Polar Biol DOI 10.1007/s00300-006-0248-9 O RI G I NAL PAPE R A new species of Pareledone (Cephalopoda: Octopodidae) from Antarctic Peninsula Waters A. L. Allcock · J. M. Strugnell · P. Prodöhl · U. Piatkowski · M. Vecchione Received: 26 May 2006 / Revised: 15 December 2006 / Accepted: 18 December 2006  Springer-Verlag 2007 Abstract During recent cruises aboard RV Polarstern in the Antarctic Peninsula region, a new species of benthic octopodid was discovered whose generic aYnities based on morphological characteristics were uncertain. Molecular sequence analysis of six mitochondrial and nuclear genes allows this species to be placed with conWdence within the genus Pareledone. The species is described herein and morphological diagnostic characters are provided for its identiWcation. Keywords Systematics · Pareledone felix · Octopodidae · Molecular phylogeny · Antarctic Peninsula A. L. Allcock · J. M. Strugnell · P. Prodöhl School of Biological Sciences, Queen’s University Belfast, 97 Lisburn Road, Belfast, BT9 7BL, UK J. M. Strugnell British Antarctic Survey, Natural Environmental Research Council, High Cross, Madingley Road, Cambridge, CB3 0ET, UK U. Piatkowski (&) FB Marine Ökologie, Leibniz-Institut für Meereswissenschaften, IFM-GEOMAR, Düsternbrooker Weg 20, 24105 Kiel, Germany e-mail: upiatkowski@ifm-geomar.de M. Vecchione NMFS National Systematics Laboratory, National Museum of Natural History, 10th and Constitution Avenue, NW, Washington, DC 20013-7012, USA Introduction During research cruises to the Antarctic Peninsula aboard RV Polarstern during 1996 (Kattner 1998), 2000 (Arntz and Brey 2001) and 2002 (Fütterer et al. 2003), a considerable number of new species of octopuses were discovered (Allcock et al. 2003a, 2003b, 2004; Allcock 2005; Vecchione et al. 2005). These were placed in genera that are either endemic to the Southern Ocean, where they are usually abundant on the shelf and continental slope (e.g., Pareledone, Megaleledone, and Adelieledone), or conWned to the deep sea (e.g., Thaumeledone, Graneledone, and Praealtus). Of these new species, one was particularly hard to place taxonomically. It displayed superWcial morphological similarities with the genus Megaleledone (in terms of overall shape and web depth), yet was internally similar to species of Pareledone. Pareledone is a genus which has radiated widely throughout the Southern Ocean (Lu and Stranks 1994; Allcock 2005). One group of species within this genus is characterized by close-set raised papillae on the dorsal mantle and sometimes other surfaces. The unplaced species was at Wrst thought to have a smooth integument but it became apparent that, when alive, this species could indeed raise small widely set papillae on its dorsal mantle surface. This diVers from other papillated species of Pareledone whose papillae remain raised at all times; although the size of some enlarged papillae (e.g., supraocular papillae) can be varied in described species of Pareledone, the papillae can never be completely Xattened. When seven new species of Pareledone were described from the region of the Antarctic Peninsula (Allcock 2005), the aberrant species was excluded as no certainty had been reached as to its taxonomic placement. Following molecular sequence analysis, it is now 13 Polar Biol clear that this species falls within Pareledone. The molecular data are presented herein, and the new species is described as Pareledone felix sp. nov. Materials and methods As part of expedition ANTARKTIS XVII/3 (Arntz and Brey 2001), RV Polarstern, the German icebreaker, Wshed