Smithsonian at the Poles: Contributions to International Polar

144 SMITHSONIAN AT THE POLES / PARK AND FERRARI
surpass the insects. Among the copepod orders, calanoid
copepods contribute more numbers of individuals to the
Earth’s biomass, primarily because of their unique success
in exploiting pelagic aquatic habitats. Calanoid copepods
also are speciose; Bowman and Abele (1982) estimated
2,300 species of calanoids, and as of this writing, 525 species
have been added. These calanoid species are placed in
313 nominal genera belonging to 45 families (F. D. Ferrari,
personal database).
Knowledge about the distribution and diversity of
calanoid copepods in the Southern Ocean has increased
signifi cantly over the past century (Razouls et al., 2000).
Most of the calanoid copepods reported from the Southern
Ocean have been collected from pelagic waters. However,
more species new to science are now being described
from waters immediately over the deep-sea fl oor of the
Southern Ocean (Bradford and Wells, 1983; Hulsemann,
1985b; Schulz and Markhaseva, 2000; Schulz, 2004,
2006; Markhaseva and Schulz, 2006a; Markhaseva and
Schulz, 2006b, 2007a, 2007b). The diversity of this benthopelagic
calanoid fauna from other oceans (Grice, 1973;
Markhaseva and Ferrari, 2006) suggests that the total calanoid
diversity from this habitat of the Southern Ocean
is signifi cantly underestimated, and many new species
are expected to be described. The present review then is
restricted to pelagic calanoid copepods because the benthopelagic
fauna has not been well surveyed and their species
not as well known as pelagic calanoids.
Pelagic calanoid copepods are numerically the dominant
species of the zooplankton community in the Southern
Ocean (Foxton, 1956; Longhurst, 1985). Beginning
with the Challenger expedition (1873– 1876), many expeditions
to the Southern Ocean have provided specimens
for taxonomic studies of the calanoids. Early works by
Brady (1883), based on the Challenger collections, Giesbrecht
(1902), based on Belgica collections, Wolfenden
(1905, 1906, 1911), based on the Gauss (German deepsea
expedition) collections, and Farran (1929), based on
the British Terra Nova collections, led to the discovery
of most of the numerically dominant and widespread
pelagic calanoid species in the Southern Ocean. Several
major national expeditions followed, such as the Meteor
expedition, 1925– 1927, the SS Vikingen expedition,
1929– 1930, and the Norvegia expedition, 1930– 1931.
However, collections obtained by these expeditions were
studied mainly to understand the vertical or seasonal
distribution or other aspects of the biology of pelagic
animals. Signifi cant publications resulting from these
studies include Hentschel (1936), Steuer and Hentschel
(1937), and Ottestad (1932, 1936). In 1925 the British
Discovery Committee launched a program of extensive
oceanographic research in the Southern Ocean, including
intensive studies of the zooplankton fauna. Publications
by Mackintosh (1934, 1937), Hardy and Gunther
(1935), and Ommanney (1936) based on the Discovery
collections are notable for their valuable contributions to
the population biology of the numerically dominant calanoid
copepods. Continued studies of the Discovery collections
led to the publication of additional papers, such
as Foxton (1956) about the zooplankton community and
Andrew (1966) on the biology of Calanoides acutus, the
dominant herbivore of the Southern Ocean.
Southern Ocean copepods became the subject of taxonomic
studies once again toward the middle of the last
century with two important monographs (Vervoort, 1951,
1957). These were the most comprehensive treatments
published on pelagic calanoids, to that time, and began a
new era of taxonomic analyses of Southern Ocean copepods.
In these two studies, many previously known species
of the Southern Ocean calanoids were completely and carefully
redescribed, confusion regarding their identity was
clarifi ed, and occurrences of these species in other oceans
were noted from the published literature. Two papers by
Tanaka (1960, 1964) appeared soon afterward, reporting
on the copepods collected by the Japanese Antarctic Expedition
in 1957 and 1959. On the basis of collections made
by the Soviet Antarctic expeditions, 1955– 1958, Brodsky
(1958, 1962, 1964, 1967) published several studies of the
important herbivorous genus Calanus. More recently, important
contributions to taxonomy of the Southern Ocean
calanoids have been made by Bradford (1971, 1981) and
Bradford and Wells (1983), reporting on calanoids found
in the Ross Sea. Additionally, invaluable contributions
have been made to the taxonomy of the important inshore
genus Drepanopus by Bayly (1982) and Hulsemann
(1985a, 1991).
Beginning in 1962, the U.S. Antarctic Research Program
funded many oceanographic cruises to the Southern
Ocean utilizing the USNS Eltanin. Samples taken with
opening-closing Bé plankton nets and Isaacs-Kidd midwater
trawls on these cruises were made available by the
Smithsonian Institution for study through the Smithsonian
Oceanographic Sorting Center. The exhaustive taxonomic
works by Park (1978, 1980, 1982, 1983a, 1983b, 1988,
1993) are based almost exclusively on the midwater trawl
samples collected during the Eltanin cruises, and these
results signifi cantly increased taxonomic understanding
of most species of pelagic calanoids. Other studies based
on the Eltanin samples include the following: Björnberg’s
(1968) work on the Megacalanidae; Heron and Bowman’s