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Smithsonian at the Poles: Contributions to International Polar

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(1971) on postnaupliar developmental stages of three species<br />

belonging <strong>to</strong> <strong>the</strong> genera Clausocalanus and Ctenocalanus;<br />

Björnberg’s (1973) survey of some copepods from<br />

<strong>the</strong> sou<strong>the</strong>astern Pacifi c Ocean; Yamanaka’s (1976) work<br />

on <strong>the</strong> distribution of some Eucalanidae, Aetideidae, and<br />

Euchaetidae; Fontaine’s (1988) on <strong>the</strong> antarctica species<br />

group of <strong>the</strong> genus Paraeuchaeta; Markhaseva’s (2001) on<br />

<strong>the</strong> genus Metridia; and Markhaseva and Ferrari’s (2005)<br />

work on Sou<strong>the</strong>rn Ocean species of Lucicutia. In addition,<br />

four exhaustive monographs have tre<strong>at</strong>ed <strong>the</strong> taxonomy of<br />

a pelagic calanoid family throughout <strong>the</strong> world’s oceans,<br />

and <strong>the</strong>se also have contributed fur<strong>the</strong>r <strong>to</strong> an understanding<br />

of <strong>the</strong> Sou<strong>the</strong>rn Ocean fauna: Damkaer’s (1975) work<br />

on <strong>the</strong> Spinocalanidae, Park’s (1995) on <strong>the</strong> Euchaetidae,<br />

Markhaseva’s (1996) on <strong>the</strong> Aetideidae, and Park’s (2000)<br />

on <strong>the</strong> Heterorhabdidae. These monographs were based<br />

on specimens from an extensive set of samples from <strong>the</strong><br />

world’s oceans. As a result, <strong>the</strong> taxonomy and geographical<br />

range of most of <strong>the</strong> widespread species of <strong>the</strong>se families<br />

are now well known.<br />

Recently described new species of Sou<strong>the</strong>rn Ocean<br />

calanoids are ei<strong>the</strong>r pelagic species th<strong>at</strong> previous authors<br />

failed <strong>to</strong> recognize as distinct from similar rel<strong>at</strong>ives, e.g.,<br />

Pleuromamma antarctica Steuer, 1931 (see Ferrari and<br />

Saltzman, 1998), or species inhabiting extraordinary<br />

habit<strong>at</strong>s seldom explored previously, like <strong>the</strong> w<strong>at</strong>er immedi<strong>at</strong>ely<br />

above <strong>the</strong> seafl oor. Bradford and Wells (1983)<br />

described <strong>the</strong> fi rst benthopelagic calanoid copepods of<br />

<strong>the</strong> Sou<strong>the</strong>rn Ocean, Tharybis magna and Xanthocalanus<br />

harpag<strong>at</strong>us, from a bait bottle. Neoscolecithrix antarctica<br />

was collected in small numbers in <strong>the</strong> Antarctic Sound adjacent<br />

<strong>to</strong> <strong>the</strong> Antarctic Peninsula by Hulsemann (1985b),<br />

who believed <strong>the</strong> species lived in close proximity <strong>to</strong> <strong>the</strong><br />

seafl oor. More recent additions <strong>to</strong> <strong>the</strong> benthopelagic calanoid<br />

fauna include Parabradyidius angelikae Schulz and<br />

Markhaseva, 2000, Paraxantharus brittae Schulz, 2006,<br />

and Sensiava longiseta Markhaseva and Schulz, 2006,<br />

each belonging <strong>to</strong> a new genus, and Brachycalanus antarcticus<br />

Schulz, 2005, Scolecitrichopsis elenae Schulz,<br />

2005, Byr<strong>at</strong>his arnei Schulz, 2006, Pseudeuchaeta arcuticornis<br />

Markhaseva and Schulz, 2006, Bradyetes curvicornis<br />

Markhaseva and Schulz, 2006, Brodskius abyssalis<br />

Markhaseva and Schulz, 2007, Rythabis assymmetrica<br />

Markhaseva and Schulz, 2007, and Omorius curvispinus<br />

Markhaseva and Schulz, 2007. These l<strong>at</strong>ter species were<br />

collected from <strong>the</strong> Weddell Sea, an arm of <strong>the</strong> Sou<strong>the</strong>rn<br />

Ocean, and from <strong>the</strong> Scotia Sea.<br />

In this paper, all relevant studies of <strong>the</strong> taxonomy of<br />

pelagic Sou<strong>the</strong>rn Ocean calanoid copepods are reviewed.<br />

Lists are compiled of species, genera, and families, and <strong>the</strong><br />

