Some Old and New Genera of Octopus

Interdisciplinary Information Sciences, Vol. 10, No. 2, pp. 99–112 (2004)

Some Old and New Genera of Octopus

Ian G. GLEADALL

Tohoku Bunka Gakuen University, 6-45-1 Kunimi, Sendai 981-8551, Japan E-mail: [email protected]

Received September 1, 2003; final version accepted July 23, 2004

Two old, one new and two current genus names are discussed. The old names are Schizoctopus Hoyle, 1886, for which a type species is here designated, and Tritaxeopus Owen, 1881, which is here associated with the ‘Octopus horridus group’ of species. The two current genus names concern two species of octopus with males hectocotylized on the third left arm: Scaeurgus unicirrhus (delle Chiaje in d’Orbigny, 1841) and Pteroctopus tetracirrhus (delle Chiaje, 1830). The type species of the genus names Scaeurgus and Pteroctopus have been identified as synonyms (rendering these genus names synonyms, also) but approval of an unchallenged redesignation of the type species of Scaeurgus provides an appropriate solution to this problem that would best maintain taxonomic stability. A new genus name is proposed for the species Octopus januarii Hoyle, 1885, which was previously placed in Benthoctopus Grimpe, 1921. This follows the recent identification of the type species of three genera (Bathypolypus, Benthoctopus and Atlantoctopus, all described by Grimpe, 1921) as belonging to a single genus (excluding Octopus januarii), of which Bathypolypus is the senior synonym. It is here observed that the species closest in morphology to Hoyle’s ‘Octopus’ januarii appears to be ‘Octopus’ longispadiceus (Sasaki, 1917), although the former has no ink sac while the latter has. The common features of these two species include close similarities in the internal structure of the male pseudophallus. Comparisons are made with structural features of the pseudophallus in Enteroctopus dofleini (Wu¨lker, 1910), ‘Octopus’ hongkongensis Hoyle, 1885, and ‘O.’ conispadiceus (Sasaki, 1917), and it is suggested that all five species may be related at the subfamily level.

Introduction

There is currently a great deal of research under way to clarify the biodiversity of octopuses in the world’s oceans. An emerging trend is that many species can now be removed from former ‘hold-all’ genera such as Octopus and Benthoctopus. Some have been identified as belonging to formerly poorly known genera and other species have been placed in new genera. One example is the old genus name Amphioctopus Fischer, 1882, which until recently was a monospecific genus but for which 11 species (formerly in genus Octopus) have now been identified (cf. Gleadall, 2002; genus redescription submitted). The present paper designates a type species for Schizoctopus Hoyle, 1886 (here shown to be a synonym of Amphioctopus), and addresses the synonymy of two genera of medium sized octopuses with males that (in contrast to other members of the Octopodidae) have their third left arm modified for reproduction. Several species from Japan have a pseudophallus with the spermatophoric duct joining close to its anterior end, rather than joining (and entering) posteriorly as in Amphioctopus and Octopus (Gleadall, 2002). The morphology and internal structure of the pseudophallus of these species is compared: all have an arch structure on the internal surface of the dorsal wall. Such an arch is found also in the inkless species ‘Octopus’ januarii Hoyle, 1885, which is here given a new genus name.

Abbreviations: AMS, Australian Museum, Sydney; BMNH, The Natural History Museum, London (formerly the British Museum of Natural History); DML, dorsal mantle length; ICZN, International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999); L, left (arm); NMV, Museum Victoria, Melbourne; NRML, National Museum of Natural History—Naturalis, Leiden; PMBC, Phuket Marine Biological Centre; R, right (arm); UMML, University of Miami Marine Laboratory (specifically the museum in the Rosenstiel School of Marine and Atmospheric Sciences); ZMUC, Zoological Museum at the University of Copenhagen; ZUMT, Zoology Department of Tokyo University Museum.

1. Schizoctopus Hoyle, 1886

‘Steenstrup’s group Schizoctopus’ Hoyle, 1886a: 81. ?Octopus (Schizoctopus) Jatta, 1889: 64; 1899: 2.

Etymology: Schizo– (Gk. ‘split’) plus octopus, presumably referring to the very short sector of interbrachial web between the most dorsal arms (arms I).

Type species: Octopus fangsiao d’Orbigny, 1841, here designated. 100 GLEADALL

Material examined. Two females, DML 44 and 37 mm, submature, collected by Reinhardt from Hong Kong on the Galathea Expedition of 1845–1847 (see Bille, 1850), ZMUC CEP-12.

