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Thaumeledone and other deep water octopodids from the Southern Ocean
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Deep-Sea Research II 51 (2004) 1883–1901 www.elsevier.com/locate/dsr2
Thaumeledone and other deep water octopodids from the Southern Ocean A.L. Allcocka,, M.A. Collinsb, U. Piatkowskic, M. Vecchioned a
School of Biology and Biochemistry, Marine Systems Research Group, Queen’s University Belfast, 97 Lisburn Road, Belfast BT9 7BL, UK b British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK c Leibniz Institut fu¨r Meereswissenschaflen, IFM-GEOMAR, Forschungsbereich Marine O¨kologie, Du¨sternbrooker Weg 20, D-24105 Kiel, Germany d NMFS Systematics Laboratory, National Museum of Natural History, Washington, DC 20560, USA Received 30 January 2004; received in revised form 7 June 2004; accepted 5 July 2004 Available online 26 October 2004
Abstract Recent trawling in the Southern Ocean has yielded an unusual and relatively large collection of deep-sea octopods, comprising four species in two genera. Several deep-sea genera, which are inadequately characterised, have been reported previously from the Southern Ocean. Within this paper, all the relevant historical type material has been examined and a full revision has been undertaken. Species previously considered to be representative of the genus Bentheledone have either been moved to Thaumeledone or are considered nomen dubium. A revised diagnosis of Thaumeledone is provided together with redescriptions of its Southern Ocean species as well as a description of a new species. A new genus has been erected to accommodate the remainder of the new specimens. r 2004 Elsevier Ltd. All rights reserved.
1. Introduction Recent trawling opportunities aboard R.V. Polarstern during The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) fish surveys, the Coastal Shelf Ecology of the Antarctic Sea Ice Zone (CS-EASIZ) and ANDEEP Corresponding author. Fax: +44-0-28-90975877.
E-mail addresses: [email protected] (A.L. Allcock), [email protected] (M.A. Collins), [email protected] (U. Piatkowski), [email protected] (M. Vecchione).
programmes, and during fisheries surveys around South Georgia have yielded an unusual collection of octopods initially identified as belonging to the deepsea genera Thaumeledone and Bentheledone. Both genera were erected by Robson (1930, 1932), but neither has ever been adequately characterised. Within the two genera, six names are available (Table 1), none of which are thought to be synonymous. Most deep-water Thaumeledone specimens from the Southern Ocean have been attributed to the species Thaumeledone brevis even though the type locality of the latter is off
0967-0645/$ - see front matter r 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.dsr2.2004.07.019
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Table 1 Species assigned to either Thaumeledone or Bentheledone prior to this publication and their revised generic placement Species and previous generic placement
Type locality
Status
Revised generic placement
T. T. T. T. B. B.
371S 531S 421S 441S 531S 641S
Type of Thaumeledone
Thaumeledone Thaumeledone Thaumeledone Thaumeledone Thaumeledone nomen dubium
brevis (Hoyle, 1885) gunteri (Robson, 1930) marshalli (O’Shea, 1999) zeiss (O’Shea, 1999) rotunda (Hoyle, 1885) albida (Berry, 1917)
541W 351W 1751E 1791E 1081E 1271E
Montevideo in the southwest Atlantic (approximately 371S). With the exception of O’Shea (1999), the genus has received little attention in recent years, possibly because the type material of some species is in extremely poor condition. Both the new material and the relevant type material have been thoroughly examined and a taxonomic revision of these deep-sea groups has been undertaken (for a summary, see Table 1).
2. Materials and methods Between 16 November and 26 December 1996, a benthic survey of the Antarctic Peninsula was undertaken by R.V. Polarstern (expedition ANTARKTIS XIV/2; Kattner, 1998). Although the majority of the cruise was dedicated to trawling in shallower depths as part of a CCAMLR fish survey, some deeper areas were also surveyed. As part of expedition ANTARKTIS XVII/3, run under the auspices of the CS-EASIZ programme (Arntz and Brey, 2001), R.V. Polarstern fished in the area of the Bransfield Straits and South Shetland Islands between 24 April and 7 May 2000. Specimens were collected predominantly with a commercial bottom trawl at depths down to 900 m. During the ANDEEP cruises (Fu¨tterer et al., 2003), several hauls were made in extremely deep water using an Agassiz trawl and specimens were obtained from depths of approximately 3000 m. In January 2003, a demersal fish survey was undertaken around South Georgia on the R.V. Dorada. The primary aim of the cruise was to investigate the distribution and ecology of the Patagonian toothfish and by-catch species caught in the fishery. The cruise yielded 25 specimens of
Type of Bentheledone
Thaumeledone gunteri (Collins et al., 2004). Several similar surveys in recent years around South Georgia have also yielded specimens of T. gunteri, a small proportion of which were fixed (see Yau et al., 2002). A full list of deep-sea specimens captured in the recent trawling activities described is given in the appendix. These data have been used to estimate the distributions and depth ranges of each species. Recently captured specimens were examined live where possible and when freshly dead. A number of specimens were preserved in 4% formalin and shipped to the UK where they have been deposited in the Natural History Museum, London (BMNH), the Smithsonian Institution (USNM) or the National Museums of Scotland, Edinburgh (NMSZ). Comparative material has been made available by the Australian Museum, Sydney (AMS), the Natural History Museum, London (formerly known as the British Museum (Natural History) abbreviated as BMNH), the National Museums and Galleries of Wales (NMWZ), the Santa Barbara Museum of Natural History (SBMNH) and the National Museums of Scotland (formerly known as the Royal Scottish Museum, now abbreviated as NMSZ). Other abbreviations and indices follow the guidelines for octopus taxonomy published by Roper and Voss (1983). Size descriptors (e.g., large, deep), where given alongside indices, follow the guidelines proposed at the taxonomy workshop at the Cephalopod International Advisory Council Symposium in Phuket, 2003.
3. Systematics A review of the historical type material showed that many of the type specimens were in excep-
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tionally poor condition. The worst of these was the holotype of Bentheledone albida (Berry, 1917), catalogue number AMS C040888, collected during the Mawson Antarctic Expedition, 14 January 1914, 641340 S 1271170 E, at approximately 3100 m. The mantle and the brachial crown are separate, and it is impossible to identify and number the arms. There is no sign of the funnel, which appears to have been lost in the separation of the mantle from the brachial crown. The web is torn at all sectors, and it is no longer possible to tell where it attached. All the tissue is extremely delicate, and it is difficult to spread out for examination for fear of tearing. It is not therefore possible to take morphometric measurements from the specimen. A pot of internal organs is enclosed with the specimen, but it is not possible to identify any useful characters. In some cases, it is not possible even to identify from which organs the material is derived. The ovary and one oviduct were identified. The oviducal gland is approximately 5 mm in diameter. The eggs in the ovary are of multiple sizes, the largest being 6 mm 2 mm, the smallest being less than 1 mm in length. The beak is in relatively good condition although the wings are curled. The radula is missing. Although some measurements were recorded by Berry (1917) (Table 5) and an illustration of the radula provided, we consider that it would be almost impossible to assign any recent specimens to this species with confidence and we therefore consider this species a nomen dubium. Although some of the other type material is also in poor condition, it has at least been possible to recognise characters and take enough useful measurements from the specimens to provide a brief diagnosis of each primary type. These are therefore dealt with under the relevant taxonomic section below.
Family: Octopodidae Orbigny, 1840. Thaumeledone Robson; 1932: Amended diagnosis Benthic octopodids, ML to 60 mm, TL to 190 mm; arms short to moderate (ALI 100–230),
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with small (ASI 3–8) uniserial suckers, third right arm of males hectocotylised with end of arm clearly differentiated into ligula and calamus, ligula large (LLI 9–17) and club shaped, ligula groove very deep and concave without transverse ridges, calamus long to very long (CaLI 30–85), arm tips not otherwise modified; web depth moderate to very deep (WDI 20–65); funnel organ VV- or W-shaped; gills poorly developed, with 4–6 lamellae per demibranch; ink sac absent; anal flaps absent; cartilaginous stylets present; diverticulum of penis coiled, spermatophores long (SpLI 120–135) and slender, mature males with 3–5 spermatophores; beak classically shaped; pigmentation deep purple, either on the papillae (T. gunteri) or on the oral surface of the web (other species). Type species: Eledone brevis Hoyle, 1885. By original designation. Included species: Eledone brevis Hoyle, 1885, T. gunteri Robson, 1930, Eledone rotunda Hoyle, 1885, Thaumeledone marshalli O’Shea, 1999, Thaumeledone peninsulae, sp. nov. Status of Thaumeledone zeiss O’Shea, 1999 is unclear. Remarks: Although the characters listed above are consistent, there is some variability in morphology in this genus. The posterior salivary glands of T. rotunda are most unusual (see later) and smaller than in other species (approximately 30% of the length of the buccal mass cf. 70–80% length of buccal mass). Of the described species of Thaumeledone, this species inhabits the deepest waters. Voss (1988) suggests that prey items in the deep sea are often small and soft bodied, and that a reduction in salivary gland size might be expected to reflect this. A certain amount of variation in radula morphology is also seen in this genus. There are 9 elements in T. zeiss, 5 elements in T. gunteri, T rotunda and T. marshalli, and a single element (rachidian) in T. brevis, as drawn by Robson (1932). It is possible that the radula of T. brevis in fact does have rows in addition to the rachidian as these would have been very hard to detect given the size of the type material, and the fact that electron microscopy was not available to Robson. Unfortunately, the radulae of the type material appear to be lost. Since the species is known only
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from the type material, it is not possible to verify the radula morphology without additional sampling in the type locality. Why T. zeiss should have 9 radular elements is unclear. Since this is the nondegenerate condition in the Coleoidea, the implication is either that T. zeiss is ancestral, that the radula varies according to some other factor and is not a systematically useful character, or that T. zeiss does not belong in Thaumeledone. O’Shea (1999, p. 255) indicates that his new species of Thaumeledone may not, in fact, be congeneric. However, similar levels of variation in radula morphology have been seen in a related deep-sea genus, Graneledone, even within a single species (Voss, 1976; Voss and Pearcy; 1990, Allcock et al., 2003) and remain unexplained although the genus and species therein are generally considered to be valid. Reduction or loss of radula teeth within the Octopoda is not uncommon (Nixon, 1998).
T. brevis (Hoyle, 1885) (Table 2) E. brevis Hoyle, 1885: 230. T. brevis (Hoyle, 1885).—Robson, 1932: 315. Material examined Lectotype and paralectotype: BMNH 1889.4.24.50-51, Stn 320, off Montevideo, 14 February 1876, 371170 S 531520 W, 600 fathoms [approx. 1100 m], green sand, Challenger Expedition. ~~. Paralectotype: NMWZ 78.14.424, Stn 320, off Montevideo,14 February 1876, 371170 S 531520 W, 600 fathoms [approx. 1100 m], green sand, Challenger Expedition. ~. Diagnosis of lectotype BMNH 1889.4.24.50 Specimen in poor condition and heavily dissected. Internal organ fragments in separate jar. Beaks and radula missing. Specimen small (dorsal mantle length 26 mm, total length 45 mm), female and immature. Arms approximately 3 mm wide and 26 mm long, 4th arms slightly shorter (22 mm). Suckers small (1 mm diameter). Approximately 24 suckers on 4th arms, 27 suckers on other arms.
Gills 3.5 mm in length with 4–5 lamellae per demibranch. Eyes are enormous (approximately 8 mm diameter cf. head width of approximately 16 mm). Funnel organ VV-shaped. Lateral limb ‘broken’ from medial limb in each V. Evidence of dissection in area of stylets, but unclear whether stylets were found. Small papillae, approximately 1 mm apart, cover the dorsal mantle surface although skin is badly torn in places apparently obscuring these. There is some evidence of deep purple pigmentation on the oral surface of the web. Remarks: The lectotype is the largest of three original syntypes. The paralectotype held by the BMNH is a slightly smaller immature male in similar condition. Notable features include a comparable funnel organ with medial and lateral limbs of each V ‘broken’ in the same fashion. The hectocotylised arm bears 21 suckers. The paralectotype held by NMGW is an even smaller immature female (dorsal mantle length 10 mm, total length 34 mm). The funnel organ is damaged although the remaining V is comparable in form to that seen in the lectotype and other paralectotype. The specimen bears papillae on the dorsal surface and has two large supraocular papillae. The type material is in too poor a condition (and too immature) to facilitate a redescription of this species: for this, trawling in the area of the type locality is required. However, it is clear that recently collected specimens from the Southern Ocean do not correspond to this species. Characters that separate it from Southern Ocean specimens include the peculiar VV-shaped funnel organ, the two large, distinct, supraocular papillae and the general papillation of the dorsal surface.