in the area of the BransWeld Strait and the South Shetland Islands between 24 April and 7 May 2000. Specimens were collected predominantly with a commercial bottom trawl. The sampling yielded 21 specimens of P. felix sp. nov. During expeditions ANTARKTIS XIV/2 (Kattner 1998) and ANTARKTIS XIX/3 (Fütterer et al. 2003), a CCAMLR Wsh survey was conducted in the region of Elephant Island, the continental shelf region north and west of the southern South Shetland Islands, and the shelf north of Joinville Island. Between 15 November and 24 December 1996, a commercial bottom trawl yielded 263 specimens of P. felix sp. nov. Between 29 January and 21 February 2002, a commercial bottom trawl yielded 126 specimens of P. felix sp. nov. Specimens were examined live where possible and when freshly dead. A small number of specimens was preserved in 4% formalin and shipped to UK where they have been deposited in the zoological collections of the National Museums of Scotland, Edinburgh (NMSZ) under catalog numbers NMSZ 2000081 and NMSZ 2002037. Prior to formalin Wxation, tissue samples were taken from these vouchered specimens and from vouchered specimens of other closely related species in the vicinity and preserved in 70% ethanol for subsequent application of molecular techniques. DNA was extracted from a selection of these tissue samples (Table 1) and used in molecular sequence analysis. The DNA extraction protocol followed that of Taggart et al. (1992). BrieXy, 375
l of 0.2 M EDTA, 0.5% sodium lauroylsarcosine (pH 8.0), and 10
l proteinase K (20 mg/ml) were added to the tissue sample (»0.1 g) and incubated overnight at 55°C. The following day 10
l RNAse (20 mg/ml) was added to each tube and incubated for 1 h at 37°C. The solution was extracted once with phenol and once with chloroform:isoamyl alcohol (99:1). DNA was precipitated using three volumes of 92% ethanol and then washed overnight in 70% ethanol before being re-suspended in 30
l of sterile TE (pH 8.0) and stored at 4°C. Primers for four mitochondrial genes (12S rDNA, 16S rDNA, COI, COIII) were taken from the literature (Simon et al. 1990, 1991; Folmer et al. 1994; Simon et al. 1994; Strugnell et al. 2004; Guizk et al. 2005) with the 12S rDNA and COI primers modiWed slightly to match cephalopod sequences on GenBank. Primers used for two nuclear genes (rhodopsin, ODH) were designed in the conserved regions of cephalopod and invertebrate sequences of these genes present on GenBank and are available from the authors on request. PCR reactions were carried out in 25
l volumes. Thermal cycling conditions consisted of a denaturation Table 1 Cephalopod tissue samples used in this study Species Specimen number RV Polarstern station number Museum catalog number Adelieldone polymorpha Megaleledone setebos Pareledone albimaculata P. albimaculata Pareledone aequipapillae P. aequipapillae Pareledone aurata P. aurata Pareledone charcoti P. charcoti Pareledone cornuta P. cornuta Pareledone panchroma Pareledone felix P. felix Pareledone serperastrata P. serperastrata Pareledone subtilis P. subtilis Pareledone turqueti P. turqueti 219 AP 136 AP 1973 AP 1992 AP 222 AP 220 AP 450 AP 459 AP 422AP 428 AP 124 AP 44 AP 1913 AP 206 AP 393 AP 1898 AP 1897 AP 194 AP 193 AP 269 AP 119 AP 61/048-1 61/045-1 61/101-1 61/103-1 61/048-1 61/048-1 61/052-1 61/052-1 61/051-1 61/051-1 61/045-1 61/044-1 61/098-1 61/048-1 61/049-1 61/093-1 61/093-1 61/047-1 61/047-1 61/048-1 56/173-1 NMSZ 2002037.029 NMSZ 2002037.037 NMSZ 2002037.010 NMSZ 2002037.011 NMSZ 2002037.002 NMSZ 2000152.002 NMSZ 2002037.003 