In a short paper and in his monograph on the Octopodinae, Robson (1929a, b) mentioned two genus names that he regarded as invalid: Amphioctopus Fischer, 1882, and Schizoctopus Hoyle, 1886. Both names are associated with species that Robson called the ‘group of Octopus aegina.’ Recently, many of these species have been included in the genus Amphioctopus (Gleadall, 2002) but no further mention has been made of Schizoctopus, for which no type species has ever been designated. Identifying two specimens as Octopus granulatus Lamarck, Hoyle (1886a) stated: ‘Like most other rough- skinned species of Octopus, this belongs to Professor Steenstrup’s group Schizoctopus, characterised by having the umbrella between the dorsal arms very short and the cutaneous sculpture continued over its inner surface.’ Steenstrup did not publish this genus group name and Hoyle did not actually apply it to any of his identifica- tions, nor did he even mention Schizoctopus again (cf. Hoyle, 1910). Jatta (1889) interpreted Hoyle’s identification as a Fig. 1. (A) Ventrolateral view of the two specimens at ZMUC Schizoctopus subgenus of Octopus, listing (without further comment) (lot CEP-12) labelled by Steenstrup, with obvious ocelli (oc). (B) Dorsal view of specimen to the right three specimens of ‘Octopus (Schizoctopus) areolatus’ in (A), showing the head patch (h). Note ‘patch-and-groove’ (locality not stated) and one specimen of ‘Octopus sculpturing of skin, particularly on dorsal mantle. Scale bars (Schizoctopus) granulatus’ from near Pernambuco (Re- 20 mm. cife, Brazil?). These specimens have not been located and are presumed to be no longer extant. Octopus areolatus is a nomen dubium fixed (along with O. ocellatus Gray, 1849, and O. brocki Ortmann, 1888) as a junior synonym of the Japanese species Octopus fangsiao by Sasaki (1920, 1929), the first reviser of this species (cf. ICZN Art. 24.2; see also Roeleveld, Goud and Gleadall, 2003). The original description of Octopus granulatus Lamarck, 1798, was very brief (‘octopus corpore tuberculis sparsis granulato, cotyledonibus crebris biserialibus’) but was soon followed by a more detailed description (Lamarck, 1799) that mentioned two syntypes. However, no measurements of the specimens were ever published, they were not illustrated, they are no longer extant (Lu, Boucher- Rodoni and Tillier, 1995), no type locality was stated and none of the descriptions in the literature is sufficient to distinguish O. granulatus from species such as Octopus vulgaris Cuvier, 1797. One of the specimens identified by Hoyle (1886a) as O. granulatus was subsequently identified as a new species: Octopus vincenti Pickford, 1955, a junior synonym of Octopus burryi Voss, 1950 (Toll and Voss, 1998; personal observation), which is one of the species included in the genus Amphioctopus (Gleadall, 2002). Lamarck (1799) suggested that O. granulatus may be a synonym of Octopus rugosus (Bosc, 1792), but since neither species has been described sufficiently to enable distinction from other species of octopus and none of the type material is extant, they are now both widely regarded as dubious species, most recently as queried junior synonyms of O. vulgaris (see, for example, Mangold, 1998; cf. Robson, 1929a, and Pickford, 1955). Octopus cf. rugosus from southern Australasia described by O’Shea (1999) appears to be related to Amphioctopus marginatus (Taki, 1964), including comparison of the form of the pseudophallus (cf. Gleadall, 2002). To summarize, there has been no previous designation of a type species for the genus group name Schizoctopus: Hoyle did not identify any of his species as either a genus or subgenus Schizoctopus (cf. ICZN Art. 67.5.1); Jatta’s identifications of two species in the group Octopus (Schizoctopus) were without comment, and are ineligible in considering a subsequent type species designation by Jatta (e.g., by monotypy: ICZN Art. 69.3, 69.4); and although Robson clearly treated Schizoctopus at the genus level, he considered it an invalid name, did not apply it in any of his identifications and did not otherwise mention any species names associated with it. However, Jatta’s action in listing O. areolatus and O. granulatus defined the included nominal species for the genus group name Schizoctopus (ICZN Art. 67.2.2), from which the type species should be selected (ICZN Art. 67.2). The material examined was obtained on loan upon requesting specimens labelled ‘Schizoctopus’ by Steenstrup (as indicated by Hoyle). The labels with the specimens state: ‘Octopus (Schizoctopus) areolatus.’ The specimens are in excellent condition and, since both have a prominent head bar and large ocelli with an iridescent annulus (Fig. 1A, B), Old and New Octopus Genera 101 they are identified without doubt as Octopus fangsiao (cf. Sasaki, 1929). Since O. areolatus was the first species listed by Jatta (1889), it could be designated as type species by position precedence (cf. ICZN Rec. 69A.10). However, since both included nominal species are nomina dubia and the specimens are no longer extant (see above), stability is best served by designating the species actually involved in the original publication of the genus group name. Therefore, the type species of Schizoctopus Hoyle, 1886, is here fixed (under Article 70.3.2 of the ICZN) as Octopus fangsiao d’Orbigny, 1841, misidentified as Octopus granulatus Lamarck, 1798, by Hoyle (1886) and mislabelled as Octopus (Schizoctopus) areolatus de Haan in d’Orbigny, 1841. Octopus fangsiao is also a nominal species included in the genus Amphioctopus (Gleadall, 2002) and, since Amphioctopus is the older name, the genus group name Schizoctopus is a junior synonym of Amphioctopus.

2. Scaeurgus Troschel, 1857, and Pteroctopus Fischer, 1882

Description of a new genus of octopus from Hong Kong (submitted) required that comparisons be made with other known genera of the Octopodinae, particularly with those including species with a ‘limus’ (a raised wine-red ridge passing horizontally around the ventral mantle; Gleadall, 2002). During that study, it was noticed that there is a problem regarding usage of the genus Scaeurgus Troschel, 1857, to describe the monospecific genus containing Octopus unicirrhus delle Chiaje in d’Orbigny, 1841 (one of the species recognized as possessing a limus). Troschel’s original description was of two species: Scaeurgus titanotus (new genus and species) and Scaeurgus coccoi (originally Octopus coccoi Ve´rany, 1846). The type species is Scaeurgus titanotus by subsequent designation (Hoyle, 1910). Robson (1929b, p. 191) had stated: ‘I retain Scaeurgus as the generic name for unicirrhus (coccoi) in spite of the fact that Hoyle (1910, p. 412) designated Scaeurgus titanotus Troschel as the type of the genus. Hoyle was apparently unaware of the fact that Fischer (1882, p. 334) had previously made Octopus tetracirrhus (titanotus) the type of his Pteroctopus. The latter genus actually does not figure in Hoyle’s list. As I think it is quite clear (cf. p. 196) that Troschel’s titanotus is delle Chiaje’s tetracirrhus, and as it is proposed to maintain Fischer’s genus, it follows that unicirrhus must be regarded as the genotype [= type species] of Scaeurgus.’ Robson had therefore attempted to change the type species designation of Scaeurgus from S. titanotus (identified as a junior synonym of O. tetracirrhus)toOctopus unicirrhus (regarded as the senior synonym of O. coccoi). Hoyle’s (1910) review of the Cephalopoda genera presumably excluded Pteroctopus Fischer because he regarded it as a junior synonym of Scaeurgus Troschel (cf. also the descriptions and diagrams of both species as Octopus (‘‘Scaeurgus’’ ) b y Naef, 1921–1923). Once validly fixed, the type species of a genus is immutable (ICZN Art. 69.1), so the type species of Scaeurgus is still Scaeurgus titanotus. Since Scaeurgus and Pteroctopus are considered to be synonyms describing Octopus tetracirrhus, the species Octopus unicirrhus (widely recognized as belonging in a genus different from Octopus as well as from Scaeurgus or Pteroctopus tetracirrhus) should therefore require a new genus name. However, since attention has not been drawn to this problem since Robson’s actions, and currently there are only two monospecific genera involved, the best solution to maintain stability is to clarify the type species of the two genera by applying to the International Commission on Zoological Nomenclature to use plenary powers to change the designation of the type species of Scaeurgus to Octopus unicirrhus as recommended by Robson (1929b). This would maintain current usage of the two binomial names Pteroctopus tetracirrhus (delle Chiaje, 1830) and Scaeurgus unicirrhus (delle Chiaje in d’Orbigny, 1841). With regard to the species names unicirrhus and tetracirrhus, note that the suffix –cirrhus (from the Greek o&, yellow; as in hepatic ‘cirrhosis’) is actually an incorrect spelling for –cirrus (from the Latin word, meaning a curl-like tuft, fringe or filament, describing the appearance of prominent papillae above the eyes of these octopuses). However, even though incorrect these spellings are in common use and should be retained (ICZN Art. 33).