T. rotunda (Hoyle, 1885) comb. nov. (Plates 1–2, Figures 1–2, 6A, 7A, 8A, Table 2) E. rotunda Hoyle, 1885: 35, Figs. 1 and 2. Bentheledone rotunda (Hoyle, 1885). —Robson, 1932: 317, Text-Figs. 73–75.
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Fig. 2. T. rotunda, NMSZ 2002038.001, #, 51 mm ML; ~, 49 mm ML. Reproductive systems. Abbreviations: ag, accessory gland; d, diverticulum; mg, mucilaginous gland; o, ovary, og, oviducal gland; sg, spermatophoric gland; ss, spermatophoric sac; t, testes; to, terminal organ; vd, vas deferens.
Other material: USNM 1020991, Stn 42/039, Southern Ocean, 25 November 1996, 601370 S 541560 W, 3213–3222 m, R.V. Polarstern, ANTARKTIS XIV/II. 2#. NMSZ 2002038.001, Stn 61/046-8, Southern Ocean, 2 February 2002, 601390 S 531580 W, 2896 m, R.V. Polarstern, ANDEEP I. 1#, 1~.
Fig. 1. T. rotunda, NMSZ 2002038.001, #, 51 mm ML. Digestive system. Abbreviations: a, anus; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
Material examined Lectotype (except beaks): BMNH 1890.1.24.6, Stn 157, Southern Ocean, 3 March 1874, 531550 S, 1081350 E, 1950 fathoms [approx. 3500 m], diatom ooze, Challenger Expedition. 1~. [‘Holotype’ of Robson (1932) taken as lectotype designation.] Lectotype (beaks only): NMWZ 78.14.432 [same data].
Diagnosis of lectotype BMNH 1890.1.24.6 Specimen in poor condition. Brachial crown separated from mantle. Parts of digestive tract and female reproductive system also separate but present (Plate 1). Other parts missing (e.g., posterior salivary glands). Mantle soft, like paper, easily torn. Beaks and radula missing. Evidence of prior dissection in area of stylets. Unclear whether stylets found. Total length approximately 150 mm, ventral mantle length (dorsal unmeasurable) 51 mm. All arms approximately 10 mm wide and 95 mm long with small (2 mm) close set suckers. No evidence of enlarged suckers. Sucker count approximately 45. Gill length 11 mm, gill count 4/4. Web deepest at sector A (45 mm). Oviducal gland approximately 8 mm in diameter. Mature ovarian eggs large (approximately 16 mm 7 mm2) with 11 plical folds. Approximately 20 eggs present. Most colour leached from specimen but evidence remains that oral surface was deep purple.
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Plate 1. T. rotunda holotype. BMNH 1890.1.24.6. Approx. 150 mm TL.
Diagnosis of species Animals reach a total length of 188 mm. Funnel organ W-shaped. Posterior salivary glands bilobed, approximately 30% of length of buccal mass. Pale external coloration with deep purple oral surface. Hectocotylised sucker count 23–27. Description of species [based on recent catches of T. rotunda, specimens detailed in Table 2] Animals small to medium sized (ML to 62 mm; TL to 188 mm; Plate 2). Mantle approximately round (MWI 96.6711.2), head narrower than mantle (HWI 71.9710.8). Funnel medium sized (FuLI 45.373.6), gently tapered; funnel organ Wshaped. Arms short (MAI 47.172.9). Arm lengths subequal, arm order usually 1=4.2.3 (ALI L1 207.1713.5; L2 201.0713.9; L3 189.4716.2; L4 207.0717.1). Suckers uniserial, small (ASI 5.870.5), without sucker enlargement. Third right
arm of males hectocotylised, shorter than opposite number (OAI 85.878.4). Hectocotylus club-like in appearance. Ligula large (LLI 14.171.9); ligula groove deeply concave, without transverse ridges (Plate 2). Calamus distinct and very large (CaLI 79.772.7). Hectocotylised arm with 23–27 suckers, opposite arm with up to 45 suckers. Web deep to very deep (WDI 49.375.7), web formula approximately A.B=C=D.E. Ink sac absent. Posterior salivary glands bilobed and extremely unusual in appearance, approximately equal to 30% of length of buccal mass. Digestive tract otherwise normal (Fig. 1). Gills with 4–5 lamellae per demibranch. The female specimen registered under NMSZ 2002038.001 appears to be fully mature from the development of the oviducal gland which is 8 mm in diameter. However the eggs are still relatively small (5 mm) and are clearly not mature (Plate 2, Fig. 2). In a developing ovary, the ovary wall is normally stretched around the eggs but, in this specimen, the ovary capsule is very tough and hangs loosely around the eggs. This implies the individual has already laid. The presence of 40 additional developing eggs, all of a similar size, suggests that this species may be an intermittent terminal spawner (Rocha et al., 2001). Penis diverticulum coiled. Reproductive systems otherwise unremarkable (Fig. 2). Spermatophores medium to long (SpLI 122.2712.5) and slender. Lower beak with blunt rostrum, perhaps due to wear; beaks otherwise unremarkable (Fig. 6A). Cartilaginous stylets present (Fig. 7A). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid (Fig. 8A). Newly captured specimens of T. rotunda are pale on the dorsal and ventral surfaces but have a deep purple coloration on the oral surfaces. After fixation, two faint protrusions are apparent above the eyes which can probably be raised as supraocular papillae in the live animal. Other scattered papillae are evident on the dorsal surface of the mantle, head and brachial crown. Type locality: Southern Ocean, 531550 S, 1081350 E, 1950 fathoms [approx. 3500 m], diatom ooze. Distribution: Probably circum-Antarctic in depths between 2900 and 3500 m.
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Table 2 Counts and measurements (mm) for T. brevis and T. rotunda Species Status Repository Catalogue number Sex Maturity
brevis Lectotype BMNH 1889.4.24.50 Female Juvenile
brevis Paralectotype NMGW 78.14.424 Female Juvenile
rotunda Holotype BMNH 1890.1.24.6 Female Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm Width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
45 16
34 10 12 12 13 5 4 4/4 vv
150
18 16 8 4 4/4 vv 7 15/ 11/ 10/ 10 26 25 24 — 22 — 27 1 3 — — 5/4 5/5 4/4 —
51
45
E20 E20 E20 — E20 — 25 1 3 — —
4/4 —
95 d 95 — 95 — 45 2 10 — — 4/ 4/ 11/ 16
rotunda
rotunda
rotunda
rotunda
NMSZ 2002038.001 Female Mature
NMSZ 2002038.001 Male Mature
USNM 1020991 Male Mature
USNM 1020991 Male Mature
153 49 45 53 40 20 13 18/18 W 59 50/50 50/52 48/49 39 101 99 98 — 109 — 41 3 10 — — 5/5 5/5 10/11 5
145 51 41 47 42 23 18 16/16 W 46 40/45 30/40 33/36 35 103 97 94 85 98 27 43 3 10 14 11 5/5 5/5 10/10 63
[172] [59] [49] [51] [36] [25] [19] —/— — [60]
[188] [62] [53] [55] [35] [28] [22] —/— — [55] [55]/
[111] [113] [103] [84] [116] [23] [39] [3] [14] [10] [8] [5]/ [4]/
[125] [127] [114] [87] [129] [24] [42] [4] [13] [12] [10] [5]/ [4]/
[79]
l/r, left/right; m/l, medial/lateral;—not measurable, e.g., male character on female specimen, or funnel organ on unfixed material. Brackets indicate measurements made on fresh tissue.
Remarks: The lectotype of rotunda was captured the other side of Antarctica to our new specimens and in slightly lower latitudes (although still just south of the Polar Frontal Zone). Although the lectotype is in poor condition (Plate 1), there is good resemblance between this and our new specimens, particularly in size at maturity, arm shape (and length and width), sucker count, and gill count and size. Unfortunately several features that would probably be diagnostic (such as the funnel organ and unusual form of the posterior salivary glands) are missing from the lectototype. Nonetheless, male
characteristics, such as the shape of the copulatory organ of the hectocotylus, the relative size of the calamus, confirm that the new specimens undoubtedly belong in the genus Thaumeledone. They do not correspond to any other described species in the Southern Ocean and the W-shaped funnel organ of our specimens confirms that they are not synonymous with T. brevis. It is well known that deep-sea species can have extensive ranges and we are therefore confident in our assertion that our new specimens correspond to Hoyle’s rotunda. The one issue that casts any doubt on our conclusion is
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Southern Ocean, 21 January 1927, 531480 3000 S 351570 0000 W, 401–411 m, Discovery Expedition. 1~. Other material: NMSZ 1999275.010-011 Event 137, Stn 80, South Georgia, Southern Ocean, September 27th 1997, 541010 S 391330 W, 386–413 m, South Georgia Fish Survey 1997. 2#. SBMNH South Georgia, Southern Ocean, March 21st 1972, R.V. John Biscoe. 1~. NMSZ 2004083.009 Event 7, South Georgia, Southern Ocean, January 13th 2003, 531250 S 371010 W, 858–964 m, F.P.V. Dorada. 1~, 1#. NMSZ 2004083.010 Event 60, South Georgia, Southern Ocean, January 30th 2003, 541520 S 341220 W, 850–900 m, F.P.V. Dorada. 1~. NMSZ 2004083.007 Event 63, South Georgia, Southern Ocean, January 31st 2003, 531480 S 351510 W, 679–719 m, F.P.V. Dorada. 1#.
Plate 2. T. rotunda. NMSZ 2002038.001. Male, 51 mm ML, 145 mm TL. Inset A. shows enlargement of copulatory organ. Ligula length 14 mm, calamus length 11 mm. Inset B. shows ovary of female, 49 mm ML, 153 mm TL with immature eggs. Egg length 5 mm.
Robson’s (1932) illustration of the radula of the lectotype, which shows 9 elements. However, this radula is now lost and we have no way of checking the accuracy of Robson’s drawing. The removal of rotunda to Thaumeledone normally would have consequences for the genus Bentheledone since rotunda was the type species of this genus. However, with the conclusion that albida is a nomen dubium, no valid species actually remain in Bentheledone. T. gunteri Robson, 1930 (Plates 3-4, Figures 3, 6b, 7b, 8b, Table 3) Material examined Holotype: BMNH 1951.4.26.50 [previously BMNH 1931.1.21.5a], Stn 158, South Georgia,
Diagnosis of holotype BMNH 1951.4.26.50 The holotype is a mature female (dorsal mantle length 35 mm, total length 80 mm) which is well preserved but which has been extensively dissected. Arms are approximately equal in length (approximately 50 mm each) and width (8 mm). Suckers are small (2 mm) with approximately 35 suckers on each arm. All sectors of the web are of similar depth (26–28 mm). The funnel is partly severed from the specimen; however the funnel organ appears to be VV-shaped with the lateral limb considerably shorter than the medial limb (6 mm cf. 11 mm). Gills are 9 mm in length and have 5 lamellae per demibranch. A small pot labelled ‘‘repro tract and stylets’’ is present but there are no stylets therein. The posterior salivary glands measure 11 mm. The specimen retains no colour but there are large, irregularly shaped papillae, which are raised approximately 1 mm from the skin surface, on all surfaces, including the oral surface. The beaks are present (in a separate jar) but the radula is missing. Diagnosis of species Animals reach a total length of 107 mm. Funnel organ VV-shaped. Posterior salivary glands approximately 80% of length of buccal mass. External coloration pale, almost white, with large irregularly shaped deep purple papillae. The oral surface of the web is pale. Hectocotylised sucker count 19–22.
