Species Diversity 21: 31–42 25 May 2016 DOI: 10.12782/sd.21.1.031

Octopus sinensis d’Orbigny, 1841 (Cephalopoda: ): Valid Species Name for the Commercially Valuable East Asian Common

Ian G. Gleadall International Fisheries Science Unit, Tohoku University Graduate School of Agricultural Science, 1-1 Amamiya, Sendai, Miyagi 981-8555, Japan E-mail: [email protected] (Received 21 July 2015; accepted 12 May 2016)

http://zoobank.org/34118987-F3F3-4FEA-BD74-03218965CF84

The East Asian common octopus has long been synonymized with the Atlantic and Mediterranean species Octopus vulgaris Cuvier, 1797. However, evidence from molecular genetic studies has firmly established that the so-called cosmo- politan common octopus is in fact a group of several biogeographically distinct populations which form a complex of spe- cies with closely similar morphology. Here, a diagnosis and brief description are provided which distinguish the East Asian common octopus from O. vulgaris, and as a suitable name for it a former junior synonym of O. vulgaris is identified as a valid species: Octopus sinensis d’Orbigny, 1841. A neotype is designated. Voucher material includes specimens collected in Japan by Philipp Franz von Siebold and deposited in the National Museum of Natural History - Naturalis - in Leiden; and others that were studied by Madoka Sasaki in preparation for the detailed description of this species (as O. vulgaris) in a monograph on Japanese Cephalopoda published in 1929. At present, all species in this complex (particularly O. vulgaris and the East Asian species here identified as O. sinensis) are highly vulnerable to overfishing, so recognizing O. sinensis as a species distinct from O. vulgaris is an important step towards improving sustainable fisheries management policies for each species in this group of commercially valuable . Key Words: hectocotylus, Octopus sinensis, Octopus vulgaris, arm length, suckers.

Octopus insularis Leite and Haimovici in Leite, Haimovici, Introduction Molina and Warnke, 2008 (q.v.). There is no extant type material for O. vulgaris (first de- Octopus is the most familiar and widely used genus name scribed from Europe) and the species has never been de- for octopus species but its has had a confusing scribed in sufficient detail to allow recognition of any small history and the names of both the genus and its type spe- differences from populations in other parts of the world. cies, O. vulgaris Cuvier, 1797, were stabilized only relatively Indeed, its widely known presence appears to have resulted recently (ICZN 1954, 1997). Octopus vulgaris is best known in a lack of interest in preserving specimens of this species as an Atlantic and Mediterranean species but this name has of sufficient maturity and quality for systematic study (pers. long been associated also with the East Asian common octo- obs.). Until this century, therefore, despite being a common pus from Japanese waters, since it was thought to be just one species there have been few preserved specimens available species with a cosmopolitan distribution. This was also the for morphological comparisons of O. vulgaris and the East conclusion of Sasaki (1929) in his monograph on the Ceph- Asian common octopus. However, Guerra et al. (2010) have alopoda of Japanese waters, which has been supported even recently reported morphological measurements for repre- by studies early this century using molecular genetic analy- sentative specimens of O. vulgaris to complement their ge- sis (e.g., Warnke et al. 2004; Kaneko et al. 2011). netic analysis. Recent studies of aquaculture techniques for However, more recent molecular genetic analyses (e.g., the East Asian common octopus in Japan have provided the Guerra et al. 2010; Amor et al. 2014) have revealed conclu- opportunity to acquire a range of new specimens sufficient sively that this apparently cosmopolitan species is in fact a for comparison with the Mediterranean-Atlantic form. This species complex, with as many as five distinct species of “O. comparison here reveals clear morphological and meristic vulgaris” in the Atlantic alone. Within this species complex, differences between the East Asian common octopus and in the western North Pacific the East Asian common oc- O. vulgaris which, together with the evidence available from topus forms a distinct genetic clade that differs from other genetic analysis, confirm that these two forms are different members of this complex at the species level (Guerra et al. species. Here, the species name Octopus sinensis d’Orbigny, 2010; Amor et al. 2014). Most species in this complex have 1841 (previously judged to be a synonym of O. vulgaris; see, yet to be distinguished using morphological and meristic for example, Sasaki 1929), is reinstated as the appropriate characters, although recently one of the Atlantic popula- valid name for the East Asian common octopus. tions off Brazil has been well described as the new species

