Polar Biol

DOI 10.1007/s00300-006-0248-9

ORIGINAL PAPER

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 Polarsternin the Antarctic Peninsula region, a new species of ben-thic octopodid was discovered whose generic aYnitiesbased on morphological characteristics were uncertain.Molecular sequence analysis of six mitochondrial andnuclear genes allows this species to be placed with con-Wdence within the genus Pareledone. The species isdescribed herein and morphological diagnostic charac-ters are provided for its identiWcation.

Keywords Systematics · Pareledone felix · Octopodidae · Molecular phylogeny · Antarctic Peninsula

Introduction

During research cruises to the Antarctic Peninsulaaboard RV Polarstern during 1996 (Kattner 1998), 2000(Arntz and Brey 2001) and 2002 (Fütterer et al. 2003), aconsiderable number of new species of octopuses werediscovered (Allcock et al. 2003a, 2003b, 2004; Allcock2005; Vecchione et al. 2005). These were placed ingenera that are either endemic to the Southern Ocean,where they are usually abundant on the shelf andcontinental slope (e.g., Pareledone, Megaleledone, andAdelieledone), or conWned to the deep sea (e.g., Thaum-eledone, Graneledone, and Praealtus). Of these new spe-cies, one was particularly hard to place taxonomically. Itdisplayed superWcial morphological similarities with thegenus Megaleledone (in terms of overall shape and webdepth), yet was internally similar to species of Parele-done. Pareledone is a genus which has radiated widelythroughout the Southern Ocean (Lu and Stranks 1994;Allcock 2005). One group of species within this genus ischaracterized by close-set raised papillae on the dorsalmantle and sometimes other surfaces. The unplaced spe-cies was at Wrst thought to have a smooth integumentbut it became apparent that, when alive, this speciescould indeed raise small widely set papillae on its dorsalmantle surface. This diVers from other papillated speciesof Pareledone whose papillae remain raised at all times;although the size of some enlarged papillae (e.g., supra-ocular papillae) can be varied in described species ofPareledone, the papillae can never be completely Xat-tened. When seven new species of Pareledone weredescribed from the region of the Antarctic Peninsula(Allcock 2005), the aberrant species was excluded as nocertainty had been reached as to its taxonomic place-ment. Following molecular sequence analysis, it is now

A. L. Allcock · J. M. Strugnell · P. ProdöhlSchool of Biological Sciences, Queen’s University Belfast, 97 Lisburn Road, Belfast, BT9 7BL, UK

J. M. StrugnellBritish 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, Germanye-mail: upiatkowski@ifm-geomar.de

M. VecchioneNMFS National Systematics Laboratory, National Museum of Natural History, 10th and Constitution Avenue, NW, Washington, DC 20013-7012, USA

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clear that this species falls within Pareledone. Themolecular data are presented herein, and the new spe-cies is described as Pareledone felix sp. nov.

Materials and methods

As part of expedition ANTARKTIS XVII/3 (Arntzand Brey 2001), RV Polarstern, the German ice-breaker, Wshed in the area of the BransWeld Strait andthe South Shetland Islands between 24 April and 7May 2000. Specimens were collected predominantlywith a commercial bottom trawl. The sampling yielded21 specimens of P. felix sp. nov.

During expeditions ANTARKTIS XIV/2 (Kattner1998) and ANTARKTIS XIX/3 (Fütterer et al. 2003),a CCAMLR Wsh survey was conducted in the region ofElephant Island, the continental shelf region north andwest of the southern South Shetland Islands, and theshelf north of Joinville Island. Between 15 Novemberand 24 December 1996, a commercial bottom trawlyielded 263 specimens of P. felix sp. nov. Between 29January and 21 February 2002, a commercial bottomtrawl yielded 126 specimens of P. felix sp. nov.

