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Smithsonian at the Poles: Contributions to International Polar

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146 SMITHSONIAN AT THE POLES / PARK AND FERRARI<br />

oceanographic expeditions, many of which routinely carried<br />

out sampling pro<strong>to</strong>cols for pelagic organisms.<br />

The Isaacs-Kidd midw<strong>at</strong>er trawls employed by <strong>the</strong><br />

U.S. Antarctic Research Program were particularly effective<br />

in collecting large, pelagic copepods and signifi cantly<br />

increased knowledge about <strong>the</strong> calanoid fauna. More than<br />

1,000 midw<strong>at</strong>er trawl samples were taken, and <strong>the</strong>se samples<br />

are believed <strong>to</strong> have collected nearly all of <strong>the</strong> pelagic<br />

calanoid copepods in <strong>the</strong> w<strong>at</strong>er column; most of <strong>the</strong>se species<br />

have been described (Park, 1978, 1980, 1982, 1983a,<br />

1983b, 1988, 1993). The trawls were not fi tted with a<br />

device <strong>to</strong> measure w<strong>at</strong>er fl ow through <strong>the</strong> mouth of <strong>the</strong><br />

trawl, so no quantit<strong>at</strong>ive measure of <strong>the</strong> amount of w<strong>at</strong>er<br />

fi ltered by <strong>the</strong> trawl can be calcul<strong>at</strong>ed for <strong>the</strong>se samples.<br />

Sampling times, ranging from one <strong>to</strong> four hours, have allowed<br />

a calcul<strong>at</strong>ion of <strong>the</strong> number of animals collected per<br />

unit time of trawl oper<strong>at</strong>ion for <strong>the</strong> more abundant species,<br />

e.g., Paraeuchaeta antarctica (see Ferrari and Dojiri, 1987,<br />

as Euchaeta antarctica), but this measure is <strong>to</strong>o coarse for<br />

<strong>the</strong> rare species th<strong>at</strong> are <strong>the</strong> primary focus of this study.<br />

The Isaacs-Kidd midw<strong>at</strong>er trawls were quickly lowered<br />

<strong>to</strong> a specifi ed deepest depth, obliquely <strong>to</strong>wed <strong>at</strong> 3-5 knots <strong>to</strong><br />

a specifi ed shallowest depth, and <strong>the</strong>n quickly retrieved <strong>to</strong><br />

<strong>the</strong> surface again. It is not possible <strong>to</strong> determine <strong>the</strong> depth<br />

of collection for <strong>the</strong> specimens captured in a sample with<br />

this pro<strong>to</strong>col. Fur<strong>the</strong>rmore, normally, only one trawl sample<br />

was collected <strong>at</strong> a st<strong>at</strong>ion, and <strong>the</strong>refore, only one depth<br />

range was sampled <strong>at</strong> a particular loc<strong>at</strong>ion. As a result of<br />

<strong>the</strong>se constraints, studies based on <strong>the</strong>se samples cannot<br />

provide direct inform<strong>at</strong>ion about <strong>the</strong> vertical distribution of<br />

<strong>the</strong> calanoid species. However, by comparing <strong>the</strong> presence<br />

or absence of a species in samples taken <strong>to</strong> different gre<strong>at</strong>est<br />

depths <strong>at</strong> different loc<strong>at</strong>ions here or by calcul<strong>at</strong>ing a frequency<br />

of occurrence for each sampled depth range rel<strong>at</strong>ive<br />

<strong>to</strong> all trawls <strong>at</strong> th<strong>at</strong> depth range (Yamanaka, 1976), it is<br />

possible <strong>to</strong> make a fi rst-order determin<strong>at</strong>ion of how deep a<br />

trawl has <strong>to</strong> be <strong>to</strong>wed in order <strong>to</strong> collect a certain species.<br />

Sou<strong>the</strong>rn Ocean pelagic calanoids are c<strong>at</strong>egorized here<br />

in several different ways. Species collected in <strong>the</strong> vicinity<br />

of continental or insular land masses are inshore species,<br />

while species collected away from continental or insular<br />

land masses have been c<strong>at</strong>egorized <strong>to</strong> vertical zones as follows:<br />

epipelagic (0– 200m), mesopelagic (200– 1,000m),<br />

and b<strong>at</strong>hypelagic (1,000– 4,000). The term “deepw<strong>at</strong>er”<br />

refers <strong>to</strong> <strong>the</strong> mesopelagic and b<strong>at</strong>hypelagic zones <strong>to</strong>ge<strong>the</strong>r.<br />