© 2016 The Japanese Society of Systematic Zoology 32 Ian G. Gleadall

fication: subsequently identified by Sasaki 1929: 37). Materials and Methods Octopus vulgaris (non Cuvier, 1797): Appellöf 1886: 7; Ort- mann 1888: 642; Robson 1929: 57 (in part); Nesis 1987: New specimens of Octopus sinensis were obtained as part 312 (in part; northwestern Pacific population); Okutani of a research project to develop aquaculture methods for et al. 1987: 157, fig. 60 A–C; Kaneko et al. 2011: 106; Ku- the East Asian common octopus. They were fixed in 10% bodera 2013: 212. formalin in seawater. Small pieces of arm muscle were first Polypus vulgaris: Wülker 1910: 5; Berry 1912: 386; Sasaki removed and preserved in absolute ethanol for molecular 1929: 35 (in part), text figs 11–12, pl. IV fig. 1, pl. IX figs analysis. After about a month, the specimens in formalin 1–3, pl. XXIX fig. 1. (Polypus is a suppressed name; ICZN were thoroughly rinsed in running tap water and preserved 1954). in 45% isopropanol. Specimens collected previously and de- Octopus cf. vulgaris (cf. O. sinensis): Gleadall and Salcedo- posited in other institutions were inspected and those iden- Vargas 2004: 119 (in part). tified as O. sinensis are included in the redescription. Meth- ods for measurements and for counting of meristic charac- Material examined. Neotype, here designated, mature ters follow Gleadall et al. (2010). All dimensions are in mm. male (ML 97 mm), Ariake Sea, Hato-no-kama Harbour, For comparison, a search was made for specimens of O. Oyano Is., Kami-amakusa City, Kumamoto Pref., Kyushu, vulgaris in good condition from recorded localities in muse- Japan, IGG 312, NSMT Mo 85659, 25.x.2013, M. Abe, I.G. um collections (listed at the end of the text), and comparisons Gleadall, K. Matsubara and S. Morii. were made also with data reported by Guerra et al. (2010). Other material examined: 1 mature male (ML 95 mm), The specimens used by Guerraet al. (2010) were, as in the loc. not stated (?Kyushu, Japan), NRML 2438, undated present study, measured following preservation. However, (?1824–1827), P.F. von Siebold; 1 immature female (ML they are now no longer extant, having been donated to the 89 mm), loc. not stated (?Kyushu, Japan), NRML 472, un- Luarca Museum on the northwestern coast of Spain, but then dated (?1824–1827), P.F. von Siebold; 1 submature male (ML lost when that museum was tragically devastated by a power- 107 mm), Kagoshima, Kyushu, Japan, ZUMT 994.CO-38 ful storm on 5 February, 2014 (A. Guerra, pers. comm.). [056], August 1903, donated by Kagoshima Prefectural Gov- Abbreviations: Arms no. 1–4, arms counted in corre- ernment; 1 male ML 106 mm (head dissected, arms miss- sponding left (L) and right (R) pairs from the dorsal to the ing), 1 submature female ML 119 mm (no collection data), ventral midline of the head (arms identified individually as ZUMT 994.CO-20 [058]; 1 submature female (ML 84 mm), L1, R3, etc.); #, number prefix indicating the position of the 1 juv. (ML 35 mm), Japan Sea, off Konoura, Yuri-gun, south- largest suckers on an arm, counting distally from the mouth; ern Akita Pref., Honshu, Japan, NSMT Mo 75758 (SHC), 17 BMNH, (British) Natural History Museum, London; CO1, October 1931; 1 male (ML 96 mm), Onahama, Iwaki-gun, Cytochrome C oxidase subunit 1; CO3, CO subunit 3; ICM, Fukushima Pref., Honshu, Japan, NSMT Mo 75750 (SHC), Instituto de Ciencias del Mar, Barcelona; ICZN, Internation- January 1932; 2 males (ML 82, 68 mm), Pacific Ocean, Send- al Commission on Zoological Nomenclature; IGG, lot pre- ai Bay, off Yuriage, Natori-gun, Miyagi Pref., Honshu, Japan, fix of specimens fixed and preserved by the author (as indi- NSMT Mo 75759 (SHC), January 1933; 1 immature male cated below, lots IGG 290, 307 and 309 are included among (ML 42 mm), Pacific Ocean, off Hayama, Sagami (Kanagawa specimens donated to Michael Amor, at La Trobe Univer- Pref.), Honshu, Japan, SMI 051, 8 June 1957; 1 male (ML sity, Melbourne, Australia, towards his study on the Octopus 56 mm), Japan Sea, Fukaura, Nishitsugaru-gun, Aomori vulgaris species complex); juv., juvenile; loc., locality; ML, Pref., Honshu, Japan, NSMT Mo 75739 (SHC), August 1963; dorsal mantle length; NRML, National Museum of Natural 1 male (ML 72 mm), Japan Sea, Kamo Aquarium (original- History - Naturalis, Leiden; NSMT, National Museum of ly caught by octopus pot), Yamagata Pref., Honshu, Japan, Nature and Science, Tsukuba; pers. comm., personal com- BMNH 1994108, Sept. 1987, I.G. Gleadall; 1 mature female munication; pers. obs., personal observation; Pref., Prefec- (ML 122 mm), Japan Sea, Toyama Bay, Uozu, (octopus pot), ture; SHC, Saitō Hō-onkai Collection (the collection of the Toyama Pref., Honshu, Japan, IGG 036, NSMT Mo 85656, Zenuemon Saito Gratitude Foundation, formerly in Sendai, August 1992, O. Inamura; 1 mature male (ML 93 mm), all specimens of which were donated to NSMT; now part Shiriyazaki, Pacific coast of Aomori, IGG 039, NSMT Mo of the NSMT collections); SMI, Shōwa Memorial Institute, 85657, 22 March 1993, I.G. Gleadall and K. Noro (né Sato); Tsukuba; ZUMT, Zoology Department of the Museum of 1 maturing female (ML 124 mm), East China Sea, 31°40.95′ the University of Tokyo. N, 127°47.09′ E, NSMT Mo 74775, 19 June 2000, H. Horika- wa, Kumamoto Maru; 1 mature male (ML 73 mm), Iyo-na- Octopus sinensis d’Orbigny, 1841 da, Seto Inland Sea, Mori, Japan, in a cage, IGG 169, NSMT [English vernacular name: Mo 85658, 21 June 2003, H. Sakaguchi; 1 submature male The East Asian common octopus] (ML 86 mm, arm L2 with bifurcated tip), Japan Sea, Kamo, [Japanese name: Madako] Yamagata Pref., Japan, from beach with rod and line, IGG (Figs 1–6, 8–11; Tables 1–2) 290 (donated to M. Amor), 22 October 2011, Y. Hino and T. Itō; 1 mature male (ML 82 mm), collected with neotype, Octopus sinensis d’Orbigny, 1841 (text): 68; d’Orbigny 1834 IGG 302, NSMT Mo 85659; 2 mature males (ML 82–111) (atlas of plates): pl. 9 (lower figure, without species identi- mm, collected with neotype, IGG lots 307 and 309 (donat- Octopus sinensis 33 ed to M. Amor); 7 mature males (ML 111–168 mm, 902– #12 and #15 (commonly #13 and/or #14) on arms 2 and 3; 2432 g; IGG 318, 324–328, 330), 5 submature females (ML length of hectocotylized arm around 80% of length of third 130–162 mm, 1542-2221 g, IGG 319–321, 323, 329), and left arm, with 119–152 suckers, ending with a minute ligula 1 immature female (ML 112 mm, 956 g, IGG 322), Pacific (length about 3 mm). Spermatophores small, approximately Ocean, Oura Port, near Onagawa, Oshika District, Miyagi 60 (each 40 mm in length) in a male of ML 92 mm. Females Prefecture, Honshu, Japan, landed from baited cage traps at lay many small eggs together as a number of branched 50–100 m [mixed catch with Enteroctopus dofleini (Wülker, strings (festoons). Eggs hatch into planktonic paralarvae 1910) of similar size], NSMT Mo 85660, 30 January 2014, F. with a ML of 1.5–2.0 mm and arms each bearing 3 suckers. Abe, I.G. Gleadall, K. Matsubara, and S. Morii. Well-developed ink sac and anal cirri present. Molecular sequences. NSMT Mo 74775, GenBank Ac- Description. Based on 23 males (14 mature, 2 sub- cession nos. AB430548 (CO1) and AB573218 (CO3), regis- mature, 1 immature, 6 maturity not assessed), 11 females tered (as Octopus vulgaris) by Kaneko et al. (2011). (2 mature, 7 submature, 2 immature), and 1 juvenile. Me- Diagnosis. Member of the Octopus vulgaris species dium- to large-sized octopus: maximum size around 3 kg, complex inhabiting the western North Pacific in the vi- males maturing around ML 70 mm (approx. 400 g), females cinity of Japan. Maximum size around 3 kg, males matur- 110 mm (approx. 900 g). Body robust, ovoid (MW approx. ing around ML 70 mm (approx. 400 g), females 110 mm two-thirds of ML); head broad, width just less than half of (approx. 900 g). Funnel organ broad, W-shaped. Ocelli ab- ML; neck broad, almost as wide as head (Figs 1–3). Eyes sent. Mature males with 1 or 2 enlarged suckers between large though not especially prominent, each with a single