NMSZ 2002037.003 NMSZ 2002037.044 NMSZ 2002037.044 NMSZ 2002037.017 NMSZ 2002037.016 NMSZ 2002037.026 NMSZ 2002037.013 NMSZ 2002037.014 NMSZ 2002037.022 NMSZ 2002037.022 NMSZ 2002037.026 NMSZ 2002037.026 NMSZ 2002037.046 NMSZ 2000081.036 13 Polar Biol step at 94°C for 2 min, followed by 35 cycles of 94°C for 40 s, 50°C for 40 s, and 72°C for 90 s. A Wnal extension step of 72°C for 10 min was added in each case. Annealing temperatures varied according to the primers used and are available from the authors on request. AmpliWed products were puriWed using the QIAGEN PCR puriWcation kit (QIAGEN Ltd., UK) following manufacturer’s instructions. PuriWed PCR products were commercially sequenced by Macrogen Inc (Korea) in both directions using the same primers used for PCR ampliWcation. DNA sequences were compiled and aligned by eye in Se-Al v2.0a11 Carbon (Rambaut 2002). It was necessary to introduce gaps to align sequences of 12S rDNA, 16S rDNA, and rhodopsin. The sequence data for each gene was concatenated into a single dataset. Of the 2,755 characters used in the analysis, 357 (13%) were found to be variable. PAUP v4.0b10 (SwoVord 1998) was used to perform full heuristic searches. Starting trees were generated by neighbor joining (NJ) (Saitou and Nei 1987). A GTR (
+ I) likelihood model incorporating rate heterogeneity was used. Branch swapping was performed using SPR (subtree pruning regrafting). Parameters were then re-estimated and Wnally branch swapping was performed using NNI (nearest-neighbor interchange). Substitution model parameter values were A = 0.33, C = 0.16, G = 0.15, T = 0.36, AMC = 1.61, AMG = 6.29, AMT = 2.53, CMG = 1.72, CMT = 14.17, GMT = 1.00, I = 0.64,
= 0.74. ML bootstrap values of clade support were generated using the above parameters. MrBayes 3 (Ronquist and Huelsenbeck 2003) was used to calculate marginal posterior probabilities using the GTR + I +
model of nucleotide substitution for each partition. Model parameter values were treated as unknown and were estimated in each analysis. Random starting trees were used and analyses were run 1 million generations, sampling the Markov chain every 100 generations. The analysis was performed twice, in each case starting from a diVerent random tree to ensure that the analyses were not trapped in a local optima. The program Tracer v1.3 (Rambaut and Drummond 2003) was used to ensure that the Markov chains had reached stationarity and to also determine the correct ‘burn-in’ for the analysis (i.e., the number of initial generations that must be discarded before stationarity is reached). For the morphological taxonomic analyses, comparative material was made available by the Australian Museum, Sydney (AM); the Museum National d’Histoire Naturelle, Paris (MNHN); the Natural History Museum, London (BMNH); the Zoologisches Museum, Berlin (ZMB); and Museum Victoria (NMV). Where indices are calculated, the values given are the mean § standard deviation. 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. Abbreviations used are those recommended in the guidelines for octopus taxonomy published by Roper and Voss (1983) and are as follows: mantle length (ML), total length (TL), mantle width index (MWI), head width index (HWI), web depth index (WDI), funnel length index (FuLI), mantle arm index (MAI), arm length index (ALI), arm sucker index (ASI), opposite arm index (OAI), ligula length index (LLI), calamus length index (CaLI), egg length index (EgLI), and spermatophore length index (SpLI). Results Sequences generated in this study are available from GenBank under