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Description [based on recent catches of T. gunteri, specimens detailed in Table 3] Animals small to medium sized (ML to 50 mm; TL to 107 mm; Plate 4). Mantle ovoid to round (MWI 91.679.6), head narrower than mantle (HWI 74.575.2). Funnel medium sized (FuLI 42.773.6), gently tapered; funnel organ VVshaped. Arms short but with considerable variation (MAI 61.6712.5). Arm lengths approximately equal (ALI L1 153.2728.9; L2 162.3740.1; L3 161.5732.9; L4 168.5736.1). Suckers uniserial, small (ASI 6.471.4), without sucker enlargement. Third right arm of males hectocotylised, shorter than opposite number (OAI 83.676.1). Hectocotylus club-like in appearance. Ligula large (LLI 16.974.4); ligula groove deeply concave, without transverse ridges (Plate 4). Calamus distinct and very large (CaLI 80.377.4). Hectocotylised arm with 19–22 suckers, opposite arm with up to 36 suckers. Web deep to very deep (WDI 40.977.7), web formula approximately A=B=C=D.E. Ink sac absent. Posterior salivary glands approximately 80% of length of buccal mass. Digestive tract otherwise normal, except for a loop in the rectum (Fig. 3). Gills with 5 lamellae per demibranch. Mature ovarian eggs large (410 mm). Penis diverticulum coiled. Spermatophores medium to long (SpLI 113.3727.2) and slender. Reproductive system unremarkable and hence not illustrated. Beaks unremarkable (Fig. 6B). Cartilaginous stylets present (Fig. 7B). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and almost acuspid (Fig. 8B). Newly captured specimens of T. gunteri are pale, almost white, with large irregularly shaped papillae on the dorsal and ventral surfaces that have deep purple pigmentation. There are several enlarged supraocular papillae. The oral surface of the web is pale. Type locality: South Georgia, Southern Ocean, 531480 3000 S 351570 0000 W, 401–411 m. Distribution: Probably restricted to South Georgia and Shag Rocks in depths of 364–964 m and probably deeper, although lack of deep fishing hampers this knowledge. Remarks: T. gunteri is easily identified by the purple pigmentation of the papillae on the back-
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ground of a pale mantle. It occurs only around South Georgia and is the only species of Thaumeledone to occur in this region. The type specimen is in good condition, and although the coloration is lost, the distinctive papillae are clearly visible (Plates 3 and 4).
T. peninsulae sp. nov. (Plate 5, Figures 4, 6C, 7C, 8C, Table 4) Material examined Holotype: NMSZ 2000081.057, Stn 178-1, Antarctic Peninsula, Southern Ocean, 2 May 2000, 611590 S 601190 W, 804–930 m, R.V. Polarstern, ANTARKTIS XVII/3. 1#. Paratypes: NMSZ 2000081.052, Stn 178-1, Antarctic Peninsula, Southern Ocean, 2 May 2000, 611590 S 601190 W, 804–930 m, R.V. Polarstern, ANTARKTIS XVII/3. 6~, 2#. Diagnosis Animals reach a total length of 120 mm. Funnel organ VV-shaped. Posterior salivary glands approximately 80% of length of buccal mass. External coloration pale purple, with large irregularly shaped papillae of a similar hue. The oral surface of the web is deep purple. Hectocotylised sucker count 22–25. Description [based on specimens detailed in Table 4] Animals small to medium sized (ML to 48 mm; TL to 120 mm; Plate 5). Mantle approximately round (MWI 96.8713.3), head narrower than mantle (HWI 72.879.1). Funnel small (FuLI 34.975.1), gently tapered; funnel organ VVshaped. Arms short (MAI 72.179.2). Arm lengths approximately equal (ALI L1 136.1717.4; L2 129.6718.4; L3 129.1717.0; L4 139.4716.7). Suckers uniserial, small (ASI 5.670.9), without sucker enlargement. Third right arm of males hectocotylised, slightly shorter than opposite number (OAI 93.771.2). Hectocotylus club-like in appearance. Ligula moderate to large (LLI 9.872.4); ligula groove very deep, without transverse ridges (Plate 5). Calamus distinct and very
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Table 3 Counts and measurements (mm) for T. gunteri
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
80 35 32 40 31 13 11/6 vv 26 25/26 26/28 25/28 25 50 48 51 — 49 — 34 2 8 — — 5/5 5/5 6
gunteri
gunteri
gunteri
gunteri
gunteri
gunteri
NMSZ 2004083.007 Male Mature
NMSZ 2004083.009 Female Submature
NMSZ 2004083.009 Male Mature
NMSZ 2004083.010 Female Spent
NMSZ 1999275 Male Mature
NMSZ SBMNH NMSZ 1999275 2004083.006 Male Female Mature Mature Juvenile
101 38 30 33 30 17 10 6/6 vv 26 21/24 24/22 23/25 20 59 55 55 49 55 19 32 3 8 11 9 5/5 5/5 6/6 50
80 34 29 33 26 15 8 7/7 vv 22 22/24 22/22 21/21 17
92 38 30 34 25 14 8 7/7 vv 20 20/19 20/19 16/19 14 46 48 57 44
107 35 29 37 28 14 10 8/7 vv 31 29/30 30/30 31/30 26 64 65 61 — 69 — 33 3 7 — — 5/5 5/5 7/6 3
44 40 36 34 21 11
50 44 40 35 22 13
25 35/30 29/22+ 28/ 27
30 32/29 30/35 27/32 18 90 91 88 78 95 21 36 3 8 11 9 5/ 5/ 13/ 41
43 42 — 43 — 32 1.5 6 — — 5/5 5/5 7/8 —
19 30 2 6 8 7 5/5 5/5 7/7 48
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen.
103 98 78 94 22 34 3 8 10 7 5/5 5/5 9/
gunteri
41 31 41 30 17 6 7/8 vv 16 19/18 20/20 24/24 21 52 56 57 — 57 — 33 2.5 7 — — 5/5 5/5 7/8
gunteri
gunteri
gunteri
NMSZ 2004083.006 Male Immature
NMSZ 2004083.006 Female Immature
37 16 13 13 15 8 6 4/4 vv E9 E9/9 E9/9 E9/9 E9 17 18 20 — 20 —
62 25 21 24 23 10 6 5/5 vv E12 E12/12 E12/12 E12/12 E10 30 32 35 30 36 20
56 25 21 20 20 11 6 6/6 vv 12 12/11 13/10 13/11 8 31
1 3 — — 5 5 4 —
2 4 4 3 5 5 5 —
1.5 4 — — 5 5 5 —
5 4
5 5 5
29 — 31 —
5 5 5
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Species Status Repository Catalogue number Sex Maturity
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tive system unremarkable and hence not illustrated. Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid. Beaks unremarkable (Fig. 6C). Cartilaginous stylets present (Fig. 7C). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid (Fig. 8C). Newly captured specimens of T. peninsulae are pale purple, with large irregularly shaped papillae on the dorsal and ventral surfaces. Unlike the papillae of T. gunteri, they are not differently coloured than the background skin. Supraocular papillae are not apparent. The oral surface of the web is deep purple. Type locality: Antarctic Peninsula, Southern Ocean, 611590 S 601190 W, 804–930 m. Distribution: Antarctic Peninsula, Southern Ocean, in depths from 377 to 1512 m. Etymology: Named after the type locality—the Antarctic Peninsula. Remarks: This species is clearly closely related to T. gunteri. It shares the large irregular papillae that cover the body surface, has similar arm and web formulae and a similar funnel organ. Both species have a loop in the rectum. Nevertheless, we feel that its distinctness in colour patterning, and in the relative size of the ligula and calamus warrants specific status. Praealtus gen. nov. Fig. 3. T. gunteri, NMSZ 2004083.009, #, 38 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
large (CaLI 90.378.7). Hectocotylised arm with 22–25 suckers, opposite arm with up to 35 suckers. Web deep to very deep (WDI 50.072.7), web formula approximately A=B=C=D.E. Ink sac absent. Posterior salivary glands approximately 80% of length of buccal mass. Digestive tract otherwise normal, except for a loop in the rectum (Fig. 4). Gills with 4–6 lamellae per demibranch. Mature ovarian eggs large (410 mm). Penis diverticulum coiled. Spermatophores medium to long (SpLI 110.1717.5) and slender. Reproduc-
Diagnosis Benthic octopodids, ML to 82 mm, TL to 380 mm; arms medium length (ALI 250–340), with small (ASI 4–8) uniserial suckers, third right arm of males hectocotylised with end of arm clearly differentiated into ligula and calamus, ligula small to medium (LLI 4–5), ligula groove round and shallow without transverse ridges, calamus very long (CaLI 50–70), arm tips not otherwise modified; web depth deep (WDI approximately 30); funnel organ VV-shaped; gills with 7–8 lamellae per demibranch; ink sac absent; anal flaps absent; posterior salivary glands tiny; stylets absent; diverticulum of penis coiled, spermatophores long (SpLI 185); beak classically shaped; radula with 7 elements, all unicuspid.
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Plate 3. T. gunteri holotype. BMNH 1951.4.26.50. 35 mm ML, approx. 80 mm TL.
Remarks: This genus is clearly differentiated from Thaumeledone by a range of characters. The arms are longer relative to mantle length (ALI 250–340 cf. 100–230 in Thaumeledone). The ligula is shorter relative to the length of the hectocotylised arm (LLI 4–5 cf. 9–17 in Thaumeledone) and the ligula groove is shallower, although this is hard to quantify numerically. There are more gill lamellae per demibranch (7–8 cf. 4–6 in Thaumeledone). The spermatophores are longer (SpLI 185 cf. 120–135 in Thaumeledone). Although there is extensive variation in the size of the posterior salivary glands in Thaumeledone, they are never tiny as they are in Praealtus. Finally the shape of the 2nd laterals in the radula is very different in Praealtus (Fig. 8). The absence of stylets in Praealtus also appears to differentiate it from Thaumeledone although it is, as yet, unclear whether stylets are present or absent in T. brevis.
Plate 4. T. gunteri. NMSZ 2004083.007. 38 mm ML, 101 mm TL. Inset shows enlargement of copulatory organ. Ligula length 11 mm, calamus length 9 mm.
P. paralbida sp. nov. (Plate 6, Figures 5, 6D, 8D, Table 5)
Type species: Praealtus paralbida sp. nov. Included species: P. paralbida sp. nov. Etymology: From the Latin, Praealtus, meaning very deep, referring to the depths at which specimens of this genus were captured.
Material examined Holotype: NMSZ 2002038.002, Stn 61/046-8, Southern Ocean, 2 February 2002, 601390 S 531580 W, 2896 m, R.V. Polarstern, ANDEEP I. 1#.
Table 4 Counts and measurements (mm) for T. peninsulae peninsulae Paratype NMSZ 2000081.052 Male Mature
peninsulae Paratype NMSZ 2000081.052 Male Mature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Mature
peninsulae Paratype NMSZ 2000081.052 Female Mature
peninsulae Paratype NMSZ 2000081.052 Female Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
104 37 30 39 29 15 7 9/9 vv 30 28/28 28/30 26/30 18 57 57 55 51 54 25 35 2.5 9 6 5 5/5 5/5 8/8 45
99 40 32 33 29 14 8 9/8 vv 25 23/25 23/25 22/25 20 49 44 45 42 48 24 34 2.5 7 3 3 6/5 5/6 6/6 36
88 32 25 31 24 13 8 7/7 vv 22 22/22 22/22 20/22 18 42 42 40 38 43 22 33 2 5 4 3.5 5/5 5/5 7/8 38
68 36 16 27 20 10 6 6/5 vv 18 18/18 18/18 18/18 17 36 36 37 — 40 — 26 1.5 6 — — 5/5 5/5 6/6 —
75 32 22 30 26 11 10 8/8 vv 23 23/22 23/22 22/22 18 43 43 42 — 44 — 34 2 7 — — 5/5 5/5 7/7 —
77 28 23 32 24 11 8 6/5 vv 21 20/23 21/22 21/22 17 42 38 42 — 46 — 33 1.5 7 — — 5/4 5/4 7/7 —
120 48 36 47 31 14 10 11/11 vv 32 33/34 32/31 32/32 24 68 54 56 — 70 — 32 2 8 — — 5/5 5/5 10/9
113 42 29 48 31 16 12 8/11 Vv 29 31/30 30/30 30/31 26 65 63 62 — 66 — 34 2.5 9 — — 5/5 5/5 10/10 13
112 44 33 40 30 13 9 6/8 vv 32 32/32 29/30 30/30 26 60 61 56 — 61 — 35 2.5 9 — — 5/5 5/5 9/8 9
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen.
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Plate 5. T. peninsulae holotype. NMSZ 2000081.057. 37 mm ML, 104 mm TL. Inset shows enlargement of copulatory organ. Ligula length 6 mm, calamus length 5 mm.
Pale external coloration with pale oral surface. Hectocotylised sucker count 38–42.
Fig. 4. T. peninsulae, NMSZ 2000081.057, #, 37 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
Other material: USNM 1021021, Stn 42/039, Southern Ocean, 25 November 1996, 601370 S 541560 W, 3213–3222 m, R.V. Polarstern, ANTARKTIS XIV/II. 5#, 1~. Comparative material: Moschites albida Berry, 1917. Holotype. AMS C40888, Stn 5, off Wilkes Land, Southern Ocean, 14 January 1914, 641340 S 1271170 E, 1700 fathoms [approx. 3100 m], Mawson Antarctic Expedition. 1~. Diagnosis Animals reach a total length of 380 mm. Funnel organ VV-shaped. Posterior salivary glands tiny.
