<I>Octopus Cyanea</I> Gray, 1849 (Mollusca: Cephalopoda) in Australian Waters: Description, Distribution and Taxonom

Home , Callistoctopus, Cyanea (jellyfish), Octopus cyanea, Point Cloates

BULLETIN OF MARINE SCIENCE, 49(1-2): 2a-38, 1991

OCTOPUS CYANEA GRAY, 1849 (MOLLUSCA: CEPHALOPODA) IN AUSTRALIAN WATERS: DESCRIPTION, DISTRIBUTION AND TAXONOMY

Mark D. Norman

ABSTRACT Octopus cyanea Gray, 1849 is widely distributed throughout tropical waters of the Indian and Pacific Oceans, O. cyanea was originally described from an unspecified locality in Aus- tralian waters as "Coast of New Holland." This paper provides detailed description and diagnostic characters for this species based on Australian specimens, original data on Aus- tralian and wider distributions, and an updated synonymy. The extant syntype "a" (BMNH 1928.2.4.1) is designated as lectotype. O. cyanea occurs in association with coral reefs in northern Australian tropical waters from the southern end of the Great Barrier Reef, north to Torres Strait and west to North West Cape, Western Australia. The misuse of the name Octopus cyanea (and cyaneus) for a species from temperate Australian waters is also resolved.

Octopus cyanea is a large, diurnally active octopus found in association with coral reefs throughout the tropical waters of the Indian Ocean and much of the Pacific Ocean. Gray described this species in 1849 from two preserved specimens in the British Museum of Natural History collection. The type locality was given as "Coast of New Holland" and the distribution of this species as "Australian Seas." The inadequate original description and imprecise type locality provided by Gray may have led to O. cyanea being designated dubious if not for the well preserved syntype (specimen "a": BMNH 1928.2.4.1) housed in the British Mu- seum of Natural History. This specimen is extant, in excellent condition and exhibits the distinctive diagnostic features of this species. The location of the second syntype is unknown. To avoid potential future confusion, the extant syntype (BMNH 1928.2.4.1) is hereby designated as lectotype for this species. Octopus cyanea has been described and treated under a number of species names from throughout its range. Major works include those of Robson (1929: As Octopus cyanea), Sasaki (1929: As Polypus marmoratus), Adam (1954: As O. cyaneus) and Taki (1964: As Callistoctopus magnocellatus). In addition, the name cyanea has been regularly and inappropriately assigned to a distinct temperate water species of Australia's east coast. This paper aims to resolve some of the confusion surrounding the identity, synonymy and distribution of O. cyanea in Australian waters by providing the first detailed description and distributional information for this species from Australian waters.

MATERIAL AND METHODS

In this study, field collection and observations were undertaken on coral islands, reefs and continental islands within the boundaries of the Great Barrier Reef, Queensland. Twenty-nine O. cyanea were encountered in the field and where possible photographed and collected for preservation and detailed examination. Specimens were fixed and preserved according to the techniques of Roper and Sweeney (1983) and are now housed in the Museum of Victoria (NMV). Additional preserved specimens were examined from the collections of Australian Museum, Sydney (AMS), Queensland Museum, Brisbane (QMB) and Western Australian Museum (WAM). Material examined is presented in Table 1. In June 1990, a brief visit to the Smithsonian Institution in Washington, D.C. enabled examination of over 100 specimens of O. cyanea from throughout the Pacific and Indian Oceans. The localities for these specimens are provided in Figure 1b. In the description and tables, measurements and indices follow Roper and Voss (1983: 56), see

20 NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 21

Table 2 for key to symbols. The following additional indices and symbols are also employed: Maximum Arm Width Index (MA WI): arm width at widest point on stoutest arm, as % of ML; Sucker Diameter Index (SDI): diameter oflargest sucker (no distinction between enlarged and normal); Ocellus Diameter Index (OcDI): diameter of ocellus (inner dark spot for O. cyanea) as percentage of ML; Stage of Maturity (StM): Immature (1m: sex indeterminate), Submature (S: reproductive organs distinct but poorly developed) and Mature (M: developed spermatophores or eggs distinct); Hectocotylized Arm Sucker Count (RASC): number of suckers on hectocotylized arm (Toll, 1988). Funnel Organ Limb Index (OrLI): length of outer limb offunnel organ as percentage of median limb length; Pallial Aperture Arc Index (PAAI): length of straightened arc of pallial aperture as percentage of circumference of body at level of aperture.