Table 1. Basic measurements and counts of neotype and voucher specimens of Octopus sinensis.

Males Females Depository & NSMT NSMT NSMT NSMT NSMT NSMT NSMT NRML BMNH NRML Registration Mo 85659 Mo 85660 Mo 85657 Mo 85659 Mo 85660 Mo 85660 Mo 85656 2438 1994108 472 code Neotype (IGG 318) (IGG 039) (IGG 302) (IGG 320) (IGG 323) (IGG 036) ML 97 168 95 92 82 72 151* 127 122 89 VML 65 116 66 74 58 52 127 97 98 62 MW 57 93 66 54 56 45 82 85 86 49 HW 47 72 45 45 38 35 69 64 49 32 NW 47 70 39 40 37 25 58 66 45 24 GC 8–9 9 7–8 9 8–9 8–9 9–10 8–9 8 8 GL 33 39 26 29 27 19 49 40 44 32 FL 26 56 33 28.5 37 27 54 44 48 33 FFL 25 48 28 17 20 14 42 26 34 23 LL 4.5 3.8 3.0 4.0 3.3 5.3 — — — — CL 2.8 1.6 1.1 1.5 1.5 1.6 — — — — AL 1 279/242 491/520 270/292 217/T 208/246 193/179 462/474 Rg/504 380/440 306/297 2 355/350 633/598 Rg/285 281/275 Rg/285 256/254 Rg/580 588/T 520/518 368/320 3 365/292 624/482 375/277 277/238 T/234 244/192 547/552 583/625 515/510 T/360 4 289/327 566/622 325/315 248/243 T/T 199/202 488/500 553/507 490/405 270/303 AF 3.2.4.1 2.3.4.1 3.4.2.1 2.3.4.1 — 2.3.4.1 2.3.4.1 2.3.4.1 2.3.4.1 2.3.1.4 SD 17.5 25 18 15 12 13.3 19 19 14.5 11.4 SD (arms) L2, L3, L3 2, 3 L3, R2 R2, L2, L3 2, 3 L3, R3 L3 L2, R2 L2, R2 2, 3 SD (#) 13, 14, 14 13 14, 13 14, 14, 14 14, 14 12–14 13 15, 15 14–22 12–15 WD A 37 78 48 44 42 40 79 67 50 42 B 52/52 108/108 50/60 51/51 50/54 58/— 109/103 107/85 70/61 49/58 C 84/74 135/152 75/75 65/64 64/70 69/— 126/134 127/117 90/75 57/70 D 86/68 155/126 80/— 65/60 64/58 64/— 124/125 124/106 90/85 60/63 E 66 120 53 54 49 46 103 99 45 46 WDF C=D.B.E.A D.C.E.B.A D.C.B.E.A C=D.E.B.A C.D.B=E.A C.D.B.E.A C.D.B=E.A C=D.B.E.A C=D.B.A.E C.D.B.E.A SC (L3/R3) 264/145 273/137 207/126 234/124 242/145 254/119 255/245 256/275** 276/276 223/206 Abbreviations: AF, arm formula (arm numbers arranged in length order, longest to shortest); AL, arm length (length of each arm pair, left/right); CL, calamus length; FL, funnel length; FFL, free funnel length; GC, gill count (number of gill lamellae per demibranch); GL, gill length; HW, head width; L1, L2, . . ., left arm 1, 2, etc.; LL, ligula length; MW, mantle width; NW, neck width; R1, R2, . . ., right arm 1, 2, etc.; Rg, regenerating arm; SC (L3/R3), sucker count (no. of suckers on 3rd arm of left/right side); SD, diameter of largest sucker (especially enlarged in males); SD (arms), arms with the largest suckers (particularly enlarged in males); SD (#), number of largest sucker counting distally from mouth (where more than one arm bears enlarged suckers, each value corresponds to respective SD arm); T, truncated arm; VML, ventral mantle length; WD, depth of interbrachial membrane (A–E, sectors of interbrachial membrane from dorsal to ventral); WDF, web depth formula (interbrachial membrane depths arranged in depth order, deepest to shallowest). *Originally 162 mm (shrinkage detected). **Arm R3 of this specimen has regenerated from an earlier trunca- tion distal to sucker #43; value given is from the next longest arm, L2. 34 Ian G. Gleadall papilla of modest size posterodorsally, a smaller one some- cream-grey, typically speckled with darker shades, chro- times visible anterodorsally (Fig. 3). matophore expansion creating deep brick-red to brown co- Skin with well-defined topology of raised ‘patches’ de- lours when undisturbed but alert, and also when fixed (Figs lineated by a system of recessed ‘grooves’ (Fig. 2), these not 2–3). Ventral mantle smoother and paler in colour (Figs 1, retained in specimens preserved following a delay between 3); sucker acetabula white to grey against an oral arm sur- death and fixation (e.g., storage on ice). Primary skin colour face of dark red (Fig. 4). Large skin papillae may be ob-