Description [based on recently captured specimens detailed in Table 5] Animals medium sized (ML to 82 mm; TL to 380 mm; Plate 6). Mantle approximately round (MWI 104.374.0), head narrower than mantle (HWI 61.5723.1). Funnel medium sized (FuLI 43.473.8), gently tapered; funnel organ VVshaped. Arms medium length (MAI 32.672.5). Arm lengths subequal, arm order usually 1=2=3.4 (ALI L1 298.0722.6; L2 307.2723.8; L3 293.9728.6; L4 278.9729.5). Suckers uniserial, small (ASI 6.171.0), without sucker enlargement. Third right arm of males hectocotylised, usually slightly shorter than opposite number (OAI 93.4711.5). Ligula small to medium (LLI 4.770.5); ligula groove round and shallow, without transverse ridges (Plate 6). Calamus distinct and large to very large (CaLI 53.477.2). Hectocotylised arm with 38–42 suckers, opposite arm with up to 66 suckers. Web deep (WDI 29.170.9), damage prevents estimation of web formula. Ink
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Table 5 Counts and measurements (mm) for E. albida (taken from Berry, 1917) and P. paralbida Species Status Repository Catalogue number Sex Maturity
albida Holotype AM C4088 Female Mature
paralbida Holotype NMSZ 2002038.002 Male Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm Width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
170 35 40 31 12
37
27 129 124 111 — 117 — 57 (R3)
paralbida
paralbida
paralbida
paralbida
paralbida
paralbida
USNM 1021021 Female Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
320 65 62 72 52 26 21 14/12 vv 65
[248] [63] [54] [64] [41] [29] [16] —/— —
[258] [64] [60] [63] [41] [27] [20] —/— —
[266] [62] [58] [65] [43] [29] [19] —/— —
[233] [61] [55] [62] [38] [23] [14] —/— —
[273] [61] [57] [66] [44] [30] [25] —/— —
[55]/ 61/
[55]/ [53]/ [55]/
[53]/ [60]/ [53]/
[60]/
61/
[250] [66] [58] [68] [4] [28] [17] —/— — [55] [55]/ [55]/ [55]/
[60]/
[50]/ [60]/ [50]/
[186] [195] [189] [150] [175] [39] [55] [4] [14] [6] [3] [8]/ [7]/
[192] [189] [180] [160] [175] [38] [61] [3] [13] [8] [4] [8]/ [8]/
[201] [204] [203] [179] [192] [38] [62] [4] [13] [8] [4] [8]/ [7]/
[162] [166] [154] [139] [147] [40] [60] [3] [12] [6] [3] [8]/ [7]/
[196] [204] [203] [179] [200] [42] [66] [4] [15] [8] [4] [7]/ [7]/
—
—
—
—
—
46
— —
220 205 220 190 39 64 5 13 11 6
6
120
[186] [189] [181] — [171] — [59] [4] [13] — — 7/8 7/7 16/16 —
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen, or funnel organ on unfixed material. Brackets indicate measurements made on fresh tissue.
sac absent. Posterior salivary glands tiny. Digestive tract otherwise normal (Fig. 5). Gills with 7–8 lamellae per demibranch. Mature females have not been captured. Penis diverticulum coiled. Spermatophores long (SpLI 184.6) and slender. Reproductive system unremarkable and hence not illustrated. Beaks unremarkable (Figs. 6D and 7). Stylets absent. Radula with 7 elements, rachidian unicuspid, 3 slightly smaller unicuspid elements on either side of rachidian. Marginal plates absent (Fig. 8D).
Newly captured specimens of P. paralbida are pale slaty grey to very pale (almost white) violet all over. There are numerous small simple papillae scattered over the dorsal surface. Supraocular papillae are not apparent. Type locality: Off the Antarctic Peninsula, Southern Ocean, 601390 S 531580 W, 2896 m. Distribution: Possibly circum-Antarctic, in depths from 2896 to 3222 m. Etymology: Paralbida means ‘like albida’ recognising the similarity of this species to albida.
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Plate 6. P. paralbida holotype. NMSZ 2002038.002. 65 mm ML, 320 mm ML. Inset shows enlargement of copulatory organ. Ligula length 11 mm, calamus length 6 mm.
Remarks: The new material of this species was described by Vecchione et al. (in press) as Bentheledone cf. albida. At that time, there were no inconsistencies between our specimens and the little that is known of B. albida, although there was, and there remains, a great deal of uncertainty about the taxonomic characters of B. albida. Although there is only Berry’s (1917) drawing on which to rely, it appears that there may be some differences in the radula of the two species although they are similar in many respects. Berry (1917) illustrates the radula of B. albida as having a large unicuspid rachidian and very small, almost rounded first laterals. The second laterals are more cuspid and the marginal teeth are of a classic cone shape (see Nixon, 1998). Berry appears to illustrate the presence of marginal plates although he does not explicitly state this. The radula of P. paralbida, as seen by light microscopy, appears to lack marginal plates. Furthermore, the first and second laterals resemble marginal teeth in their appearance (Fig. 8D). As commented previously, high levels of variation in radulae of deep-sea species is
Fig. 5. P. paralbida, NMSZ 2002038.002, #, 65 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
not uncommon, however, it is impossible to conclude that B. albida and P. paralbida are synonymous despite their other similarities. The status of albida can only be established by collection of new material in the region of the type locality and we currently consider albida to be a nomen dubium. Should subsequent trawling show albida and paralbida to be conspecific, albida will take precedence and become the type species of Praealtus, thus maintaining taxonomic stability.
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Fig. 8. Radula of (A) T. rotunda, NMSZ 2002038.001, #, 51 mm ML; (B) T. gunteri, NMSZ 2004083.009, #, 38 mm ML; (C) T. peninsulae, NMSZ 2000081.057, #, 37 mm ML; (D) P. paralbida, NMSZ 2002038.002, #, 65 mm ML. Fig. 6. Lower and upper beaks of (A) T. rotunda, 2002038.001, #, 51 mm ML; (B) T. gunteri, 2004083.009, #, 38 mm ML; (C) T. peninsulae, 2000081.057, #, 37 mm ML; (D) P. paralbida, 2002038.002, #, 65 mm ML.
NMSZ NMSZ NMSZ NMSZ
Fig. 7. Cartilaginous stylets of (A) T. rotunda, NMSZ 2002038.001, #, 51 mm ML; (B) T. gunteri, NMSZ 2004083.009, #, 38 mm ML; (C) T. peninsulae, NMSZ 2000081.057, #, 37 mm ML.
Acknowledgements We would like to thank the organisers and leaders of all the cruises listed in the appendix, particularly Wolf Arntz, Karl Herman Koch and Angelika Brandt. The Alfred Wegener Institute kindly provided sea-time aboard R.V. Polarstern; the South Georgia Government, British Antarctic Survey and MRAG kindly facilitated collections during the South Georgia Fish Surveys. Curators at the Australian Museum, the National Museums and Galleries of Wales and the British Museum (Natural History) kindly provided access to specimens. Particular thanks to Amelia MacLellan. The early part of this work was undertaken at the National Museums of Scotland (NMS) with extensive support from Sankurie Pye. NMS’s help in financing cruise participation is much appreciated. Illustrations are by Ian Rendall. We are grateful to F.G. Hochberg for additional input and Vladimir Laptikhovsky and two anonymous referees for their helpful comments on the manuscript.
Appendix See Table A1.
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Table A1 A full list of deep-sea specimens captured in the recent trawling activities Cruise
Location
Depth
No. of specimens
Catalogue number
T. rotunda ANDEEP I ANTARKTIS XIV/2
601390 S 531580 W 601370 S 541560 W
2896 m 3213–3222 m
2 2
NMSZ 2002038.001 USNM 1020991
541010 S 541010 S 531250 S 531460 S 531470 S 531480 S 531530 S 541520 S 541530 S 541570 S 551210 S 551220 S
391330 W 391330 W 371010 W 401300 W 391190 W 311510 W 351470 W 341220 W 341220 W 341160 W 361040 W 361170 W
364–394 m 386–413 m 858–964 m 840–845 m 452–464 m 679–719 m 507–564 m 850–900 m 441–475 m 589–624 m 434–442 m 550–567 m
3 2 2 1 2 2 8 1 2 3 1 4
NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ
611190 S 611350 S 611380 S 611410 S 611590 S
571020 W 581450 W 581470 W 591100 W 601190 W
1440–1512 m 766–824 m 377–429 m 792–862 m 804–930 m
1 3 1 2 9
NMSZ 2000081.052/057
601390 S 531580 W 601370 S 541560 W
2896 m 3213–3222 m
1 6
NMSZ 2002038.002 USNM 1021021
T. gunteri South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia
Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries
T. peninsulae ANTARKTIS ANTARKTIS ANTARKTIS ANTARKTIS ANTARKTIS
XIV/2 XIV/2 XIV/2 XIV/2 XVII/3
P. paralbida ANDEEP I ANTARKTIS XIV/2
Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey
1994 1997 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003
1999275.010-011 2004083.009 2004083.008 2004083.003 2004083.007 2004083.004 2004083.010 2004083.002 2004083.006 2004083.001 2004083.005
These data have been used to estimate the distributions and depth ranges of each species.
References Allcock, A.L., Collins, M.A., Vecchione, M., 2003. A redescription of Graneledone verrucosa (Verrill, 1881) (Octopoda: Octopodidae). Journal of Molluscan Studies 69, 135–143. Arntz, W.E., Brey, T. (Eds.), 2001. The expedition ANTARKTIS XVII/3 (EASIZ III) of R.V. Polarstern in 2000. Berichte zur Polar- und Meeresforschung 402, pp. 1–181. Berry, S.S., 1917. Cephalopoda. Scientific reports. Australian Antarctic Expedition. Series C. Zoology and Botany 4 (2), pp. 1–38. Collins, M.A., Allcock, A.L., Belchier, M., 2004. Cephalopods of the South Georgia slope. Journal of the Marine Biological Association of the UK 84, 415–419. Fu¨tterer, D.K., Brandt, A., Poore, G.C.B. (Eds.), 2003. The Expeditions ANTARKTIS XIX/3–4 of the Research Vessel POLARSTERN in 2002. Berichte zur Polar- und Meeresforschung 470, pp. 1–174.
Hoyle, W.E., 1885. Diagnoses of new species of cephalopoda collected during the Cruise of HMS challenger. Part 1. The octopoda. Annals and Magazine of Natural History Series V 15, 222–236. Kattner, G. (Ed.), 1998. The expedition ANTARKTIS XIV/2 of R.V. Polarstern in 1996/7. Berichte zur Polarforschung 274, pp. 1–87. Nixon, M., 1998. The radulae of Cephalopoda. In: Voss, N.A., Vecchione, M., Toll, R.B., Sweeney, M.J. (Eds), Systematics and Biogeography of Cephalopods. vol 1. Smithsonian Contributions to Zoology 586, pp. 39–54. O’Shea, S., 1999. The marine fauna of New Zealand: octopoda (mollusca: cephalopoda). NIWA Biodiversity Memoir 112, 1–280. Robson, G.C., 1930. Cephalopoda. I. Octopoda. Discovery Reports 2, pp. 371–402. Robson, G.C., 1932. A Monograph of the Recent Cephalopoda Based on the Collections in the British Museum (Natural History). Part II. The Octopoda (Excluding the Octopodinae). British Museum, London.
ARTICLE IN PRESS A.L. Allcock et al. / Deep-Sea Research II 51 (2004) 1883–1901 Rocha, F., Guerra, A., Gonzalez, A.F., 2001. A review of reproductive strategies in cephalopods. Biological Reviews 76, 291–302. Roper, C.F.E., Voss, G.L., 1983. Guidelines for taxonomic descriptions of cephalopod species. Memoirs of the National Museum of Victoria 44, 13–27. Vecchione, M., Allcock, A.L., Piatkowski, U. Unusual incirrate octopods from the South Shetland Islands, Antarctica, including Bathypurpurata profunda, a newly discovered genus and species of deepwater pygmy octopod. Phuket Marine Biological Center Research Bulletin, in press. Voss, G.L., 1976. Two new species of octopods of the genus Graneledone (mollusca: cephalopoda) from the Southern
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Ocean. Proceedings of the Biological Society of Washington 88, 447–458. Voss, G.L., 1988. Evolution and phylogenetic relationships of deep-sea octopods (cirrata and incirrata). In: Clarke, M.R., Trueman, E.R. (Eds.)., The Mollusc. Volume 12. Paleontology and Neontology of Cephalopods. Academic Press, London, pp. 253–276. Voss, G.L., Pearcy, W.G., 1990. Deep-water octopods (mollusca: cephalopoda) of the northeastern pacific. Proceedings of the California Academy of Sciences 47, 47–94. Yau, C., Allcock, L., Daly, H.I., Collins, M.A., 2002. Distribution of Pareledone spp. (octopodidae: eledoninae) around South Georgia. Bulletin of Marine Science 71 (2), 993–1002.