Octopus cyanea Gray, 1849

Octopus cyanea Gray, 1849, Catalogue of the Mollusca in the collection of the British Museum, Part I: Cephalopoda Antepedia, London: 15. Lectotype: BMNH 1928.2.4.1: 112 mm ML male. Type locality: "Coast of New Holland." Tryon, 1879: 124; Cox, 1882: 788; Hoyle, 1886a: 221; Robson, 1929: 94, textfigs. 21-23; Adam, 1937: 74, fig. 28; Boone, 1938: 354, pI. 149; Rees, 1950: 99; Rees and Stuckey, 1954: 190; Van Heukelem, 1966; 1970: 48; 1973: 299; 1976: 1; 1979: 123; 1983: 267; Maginnis and Wells, 1969: 607; Yarnall, 1969: 747; Wells and Wells, 1970: 65,2 figs.; 1972a: 71; 1972b: 293; Boucher-Rodoni, 1973: 237; Houck, 1982: 152, fig. I; Okutani et aI., 1987: 159 figs. 6Ia-b; Roper and Hochberg, 1987: 18; 1988: 157, figs. 2-6, 31. Polypus cyanea: as Hoyle, 1909: 259 (see discussion of "Common Sydney Octopus.") Octopus cyaneus: as Adam, 1938 (in part): 5, fig. 2; 1939: 79; 1942; 15; 1946: 6, fig. 3; 1954: 171; 1955: 191; 1959: 172; 1960: 17; 1973: 38; Pickford, 1955:' 91; Roper, 1983: 19; Roper, Sweeney and Nauen, 1984: 196, textfigs. Callistoctopus cyanea: as Taki and Igarashi, 1967: 22; Taki, 1981: 253. Octopus tonganus Hoyle, (in part) 1885a, Ann. Mag. Nat. Hist., ser. 5, vol. 15: 225. Syntypes: BMNH; three types of which third specimen (90.12.28.83) is a small female O. cyanea. Type locality: Tongatabu, Tonga Islands. Hoyle, (in part) 1885b: 100; Robson, (in part) 1929: 77; Adam, 1954 (in part): 172. Octopus marmoratus Hoyle, 1885a, Ann. Mag. Nat. Hist., ser. 5, vol. XV: 227. Type: BMNH 1889.4.24.25-26: one male, one female (originally described from I male, 2 females: one female lost 7). Type locality: Honolulu, "Sandwich Islands" (=Hawaii). Hoyle, 1885b: 102; 1886a: 217; 1886b: 85, pI. VI; Ortmann, 1891: 671. Polypus marmoratus: as Brock, 1887: 610; Hoyle, 1905: 978; 1909: 260; Berry, 1909: 418; 1912: 291, pI. XLV, pI. XLVIII, fig. 6, textfig. 13; Wiilker, 1913: 457; Sasaki, 1929: 47, pI. I, fig. 10; pI. V, fig. 4; pI. IX, figs, 19-24; textfig. 17. Octopus vulgaris (non Lamarck, 1798) Goodrich, 1896: 19. Octopus horsti Joubin, 1898, Notes Leyden Mus. vol. xx: 23. Type: Musee royal de Leyde: one 7 sex. Type locality: Djeddah, Red Sea. Hoyle, 1907: 451, pI. xx, figs. 1-3, 12, textfig. 128; 1909: 259. Polypus horsti: as Weindl, 1912: 270; Wiilker, 1920: 51. Polypus herdmani Hoyle, 1904, Ceylon Pearl Oyster Fisheries, Supplementary Reports, No. XIV, "Cephalopoda": 187, pI. 1. Type: location of type specimen unknown: 95 mm ML female. Type locality: Galle, Ceylon (=Sri Lanka). Hoyle, 1907: 454; 1909: 259; Massy, 1916: 206. Octopus herdmani: as Winckworth, 1926: 326; Robson, 1929: 86. Polypus glaber "Riippell MS," (see Wiilker, 1920: 51). Original manuscript not found. Polypusfontanianus (non d'Orbigny, 1835) Robson, 1921: 437, pI. 66, fig. 5a-e. Octopus (Octopus) cyanea var. gracilis Robson, 1929, A monograph of the recent Cephalopoda. Part I: Octopodinae. Richard Clay and Sons, Suffolk: 98. Type: BMNH 1897.9.28.1 (not 1907.9.28.1 as provided by Robson: see Pickford, 1955:' 92). One female: "maximum size 31O±mm". Type locality: Madras. Callistoctopus magnocellatus Taki, 1964, J. Faculty Fish. Anim. Husb., Hiroshima University, vol. 5 (2): 298, pI. 4, figs. 1-2, textfigs. 42-46. Types: Holotype: 147 mm ML male; paratypes: 120, 125,140 mm ML males, location of type specimens unknown. Type locality: Iyeno-kushi, Uchi- umi Village, Minami-Uwa County, Ehime Prefecture, Japan. Octopus bimaculatus (non Verrill, 1883) Dong, 1987: 165, fig. 109.

I Pickford, G. E. A revision orthe Indo-Malayan Octopodinae based on the Collections ofthe British Museum, Bingham Oceanographic Laboratory, Yale University. Unpublished manuscript. 22 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, \99\

Table 1. Material examined: Octopus cyanea

Sex ML(mm) Source Reg. No. Location (date, depth)

