Check-List of the Pycnogonids from Antarctic and Sub-Antarctic Waters

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Check-List of the Pycnogonids from Antarctic and Sub-Antarctic Waters 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 TOMA´ S MUNILLA and ANNA SOLER MEMBRIVES Unidad de Zoologı´a, Universidad Auto´noma de Barcelona, 08193 Bellaterra, Barcelona, Spain [email protected] 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 sub- Antarctic, 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 1998, Pushkin 1988, Munilla 1989, 1991, 2000, 2001b, Pycnogonids (Chelicerata, Arthropoda) from Antarctic and 2002, 2005, Stiboy-Risch 1992, 1993, 1994, Bamber 1995, sub-Antarctic waters have been studied more extensively than 2007, Bamber et al. 2001, Chimenz & Gravina 2001), were those from any other ocean of a similar size. Exploration of used to compile the species list. The main zoogeographical this area began with the American expedition of Nathaniel works about pycnogonids are those of Fry (1964), Fry & Palmer whose naturalist, James Eights (1835), described and Hedgpeth (1969), Hedgpeth (1969a, 1969b, 1971) and drew the first Antarctic pycnogonid Decolopoda australis on Munilla 2001a. Other works (Clarke & Crame 1989, 1997, a serolid and some fossils. Pycnogonids have been collected Barnes & De Grave 2000, Clarke & Johnston 2003, Arntz by many scientists but the main monographs are by Hoek et al. 2005, 2006, Barnes 2005, 2006, Clarke et al. 2005, (1881), Mo¨bius (1902), Hodgson (1907, 1927), Bouvier Moyano 2005, Thatje et al. 2005, Gili et al. 2006, Linse et al. (1913), Calman (1915), Gordon (1932, 1938, 1944), Fry & 2006), contain particular zoogeographical reviews of some Hedgpeth (1969), Pushkin (1993) and Child (1994a, 1994b, zoological groups or global and evolutionary reviews of 1995a, 1995b, 1995c). The most detailed information about benthos. A total of 98 papers were consulted. 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 Results and discussion Antarctic species to be described are Ammothe bigibossa Munilla, 2005 and Ammothea victoriae Cano & Lo´pez- Historical research and specific richness Gonzalez, 2007 from the Antarctic Peninsula and the Ross This analysis of the austral pycnogonids covers 172 years, Sea respectively. The aim of this paper is to provide a 16 countries and more than 42 ships or expeditions. So far complete up-to-date list of austral pycnogonids and, using 40 000 specimens have been found in the Antarctic and zoogeographical information consider hypotheses describing sub-Antarctic waters, (termed the Austral Ocean by Jacques their geographical distribution. & 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 Material and methods localities between this front and the Subtropical This paper is an analysis of published data. The works Convergence (D. Barnes, personal communication 2007). mentioned above, as well as the Mu¨ller catalogue (1993) and This sub-Antarctic zone contains many islands with many others (Hoek 1898, Hodgson 1902, 1904, 1908, 1914, pycnogonids (see legend, Table I) and the South American 1915, Bouvier 1905, 1906, Loman 1923, Calman 1933, zone (Magellan and Falkland zones). Bouvet Island is just Stephensen 1947, Hedgpeth 1950, Fage 1952a, 1952b, south of the Polar Front, but given its isolation and that it Utinomi 1959, Stock 1965, Arnaud 1972a, 1972b, Pushkin acts as a link between sub-Antarctic and Antarctic fauna 1974, 1975a, 1975b, 1976, 1977, 1982, 1984a, 1984b, 1990, (Arntz et al. 2006), I consider it as another sub-Antarctic Turpaeva 1974, 1990, 1998, 2000, Krapp 1980, Child 1987, island. Moreover, the general composition of the actual 1 2 TOMA´ S MUNILLA & ANNA SOLER MEMBRIVES 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 ¼ number of world species. Antarctic waters: C ¼ circumpolar, Sc ¼ Scotia Sea, p ¼ Antarctic Peninsula, a ¼ Amundsen Sea, b ¼ Bellingshausen Sea, r ¼ Ross Sea, w ¼ Weddell Sea, e ¼ East Antarctic zone. sub-Antarctic waters: s ¼ South America, k ¼ Iles Kerguelen, c ¼ Ile Crozet, n ¼ New Zealand Plateau, t ¼ Tristan da Cunha, b ¼ Bouvet