3. Tritaxeopus Owen, 1881

?Octopus ‘body granular’ group of Gray, 1849: 8 (in part). Tritaxeopus Owen, 1881: 131, Pl. 23; Fischer, 1882: 56; 1887: 334. Octopus (Tritaxeopus) Robson, 1929a: 607; 1929b: 172. Octopus horridus species group of Norman and Sweeney, 1997: 96, 108; Norman, 1998: 800. Octopus (Abdopus) Norman and Finn, 2001: 14.

Etymology: Trit (Gk. ‘three’) + axeo (Gk. ‘rank’) + pus (Gk. ‘foot’).

Type species: Tritaxeopus cornutus Owen, 1881, by monotypy.

Genus Tritaxeopus was described from a single specimen, identified as Tritaxeopus cornutus Owen, 1881 (the 102 GLEADALL species name being derived from the presence of eye Table 1. Measurements (mm)1 of the holotype of Tritaxeopus papillae ‘so large and pointed as to simulate horns’). This cornutus Owen, 1881. holotype by monotypy apparently is no longer extant and Mantle length (‘length of the body’) 95 the only information about the locality is that it was from Mantle width 66 ‘Australia.’ The distinguishing feature of the genus is the Head width 57 presence of suckers in three rows ‘along more or less of Length of shortest arm (arm 1)2 356 each arm.’ Owen expanded on this description as follows. Length of longest arm (arm 3)2 584 The suckers begin as a single series at the base of the arm Arm diameter 25 then, after the third or fourth sucker, ‘assume the ordinary Depth of interbrachial web sector B 38 biserial arrangement; then the two series diverge after a Depth of interbrachial web sector C 64 short course to make way for a third supplementary row, 1Converted from the original imperial units. 2Owen did not state ...’ which continues along the arm to the attenuated last whether arm measurements referred to arms of one or both sides. quarter of the arm, from which the suckers fall again into two rows. Finally, at ‘the filamentary terminations of the arms’ the suckers are arranged singly once more. Other details in Owen’s description included arm length order 3.2.4.1; interbrachial membrane about two- thirds of the mantle length. Body surface with scattered wart-like prominences, particularly dorsally, ‘of which four or five of the largest affect a longitudinal disposition’ (cf. Fig. 2). Eyes with prominent, horn-like papillae above each eye. Longest arm six times mantle length (other measurements provided by Owen are listed in Table 1). Arms semicircular in cross-section, with the suckers on the flat side; suckers sessile with soft, thick border and cushions divided by conspicuous folds. Mantle with wide aperture terminating close to the posterior of the eyes and with a robust free margin. Funnel free for much of its length. The living animal was said to emerge from the water frequently, inhabiting recesses on the shore during low water. Colouration dullish pink; arm crown a ‘subviolet tint.’ When alarmed, colour changes include bluish red to deep violet. Inner surface of interbrachial membrane (‘coronal membrane’) lighter; whitish inner lip (at mouth). Owen’s account describes a long-armed octopus similar to a species common throughout warm waters of the Pacific in littoral and sublittoral habitats, typically found occupying cavities among coral rubble in the intertidal zone. Specimens of this common species inspected by the present author include: the holotype of ‘Octopus’ inconspicuus Brock, 1887, from Ambon, Indonesia (see Fig. 2; Tables 2, 3); specimens collected at first hand from the shores of the Andaman Sea on sandy and coral rubble beaches in Phuket, Thailand, in 1996 and 2002 (Figs. 3A, B, 4; Table 3); and a specimen (misidentified by the vernacular name ‘madako’) ob- served in video sequences taken off Kashiwa Island in southeastern Shikoku, Japan (in the television programme Kuroshio Kaichuˆ Sanpoˆ, NHK [Japanese Broadcasting Corporation], January 2003). Resting living specimens show a chronic ‘rugose general resemblance pattern’ (Hanlon et al., 1999) which involves extreme papillation of skin over the eyes, including several long, branching papillae (personal observations; cf. Norman and Sweeney, Fig. 2. Illustration of the holotype of ‘Octopus’ inconspicuus 1997; Norman and Finn, 2001; cf. Owen’s illustration of (Brock, 1887). Male, DML 45 mm, locality Amboina T. cornutus, here reproduced as Fig. 3C). One small [Banda Sea, Indonesia, Maluku, Ambon], ZMUG 116. The differences in arm lengths and widths obviously reflect difference from the description of Owen’s species is that the various states of regeneration of the arms following the suckers are paired to the extremity of the arms earlier autotomy (see text; cf. Brock, 1887; Norman and (although the distal suckers are so small that a microscope Finn, 2001). p, primary papillae; n, network of dark lines. Old and New Octopus Genera 103 is required to confirm this). This common, small-egged Table 2. Measurements (mm) of the holotype of ‘Octopus’ species from Asian waters is identified as ‘Octopus’ inconspicuus Brock, 1887. aculeatus d’Orbigny, 1841 (Table 3; cf. Norman and Dorsal mantle length 45 Finn, 2001). Ventral mantle length 31 Three out of 12 fixed specimens of ‘Octopus’ aculeatus Mantle width 26 recently obtained from the Andaman Sea have some Head width 20 suckers arranged in a series of apparent triplets along Head length 11.5 sections of one or more arms (Figs. 3A, B). Inspection Neck width 15 reveals that this is an illusion probably caused by Funnel length 17 distortion of doublet sucker positions during fixation. It Free funnel length 8 has not been observed in the living animals, nor has the Length of arms 1 left/right 187 (136) present author seen this phenomenon in any other kind of 2 242* (157) octopus (following inspection of hundreds of museum 3 (134) 202 specimens; cf. also Table 5). Inspection of Owen’s 4 165 215 original Figure 1 of his Plate 23 (reproduced in Fig. 3C Web depth sector A 17 of the present paper) reveals that the extent of apparent B left/right 22 20 triplets of suckers in T. cornutus resembles that observed C2421 in the recent specimens, rather than the extent implied by D2321 Owen’s statement (compare the extent of triplets in E18 Figs. 3A–C, but cf. Fig. 3D). Width of largest arm 7 In his account of Tritaxeopus (as a subgenus of Diameter of largest sucker, number 12 on arm R2 6.3 Octopus), Robson (1929b) stated ‘This genus was (numerals in parentheses are measurements of truncated arms). As in founded by Gray.’ However, no reference was provided specimens of ‘Octopus’ aculeatus, the suckers are in tightly crowded and it is presumed that the statement was an error. It is pairs right to the end of the arms. *Arm L2 is the longest intact arm interesting to note, though, that Gray published an but it is clearly regenerating beyond the 47th sucker, where this arm narrows abruptly.

