Antarctic Science page 1 of 13 (2008) & Antarctic Science Ltd 2008

doi:10.1017/S095410200800151X

Check-list of the pycnogonids from Antarctic and sub-Antarctic waters: zoogeographic implications
S MUNILLA and ANNA SOLER MEMBRIVES TOMA
a, Universidad Auto noma de Barcelona, 08193 Bellaterra, Barcelona, Spain Unidad de Zoolog tomas.munilla@uab.es

Abstract: This study contains the current list of the austral pycnogonids together with details of their depth range and distribution. To date 264 species have been recorded, accounting for 19.6% of the 1344 species recorded worldwide. One hundred and eight species are endemic to Antarctic waters, 62 to the subAntarctic, 63 are common in both regions, and 55 are circumpolar. The richest genus is Nymphon, with 67 species and the richest area is the Scotia Sea. Comparing species lists between the years 2000 and 2007 shows that increased expeditions with more sampling has increased the circumpolarity of species and decreased zonal endemicity. The benthic insular refuge hypothesis is proposed as an explanation for the southern distribution of the present pycnogonid fauna, with an origin in the Scotia Arc.
Received 2 May 2007, accepted 13 March 2008

Key words: benthic insular refuge hypothesis, biogeography, circumpolarity, endemicity Introduction Pycnogonids (Chelicerata, Arthropoda) from Antarctic and sub-Antarctic waters have been studied more extensively than those from any other ocean of a similar size. Exploration of this area began with the American expedition of Nathaniel Palmer whose naturalist, James Eights (1835), described and drew the rst Antarctic pycnogonid Decolopoda australis on a serolid and some fossils. Pycnogonids have been collected by many scientists but the main monographs are by Hoek (1881), Mo bius (1902), Hodgson (1907, 1927), Bouvier (1913), Calman (1915), Gordon (1932, 1938, 1944), Fry & Hedgpeth (1969), Pushkin (1993) and Child (1994a, 1994b, 1995a, 1995b, 1995c). The most detailed information about the historical background of several families is contained in the Child papers, whilst the Pushkin monograph shows the geographical distribution of many species. The two latest Antarctic species to be described are Ammothe bigibossa Munilla, 2005 and Ammothea victoriae Cano & Lo pezGonzalez, 2007 from the Antarctic Peninsula and the Ross Sea respectively. The aim of this paper is to provide a complete up-to-date list of austral pycnogonids and, using zoogeographical information consider hypotheses describing their geographical distribution. Material and methods This paper is an analysis of published data. The works mentioned above, as well as the Mu ller catalogue (1993) and many others (Hoek 1898, Hodgson 1902, 1904, 1908, 1914, 1915, Bouvier 1905, 1906, Loman 1923, Calman 1933, Stephensen 1947, Hedgpeth 1950, Fage 1952a, 1952b, Utinomi 1959, Stock 1965, Arnaud 1972a, 1972b, Pushkin 1974, 1975a, 1975b, 1976, 1977, 1982, 1984a, 1984b, 1990, Turpaeva 1974, 1990, 1998, 2000, Krapp 1980, Child 1987, 1 1998, Pushkin 1988, Munilla 1989, 1991, 2000, 2001b, 2002, 2005, Stiboy-Risch 1992, 1993, 1994, Bamber 1995, 2007, Bamber et al. 2001, Chimenz & Gravina 2001), were used to compile the species list. The main zoogeographical works about pycnogonids are those of Fry (1964), Fry & Hedgpeth (1969), Hedgpeth (1969a, 1969b, 1971) and Munilla 2001a. Other works (Clarke & Crame 1989, 1997, Barnes & De Grave 2000, Clarke & Johnston 2003, Arntz et al. 2005, 2006, Barnes 2005, 2006, Clarke et al. 2005, Moyano 2005, Thatje et al. 2005, Gili et al. 2006, Linse et al. 2006), contain particular zoogeographical reviews of some zoological groups or global and evolutionary reviews of benthos. A total of 98 papers were consulted.

Results and discussion Historical research and specic richness This analysis of the austral pycnogonids covers 172 years, 16 countries and more than 42 ships or expeditions. So far 40 000 specimens have been found in the Antarctic and sub-Antarctic waters, (termed the Austral Ocean by Jacques & Treguer 1986, p. 133) at about 2100 sampled stations. The Southern Ocean usually contains the Antarctic waters south of the Polar Front and does not include sub-Antarctic localities between this front and the Subtropical Convergence (D. Barnes, personal communication 2007). This sub-Antarctic zone contains many islands with pycnogonids (see legend, Table I) and the South American zone (Magellan and Falkland zones). Bouvet Island is just south of the Polar Front, but given its isolation and that it acts as a link between sub-Antarctic and Antarctic fauna (Arntz et al. 2006), I consider it as another sub-Antarctic island. Moreover, the general composition of the actual