Specimens were examined live where possible andwhen freshly dead. A small number of specimens waspreserved in 4% formalin and shipped to UK wherethey have been deposited in the zoological collectionsof the National Museums of Scotland, Edinburgh(NMSZ) under catalog numbers NMSZ 2000081 andNMSZ 2002037. Prior to formalin Wxation, tissue

samples were taken from these vouchered specimensand from vouchered specimens of other closely relatedspecies in the vicinity and preserved in 70% ethanol forsubsequent application of molecular techniques. DNAwas extracted from a selection of these tissue samples(Table 1) and used in molecular sequence analysis.

The DNA extraction protocol followed that of Tag-gart et al. (1992). BrieXy, 375 �l of 0.2 M EDTA, 0.5%sodium lauroylsarcosine (pH 8.0), and 10 �l proteinaseK (20 mg/ml) were added to the tissue sample (»0.1 g)and incubated overnight at 55°C. The following day10 �l RNAse (20 mg/ml) was added to each tube andincubated for 1 h at 37°C. The solution was extractedonce with phenol and once with chloroform:isoamylalcohol (99:1). DNA was precipitated using three vol-umes of 92% ethanol and then washed overnight in70% ethanol before being re-suspended in 30 �l ofsterile TE (pH 8.0) and stored at 4°C.

Primers for four mitochondrial genes (12S rDNA,16S rDNA, COI, COIII) were taken from the litera-ture (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 modiWedslightly to match cephalopod sequences on GenBank.Primers used for two nuclear genes (rhodopsin,ODH) were designed in the conserved regions ofcephalopod and invertebrate sequences of thesegenes present on GenBank and are available from theauthors 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 219 AP 61/048-1 NMSZ 2002037.029Megaleledone setebos 136 AP 61/045-1 NMSZ 2002037.037Pareledone albimaculata 1973 AP 61/101-1 NMSZ 2002037.010P. albimaculata 1992 AP 61/103-1 NMSZ 2002037.011Pareledone aequipapillae 222 AP 61/048-1 NMSZ 2002037.002P. aequipapillae 220 AP 61/048-1 NMSZ 2000152.002Pareledone aurata 450 AP 61/052-1 NMSZ 2002037.003P. aurata 459 AP 61/052-1 NMSZ 2002037.003Pareledone charcoti 422AP 61/051-1 NMSZ 2002037.044P. charcoti 428 AP 61/051-1 NMSZ 2002037.044Pareledone cornuta 124 AP 61/045-1 NMSZ 2002037.017P. cornuta 44 AP 61/044-1 NMSZ 2002037.016Pareledone panchroma 1913 AP 61/098-1 NMSZ 2002037.026Pareledone felix 206 AP 61/048-1 NMSZ 2002037.013P. felix 393 AP 61/049-1 NMSZ 2002037.014Pareledone serperastrata 1898 AP 61/093-1 NMSZ 2002037.022P. serperastrata 1897 AP 61/093-1 NMSZ 2002037.022Pareledone subtilis 194 AP 61/047-1 NMSZ 2002037.026P. subtilis 193 AP 61/047-1 NMSZ 2002037.026Pareledone turqueti 269 AP 61/048-1 NMSZ 2002037.046P. turqueti 119 AP 56/173-1 NMSZ 2000081.036

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step at 94°C for 2 min, followed by 35 cycles of 94°Cfor 40 s, 50°C for 40 s, and 72°C for 90 s. A Wnal exten-sion step of 72°C for 10 min was added in each case.Annealing temperatures varied according to the prim-ers used and are available from the authors on request.AmpliWed products were puriWed using the QIAGENPCR puriWcation kit (QIAGEN Ltd., UK) followingmanufacturer’s instructions. PuriWed PCR productswere commercially sequenced by Macrogen Inc(Korea) in both directions using the same primers usedfor PCR ampliWcation.

DNA sequences were compiled and aligned by eyein Se-Al v2.0a11 Carbon (Rambaut 2002). It was neces-sary to introduce gaps to align sequences of 12S rDNA,16S rDNA, and rhodopsin. The sequence data for eachgene was concatenated into a single dataset. Of the2,755 characters used in the analysis, 357 (13%) werefound to be variable.