Table 3. Comparison of Owen’s measurements and counts of Tritaxeopus cornutus with corresponding measurements for specimens of Octopus’ aculeatus, ‘Octopus’ inconspicuus, and ‘Octopus’ sp. from Phuket (here proposed as congeneric, and possibly conspecific) and representative species of three other genera: Octopus (O. cyanea), Amphioctopus (A. fangsiao) and the ‘Octopus macropus group’ member ‘Octopus’ minor (all measurements expressed as % DML).

T. cornutus1 ‘O.’ aculeatus2 ‘O.’ inconsp.3;5 ‘Octopus’ sp.4;5 O. cyanea6 A. fangsiao7 ‘O.’ minor8 Mantle width 69 66 58 63 71 83 49 Head width 60 51 44 45 56 46 27 Shortest arm (arm #1) 374 410 416y 429* 419 259 231 (L4) Longest arm 615 (arm #3) 668 (arm L3) 538 (L2)y 739 (L3)* 554 (R3) 322 (L4) 538 (L1) Arm diameter 26 22 16 17 16 15 12 Web sector B 40 — 47 63* 63 61 54 Web sector C 67 78 51 79* 71 68 45 No. of suckers on longest arm) 268 225 214y 20310 397 (L3) 150 (L3) 194 (L1) (190–262)9 No. of suckers on hectocotylus not stated; damaged 162 150 (female) 89 49 sex unknown (138–175) The species here proposed as congeneric are to the left of the vertical line near the centre of the table. 1see Table 1; 2based on male specimen, DML 41 mm (NMV F67015; fide Norman and Finn, 2001), with sucker count ranges from the data of Norman and Finn in parentheses for Australian and Philippine populations combined; 3see Table 2 (DML 45 mm); 4male, DML 38 mm (author’s specimen 1996.2.18.3); 5data fall within ranges of those for ‘O.’ aculeatus (see Norman and Finn, 2001); 6Octopus cyanea Gray, 1849, female, DML 68 mm (NRML 488; cf. also measurements of O. cyanea in Norman, 1992a, including a maximum recorded arm length of 582% of DML, for arm R3 of specimen AMS C162598); 7Amphioctopus fangsiao (d’Orbigny, 1841), male, DML 41 mm (ZUMT 994.CO-03); 8‘Octopus’ minor (Sasaki, 1920), male, DML 84 mm (ZUMT 996.3.9). 9Largest value in range is for a specimen of DML 58 mm (NMV F67014; Norman and Finn, 2001). 10Including approx. 20 suckers on regenerating arm tip. yShortest arm: arm L1 is the shortest arm of ‘O.’ inconspicuus that does not appear to be regenerating. Longest arm: note that left arm #3 is damaged (see Table 2) and right arm #3 is hectocotylized (and therefore much shorter than arm L3), hence the longest arm available is the relatively much shorter left arm #2, which is regenerating (see footnote in Table 2; cf. data for ‘O.’ aculeatus in Norman and Finn, 2001). *Possibly artifactually long arm and web sector measurements because arms are tightly curled back. account of a single species (‘Octopus rugosus’) in his subgroup ‘body granular’ (which was part of a higher grouping of octopuses, including O. vulgaris, with suckers of similar size, far apart and in one row near the tip of the arm; Gray, 1849). This description included the characters: body oval, purse-shaped, large; head, arms and upper part of body covered with rounded tubercles; head short, warty, ocular beard one, elongated; arms short, thick, conical, order 4.3.2.1; suckers large, somewhat smaller towards the base of the arm and towards the tip; web short; when alive, violet- brown, white beneath; side of the arms netted with brown lines. The only locality given by Gray was Valparaiso, Chile. 104 GLEADALL

Fig. 3. Comparison of specimens with suckers in a triple row (apparently) in recently preserved ‘Octopus’ sp., collected from the Andaman Sea, Thailand (A, B) in comparison with the original illustrations of Tritaxeopus cornutus Owen (C, D). (A) Submature female, DML 45 mm, collected on Phanwa Beach, Phuket, by Tah, Vongpanich and Gleadall, 5.iii.2002 (removed from a hole in coral rubble). (B) Arm L4 of a mature female, DML 59mm, collected from inshore sand flats on Rawai Beach, Phuket, by Chotiyaputta and Gleadall, 18.ii.1996. (Both specimens will be deposited in BMNH). (C) Owen’s original Plate 23, Figure 1, a somewhat stylized illustration of the dorsal view of T. cornutus. (D) Owen’s Plate 23, Figure 2, a diagram of the proximal suckers on the arm crown of T. cornutus. In Figs. A–C, apparent triplets are each indicated between arrowheads.