In terms of abundance, species are c<strong>at</strong>egorized by <strong>the</strong><br />

number of specimens collected or known from o<strong>the</strong>r expeditions:<br />

CC, very common (over 100 specimens found);<br />

C, common (between 99 and 50 specimens found); R, rare<br />

(between 49 and 10); and RR, very rare (less than 10 specimens<br />

found). These are not counts per sample, but are all<br />

specimens known <strong>to</strong> science.<br />

A large number of endemic species are found in <strong>the</strong><br />

Sou<strong>the</strong>rn Ocean, and <strong>the</strong>se discoveries are placed within<br />

<strong>the</strong> context of endemicity throughout <strong>the</strong> world’s oceans<br />

for two well-studied families, Euchaetidae and Heterorhabdidae.<br />

To facilit<strong>at</strong>e comparisons, four noncontiguous<br />

areas of interest were defi ned among <strong>the</strong> world’s oceans:<br />

<strong>the</strong> Sou<strong>the</strong>rn Ocean, <strong>the</strong> Arctic (including adjacent boreal<br />

seas of <strong>the</strong> Atlantic and Pacifi c oceans), <strong>the</strong> eastern Pacifi c<br />

Ocean (along <strong>the</strong> Pacifi c coasts of <strong>the</strong> Americas, including<br />

<strong>the</strong> boundary currents), and <strong>the</strong> Indo-West Pacifi c Ocean<br />

(in and around <strong>the</strong> Malay Archipelago). The l<strong>at</strong>ter three<br />

were chosen as areas of interest because most of <strong>the</strong> endemic<br />

species of Euchaetidae and Heterorhabdidae not<br />

found in <strong>the</strong> Sou<strong>the</strong>rn Ocean occur in one of <strong>the</strong>m. So,<br />

for example, <strong>the</strong> Atlantic Ocean was not considered an<br />

area of interest because its endemic species occur mainly<br />

<strong>to</strong>ward its nor<strong>the</strong>rn and sou<strong>the</strong>rn boundaries, and <strong>the</strong>se<br />

endemic species could be included in <strong>the</strong> Arctic Ocean and<br />

Sou<strong>the</strong>rn Ocean areas, respectively. There are no a priori<br />

biological hypo<strong>the</strong>ses th<strong>at</strong> support <strong>the</strong>se utilitarian areas<br />

of interest, although endemicity is discussed in <strong>the</strong> context<br />

of high primary and secondary productivity.<br />

Three broad feeding c<strong>at</strong>egories, herbivory, carnivory,<br />

and detritivory, are recognized for many pelagic calanoids.<br />

Food preferences of calanoid copepods have not<br />

been studied system<strong>at</strong>ically, but <strong>the</strong> following public<strong>at</strong>ions<br />

supported <strong>the</strong> general feeding c<strong>at</strong>egories of <strong>the</strong>se<br />

taxa: Mullin’s (1967) work on <strong>the</strong> herbivory of Calanidae<br />

and Eucalanidae, Yen’s (1983) on <strong>the</strong> carnivory of<br />

Euchaetidae, Ohtsuka et al.’s (1997) on <strong>the</strong> carnivory of<br />

derived Heterorhabdidae, Nishida and Ohtsuka’s (1997)<br />

on <strong>the</strong> detritivory of Scolecitrichidae, and I<strong>to</strong>h’s (1970)<br />

and M<strong>at</strong>suura and Nishida’s (2000) on <strong>the</strong> carnivory of<br />

Augaptilidae. Studies of a few species of <strong>the</strong> large family<br />

Aetideidae (Robertson and Frost, 1977; Auel, 1999)<br />

have suggested th<strong>at</strong> <strong>the</strong>se species may be omnivores, but<br />

on <strong>the</strong> basis of <strong>the</strong> morphology of <strong>the</strong>ir mouthparts, <strong>the</strong>y<br />

are considered carnivores here. Feeding is connected <strong>to</strong><br />

environment through areas of high primary and secondary<br />

productivity, and here high primary and secondary<br />

productivity is equ<strong>at</strong>ed with permanent, annually episodic<br />

upwelling areas. These areas of upwelling are associ<strong>at</strong>ed<br />

with western boundary currents adjacent <strong>to</strong> all continents<br />

or are associ<strong>at</strong>ed with three oceanic bands: trans-Sou<strong>the</strong>rn<br />

Ocean, worldwide equ<strong>at</strong>orial, and boreal Pacifi c (LaFond,<br />

1966; Huber, 2002).

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