PAUP v4.0b10 (SwoVord 1998) was used to per-form full heuristic searches. Starting trees were gener-ated by neighbor joining (NJ) (Saitou and Nei 1987).A GTR (� + I) likelihood model incorporating rateheterogeneity was used. Branch swapping was per-formed using SPR (subtree pruning regrafting).Parameters were then re-estimated and Wnally branchswapping was performed using NNI (nearest-neigh-bor interchange). Substitution model parameter val-ues 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. MLbootstrap values of clade support were generatedusing the above parameters.

MrBayes 3 (Ronquist and Huelsenbeck 2003) wasused to calculate marginal posterior probabilitiesusing the GTR + I + � model of nucleotide substitu-tion for each partition. Model parameter values weretreated as unknown and were estimated in each anal-ysis. Random starting trees were used and analyseswere run 1 million generations, sampling the Markovchain every 100 generations. The analysis was per-formed twice, in each case starting from a diVerentrandom tree to ensure that the analyses were nottrapped in a local optima.

The program Tracer v1.3 (Rambaut and Drummond2003) was used to ensure that the Markov chains hadreached stationarity and to also determine the correct‘burn-in’ for the analysis (i.e., the number of initialgenerations that must be discarded before stationarityis reached).

For the morphological taxonomic analyses,comparative material was made available by theAustralian Museum, Sydney (AM); the MuseumNational d’Histoire Naturelle, Paris (MNHN); the

Natural History Museum, London (BMNH); theZoologisches Museum, Berlin (ZMB); and MuseumVictoria (NMV). Where indices are calculated, thevalues given are the mean § standard deviation.Size descriptors (e.g., large, deep), where givenalongside indices, follow the guidelines proposed atthe taxonomy workshop at the Cephalopod Interna-tional Advisory Council Symposium in Phuket,2003. Abbreviations used are those recommended inthe guidelines for octopus taxonomy published byRoper and Voss (1983) and are as follows: mantlelength (ML), total length (TL), mantle width index(MWI), head width index (HWI), web depth index(WDI), funnel length index (FuLI), mantle armindex (MAI), arm length index (ALI), arm suckerindex (ASI), opposite arm index (OAI), ligulalength index (LLI), calamus length index (CaLI),egg length index (EgLI), and spermatophore lengthindex (SpLI).

Results

Sequences generated in this study are available fromGenBank under accession numbers EF102113-EF102235. The phylogenetic tree is rooted usingAdelieledone polymorpha as previous phylogeneticstudies (Carlini et al. 2001; Strugnell et al. 2004) haveconWrmed its position to be basal to Pareledone(Fig. 1). Pareledone turqueti is basal within Parele-done and is supported by a posterior probability (PP)of 0.98 and a bootstrap (BS) of 58%. The rest of thePareledone taxon is divided into two clades(PP = 0.99, BS = 55) (Fig. 1). The Wrst of these cladesis highly supported (PP = 1.00, BS = 97) and also con-tains highly supported relationships between Parele-done 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 withinthe clade is less well supported (PP = 0.51). The sec-ond clade is only supported by Bayesian posteriorprobabilities (0.81) and contains a sister taxa relation-ship between Pareledone aurata and Pareledone sub-tilis (PP = 0.92, BS = 52) and a less well supportedrelationship between Pareledone panchroma and Par-eledone serperastrata (PP = 0.64).