Deep-Sea Research II 51 (2004) 1883–1901 www.elsevier.com/locate/dsr2
Thaumeledone and other deep water octopodids from the Southern Ocean A.L. Allcocka,, M.A. Collinsb, U. Piatkowskic, M. Vecchioned a
School of Biology and Biochemistry, Marine Systems Research Group, Queen’s University Belfast, 97 Lisburn Road, Belfast BT9 7BL, UK b British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK c Leibniz Institut fu¨r Meereswissenschaflen, IFM-GEOMAR, Forschungsbereich Marine O¨kologie, Du¨sternbrooker Weg 20, D-24105 Kiel, Germany d NMFS Systematics Laboratory, National Museum of Natural History, Washington, DC 20560, USA Received 30 January 2004; received in revised form 7 June 2004; accepted 5 July 2004 Available online 26 October 2004
Abstract Recent trawling in the Southern Ocean has yielded an unusual and relatively large collection of deep-sea octopods, comprising four species in two genera. Several deep-sea genera, which are inadequately characterised, have been reported previously from the Southern Ocean. Within this paper, all the relevant historical type material has been examined and a full revision has been undertaken. Species previously considered to be representative of the genus Bentheledone have either been moved to Thaumeledone or are considered nomen dubium. A revised diagnosis of Thaumeledone is provided together with redescriptions of its Southern Ocean species as well as a description of a new species. A new genus has been erected to accommodate the remainder of the new specimens. r 2004 Elsevier Ltd. All rights reserved.
1. Introduction Recent trawling opportunities aboard R.V. Polarstern during The Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) fish surveys, the Coastal Shelf Ecology of the Antarctic Sea Ice Zone (CS-EASIZ) and ANDEEP Corresponding author. Fax: +44-0-28-90975877.
E-mail addresses: [email protected] (A.L. Allcock), [email protected] (M.A. Collins), [email protected] (U. Piatkowski), [email protected] (M. Vecchione).
programmes, and during fisheries surveys around South Georgia have yielded an unusual collection of octopods initially identified as belonging to the deepsea genera Thaumeledone and Bentheledone. Both genera were erected by Robson (1930, 1932), but neither has ever been adequately characterised. Within the two genera, six names are available (Table 1), none of which are thought to be synonymous. Most deep-water Thaumeledone specimens from the Southern Ocean have been attributed to the species Thaumeledone brevis even though the type locality of the latter is off
0967-0645/$ - see front matter r 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.dsr2.2004.07.019
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Table 1 Species assigned to either Thaumeledone or Bentheledone prior to this publication and their revised generic placement Species and previous generic placement
Type locality
Status
Revised generic placement
T. T. T. T. B. B.
371S 531S 421S 441S 531S 641S
Type of Thaumeledone
Thaumeledone Thaumeledone Thaumeledone Thaumeledone Thaumeledone nomen dubium
brevis (Hoyle, 1885) gunteri (Robson, 1930) marshalli (O’Shea, 1999) zeiss (O’Shea, 1999) rotunda (Hoyle, 1885) albida (Berry, 1917)
541W 351W 1751E 1791E 1081E 1271E
Montevideo in the southwest Atlantic (approximately 371S). With the exception of O’Shea (1999), the genus has received little attention in recent years, possibly because the type material of some species is in extremely poor condition. Both the new material and the relevant type material have been thoroughly examined and a taxonomic revision of these deep-sea groups has been undertaken (for a summary, see Table 1).
2. Materials and methods Between 16 November and 26 December 1996, a benthic survey of the Antarctic Peninsula was undertaken by R.V. Polarstern (expedition ANTARKTIS XIV/2; Kattner, 1998). Although the majority of the cruise was dedicated to trawling in shallower depths as part of a CCAMLR fish survey, some deeper areas were also surveyed. As part of expedition ANTARKTIS XVII/3, run under the auspices of the CS-EASIZ programme (Arntz and Brey, 2001), R.V. Polarstern fished in the area of the Bransfield Straits and South Shetland Islands between 24 April and 7 May 2000. Specimens were collected predominantly with a commercial bottom trawl at depths down to 900 m. During the ANDEEP cruises (Fu¨tterer et al., 2003), several hauls were made in extremely deep water using an Agassiz trawl and specimens were obtained from depths of approximately 3000 m. In January 2003, a demersal fish survey was undertaken around South Georgia on the R.V. Dorada. The primary aim of the cruise was to investigate the distribution and ecology of the Patagonian toothfish and by-catch species caught in the fishery. The cruise yielded 25 specimens of
Type of Bentheledone
Thaumeledone gunteri (Collins et al., 2004). Several similar surveys in recent years around South Georgia have also yielded specimens of T. gunteri, a small proportion of which were fixed (see Yau et al., 2002). A full list of deep-sea specimens captured in the recent trawling activities described is given in the appendix. These data have been used to estimate the distributions and depth ranges of each species. Recently captured specimens were examined live where possible and when freshly dead. A number of specimens were preserved in 4% formalin and shipped to the UK where they have been deposited in the Natural History Museum, London (BMNH), the Smithsonian Institution (USNM) or the National Museums of Scotland, Edinburgh (NMSZ). Comparative material has been made available by the Australian Museum, Sydney (AMS), the Natural History Museum, London (formerly known as the British Museum (Natural History) abbreviated as BMNH), the National Museums and Galleries of Wales (NMWZ), the Santa Barbara Museum of Natural History (SBMNH) and the National Museums of Scotland (formerly known as the Royal Scottish Museum, now abbreviated as NMSZ). Other abbreviations and indices follow the guidelines for octopus taxonomy published by Roper and Voss (1983). Size descriptors (e.g., large, deep), where given alongside indices, follow the guidelines proposed at the taxonomy workshop at the Cephalopod International Advisory Council Symposium in Phuket, 2003.
3. Systematics A review of the historical type material showed that many of the type specimens were in excep-
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tionally poor condition. The worst of these was the holotype of Bentheledone albida (Berry, 1917), catalogue number AMS C040888, collected during the Mawson Antarctic Expedition, 14 January 1914, 641340 S 1271170 E, at approximately 3100 m. The mantle and the brachial crown are separate, and it is impossible to identify and number the arms. There is no sign of the funnel, which appears to have been lost in the separation of the mantle from the brachial crown. The web is torn at all sectors, and it is no longer possible to tell where it attached. All the tissue is extremely delicate, and it is difficult to spread out for examination for fear of tearing. It is not therefore possible to take morphometric measurements from the specimen. A pot of internal organs is enclosed with the specimen, but it is not possible to identify any useful characters. In some cases, it is not possible even to identify from which organs the material is derived. The ovary and one oviduct were identified. The oviducal gland is approximately 5 mm in diameter. The eggs in the ovary are of multiple sizes, the largest being 6 mm 2 mm, the smallest being less than 1 mm in length. The beak is in relatively good condition although the wings are curled. The radula is missing. Although some measurements were recorded by Berry (1917) (Table 5) and an illustration of the radula provided, we consider that it would be almost impossible to assign any recent specimens to this species with confidence and we therefore consider this species a nomen dubium. Although some of the other type material is also in poor condition, it has at least been possible to recognise characters and take enough useful measurements from the specimens to provide a brief diagnosis of each primary type. These are therefore dealt with under the relevant taxonomic section below.
Family: Octopodidae Orbigny, 1840. Thaumeledone Robson; 1932: Amended diagnosis Benthic octopodids, ML to 60 mm, TL to 190 mm; arms short to moderate (ALI 100–230),
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with small (ASI 3–8) uniserial suckers, third right arm of males hectocotylised with end of arm clearly differentiated into ligula and calamus, ligula large (LLI 9–17) and club shaped, ligula groove very deep and concave without transverse ridges, calamus long to very long (CaLI 30–85), arm tips not otherwise modified; web depth moderate to very deep (WDI 20–65); funnel organ VV- or W-shaped; gills poorly developed, with 4–6 lamellae per demibranch; ink sac absent; anal flaps absent; cartilaginous stylets present; diverticulum of penis coiled, spermatophores long (SpLI 120–135) and slender, mature males with 3–5 spermatophores; beak classically shaped; pigmentation deep purple, either on the papillae (T. gunteri) or on the oral surface of the web (other species). Type species: Eledone brevis Hoyle, 1885. By original designation. Included species: Eledone brevis Hoyle, 1885, T. gunteri Robson, 1930, Eledone rotunda Hoyle, 1885, Thaumeledone marshalli O’Shea, 1999, Thaumeledone peninsulae, sp. nov. Status of Thaumeledone zeiss O’Shea, 1999 is unclear. Remarks: Although the characters listed above are consistent, there is some variability in morphology in this genus. The posterior salivary glands of T. rotunda are most unusual (see later) and smaller than in other species (approximately 30% of the length of the buccal mass cf. 70–80% length of buccal mass). Of the described species of Thaumeledone, this species inhabits the deepest waters. Voss (1988) suggests that prey items in the deep sea are often small and soft bodied, and that a reduction in salivary gland size might be expected to reflect this. A certain amount of variation in radula morphology is also seen in this genus. There are 9 elements in T. zeiss, 5 elements in T. gunteri, T rotunda and T. marshalli, and a single element (rachidian) in T. brevis, as drawn by Robson (1932). It is possible that the radula of T. brevis in fact does have rows in addition to the rachidian as these would have been very hard to detect given the size of the type material, and the fact that electron microscopy was not available to Robson. Unfortunately, the radulae of the type material appear to be lost. Since the species is known only
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from the type material, it is not possible to verify the radula morphology without additional sampling in the type locality. Why T. zeiss should have 9 radular elements is unclear. Since this is the nondegenerate condition in the Coleoidea, the implication is either that T. zeiss is ancestral, that the radula varies according to some other factor and is not a systematically useful character, or that T. zeiss does not belong in Thaumeledone. O’Shea (1999, p. 255) indicates that his new species of Thaumeledone may not, in fact, be congeneric. However, similar levels of variation in radula morphology have been seen in a related deep-sea genus, Graneledone, even within a single species (Voss, 1976; Voss and Pearcy; 1990, Allcock et al., 2003) and remain unexplained although the genus and species therein are generally considered to be valid. Reduction or loss of radula teeth within the Octopoda is not uncommon (Nixon, 1998).
T. brevis (Hoyle, 1885) (Table 2) E. brevis Hoyle, 1885: 230. T. brevis (Hoyle, 1885).—Robson, 1932: 315. Material examined Lectotype and paralectotype: BMNH 1889.4.24.50-51, Stn 320, off Montevideo, 14 February 1876, 371170 S 531520 W, 600 fathoms [approx. 1100 m], green sand, Challenger Expedition. ~~. Paralectotype: NMWZ 78.14.424, Stn 320, off Montevideo,14 February 1876, 371170 S 531520 W, 600 fathoms [approx. 1100 m], green sand, Challenger Expedition. ~. Diagnosis of lectotype BMNH 1889.4.24.50 Specimen in poor condition and heavily dissected. Internal organ fragments in separate jar. Beaks and radula missing. Specimen small (dorsal mantle length 26 mm, total length 45 mm), female and immature. Arms approximately 3 mm wide and 26 mm long, 4th arms slightly shorter (22 mm). Suckers small (1 mm diameter). Approximately 24 suckers on 4th arms, 27 suckers on other arms.
Gills 3.5 mm in length with 4–5 lamellae per demibranch. Eyes are enormous (approximately 8 mm diameter cf. head width of approximately 16 mm). Funnel organ VV-shaped. Lateral limb ‘broken’ from medial limb in each V. Evidence of dissection in area of stylets, but unclear whether stylets were found. Small papillae, approximately 1 mm apart, cover the dorsal mantle surface although skin is badly torn in places apparently obscuring these. There is some evidence of deep purple pigmentation on the oral surface of the web. Remarks: The lectotype is the largest of three original syntypes. The paralectotype held by the BMNH is a slightly smaller immature male in similar condition. Notable features include a comparable funnel organ with medial and lateral limbs of each V ‘broken’ in the same fashion. The hectocotylised arm bears 21 suckers. The paralectotype held by NMGW is an even smaller immature female (dorsal mantle length 10 mm, total length 34 mm). The funnel organ is damaged although the remaining V is comparable in form to that seen in the lectotype and other paralectotype. The specimen bears papillae on the dorsal surface and has two large supraocular papillae. The type material is in too poor a condition (and too immature) to facilitate a redescription of this species: for this, trawling in the area of the type locality is required. However, it is clear that recently collected specimens from the Southern Ocean do not correspond to this species. Characters that separate it from Southern Ocean specimens include the peculiar VV-shaped funnel organ, the two large, distinct, supraocular papillae and the general papillation of the dorsal surface.