15 19.2 AMS CI56385 Osterland Reef, Qld., 15°33'S, 145°32'E (30/VII 1988 intertidal) 15, I <;> 21.6,27.4 WAM 2329.84 Scott Reef, Western Australia, -14000'S, -121°55'E (7/1X!1984, 8-18 m) I 'i' 22.4 WAM 2944-84 Seringapatam Reef, Western Australia, -13°40'S, -122005'E (12/1X!1984) 1<;> 27.8 AMS CI62595 Michaelmas Cay, offCims, Qld., -16°35'S, -146000'E (8/VII/ 1975, intertidal) 1<;> 30.2 AMS C36582 Murray Is., Torres Strait, Qld., -lOolO'S, -142°15'E (1907) 15 32.7 NMV F57907 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, I 52005'E (16/X/1989, 1 m) 15 33.4 WAM 290-88 Point Cloates, Ningaloo Reef, Western Australia, 22°42'S, 113°39'E (29/VIII 1968) I'i' 37.0 AMS CI62596 Escape Reef, Qld., 15°47'S, 145°47'E (6/XII 1981) 15 43.9 AMS CI62597 Murray Is., Torres Strait, Qld., -lOolO'S, -142°I5'E (1907) 15 49.8 NMV F57906 Crystal Cove, Lizard Is., Qld., 14°38'S, 145°24'E (22/XI/1989, 1.2 m) 25 66.8,93.1 AMS C36591 Murray Is., Torres Strait, Qld., -lOolO'S, -142°15'E (1907) 1<;> 71.1 QMB Green Is., off Cairns, Qld., 16°45'S, 145°58'E (no collection date or depth) 1 <;> 73.9 AMS CI62598 Mellish Reef, Coral Sea, 17°25'S, 155°50'E (VIII 1972) 1 \? 76.3 AMS C162599 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, I52"05'E (IX! 1971) 1 'i' 76.8 AMS C162600 Michaelmas Cay, off Cairns, Qld., -16°35'S, - I46000'E (V-VII 1926) 1 'i' 77.3 WAM 1861.67 Aacourt Bay, Barrow Is., Western Australia, 20046'S, 15°21'E (30/VIIIII966, intertidal) 1 \? 77.5 AMS CI62601 Tarauma, Papua New Guinea, -9°30'S, 150"00'E (20/V1/1970) 15 77.7 AMS CI56386 Undine Reef, Qld., 16008'S, 145°40'E (2/VIII 1988, intertidal) I\? 84.6 WAM 395-88 Barrow Is., Western Australia, 20046'S, 115°21 'E (VIII-IX! 1966) 1 <;> 86.7 NMV F57903 Heron Is., Capricorn Bunker Group, Qld., 23°50'S, 152°25'E (1I/X/1989, 0.2 m) 15 88.1 AMS C162604 Heron Is., Capricorn Bunker Group, Qld., 23°50'S, I 52°25'E (no date or depth) I d 100.0 NMV F57909 "Bindoy," Phillipines (XI/l979) 1<;> 101.4 NMV F57905 Tryon Is., Capricorn Bunker Group, Qld., 23°15'S, 151°47'E (27/X/1989, 0.2 m) I d 103.9 NMV F57904 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, I 52005'E (15/X/1989, 3 m) 1 <;> 105.1 NMV F57901 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, I 52"05'E (15/X/1989, 2 m) 15 105.6 NMV F57902 Palfrey Is., Lizard Island Group, Qld., 14°38'S, 145°24'E (17/XIII989, 3 m) I\? 108.2 NMV F57908 Panglao Is., Phillipines, -9°35'N, -123°50'E (XI/I979) 1 d 127.1 NMV F57900 Watsons Bay, Lizard Is., Qld., 14°38'S, 145°24'E (17/XIII989, 15 m) 1 <;> 139.5 AMS C162602 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, 152005'E (27/XI/l966, intertidal) 1 d 150.0 QMB Mo316 Moreton Bay, Qld., -27°23'S, -153°15'E (14/VI 1915) I \? 158.0 AMS C162603 One Tree Is., Capricorn Bunker Group, Qld., 23°30'S, 152005'E (8/VII/l966) NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 23

a 0' •

b. 10'S

•b.

20'S ------~--- ,//' 3O'S ~/ •...--- •..• -

• • V

1000E 120'E 1400E 160'E

Figure I. Octopus cyanea: a) Distribution in Australasian waters: & = material examined in detail; t:. = material examined in visit to Smithsonian Institution, Washington, D.C.;. = published records; ----- = 20"C winter isocryme (mean minimum sea water temperature). b) Global distribution: & = material examined in detail; t:. = material examined in visit to Smithsonian Institution, Washington, D.C.;. = published records; ----- = 20"C winter isocryme (mean minimum sea water temperature).

Diagnosis. -Distinctive ocellus present on arm crown on base of third arms, located one third of distance from eye to web margin. Ocellus consists of dark central ovate spot (approximately one seventh ML), surrounded by pale ring and fine dark outer ring. Ring of iridescent tissue within ocellus absent. Dark bars on ventral arm faces oflarger specimens (> -60 mm ML), forming stripes in alter- nation with base of suckers. Three to seven rows of cream spots on aboral arm surfaces from web margin to tips, containing small erectile skin ridges. Reticu- 24 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

Table 2. Measurements (mm) and indices of female Octopus cyanea from Australian waters

