Biology of the Antarctic Seas XIX

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Biology of the Antarctic Seas XIX
Physical Sciences UPPER ATMOSPHERE RESEARCH IN ANTARCTICA
L. J. Lanzerotti and C. G. Park, Editors
ANTARCTIC OCEANOLOGY THE ROSS ICE SHELF: GLACIOLOGY AND GEOPHYSICS
Joseph L. Reid, Editor C. R. Bentley and D. E. Hayes, Editors
ANTARCTIC OCEANOLOGY II: THE AUSTRALIAN-
NEW ZEALAND SECTOR Biological and Life Sciences
Dennis E. Hayes, Editor
BIOLOGY OF THE ANTARCTIC SEAS
ANTARCTIC SNOW AND ICE STUDIES Milton O. Lee, Editor
Malcolm Melior, Editor BIOLOGY OF THE ANTARCTIC SEAS II
ANTARCTIC SNOW AND ICE STUDIES II George A. Llano, Editor
A. P. Crary, Editor BIOLOGY OF THE ANTARCTIC SEAS III
George A. Llano and Waldo L. Schmitt, Editors
ANTARCTIC SOILS AND SOIL FORMING PROCESSES BIOLOGY OF THE ANTARCTIC SEAS IV
J. C. F. Tedrow, Editor George A. Llano and I. Eugene Wallen, Editors
DRY VALLEY DRILLING PROJECT BIOLOGY OF THE ANTARCTIC SEAS V
L. D. McGinnis, Editor David L. Pawson, Editor
GEOLOGICAL INVESTIGATIONS IN NORTHERN BIOLOGY OF THE ANTARCTIC SEAS VI
VICTORIA LAND David L. Pawson, Editor
Edmund Stump, Editor BIOLOGY OF THE ANTARCTIC SEAS VII
GEOLOGY AND PALEONTOLOGY OF THE ANTARCTIC David L. Pawson, Editor
Jarvis B. Hadley, Editor BIOLOGY OF THE ANTARCTIC SEAS VIII
GEOLOGY OF THE CENTRAL TRANSANTARCTIC MOUNTAINS David L. Pawson and Louis S. Kornicker, Editors
Mort D. Turner and John F. Splettstoesser, Editors BIOLOGY OF THE ANTARCTIC SEAS IX
GEOMAGNETISM AND AERONOMY Louis S. Kornicker, Editor
A. H. Waynick, Editor BIOLOGY OF THE ANTARCTIC SEAS X
METEOROLOGICAL STUDIES AT PLATEAU STATION, Louis S. Kornicker, Editor
ANTARCTICA BIOLOGY OF THE ANTARCTIC SEAS XI
Joost A. Businger, Editor Louis S. Kornicker, Editor
OCEANOLOGY OF THE ANTARCTIC CONTINENTAL SHELF BIOLOGY OF THE ANTARCTIC SEAS XII
Stanley S. Jacobs, Editor David L. Pawson, Editor
STUDIES IN ANTARCTIC METEOROLOGY BIOLOGY OF THE ANTARCTIC SEAS XIII
Morton J. Rubin, Editor Louis S. Kornicker, Editor
ANTARCTIC
American Geophysical Union RESEARCH
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BIOLOGY OF THE ANTARCTIC SEAS XIV ANTARCTIC ASCIDIACEA
Louis S. Kornicker, Editor Patricia Kott
BIOLOGY OF THE ANTARCTIC SEAS XV ANTARCTIC BIRD STUDIES
Louis S. Kornicker, Editor Oliver L. Austin, Jr., Editor
BIOLOGY OF THE ANTARCTIC SEAS XVI ANTARCTIC PINNIPEDIA
Louis S. Kornicker, Editor William Henry Burt, Editor
BIOLOGY OF THE ANTARCTIC SEAS XVII ANTARCTIC CIRRIPEDIA
Louis S. Kornicker, Editor William A. Newman and Arnold Ross
BIOLOGY OF THE ANTARCTIC SEAS XVIII BIRDS OF THE ANTARCTIC AND SUB-ANTARCTIC
Louis S. Kornicker, Editor George E. Watson
BIOLOGY OF THE ANTARCTIC SEAS XIX ENTOMOLOGY OF ANTARCTICA
Louis S. Kornicker, Editor J. Linsley Gressitt, Editor
HUMAN ADAPTABILITY TO ANTARCTIC CONDITIONS
E. K. Eric Gunderson, Editor
POLYCHAETA ERRANTIA OF ANTARCTICA
ANTARCTIC TERRESTRIAL BIOLOGY alga Hartman
George A. Llano, Editor POLYCHAETA MYZOSTOMIDAE AND SEDENTIARIA OF
TERRESTRIAL BIOLOGY II ANTARCTICA
Bruce Parker, Editor alga Hartman
TERRESTRIAL BIOLOGY III RECENT ANTARCTIC AND SUBANTARCTIC BRACHIOPODS
Bruce, Parker, Editor' Merrill W. Foster
ANTARCTIC
Volume 4 7 RESEARCH
SERIES
Biology of the
Antarctic Seas XIX
Louis S. Kornicker, Editor
(cid:127) AmericaGne ophysUicnailo n
Washington, D.C.
