows of spots on dorsal mantle Pacific centred in the tropical waters of south- and display of black spots on oral surface of dor- east Asia and tropical Australia; and ii) the sal web. Americas, centred on the cast and west coasts of Octopus mototi has been collected from shal- central America. low subtidal depths (< 1 m) down to 54 m, on Fifteen species and three subspecies of ocel- sandy substrates near coral or coral rubble in late species have been described from lndo-West clear water. Pacific waters (Table 7). Of these many arc inad- Life history. Octopus mototi appears to favour equately described, lack type malerial or are sandy substrata, often associated with coral synonymous with others. The list can be reduced heads or rubble. Deep lairs arc excavated under to eight apparently valid taxa from these waters coral heads or coral rubble. From limited obser- (including the two new species described here): vations, this species appears to have crepuscular "iidako" (O. areolatus d'Orbigny, 1839 / O activity patterns. tangsiao d'Orbigny. 1839 / O. ocellatus Gray.

Examination of stomach and intestine con- 1 849); O. pohzenia Gray, 1 849: O. cyanea Gray. tents found a high proportion of crustacean 1 849; O. oxulum Sasaki. 191 7; O. rabsoni Stem. OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF, AUSTRALIA 339

a, a

S S

o -a c a s s o o

I I >> I I I I ll>

Cc o a X) C o

z 3 * z, 'i3 nj o S '3 M 3 — s a « c cd . (9 Cn „ ffl o. n o. o,-? 0.0.0.3," ' O M a B P3 (D (l> J™ Oh < XoiuS 'ad?

CO '£ 00 3 [ 00 t/) - 00 —. cd o 8 1 c «o5 OO — o —

o 2§" 2 g= so D <3 a

JT3

r- 3 oo — ** , ' is a 5 i S 5 O-' —. * 3 3 a a aa G G b "5 y y > i1 .^ oOOO S 00000 w •SO tZ) < S'SS -Q Si -° ^ "3 a. 1 1 g &»1 6*3 tO § 30 3 3 S 3 =3 & & s- a it -a oor 00 OOO£3 O U i 1

(40 MAKh I) NIIKMAN

1941; taivnat Oommen, 1971; exannu Polypus fang uao lypicus Sasaki, 1929; Polypus latin sp, nov., and motoU sp, oov. fang itao rtchuanus Sasaki, 1929. is I lu- presence of on iridescent ring(ie, metallic Octopus poiyzenia similar to "iidako*' in blue, gold, lilvei oi green colours produced by sharing large eggs, iridesceni i ing within ocellus iridophores) within the ocellus oi some species and moderate!) low gill counl (6 / vs 7 8), has been noted by many workers (eg., Brock, Howevet ii differs in possessing widely spaced l887;Robson, 1929; Sasaki, 1929; Adam. 1959, transverse bant on ail anus and lacks the Okutanietul I98T) Voss( 1 963) suggested thai gold/brown dumbbell-shaped patch on the head ihc presence of this i Ing depended on the nature between the eyes, characteristic of "iidako" .Hid duration ol preset vailon Adam (1973) dis' fhess loiiini patterns are consistenl on all speci-

hi i re! this Hi is mens examined >>i pofy enio i and proposed thai charactei was both and constam within and between species, Adam's "iidako" (OKI )0 of the laik-i species have been proposal holds truefot the material examined fa examined in nmv, am. QMB, nmnii, CAS, ihis study I here were no inconsistencies in the itMNiiiiiii MNHN collections) Octopus pofy several hundred specimens examined, The iri« in i,i is also distinct from this taxon In havings descent ring found in the ocellus of species such lowci hectocot) lized arm suckei counl (about 50

s i hsO poty (7i/w,iinif matoli was always visible \ 'Hi ) ami in possessing enlarged suckers on all even in poorly preserved material Similarly, ainis.ii the level ol the seventh rovi (vs. enlarged more than !00 spc< imens of tyttnta and suckers on amis ,' and 5 onl) at the third row in i'sonnulatits were examined and none possessed "iidako"), "iidako" occurs in northern hemi .in iridesi enl ring sphere warm temperate waters and conse

l its is he Absence oft he iridescent ring in > vanca quently distribution unJikelj to extend to and L'xannulalus is sufficient to distinguish Australia

Ihcse inxa from those listed above, as all ol the (2) Octopus memhranacvui Quo) and < lalmard, remaining species possess this iridescent ring 1832 has been reported widelj throughout the iniioWi'si is Several laxa require additional comment Pacific region, I he type in the