T. rotunda (Hoyle, 1885) comb. nov. (Plates 1–2, Figures 1–2, 6A, 7A, 8A, Table 2) E. rotunda Hoyle, 1885: 35, Figs. 1 and 2. Bentheledone rotunda (Hoyle, 1885). —Robson, 1932: 317, Text-Figs. 73–75.
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Fig. 2. T. rotunda, NMSZ 2002038.001, #, 51 mm ML; ~, 49 mm ML. Reproductive systems. Abbreviations: ag, accessory gland; d, diverticulum; mg, mucilaginous gland; o, ovary, og, oviducal gland; sg, spermatophoric gland; ss, spermatophoric sac; t, testes; to, terminal organ; vd, vas deferens.
Other material: USNM 1020991, Stn 42/039, Southern Ocean, 25 November 1996, 601370 S 541560 W, 3213–3222 m, R.V. Polarstern, ANTARKTIS XIV/II. 2#. NMSZ 2002038.001, Stn 61/046-8, Southern Ocean, 2 February 2002, 601390 S 531580 W, 2896 m, R.V. Polarstern, ANDEEP I. 1#, 1~.
Fig. 1. T. rotunda, NMSZ 2002038.001, #, 51 mm ML. Digestive system. Abbreviations: a, anus; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
Material examined Lectotype (except beaks): BMNH 1890.1.24.6, Stn 157, Southern Ocean, 3 March 1874, 531550 S, 1081350 E, 1950 fathoms [approx. 3500 m], diatom ooze, Challenger Expedition. 1~. [‘Holotype’ of Robson (1932) taken as lectotype designation.] Lectotype (beaks only): NMWZ 78.14.432 [same data].
Diagnosis of lectotype BMNH 1890.1.24.6 Specimen in poor condition. Brachial crown separated from mantle. Parts of digestive tract and female reproductive system also separate but present (Plate 1). Other parts missing (e.g., posterior salivary glands). Mantle soft, like paper, easily torn. Beaks and radula missing. Evidence of prior dissection in area of stylets. Unclear whether stylets found. Total length approximately 150 mm, ventral mantle length (dorsal unmeasurable) 51 mm. All arms approximately 10 mm wide and 95 mm long with small (2 mm) close set suckers. No evidence of enlarged suckers. Sucker count approximately 45. Gill length 11 mm, gill count 4/4. Web deepest at sector A (45 mm). Oviducal gland approximately 8 mm in diameter. Mature ovarian eggs large (approximately 16 mm 7 mm2) with 11 plical folds. Approximately 20 eggs present. Most colour leached from specimen but evidence remains that oral surface was deep purple.
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Plate 1. T. rotunda holotype. BMNH 1890.1.24.6. Approx. 150 mm TL.
Diagnosis of species Animals reach a total length of 188 mm. Funnel organ W-shaped. Posterior salivary glands bilobed, approximately 30% of length of buccal mass. Pale external coloration with deep purple oral surface. Hectocotylised sucker count 23–27. Description of species [based on recent catches of T. rotunda, specimens detailed in Table 2] Animals small to medium sized (ML to 62 mm; TL to 188 mm; Plate 2). Mantle approximately round (MWI 96.6711.2), head narrower than mantle (HWI 71.9710.8). Funnel medium sized (FuLI 45.373.6), gently tapered; funnel organ Wshaped. Arms short (MAI 47.172.9). Arm lengths subequal, arm order usually 1=4.2.3 (ALI L1 207.1713.5; L2 201.0713.9; L3 189.4716.2; L4 207.0717.1). Suckers uniserial, small (ASI 5.870.5), without sucker enlargement. Third right
arm of males hectocotylised, shorter than opposite number (OAI 85.878.4). Hectocotylus club-like in appearance. Ligula large (LLI 14.171.9); ligula groove deeply concave, without transverse ridges (Plate 2). Calamus distinct and very large (CaLI 79.772.7). Hectocotylised arm with 23–27 suckers, opposite arm with up to 45 suckers. Web deep to very deep (WDI 49.375.7), web formula approximately A.B=C=D.E. Ink sac absent. Posterior salivary glands bilobed and extremely unusual in appearance, approximately equal to 30% of length of buccal mass. Digestive tract otherwise normal (Fig. 1). Gills with 4–5 lamellae per demibranch. The female specimen registered under NMSZ 2002038.001 appears to be fully mature from the development of the oviducal gland which is 8 mm in diameter. However the eggs are still relatively small (5 mm) and are clearly not mature (Plate 2, Fig. 2). In a developing ovary, the ovary wall is normally stretched around the eggs but, in this specimen, the ovary capsule is very tough and hangs loosely around the eggs. This implies the individual has already laid. The presence of 40 additional developing eggs, all of a similar size, suggests that this species may be an intermittent terminal spawner (Rocha et al., 2001). Penis diverticulum coiled. Reproductive systems otherwise unremarkable (Fig. 2). Spermatophores medium to long (SpLI 122.2712.5) and slender. Lower beak with blunt rostrum, perhaps due to wear; beaks otherwise unremarkable (Fig. 6A). Cartilaginous stylets present (Fig. 7A). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid (Fig. 8A). Newly captured specimens of T. rotunda are pale on the dorsal and ventral surfaces but have a deep purple coloration on the oral surfaces. After fixation, two faint protrusions are apparent above the eyes which can probably be raised as supraocular papillae in the live animal. Other scattered papillae are evident on the dorsal surface of the mantle, head and brachial crown. Type locality: Southern Ocean, 531550 S, 1081350 E, 1950 fathoms [approx. 3500 m], diatom ooze. Distribution: Probably circum-Antarctic in depths between 2900 and 3500 m.
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Table 2 Counts and measurements (mm) for T. brevis and T. rotunda Species Status Repository Catalogue number Sex Maturity
brevis Lectotype BMNH 1889.4.24.50 Female Juvenile
brevis Paralectotype NMGW 78.14.424 Female Juvenile
rotunda Holotype BMNH 1890.1.24.6 Female Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm Width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
45 16
34 10 12 12 13 5 4 4/4 vv
150
18 16 8 4 4/4 vv 7 15/ 11/ 10/ 10 26 25 24 — 22 — 27 1 3 — — 5/4 5/5 4/4 —
51
45
E20 E20 E20 — E20 — 25 1 3 — —
4/4 —
95 d 95 — 95 — 45 2 10 — — 4/ 4/ 11/ 16
rotunda
rotunda
rotunda
rotunda
NMSZ 2002038.001 Female Mature
NMSZ 2002038.001 Male Mature
USNM 1020991 Male Mature
USNM 1020991 Male Mature
153 49 45 53 40 20 13 18/18 W 59 50/50 50/52 48/49 39 101 99 98 — 109 — 41 3 10 — — 5/5 5/5 10/11 5
145 51 41 47 42 23 18 16/16 W 46 40/45 30/40 33/36 35 103 97 94 85 98 27 43 3 10 14 11 5/5 5/5 10/10 63
[172] [59] [49] [51] [36] [25] [19] —/— — [60]
[188] [62] [53] [55] [35] [28] [22] —/— — [55] [55]/
[111] [113] [103] [84] [116] [23] [39] [3] [14] [10] [8] [5]/ [4]/
[125] [127] [114] [87] [129] [24] [42] [4] [13] [12] [10] [5]/ [4]/
[79]
l/r, left/right; m/l, medial/lateral;—not measurable, e.g., male character on female specimen, or funnel organ on unfixed material. Brackets indicate measurements made on fresh tissue.
Remarks: The lectotype of rotunda was captured the other side of Antarctica to our new specimens and in slightly lower latitudes (although still just south of the Polar Frontal Zone). Although the lectotype is in poor condition (Plate 1), there is good resemblance between this and our new specimens, particularly in size at maturity, arm shape (and length and width), sucker count, and gill count and size. Unfortunately several features that would probably be diagnostic (such as the funnel organ and unusual form of the posterior salivary glands) are missing from the lectototype. Nonetheless, male
characteristics, such as the shape of the copulatory organ of the hectocotylus, the relative size of the calamus, confirm that the new specimens undoubtedly belong in the genus Thaumeledone. They do not correspond to any other described species in the Southern Ocean and the W-shaped funnel organ of our specimens confirms that they are not synonymous with T. brevis. It is well known that deep-sea species can have extensive ranges and we are therefore confident in our assertion that our new specimens correspond to Hoyle’s rotunda. The one issue that casts any doubt on our conclusion is
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Southern Ocean, 21 January 1927, 531480 3000 S 351570 0000 W, 401–411 m, Discovery Expedition. 1~. Other material: NMSZ 1999275.010-011 Event 137, Stn 80, South Georgia, Southern Ocean, September 27th 1997, 541010 S 391330 W, 386–413 m, South Georgia Fish Survey 1997. 2#. SBMNH South Georgia, Southern Ocean, March 21st 1972, R.V. John Biscoe. 1~. NMSZ 2004083.009 Event 7, South Georgia, Southern Ocean, January 13th 2003, 531250 S 371010 W, 858–964 m, F.P.V. Dorada. 1~, 1#. NMSZ 2004083.010 Event 60, South Georgia, Southern Ocean, January 30th 2003, 541520 S 341220 W, 850–900 m, F.P.V. Dorada. 1~. NMSZ 2004083.007 Event 63, South Georgia, Southern Ocean, January 31st 2003, 531480 S 351510 W, 679–719 m, F.P.V. Dorada. 1#.
Plate 2. T. rotunda. NMSZ 2002038.001. Male, 51 mm ML, 145 mm TL. Inset A. shows enlargement of copulatory organ. Ligula length 14 mm, calamus length 11 mm. Inset B. shows ovary of female, 49 mm ML, 153 mm TL with immature eggs. Egg length 5 mm.
Robson’s (1932) illustration of the radula of the lectotype, which shows 9 elements. However, this radula is now lost and we have no way of checking the accuracy of Robson’s drawing. The removal of rotunda to Thaumeledone normally would have consequences for the genus Bentheledone since rotunda was the type species of this genus. However, with the conclusion that albida is a nomen dubium, no valid species actually remain in Bentheledone. T. gunteri Robson, 1930 (Plates 3-4, Figures 3, 6b, 7b, 8b, Table 3) Material examined Holotype: BMNH 1951.4.26.50 [previously BMNH 1931.1.21.5a], Stn 158, South Georgia,
Diagnosis of holotype BMNH 1951.4.26.50 The holotype is a mature female (dorsal mantle length 35 mm, total length 80 mm) which is well preserved but which has been extensively dissected. Arms are approximately equal in length (approximately 50 mm each) and width (8 mm). Suckers are small (2 mm) with approximately 35 suckers on each arm. All sectors of the web are of similar depth (26–28 mm). The funnel is partly severed from the specimen; however the funnel organ appears to be VV-shaped with the lateral limb considerably shorter than the medial limb (6 mm cf. 11 mm). Gills are 9 mm in length and have 5 lamellae per demibranch. A small pot labelled ‘‘repro tract and stylets’’ is present but there are no stylets therein. The posterior salivary glands measure 11 mm. The specimen retains no colour but there are large, irregularly shaped papillae, which are raised approximately 1 mm from the skin surface, on all surfaces, including the oral surface. The beaks are present (in a separate jar) but the radula is missing. Diagnosis of species Animals reach a total length of 107 mm. Funnel organ VV-shaped. Posterior salivary glands approximately 80% of length of buccal mass. External coloration pale, almost white, with large irregularly shaped deep purple papillae. The oral surface of the web is pale. Hectocotylised sucker count 19–22.