1988
ANTARCTIC
Volume 4 7 RESEARCH
SERIES
BIOLOGY OF THE ANTARCTIC SEAS XIX
LOUIS S. KORNICKER, Editor
Published under the aegis of the
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CONTENTS
The Antarctic Research Series' Statement of Objectives
Board of Associate Editors ix
Calanoid Copepodso f the Genus Haloptilus From Antarctic and
Subantarctic Waters
Taisoo Park
Taxonomy and Distribution of the antarctica Species Group of the
Genus Euchaeta (Copepoda, Calanoida)
Marion Fontaine 27
Studies on the Zoarcidae (Teleostei' Perciformes) of the Southern
Hemisphere, I. The Antarctic and Subantarctic Regions
M. Eric Anderson 59
vii
THE ANTARCTIC RESEARCH SERIES:
STATEMENT OF OBJECTIVES
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BOARD OF ASSOCIATE EDITORS
ANTARCTIC RESEARCH SERIES
ix
Biology of the Antarctic Seas XIX
Antarctic Research Series, Volume 47, Pages 1-25
CALANOlD COPEPODS OF THE GENUS HALOPTILUS
FROM ANTARCTIC AND SUBANTARCTIC WATERS
Taisoo Park
Department of Marine Biology, Texas A&M University, Galveston, Texas 77550
Abstract. A systematic study was made of the study, only four species ((cid:127). fons, (cid:127).
the calanoid copepod genus Haloptilus found in ocellatus, (cid:127). oxycephalus, and (cid:127). longicirrus)
Isaacs-Kidd midwater trawl and Bongo plankton occurred in the Antarctic waters, of which
Subantarctic seas and adjacent waters. A to- only (cid:127). ocellatus was truly endemic to the
tal of 799 adult copepods representing eight Antarctic and found exclusively in waters
species of Haloptilus were found, of which south of the Antarctic Convergence. (cid:127). fons
only one species ((cid:127). ocellatus) was truly and (cid:127). oxycephalus occurred throughout the
endemic to the Antarctic, occurring exclusive- study area and have been recorded widely in
ly in waters south of the Antarctic Conver- the world oceans. The findings of (cid:127). longi-
gence. Three other species that were also cirrus in the present study are the first
found in the Antarctic were (cid:127). fons, (cid:127). oxy- records of the species from the South Atlantic
cephalus, and (cid:127). longicirrus, of which the and the Antarctic.
last is a new record and the first two are The remaining four species ((cid:127). spiniceps,
known as more or less cosmopolitan species. (cid:127). ornatus, (cid:127). longicornis, and (cid:127). paralongi-
Four warmwater species found at stations just cirrus) were found during the study only in
north to the Subantarctic were (cid:127). spiniceps, temperate waters north of the Subantarctic,
(cid:127). ornatus, (cid:127). longicornis, and (cid:127). paralongi- of which (cid:127). spiniceps, (cid:127). ornatus, and (cid:127).
cirrus, of which the last is a new record. longicornis have been known to occur widely
All species are redefined with pertinent de- in warm waters of the world oceans. The
scriptions and illustrations. A key is pre- finding of (cid:127). paralongicirrus in the present
sented for identification of the species. study, however, constitutes the first record
of the species from the southern hemisphere.