Is., M þ PE ¼ Marion & Prince Edwards Is., St.P þ A ¼ Saint Paul & Amsterdam Is. Genera/species W. A þ S sp. & Depth Antarctic sub-Antarctic Remarks and synonyms sp. %ofW.sp (m) zone zone Achelia 80 12 (15.4) A. assimilis (Haswell, 1875) 0–903 s,n also west Pacific and Australia A. communis (Bouvier, 1906) 0–714 C s,t A. brucei Calman, 1915 A. dorhni (Thompson, 1884) 0–192 n A. hoekii (Pfeffer, 1889) 5–256 Sc,p,b s A. lagena Child, 1994a 23–137 s A. megacephala (Hodgson, shallow water k 1915) A. parvula (Loman, 1923) 0–267 p s also in Peru-Ecuador-Argentina A. quadridentata (Hodgson, 0–21 St.Pl þ A 1910) A. serratipalpis (Bouvier, 1911) 64–361 Sc,p,b,e. also on Angola coast A. spicata (Hodgson, 1915) 0–1138 C described as Austrothea A. sufflata Gordon, 1944 0–300 w,e e: 40–100ºE A. transfuga Stock, 1954 2–10 n muddy bottom Ammothea 40 25 (62.5) Magnammothea and Biammothea A. adunca Child, 1994a 185–800 w k A. allopodes Fry & Hedgpeth, 210–2000 C b A. bicorniculata Stiboy-Risch, 1992 1969 A. antipodensis Clark, 1972 0–24 n A. armentis Child, 1994a 230–380 k A. bentartica Munilla, 2001 167–335 Sc Livingston. Island, mud A. bigibbosa Munilla, 2005 517 p A. calmani Gordon, 1932 99–1408 Sc,p,b,r,w A. carolinensis Leach, 1814 3–670 C b A. clausi Pfeffer, 1889 3–860 C s A. cooki Child, 1987 1463–2992 n A. dubia (Hedgpeth, 1950) 106 r described as Boehmia A. gibbosa Bouvier, 1913 439–567 b recorded as Colossendeis gibbosa Mo¨bius, 1902 A. gigantea Gordon, 1932 99–1116 C 145ºE–178ºW. Magnammothea gigantea Fry & Hedgpeth, 1969 A. glacialis (Hodgson, 1907) 0–640 C A. gordonae Child, 1994a) 348–732 Sc,r A. hesperidensis Munilla, 2000 30–439 Sc Livingston. Island, mud A. longispina Gordon, 1932 57–1454 C s A. meridionalis (Hodgson, 1915) 10–454 C A. minor (Hodgson, 1907) 8–473 C A. sextarticulata Munilla, 1989 5–516 C also on South Georgia, Biammothea brevipalpa Puskin, 1993 A. spinosa Hodgson, 1907 76–1679 Sc,p,w,r s also in Argentine Basin A. striata Mo¨bius, 1902 72–567 C b A. stylirostris Gordon, 1932 165–494 Sc,w,p s A. tetrapora Gordon, 1932 105–303 Sc s A. tibialis Munilla, 2002 710 Sc A. victoriae Cano & Lo´pez, 2007 360–366 r Ascorhynchus 75 6 (8.0) A. antipodus Child, 1987 5340 n A. cooki Child, 1987 1463–2992 n A. cuculus Fry & Hedgpeth, 993–4008 Sc also in Argentine Basin 1969 A. hedgpheti Turpaeva, 1974 3700–3910 Sc A. inflatum Stock, 1963 2743–6070 Sc also in Peru–Chile trench, South Africa, Kurile trench A. ornatum (Helfer, 1938) 90–108 k also in South Africa Continued CHECK-LIST OF ANTARCTIC AND SUB-ANTARCTIC PYCNOGONIDS 3 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 ¼ number of world species. Antarctic waters: C ¼ circumpolar, Sc ¼ Scotia Sea, p ¼ Antarctic Peninsula, a ¼ Amundsen Sea, b ¼ Bellingshausen Sea, r ¼ Ross Sea, w ¼ Weddell Sea, e ¼ East Antarctic zone. sub-Antarctic waters: s ¼ South America, k ¼ Iles Kerguelen, c ¼ Ile Crozet, n ¼ New Zealand Plateau, t ¼ Tristan da Cunha, b ¼ Bouvet Is., M þ PE ¼ Marion & Prince Edwards Is., St.P þ A ¼ Saint Paul & Amsterdam Is. Genera/species W. A þ S sp. & Depth Antarctic sub-Antarctic Remarks and synonyms sp. %ofW.sp (m) zone zone Austroraptus 5 5 (100) A. calcaratus Gordon, 1944 143–219 r,e e: 65ºE A. juvenilis Calman, 1915 3–500 C e: 92ºE, 145ºE A. polaris Hodgson, 1907 10–569 Sc,p,b,r,e e: 78ºE. Possibly C A. praecox Calman, 1915 6–260 C A. sicarius Fry & Hedgpeth, 220–380 Sc,r 1969 Cilunculus 31 4 (12.9) C. acanthus Fry & Hedgpeth, 2440–2818 b also in Argentine Basin 1969 C. cactoides Fry & Hedgpeth, 38–540 Sc,p,e n 1969 C. kravcovi Pushkin, 1973 255–309 c,M þ PE C. spinicristus Child, 1987 476–540 n Dromedopycnon 2 1 (50) D. acanthus Child, 1982 124–903 s also in Brazilian slope Eurycyde 21 1 (4.8) E. antarctica Child, 1987 527–714 r Sericosura 6 1 (16.7) S. mitrata (Gordon, 1944) 106–2154 r also in Walvis Ridge (South Africa) Tanystylum 45 10 (22.2) T. antipodum Clark, 1977 shallow water n T. brevicaudatum (Fage & Stock, 0–15 St.P þ A also in Cape Verde Island 1966) T. brevipes (Hoek, 1881) 45–100 p,St þ A also in South Africa T.
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