accession numbers EF102113EF102235. The phylogenetic tree is rooted using Adelieledone polymorpha as previous phylogenetic studies (Carlini et al. 2001; Strugnell et al. 2004) have conWrmed its position to be basal to Pareledone (Fig. 1). Pareledone turqueti is basal within Pareledone and is supported by a posterior probability (PP) of 0.98 and a bootstrap (BS) of 58%. The rest of the Pareledone taxon is divided into two clades (PP = 0.99, BS = 55) (Fig. 1). The Wrst of these clades is highly supported (PP = 1.00, BS = 97) and also contains highly supported relationships between Pareledone cornuta and P. albimaculata (PP = 0.99, BS = 85) and Pareledone charcoti and P. felix (PP = 1.00, BS = 88). The placement of P. aequipapillae within the clade is less well supported (PP = 0.51). The second clade is only supported by Bayesian posterior probabilities (0.81) and contains a sister taxa relationship between Pareledone aurata and Pareledone subtilis (PP = 0.92, BS = 52) and a less well supported relationship between Pareledone panchroma and Pareledone serperastrata (PP = 0.64). Systematics Octopodidae Orbigny 1840 Pareledone Robson 1932 P. felix sp. nov (Figs. 2, 3, 4, and 5; Tables 2 and 3) 13 Polar Biol Fig. 1 Maximum likelihood (ML) tree depicting the phylogenetic relationship of 12 species (21 individuals) of Octopoda. The analysis employed four mitochondrial (12S rDNA, 16S rDNA, COI, COIII) and two nuclear genes (rhodopsin, ODH). Bayesian support values are indicated above the nodes, ML bootstrap values with 50% support or greater are indicated below the nodes Material examined Holotype: NMSZ 2002037.013, RV Polarstern, cruise ANT XIX/3, Elephant Island, Stn 61/048-1, 61°10⬘S, 54°41⬘W, 278–343 m, one mature #. Paratypes: NMSZ 2000081.034, RV Polarstern, cruise ANT XVII/3, West of Deception Island, Stn 56/173-1, 63°01⬘S, 61°09⬘W, 352–379 m, one submature #, one juvenile $, two immature $, one submature $; NMSZ 2000081.039, RV Polarstern, cruise ANT XVII/3, West of Deception Island, Stn 56/174-1, 63°01⬘S, 61°09⬘W, 311–365 m, one submature $; NMSZ 2000081.044, RV Polarstern, cruise ANT XVII/3, west of Deception Island, Stn 56/ 177-1, 62°50⬘S, 60°49⬘W, 200–202 m, three mature #, one immature $, two submature $, two mature $; NMSZ 2000081.056, RV Polarstern, cruise ANT XVII/3, Drake 13 Passage, north of Livingstone Island, Stn 56/183-1, 62°07⬘S, 60°22⬘W, 200–204 m, one immature #, two mature #, one mature $; NMSZ 2002037.012, RV Polarstern, cruise ANT XIX/3, Elephant Island, Stn 61/ 045-1, 60°59⬘S, 55°11⬘W, 196–269 m, one immature #, one mature #, one submature $; NMSZ 2002037.014, RV Polarstern, cruise ANT XIX/3, Elephant Island, Stn 61/049-1, 61°12⬘S, 54°41⬘W, 272–306 m, one immature #, two submature #, four mature #, one immature $, two submature $; NMSZ 2002037.015, RV Polarstern, cruise ANT XIX/3, Elephant Island, Stn 61/0631, 60°52⬘S, 55°31⬘W, 279–292 m, one juvenile $. Other material: NMSZ 2000081.026, RV Polarstern, cruise ANT XVII/3, BransWeld Strait, Stn 56/165-1, 63°01⬘S, 59°07⬘W, 615–621 m, one immature #; NMSZ 20000 81.029, RV Polarstern, cruise ANT XVII/3, BransWeld Polar Biol Fig. 2 Pareledone felix sp. nov. Whole animal. NMSZ 20000 81.029, 45 mm ML Strait, Stn 56/166-1, 63°02⬘S, 59°10⬘W, 666 m, one mature #, one mature $; NMSZ 2000081.049, RV Polarstern, cruise ANT XVII/3, Drake Passage, north of Livingstone Island, Stn 56/178-2, 61°59⬘S, 60°19⬘W, 804–930 m, one immature #, one mature #. Comparative material examined Moschites aurorae Berry 1917, holotype [AM C40891], Mawson Antarctic Expedition stn 8, oV Queen Mary Land, 66°08⬘S, 94°17⬘E, 219 m. Pareledone framensis Lu and Stranks 1994, paratype [NMV F65667], RV Aurora australis stn AA91-100, oV MacRobertson Land, 67°27.35⬘S, 68°50.34⬘E, 145–150 m. Pareledone prydzensis Lu and Stranks 1994, paratype [NMV F65625], RV Aurora australis stn Fig. 3 Pareledone felix sp. nov. Digestive tract with inset of upper and lower beaks to same scale. NMSZ 2000081.029, 45 mm ML. 