Introduction In view of the large number of samples
examined and the extent of the area covered,
This work is the sixth in a series of sys-
it is believed that the species found in the
tematic studies on the Antarctic and Subant-
present study represent about the entire fauna
arctic calanoid copepods started in 1967. The
of the genus Haloptilus in the Antarctic and
samples used for this study are the 74 Isaacs-
Subantarctic seas. The most common species of
Kidd midwater trawl and Bongo plankton net
Haloptilus in the Antarctic and Subantarctic
samples previously studied [Park, 1978, 1980,
waters was (cid:127). oxycephalus and 388 specimens,
1982, 1983a, b] for the families Aetideidae,
or 48.6% of the total found in the study,
Euchaetidae, Scolecithricidae, and Phaennidae.
belonged to this species. The second most
The collection data of the samples including
common species in the Antarctic waters was (cid:127).
the date, local time, position, and depth and
ocellatus, represented by 152 specimens, which
a map showing the sampling stations have been
accounted for about 19% of the total found in
given by Park [1980]. Except for three
the entire study area. In the area north of
Isaacs-Kidd midwater trawl samples that were
the Subantarctic, (cid:127). longicornis was repre-
collected on Atlantis II cruise 31, all of
sented by the highest number -- 220 specimens,
these samples were obtained on USNS Eltanin
or 27.6% of the total found in the entire
cruises sponsored by the U.S. Antarctic Re-
study area -o but nearly 99% of them occurred
search Program and were sorted and curated by
at a single station. All species found in
the Smithsonian Oceanographic Sorting Center.
the study are fully redefined in the light of
The procedures and methods of the study are
the anatomical details observed of the speci-
consistent with those employed in the previous
mens and are fully illustrated.
studies.
In the present study a total of 799 adult Genus Haloptilus Giesbrecht, 1898
copepods representing eight species of the
genus Haloptilus were found. Adult males Hemicalanus Claus, 1863, pp. 176-178.
were, however, found only in one sample (E1- Haloptilus; Giesbrecht and Schmeil, 1898,
tanin cruise 46, station 4, 1000 m), and all p. 177.
belonged to a single species, (cid:127). oxycephalus. The genus Hemicalanus was erected by Claus
Of the eight species of Haloptilus found in [1863] to accommodate five new species
Copyright 1988
by the American Geophysical Union.
2 BIOLOGY OF THE ANTARCTIC SEAS XIX
copepods (H. plumosus, H. mucronatus, H. fili- cephalosomal appendages and first 4 pairs of
gerus, H. longicornis, and H. longicaudatus) legs similar to those of female. Fifth pair
found in the Mediterranean Sea. Claus did not of legs only slightly asymmetrical, with a
designate a type species, but since H. plu- fingerlike process internally on left coxa
mosus was described first, it should be re- and a better developed right exopod. In both
garded as the type species of the genus. legs, basis with a long outer seta; endopod
Giesbrecht [1892] subsequently transferred H. 3-segmented, with 0+1+6 setae; exopod also 3-
filigerus and H. longicaudatus to the genus segmented, with 1+1+2 outer spines and a ter-
Augaptilus, and as the generic name Hemicala- minal spine and without inner setae.
nus was previously used by Dana [18(cid:127)53] for a Remarks. The species of Haloptilus show a
different group of copepods, he (in Giesbrecht progressive reduction in the number of teeth
and Schmeil [1989]) proposed the new name on the mandibular blade and setae on the max-
Haloptilus. Matthews [1972] listed 28 nominal illule and maxilla. The most primitive spe-
species of Haloptilus. However, many of these cies of the genus is (cid:127). fons Ferran, 1908,
species still remain poorly known. which has five teeth and a basal spine on the
mandibular blade, a three-segmented endopod
Redefinition of Haloptilus on the maxillule, four setae on the second
inner lobe of the maxillule, and three setae
Female. Foreheadr ound or producedi nto a on each of the six protopodal lobes of the
cephalic'spine of varying size. Rostrumc on- maxilla. (cid:127). angusticepsS ars, 1907, (cid:127). carib-
sisting of 2 filaments, well developedi n most beanensisP ark, 1970, (cid:127). furcatus Sars, 1920,
species but very small or nearly invisible in and (cid:127). major Wolfenden,1 911, are similar to
certain species. First metasomal segment sep- -H . fons in the structure of the mandibular
arate from cephalosome'f ourth and fifth meta- blade and, to somee xtent, in the numbero f
somal segments fused. Urosome 4-segmented. setae of the maxillule and maxilla, and they
Antennule 25-segmented; all segments fully could be grouped with H. fons as the primitive
separate. Exopod of antenna 8-segmented; members of the genus.