1 1 I lu-ir i" 1 "iidako" appears be a single t om Museum d'Histoire Naturelle, Paris imniin mini occflntc species hi shallow coastal waters i '-922) and was examined in Novembct 1991

i.ip.in i it is a from to long Kong, which in Japan goes small immature female ( 1.0 mm Ml ) in

1 n, in iius 1 the common name "iidako" n ischarac pooi condition Despite condition, an iri ii'iim',1 in ,i large green to gold iridesceni ring descent ring within each ocellus is sun clearly within each ocellus, ,i dumbbell-shaped head msi Mi- i in- inadequate original description and patch, ' s gill lamellae and large eggs (7 in the badlj damaged type ii" not allow further

t ;i,-,u i.iii ,iini nun) Naggs ( 1991 1 proposed thut clarification ol this species, I lu- name is here this species should in- assigned the name (> fang proposed as nomen dublum, \li previous reports 1 1 i- 1 'i 'i >ii' oi *tao n \ is 19 .u i.i ui 1 'i iii>ii',n\ derived this species and, where possible, the original from earlici Japanese descriptions of this i||s specimens need la be re-examined fhe name lias lincUve species (TentjimB, i '13; Katsuma been used in Japan and I long Kong, applied i '62) No type material was designated Octapth to "iidako" (Tryon, 1879 Cox, 1882 Vossand l) v. tuvolatus also was deset ibed bj d'< (rbigtij in the Williamson, 1971; I am and < "hiu, i s Roper, same »>nk (1839) ,nni has page prioritj rhc Sweenc; and Nauen, 1984; Khromov, 1990), original description is inadequate, having been Voss (1963) reported tins species from the

mi .i (Irs, 1 pi urn -i Philippines based a based 1 pro> ided in Ictlct from on single female specimen

ii. i. in inin s Dc but the two syntypes of arrolatiis ( 10.0 Ml . hsnm (5405) fhis specimen collected b) Oe Haan from Japan are In the wascxamined in theUSNM inOciobw 1990 ii

Royal Museum in I eiden (collection numbers lacks the iridesceni ring in the ocellus and 490 .mil 2438) rhesc types Bfe cleurl) speci appears to be an iindescribed species, mens ol 'iidako" rhc Inttet name is therefore iiu-u- arc two reports of <> nwntbwnoci the valid scntot synonym but the status of these from \ustrolian waters, Odhner (191 7: 12, *0) two names requires further clarification In this reported ilus species from the north west coast of discussion, iius taxon is referred to bj the vet Western Australia on the basis of three small naculai n.iiur. "ii,i,iki>" flic following nominal ocellated specimens, Utc largesi with a mantle i.iv.i ,iu- considered synonymous length of 1 1 mm No description «as provided

i, ' r iofi m sri itlattis d'( Irbigii) 1 8 19: I i and the specimens have hot been traced to date

>i ', , \i Uao l >rbignj 1839 I ., ipm i h least tout species of ocellated octopus occui

>, "'. tiu-si- i iray, 1849; i u>put h ( hinnmn. 1888 m waters and OiiiiiK-r's specimens ma\ (KM I All OCTOPUSES OF I 111 GREAT BARRIER REEF, AUSTRA1 IA Ml

belong lo one or more of these laxa. It is unlikely that this record will be resolved unless the orig- E ^ inal material surfaces. In and Phillips) 1985: 33) O | ._; o reported (>. mcinlininatcus from Morelon Hay |°2 and the Gulfof Carpentaria. These reports refer 2§E2 | | to specimens of the new species, (I exannuia

Ills. | o o

1 (3) Octopus oruluin Sasaki. I ) 17 was described 9 i o 7 6 o o as a small-egg species based on material pur- w (S >.0O — ^O Tt OO t chased from Tokyo fish Market. No subsequent records of a small-egg ocellate species in