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 AA91-89 (2), oV Amery Iceshelf, 66°47⬘S, 72°36⬘W, 526–532 m. Eledone turqueti Joubin 1905, lectotype [MNHN 5.7.1089], ‘Charcot’ Antarctic Expedition, Ile Wandel, 65°05⬘S, 25 m. Moschites harrissoni Berry 1917, holotype [AM C40892], Mawson Antarctic Expedition stn 10, oV Shackleton Glacier, 65°06⬘S, 96°13⬘E, 494–595 m. Moschites antarcticus Thiele 1920, lectotype [ZMB Moll 110001a], Kaiser Wilhelm II Land, 66°47⬘S, 89°20⬘E, 385 m. 13 Polar Biol Fig. 4 Pareledone felix sp. nov. Reproductive anatomy. NMSZ 2000081.044, 42 mm ML, # and NMSZ 2000081.056, 60 mm ML, $. a Calamus (4 mm) and ligula (8 mm). b Male reproductive tract. ag accessory gland; d diverticulum; mg mucilaginous gland; sg spermatophoric gland; ss spermatophoric sac; t testes; to terminal organ; vd vas deferens. c Female reproductive tract. o ovary; og oviducal gland; po posterior oviduct. d Transverse section through egg (8 mm diameter) Fig. 5 Pareledone felix sp. nov. Radula from specimen with ML 51 mm. Scale bar represents 0.5 mm Moschites adelieana Berry 1917, holotype [AM C40889], Mawson Antarctic Expedition stn 2, oV Mertz Glacier, Adelieland, 66°55⬘S, 145°21⬘E, 450– 549 m. Graneledone polymorpha Robson 1930, holotype [BMNH 1951.4.26.26], Discovery Expedition stn 42, South Georgia, 120–204 m. P. aequipapillae Allcock 2005, holotype [NMSZ 2003152.002], RV Polarstern, stn 61/048-1, Elephant Island, 61°10⬘S, 54°34⬘W, 278–343 m. P. albimaculata Allcock 2005, holotype [BMNH 1996192], RV Polarstern, stn 42/021, Elephant Island, 61°10⬘S, 56°04⬘W, 169–175 m. P. aurata Allcock 2005, holotype [NMSZ 2003152.001], RV Polarstern, stn 61/052-1, Elephant Island, 61°21⬘S, 55°14⬘W, 264–270 m. 13 P. cornuta Allcock 2005, holotype [BMNH 1996196], RV Polarstern, stn 42/004, Elephant Island, 61°10⬘S, 56°04⬘W, 169–175 m. P. panchroma Allcock 2005, holotype [NMSZ 2003152.005], RV Polarstern, stn 56/178-2, Elephant Island, 61°59⬘S, 60°19⬘W, 904–930 m. P. serperastrata Allcock 2005, holotype [NMSZ 2003152.003], RV Polarstern, stn 61/092-1, Elephant Island, 61°01⬘S, 55°46⬘W, 123–159 m. P. subtilis Allcock 2005, holotype [NMSZ 2003152.004], RV Polarstern, stn 61/047-1, Elephant Island, 61°04⬘S, 54°36⬘W, 308–399 m. Graneledone setebos Robson 1932, holotype [BMNH 1919.12.30.27], British Antarctic ‘Terra Nova’ Expedition 1911–1912, stn 325, McMurdo Sound, oV Cape Evans, 77°40⬘S, 166°30⬘W. In addition, all specimens listed in Table 1 were examined and compared to the type material listed to check identiWcations. Diagnosis: Small benthic incirrate octopod. Mantle length 60 mm, total length to 150 mm. Stylets present. Arms with uniserial row of suckers. Arms approximately 1.5 times the length of body. Right third arm of males hectocotylized; copulatory organ with clearly diVerentiated ligula and calamus. Ligula simple, without transverse ridges. Arm tips not otherwise modiWed. Suckers small; distinct enlarged suckers absent. Web deep to very deep. Gills with 7–9 lamellae per demibranch. Ink sac present, anal Polar Biol Table 2 Pareledone felix sp. nov: counts and measurements (mm) of male specimens Status Holotype