However, he also included the ‘habitats’ Atlantic Ocean and Indian Ocean. Gray’s account appears to have confounded descriptions of at least two different genera but it seems to include a description of specimens attributable to the genus Tritaxeopus from the Indian Ocean. Octopus rugosus (Bosc, 1792) is a dubious species for which there are no extant types (Lu et al., 1995) and which is widely regarded as a junior synonym of O. vulgaris Cuvier (see Section 1, above). A network of brown lines on the arms, mentioned by Gray, is clearly visible in the holotype of ‘Octopus’ inconspicuus (Fig. 2), which is here included among the specimens identified as ‘Octopus’ aculeatus. This network of lines is possibly related to the intricate pattern of the startle response (Fig. 4). It is possible that Owen’s Tritaxeopus cornutus, too, is a synonym of ‘Octopus’ aculeatus. Its exact affinities at the species level cannot be confirmed at the present time because there is no extant type material and more than one species in this group is extant in Australia. However, the description and measurements given by Owen, summarized here (Tables 1, 3), leave little doubt that T. cornutus is congeneric with species in the genus group Abdopus Norman and Finn, 2001 (cf. their measurements and description, including the presence of a papilla above each eye that can be raised in distinct ‘horns’ in ‘Octopus’[=Tritaxeopus] aculeatus). In comparing maximum arm lengths, the longest arm for a member of the genus Octopus sensu stricto (i.e., Octopus Cuvier) appears to be less than 6 times DML for O. cyanea, while in species including Tritaxeopus cornutus, the longest arm (in intact specimens) is 6 to 7 times DML. Norman and Sweeney (1997) have quoted an arm length range of 4–7 times DML for the Octopus vulgaris species group of Robson, 1929b (Octopus sensu stricto), although the only species of Octopus sensu stricto included in their study of octopuses from the Philippines was O. cyanea, and the maximum arm length for that species (stated in Table 3 Old and New Octopus Genera 105

Fig. 4. The startle response (dymantic or deimatic display) of ‘Octopus’ sp. from Phuket, based on digital movie and still images. of the present paper; cf. Norman, 1992a) was rounded up from 5.8 to 6 times DML. Also, as hinted at in the footnotes to Table 3, it would be useful to have more detail about the extent of regeneration in the arms of specimens used to redescribe species of the O. horridus group in order to clarify differences in the relationship between mantle and arm lengths among these two groups of species. A redescription of the group of Octopus sensu stricto is required in order to confirm whether or not there is a significant difference in the relative lengths of undamaged arms. The inkless species Ameloctopus litoralis, too, has very long arms but, in addition to lacking an ink sac, its general morphology and body patterning (Norman, 1992b) are very different from the description of Tritaxeopus cornutus furnished by Owen, so it is not in contention as a possible alternative identification. Species in other octopodine genera (here represented by Amphioctopus fangsiao and ‘Octopus’ minor), also, are easily distinguished from Tritaxeopus (Table 3). The much larger size of Owen’s T. cornutus in comparison with specimens of ‘Octopus’ aculeatus recorded by Norman and Finn is not regarded as problematic: Mangold (1998), for example, noted a DML range of 20–250 mm for Octopus vulgaris; and mature specimens of Amphioctopus fangsiao have been preserved over the range 29–77 mm (personal observations). Owen’s description of sucker triplets extending along a more substantial length of the arms suggests that either the extent of the triplet artifact was unusual in his type specimen (perhaps related to its large size) or Owen’s description of it may have been overstated (compare Figs. 3C, D; see above). The occurrence of similar sucker triplet artifacts on the arms of recent specimens and comparisons with Owen’s distinctive morphological measurements and sucker count, detailed description of the body colouration, morphology of the arms and suckers, and the intertidal habit, leave little doubt that Tritaxeopus is a genus name describing the ‘Octopus horridus group’ of octopuses.

4. Muusoctopus, new genus

Family Octopodidae d’Orbigny, 1839 Subfamily (to be determined) Genus Muusoctopus, gen. nov.

Diagnosis. Mantle globose, head broad, eyes relatively large. Arms long, slender, cylindrical, 3–4 times DML, arms 1 and 2 longer than 3 and 4. Suckers in two moderately spaced rows. Interbrachial membranes of moderate, subequal depth (just less than DML), a little shallower between arms 3 and 4; pouches absent. Males with hectocotylus on third right arm. Pseudophallus relatively large, with moderately muscular wall and comprising two chambers, one anteromedial and one posterolateral; spermatophoric duct joins anteriorly, entering along dorsal surface of anteromedial chamber; dorsal arch present, located posterior to entrance of spermatophoric duct, forming passage 106 GLEADALL between the two chambers. Funnel large, free for half its length. Posterior salivary glands of modest size, flat, triangular, almost equilateral. Intestine with hairpin loop to right side. Skin without well defined patch-and-groove system. Arm autotomy absent.

Type species: Octopus januarii Hoyle, 1885 (other included species yet to be determined).

Etymology: Muus– (surname prefix) + octopus, after Bent Muus, the reviser of genus Bathypolypus.

Muusoctopus januarii (Hoyle, 1885), new combination (Figs. 5, 6A, 7A; Tables 4, 5)

Octopus januarii Hoyle, 1885a: 229 (in part); 1885b: 105 (in part); 1886a: 97 (in part), Pl. 7 Figs. 1–4; 1886b: 220 (in part?). (not O. januarii—Goodrich, 1896: 19). Not Polypus januarii—Hoyle, 1904: 18; 1909: 260; Berry, 1912: 392; Massy, 1916: 199; Sasaki, 1920: 172; 1929: 61. Benthoctopus januarii—Robson, 1929b: 41 (in part); 1932: 235, Figs. 33a, 40, 41; Voss, 1968: 656; Toll, 1981: 84, Figs. 1A–J, 3A, D–G; Roper, Sweeney and Nauen, 1984: 223; Nesis, 1987: 320, Figs. 84O, P; Lipinski, Naggs and Roeleveld, 2000: 111.