Japanese waters have emerged and it is possible E as that Sasaki's material originated from outside JL >' 9 9*" ° '~. oo u I Pl Japanese waters. The type material may have o I been collected from tropical Indo-West Pacific waters and transported to Japan lor sale. So Sasaki's description of this species shows some O I °?,'< similarities with an undeseribed species I mm 7 O O i O Thai waters. Further study will be required lo Ov — I m:> resolve the status of this laxon. (4) Octopus rohsom Adam, 1941 was described from the Red Sea. The type material is in the 212P Museum National d'llisloirc Nalurelle, Paris and was examined in November 19') I. This o moderately small species is aligned with the

small Indo-Malayan ocellates. It lays moderately small eggs (ovarian eggs about 5 mm long}, has O oo O i- > > © t - ^> c i a. o o 10 gill lamellae per demi branch and a fine purple " 7 i <*: ": 7 iridescent ring in the ocellus. Mature males pos- o i mi "~> ~ -II rr\ sess a very long fine copnlalory organ (1 II 9.0) o and two to three enlarged suckers at the level ol Pi the third sucker pair on arms 2 and 3. with slightly enlarged suckers on arm 4. This combi- nation of characters distinguishes this species from the oecllale species of the (ileal Itarricr rl Reef. 2 (5) Octopus vannim- Ooromen, 1971 was described from four males ami two females col- o — Q a —• q in from 125-135 m oil' the west coast of lected O ' Is ° A 4 India. This species is known only from the orig- o9 o£ inal description. It possesses an ocellus with an iridescent ring, lays small eggs (ovarian eggs about 2 mm long), has 10 gill lamellae per demibranch and a long narrow copnlalory organ. No material of this species has been examined by u the author. Octopus niniiiur shows many simi-

- 1 with 0. rabsonl and further study is U -5 larities in w— Sb required to delineate these forms. II is distinct, c * —If from (deal Harrier Reef ocellate octo- ox sj Sl/1 f Table X summarises the character slates ol the •c - E-ga Reef ocellate octopuses and com •s s g e — on O ( ircat Harrier 1 , characters for other N

tionsriidako" from Sasaki. I 929 as O, RtngSion 342 MARK D. NORMAN

var. lypicus) and supplemented with COUJUS and species may prove to have a distribution restric- measurements taken from the type of O. lohsoni ted to the rim of Ihe Pacific Plate as reported for and Specimens at "iidako" IVoni NMNH and certain fish species in Springer (1 982). The eggs CAS. of O. ntototi are moderately small (ovarian eggs Great Barrier Reef ocellate octopuses fall into lo 5 mm, ELI 7.8) indicating thai hatchlings are two groups of potentially different origin. Octo- planktonic. pus polyzcnia, O. cxannulatus and O. moloti Octopus cxannulatus and O, polyzcnia exhibit show close affinities with all other Indo-Mal- distributions restricted to Ihe tropical coastal ayan ocellales in sharing: small body size, short walers of northern Australia, referred to as the arms (AMI about 200-280). body patterns that "Northern Tropical Zone" or "Northern Aus- often include four to six wide longitudinal tralian Region" by Wilson and Allen (1987). stripes on the dorsal mantle and arm crown, Many other tropical Australian marine biota modcratclv low sucker counts (SC about 100- exhibit this distribution, including fishes 200, I1ASC' about 50-100), and medium sized (Wilson and Allen, 1987), echinoderms (Rowe,