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Description [based on recent catches of T. gunteri, specimens detailed in Table 3] Animals small to medium sized (ML to 50 mm; TL to 107 mm; Plate 4). Mantle ovoid to round (MWI 91.679.6), head narrower than mantle (HWI 74.575.2). Funnel medium sized (FuLI 42.773.6), gently tapered; funnel organ VVshaped. Arms short but with considerable variation (MAI 61.6712.5). Arm lengths approximately equal (ALI L1 153.2728.9; L2 162.3740.1; L3 161.5732.9; L4 168.5736.1). Suckers uniserial, small (ASI 6.471.4), without sucker enlargement. Third right arm of males hectocotylised, shorter than opposite number (OAI 83.676.1). Hectocotylus club-like in appearance. Ligula large (LLI 16.974.4); ligula groove deeply concave, without transverse ridges (Plate 4). Calamus distinct and very large (CaLI 80.377.4). Hectocotylised arm with 19–22 suckers, opposite arm with up to 36 suckers. Web deep to very deep (WDI 40.977.7), web formula approximately A=B=C=D.E. Ink sac absent. Posterior salivary glands approximately 80% of length of buccal mass. Digestive tract otherwise normal, except for a loop in the rectum (Fig. 3). Gills with 5 lamellae per demibranch. Mature ovarian eggs large (410 mm). Penis diverticulum coiled. Spermatophores medium to long (SpLI 113.3727.2) and slender. Reproductive system unremarkable and hence not illustrated. Beaks unremarkable (Fig. 6B). Cartilaginous stylets present (Fig. 7B). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and almost acuspid (Fig. 8B). Newly captured specimens of T. gunteri are pale, almost white, with large irregularly shaped papillae on the dorsal and ventral surfaces that have deep purple pigmentation. There are several enlarged supraocular papillae. The oral surface of the web is pale. Type locality: South Georgia, Southern Ocean, 531480 3000 S 351570 0000 W, 401–411 m. Distribution: Probably restricted to South Georgia and Shag Rocks in depths of 364–964 m and probably deeper, although lack of deep fishing hampers this knowledge. Remarks: T. gunteri is easily identified by the purple pigmentation of the papillae on the back-
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ground of a pale mantle. It occurs only around South Georgia and is the only species of Thaumeledone to occur in this region. The type specimen is in good condition, and although the coloration is lost, the distinctive papillae are clearly visible (Plates 3 and 4).
T. peninsulae sp. nov. (Plate 5, Figures 4, 6C, 7C, 8C, Table 4) Material examined Holotype: NMSZ 2000081.057, Stn 178-1, Antarctic Peninsula, Southern Ocean, 2 May 2000, 611590 S 601190 W, 804–930 m, R.V. Polarstern, ANTARKTIS XVII/3. 1#. Paratypes: NMSZ 2000081.052, Stn 178-1, Antarctic Peninsula, Southern Ocean, 2 May 2000, 611590 S 601190 W, 804–930 m, R.V. Polarstern, ANTARKTIS XVII/3. 6~, 2#. Diagnosis Animals reach a total length of 120 mm. Funnel organ VV-shaped. Posterior salivary glands approximately 80% of length of buccal mass. External coloration pale purple, with large irregularly shaped papillae of a similar hue. The oral surface of the web is deep purple. Hectocotylised sucker count 22–25. Description [based on specimens detailed in Table 4] Animals small to medium sized (ML to 48 mm; TL to 120 mm; Plate 5). Mantle approximately round (MWI 96.8713.3), head narrower than mantle (HWI 72.879.1). Funnel small (FuLI 34.975.1), gently tapered; funnel organ VVshaped. Arms short (MAI 72.179.2). Arm lengths approximately equal (ALI L1 136.1717.4; L2 129.6718.4; L3 129.1717.0; L4 139.4716.7). Suckers uniserial, small (ASI 5.670.9), without sucker enlargement. Third right arm of males hectocotylised, slightly shorter than opposite number (OAI 93.771.2). Hectocotylus club-like in appearance. Ligula moderate to large (LLI 9.872.4); ligula groove very deep, without transverse ridges (Plate 5). Calamus distinct and very
1892
Table 3 Counts and measurements (mm) for T. gunteri
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
80 35 32 40 31 13 11/6 vv 26 25/26 26/28 25/28 25 50 48 51 — 49 — 34 2 8 — — 5/5 5/5 6
gunteri
gunteri
gunteri
gunteri
gunteri
gunteri
NMSZ 2004083.007 Male Mature
NMSZ 2004083.009 Female Submature
NMSZ 2004083.009 Male Mature
NMSZ 2004083.010 Female Spent
NMSZ 1999275 Male Mature
NMSZ SBMNH NMSZ 1999275 2004083.006 Male Female Mature Mature Juvenile
101 38 30 33 30 17 10 6/6 vv 26 21/24 24/22 23/25 20 59 55 55 49 55 19 32 3 8 11 9 5/5 5/5 6/6 50
80 34 29 33 26 15 8 7/7 vv 22 22/24 22/22 21/21 17
92 38 30 34 25 14 8 7/7 vv 20 20/19 20/19 16/19 14 46 48 57 44
107 35 29 37 28 14 10 8/7 vv 31 29/30 30/30 31/30 26 64 65 61 — 69 — 33 3 7 — — 5/5 5/5 7/6 3
44 40 36 34 21 11
50 44 40 35 22 13
25 35/30 29/22+ 28/ 27
30 32/29 30/35 27/32 18 90 91 88 78 95 21 36 3 8 11 9 5/ 5/ 13/ 41
43 42 — 43 — 32 1.5 6 — — 5/5 5/5 7/8 —
19 30 2 6 8 7 5/5 5/5 7/7 48
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen.
103 98 78 94 22 34 3 8 10 7 5/5 5/5 9/
gunteri
41 31 41 30 17 6 7/8 vv 16 19/18 20/20 24/24 21 52 56 57 — 57 — 33 2.5 7 — — 5/5 5/5 7/8
gunteri
gunteri
gunteri
NMSZ 2004083.006 Male Immature
NMSZ 2004083.006 Female Immature
37 16 13 13 15 8 6 4/4 vv E9 E9/9 E9/9 E9/9 E9 17 18 20 — 20 —
62 25 21 24 23 10 6 5/5 vv E12 E12/12 E12/12 E12/12 E10 30 32 35 30 36 20
56 25 21 20 20 11 6 6/6 vv 12 12/11 13/10 13/11 8 31
1 3 — — 5 5 4 —
2 4 4 3 5 5 5 —
1.5 4 — — 5 5 5 —
5 4
5 5 5
29 — 31 —
5 5 5
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Species Status Repository Catalogue number Sex Maturity
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tive system unremarkable and hence not illustrated. Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid. Beaks unremarkable (Fig. 6C). Cartilaginous stylets present (Fig. 7C). Radula with 5 elements, rachidian unicuspid, 1st lateral low and conical, 2nd lateral broad and unicuspid (Fig. 8C). Newly captured specimens of T. peninsulae are pale purple, with large irregularly shaped papillae on the dorsal and ventral surfaces. Unlike the papillae of T. gunteri, they are not differently coloured than the background skin. Supraocular papillae are not apparent. The oral surface of the web is deep purple. Type locality: Antarctic Peninsula, Southern Ocean, 611590 S 601190 W, 804–930 m. Distribution: Antarctic Peninsula, Southern Ocean, in depths from 377 to 1512 m. Etymology: Named after the type locality—the Antarctic Peninsula. Remarks: This species is clearly closely related to T. gunteri. It shares the large irregular papillae that cover the body surface, has similar arm and web formulae and a similar funnel organ. Both species have a loop in the rectum. Nevertheless, we feel that its distinctness in colour patterning, and in the relative size of the ligula and calamus warrants specific status. Praealtus gen. nov. Fig. 3. T. gunteri, NMSZ 2004083.009, #, 38 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
large (CaLI 90.378.7). Hectocotylised arm with 22–25 suckers, opposite arm with up to 35 suckers. Web deep to very deep (WDI 50.072.7), web formula approximately A=B=C=D.E. Ink sac absent. Posterior salivary glands approximately 80% of length of buccal mass. Digestive tract otherwise normal, except for a loop in the rectum (Fig. 4). Gills with 4–6 lamellae per demibranch. Mature ovarian eggs large (410 mm). Penis diverticulum coiled. Spermatophores medium to long (SpLI 110.1717.5) and slender. Reproduc-
Diagnosis Benthic octopodids, ML to 82 mm, TL to 380 mm; arms medium length (ALI 250–340), with small (ASI 4–8) uniserial suckers, third right arm of males hectocotylised with end of arm clearly differentiated into ligula and calamus, ligula small to medium (LLI 4–5), ligula groove round and shallow without transverse ridges, calamus very long (CaLI 50–70), arm tips not otherwise modified; web depth deep (WDI approximately 30); funnel organ VV-shaped; gills with 7–8 lamellae per demibranch; ink sac absent; anal flaps absent; posterior salivary glands tiny; stylets absent; diverticulum of penis coiled, spermatophores long (SpLI 185); beak classically shaped; radula with 7 elements, all unicuspid.
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Plate 3. T. gunteri holotype. BMNH 1951.4.26.50. 35 mm ML, approx. 80 mm TL.
Remarks: This genus is clearly differentiated from Thaumeledone by a range of characters. The arms are longer relative to mantle length (ALI 250–340 cf. 100–230 in Thaumeledone). The ligula is shorter relative to the length of the hectocotylised arm (LLI 4–5 cf. 9–17 in Thaumeledone) and the ligula groove is shallower, although this is hard to quantify numerically. There are more gill lamellae per demibranch (7–8 cf. 4–6 in Thaumeledone). The spermatophores are longer (SpLI 185 cf. 120–135 in Thaumeledone). Although there is extensive variation in the size of the posterior salivary glands in Thaumeledone, they are never tiny as they are in Praealtus. Finally the shape of the 2nd laterals in the radula is very different in Praealtus (Fig. 8). The absence of stylets in Praealtus also appears to differentiate it from Thaumeledone although it is, as yet, unclear whether stylets are present or absent in T. brevis.
Plate 4. T. gunteri. NMSZ 2004083.007. 38 mm ML, 101 mm TL. Inset shows enlargement of copulatory organ. Ligula length 11 mm, calamus length 9 mm.
P. paralbida sp. nov. (Plate 6, Figures 5, 6D, 8D, Table 5)
Type species: Praealtus paralbida sp. nov. Included species: P. paralbida sp. nov. Etymology: From the Latin, Praealtus, meaning very deep, referring to the depths at which specimens of this genus were captured.
Material examined Holotype: NMSZ 2002038.002, Stn 61/046-8, Southern Ocean, 2 February 2002, 601390 S 531580 W, 2896 m, R.V. Polarstern, ANDEEP I. 1#.
Table 4 Counts and measurements (mm) for T. peninsulae peninsulae Paratype NMSZ 2000081.052 Male Mature
peninsulae Paratype NMSZ 2000081.052 Male Mature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Submature
peninsulae Paratype NMSZ 2000081.052 Female Mature
peninsulae Paratype NMSZ 2000081.052 Female Mature
peninsulae Paratype NMSZ 2000081.052 Female Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
104 37 30 39 29 15 7 9/9 vv 30 28/28 28/30 26/30 18 57 57 55 51 54 25 35 2.5 9 6 5 5/5 5/5 8/8 45
99 40 32 33 29 14 8 9/8 vv 25 23/25 23/25 22/25 20 49 44 45 42 48 24 34 2.5 7 3 3 6/5 5/6 6/6 36
88 32 25 31 24 13 8 7/7 vv 22 22/22 22/22 20/22 18 42 42 40 38 43 22 33 2 5 4 3.5 5/5 5/5 7/8 38
68 36 16 27 20 10 6 6/5 vv 18 18/18 18/18 18/18 17 36 36 37 — 40 — 26 1.5 6 — — 5/5 5/5 6/6 —
75 32 22 30 26 11 10 8/8 vv 23 23/22 23/22 22/22 18 43 43 42 — 44 — 34 2 7 — — 5/5 5/5 7/7 —
77 28 23 32 24 11 8 6/5 vv 21 20/23 21/22 21/22 17 42 38 42 — 46 — 33 1.5 7 — — 5/4 5/4 7/7 —
120 48 36 47 31 14 10 11/11 vv 32 33/34 32/31 32/32 24 68 54 56 — 70 — 32 2 8 — — 5/5 5/5 10/9
113 42 29 48 31 16 12 8/11 Vv 29 31/30 30/30 30/31 26 65 63 62 — 66 — 34 2.5 9 — — 5/5 5/5 10/10 13
112 44 33 40 30 13 9 6/8 vv 32 32/32 29/30 30/30 26 60 61 56 — 61 — 35 2.5 9 — — 5/5 5/5 9/8 9
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen.
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Plate 5. T. peninsulae holotype. NMSZ 2000081.057. 37 mm ML, 104 mm TL. Inset shows enlargement of copulatory organ. Ligula length 6 mm, calamus length 5 mm.
Pale external coloration with pale oral surface. Hectocotylised sucker count 38–42.