first 7 segments each with a seta; eighth The specialized species of Haloptilus are
segment with 4 setae. Endopod 2-segmented, characterized by the presence on the mandibu-
longer than exopod. Mandible with a strong lar b%ade of two large teeth separated by a
masticatory blade bearing a varying number wide gap, a one-segmented endopod on the max-
of teeth' endopod2 -segmentede' xopod5 - illule, a single seta on the secondi nner
segmented.M axillule with an elongate exopod, lobe of the maxillule, and two setae on each
a relatively small endopod,a nd 3 well- of the secondt o fifth protopodal lobes of
developed inner lobes. Maxilla with 6 proto- the maxilla. These specialized species can
podal lobes and a small endopod bearing 7 be further subdivided according to the pres-
setae. Coxa of maxilliped with 3 lobes bear- ence or absence of a cephalic spine and the
ing 2, 3, and 3 setaeo Basis with 2 middle relative length of the antennule:
and 2 distal setae. Endopod 5-segmented, with
4+4+3+3+4 setae. In all 5 pairs of legs, both I. Species with a conspicuous cephalic spine
endopod and exopod 3-segmented. First leg (cid:127). acutifrons (Giesbrecht, 1892)
with an inner seta on coxa and an outer seta Synonym' (cid:127). spinifrons (Sars, 1900)
on basis. Endopod with 1+2+5 setae. Exopod (cid:127). mucronatus (Claus, 1863)
with 1+1+4 inner setae, 1+1+2 outer spines, (cid:127). ocellatus Wolfenden, 1905
and a terminal spine. Second to fourth legs (cid:127). oxycephatus (Giesbrecht, 1889)
similar in having an inner seta on coxa; 1 and (cid:127). pseudooxycephalus Brodsky, 1950
2 setae on first and second endopodal segment, (cid:127). spiniceps (Giesbrecht, 1892)
respectively; 1+1+5 inner setae, 1+1+3 outer II. Species without a conspicuous cephalic
spines, and a terminal spine on exopod. Basis spine
with a long outer seta in fourth leg. Third A. Species with a relatively short anten-
endopodal segment with 6 or 7 setae in second nule
leg, 7 or 8 setae in third, and 7 setae in (cid:127). ornatus (Giesbrecht, 1892)
fourth. Fifth leg with a long outer seta on (cid:127). plumosus (Claus, 1863)
basis. Inner seta of coxa small or entirely (cid:127). tenuis Farran, 1908
missing. Endopod with 1+1+6 setae. Exopod B. Species with a relatively long anten-
with 0+1+3 inner setae, 1+1+2 outer spines, nule
and a terminal spine. Inner seta of second (cid:127). longicirrus Brodsky, 1950
exopodal segment small and often missing. Synonym' (cid:127). setuliger Tanaka, 1964
Male. Forehead round, without a cephalic (cid:127). longicornis (Claus, 1896)
spine. First metasomal segment separate from (cid:127). paralongicirrus Park, 1970
cephalosome. Fourth and fifth metasomal seg-
ments fused. Urosome 5-segmented. Left an- Furthermore, there are species that are
tennule geniculate, with a knee joint between neither obviously primitive nor fully special-
eighteenth and nineteenth segments. Other ized with regard to these characters.
PARK' CALANOID COPEPODS, GENUS HALOPTILUS 3
transitional species have two setae on the 4. Mandibular blade with 5 teeth of simi-
second inner lobe of the maxillule, as in some lar size in addition to a basal spine
of the primitive species, but their dentition (Figure 2a) ø in maxillule, second inner
of the mandible and setation of the maxillule lobe with 4 setae' endopod 3-segmented
and maxilla show a varying degree of reduction (Figure 2b)' all protopodal lobes of
toward the conditions found in the specialized maxilla each with 3 setae (Figure 2c) .....