Description. Small to medium sized species (DML to Fig. 5. Illustration of Muusoctopus januarii (Hoyle, 1885). 60 mm). Mantle globose, aperture wide; limus absent. Funnel robust, tapering, free for half its length; funnel organ not preserved. Eyes large (head length almost half DML). Arms long and slender, cylindrical cross-section; 3–4 times mantle length, in the order 1.2.3.4: arms 1 and 2 markedly longer than 3 and 4 (Fig. 5, 6A, Table 4). Interbrachial membranes a little shorter than (up to 85% of) DML, subequal, slightly shorter in sectors D and E. Suckers small, with small infundibulum, in two moderately spaced rows directly from mouth; none especially enlarged. Hectocotylized third right arm (two-thirds the length of third left arm) with approximately 80 suckers (83 in lectotype, 71–91 in other specimens; Table 5). Longest unmodified arms with sucker count of approximately 180 (176 on right arm 1 of lectotype). Ligula modest in size (approx. 8% of length of third right arm), with distinct margin surrounding wide, flat, shallow inner surface, tapering evenly to an acute point; approximately 20 weakly developed transverse ridges; calamus well defined, sharply pointed (illustrated by Toll, 1981; that of lectotype illustrated by Robson, 1932: 223 Fig. 33a); spermatophoric groove well developed. Pseudophallus large (similar length as ligula, approximately 20% of DML), spermatophoric duct joining close to anterior, turning in posterior direction as it joins. Spermatophores in spermatophore sac 10–15 (14 in lectotype), slim, approximately 85 mm in length. Gill lamellae 7–8 per demibranch. For other measurements, see Table 4 and Toll (1981). Skin without well defined patch-and-groove system. Ink sac and anal cirri absent. Colouration in ethanol uniform pinkish grey, slightly paler beneath. Some specimens show evidence of approximately nine rows of short, thin, straight dark lines arranged longitudinally on the dorsal mantle (Figs. 5, 6A).

Type material: Lectotype by subsequent designation (Robson, 1932: 235) 1 mature male, DML 49 mm, BMNH 1889.4.24.41. Collected on the Challenger Expedition, 10.ix.1873. (One other former syntype, locality Yokohama, Japan, identified as a different species; see below).

Other material examined: 1 male, 60 mm DML, Oregon Sta. 3670, 400 shrimp trawl, 30.vii.1962, Gulf of Mexico, off New Orleans, 732 m, UMML 31.442; 1 male, 41 mm DML, Oregon Sta. 3739, 1000 shrimp trawl, 26.viii.1962, Gulf of Mexico, off New Orleans, 494 m, UMML 31.443; 1 male, 39 mm DML, Silver Bay Sta. 1203, 400 flat trawl, 11– 12.vi.1959, Gulf of Mexico, off New Orleans, 458 m, UMML 31.463; 2 males, 50 mm and approx. 42 mm DML, Oregon Sta. 3565, 400 shrimp trawl, 21.v.1962, Caribbean Sea, off Nicaragua, between Cayos Miskito and Quita Suen˜o Bank, 439–458 m, UMML 31.923; 1 male, 46 mm DML, Oregon Sta. 2771, 400 shrimp trawl, 15.iv.1960, Caribbean Sea, Lesser Antilles, off Venezuela, between Grenada and Los Testigos, 403 m, UMML 31.924; 1 male, 40 mm DML, Old and New Octopus Genera 107

Table 4. Measurements (mm) and counts for the lectotype of Muusoctopus januarii (Hoyle, 1885) in comparison with a male specimen at UMML identified as Benthoctopus januarii by Toll (1981) and a ZUMT specimen of ‘Octopus’ longispadiceus.

UMML ZUMT Lectotype 31.3199 000.3.31.1 Dorsal mantle length 49 46 50 Ventral mantle length 40 41 48 Mantle width 44 29 40 Mantle thickness 2.2 1.4 1.2 Head width 31 27 30 Head length 22 16 16 Neck width 28 19 29 Funnel length 20 14 29 Free funnel length 9 3 12 Mantle aperture 25 19 27 Length of free mantle edge 42 27 55 Mantle circumference 135 93 110 Length of arms 1 left/right (204) 227 D 202 225 D 2 219 201 D 207 219 204 3 (180) 123 D 109 174 186 4 161 159 D 143 179 161 Web depth sector A 42 26 43 B left/right 40 40 — 32 48 44 C 42 41 — 37 46 42 D 33 38 — 33 44 41 E382539 Ligula length 10.5 9.3 15 Calamus length 4.0 2.2 2.3 Arm diameter 8 6 9 Largest sucker 3.81 —102 Gill length 13 D 16 Length of posterior salivary gland 103 —103 No. of suckers on third right arm 83 914 1005 No. of gill lamellae (per demibranch) 8 — 9–11 D, damaged or truncated. (Arms with lengths in parentheses have lost their tip). 1Sucker #19 on arm R1. 2#21 on arm L2. 3Shape of posterior salivary gland: triangular, flattened. 4See Table 5. 5Suckers all normal: cf. Table 5, footnote 2.

Table 5. Details of male specimens recently examined at UMML, some of which Toll (1981) used to redescribe ‘Benthoctopus’ januarii.