WAM2944-84 WAM 2329-84 AMSCI62598 AMS CI62599 AMSC162600 ML 22.4 27.4 73.9 76.3 76.8 TL 92 126 515 476 443 MWI 58,5 61.7 67.9 79.9 56.3 HWI 64.7 61.7 62.1 62.0 41.9 MAl 32.9 28.8 17.2 19.8 21.5 ALI L R L R L R L R L R 196.4 285.7 281.0 273.7 491.2 504.7 437.7 426.0 341.1 D 263.4 D 306.6 317.5 473.6 487.1 478.4 484.9 D 411.5 D 303.6 346.7 313.9 518.3 581.9 500.7 505.9 440.1 445.3 D 232.1 288.3 324.8 548.0 564.3 465.3 498.0 466.1 414.1 MAW! 17.9 16.1 N.R. N.R. N.R. sm· 9.4 8.0 10.0 10.9 7.4 WDI 22.4 24.2 14.7 14.2 17.0 WF CDB=EA C=DB=EA C=DB=EA B=C=DEA DCBEA GiLC 10/10 9/9 9/10 10/9 9/9 9/9 9/9 9/9 10/10 10/10 OeDI 13.4 15.3 N.R. N.R. N.R. StM 1m 1m S S S EgLl EgWI FuLl 34.4 38.0 37.3 37.2 35.8 FFul 21.4 16.1 26.0 21.9 17.4 OrLl 88.6 83.6 94.0 89.1 98.4 PAAI 38.5 36.4 42.3 40.5 53.6 Explanation of symbols (all tables): ML.: mantle length; TL: total length; MWI: mantle width index; HWI: head width index; MAl: mantle ann index; AU: ann length index; MA WI: maximum arm width index; SDI: sucker diameter index; WDI: web depth index; WF: web formula; GiLC: gilllameUae count (per demibrancb); OeDI: oceUus diameter index measured frnm central dark spot for O. cyanea; SIM: stage of maturity (1m = immature; S = submature; M = mature); EgLI: egg length index; EgWI: egg width index; HeAl: hectocotylized arm index; OAl: opposite arm index; HASC: hectocotylized ann sucker count; LU: ligula length index; CaU: calamus length index; PU: penis length index; PdU: penis diverticulum length index; SpLl: spermatophore length index; SpWI: spermatophore width index; SpRI: sperm reservoir index; FuLl: funnel length index; FFul: free funnel index; OrLl: organ limb index; PAAI: pallial aperture arc index. D = damaged. N.R. ~ not recorded, • = for largest sucker only. In.D. = indistinct. lations over body enclosing roughly circular warts, three large ones pronounced on dorsal arm crown below eyes, especially in younger animals. Description. -Medium to large robust species (Fig. 2a): ML to at least 160 mm for both sexes, TL to at least 900 mm. Mantle broadly ovoid, muscular and thick walled (MWI 53.8-67.4-80.5) with well developed cartilaginous stylets. Pallial aperture moderately wide (PAAI 35.4-49.4-61.9). Funnel broad based and muscular (FuLl 29.5-38.2-48.4), free portion approximately halffunnellength (FFuI 12.3-21.6-26.6). Funnel organ W shaped with broad limbs (Fig. 2c), outer limbs slightly shorter to slightly longer than median ones (OrLl 83.0-93.1-106.8). Head slightly narrower than mantle (HWI 41.8-54.6-66.7), neck not greatly pronounced. Eyes large, prominence varies with state of preservation. Arms relatively long (MAl 17.2-23.5-32.9), robust (MA WI 11.1-15.9-20.8), tapering to fine tips. Arms approximately equal in length (AF equal or IV = III = 11.1),tips often damaged in larger specimens. Suckers biserial (first four from mouth uniserial), moderate size (SDI5 7.4-9.8-12.4; SDI2 7.4-9.3-11.1). Two to four enlarged suckers visible on mature males at approximately 6th or 7th sucker row on arms II and III. Webs well developed (WDI 14.2-19.1-29.0), deepest between lateral arms, narrowest between dorsal arms (WF mainly CB = D = EA or B = C = D = EA). Web remnants extend along ventral edges of arms as narrow membrane to arm tips (Fig. 3a). Third right arm in males hectocotylized, slightly shorter than opposite arm (OAI 70.9-81.5-87.9; HcAI 296.3-351.4-435.7). Ligula very small (LLl 0.4-1.2- NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 25

Table 2. Continued

NMV FS7903 NMV FS790S NMV FS7901 AMSCI62602 AMS CI62603 86.7 101.4 105.1 139.5 158.0 521 656 582 972 890 78.4 61.7 80.5 77.4 68.9 58.9 45.4 56.0 57.7 43.6 21.3 18.8 23.1 22.9 23.1 L R L R L R L R L R 412.9 436.0 420.1 422.1 414.8 412.0 D 369.9 433.5 D 469.4 395.6 D 454.6 428.2 411.0 377.1 377.1 D 323.0 459.1 D 530.6 495.1 432.0 375.8 417.9 457.3 D 371.5 459.6 446.4 509.9 479.3 427.2 425.3 437.3 425.8 D D 20.3 16.1 20.6 N.R. 17.5 D 8.8 9.2 11.1 9.0 17.7 19.0 17.6 22.4 16.4 C=DB=EA DC=EBA CB=DAE DC=EBA C=DB=EA 10/10 10/10 10/10 10/10 11/11 11/11 10/10 10/10 9/9 9/9 21.2 14.4 14.1 N.R. 10.9 S S S M M 1.5 0.2 46.3 39.3 47.5 33.8 29.5 20.9 21.5 24.6 20.2 12.3 90.0 99.1 85.2 84.8 In.D. 52.6 47.4 52.5 54.8 55.6