Paratype Paratype Paratype Paratype Repository NMSZ NMSZ NMSZ NMSZ NMSZ Catalog number 2002037.013 2000081.044 2000081.044 2000081.056 2000081.056 2000081.029 2002037.012 2002037.014 Vessel Polarstern Polarstern Polarstern Polarstern Polarstern Polarstern Polarstern Polarstern Station number 61/048-1 56/177-1 56/177-1 56/183-1 56/183-1 56/166-1 61/045-1 61/049-1 Depth (m) 278–343 200–202 200–202 200–204 200–204 666 196–269 272–306 Sex M M M M M M M M Maturity Mature Mature Mature Mature Mature Mature Mature Mature 42 41 39 41 44 44 43 45 115 39 31 20 115 38 33 17 115 41 35 17 116 39 31 17 116 42 32 19 124 44 35 20 127 43 32 23 144 43 34 23 10 10 10 12 13 10 11 14 12 14 9 12 12 12 12 12 27 40 25 40 27 40 28 45 27 45 30 45 28 45 27 45 30 34 29 30 29 32 33 32 30/30 34/32 31/30 28/34 32/34 35/36 33/38 36/30 33/34 32/33 30/30 28/34 32/34 37/37 33/38 38/34 33/32 31/31 32/31 26/31 32/34 34/37 33/35 35/30 26 31 27 26 D 27 30 63/62 70/60 67/63 61/64 d/60 64/62 d/77 77/78 59/57 63/55 62/66 62/65 62/67 66/67 67/75 81/74 61/60 54/55 63/62 64/63 64/62 65/66 66/73 85/67 64/66 55/62 64/63 66/70 67/72 67/72 77/73 85/84 7 3 38/28 7 2 36/26 7 2.5 38/25 7 2.5 39/27 8 2.5 38/27 7 3 40/26 8 3.5 33/25 8 3 38/27 13/12 8/8 10/10 8/8 10/10 8/8 9/10 8/8 11/11 8/8 11/11 8/8 11/12 8/7 12/12 9/8 8/8 9/8 8/8 8/8 8/8 8/8 7/7 8/8 6 2 61 8 4 70 8 5 70 7 3 67 8 3 67 6 1 55 7 2.5 66 7 2 68 1+1 4+1 5+1 2+1 1+1 1+1 3+1 Mantle length (dorsal) Total length Mantle width Head width Full funnel length Free funnel length Funnel organ length Pallial aperture Pallial aperture (%) 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/R1 Arm length L2/R2 Arm length L3/Hc Arm length L4/R4 Arm width Sucker diameter Sucker count L3/Hc Gill length (l/r) Gill lamellae: inner (l/r) Gill lamellae: outer (l/r) Ligula length Calamus length Spermatophore length No. of spermatophores NMSZ D 5+0 Paratype Paratype NMSZ NMSZ l left; r right; d damaged 13 Polar Biol Table 3 Pareledone felix sp. nov: counts and measurements (mm) of female specimens Status Paratype Paratype Paratype Paratype Repository NMSZ NMSZ NMSZ NMSZ NMSZ NMSZ Catalog number 2000081.44 2000081.44 2000081.44 2000081.056 2000081.029 2002037.014 Vessel Polarstern Polarstern Polarstern Polarstern Polarstern Polarstern Station number 56/177-1 56/177-1 56/177-1 56/183-1 56/166-1 61/049-1 Depth (m) 200–202 200–202 200–202 200–204 666 272–306 Sex F F F F F F Maturity Mature Mature Mature Mature Mature Mature 43 41 40 60 45 48 120 44 33 20 109 41 32 18 106 44 29 17 149 58 33 26 132 47 34 24 124 40 29 21 12 10 8 16 14 9 12 12 9 16 17 14 28 45 32 28 45 28 28 45 28 29 45 33 33 45 43 26 45 32 34/33 28/28 31/29 37/38 42/41 32/34 38/34 28/29 32/31 37/38 47/42 31/35 35/34 28/29 31/31 34/37 44/45 33/35 33 26 30 38 29 59/56 57/55 56/52 d/70 68/74 62/63 61/59 57/57 57/60 75/75 75/72 62/59 64/60 59/58 55/60 71/75 75/75 62/65 66/65 62/61 56/61 76/74 76/79 64/66 8 3 38/38 7 3 37/39 7 3 40/38 7 4.5 36/38 8 4 36/38 8 3 37/37 12/11 9/10 9/9 13/11 13/14 12/12 9/9 8/9 8/8 8/8 8/8 9/8 8/9 8/9 8/8 8/8 8/8 8/8 14 4 27 12 4 22 10 3 31 22 8 54 12 4 42 9 4 39 Mantle length (dorsal) Total length Mantle width Head width Full funnel length Free funnel length Funnel organ length Pallial aperture Pallial aperture (%) 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/R1 Arm length L2/R2 Arm length L3/R3 Arm length L4/R4 Arm width Sucker diameter Sucker count L3/R3 Gill length (l/r) Gill lamellae: inner (l/r) Gill lamellae: outer (l/r) Egg length Egg width Number of eggs 30 Paratype l left; r right; d damaged Xaps present. Integument covered in small, widely spaced papillae that appear as small, pale dots on the mantle. 