Table 2. Comparison of basic measurement indices for male and female specimens of Octopus sinensis and O. vulgaris.

Males Females Index O. sinensis O. vulgaris O. sinensis O. vulgaris ML range 72–168 mm 135–210 mm 89–151 mm 140–190 mm MWI 62±2.3 66±2.7 62±4.1 61±1.9 HWI 47±0.9 29±1.7 43±3.2 29±1.6 ALI 1 285±11.6 394±8.8 354±17.3 379±14.2 ALI 2 342±13.0 474±18.4 422±16.3 472±13.1 ALI L3 356±16.5 457±20.4 414±28.2 490±26.1 ALI R3 282±6.5 360±10.1 329±27.2 485±16.3 ALI 4 312±19.1 408±15.9 377±24.9 407±15.6 OAI 79±2.0 80±4.2 102±2.5 99±2.8 WDI max 86±3.6 89±3.4 84±5.7 93±4.2 WDI min 48±2.4 46±2.1 48±2.8 44±3.7 FLI 35±2.5 37±2.4 37±1.0 37±2.3 FFLI 72±7.8 65±5.5 69±3.9 70±1.4 LLI 1.5±0.3 3.3±0.2 — — CLI 42±4.5 38±3.5 — — GLI 29±1.7 26, 30 43±3.7 — GC range 7–9 9–10 8–10 9–10 SDI 16.9±0.8 19.8±0.9 13.1±0.7 13.9±1.1 SC (N) range1 207–273 322–350 223–276 320–342 SC (R3) range 119–152 154–192 — — Values are either mean (±standard error) or range (min.–max.). For O. sinensis, values are from all male specimens in Table 1; and for O. vulgaris include the male specimens listed as comparative material, and data tabulated by Guerra et al. (2010); 1, data for heavily damaged specimens omitted. Abbreviations as in Table 1: suffix ‘I’ indicates an index of measurement expressed as % ML, except for CLI (CL expressed as % LL), FFLI (FFL expressed as % FL), LLI (LL expressed % length of arm R3) and OAI (opposite arm index: length of arm R3 as % of that of arm L3). SC (N) range, sucker count range for normal arms (taking the largest value for each specimen); SC (R3) range, range of number of suckers on hectocotylus (males only, for all males inspected).