species. The species belonging to this cate- .................................. H. fons
gory are as follows: 4. Mandibular blade with 2 large teeth,
without a basal spine (Figure 9i)' in
H. austini Grice, 1959 maxillule, second inner lobe with a
Synonym' H. longiceps Tanaka, 1964 single seta' endopod 1-segmented (Fig-
H. chierchiae (Giesbrecht, 1889) ure 1Oa)' second to fifth protopodal
H. validus Sars, 1920 lobes of maxilla each with 2 setae
(Figure lob) ............................ 5
The first two species are characterized by 5. Rostral filaments well developed (Fig-
the presence of three large teeth on the man- ure 11c)' antennule nearly twice length
dibular blade, while the last species allies of body (Figure 11a)' endopod of maxil-
with the specialized species in having two lule with 5 setae (Figure 12a)' third
large mandibular teeth, a one-segmented maxil- endopodal segment of third leg with 7
lular endopod, and two setae on the second to setae (Figure 12f) ...................... 6
fifth protopodal lobes of the maxilla. 5. Rostral filaments missing' antennule
(cid:127). fertilis (Giesbrecht, 1892) has been less than 1.5 times length of body'
described solely from the male and, according endopod of maxillule with 2 setae (Fig-
to the morphological features of the append- ure 1Oa)' third endopodal segment of
ages, seems to be very closely related to (cid:127). third leg with 8 setae ......... (cid:127). ornatus
spiniceps. Of the remaining species, (cid:127). bul- 6. Ventrally, spermathecae of genital seg-
liceps Farran, 1926, was originally described ment close to each other (Figure 11f)'
from immature specimens, and the adults still caudal ramus about 1.5 times as long as
remain unknown. (cid:127). aculaetus (Brady, 1883), it is wide (Figure lie) .... (cid:127). long(cid:127)cornis
5. orientalis (Brady, 1883), and 5. pacificus 6. Ventrally, spermathecae of genital seg-
Chiba, 1956, have been described either so ment widely separated from each other
briefly or poorly that their future identifi- (Figures 13b and 14b)' caudal ramus at
cation seems nearly impossible. Wolfenden least 2 times as long as it is wide ..... 7
[1906] described a new species under the name 7. Dorsally, forehead with a conical an-
of Haloptilus longimanus, which he [Wolfenden, terior projection (Figure 13a)' ven-
1911] subsequently transferred to the genus trally, spermathecae relatively large,
Pseudhaloptilus Wolfenden, 1911. expanded in an anterodistal direction
(Figure 13b) ............... (cid:127). longScirrus
Key to Species of Haloptilus 7. Dorsally, forehead without a conical
in This Study (Female) anterior projection (Figure 14a) ø ven-
trally, spermathecae relatively small,
1. Forehead with a cephalic spine (Fig- somewhat elongated in a lateral direc-
ure 3a) ................................. 2 tion (Figure 14c) ..... 5. paralong(cid:127)cirrus
1. Forehead without a cephalic spine (Fig-
ure lb) ................................ 4 Haloptilus fons Farran, 1908
2. Cephalic spine long, more or less Figs. 1 and 2
straight (Figure 3b); posterior tooth
Haloptilus fons Farran, 1908, pp. 69-71,
of mandibular blade divided distally
pl. 7, figs. 11-15.--Sars, 1925, pp. 245,
into 3 spiniform processes of nearly
246, pl. 71, figs. 1-14.--Rose, 1933,
equal size (Figure 3i); endopod of max-
pp. 211, 212, fig. 252.--Wilson, 1950,
illule with 4 or 5 setae (Figure 4a) .... 3
p. 236.--Matthews, 1972, p. 49.--Tanaka
2. Cephalic spine short, strongly curved
and Omori, 1974, pp. 264-266, fig. 33.
ventrally (Figure 7d); posterior tooth
of mandibular blade tapering into a Occurrence. The following station list
large single point with 3 small lateral shows the occurrence of (cid:127). fons Farran, 1908
spiniform processes (Figures 7h and (hereafter F, female' M, male' PL, prosome
7i); endopod of maxillule with 3 setae length' BL, body length)'
(Figure 8a) .................. (cid:127). spiniceps
3. Body 8.25ñ0.30 mm long; sixth proto- Eltanin Cruise 21
podal lobe of maxilla with 3 setae
(Figure 4b) .................. H. ocellatus Sta. 198, 2972-0 m, iF (PL=4.87 mm;
3. Body 4.43ñ0.27 mm long; sixth proto- BL=6.00 mm)
podal lobe of maxilla with 4 setae Sta. 240, 2470-0 m, iF (PL=5.00 mm;
(Figure 5i) ................ H. oxycephalus BL=6.16