Systematics

Octopodidae Orbigny 1840Pareledone Robson 1932P. felix sp. nov (Figs. 2, 3, 4, and 5; Tables 2 and 3)

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Material examined

Holotype: NMSZ 2002037.013, RV Polarstern, cruiseANT XIX/3, Elephant Island, Stn 61/048-1, 61°10�S,54°41�W, 278–343 m, one mature #. Paratypes: NMSZ2000081.034, RV Polarstern, cruise ANT XVII/3, Westof Deception Island, Stn 56/173-1, 63°01�S, 61°09�W,352–379 m, one submature #, one juvenile $, twoimmature $, one submature $; NMSZ 2000081.039, RVPolarstern, cruise ANT XVII/3, West of DeceptionIsland, Stn 56/174-1, 63°01�S, 61°09�W, 311–365 m, onesubmature $; 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 $; NMSZ2000081.056, RV Polarstern, cruise ANT XVII/3, Drake

Passage, north of Livingstone Island, Stn 56/183-1,62°07�S, 60°22�W, 200–204 m, one immature #, twomature #, one mature $; NMSZ 2002037.012, RVPolarstern, 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 $; NMSZ 2002037.014,RV Polarstern, cruise ANT XIX/3, Elephant Island,Stn 61/049-1, 61°12�S, 54°41�W, 272–306 m, one imma-ture #, two submature #, four mature #, one immature$, two submature $; NMSZ 2002037.015, RV Polar-stern, cruise ANT XIX/3, Elephant Island, Stn 61/063-1, 60°52�S, 55°31�W, 279–292 m, one juvenile $. Othermaterial: NMSZ 2000081.026, RV Polarstern, cruiseANT XVII/3, BransWeld Strait, Stn 56/165-1, 63°01�S,59°07�W, 615–621 m, one immature #; NMSZ 2000081.029, RV Polarstern, cruise ANT XVII/3, BransWeld

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 boot-strap values with 50% support or greater are indicated below the nodes

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Strait, Stn 56/166-1, 63°02�S, 59°10�W, 666 m, onemature #, one mature $; NMSZ 2000081.049, RVPolarstern, cruise ANT XVII/3, Drake Passage, northof 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 [AMC40891], Mawson Antarctic Expedition stn 8, oVQueen Mary Land, 66°08�S, 94°17�E, 219 m.Pareledone framensis Lu and Stranks 1994, para-type [NMV F65667], RV Aurora australis stnAA91-100, oV MacRobertson Land, 67°27.35�S,68°50.34�E, 145–150 m.Pareledone prydzensis Lu and Stranks 1994, para-type [NMV F65625], RV Aurora australis stn

AA91-89 (2), oV Amery Iceshelf, 66°47�S,72°36�W, 526–532 m.Eledone turqueti Joubin 1905, lectotype [MNHN5.7.1089], ‘Charcot’ Antarctic Expedition, Ile Wan-del, 65°05�S, 25 m.Moschites harrissoni Berry 1917, holotype [AMC40892], Mawson Antarctic Expedition stn 10, oVShackleton Glacier, 65°06�S, 96°13�E, 494–595 m.Moschites antarcticus Thiele 1920, lectotype [ZMBMoll 110001a], Kaiser Wilhelm II Land, 66°47�S,89°20�E, 385 m.

Fig. 2 Pareledone felix sp. nov. Whole animal. NMSZ 20000 81.029,45 mm ML

Fig. 3 Pareledone felix sp. nov. Digestive tract with inset of upperand lower beaks to same scale. NMSZ 2000081.029, 45 mm ML. aanus; asg anterior salivary gland; bm buccal mass; cae caecum; crocrop; dg digestive gland; oes oesophagus; psg posterior salivarygland; r rectum; sto stomach

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Moschites adelieana Berry 1917, holotype [AMC40889], Mawson Antarctic Expedition stn 2, oVMertz Glacier, Adelieland, 66°55�S, 145°21�E, 450–549 m.Graneledone polymorpha Robson 1930, holotype[BMNH 1951.4.26.26], Discovery Expedition stn42, South Georgia, 120–204 m.P. aequipapillae Allcock 2005, holotype [NMSZ2003152.002], RV Polarstern, stn 61/048-1,Elephant Island, 61°10�S, 54°34�W, 278–343 m.P. albimaculata Allcock 2005, holotype [BMNH1996192], RV Polarstern, stn 42/021, ElephantIsland, 61°10�S, 56°04�W, 169–175 m.P. aurata Allcock 2005, holotype [NMSZ2003152.001], RV Polarstern, stn 61/052-1, Ele-phant Island, 61°21�S, 55°14�W, 264–270 m.