Fig. 1. Ventral view of a mature male specimen of Octopus sinensis (NSMT Mo 85658), illustrating the paler ventral surface (cf. dorsal sur- face of the same specimen visible in Figs 2 and 3) and the size and shape of the funnel (F, held in place with a piece of white cord). A sample (S) of muscle tissue was removed prior to fixation, leaving a circular wound just anterior to the funnel. Fig. 2. Dorsal view of the specimen in Fig. 1, showing the difference in lengths of arms 2 and 3 (longer, ending within the region labelled ‘Arms 2 and 3’ bracketed to the left) versus 1 and 4 (clearly shorter, ending within the adjacent bracketed region labelled ‘Arms 1 and 4’). The longer arms include arm R3 (the hectocotylus, H), which (typically much shorter than its counterpart, L3) is only just a little longer than arm pairs 1 and 4 (cf. Table 1, and OAI in Table 2). Note also the relative size and shape of the head, body and arms, and the rugose texture caused by the fixed appearance of the patch-and-groove topology of the skin. Fig. 3. View from the right side (body to left; arms to right) of the specimen in Figs 1 and 2. The funnel (F) was fixed in an unusual posi- tion (here seen pointing ventrally, perpendicular to the main body axis) as a result of damage following removal (before fixation) of a tissue sample (S) for molecular analysis (cf. Fig. 1). The broad extent of the mantle opening can be seen ventrally. Dorsally there is a large, contract- ed posterodorsal papilla (Ppd) above the right eye, with a smaller one anterodorsally (Pad). The anterodorsal papilla typically is not obvious except in well-fixed specimens. Fig. 4. An example of the appearance of the oral surface of a male Octopus sinensis to show the position of specially enlarged suckers on arms 2 and 3 of each side (most obvious on arms 2, as indicated: suckers #12 and 13 on the left, and #13 and 14 on the right). Anterior of (Ant.) is to the left, so the left arms are in the upper half of the figure and the right arms in the lower half. (Neotype specimen NSMT Mo 85659). Fig. 5. End of arm R3 of the specimen in Figs 1–3, showing the robust spermatophore groove (SGr; cf. Fig. 2, H) and small ligula (L). Fig. 6. Diagram of the tip of arm R3 of Octopus sinensis (lot NSMT Mo 85660 specimen IGG 318, ML 168 mm), illustrating the form of the ligula in more detail. Note modest calamus (Cal.) nestled in the hilum at the base of the ligula, and protuberance (Pr.) on the anterior side closely apposed to the region at the end of the spermatophore groove (SGr), covering the base of both ligula and calamus. (Line drawing by Aki Hamamoto). Fig. 7. Diagram of the tip of arm R3 of Octopus vulgaris specimen BMNH 98.5.21.346 (ML 96 mm), showing that the form and size of the ligula are similar to those of the ligula of O. sinensis (as shown in Figs 5 and 6). (Line drawing by Aki Hamamoto). Octopus sinensis 35 36 Ian G. Gleadall served on the dorsal mantle, including 4 papillae arranged length approximately half of ML, anterior section just over in a diamond configuration (two in the midline, and a later- a quarter of total stylet length. Gills typically with 8 or 9 al papilla on each side forming a symmetrically placed pair). (range 7–10) lamellae per demibranch; relatively long (GLI Hints seen of a pair of frontal white spots on the bases of 30–45; Table 2). Ink sac and anal cirri well-developed. arms 1 (no other conspicuous white spots observed). Spermatophores small, length around half of ML (approx- Arms long, longest around 4 times ML, shortest 3 to 3.5 imately 40 mm long in a male of ML 92 mm); released from times ML (growing allometrically throughout life, relatively a pseudophallus of modest size (length approximately 12% longer in larger ). Arms 2 and 3 subequal in length, of ML) with spermatophore duct entering posteriorly close longer than subequal arms 1 and 4, which are about 1 ML to dorsomedial bulbous expansion (occupying approximate- shorter than arms 2 and 3 (Fig. 2). Arm formula most com- ly 30% of total pseudophallus length; cf. Sasaki 1929, pl. IX monly 2.3.4.1 (Table 1). Arms robust and muscular, broader fig. 2a, b). Females lay small eggs in festoons made by en- than mantle when placed together (Figs 1–2); width of each twining egg stalks and cementing them to hang down from arm approximately 15% of ML at broadest extent (which roof of den. Eggs hatch into planktonic paralarvae of ML corresponds approximately to deepest extent of interbrachi- 1.5–2.0 mm and 3 suckers per arm (personal observations al membrane; Figs 1, 2, 4), narrowing gradually until distal from aquaculture experiments). third, then more abruptly to slender tips (Figs 1–2). Autot- Distribution. Known to occur in Japan as far north omy absent. Males with right arm 3 hectocotylized, shorter as southern Hokkaido and in the East China Sea off Japan than opposite arm by around 20% (OAI around 80%; Table (Sasaki 1929; Kaneko et al. 2011) and Taiwan (cf. Sasaki 2; cf. Fig. 2). Ligula of copulatory organ minute (Figs 5–6; 1929; Lü et al. 2013; Reid and Wilson, 2015), although the LLI 1.7, Table 2; cf. Sasaki 1929, pl. IX fig. 1), closely re- extent of its range southwards beyond Taiwan is currently sembling that of O. vulgaris (Fig. 7), with a relatively broad uncertain and requires further research. fleshy protuberance on anterior side closing ligula base (Fig. Remarks. The species was namedsinensis (Latin, mean- 6, Pr.; cf. Fig. 7); calamus length around 40% of that of ligula ing “from China”) even though the specimens upon which (Fig. 5; CLI 42%, Table 2), rounded, somewhat inconspicu- d’Orbigny’s description was based are known to have come ous, not raised from surface of ligula, and with closed me- from Japan (see further discussion below). d’Orbigny him- dian groove (Fig. 6, Cal.). Spermatophore guide with pale self mentioned Japan several times during the descrip- internal surface, present as a well-defined muscular edge tion and discussion, quoting from an original pre-Linnean along the marginal membrane of R3 (Fig. 2, H; Fig. 5, SGr). Japanese text clearly identified subsequently by Sasaki Arms bearing up to 276 suckers per normal arm in ma- (1929). The confusion with China appears to have aris- ture individuals (Tables 1–2); hectocotylized arm of males en from the use of Parisian scholars of Chinese hired by with 119–152 suckers (Table 2). Suckers robust, broad, 1 d’Orbigny (1839–1848) to translate the original text, which or 2 conspicuously enlarged in males between #12 and #15 was written in Chinese style characters (Takigawa, pers. (commonly #13 and/or #14) on arms 2 and 3 (cf. Fig. 4); in- comm.). fundibulum shallow, flared, with well-defined, scalloped rim The date of publication of the original description and about 32 well-developed cushions, coalescing to around (d’Orbigny 1841) was elucidated by Tillier and Boucher- 20 centrally; acetabulum moderately deep with small open- Rodoni (1994), who determined that the livraison contain- ing (Fig. 4). ing O. sinensis was published between 1839 and 1841. The Interbrachial membranes (arm webs) substantial but shal- earliest confirmed year for the publication of this descrip- low between all arms, deepest (sectors B, C and D) 80–90% tion is therefore 1841 (ICZN 1999: Article 21.6; Gleadall of ML (WDI max, Table 2), shallowest (sectors A and E) 45– and Salcedo-Vargas 2004). The original description by 50% of ML (WDI min, Table 2; Figs 1, 2 and 4); web depth d’Orbigny was inadequate and devoid of type or local- formula typically C.D.B.E.A (Table 1). ity designations but any confusion was resolved by Sasaki Funnel robust, modest in length, up to 40% of ML (FLI, (1929), who provided a thorough redescription with clari- Table 2; Figs 1, 3), moderately narrow, tapering abruptly fication of d’Orbigny’s ambiguities including confirmation (Fig. 1), free for about two-thirds of its length (FFLI, Table that the species in question was described from Japanese 2; Figs 1, 3); median basal notch approximately 12% of FL. specimens (despite the specific name alluding to China). Funnel attachment is directly beneath posterior region of There are no extant type specimens but it is deduced head, with free funnel extending just to anterior margin of (ICZN 1999: Article 73.1.4) that the holotype is that illus- eyes, well short of interbrachial membrane (Figs 1, 3). Fun- trated by d’Orbigny in his atlas of plates (d’Orbigny 1834– nel organ ‘W’ shaped, robust, with broad limbs, lateral limbs 1842; dated 1834 by Tillier and Boucher-Rodoni 1994), 20–25% shorter than median, occupying about half funnel Plate 9 (lower figure, without species identification; sub- length, positioned close to funnel tip. sequently identified by Sasaki 1929). The actual specimen Mantle robust, muscular, with moderately thick wall; (holotype by illustration) has never been identified. In view opening wide (up to 25% wider than neck width; mantle of the confusion over the number of species in the Octopus edge length approximately 2.5 times direct width across vulgaris complex, and the lack of extant type material for O. mantle aperture; cf. Fig. 1). A pair of non-calcareous stylets sinensis, a neotype is here designated to stabilize this species present embedded in dorsolateral mantle musculature, sup- name. The vouchers for O. sinensis listed above provide a porting origin of funnel retractor muscle of each side; stylet range of material for reference in museums in Australia, The Octopus sinensis 37