1.7), conical and bluntly pointed (Fig. 2d) with slightly raised short blunt calamus (CaLI 35.3-38.3-41.4). Ligular groove shallow with approximately 10 fine transverse ribs. Spermatophore groove well developed, wide and thin with fine transverse ridges. Spermatophore guide small but distinct with no obvious papillae. Around 200 suckers on hectocotylized arm (RASC 160-229, N = 10). Gills possess 9-11 lamellae on both inner and outer demibranchs, plus the terminal lamella. Digestive tract (Fig. 4a). Anterior salivary glands extend along approximately one third of buccal mass from posterior margin on dorsal surface. Posterior salivary glands attached to lateral faces of crop. Salivary ducts from each posterior gland joining close to crop diverticulum, continuing as a single duct to buccal mass. Oesophagus elongate, approximately equal in length to crop. Well developed crop diverticulum demarked from posterior crop by distinct bend. Stomach typ- ically bipartite. Caecum coiled in two to three whorls. Digestive gland duct passes from midline of posterior end of digestive gland, branching midway to attach on either side of base of caecum. Intestine undifferentiated with tight coil one third of length from stomach. Ink sac very large, most developed at anterior end of digestive gland extending along bulk of ventral surface. Thick walled ink duct recurving near posterior end of digestive gland, attaches to intestine close to anus. Anal flaps present. Upper beak with short hooked rostrum, narrow hood, curved crest and large lateral walls with marked concave posterior margin (Fig. 4b). Lower beak with short blunt rostrum, narrow hood, narrow and widely spread wings, and narrow and widely spread lateral walls (Fig. 4c). Radula with seven transverse rows of teeth (Fig. 4d). Rhachidian tooth has symmetrical seriation of A4-s type with 1- 2 small lateral cusps on either side. First lateral teeth small and unicuspidate; 26 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

a b

;: ::" '. " ~" " t '., ~, 10mm

d e

>----< 20mm !lmm Figure 2. Octopus cyanea: a) Dorsal view ofNMV F57904 (108.9 mm ML male), illustrating position off our primary papillae; b) Ocellus of AMS CI62603 (158.0 mm ML female); c) Funnel organ of NMV F57900 (127.1 mm ML male) and NMV F57906 (49.8 mm ML male); d) Hectocotylus of NMV F57902 (105.6 mm ML); e) Female reproductive organs of AMS CI62603 (158.0 mm ML); DO-distal oviduct; OB-oviducal gland; PO-proximal oviduct; OV -ovary. NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 27

Figure 3. Octopus cyanea: a-d) Live animal color patterns; a) Warning posture with webs flared and ocelli pronounced (NMV F57903; 86.7 mm ML female); b) "Spiked" posture; c) Chocolate brown coloration showing start of medial stripe; d) Medial stripe coloration; e-f) Skin sculpturing of WAM 290-88 (33.4 mm ML male); e) Dorsal arm crown exhibiting circular warts; f) Lateral view exhibiting ocellus and circular warts; g) Dark barring on ventral arm faces (NMV F57909: 100.0 mm ML male; NMV F57904: 103.9 mm ML male); h) Aboral arm spots (NMV F57904: 103.9 mm ML male). 28 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

a c

10mm

CJ ,"cj 6J\.~ CJ 0"~ 6 6~o

20mm Figure 4. Octopus cyanea: a) Digestive tract of NMV F57902 (105.6 mm ML male); BM-buccal mass; ASG-anterior salivary glands; O-oesophagus; CD-crop diverticulum; PSG-posterior salivary gland; PC-posterior crop; S-stomach; C-caecum; I-intestine; DG-digestive gland; IS-ink sac, b) Upper beak ofNMV F57904 (103.9 mm ML male); c) Lower beak of NMV F57904 (103.9 mm ML male); d) Radula ofNMV F57905 (lOlA mm ML female).

second lateral teeth long with curved base; third lateral teeth long, slightly curved; marginal plates oblong, plain. Male genitalia (Fig. Sa). Penis in mature males long and narrow (PLI 3,9-13.8- 27.3). Genital aperture subterminal inside curve of tip of penis. Diverticulum with single loop approximately one third penis length (PdLI 3.7-5.2-6.9). L shaped Needham's sac tapers to fine point at free end. Anterior end of Needham's sac attaches to penis at level of diverticulum. Tapering portion branches off Need- ham's sac from junction of accessory gland, extending anteriorly to lie against base of penis. Accessory gland long and robust with blind coil at tip. Spermato- phoral gland long and thin with large recurved coil approximately 3f4 along length. Distal portion enlarged at attachment of vas deferens. Vas deferens fine, very long and tightly coiled (over 470 mm in one 127.1 mm ML specimen), attaching to testis in raised centre of anterior face. Spermatophores (Fig. 5b-e) very numerous (337 in a specimen of 105.6 mm ML), relatively short (SpLI 31.9-39.8-49.8) and very fine (SpWI 1.3-1.5-1.6). Spermatophore surface unarmed. Oral cap simple, slightly expanded, bearing a long cap thread. Ejaculatory organ consists of spiralled tube extending along two thirds of spermatophore length, narrowing in last third before cement body. Small clear cement body attached to tapered ends of both ejaculatory apparatus and sperm reservoir. Sperm reservoir approximately one third of spermatophore length (SpRI 29.6-33.3-38.4), spirally wound with round- ed aboral end. Slightly enlarged aboral end forms widest section of spermatophore. Female genitalia (Fig. 2e). Ovary roundly triangular, proximal oviducts short and straight prior to entering membrane enclosing ovary, merging below mem- NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 29