S MUNILLA & ANNA SOLER MEMBRIVES TOMA

Table I. Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species Achelia A. assimilis (Haswell, 1875) A. communis (Bouvier, 1906) A. dorhni (Thompson, 1884) A. hoekii (Pfeffer, 1889) A. lagena Child, 1994a A. megacephala (Hodgson, 1915) A. parvula (Loman, 1923) A. quadridentata (Hodgson, 1910) A. serratipalpis (Bouvier, 1911) A. spicata (Hodgson, 1915) A. sufata Gordon, 1944 A. transfuga Stock, 1954 Ammothea A. adunca Child, 1994a A. allopodes Fry & Hedgpeth, 1969 A. antipodensis Clark, 1972 A. armentis Child, 1994a A. bentartica Munilla, 2001 A. bigibbosa Munilla, 2005 A. calmani Gordon, 1932 A. carolinensis Leach, 1814 A. clausi Pfeffer, 1889 A. cooki Child, 1987 A. dubia (Hedgpeth, 1950) A. gibbosa Bouvier, 1913 A. gigantea Gordon, 1932 A. glacialis (Hodgson, 1907) A. gordonae Child, 1994a) A. hesperidensis Munilla, 2000 A. longispina Gordon, 1932 A. meridionalis (Hodgson, 1915) A. minor (Hodgson, 1907) A. sextarticulata Munilla, 1989 A. spinosa Hodgson, 1907 A. striata Mo bius, 1902 A. stylirostris Gordon, 1932 A. tetrapora Gordon, 1932 A. tibialis Munilla, 2002 A. victoriae Cano & Lo pez, 2007 Ascorhynchus A. antipodus Child, 1987 A. cooki Child, 1987 A. cuculus Fry & Hedgpeth, 1969 A. hedgpheti Turpaeva, 1974 A. inatum Stock, 1963 A. ornatum (Helfer, 1938) 75 6 (8.0) 5340 14632992 9934008 37003910 27436070 90108 n n Sc Sc Sc k also in Argentine Basin W. sp. 80 A S sp. & % of W.sp 12 (15.4) 0 903 0 714 0 192 5 256 23137 shallow water 0 267 0 21 64361 0 1138 0 300 2 10 40 25 (62.5) 185800 2102000 0 24 230380 167335 517 991408 3 670 3 860 14632992 106 439567 991116 0 640 348732 30439 571454 10454 8 473 5 516 761679 72567 165494 105303 710 360366 w C k b n k Sc p Sc,p,b,r,w C C r b C C Sc,r Sc C C C C Sc,p,w,r C Sc,w,p Sc Sc r Livingston. Island, mud C Sc,p,b s,n s,t n s s k s St.Pl A also west Pacic and Australia A. brucei Calman, 1915 Depth (m) Antarctic zone sub-Antarctic zone Remarks and synonyms

also in Peru-Ecuador-Argentina

Sc,p,b,e. C w,e n

also on Angola coast described as Austrothea e: 40100E muddy bottom Magnammothea and Biammothea A. bicorniculata Stiboy-Risch, 1992

b s n described as Boehmia recorded as Colossendeis gibbosa Mo bius, 1902 145E178W. Magnammothea gigantea Fry & Hedgpeth, 1969

Livingston. Island, mud s

s b s s

also on South Georgia, Biammothea brevipalpa Puskin, 1993 also in Argentine Basin

also in PeruChile trench, South Africa, Kurile trench also in South Africa Continued

CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species Austroraptus A. calcaratus Gordon, 1944 A. juvenilis Calman, 1915 A. polaris Hodgson, 1907 A. praecox Calman, 1915 A. sicarius Fry & Hedgpeth, 1969 Cilunculus C. acanthus Fry & Hedgpeth, 1969 C. cactoides Fry & Hedgpeth, 1969 C. kravcovi Pushkin, 1973 C. spinicristus Child, 1987 Dromedopycnon D. acanthus Child, 1982 Eurycyde E. antarctica Child, 1987 Sericosura S. mitrata (Gordon, 1944) Tanystylum T. antipodum Clark, 1977 T. brevicaudatum (Fage & Stock, 1966) T. brevipes (Hoek, 1881) T. bueroisi Arnaud, 1974 T. cavidorsum Stock, 1957 T. pfefferi Loman, 1923 T. neorhetum Marcus, 1940 T. oedinotum Loman, 1923 T. ornatum Flynn, 1928 T. styligerum Myers, 1875 Austrodecus A. breviceps Gordon, 1938 A. calcaricauda Stock, 1957 A. cestum Child, 1994b A. crenatum Child, 1994b A. curtipes Stock, 1957 A. elegans Stock, 1957 A. fagei Stock, 1957 A. fryi Child, 1994b A. glabrum Stock, 1957 A. glaciale Hodgson, 1907 A. goughense Stock, 1957 A. kelpi Pushkin, 1977 A. longispinum Stock, 1957 A. macrum Child, 1994b A. profundum Stock, 1957 A. pushkini Child, 1994b A. serratum Child, 1994b A. simulans Stock, 1957 A. sinuatum Stock, 1957 A. tristanense Stock, 1955 A. varum Child, 1994b Pantopipetta W. sp. 5 A S sp. & % of W.sp 5 (100) 143 219 3 500 10569 6 260 220 380 31 4 (12.9) 24402818 38540 255 309 476 540 2 21 6 45 1 (50) 124 903 1 (4.8) 527 714 1 (16.7) 106 2154 10 (22.2) shallow water 0 15 45100 80100 0 245 2 100 0 410 0 183 46560 0 200 42 22 (52.4) 0 298 731373 86207 1 360 0 903 99606 263400 112 859 18277 0 2100 42120 shallow water 91325 14422350 920 60903 79124 91545 shallow water 0 70 443 549 20 4 (20.0) Continued Sc Sc,p Sc,p Sc,w C n Sc C St.P A Sc k r p s n k n M PE,t n mud & stones, Graham Land southern Argentina Macquarie Island also in Gough Island among kelps k,n s n s,k M PE n St.P A p,St A St.P A. s,M PE,c,n s,k,n,t s,k M P.E s,k,n also in Cape Verde Island also in South Africa also in southern Chile T. dohrni Schimkewitsch, 1889 r also in Walvis Ridge (South Africa) r s also in Brazilian slope b Sc,p,e n c,M PE n also in Argentine Basin r,e C Sc,p,b,r,e C Sc,r e: 65E e: 92E, 145E e: 78E. Possibly C Depth (m) Antarctic zone sub-Antarctic zone Remarks and synonyms