PELAGIC CALANOID COPEPODS OF THE SOUTHERN OCEAN 145<br />

geographical range within <strong>the</strong> Sou<strong>the</strong>rn Ocean and rel<strong>at</strong>ive<br />

abundance of each species are noted. Morphological<br />

differences are used <strong>to</strong> suggest evolutionary rel<strong>at</strong>ionships<br />

among species. The distribution of all species is reviewed,<br />

and several generalized p<strong>at</strong>terns are hypo<strong>the</strong>sized. Of particular<br />

interest here are <strong>the</strong> species for which fewer than<br />

50 specimens have been collected during <strong>the</strong> extensive<br />

his<strong>to</strong>ry of surveys of <strong>the</strong> Sou<strong>the</strong>rn Ocean pelagic fauna.<br />

The distribution of <strong>the</strong>se rare, pelagic calanoids, almost<br />

all deepw<strong>at</strong>er species, contributes favorably <strong>to</strong> an understanding<br />

of p<strong>at</strong>terns of distribution and of speci<strong>at</strong>ion in<br />

<strong>the</strong> Sou<strong>the</strong>rn Ocean.<br />

METHODS<br />

Traditionally, <strong>the</strong> Sou<strong>the</strong>rn Ocean has been described<br />

physiographically as including <strong>the</strong> ocean basins adjacent<br />

<strong>to</strong> <strong>the</strong> continent of Antarctica plus <strong>the</strong> following adjoining<br />

seas: Amundsen Sea, Bellingshausen Sea, Ross Sea, Weddell<br />

Sea, and <strong>the</strong> sou<strong>the</strong>rn part of <strong>the</strong> Scotia Sea. In this<br />

review of pelagic calanoid copepods, <strong>the</strong> sou<strong>the</strong>rn boundary<br />

of <strong>the</strong> Sou<strong>the</strong>rn Ocean is defi ned physiographically<br />

by <strong>the</strong> Antarctic continent, but <strong>the</strong> nor<strong>the</strong>rn boundary is<br />

defi ned hydrographically, by <strong>the</strong> average position of <strong>the</strong><br />

Subtropical Convergence. The Subtropical Convergence<br />

is loc<strong>at</strong>ed around 40ºS (Deacon 1934, 1937), where <strong>the</strong><br />

surface temper<strong>at</strong>ure of <strong>the</strong> sea drops sharply from about<br />

18°C <strong>to</strong> 10°C. In this review, <strong>the</strong>n, <strong>the</strong> Sou<strong>the</strong>rn Ocean<br />

includes both <strong>the</strong> Antarctic region and <strong>the</strong> subantarctic region.<br />

Antarctic and subantarctic regions are separ<strong>at</strong>ed by<br />

<strong>the</strong> Antarctic Convergence, which is loc<strong>at</strong>ed around 55ºS,<br />

where <strong>the</strong> sea surface temper<strong>at</strong>ure drops 3°C <strong>to</strong> 5°C over<br />

about 30 miles (48.3 km). Although <strong>the</strong> l<strong>at</strong>itudinal positions<br />

of both <strong>the</strong> Subtropical Convergence, among <strong>the</strong> Atlantic,<br />

Pacifi c and Indian oceans, and <strong>the</strong> Antarctic Convergence,<br />

among <strong>the</strong> Atlantic, Pacifi c and Indian sec<strong>to</strong>rs of<br />

<strong>the</strong> Sou<strong>the</strong>rn Ocean, may vary signifi cantly, locally, <strong>the</strong>se<br />

convergences seldom vary more than a degree of l<strong>at</strong>itude<br />

from <strong>the</strong>ir mean position.<br />

Studies of <strong>the</strong> distribution of organisms are one of <strong>the</strong><br />

primary purposes of <strong>the</strong> discipline of taxonomy, and <strong>the</strong><br />

scope and effectiveness of taxonomic studies is dict<strong>at</strong>ed by<br />

<strong>the</strong> availability of specimens. Like most pelagic organisms,<br />

calanoid copepods in <strong>the</strong> Sou<strong>the</strong>rn Ocean have been collected<br />

mainly with <strong>to</strong>w nets oper<strong>at</strong>ed aboard oceangoing<br />

ships, which usually sail <strong>to</strong> a preselected set of geographic<br />

positions in <strong>the</strong> ocean. Because of <strong>the</strong> physical isol<strong>at</strong>ion<br />

of <strong>the</strong> Sou<strong>the</strong>rn Ocean, studies of its pelagic calanoid<br />

copepods have depended primarily on efforts of n<strong>at</strong>ional

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