a b c d p AG

lmm CB Figure 5. Octopus cyanea: a) Male reproductive organs ofNMV F57900 (127.1 mm ML); P-penis; AG-accessory gland; NS-Needham's sac; SG-spermatophoral gland; VD-vas deferens; T -testes; b-e) Spermatophore of NMV F57900 (127.1 mm ML); b) Whole spermatophore; c) Aboral end of spermatophore; d) Oral cap and cap thread; e) Spermatophore midsection; CB-cement body. brane to form common oviduct. Elongate distal oviducts attached along length to surface of visceral mass, extending to either side of anus. Only one female (158.1 mm ML) examined possessed mature eggs in ovary: eggs small (EgLI 1.7, EgWI 0.3) and very numerous (> 100,000). Counts and indices for the material examined are presented in Tables 2-4. Skin sculpture consists of fine reticulations forming patch and groove system over body, arm crown and along bulk of arm length, sometimes obscured by dark base color in preservation. Four large primary papillae in diamond arrangement on dorsal mantle (Fig. 2a). Patches enclosed in reticulations roughly circular, containing papillae, unbranched when erected. "Spiked" appearance produced through erection of four primary and smaller secondary papillae over bulk of body and arm crown (Fig. 3b). Three distinctive larger circular patches, each containing central papilla, present on dorsal arm crown below eyes. One medial between and slightly below eyes, pair midway between eyes and dorsal web, often more pronounced in young animals (Fig. 3e, t). Single large papilla above and slightly behind each eye, erected as distinct "horns" in some color patterns. Three to four additional smaller papillae also visible above eye in some specimens. Oral surface of dorsal and dorsolateral webs pigmented with fine reticulations, re- maining webs smooth on oral surface. Skin on ventral surface of head and between ventral arms smooth, sometimes bordered by a raised skin ridge. In life, Octopus cyanea can produce a wide range of color patterns ranging from completely dark chocolate brown, through mottled and sculptured cryptic patterns, to even grey white with marked black ocelli. Several examples of body patterns are provided in Figure 3a-d (also see Roper and Hochberg, 1988: figs. 30 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

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<"IOVV ~;j~~~ - <"I<"I<"IV <:s: \1.l V <"I"" ~ t-: t-: II "-;0"-;";0 .0Vo "'r-\OVN.,., Z--II U "'0 t-: ~O ..lOOOO 00 o-"- ~~o~ -

'2 ;:l .5 E uo 32 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

Table 4. Measurements (mm) and indices of type specimen (BMNH 1928.2.4.1) and combined ranges, means and standard deviations for Australian material

Type (d) BMNH 1928.2.4.1 Australian specimens (1] ct, 102) Range, mean, SD

ML 106 TL 596 MWI 74.3 53.8-67.4-80.5 (8.0) HWI 54.0 41.8-54.6-66.7 (8.0) MAl 22.2 17.2-23.5-32.9 (4.5) ALI L R I 371.0 360.2 196.4-370.4-504.7 (75.0) II 395.8 450.6 263.4-392.1-549.2 (71.9) III 390.7 290.4-435.6-581.9 (75.5) H 350.0H 296.3-351.4-435.7 (41.7) IV 406.4 407.9 232.1-414.6-564.3 (79.1) MAwr N.R. r 1.1-15.9-20.8 (2.7) SDI· 14.7 7.4-9.6-12.4 (I.4) WDI 18.5 14.2-19.1-29.0 (4.1) GiLe !O/N.R. !!/N.R. 9-11 OcDl 15.9 10.9-15.0-21.2 (2.9) HeAl :149.8 296.3-351.4-435.7 (41.7) OAI 89.5 70.9-81.5-87.9 (5.8) HASC 197 160-229 LLI 1.1 0.4-1.2-1.7 (0.38) CaLI 47.5 35.3-38.3-41.4 (2,2) PLI 15.8 3.9-13.8-27.3 (7.0) PdLl 4.8 3.7-5.2-6.9 (1.2) SpLl 32.6 31.9-39.8-49.8 (7.8) SpWI 1.1 1.3-1.5-1.6 (0.1) SpRI 41.5 29.6-33.3-38.4 (3.3) EgLl 1.7 from mature female: 158.1 mm ML EgWI 0.3 from mature female: 158.1 mm ML FuLl 45,1 29.5-38.2-48.4 (5.3) FFul 29.0 12.3-21.6-26.6 (3.6) OrLl N.R. 83.0-93.1-106.8 (6.9) PPAI N.R. 35.4-49.4-61.9 (8.0)

2-6,31). Active animals most often display cryptic coloration and texturing, while animals in lairs are often cream to white with two red longitudinal bands running from body through eyes and down dorsal arm crown. Three color pattern features are visible in all live animals: 1). Ocelli: One large ovate ocellus on either side of arm crown over base of third arm, approximately one third distance from eye to web margin (visible in Fig. 3a, f). Ocellus with central dark spot approximately 'l-,thmantle length (OcDI 11.4-15.4-21.2) surrounded by pale ring and fine outer dark ring (Fig. 2b). Iridescent ring or tissue not present. Pale ring often raised to form a circular ridge of skin. In certain live color patterns, the ocellus is only visible as a circular ring of raised skin. 2). Zebra markings on ventral arm faces: Larger Octopus cyanea (ML > -60 mm) exhibit a dark barring pattern over a cream base color on the ventral faces of all arms. This pattern varies from distinct "zebra stripes," one stripe between each pair of suckers, to a mottled dark band containing rows oflight spots (Fig. 3g). 3). Aboral arm spots: Three to seven rows (varying along length of arm) of cream colored elliptical spots run along aboral surfaces of all arms from web margins to arm tips, often pronounced over dark base color (Fig. 3a, h). Spots contain erectile skin ridges. In some live animal color patterns and preserved specimens aboral arm spots are only visible as series of raised skin ridges. NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 33