UMML DML R3 Notes Reg. No. (mm) suckers1 31.442 60 71 Some suckers missing; in triple row near ligula. Spermatophoric groove mostly stripped away. 31.443 41 78 — 31.463 39 82 Condition fair to poor; ligula disintegrating. 31.923 50 912 Reproductive system badly damaged. Spermatophores approx. 84 mm long, 1.6 mm wide (cf. Toll, 1981). 31.923 42 81 Poor condition. Reproductive system badly damaged. 31.924 46 82 Very pale, head badly damaged. Not listed by Toll, 1981. 31.1749 40 — Specimen disintegrating. 31.3199 46 912 Not listed by Toll, 1981. Digestive system damaged.3 Note that two specimens from lot 31.923 are included. 1Number of suckers on normal part of hectocotylized third right arm. 2Abnormal specimens: sucker no. artificially high because smallest distal suckers close to ligula appear to be in three or four abnormally developed rows. (This is clearly a different phenomenon from the illusory triplet suckers seen in specimens of Tritaxeopus; see text, Section 3). 3A label with specimen states ‘Det. R. Toll’.

Oregon Sta. 1886, 400 shrimp trawl, 23.viii.1957, Caribbean Sea, midway between Honduras and Jamaica, 500 m, UMML 31.1749; 1 male, 46 mm DML, Oregon Sta. 4427, 400 shrimp trawl, 6.x.1963, Caribbean Sea, midway between Venezuela and Curac¸ao, 549 m, UMML 31.3199.

Type locality: Western South Atlantic Ocean, off Brazil, Barra Grande, Challenger Station 122, 950 S., 34500 W., 350 fm (640 m). 108 GLEADALL

Remarks. The type species of Muusoctopus was originally described by Hoyle as ‘Octopus januarii Steenstrup, MS.’ However, there is no record of any published description by Steenstrup, who apparently coined the name, so the author of the original published description is Hoyle (1885a). The name O. januarii was based on two specimens, one from the Atlantic Ocean off northeastern Brazil and the other from the Pacific, off Honshu, Japan. Robson (1932) removed the latter from the type series, identifying it as a new species, Benthoctopus profundorum, and used the Brazilian specimen alone to redescribe Benthoctopus januarii, thereby designating the lectotype (ICZN Art. 74.5). Toll (1981) provided a redescription of species januarii but apparently did so without inspecting the lectotype and without mentioning features such as the number of suckers on the hectocotylus. The description in the present paper includes these features, following examination of the lectotype, six of the specimens examined by Toll (1981) and two additional males (see Tables 4, 5). Many of the specimens examined in the present study are in poor condition, probably because of delay between death and fixation (due to the depths from which the specimens were taken) and accelerated conditions of decay in their tropical localities. The skin of the lectotype appears to have been mostly lost, with the remains forming small greyish patches. Also, many of the specimens have been damaged during dissection (see notes in Table 5). The internal organs of the lectotype, for example, are in very poor condition: pieces of the reproductive system and one of the posterior salivary glands were found free inside the specimen jar, the crop region of the digestive tract was found severed and the region of the anus has been mutilated. The remains of the reproductive system, the digestive system and the disintegrating digestive gland were therefore dissected free and placed in separate vials with the specimen. Despite the condition of the material examined, all these specimens are identified as conspecific, supporting the redescription of the species by Toll (1981). There is a large distance between the northeastern Brazilian type locality and the central American localities of Toll’s specimens but all the localities lie within a common faunal zone, the southern limit of which is a broad transition region off southern Brazil (see review by Palacio, 1977, for cephalopods of the central and southern Atlantic). Grimpe (1921) erected three new octopod genera: Bathypolypus (type species Octopus arcticus Prosch, 1849), Benthoctopus (type species Octopus piscatorum Verrill, 1879) and Atlantoctopus (type species Octopus lothei Chun, 1914). In a thorough review of species in the genus Bathypolypus by Muus (2002), the genus name Benthoctopus Grimpe was identified as a junior synonym of Bathypolypus because Muus identified O. piscatorum (the type species of Benthoctopus) as a junior synonym of Bathypolypus bairdii (Verrill, 1873). It was suggested that current usage of the genus Benthoctopus be retained by requesting the ICZN to allow redesignation of a different type species (Muus, 2002). This follows a similar plea by Voss and Pearcy (1990), who also had tentatively identified Octopus piscatorum as a species of Bathypolypus. However, this is a very different proposal from that mentioned above (see Section 2), where a new type species designation is clearly in the interest of stability. Until the work by Muus, Benthoctopus was used as a ‘hold all’ genus that has never been satisfactorily described, so retaining its usage as a non-Bathypolypus genus is unlikely to confer any stability since its member species are almost certainly polyphyletic. Although not mentioned by Muus (2002), the poorly described O. lothei (type species of Atlantoctopus; type specimen no longer extant) was identified by Robson (1932: 231) as a possible junior synonym of O. ergasticus Fischer and Fischer, 1892, which is another of the species identified by Muus as belonging to the genus Bathypolypus. Therefore, all three of Grimpe’s genera have now been synonymized under Bathypolypus (which is the senior synonym by position precedence). The genera Benthoctopus and Atlantoctopus are still available as subgenus names of Bathypolypus, should it be deemed necessary in the future to make distinctions between different species groups each containing Bathypolypus arcticus, B. bairdii and B. ergasticus. It is clear that the genus assignations of a number of species formerly identified as Benthoctopus require revision, so there is nothing to be gained from reassigning type species for one or more of the genera now synonymized under Bathypolypus. Following the work of Muus (particularly his paper of 2002), it is now clear that species in the genus Bathypolypus are very different from M. januarii, which had been suggested as an alternative type species for genus Benthoctopus (Voss and Pearcy, 1990; Muus, 2002). Species of Bathypolypus are smaller, have short arms, a massive ligula with prominent transverse ridges, an elongate posterior salivary gland, and (in five out of six species) only 26–50 suckers on the hectocotylus. The relationship between Muusoctopus januarii and other nominal species without an ink sac, previously identified as species of ‘Benthoctopus,’ must await thorough revision and redescriptions of other such octopodid species living in deeper waters that currently have not been identified as belonging to the genus Bathypolypus.