copulalory organs (1X1 about 5+). 1 985) and another octopus species. Amcloctopus Octopus cyanca, despite possessing ocelli, is liloralis Norman, 1992 (Norman, 1992a). Both clearly of different origin from the above ocel- O polyzenia and O. cxannulatus occur in shal- late species. If shares greater affinities with O low and muddy coastal waters, on sand and/or vulgaris Lamarck. 1798 and several ocellate mud substrates. These habitats are continuous species of the Americas \0. himacitlalus Vcrrill, from southern Queensland to Shark Bay, West- 1883, O. himuculoiclcs Pickford and McCon- ern Australia. Octopus cxannulatus lays moder- naughy, 1949; O, mava Vossand Solis Ramirez, ately small eggs (ovarian eggs to 4 mm long, ELI 1966 and O oculifer Hoylc, 1904 (senior syn- 7.3) suggesting that hatchlings spend at least onym of 0- rooscvclti Stuart. 1941. juk some time in the plankton. This species could Hochberg, in prep.)|. Octopus cyanca shares the potentially disperse across its range in the plank- following characters wilh many of these species: ton, and may prove to also occur along the larger body size, long arms (AMI about 300- southern Papua New Guinea coastline where 500), absence of four to six wide longitudinal similar habitats exist. Octopus polyzenia lays stripes in body patterns, high sucker counts (SC relatively large eggs (spawned egg capsules to 8 •400, HASC about 200), and liny copulatory mm long, ELI :-20) indicating hatchlings spend organ (LLI ~-2). liltle time in the plankton. In this species, dis- persal must be limited to Biageogmphy. Distributions of the four ocellate the distribution of shallow mud and sand substrata in tropical octopuses in the Great Harrier Reef region fall Aus- tralian waters. into three patterns. Octopus cvauca is a widely-distributed species Acknowledgments found in clear tropical waters of the Pacific and Indian oceans, from Hawaii in the east to the This research forms part of a PhD research African coast in the wesl (fig. 15a). This distri- program inlooctopodsofthe Great Barrier Reef bution spans the Indo-West Pacific region and northern Australia, being carried out in (BriggS, 1974). Sued a distribution is seen in a association with the Museum of Victoria and the wide range of tropical marine organisms and is University of Melbourne. Sincere thanks lo: characteristic fauna or flora of with a lengthy C.C. Lu, F,G. Hochberg, G.F. Watson and T. planktonic phase in life their cycles (Ekman. Stranks for assistance with Ihe manuscript and 1967; Briggs, 1974). Octopus cyanca lays very project; R. Fenwick, M. Turner, S. Troy. C. small eggs (2 mm long, EI,1 1,7) and the young Davies, J. Martin, J. Swartz, G. Greenwood and enter the plankton an hatching (Van lleukelem. the Breidahl family for field assistance; G. 1976). The duration of this planktonic phase is Paulay (USNM) for information on Rapa I. not known bul must he sullieienlly long bo material; M. Potter and M. Dunning, Queens- enable hafchlings to traverse extensive the areas land Department of Primary Industries for of open ocean island between groups. provision of material from Queensland waters; The distribution c\i' 0. niotott is also wide I. Loch, P. Colman. B. Rudman (AM), F. Naggs rangingdig. I5d). apparently limited to the trop- (BMNH), I. Goslincr (CAS). R. and G. ical Pacific Ocean. In the east, specimens have Bout'her-Rodoni (MN11N), L. Vail, D. White been collected from the South Pacific, the east (NTM), J. Stanisic (QMB), C. Roper and M. coast of Australia and south of Japan. This Sweeney (NMNH) and S. Slack-Smith (WAM) OCELLATE OCTOPUSES OF THE GREAT BARRIER REEF. AUSTRALIA 343

for assistance and access to collections; J. Hoey. F.kman. S.. 1967. Zoogeography of the Sea. Sidgwick Townsville Reef Wonderland Aquarium for and Jackson: London. 417 pp. access to aquarium facilities: D. Paul and Flecker. H. and Cotton, B.C., 1955. Fatal bite from C. Rowley for assistance with photographic octopus. Medical Journal of Australia 27 Aug 1955:329-332.2 plates; N. Coleman for access to his photo- figs. graphic index (AMP1). Gleadall, EG. and Naggs, F.C., 1991. The Asian occl- late octopuses. IE The validity of Octopus fang- This research was made possible through sup- siao d*Orbigny. Annals of Applied Informalion port grants from the Museum of Victoria, Vic- Sciences 16(2): 173-180, 2 figs. torian Institute of Marine Sciences. Australian Gray. J.E.. 1849. Catalogue of the Mollusca in the

Museum (Postgraduate and Keith Sutherland collection of Ihe British Museum. I. Cephalopoda Awards), Great Barrier Reef Marine Park Auth- Anlepedia. London. 164 pp. ority, University of Melbourne M.A. liartlctt Hoylc, W.E.. 1886. A catalogue of recent Cephalo- Travel Grant and Write-up Award, Joyce W. poda. Proceedings ofthe Roval Physical Society of 9: 205-267. Vickery Research Fund (Linnean Society of Edinburgh Hoylc. W.E.. 1907. The marine fauna of Zanzibar and NSW). Ethel Mary Read Fund (Royal Zoologi- East Africa, from collections made by Cvril Cross- cal Society of New South Wales), Potter Foun- land in 1901-1902. The Cephalopoda. Proceed- dation, Hawaiian Malacological Society, Royal ings of the 'Zoological Society of London 00: Society of Victoria and Western Society of Mal- 450-461. pi. XX, textfigs. 128. 137. acologists. Certain Museum of Victoria material Kalsuma. R.. 1762. Oozukushii ("Just Pishes"). Manu- referred to in this paper was collected with the script, National Diet Library, Tokyo (/ide Glead- assistance of the captain and crew of FV all and Naggs, 1991). "Susan Wright" of K.F.V Fisheries, Townsville, Khromov, D.N., 1990. Cephalopods of the Viet- namese waters: Ihe launu and distribution through financial support bv FIRTA to Dr C.C. pal- terns. 1- -9, Lu (NMV). ICES IWO/Shell, Paper 12: I fig. Lam, V.W.W. and Chiu, S.T., 1 983. Prey selection and References feeding behaviour of Octopus membranaccus