Preserved specimens exhibit a wide range of color patterns from drab slate grey or cream with ocelli and other markings visible, through to dark purple/black. The latter form often appears to obscure diagnostic markings, however these characters can still be discerned on close examination. No live or preserved color patterns display the color blue, as early interpretations of the species name sup- posed. Integument sculpturing ranges from absent to pronounced dependent on fixation and preservation. Frozen specimens lose all sculpturing. Sexual dimorphism is not marked in Octopus cyanea, with both sexes attaining approximately equal maximum size. Males possess shorter third right arms and, in mature individuals, two to four enlarged suckers are present on arms II and III.

Distribution. - In Australia, Octopus cyanea is distributed in northern tropical waters (Fig. la), primarily in association with shallow coral reefs. It occurs along the entire length of the Great Barrier Reef, north into the waters of Papua New Guinea and west through Torres Strait to north west Western Australia. In eastern Australia, the southern limit coincides with the southern end of the Great Barrier Reef (approximately 25"8, l53°E). One specimen (QMB M0316) was collected from the warm temperate waters of Moreton Bay, southern Queensland (27°23'S, 153°15'E); however, this is probably a vagrant from tropical waters. In Western Australia, O. cyanea is confirmed as far south as Point Cloates on North West Cape (22°42'S, 113°39'E). This species has also been collected from outer shelf reefs north of Western Australia, namely Seringapatam Reef (13°40'S, 122°05'E) and Scott Reef (14°00'S, 12l055'E). O. cyanea has been collected throughout the tropical waters of the Indian Ocean and much of the Pacific Ocean. Figure 1b illustrates the distribution of this species based on material examined and published records.

Live Animal Observations. - Twenty-nine live Octopus cyanea were encountered on offshore islands and reefs along the length ofthe Great Barrier Reef, from One Tree Island in the south (23°30'S, 152°05'E) to Lizard Island in the north (14°38'S, 145°24'E). O. cyanea seems to occur in low numbers on the Great Barrier Reef, however the cryptic nature and camouflage capabilities of this species make de- tection difficult and densities impossible to estimate. Animals were encountered from intertidal reef flats to 15 m deep, occupying lairs in coral bedrock, live and dead coral heads and excavations in sand and rubble. Established lairs were often discernible by the scattering of empty crab carapaces surrounding the opening. Two multiple lairs were encountered with more than one animal present. One pair of lairs was excavated in coral rubble, only 20 cm apart and contained the two largest specimens encountered in this study (ML > 160 mm). One specimen was male, the sex of the other was not determined. It is possible these double lairs contain breeding pairs. One large male was removed from a separate triple lair. The following day two additional specimens had occupied the lair system. Prey remains were collected from around established lairs where occupants were present. These consisted solely of crab remains. Carapaces to 120 mm wide were found around lairs of large octopuses. Captive animals readily fed on dead prawns and fish. There was no evidence of inclusion of bivalves or gastropods in the diet. No mollusc shells were observed around lairs and captive animals would not feed on live or dead molluscs. Van Heukelem (1976) examined stomach contents of 49 O. cyanea from Hawaiian waters and found 89% crab remains, 41% stomatopods, 27% alpheid shrimps, 10% fish bones and in one individual, remains of a small moray eel. 34 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

Individuals of O. cyanea were mostly encountered in the mouths oflairs, alert and half emerged throughout daylight hours. Animals were similarly half emerged from lairs when encountered on night dives but may have been influenced by approach of divers and lights. Only six octopuses were observed active away from lairs. Five large specimens were observed on three occasions out of their lairs moving over live coral, active through the middle of the day (1035 hr, 1040 hr, three observed together 1230 hr). The latter three were encountered together in shallow water on open ground and may have been involved in courtship behavior. One small O. cyanea (NMV F57907: 32.7 mm ML) was encountered actively foraging over coral rubble at 0710 hr. Further study of Australian animals would be required to determine whether they exhibit the crepuscular peaks in activity as observed by Houck (1982) for O. cyanea in Hawaiian waters. The life cycle of Octopus cyanea is well documented, mainly through the studies of Van Heukelem in Hawaii (1966, 1970, 1973, 1976, 1979). Other life history studies on this species include Yarnall (1969), Wells and Wells (1970, 1972b) and Houck (1982).