P. cornuta Allcock 2005, holotype [BMNH1996196], RV Polarstern, stn 42/004, ElephantIsland, 61°10�S, 56°04�W, 169–175 m.P. panchroma Allcock 2005, holotype [NMSZ2003152.005], RV Polarstern, stn 56/178-2, Ele-phant Island, 61°59�S, 60°19�W, 904–930 m.P. serperastrata Allcock 2005, holotype [NMSZ2003152.003], RV Polarstern, stn 61/092-1, Ele-phant Island, 61°01�S, 55°46�W, 123–159 m.P. subtilis Allcock 2005, holotype [NMSZ2003152.004], RV Polarstern, stn 61/047-1, Ele-phant Island, 61°04�S, 54°36�W, 308–399 m.Graneledone setebos Robson 1932, holotype[BMNH 1919.12.30.27], British Antarctic ‘TerraNova’ Expedition 1911–1912, stn 325, McMurdoSound, oV Cape Evans, 77°40�S, 166°30�W.

In addition, all specimens listed in Table 1 were exam-ined and compared to the type material listed to checkidentiWcations.

Diagnosis: Small benthic incirrate octopod. Man-tle length 60 mm, total length to 150 mm. Styletspresent. Arms with uniserial row of suckers. Armsapproximately 1.5 times the length of body. Rightthird arm of males hectocotylized; copulatory organwith clearly diVerentiated ligula and calamus. Ligulasimple, without transverse ridges. Arm tips nototherwise modiWed. Suckers small; distinct enlargedsuckers absent. Web deep to very deep. Gills with7–9 lamellae per demibranch. Ink sac present, anal

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 sper-matophoric gland; ss sperma-tophoric sac; t testes; to terminal organ; vd vas defer-ens. 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 ML51 mm. Scale bar represents 0.5 mm

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Table 2 Pareledone felix sp. nov: counts and measurements (mm) of male specimens

l left; r right; d damaged

Status Holotype Paratype Paratype Paratype Paratype Paratype Paratype

Repository NMSZ NMSZ NMSZ 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

Mantle length (dorsal)

42 41 39 41 44 44 43 45

Total length 115 115 115 116 116 124 127 144Mantle width 39 38 41 39 42 44 43 43Head width 31 33 35 31 32 35 32 34Full funnel

length20 17 17 17 19 20 23 23

Free funnel length

10 10 10 12 13 10 11 14

Funnel organ length

12 14 9 12 12 12 12 12

Pallial aperture 27 25 27 28 27 30 28 27Pallial aperture

(%)40 40 40 45 45 45 45 45

Web depth sector A

30 34 29 30 29 32 33 32

Web depth sector B (l/r)

30/30 34/32 31/30 28/34 32/34 35/36 33/38 36/30

Web depth sector C (l/r)

33/34 32/33 30/30 28/34 32/34 37/37 33/38 38/34

Web depth sector D (l/r)

33/32 31/31 32/31 26/31 32/34 34/37 33/35 35/30

Web depth sector E

26 31 27 26 D D 27 30

Arm length L1/R1

63/62 70/60 67/63 61/64 d/60 64/62 d/77 77/78

Arm length L2/R2

59/57 63/55 62/66 62/65 62/67 66/67 67/75 81/74

Arm length L3/Hc

61/60 54/55 63/62 64/63 64/62 65/66 66/73 85/67

Arm length L4/R4

64/66 55/62 64/63 66/70 67/72 67/72 77/73 85/84

Arm width 7 7 7 7 8 7 8 8Sucker diameter 3 2 2.5 2.5 2.5 3 3.5 3Sucker count

L3/Hc38/28 36/26 38/25 39/27 38/27 40/26 33/25 38/27

Gill length (l/r) 13/12 10/10 10/10 9/10 11/11 11/11 11/12 12/12Gill lamellae:

inner (l/r)8/8 8/8 8/8 8/8 8/8 8/8 8/7 9/8

Gill lamellae: outer (l/r)