Sc Sc Sc

b,w

Macquarie Island

S MUNILLA & ANNA SOLER MEMBRIVES TOMA

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species P. P. P. P. australis (Hodgdon 1914) buccina Child 1994b lata Stock 1981 longituberculata Turpaeva, 1985 11 10 (90.9) 85920 3 280 1632955 3 1180 1042100 45645 12001280 50580 5801180 45550 35 7 6 1 (2.9) 73578 1 (14.3) 5813777 3 (50) 3 415 1122612 1586 16 2 (12.5) 250 320 6 86 1 (16.7) 3 30 18 (20.9) 35383 104830 0 430 65600 1461530 115260 22732421 117430 15275 12283060 1001138 0 400 3 4540 Sc Sc,w,p,e Sc Sc w Sc,p r Sc Sc,w,e Sc Sc,p,r,w C s also in waters of Uruguay, Brazil and Argentine e: 69S 14E also in Surinam & W. Atlantic k Sc c k,n n n P,w p s also at 23S (off Brazil) C Sc,e C C C C Sc C e Sc,r A. spicata Hodgson, 1915 Pseudopallene brachyura Bouvier, 1911 holotype without gures b,c W. sp. A S sp. & % of W.sp Depth (m) 6805340 31933423 5233655 5676700 Antarctic zone Sc,r Sc b,w Sc sub-Antarctic zone M PE,n also in Concepcio n waters, Chile also in Cape Basin, South Africa also in SW Africa, Pacic & all Atlantic P. brevicauda Stock, 1963 Austropallene A. brachyura Bouvier, 1913 A. A. A. A. A. A. A. A. A. bucera Pushkin, 1993 calmani Gordon, 1944 cornigera (Mo bius, 1902) cristata Bouvier, 1911 gracilipes Gordon, 1944 spinicornis Pushkin, 1993 tcherniai Fage, 1952 tenuicornis Pushkin, 1993 tibicina Calman, 1915 Remarks and synonyms

holotype without gures holotype without gures n

Callipallene C. margarita Gordon, 1932 Cheilopallene Ch. gigantea Child, 1987 Oropallene O. dimorpha (Hoek, 1898) O. dolichodera Child, 1995c O. metacaula Child, 1995c Pseudopallene P. centrotus Pushkin, 1990 P. glutus Pushkin, 1975 Seguapallene S. insignatus Pushkin, 1975 Pallenopsis P. boehmi Schimkewitsch, 1930 P. P. P. P. P. P. P. P. P. P. P. P. buphthalmus Pushkin, 1993 candidoi Mello-Leitao, 1949 gurjanovi Pushkin, 1993 kupei Clark, 1971 latefrontalis Pushkin, 1993 lateralia Child, 1995c lattina Pushkin, 1993 leiopus Pushkin, 1993 longiseta Turpaeva, 1958 macronix Bouvier, 1911 oblicua Thompson, 1884 patagonica (Hoek, 1881)

Holoptype with gures n n s also Subarctic & Gulf of Panama P. knipovitchi Turpaeva, 1974 rock & algal bottoms described as Phoxichilidium patagonicum P. hiemalis Hodgson, 1907 P. glabra Mo bius, 1902 P. mo biusi Pushkin, 1975 P. meridionalis Pushkin, 1975 P. hodgsoni Gordon, 1938 Cheilopallene spicata Stock, 1955, Clavigeropallene spicata Pushkin, 1974 also in Uruguay, Brazil, Argentina P. gau ssiana Hodgson, 1914 Continued