DISCUSSION Description, measurements and counts for the type specimen of Octopus cyanea (BMNH 1928.2.4.1) were provided by T. Stranks (NMV), and photographs of the type were provided by C. C. Lu (NMV). The type specimen exhibits the characteristic ocellus and markings on the ventral arm faces, and matches counts and indices of Australian material examined (Table 4). There is no doubt that the Australian specimens examined in this study are all O. cyanea. It was not possible in this study to examine type material first hand. Much of the synonymy is based on the conclusions ofthree authors, Robson (1929), Pick- ford (unpubl.) and Adam (1954, etc.). The first two authors carried out detailed studies of octopods primarily from the collections in the British Museum of Natural History and examined the type material for listed synonyms. The conclusions of these authors are further supported by references in many of the original descriptions of the synonyms to characters diagnostic for O. cyanea. Examination of the USNM collection and confirmation of the wide distribution of this species also supports the validity of the synonymy presented. In 1989 Dong Zhengzhi visited the Museum of Victoria and identified a number of specimens of O. cyanea from northwest Western Australia as O. bimaculatus, Verrill, 1883. Dong's iden- tifications, the restriction of Octopus bimaculatus to the west coast of North and Central Americas, and the occurrence of O. cyanea in regions neighboring the South and East China Seas, all indicate that O. bimaculatus reported by Dong (1987) is in fact O. cyanea. The correct form of the specific name for this species is "cyanea" named after Cyane a nymph from Greek mythology (Roper and Hochberg, 1988). Incorrect interpretation of the specific name as originating from "cyane" meaning blue (male gender) led Adam (1938) to change the specific name to "cyaneus" and employ this name in subsequent papers (1939; 1942; 1946; 1954; 1955; 1959; 1960; 1973), as did other authors (Pickford, 1955;1 Roper, 1983; Roper et al., 1984). O. cyanea is capable of a wide range of body colorations and patterns, none of which are blue. In Australian waters, the name O. cyanea has been regularly and inappropriately assigned to another species, the "Common Sydney octopus." Despite many references in the literature to O. cyanea in Australian waters, the only genuine locality records prior to this study are those of Roper and Hochberg (1987; 1988) from Lizard Island at the n0l1hern end of the Great Barrier Reef. The remainder refer NORMAN: OCTOPUS CYANEA FROM AUSTRALIAN WATERS 35 to the Common Sydney octopus. This distinct temperate species occurs along the New South Wales coast of Australia, and extends no further north than Moreton Bay in southern Queensland (-27°30'S, 153°15'E). It differs from O. cyanea in that it lacks ocelli, has much finer reticulations over the skin and lacks the zebra stripes on the ventral surfaces of the arms. The following papers refer to the Common Sydney octopus and not Octopus cyanea: Octopus cyanea Brazier, 1892: 7. In addition to accurate records, Hoyle (1909: 259) provides New South Wales and Moreton Bay, Queensland records for Polypus cyanea. These are taken from Brazier and refer to the "Common Sydney octopus." Octopus cyaneus Le Souef and Allan, 1933: 373; 1937: 64; Cotton, 1942: 84; Dew, 1959: 44, figs. 1,2; Allan, 1959: 457, textfig. 110, fig. 7; Dakin, 1960: 309, 352, pI. 78; Ireda1e and McMichael, 1962: 99; Loch, 1980: 3; Dakin, 1987: 333, 338, pi.; Loch, 1987: 8; 1988: 9. The identity of the Common Sydney octopus is currently being investigated by the author and T. Stranks of the Museum of Victoria. There is no commercial fishery exploiting Octopus cyanea in Australian waters. The cryptic habits of this species on coral reefs exclude it from trawl fisheries, and no small scale local exploitation has been noted. Its distribution on offshore coral islands and reefs makes it unlikely to be taken in any numbers by mainland Aboriginal communities. Octopus cyanea exhibits a classic "Indo West Pacific" distribution throughout the tropical waters of the Indian Ocean and much of the Pacific Ocean. This distribution is exhibited by many tropical marine organisms and is well documented in the literature (Ekman, 1967; Briggs, 1974). This wide distribution is presumably the product of the juvenile planktonic phase of this species, of which little is known. Van Heukelem (1976) found that captive hatchlings remained in the water column for at least 21 days. The duration of this stage would need to be considerably longer to account for dispersal and maintenance of gene flow over such a great range, especially to isolated island groups such as the Hawaiian Islands.

ACKNOWLEDGMENTS

The findings presented in this paper form part of a broader Ph.D. research program into the octopods of the Great Barrier Reef and northern Australia, being carried out in association with the Museum of Victoria and the University of Melbourne. Sincere thanks to Dr. C. C. Lu and T. Stranks of the Museum of Victoria for invaluable assistance on this paper and the wider research program; Dr. G. F. Watson for support and assistance with the manuscript; M. Turner, C. Davies, J. Martin, J. Swartz, G. Knight and the Breidahl family for field assistance; J. Whyte for photography; D. Paul, R. Plant and C. Rowley for preparation of figures. Warm thanks to C. Roper and M. Sweeney (USNM) and E. Hochberg (SBMNH). Many thanks to S. Troy for support and encouragement. This research was made possible through support grants from Museum of Victoria, Victoria Institute of Marine Sciences, Linnean Society of New South Wales, Keith Sutherland Trust, Australian Museum, and M. A. Bartlett Travel Grant, University of Melbourne.

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DATEACCEPTED: December 3, 1990.

ADDRESSES:Department of Invertebrate Zoology, Museum of Victoria, 285 Russell St., Melbourne, Vic. 3000 and Department of Zoology, University of Melbourne, Parkville, Vic. 3052, Australia.