8/8 9/8 8/8 8/8 8/8 8/8 7/7 8/8

Ligula length 6 8 8 7 8 6 7 7Calamus length 2 4 5 3 3 1 2.5 2Spermatophore

length61 70 70 67 67 55 66 68

No. of spermatophores 1+1 4+1 5+1 2+1 1+1 5+0 1+1 3+1

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Xaps present. Integument covered in small, widelyspaced papillae that appear as small, pale dots on themantle.

Description [based on eight mature male and sixmature female specimens for whom counts and measure-ments are given in Tables 2 and 3]: Animals small to

Table 3 Pareledone felix sp. nov: counts and measurements (mm) of female specimens

l left; r right; d damaged

Status Paratype 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

Mantle length (dorsal)

43 41 40 60 45 48

Total length 120 109 106 149 132 124Mantle width 44 41 44 58 47 40Head width 33 32 29 33 34 29Full funnel

length20 18 17 26 24 21

Free funnel length

12 10 8 16 14 9

Funnel organ length

12 12 9 16 17 14

Pallial aperture 28 28 28 29 33 26Pallial aperture (%) 45 45 45 45 45 45Web depth

sector A32 28 28 33 43 32

Web depth sector B (l/r)

34/33 28/28 31/29 37/38 42/41 32/34

Web depth sector C (l/r)

38/34 28/29 32/31 37/38 47/42 31/35

Web depth sector D (l/r)

35/34 28/29 31/31 34/37 44/45 33/35

Web depth sector E

33 26 30 30 38 29

Arm length L1/R1

59/56 57/55 56/52 d/70 68/74 62/63

Arm length L2/R2

61/59 57/57 57/60 75/75 75/72 62/59

Arm length L3/R3

64/60 59/58 55/60 71/75 75/75 62/65

Arm length L4/R4

66/65 62/61 56/61 76/74 76/79 64/66

Arm width 8 7 7 7 8 8Sucker diameter 3 3 3 4.5 4 3Sucker count

L3/R338/38 37/39 40/38 36/38 36/38 37/37

Gill length (l/r)

12/11 9/10 9/9 13/11 13/14 12/12

Gill lamellae: inner (l/r)

9/9 8/9 8/8 8/8 8/8 9/8

Gill lamellae: outer (l/r)

8/9 8/9 8/8 8/8 8/8 8/8

Egg length 14 12 10 22 12 9Egg width 4 4 3 8 4 4Number of eggs 27 22 31 54 42 39

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medium sized (ML 60 mm; TL 149 mm). Mantle spherical(MWI 98.1 § 6.6), head narrower than mantle (HWI74.3 § 8.3). Funnel medium sized (FuLI 45.8 § 4.1), gen-tly tapered; funnel organ W or VV shaped, usually VV.Arms short (MAI 62.4 § 7). Arm lengths approximatelyequal (ALI L1 150.4 § 14.8; L2 148.2 § 14.5; L3148.2 § 17.4; L4 154.1 § 17.5). Suckers uniserial, small(ASI 6.9 § 1), without sucker enlargement. Third rightarm of males hectocotylised, approximately equal inlength to opposite number (OAI 98.1 § 8.9). Ligulamedium 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). Hectocotylisedarm with 25–28 suckers, opposite arm with up to 40 suck-ers. Web deep to very deep (WDI 50.5 § 4.2), web sectorsapproximately equal in depth although some variation isapparent with sector E most commonly shallowest (butonly by a couple of mm) and sector A sometimes shal-lower than sectors B, C, and D. Ink sac present. Posteriorsalivary glands medium sized, slightly shorter than lengthof buccal mass (Fig. 3). Typical octopodid radula withnine elements (Fig. 5). Gills with 7–9 lamellae per innerdemibranch and 7–9 lamellae per outer demibranch. Ros-tral tip of lower beak rounded (Fig. 3). Mature ovarianeggs large, up to 22 mm, equivalent to an EgLI of 36.7with approximately 12 follicular folds (Fig. 4d). Penisdiverticulum coiled (Fig. 4b). Spermatophores long, up to70 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 togreen (Fig. 6). Papillae, which are scattered and wide-spread over the dorsal surface, are pale. The papillaeare small, and are not permanently raised as, for exam-ple, in P. charcoti. In dead specimens the integumentoften appears completely smooth, the pale dots beingthe 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, includingoV Elephant Island, south into the BransWeld Strait andnorth onto the slope of the Drake Passage. Found in awide range of depths from approximately 200 to 800 m.