P. pilosa (Hoek, 1881) P. spicata Hodgson, 1914 P. tumidula Loman, 1923 P. vanho ffeni Hodgson, 1915

253650 25549 42270 3 889

C C Sc C

CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species W. sp. A S sp. & % of W.sp Depth (m) Antarctic zone sub-Antarctic zone Remarks and synonyms P. setigera Hodgson, 1914 P. villosa Hodgson, 1907 Colossendeis C. adelpha Child, 1998 C. angusta Sars, 1877 C. arundorostris (Fry & Hedpeth, 1969) C. australis Hodgson, 1907 C. avidus Pushkin, 1970 C. belekurovi Pushkin, 1993 C. brevirostris Child, 1995b C. colossea Wilson, 1881 C. concedis Child, 1995b C. drakei Calman, 1915 C. elephantis Child, 1995b C. enigmatica Turpaeva, 1974 C. ensifer Child, 1995b C. fragilis Pushkin, 1992 C. grassus Pushkin, 1993 C. hoecki Gordon, 1944 C. insolitus Pushkin, 1993 C. korotkevischi Pushkin, 1984 C. kurtchatovi Turpaeva, 1993 C. leniensis Pushkin, 1993 C. lepthorynchus Hoek, 1881 C. longirostris Gordon, 1932 C. macerrima Wilson, 1881 C. media Hoek, 1881 C. megalonix Hoek, 1881 75 36(48.0) 333 341 225480 610 153935 270 426 150 377 5449604 425 4140 22482907 3 3000 23844795 315 335 32503285 3 830 315 435 120 3112 132 660 4700 250 432 561 3675 2 3700 20102100 33865798 7 4900 e Sc r C p,w,e a Sc,w Sc,r C Sc,e Sc Sc Sc,e Sc Sc,w,r w k c cosmopolitan s also in southern Tasmania C. smirnovi Pushkin, 1988 also in Chile & Argentina C. acuta Stiboy-Rich, 1993 Prydz Bay cosmopolitan C. gracilis Hoek, 1881 160 2804 C

s k,n s k,c s 52S 43E M PE,s n n

47S, Argentina

Sc C s,k,n

C. mica Pushkin, 1970 C. notialis Child, 1995b C. pseudochelata Pushkin, 1993 C. robusta Hoek, 1881 C. glacialis Hodgson, 1907 C. scoresbii Calman, 1915 C. scotti Calman, 1915 C. stramendi Fry & Hedgpeth, 1969 C. tenuipedis Pushkin, 1993 C. tethya Turpaeva, 1974 C. tortipalpis Gordon, 1932 C. wilsoni Calman, 1915 Decolopoda D. australis Eights, 1835 D. quasami Sree et al., 1993 2 2 (100)

1400 260 380 125 180 0 3610

378S 228E k Sc,e C b,k

south of Iles Crozet cosmopolitan C. pennata Pushkin, 1970 also in southern Tasmania cosmopolitan. C. japonica, Hoek, 1898, C. spei also in Argentina C. brevipes Hoek, 1881 also in South Africa, Madagascar & E. Argentina C. rugosa Hodgson, 1907 C. frigida Hodgson, 1907 C. orcadense Stock, 1963 Sub-Antarctic Indian Ocean e: 69S, 11E C. lilliei Calman, 1915 C. gracillipes Bouvier, 1911 C. rostrata Turpaeva, 1994

130 5227 35352 645 3806 250 860 318 160 4026 60801 0 1890 150

Sc,w,r C

s s

C Sc,w C Sc,w,r Sc,p,b,w,r e

s s,k

also in E. Argentina

also in Ade lie Land k possibly C D. antarctica Bouvier, 1905

Continued

S MUNILLA & ANNA SOLER MEMBRIVES TOMA

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species Dodecolopoda D. mawsoni Calman & Gordon, 1933 Heteronymphon H. exiguum (Hodgson, 1927) Nymphon N. adareanum Hodgson, 1907 N. andriashevi Puskin, 1993 N. arcuatum Child, 1995a N. articulare Hodgson, 1908 N. australe Hodgson, 1902 N. stylops Bouvier, 1913 N. biarticulatum Hodgson, 1907 N. bouvieri Gordon, 1932 N. brachyrhynchum Hoek, 1881 N. brevicaudatum Miers, 1875 N. bucuspidum Child, 1995a N. chaetodir Utinomi, 1971 N. charcoti Bouvier, 1911 N. clarencei Gordon, 1932 N. compactum Hoek, 1881 N. eltaninae Child, 1995a N. forticulum Child, 1995a N. frigidum Hodgson, 1907 N. galatheae Fage, 1956 N. gerlachei Giltay, 1937 N. glabrum Child, 1995a N. gracilipes Miers, 1875 N. gruzovi Pushkin, 1993 N. hadale Child, 1982 N. hamatum Hoek, 1881 N. hiemale Hodgson, 1907 N. inferum Child, 1995a N. inornatum Child, 1995a N. isabellae Turpaeva, 2000 N. isaenki Pushkin, 1993 N. lanare Hodgson, 1907 N. lomani Gordon, 1944 N. longicolum Hoek, 1881 N. longicoxa Hoek, 1881 N. longisetosum Hodgson, 1915 N. macquarensis Child, 1995a N. macrochelatum Pushkin, 1993 N. mendosum (Hodgson, 1907) N. microgracilipes Pushkin, 1993 N. monotrix Child, 1995a N. multidens Gordon, 1944 N. multituberculatum Gordon, 1944 N. neelovi Pushkin, 1993 N. neumayeri Gordon, 1932 N. orcadense (Hodgson, 1908) W. sp. 1 A S sp. & % of W.sp 1 (100) 160549 7 268 1 (14.3) 3 415 67 (25.0) 1 903 116135 38157 18910 8 4136 Sc,p,r,e p Sc,p,w C s s s s,n,b C n described as Nymphon Chaetonymphon also in SE Argentina, possibly C C Depth (m) Antarctic zone sub-Antarctic zone Remarks and synonyms

also Indian Ocean & off ArgentineChilean coasts

35889 158583 82430 271100 1262 9951110 3 1200 65342 7313246 4671233 438548 227 31115798 460578 55 3 3055 250 30105798 25023400 301435 24503873 513 333571 500700 60848 112714 684600 3182998 3852450 112124 540 15555 150309 34953514 40260 180640 65240 160403 18163