Netherlands, the UK and Japan for future comparisons with sinensis. other closely related species of the O. vulgaris complex. There is evidence of other closely related O. vulgaris-like It is noted that the collection of Japanese octopus speci- species in East Asian waters (Gleadall unpublished; Reid mens by Philipp Franz von Siebold and their deposition in and Wilson 2015) but this requires further research. The Leiden was probably communicated to d’Orbigny by De species described here as O. sinensis relates particularly to Haan, and that descriptions in Japanese were also pres- the main islands of Japan where it is a well-known, popular ent in the extensive libraries of Japanese works taken from and sought-after seafood item fetching a high market value. Japan to Leiden (by von Siebold) and to Paris (d’Orbigny Surveys of other localities around Japan have not detected 1841; Sasaki 1929; Gleadall and Goud 1993; Takigawa pers. the presence of any other species that could be confused comm.). Attempts have been made to locate correspondence with O. sinensis. With the description and sequences provid- among the early nineteenth century scientists concern- ed by the present study, which includes specimens also iden- ing specimens of Japanese Cephalopoda but none has been tified as the Japanese common octopus by Sasaki (1929), O. found (extensive enquiries of staff at BMNH, NRML, the sinensis is now a clearly described species with which other National Museum of Natural History, Paris, and the Natural closely related species can be compared. History Museum of La Rochelle). Japanese octopus speci- mens deposited in Leiden by von Siebold have been identi- fied in the present study as O. sinensis (see list of material Discussion examined, above), corresponding to the species described by d’Orbigny (1841) as the common octopus of Japan and The complex to which Octopus sinensis belongs is com- subsequently redescribed by Sasaki (1929). posed of species that are closely similar in appearance to O. Sasaki (1929) included Taiwan in his list of localities vulgaris, which has been described and illustrated by pre- for O. sinensis but the exact distribution of this species in vious authors (e.g., Naef 1923; Robson 1929; Nesis 1987), Chinese waters requires further research. The presence of although not in sufficient detail to recognize small differ- O. sinensis in Taiwanese waters is confirmed from identi- ences among the member species, even as population dif- cal gene sequences obtained recently from specimens col- ferences (but see Guerra et al., 2010). The major characters lected off Japan and Taiwan (compare for example sequenc- of this species complex are: medium to large octopuses with es from specimens identified as O. vulgaris in the studies an ovoid mantle; dorsal skin surfaces with a well-defined of Kaneko et al. 2011 and Lü et al. 2013). Sequences of the ‘patch-and-groove’ topology; prominent but modestly sized CO1 gene obtained from molecular analysis of tissue from eyes; adult specimens with 200–350 prominent suckers in the Japanese specimens listed here (Nishitani and Gleadall, two columns down each arm; arms robust, muscular; arm unpublished data) were all identical to those reported else- pairs 2 and 3 (dorso- and ventro-lateral pairs) and 1 and 4 where for specimens of the O. vulgaris complex from lo- (dorsal and ventral pairs) subequal, the former longer than calities off East Asia (cf. Guerra et al. 2010; Kaneko et al. the latter by a margin of around one mantle length; and hec- 2011; Amor et al. 2014). These will be submitted to a gene tocotylized arm (R3) length about 20% shorter than that of sequence database subsequently under the species name O. its partner (L3) on the left side (OAI in Table 2).

Fig. 8. Scatter diagram comparing the distribution of sucker number on the hectocotylus (third right arm) of males versus mantle length for preserved museum specimens of Octopus sinensis (white circles) and specimens of O. vulgaris (black discs), including those reported by Guerra et al. (2010) and also BMNH 1898.5.21.346 and ICM C468/1991. 38 Ian G. Gleadall

Fig. 9. Scatter diagram comparing longest and shortest arm lengths versus mantle length for Octopus sinensis (Osn) and O. vulgaris (Ovs) males (M) and females (F) as measured in the present study and (for O. vulgaris) in the study by Guerra et al. (2010). Values recorded are the longest of the longest four arms (2 and 3) and of the shortest four (arms 1 and 4) of each specimen (specimens as in Tables 1 and 2).