13 Description [based on eight mature male and six mature female specimens for whom counts and measurements are given in Tables 2 and 3]: Animals small to Polar Biol medium sized (ML 60 mm; TL 149 mm). Mantle spherical (MWI 98.1 § 6.6), head narrower than mantle (HWI 74.3 § 8.3). Funnel medium sized (FuLI 45.8 § 4.1), gently tapered; funnel organ W or VV shaped, usually VV. Arms short (MAI 62.4 § 7). Arm lengths approximately equal (ALI L1 150.4 § 14.8; L2 148.2 § 14.5; L3 148.2 § 17.4; L4 154.1 § 17.5). Suckers uniserial, small (ASI 6.9 § 1), without sucker enlargement. Third right arm of males hectocotylised, approximately equal in length to opposite number (OAI 98.1 § 8.9). Ligula medium to large (LLI 11.3 § 1.9); ligula groove long, well-marked and shallow, without transverse ridges (Fig. 4a). Calamus distinct and large (CaLI 38.4 § 13.8) although quite variable in length (Fig. 4a). Hectocotylised arm with 25–28 suckers, opposite arm with up to 40 suckers. Web deep to very deep (WDI 50.5 § 4.2), web sectors approximately equal in depth although some variation is apparent with sector E most commonly shallowest (but only by a couple of mm) and sector A sometimes shallower than sectors B, C, and D. Ink sac present. Posterior salivary glands medium sized, slightly shorter than length of buccal mass (Fig. 3). Typical octopodid radula with nine elements (Fig. 5). Gills with 7–9 lamellae per inner demibranch and 7–9 lamellae per outer demibranch. Rostral tip of lower beak rounded (Fig. 3). Mature ovarian eggs large, up to 22 mm, equivalent to an EgLI of 36.7 with approximately 12 follicular folds (Fig. 4d). Penis diverticulum coiled (Fig. 4b). Spermatophores long, up to 70 mm (SpLI 155.1 § 16.6) and slender. The integument is delicate and prone to damage, especially on the web. Live animals are a pale blue to green (Fig. 6). Papillae, which are scattered and widespread over the dorsal surface, are pale. The papillae are small, and are not permanently raised as, for example, in P. charcoti. In dead specimens the integument often appears completely smooth, the pale dots being the only clue to where the papillae are situated. Type locality: Antarctica, oV Elephant Island, 61°10⬘S, 54°41⬘W, 278–343 m. Distribution: OV South Shetland Islands, including oV Elephant Island, south into the BransWeld Strait and north onto the slope of the Drake Passage. Found in a wide range of depths from approximately 200 to 800 m. Etymology: This name is in honor of Louise Allcock’s son Felix and Mike Vecchione’s father and grandfather, both Felixes. Discussion Some initial diYculties were encountered in placing this species. In cruise reports it has been referred to as Pareledone (as P. sp. 1; Piatkowski et al. 1998: 44) and Fig. 6 Pareledone felix sp. nov. Live animal to Megaleledone? (as M. sp. 17; Piatkowski et al. 2003: 33). There are only three genera of shallow water octopodids in the Southern Ocean: Adelieledone, Pareledone and Megaleledone. All these taxa share certain morphological features in common, for example, they have similarly sized uniserial suckers, a relatively small number of large eggs, and a large calamus. Some of the features commonly recorded in octopus taxonomy are of no use in diVerentiating between these taxa. For example, in these taxa, the presence of an ink sac appears to be depth dependant. In the shallower A. polymorpha, the ink sac is clearly deWned whereas, in the deeper A. piatkowski, the inksac is reduced (Allcock et al. 