C Sc,p,e C

k k k n n also in Heard Island

C Sc Sc Sc,r r w p,b Sc w,e p,w Sc Sc C Sc, Palmer Is. w w C C C Sc,b,r e 68S 32E,w C B r Sc,p,b w,Sc,e Sc Sc,p Sc,p

n s

also in South Africa s: SE Argentine

also in Kermadec Trench, (SW Pacic Ocean) holotype is a juvenile also in Argentine basin N. gracillimum Calman, 1915

k n n n n

also in Chilean Basin & New Zealand also in Argentine Basin

c: 46S 49E

b e: 20140E c s s Continued

CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species N. pagophylum Child, 1995a N. paucidens Gordon, 1932 N. paucituberculatum Gordon, 1944 N. pfefferi Loman, 1923 N. phasmatoides Bo hm, 1879 N. polare Hodgson, 1915 N. premordicum Child, 1995a N. proceroides Bouvier, 1913 N. procerum Hoek, 1881 N. proximum Calman, 1915 N. pseudogracilipes Pushkin, 1993 N. punctum Child, 1995a N. rybakovi Pushkin, 1993 N. sabellum Child, 1995 N. scotiae Stock, 1981 N. subtile Loman, 1923 N. tenuimanum Hodgson, 1914 N. tenuipes Bouvier, 1911 N. trituberculum Child, 1995a N. typhops (Hodgson, 1915) N. unguiculatum Hodgson, 1927 N. villosum (Hodgson, 1915) N. zundiamum Pushkin, 1993 Pentanymphon P. antarcticum Hodgson, 1904 Sexanymphon S. mirabilis Hedgpeth & Fry, 1964 Anoplodactylus A. australis (Hodgson, 1914) A. californicus Hall, 1912 A. laciniosus Child, 1995c A. laminifer Arnaud, 1974 A. petiolatus (Kro yer, 1844) A. speculus Child, 1995c A. typhlops Sars, 1888 1 1 1 (100) 3 3227 1 (100) 16872897 140 9 (6.4) 15616 0 100 456 540 80100 0 1180 15861640 915 3620 C t s n St.P A s n c,n,PE also in Tasmania cosmopolitan; C. projectus Hilton, 1942, C. portus Sawaya 1950 also in Antipodes Island also in Atlantic Mediterranean cosmopolitan A. pelagicus Flynn, 1908 A. neglectus Hoek, 1898 also in Atlantic Mediterranean Sc,p C P. minutum Gordon, 1944 W. sp. A S sp. & % of W.sp Depth (m) 265 1170 22334 180 219 12250 1394 350 25973215 911180 24506135 401555 195 216 415 220 28722928 29602980 13304 19033398 122 1180 32003259 24502815 168 450 13636 160 Antarctic zone Sc,p,w Sc w,e Sc p,w e Sc Sc,p,e Sc C k n Sc,w r Sc s,k r C e Sc,p,w,e Sc C s 62S160W N. stocki Turpaeva, 1974 also SE Argentine also in S. Australia N. soyae Utinomi, 1953 sub-Antarctic zone c Remarks and synonyms

also SE Argentina

s also in South Africa; P. capense Hodgson, 1908

cosmopolitan N. banzare Gordon, 1944

also 10E s: near Falkland Islands

Sc

A. vemae Child, 1982 A. virescens (Hodge, 1864) Phoxichilidium P. pigordum Child, 1995c Endeis E. australis (Hodgson, 1907) E. viridis Pushkin, 1976 Pentapycnon P. bouvieri Puskin, 1993 P. charcoti Bouvier, 1910 Pycnogonum 69 10 (14.5) 14 17 1 (7.1)