Gaimard. 1 ). 66 1 -68 , (Quoy and 832 Pp. 1 6 pis. 8 Adam. W„ 1939. The Cephalopoda in the Indian figs in: Morton. B. and Dudgeon, D. (cds). Pro- Museum. Calcutta. Records ofthe Indian Museum ceedings ofIhe Second International Workshop on 41: 61-110. the Malucofauna of Hong Kong and Southern Adam. W., 1941. Notes sur les Cephalopodes. XVI. - China. Hong Kong University Press: Hong Sur unc nouvelle espece de Cephalopodc [Octopus Kong. rohsoni sp. nov.) de la Mer Rouge. Bulletin du Loch, L. 1987. An eight armed, blue ringed, fusino- Muscc royal d'llisioue natwellc de Belgique 17 philic. Australian Shell News 57: 8-9, 2 pis.

(52): 1-5. Lu. C.C. and Phillips, J.U., 1 985. An annotated check- Adam. W., 1959. Les Cephalopodes de la Mer Rouge. list of the Cephalopoda from Australian waters. Result scientifique de Mission de Robert Ph Doll- Occasional Papers from the Museum o)'Victoria 2:

fits en Egyple /927-/W9 (3) 28: 125-193. 21-36, I fig. Adam. W.. 1973. Cephalopoda from the Red Sea. Norman. M.D.. 1991. Octopus cyanea Gray, 1849 Contributions to Ihe knowledge of the Red Sea, (Mollusca: Cephalopoda) in Australian waters: No. 47. liidleim of the Sea Fisheries Research description, distribution and taxonomy. Bulletin Station, Haifa (60): 9-47. of Marine Science 49 (1-2): 20-38, 5 figs. Berry, S.S., 1912. On Ihe cephalopods new to Cali- Norman. M.D., 1992a. Ameloclopus lilorafis, gen. ct fornia with a note on another species. Annual sp. nov. (Cephalopoda: Octopodidae). a new shal- report ofl.aguna Marine Laboratory: 83-87. text- low water octopus from tropical Australian figs 44-48. waters. Invertebrate Taxonomy 6 (5): 567-582, 5 Berry, S.S., 1912a. A catalogue of Japanese Cephalo- figs. poda. Proceedings ofthe Academy of Natural Sci- Norman. M.D., 1992b. Four new octopus species of ences of Philadelphia 380-444. textltgs 1-4, pis the Octopus macropus group (Cephalopoda: Octo- V-IX. podidae) from the Great Barrier Reef. Australia.

Brazier, J.. 1892. Catalogue or the marine shells of Memoirs of ihe Museum of Victoria, 53: 267- Australia and Tasmania. Part I. Cephalopoda. 308. Australian Museum Cutalogue 15: 1-19. Odhner, N.HJ., 1917. Results of Dr. E. MjObergs Briggs. J.C., 1974. Marine Zoogeography. McGraw- Swedish scientific expeditions to Australia. 1910- Hill: New York. 475 pp. 1913. Part XVII. Mollusca. Kitnliga Svenska Velenskapsakademiens, llaiullingar Brock, J., 1887. Indische Cephalopodcn. toologisM (4th ser.) 52 Jahrhchen 2: 591-614, pi. 16. figs 1-4. (16): 1-115,3 pis, 51 figs. Proceedings Okutani. T„ Tagawa. M. and llorikawa, H., 1987. Cox, J.C., 1 882. Australian Octopodidae. Wales**: ITi- Cephalopods of continental shelf and slope ofthe I innean Snciet v ofNew South 789. around Japan. Japan Fisheries Resource Corner- 344 MARK D. NORMAN