2003a). Likewise in the shallower species of papillated Pareledone from the Antarctic Peninsula (P. charcoti, P. aequipapillae, P. albimaculata, P. aurata, P. cornuta, P. serperastrata and P. subtilis), which extend to depths of about 500 m, the ink sac is present, whereas in P. panchroma, which inhabits depths from about 450 to below 800 m, the ink sac is absent (Allcock 2005). Other characters which are of little use include the funnel organ whose shape varies within taxa as well as between taxa, and characters such as relative arm length show a continuum of variation between the taxa. A comparison of characters between taxa (Table 4) shows, however, that there are clear diVerences between P. felix sp. nov and other genera. Adelieledone diVers from all other taxa in that 13 Polar Biol Table 4 Comparison of characters between shallow water taxa of Antarctic octopods Character Adelieledone Megaleledone Pareledone (papillated species) Pareledone (smooth species) P. felix sp. nov. Mantle length (mm) Total length (mm) Arm length Sucker size Ligula shape to 90 to 250 to 63 to 120 to 60 to 250 to 900 to 190 to 350 to 150 ALI to 190 ASI 4–10 With transverse ridges LLI 10–16 Large, CaLI 35–50 ALI to 300 ASI 6–12 No transverse ridges LLI 3–6 Large, CaLI 30–40 ALI to 220 ASI 5–12 No transverse ridges LLI 7–13 Large, CaLI 25–50 ALI to 250 ASI 5–10 No transverse ridges LLI 6–9 Large, CaLI 30–40 WDI 20–40 W WDI 40–60 VV WDI 20–45 VV or W WDI 20–30 VV ALI to 190 ASI 5–9 No transverse ridges LLI 9–15 Large, CaLI 30–60 WDI 45–60 VV or W 6–8 10–13 6–9 8–11 7–9 Present or reduced Present Twice length of buccal mass Present Present or absent Present or reduced Present Absent 70–80% length of buccal mass 16 Large and very robust with blunt rostrum Present Short to medium, SpLI 40–60 41 Present or absent Approximately equal to length of buccal mass Medium with blunt rostrum Present Long, SpLI, 100–200 20 Present Approximately equal to length of buccal mass Medium with blunt rostrum Present Long, SpLI 110–150 19 Present Slightly shorter than length of buccal mass Medium with blunt rostrum Present Long, SpLI 120–180 22 200 130 70 130 54 Ligula size Calamus size Web depth Funnel organ shape Gill lamellae count Ink sac Anal Xaps Size of posterior salivary glands Beak size/shape Stylets Spermatophore length Max egg size (mm) Max number of eggs Small and delicate with sharp rostral tip Absent Medium, SpLI 60–80 it has no stylets, has very large posterior salivary glands, has transverse ridges on its ligula, and has a very distinctive lower beak shape with a sharp rostral tip. P. felix diVers from Megaleledone in that the ligula in Megaleledone is very small (LLI 3–6 vs. LLI 9–15 in P. felix) as are the spermatophores (SpLI 40–60 vs. SpLI 120–180 in P. felix). The gill lamellae count in Megaleledone is much higher. Oddly, Megaleledone has no anal Xaps (thought to be used in direct ink) despite the presence of an ink sac. There is superWcial resemblance between P. felix and Megaleledone; however, in that they both have very deep webs (WDI 45–60 in P. felix, 40–60 in Megaleledone). In fact, juvenile Megaleledone and P. felix (which are approximately equal in size) can be hard to tell apart without internal examination if they are in poor condition. P. felix shares with other species of Pareledone similar beak morphology and long spermatophores, and it aligns slightly more closely with the group of papillated Pareledone sharing characters such as a long ligula and similar body size at maturity. 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