90676 0 16 79124 2 (11.8) 3 1570 3 377 3 2 (66) 90419 240 1420 Sc,w Sc,p,r C

s St.P A n n,b k,c,M PE

P. bouvieri Child (1993) P. magnum Stiboy-Rish (1994)

Continued

S MUNILLA & ANNA SOLER MEMBRIVES TOMA

Table I. (Continued) Genera and species recorded in Antarctic (A) and sub-Antarctic (S) waters with regard to their depth and various geographical zones. W.sp number of world species. Antarctic waters: C circumpolar, Sc Scotia Sea, p Antarctic Peninsula, a Amundsen Sea, b Bellingshausen Sea, r Ross Sea, w Weddell Sea, e East Antarctic zone. sub-Antarctic waters: s South America, k Iles Kerguelen, c Ile Crozet, n New Zealand Plateau, t Tristan da Cunha, b Bouvet Is., M PE Marion & Prince Edwards Is., St.P A Saint Paul & Amsterdam Is. Genera/species P. P. P. P. P. P. P. P. P. calculum Bamber, 1955 rhinoceros Loman, 1923 gaini Bouvier, 1910 gordonae Pushkin, 1984 magellanicum Hoek, 1898 magniroste Mo bius, 1902 paragaini Munilla, 1989 platylophum Loman, 1923 sivertseni Stock, 1955 19 4 (21.1) 60900 100140 0 101 0 77 C s,k,n k S,n s described as Austrodecus also in Mexico also in Caribbean Sea, E. Pacic Ocean, Azores, Mediterranean Sea W. sp. A S sp. & % of W.sp Depth (m) littoral 301115 242495 219400 85548 3 309 205440 0 903 102141 Antarctic zone Sc C C w,e sub-Antarctic zone s k s k,c Sc Sc,e k,c t Remarks and synonyms rock with algae P. diceros Marcus, 1940

Rhynchothorax R. australis Hodgson, 1907 R. oblongus (Pushkin, 1977) R. percivali Clark, 1976 R. philopsammum Hedgpeth, 1951 Total species

264

192

137

Bouvet fauna is more similar to the Magellan area than the high Antarctic region (Arntz et al. 2006). These individuals belong to 31 genera and 264 different species of pycnogonids (Table I), out of a total number of species worldwide of 1344. They thus represent 19.6% of the actual world species that have been recorded in 21% of the ocean areas (Jacques & Treguer 1986). Figure 1 shows the richness of species and genera for each family. Nymphonidae is the most abundant family (71 species), with Nymphon the major genus (67 species), and N. australe the most frequently recorded species. Of these 264 species, 108 are endemic in the Antarctic area, 62 are present only in the sub-Antarctic zone and 63 are common to both. Table I shows the current list of the Antarctic and sub-Antarctic species, their synonyms (47), the percentage

of species per genus with respect to their number worldwide, their geographical distribution and bathymetric range. The majority of synonyms have been proposed or recorded in the Child (1994a, 1994b, 1995a, 1995b, 1995c), Pushkin (1993) and Mu ller (1993) papers. The richest zone, with 89 recorded non-circumpolar species, is the Scotia Sea, followed by the sub-Antarctic islands with 64 species. The circumpolarity criterion is that a species
Table II. Variation in the austral pycnogonid species between 2000 and 2007. Differences between seven years Species in the world Species in the Southern Ocean (sensu lato) Species reported in Antarctic waters Species reported in sub-Antarctic waters Endemic species in Antarctic waters Endemic species in sub-Antarctic waters Common species Circumpolar species Circumpolar genera Cosmopolitan species Endemicity of species from Antarctic zones Scotia Sea Antarctic Peninsula Bellingshausen Sea Amundsen Sea Ross Sea Weddell Sea East Antarctica Total Endemicity of species from sub-Antarctic zones South America (Magellan region) New Zealand Plateau sub-Antarctic Islands Bouvet Island 2000 1165 251 180 131 101 59 60 45 13 5 2000 26 4 1 1 8 4 8 52 10 21 24 2007 1344 264 192 138 108 62 63 55 15 7 2007 22 3 0 1 9 3 4 42 10 23 24

Fig. 1. Richness of species and genera from austral pycnogonid families.

CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS

was recorded on one or more occasions in each of the waters to the north, south, east and west of the Antarctic continent, with the east zone the largest one and least sampled (it contains 24 species of which only four are endemic).

Zoogeography The Antarctic benthos has evolved as a consequence both of the abiotic environmental conditions in the past and of biotic interactions (Arntz et al. 1994). The distribution of most of the benthic Antarctic fauna is considered as circumpolar (Hedgpeth 1971, Arntz & Gallardo 1994, Clarke & Crame 1997), almost certainly due to the powerful Antarctic Circumpolar Current (Clarke & Johnston 2003). The circumantarctic element is also the most frequent pattern for pycnogonids (Fry & Hegdpeth 1969, Hegdpeth 1969a, Munilla 2001a), since at present 55 of the 192 (28.7%) Antarctic recorded species are circumpolar. The comparative data in Table II, based on cruise data and literature dealing with austral pycnogonids over a period of seven years, shows that the number of endemic species for each Antarctic zone is low, with the exception of the Scotia Sea, which could be considered as a sub-centre of speciation. Moreover, this table shows that the circumantarctic pattern for the pycnogonid species has increased over the seven years, and the endemicity of the species from each zone has consequently decreased. In other words, increased sampling has shown more circumpolarity and less zonal endemicity. Only 10 genera are exclusively from austral waters and four of them (Dodecolopoda, Pentanymphon, Sexanymphon, Austroraptus) are endemic to Antarctic waters. Other genera (Decolopoda, Austropallene) which were considered by Hegdpeth (1969b) as typically Antarctic, have already been found in sub-Antarctic waters, including in the Kuril Islands (Austropallene likinii Turpaeva, 2002); the same is true for some species. The genera with the most species in austral