Fig. 4. T. peninsulae, NMSZ 2000081.057, #, 37 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
Other material: USNM 1021021, Stn 42/039, Southern Ocean, 25 November 1996, 601370 S 541560 W, 3213–3222 m, R.V. Polarstern, ANTARKTIS XIV/II. 5#, 1~. Comparative material: Moschites albida Berry, 1917. Holotype. AMS C40888, Stn 5, off Wilkes Land, Southern Ocean, 14 January 1914, 641340 S 1271170 E, 1700 fathoms [approx. 3100 m], Mawson Antarctic Expedition. 1~. Diagnosis Animals reach a total length of 380 mm. Funnel organ VV-shaped. Posterior salivary glands tiny.
Description [based on recently captured specimens detailed in Table 5] Animals medium sized (ML to 82 mm; TL to 380 mm; Plate 6). Mantle approximately round (MWI 104.374.0), head narrower than mantle (HWI 61.5723.1). Funnel medium sized (FuLI 43.473.8), gently tapered; funnel organ VVshaped. Arms medium length (MAI 32.672.5). Arm lengths subequal, arm order usually 1=2=3.4 (ALI L1 298.0722.6; L2 307.2723.8; L3 293.9728.6; L4 278.9729.5). Suckers uniserial, small (ASI 6.171.0), without sucker enlargement. Third right arm of males hectocotylised, usually slightly shorter than opposite number (OAI 93.4711.5). Ligula small to medium (LLI 4.770.5); ligula groove round and shallow, without transverse ridges (Plate 6). Calamus distinct and large to very large (CaLI 53.477.2). Hectocotylised arm with 38–42 suckers, opposite arm with up to 66 suckers. Web deep (WDI 29.170.9), damage prevents estimation of web formula. Ink
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Table 5 Counts and measurements (mm) for E. albida (taken from Berry, 1917) and P. paralbida Species Status Repository Catalogue number Sex Maturity
albida Holotype AM C4088 Female Mature
paralbida Holotype NMSZ 2002038.002 Male Mature
Total length Mantle length (dorsal) Mantle length (ventral) Mantle width Head width Full funnel length Free funnel length Funnel organ length (m/l) Funnel organ shape Web depth sector A Web depth sector B (l/r) Web depth sector C (l/r) Web depth sector D (l/r) Web depth sector E Arm length L1 Arm length L2 Arm length L3 Arm length Hc Arm length 4 Sucker count (Hc) Sucker count (L3) Sucker diameter Arm Width Ligula length Calamus length IGLC (l/r) OGLC (l/r) Gill length (l/r) Gamete length
170 35 40 31 12
37
27 129 124 111 — 117 — 57 (R3)
paralbida
paralbida
paralbida
paralbida
paralbida
paralbida
USNM 1021021 Female Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
USNM 1021021 Male Immature
320 65 62 72 52 26 21 14/12 vv 65
[248] [63] [54] [64] [41] [29] [16] —/— —
[258] [64] [60] [63] [41] [27] [20] —/— —
[266] [62] [58] [65] [43] [29] [19] —/— —
[233] [61] [55] [62] [38] [23] [14] —/— —
[273] [61] [57] [66] [44] [30] [25] —/— —
[55]/ 61/
[55]/ [53]/ [55]/
[53]/ [60]/ [53]/
[60]/
61/
[250] [66] [58] [68] [4] [28] [17] —/— — [55] [55]/ [55]/ [55]/
[60]/
[50]/ [60]/ [50]/
[186] [195] [189] [150] [175] [39] [55] [4] [14] [6] [3] [8]/ [7]/
[192] [189] [180] [160] [175] [38] [61] [3] [13] [8] [4] [8]/ [8]/
[201] [204] [203] [179] [192] [38] [62] [4] [13] [8] [4] [8]/ [7]/
[162] [166] [154] [139] [147] [40] [60] [3] [12] [6] [3] [8]/ [7]/
[196] [204] [203] [179] [200] [42] [66] [4] [15] [8] [4] [7]/ [7]/
—
—
—
—
—
46
— —
220 205 220 190 39 64 5 13 11 6
6
120
[186] [189] [181] — [171] — [59] [4] [13] — — 7/8 7/7 16/16 —
l/r, left/right; m/l, medial/lateral; — not measurable, e.g., male character on female specimen, or funnel organ on unfixed material. Brackets indicate measurements made on fresh tissue.
sac absent. Posterior salivary glands tiny. Digestive tract otherwise normal (Fig. 5). Gills with 7–8 lamellae per demibranch. Mature females have not been captured. Penis diverticulum coiled. Spermatophores long (SpLI 184.6) and slender. Reproductive system unremarkable and hence not illustrated. Beaks unremarkable (Figs. 6D and 7). Stylets absent. Radula with 7 elements, rachidian unicuspid, 3 slightly smaller unicuspid elements on either side of rachidian. Marginal plates absent (Fig. 8D).
Newly captured specimens of P. paralbida are pale slaty grey to very pale (almost white) violet all over. There are numerous small simple papillae scattered over the dorsal surface. Supraocular papillae are not apparent. Type locality: Off the Antarctic Peninsula, Southern Ocean, 601390 S 531580 W, 2896 m. Distribution: Possibly circum-Antarctic, in depths from 2896 to 3222 m. Etymology: Paralbida means ‘like albida’ recognising the similarity of this species to albida.
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Plate 6. P. paralbida holotype. NMSZ 2002038.002. 65 mm ML, 320 mm ML. Inset shows enlargement of copulatory organ. Ligula length 11 mm, calamus length 6 mm.
Remarks: The new material of this species was described by Vecchione et al. (in press) as Bentheledone cf. albida. At that time, there were no inconsistencies between our specimens and the little that is known of B. albida, although there was, and there remains, a great deal of uncertainty about the taxonomic characters of B. albida. Although there is only Berry’s (1917) drawing on which to rely, it appears that there may be some differences in the radula of the two species although they are similar in many respects. Berry (1917) illustrates the radula of B. albida as having a large unicuspid rachidian and very small, almost rounded first laterals. The second laterals are more cuspid and the marginal teeth are of a classic cone shape (see Nixon, 1998). Berry appears to illustrate the presence of marginal plates although he does not explicitly state this. The radula of P. paralbida, as seen by light microscopy, appears to lack marginal plates. Furthermore, the first and second laterals resemble marginal teeth in their appearance (Fig. 8D). As commented previously, high levels of variation in radulae of deep-sea species is
Fig. 5. P. paralbida, NMSZ 2002038.002, #, 65 mm ML. Digestive system. Abbreviations: a, anus; asg, anterior salivary gland; bm, buccal mass; cae, caecum; cro, crop; dg, digestive gland; oes, oesophagus; psg, posterior salivary gland; r, rectum; sto, stomach.
not uncommon, however, it is impossible to conclude that B. albida and P. paralbida are synonymous despite their other similarities. The status of albida can only be established by collection of new material in the region of the type locality and we currently consider albida to be a nomen dubium. Should subsequent trawling show albida and paralbida to be conspecific, albida will take precedence and become the type species of Praealtus, thus maintaining taxonomic stability.
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Fig. 8. Radula of (A) T. rotunda, NMSZ 2002038.001, #, 51 mm ML; (B) T. gunteri, NMSZ 2004083.009, #, 38 mm ML; (C) T. peninsulae, NMSZ 2000081.057, #, 37 mm ML; (D) P. paralbida, NMSZ 2002038.002, #, 65 mm ML. Fig. 6. Lower and upper beaks of (A) T. rotunda, 2002038.001, #, 51 mm ML; (B) T. gunteri, 2004083.009, #, 38 mm ML; (C) T. peninsulae, 2000081.057, #, 37 mm ML; (D) P. paralbida, 2002038.002, #, 65 mm ML.
NMSZ NMSZ NMSZ NMSZ
Fig. 7. Cartilaginous stylets of (A) T. rotunda, NMSZ 2002038.001, #, 51 mm ML; (B) T. gunteri, NMSZ 2004083.009, #, 38 mm ML; (C) T. peninsulae, NMSZ 2000081.057, #, 37 mm ML.
Acknowledgements We would like to thank the organisers and leaders of all the cruises listed in the appendix, particularly Wolf Arntz, Karl Herman Koch and Angelika Brandt. The Alfred Wegener Institute kindly provided sea-time aboard R.V. Polarstern; the South Georgia Government, British Antarctic Survey and MRAG kindly facilitated collections during the South Georgia Fish Surveys. Curators at the Australian Museum, the National Museums and Galleries of Wales and the British Museum (Natural History) kindly provided access to specimens. Particular thanks to Amelia MacLellan. The early part of this work was undertaken at the National Museums of Scotland (NMS) with extensive support from Sankurie Pye. NMS’s help in financing cruise participation is much appreciated. Illustrations are by Ian Rendall. We are grateful to F.G. Hochberg for additional input and Vladimir Laptikhovsky and two anonymous referees for their helpful comments on the manuscript.
Appendix See Table A1.
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Table A1 A full list of deep-sea specimens captured in the recent trawling activities Cruise
Location
Depth
No. of specimens
Catalogue number
T. rotunda ANDEEP I ANTARKTIS XIV/2
601390 S 531580 W 601370 S 541560 W
2896 m 3213–3222 m
2 2
NMSZ 2002038.001 USNM 1020991
541010 S 541010 S 531250 S 531460 S 531470 S 531480 S 531530 S 541520 S 541530 S 541570 S 551210 S 551220 S
391330 W 391330 W 371010 W 401300 W 391190 W 311510 W 351470 W 341220 W 341220 W 341160 W 361040 W 361170 W
364–394 m 386–413 m 858–964 m 840–845 m 452–464 m 679–719 m 507–564 m 850–900 m 441–475 m 589–624 m 434–442 m 550–567 m
3 2 2 1 2 2 8 1 2 3 1 4
NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ
611190 S 611350 S 611380 S 611410 S 611590 S
571020 W 581450 W 581470 W 591100 W 601190 W
1440–1512 m 766–824 m 377–429 m 792–862 m 804–930 m
1 3 1 2 9
NMSZ 2000081.052/057
601390 S 531580 W 601370 S 541560 W
2896 m 3213–3222 m
1 6
NMSZ 2002038.002 USNM 1021021
T. gunteri South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia South Georgia
Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries Fisheries
T. peninsulae ANTARKTIS ANTARKTIS ANTARKTIS ANTARKTIS ANTARKTIS
XIV/2 XIV/2 XIV/2 XIV/2 XVII/3
P. paralbida ANDEEP I ANTARKTIS XIV/2
Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey Survey
1994 1997 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003
1999275.010-011 2004083.009 2004083.008 2004083.003 2004083.007 2004083.004 2004083.010 2004083.002 2004083.006 2004083.001 2004083.005
These data have been used to estimate the distributions and depth ranges of each species.
References Allcock, A.L., Collins, M.A., Vecchione, M., 2003. A redescription of Graneledone verrucosa (Verrill, 1881) (Octopoda: Octopodidae). Journal of Molluscan Studies 69, 135–143. Arntz, W.E., Brey, T. (Eds.), 2001. The expedition ANTARKTIS XVII/3 (EASIZ III) of R.V. Polarstern in 2000. Berichte zur Polar- und Meeresforschung 402, pp. 1–181. Berry, S.S., 1917. Cephalopoda. Scientific reports. Australian Antarctic Expedition. Series C. Zoology and Botany 4 (2), pp. 1–38. Collins, M.A., Allcock, A.L., Belchier, M., 2004. Cephalopods of the South Georgia slope. Journal of the Marine Biological Association of the UK 84, 415–419. Fu¨tterer, D.K., Brandt, A., Poore, G.C.B. (Eds.), 2003. The Expeditions ANTARKTIS XIX/3–4 of the Research Vessel POLARSTERN in 2002. Berichte zur Polar- und Meeresforschung 470, pp. 1–174.
Hoyle, W.E., 1885. Diagnoses of new species of cephalopoda collected during the Cruise of HMS challenger. Part 1. The octopoda. Annals and Magazine of Natural History Series V 15, 222–236. Kattner, G. (Ed.), 1998. The expedition ANTARKTIS XIV/2 of R.V. Polarstern in 1996/7. Berichte zur Polarforschung 274, pp. 1–87. Nixon, M., 1998. The radulae of Cephalopoda. In: Voss, N.A., Vecchione, M., Toll, R.B., Sweeney, M.J. (Eds), Systematics and Biogeography of Cephalopods. vol 1. Smithsonian Contributions to Zoology 586, pp. 39–54. O’Shea, S., 1999. The marine fauna of New Zealand: octopoda (mollusca: cephalopoda). NIWA Biodiversity Memoir 112, 1–280. Robson, G.C., 1930. Cephalopoda. I. Octopoda. Discovery Reports 2, pp. 371–402. Robson, G.C., 1932. A Monograph of the Recent Cephalopoda Based on the Collections in the British Museum (Natural History). Part II. The Octopoda (Excluding the Octopodinae). British Museum, London.
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