Etymology: This name is in honor of Louise All-cock’s son Felix and Mike Vecchione’s father andgrandfather, both Felixes.

Discussion

Some initial diYculties were encountered in placingthis species. In cruise reports it has been referred to asPareledone (as P. sp. 1; Piatkowski et al. 1998: 44) and

to Megaleledone? (as M. sp. 17; Piatkowski et al. 2003:33). There are only three genera of shallow wateroctopodids in the Southern Ocean: Adelieledone,Pareledone and Megaleledone. All these taxa share cer-tain morphological features in common, for example,they have similarly sized uniserial suckers, a relativelysmall number of large eggs, and a large calamus. Someof the features commonly recorded in octopus taxon-omy are of no use in diVerentiating between these taxa.For example, in these taxa, the presence of an ink sacappears to be depth dependant. In the shallowerA. 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 spe-cies of papillated Pareledone from the Antarctic Penin-sula (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 ispresent, whereas in P. panchroma, which inhabitsdepths from about 450 to below 800 m, the ink sac isabsent (Allcock 2005). Other characters which are oflittle use include the funnel organ whose shape varieswithin taxa as well as between taxa, and characterssuch as relative arm length show a continuum of varia-tion between the taxa. A comparison of charactersbetween taxa (Table 4) shows, however, that there areclear diVerences between P. felix sp. nov and othergenera. Adelieledone diVers from all other taxa in that

Fig. 6 Pareledone felix sp. nov. Live animal

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it has no stylets, has very large posterior salivaryglands, has transverse ridges on its ligula, and has avery distinctive lower beak shape with a sharp rostraltip. P. felix diVers from Megaleledone in that the ligulain Megaleledone is very small (LLI 3–6 vs. LLI 9–15 inP. felix) as are the spermatophores (SpLI 40–60 vs.SpLI 120–180 in P. felix). The gill lamellae count inMegaleledone is much higher. Oddly, Megaleledone hasno anal Xaps (thought to be used in direct ink) despitethe presence of an ink sac. There is superWcial resem-blance between P. felix and Megaleledone; however, inthat they both have very deep webs (WDI 45–60 inP. felix, 40–60 in Megaleledone). In fact, juvenile Mega-leledone and P. felix (which are approximately equal insize) can be hard to tell apart without internal exami-nation if they are in poor condition. P. felix shares withother species of Pareledone similar beak morphologyand long spermatophores, and it aligns slightly moreclosely with the group of papillated Pareledone sharingcharacters such as a long ligula and similar body size atmaturity. The analysis of the molecular sequence data,however, allows conWdent placement of P. felix withinthis group.

Acknowledgments We would like to thank the Alfred WegenerInstitute for extensive sea time aboard RV Polarstern. Manythanks to Iain Barratt for information on egg counts in Adeliele-done and P. turqueti. JS is supported by a Natural EnvironmentResearch Council Antarctic Funding Initiative grant (NE/C506321/1) awarded to LA.

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Total length (mm)

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