waters are: Ammothea (25 species out of 40 in the world; 62.5%), Colossendeis (36 out of 75; 48%), Austrodecus (22 out of 42; 52.4%), Nymphon (67 out of 268; 25.0%) and Pallenopsis (18 out of 86; 20.9%). There are not endemic families. The sub-Antarctic pycnogonid fauna shows origins in the Antarctic fauna at genus level (Arnaud & Bamber 1987). For example, this is true of Colossendeis and Ammothea, two genera with more than half of their species in the Austral Ocean. Like other genera with abundant species, both have more species in Antarctic than in sub-Antarctic waters. We therefore view the Southern Ocean as a centre of speciation (suggested by Hedgpeth 1969b, and Munilla 2001a) but also of geographic dispersion and evolutive radiation, because of its high relative endemicity (108 Antarctic species versus 62 of sub-Antarctic ones). The dendrogram (Fig. 2), based on the presence-absence data of the 264 austral species, shows that the Antarctic species form a large zoogeographic group linked to circumpolarity (55 species). Three trends are clear: a) The Scotia Sea is closely linked to other southern zones (60.75% of similarity), indicating some peculiarity of the former. b) Two branches of the Circumpolar Current (with 71% of similarity) have been differentiated: the north-eastern (Antarctic Peninsula Weddell-East zones) and the southern (Bellingshausen Ross zones). This supports the geographical proximity of the species distribution to the direction of the Circumpolar Current. c) Each sub-Antarctic zone is separate, and the three zones (Magellan region, New Zealand and sub-Antarctic islands), present low levels of similarity ( , 30%, Fig. 2) to Antarctic waters. The suites of organisms in the seas surrounding the sub-Antarctic islands, have long been considered sufciently dissimilar to

Fig. 2. Similarity of Antarctic and subAntarctic zones (Bray-Curtis Index, complete linkage), based in presenceabsence data. CircumAntarctic species are included. n New Zealand Plateau, s South America, s.Is subAntarctic islands, Sc Scotia Sea, R Ross Sea, B Bellingshausen Sea, Pn Antarctic Peninsula, W Weddell Sea, E East zone.

10

S MUNILLA & ANNA SOLER MEMBRIVES TOMA

constitute a different zone (Hedgepth 1969a), with 63 species at present. Recent multivariate analysis of the bryozoan component of benthos south of 47S supports the categorization of Patagonia, the subAntarctic islands and Antarctica into separate zones (Barnes & De Grave 2000). Two complementary hypotheses (Clarke & Johnston 2003) may explain the possible origin of todays Antarctic benthic fauna: 1. It comes from an in situ stock in Cretaceous waters (141 65 Ma), from the coastal fauna of Gondwana, when the present Antarctic continent was part of the supercontinent. This is supported by the Gasteropoda (Clarke 1990), two Isopoda families (Brandt 1992) and some sessile groups (Alcyonarians and sponges, Gili et al. 2006) among others. 2. There was subsequent interchange with the deep fauna of the contiguous oceans, as is the case with Tanaidacea and Amphypoda (Brandt 1999). One possibility is that the Magellan region provided Antarctic benthic fauna across the Drake Passage or the Scotia Arc. This may be the case in some groups such as Serolidae (Held 2000), Polychaeta (Montiel et al. 2004) and Bryozoa (Moyano 2005, Barnes 2006). The Scotia Arc and Bouvet Island are clearly undersampled, if they are considered as transitory areas between the Magellan region and the Antarctic Peninsula or the Weddell Sea (Arntz et al. 2005). Modern Antarctic communities are thus composed of a mixture of Palaeozoic taxa, which migrated from the deep ocean during interglacial periods, and a component of fauna that evolved from common Gondwana Cretaceous ancestors (Gili et al. 2006). The nal connection between South America and the Antarctica was broken just over 25 Ma ago. The result was the formation of the Circumpolar Current, causing the oceanographic and geographical isolation of the Antarctic continent. The continental remains of the ancient isthmus today form many of the islands adjacent to the Antarctic

Table IV. Number of species recorded in each zone (in bold) and common species between different zones. Circumpolar species are excluded. S s Sc Pn R W B E n s.Is 46 14 10 3 5 1 1 8 10 Sc 14 89 24 18 20 8 16 11 12 Pn 10 24 35 7 14 5 8 2 3 R 3 18 7 27 8 3 4 4 3 W 5 20 14 8 35 4 11 2 7 B 1 8 5 3 4 12 2 1 3 E 1 16 8 4 11 2 24 0 2 n 8 11 2 4 2 1 0 47 11 s.Is. 10 12 3 3 7 3 2 11 64 Tot 52 123 73 50 71 27 44 39 49

s South America, Sc Scotia Sea, Pn Antarctic Peninsula, R Ross Sea, W Weddell Sea, B Bellingshausen Sea, E East zone, n New Zealand Plateau, s.Is sub-Antarctic islands. Tot sum of the common species for each zone and the remaining ones.