O'Connor. B.D.S. (eds), Proceedings of the 5th valion Association 1 94 pp, textfigs. Oommen, V.P., 1971. Octopus varunac, a new species Echinoderm Conference, Ga/way, Ireland, 24-29 from the west coasl of India. Bulletin of the September, 1984. Rotterdam: Balkema. Department of Marine Biology and Oceanogra- Sasaki, M., 1917. Notes on the Cephalopoda. Annola- 361-367. phy, University of Cochin 5: 69-76, 6 figs. tiones Zoologicae Japonenses 9 (3): Ortmann, A., 1888. Japanische Cephalopoden. Sasaki, M., 1929. A monograph of the dibranchiate Zoologischc Jahrhiicher Abtheilung fur System- cephalopods of the Japanese and adjacent waters. atik. Geographic unci Biologic tier Thiere 3: 639- Journal of the Faculty of Agriculture, Hokkaido 670, pis 20-25. Imperial University 20 (supplement) 357 pp, Ortmann, A., 1891. Cephalopoden von Ceylon. Zooi- 30 pis. ogische Jahrhiicher, Abtheilung fur Syslcmatik. Smith. E.A.. 1884. Mollusca. Report on the zoological Okologie unci Geographic der Thiers 5: 669-679, collections made in the Indo-Pacifk Ocean during ' pis XLV1. the voyage of the HMS 'Alert" 1881-2 00: Pickford, Ci.E. and McConnaughcy, B.H., 1949. The 34- 116,1 pi.

Octopus himaculatus problem: a study in sibling Springer, V.G., 1 982. Pacific Plate biogeography. with species. Bulletin of the Bingham Occanographic special reference to shorefishes. Smithsonian Collection 12: 1-66. Contributions to Zoology 367: 1-182. Rees, W.J., 1950. The Cephalopods of the Cocos- Terajima, Y., 1713. Wakan Sansui Zue ("Japanese Keeling Islands collected by C.A. Gibson-Hill. Illustrated Encyclopedia"), 51: 17-18, Manu- Bulletin of the Raffles Museum, Singapore 22: script, National Diet Library, Tokyo (fide Glead- 99-100. alland Naggs, 1991). Robson, Ci.C, 1929. A monograph of the Recent Tryon.G.W., 1879. Volume 1: Cephalopoda. Manual ' Cephalopoda. I. Octopodinae. British Museum ofConchology I: 1-316, pis 1-111. (Natural History): London. 236 pp. Van Heukelem, W.F., 1976. Growth, bioencrgetics Roper, C.F.E. and llochberg, F.G., 1987. Cephalo- and life span of Octopus cyanea and Octopus

pods of Lizard 1., Great Barrier Reef. Australia. muya. PhD dissertation, University of Hawaii. Occasional Papers from the Museum of Victoria 3: 224 pp. 15-20. Van Heukelem, W.F., 1983. Octopus cyanea Pp. 267- Roper, C.F.E. and llochberg, E.G., 1988. Behaviour 276 in: Boyle, P.R. (cd.), Cephalopod Life Cycles,

and systematic? of cephalopods from Lizard Vol. I, Species Accounts. London: Academic Island. Australia based on colour and body pat- Press. terns. Malacologia 29 (I): 153-193. Voss, G.L., 1963. Cephalopods of the Philippines. Roper, C.I'.L. and Sweeney, M.J., 1983. Techniques Smithsonian Institution Bulletin 234: 1-180, for fixation, preservation, and curation of cepha- 36 figs. lopods. Memoirs of the Museum ol Victoria 44: Voss, G.L. and Solis Ramirez, M., 1966. Octopus 29-47. maya, a new species from the Bay of Campeche, Roper, C.F.E., Sweeney, M.J. and Nauen, C.E., 1984. Mexico. Bulletin of Marine Science 16 (3): 615- FAO Species Catalogue, Volume 3. Cephalopods 625, 2 figs. of the World. FAO Fisheries Synopses. (125) 3: Voss. G.L. and Williamson, G.R., 1971. Cephalopods 196 pp, textfigs. of Hong Kong. Hong Kong Government Press:

Roper, C.F.E. and Voss, G.L., 1983. Guidelines for Hong Kong. 1 38 pp, 35 pis, 68 figs. taxonomic descriptions of cephalopod species. Wilson, B.R. and Allen. G.R., 1987. Major compon- Memoirs of the Museum of Victoria 44: 49-63. ents and distribution of marine fauna Pp. 43-68. Rowe, F.W.E., 1985. Preliminary analysis of distri- in: Dyne. G.R. and Watson, D.R. (eds). Fauna of bution patterns of Australia's non-endemic, trop- Australia. General articles Vol. IA. Australian ical echinoderms. Pp. 91-98 in: Kccgan, B.F. and Government Publishing Service: Canberra.