The description of O. sinensis (as Polypus vulgaris) by Sasaki (1929) is particularly appropriate because it includ- ed: detailed measurements; diagrams of the internal or- gans; specimens here identified as O. sinensis; and analysis following inspection also of specimens of O. vulgaris while Sasaki was in Europe. The original description of O. sinensis by d’Orbigny (1841) discussed several points of similarity with the European O. vulgaris. These points were highlight- ed in footnotes to a quotation of a direct translation from a Japanese text, which was identified clearly by Sasaki (1929) and confirmed also directly by the present author (Gleadall and Naggs 1991). Sasaki described the species in Japan as the commonest octopus of central and southern regions but identified it as O. vulgaris, with O. sinensis as a junior syn- onym. One of the major meristic characters distinguishing O. Fig. 10. Bar chart illustrating the difference in relative length sinensis from O. vulgaris is the number of suckers on the (expressed as %ML) between the longest and shortest arms for hectocotylus (the third right arm of males, which is modi- Octopus sinensis and O. vulgaris (bars represent means and stan- fied for transferring spermatophores during copulation). dard errors; measurement data and categories as in Fig. 9). As Fig. 8 shows, values for the two species comprise two mutually exclusive clusters which are independent of body end of the present paper). The total number of suckers on size: 119–152 in O. sinensis (cf. also Sasaki 1929; pers. obs. normal arms is also much fewer, by around 80 suckers, in O. of n=16 male specimens, ML 72–168 mm) vs. 154–192 in sinensis than O. vulgaris (Table 2). O. vulgaris (162–192 for n=8 intact specimens, ML 135– Enlarged suckers (1, or 2 adjacent) on arms 2 and 3 of 210 mm, listed by Guerra et al. 2010; 154, 160, pers. obs. mature males tend to be more proximal in O. sinensis (Fig. of voucher specimens listed as comparative material at the 4), between #12 and #14. This compares with around #14– Octopus sinensis 39

16 in O. vulgaris (even #17 or #18; Naef 1923). Images of sexual characters of males, as reported here for O. sinensis the world’s best known example of O. vulgaris (the mature and O. vulgaris. Identification of females is even more diffi- male known as Paul, associated with the 2010 FIFA World cult and at the present time no obvious distinguishing char- Cup football competition) show that specially enlarged acters have been reported to discriminate among females suckers on the arms of his left side were #14 and #15 on of species within (or closely related to) the O. vulgaris com- arm 2; #15 and #16 on arm 3 (suckers on his right side are plex, unless they possess ocelli (e.g., O. bimaculatus Verrill, not all visible in the images available on the Internet; pers. 1883, and O. bimaculoides Pickford and McConnaughey, obs.). The range in the position of enlarged suckers for O. 1949) or large eggs that do not have a planktonic stage of sinensis is similar to that reported for the closely related O. development (e.g., O. bimaculoides and O. maya Voss and insularis, a species confined to the eastern and central South Solís Ramírez, 1966; Forsythe and Hanlon 1988, and van Atlantic off Brazil (Leite et al. 2008) but apparently different Heukelem 1983, respectively). Further data are required to to that of O. mimus Gould, 1852, from the eastern Pacific clarify the distribution of populations currently referable to (e.g., Guerra et al. 1999). However, to test whether or not the species complex including O. sinensis, O. tetricus Gould, there are any statistically significant differences in the po- 1852 (either including or excluding O. gibbsi O’Shea, 1999), sition of specially enlarged suckers among members of the O. jollyorum Reid and Wilson, 2015, O. mimus, O. insularis O. vulgaris species complex will require a study of far larger and O. vulgaris; and to better identify the range of morpho- samples of each of the recognized species. logical and meristic differences among these and closely re- A summarized comparison of data for male and female lated species yet to be identified and named. specimens of O. sinensis and O. vulgaris (Table 2) reveals (in Considering relationships among these other closely re- addition to the differences mentioned above) the following lated species, it is noted that differences in the range of suck- apparent differences in morphology: O. sinensis has a much er counts on the male third right arm between specimens broader head, shallower maximum WD and deeper mini- of O. sinensis and O. vulgaris correspond in a similar way mum WD than O. vulgaris; and males have a shorter ligula to differences reported for eastern AntipodeanO. tetricus and smaller specially enlarged suckers. Other apparent dif- and Western Australian O. aff. tetricus, respectively (Amor ferences can probably be explained by the relatively smaller et al. 2014). This is interesting in view of the identification size range of the O. sinensis specimens obtained compared by Guerra et al. (2010) of specimens from the Amsterdam with those of O. vulgaris. For example, inspection of Table and St. Paul Islands in the Indian Ocean as O. vulgaris: these 2 suggests that the O. sinensis males, in particular, appear two islands are roughly midway between South Africa and to have relatively much shorter arms (2.8–3.5 vs. around Western Australia, slightly closer to the latter, lying within 4.0–4.9 times ML, respectively). Also, both male and fe- the prevailing currents moving eastward towards, and then male O. sinensis appear to have less of a difference in length northward along, the Western Australian coast. However, among the shortest and longest arms than O. vulgaris (Fig. according to the study by Amor et al. (2014), the two cur- 9): comparison of this difference (Fig. 10) shows a tendency rently recognized Australian species form a sister clade to for the difference to be less than 1 ML in O. sinensis, versus the East Asian species, here identified as O. sinensis, and are more than 1 ML in O. vulgaris (although the effects of the more distantly related to O. vulgaris. difference in the sizes of specimens sampled for these two Mangold (1983) presented a distribution map for O. species may partly explain this difference). Analysis and vulgaris which, for Asia, included India, Sri Lanka, Singa- presentation of data comparing the results of measurements pore, Southeast Asia and the Gulf of Thailand; and, for the (Tables 1–2; Figs 8–10) include the use of longest values for Atlantic, the Caribbean and eastern American coast con- arm lengths (Figs 9–10) in an attempt to minimize varia- tinuing along Central and South America as far as south- tion among arm lengths due to loss of arm tips following ern Brazil. Following the gradual splitting of this distribu- escape from attempted predation, with or without obvious tion into endemic regions, recognition of different species regeneration. This does not take into account any possible in the O. vulgaris complex begun by the study of Söller et asymmetry in arm length between the right and left sides, al. (2000) has now progressed to confirm as distinct the which have been reported occasionally for octopuses (e.g., East Asian population, here identified as O. sinensis. Figure Gleadall 1989; O’Shea 1999). However, no such asymmetry 11 summarizes the distribution of the species reported so was noted in the present study. far, although much research remains to be done in order to Details of the internal organs of species in this complex identify each of the species, and to clarify their distribution, have yet to be investigated and are beyond the scope of the particularly in the central West Atlantic, the western Pacific present study, the main objective of which is to demonstrate and the Indian Ocean. In the Caribbean area, the presence that O. sinensis is a different species from O. vulgaris based of Octopus vulgaris-like species has long been recognized on external morphology. A detailed study is currently under (e.g., Robson 1929; Pickford 1945, 1946) but the number of way comparing a number of different members of the O. species and their exact distribution is unknown. Other con- vulgaris complex, including O. sinensis (Amor and Strugnell, troversies still to be settled include the number and identity pers. comm.). of Antipodean species. For example, Amor et al. (2014) in- As with many animals, particularly , distin- terpreted their data to suggest that O. gibbsi is part of the O. guishing among closely related species using morphological tetricus population; and Reid and Wilson (2015) consider characters is often difficult and typically involves secondary O. jollyorum collected from the Kermadec Islands to be yet 40 Ian G. Gleadall