Peninsula, but the main topographical obstacle between South America and Antarctica is the Scotia Arc, through which the Circumpolar Current passes. Two main probable dispersion routes of pycnogonids are proposed, coming from the ancient Cretaceous fauna and principally along the bottom (since they have benthic larvae): 1. From Western Antarctica to the Eastern zone by means of the Circumpolar Current. In support of this, there are 15 common non-circumpolar species between the Scotia ArcAntarctic Peninsula couplet and the Eastern Antarctic zone. Moreover, a branch of the dendrogram (PnWE, Fig. 2) also supports this suggestion. 2. From South America to western Antarctica going along the route from the Scotia Arc. This is supported by 17 common non-circumpolar species between the Scotia Arc Antarctic Peninsula couplet and the Magellan zone; moreover, 13 of 48 austral species recorded in other waters have been also found in the Argentine and Brazilian zones (the Brazil current versus circumpolar one).

The benthic insular refuge hypothesis The submerged zones of the Peter I and Thurston islands in the Bellingshausen Sea are sheltered areas and optimal zones for the settlement and protection of pycnogonid fauna from the surrounding open seas, similar to oases in the desert. They are species rich and the densities of animals are here higher than on open bottoms (Table III), probably because of more feeding possibilities. Animals can be transported from them by deep currents, including by hitchhiking on a moving animal or debris or by simply drifting to other waters. This trend, observed in the Bellingshausen Sea, is extensive in other waters, and occurs in Bouvet Island, which act as stepping stones (Arntz et al. 2006) to Antarctic waters.

Table III. Specimens and species of pycnogonids from islands and the open Bellingshausen Sea, in the Bentart-03 Cruise. Zones Bentart-03 Stations (st.) Latitude S n S n/S mean S/st. mean n/st. mean Depth (m) Thurston Peter I islands 8 6870 187 13 14.4 1.6 23.4 86726 Bellingshausen Sea 9 6870 12 10 1.2 1.1 1.3 492 1947

n number of specimens, S number of species

CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS

11

The benthic insular refuge hypothesis suggests that the islands serve as a home, accumulating a rich benthic fauna, and subsequently acting as migration points. This is similar to the reserve effect of a marine protected zone. This theory needs to be conrmed using more quantitative data about species richness, densities and biomass in pycnogonids and many other zoological groups. Far from any island, deep waters of under 1000 m, have few species and specimens (Munilla 2001a). Moreover, no circumpolar species have been recorded (Table IV) and the number of endemic species (Table II) in each austral zone is much more important at islands (Scotia Sea, New Zealand Plateau and sub-Antarctic ones) than at the benthic bottoms of the open seas. There also seems to be no decreasing latitudinal decline of species richness if the sub-Antarctic islands are included (Table II and IV), because there are more species in Antarctic waters than in sub-Antarctic ones.

large number of common species between the Scotia Sea Arc and remaining zones (123, Table IV) support this movement of pycnogonids. The low water temperature is the main factor that isolates the Antarctic species, leading to high endemism in various groups (108 of 192, 56.3% in pycnogonids). With the warming of global oceans, the colonization of the Antarctic waters will be greater in the future. The number of species will increase and the relative Antarctic endemicity will decrease. This trend will be favoured by the increasing passive transport of animals or algae on various ships, swimming animals and oating debris. Eddies and currents are factors that also contribute to passive transport (Barnes et al. 2006).

Acknowledgements This work was carried out under the auspices of two MEC grants (CGL 2004 01856/ANT, REN 2001-1074/ANT) of the Spanish Council of Scientic and Technical research. I would like to express my gratitude to the scientic colleagues of the Bentart cruises aboard the Hesperides oceanographic Spanish vessel. Likewise, we are grateful to the reviewers and the Editor for their constructive suggestions which improved the paper.

Possible stages in the origin and dispersion of the Antarctic pycnogonids 1. In situ origin (Munilla 2001a), from the Cretaceous Gondwana fauna (141 65 Ma). This possibility is supported by the two most ancient families of the sea spiders (Colossendeidae and Austrodecidae, (Arango & Wheeler 2007, Bamber 2007), morphological and molecular data, having 48% and 42% respectively of their species in southern waters. This hypothesis has been suggested previously for Austrodecidae genera (Stock 1957, Child 1995b). 2. Many archipelagos in the Scotia Arc are the tips of an almost continuous subsurface mountain chain linking the Andes and the Antarctic Peninsula (Barnes 2005). All the Scotia Sea islands sheltered the existing fauna at the time of its creation and they still retain the ancient Cretaceous fauna that the Antarctic Circumpolar Current (ACC) subsequently carries, because the Scotia Arc is the only major barrier to the circulation of this current. Many more species remain on its islands than in other waters, as shown by the large number of species that have been captured in this zone (Table IV): 89 non-circumpolar species plus 55 circumpolar ones, that is the 75% of the 192 recorded Antarctic species. 3. From the Scotia Arc waters, the fauna was, and today still is, actually exported towards the East Antarctic zone thanks to the Circumpolar Current, which also distributes some species to the sub-Antarctic islands. A similar trend happens with the benthic larvae of cheilostome bryozoa (Bouvet Island, Barnes 2006). Moreover, the Scotia Arc pycnogonid fauna also arrive at the southern branch of the Circumpolar Current (Bellinghausen Ross Sea, Fig 2). This link is shown in the cluster in Fig. 2, where the Scotia Sea branch is closely related to other Antarctic zones. The

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