5. The Pseudophallus of Muusoctopus januarii: the Japanese Connection

During the study on Muusoctopus januarii, it was noticed that there are close morphological similarities between this species and ‘Octopus’ longispadiceus (Sasaki, 1917) (Table 4 and Fig. 6), more so than with other species of ‘Benthoctopus’ such as species yaquinae (Voss and Pearcy, 1990). Comparison of the internal structure of the pseudophallus of M. januarii and ‘O.’ longispadiceus reveals that they are strikingly similar (Figs. 7A and 7B). The pseudophallus is divided into anteromedial and posterolateral chambers separated from each other by an archway on Old and New Octopus Genera 109

Fig. 6. Photographs from the dorsal aspect of (A) Muusoctopus januarii (UMML 31.3199, DML 46 mm) and (B) ‘Octopus’ longispadiceus (ZUMT 000.3.31.1, DML 50 mm).

Fig. 7. The form of the pseudophallus in Muusoctopus januarii (A) in comparison with (B) ‘Octopus’ longispadiceus (Sasaki, 1917) (ZUMT 000.3.31.1, DML 50 mm); (C) ‘Octopus’ conispadiceus (Sasaki, 1917) (BMNH 20020534, DML 166 mm); (D) Enteroctopus dofleini (Wulker, 1910) (BMNH 20020532, DML 298 mm); (E) ‘Octopus’ hongkongensis Hoyle, 1885 (author’s reg. 1994.4.4, DML 153 mm); and (F) Bathypolypus bairdii (Verrill, 1873) (UMML 31.38A, DML 30 mm). All drawings are from the ventral aspect, anterior direction towards top of page. The pseudophallus of M. januarii is based on specimens UMML 31.443 and 31.463. Note that the pseudophallus of ‘Octopus’ hongkongensis is very thick and both its outer and inner walls have been delineated (whereas in the other diagrams, the wall of the pseudophallus is represented by a single line). Abbreviations: a, anteromedial chamber of pseudophallus; ap, aperture of spermatophoric duct; ar, dorsal arch; g, spermatophore guides; m, membranous extension of lateral spermatophore guide; p, posterolateral chamber of pseudophallus; t, transverse septum on dorsal wall. Scale bars 10 mm. the dorsal wall, and the spermatophoric duct opens into the anteromedial chamber through a triangular orifice anterior to this arch. It is interesting that M. januarii lacks an ink sac, while ‘O.’ longispadiceus has one, and even more interesting that the ZUMT specimen of ‘O.’ longispadiceus lacks anal cirri (cf. ‘anal valves minute’; Sasaki, 1929). While there is no doubt that they are different species, investigations are presently under way to acquire more 110 GLEADALL information about their phylogeny, and to provide a more detailed redescription of ‘Octopus’ longispadiceus (see also Takeda, 2003). A dorsal arch is present also in the pseudophallus of three other species found in cold temperate waters off Japan: ‘Octopus’ conispadiceus (Sasaki, 1917), Enteroctopus dofleini (Wulker, 1910) and ‘Octopus’ hongkongensis Hoyle, 1885 (see Figs. 7C, D, E, respectively). However, in these species there is only one chamber, with the spermatophoric duct opening into the pseudophallus via the dorsal arch. As Fig. 7E shows, the pseudophallus of ‘O.’ hongkongensis has an unusually thick muscular wall and the arch appears to be incompletely joined at its apex. In E. dofleini (Fig. 7D), there is a complex set of spermatophore guides, the most lateral of which expands into a substantial membrane, and there is a transverse septum extending from the dorsal wall just anterior to the dorsal arch: these are involved in guiding the very long spermatophore through the three 180 turns required to accommodate it within the pseudophallus to await ejection at copulation. The pseudophallus of Bathypolypus bairdii (Fig. 7F) is included for comparison, showing a relatively simple thin-walled structure, lacking a dorsal arch and apparently more similar to that of the Octopodinae. In B. bairdii, however, the pseudophallus is massive, occupying most of the left ventral region of the mantle cavity. The absence of an ink sac (as in M. januarii) is one criterion that has been used in the past as the basis for making distinctions at the subfamily level. However, it has long been suspected that the presence or absence of an ink sac is an inappropriate taxonomic character for distinguishing subfamilies of the Octopodidae (cf. previous classifications involving Bathypolypus and Muusoctopus (as Benthoctopus) and the Antarctic eledonids; Voight, 1993; Allcock and Piertney, 2002). This will be very clearly underlined if ‘Octopus’ longispadiceus is subsequently identified as a species of Muusoctopus. The observations reported here on the pseudophallus of M. januarii, ‘Octopus’ longispadiceus, ‘O.’ hongkongensis, ‘O.’ conispadiceus and Enteroctopus dofleini suggest that the presence of a dorsal arch in the pseudophallus may be a key taxonomic character to distinguish octopuses at the subfamily level. However, further investigation is required and it is desirable that, if this distinction is confirmed, other characters be found so that reliance for such an important distinction does not rest solely on the internal anatomy of the male reproductive system. Research is currently under way to redescribe ‘O.’ longispadiceus, ‘O.’ hongkongensis and ‘O.’ conispadiceus, and to designate appropriate genera, since they certainly are not members of the genus Octopus Cuvier. Acknowledgements I thank the following individuals for their helpful co-operation, discussions and support during this study: Ryoˆichi Arai (ZUMT), Somchai Bussarawit (PMBC), Cherdchinda Chotiyaputta (Department of Marine and Coastal Resources, Ministry of Natural Resources and Environment, Bangkok), Amelia MacLellan (BMNH), Peter Mordan (BMNH), Fred Naggs (BMNH), Anuwat Nateewathana (PMBC), Kazuo Sakamoto (ZUMT), Rei Ueshima (ZUMT), Vararin Vongpanich (PMBC), Nancy Voss (UMML) and Kathie Way (BMNH). I am also very grateful to Andrew Wakeham-Dawson (BMNH) for useful advice in approaching some of the taxonomic problems included in this paper, and to two anonymous referees for their incisive comments on, and meticulous suggestions for improvements to this paper. Many thanks also to Aki Chida for her expert help with the drawings and Ken Schmidt for help with image preparation and software. This work was supported in part by a grant from the Fujiwara Natural History Foundation, and by my research grant at Tohoku Bunka Gakuen University.

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