Fig. 11. Map showing the approximate distribution of Octopus vulgaris and closely related species. Abbreviations: O. bimac., O. bimacu- latus and O. bimaculoides (note that these two species are sympatric, with some segregation by depth, except that O. bimaculoides does not occur in the Sea of Cortez). Range for O. bimaculatus based on Ambrose (1997); O. bimaculoides, on Lang (1997); O. gibbsi, on O’Shea (1999); O. insularis, on Leite et al. (2008) and Amor et al. (2015); O. jollyorum, on Reid and Wilson (2015); O. maya, on Rosas et al. (2014); O. mimus, on Zúñiga et al. (2014); O. sinensis, on Sasaki (1929); O. tetricus and O. aff. tetricus, on Joll (1983) and Amor et al. (2014); and O. vulgaris, on Guerra et al. (2010). Note that the latter includes two distributions: one in the eastern North Atlantic and Mediterranean; and the other, here indicated as Octopus vulgaris (south), around the coast of South Africa and including Tristan da Cunha and the Amsterdam and St Paul Islands. The distribution of Octopus aff. vulgaris is based on that of the South American O. vulgaris clade sensu Leite et al. 2008. “?” indicates regions (southeast Asian waters; Indian Ocean; broader Caribbean) where the identity of species in the O. vulgaris complex is un- certain and requires further sampling and research. (Base map from www.geomapapp.org; Ryan et al. 2009). another separate species in the region, conspecific with O. Morocco and Mauritania (populations now collapsed and sinensis and closely related to O. tetricus (they did not men- over-exploited, respectively). Hopefully, the overt indepen- tion O. gibbsi). However, mitochondrial DNA sequencing dence of the O. sinensis population will draw more attention also seems to indicate that the eastern North Atlantic/Medi- to the need for focus on regular monitoring and the devis- terranean and South African populations comprise a single ing of appropriate strategies for its sustainable management, species (O. vulgaris, fide Guerra et al. 2010), yet they are ob- not only in Japanese waters but throughout the range of this viously allopatric (diverging?) forms (as depicted in Fig. 11). East Asian species. Equally, better controls over the exploita- The relationship between O. jollyorum and O. sinensis may tion of O. vulgaris are clearly required, since we now know prove to be similar, particularly in view of their wide geo- for certain that it is by no means the cosmopolitan species it graphic separation and difference in the number of suckers was once considered to be. on the hectocotylus (178–185 vs. 119–152, respectively). The Comparative material (Octopus vulgaris). 1 mature presence of a second East Asian species in the O. vulgaris male, ML 73 mm, Tyrrhenian Sea, Italy, Gulf of Naples, complex attributed to a personal communication from the BMNH 1898.5.21.346, undated, A. M. Norman. 1 mature present author by Reid and Wilson (2015) does not affect male, ML 96 mm, Eastern Central Atlantic, Fisheries Area the status of O. sinensis (since the distribution of this second 34, Canary Islands, ICM C468/1991, March, 1991. species does not include Japan) and will be reported on in a separate publication (Gleadall, in prep.). The significance of recognizing O. sinensis as a species Acknowledgements separate from O. vulgaris and other closely related species is profound with regard to fisheries management of this group I thank the people who helped with specimen collection, of octopuses, because it is now clear that many of these particularly Masami Abe (Gurumeito Co. Ltd., Ishinoma- species show a previously unsuspected level of endemism ki), Keiji Matsubara (Hotland and Gurumeito Octopus Re- (see Fig. 11) and will require careful monitoring and man- search Centre, Ishinomaki, and Kami-Amakusa Research agement if they are each to be harvested sustainably. More Centre, Iwajima, Kumamoto), Hiroto Teranaka and Hiroki than 30 years ago, Mangold (1983) noted that already there Takahashi (Kami-Amakusa City Hall), Hideo Sakaguchi had been a decades-long decline in the population of O. (Ehime Research Institute of Agriculture, Forestry and sinensis (which at that time she referred to as the Japanese Fisheries, Fisheries Research Centre), and my former stu- population of O. vulgaris). During the last 30 years, this de- dents Yuichirō Hino and Tomoyuki Itō. I thank Jonathan cline has been masked in Japan because of the importing Ablett (BMNH) and Roger Villanueva (ICM) for access to and consumption of large quantities of O. vulgaris from off preserved specimens; and Tsunemi Kubodera for arrang- Octopus sinensis 41 ing deposition of new material at NSMT. My thanks also go the name ‘Octopodia’ Schneider, 1784 (Class Cephalopoda), and of certain reputed names published by the same author in 1784. to Angel Guerra (CSIC Instituto de Investigaciones Mari- Opinions and Declarations of the International Commission on nas, Vigo) for information about the specimens of Octopus Zoological Nomenclature 4: 275–296. vulgaris used in the study reported by Guerra et al. (2010); ICZN 1997. Opinion 1861. Octopus vulgaris Cuvier, [1797] and Loligo to Michael Amor and Álvaro Roura (La Trobe University) vulgaris Lamarck, 1798 (, Cephalopoda): specific names for useful discussions; and to Aki Hamamoto for drawing conserved. 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