Biology of the Antarctic Seas XVII

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Biology of the Antarctic Seas XVII
ANTARCTIC
American Geophysical Union RESEARCH
SERIES
Physical Sciences BIOLOGY OF THE ANTARCTIC SEAS VIII
David L. Pawson and Louis S. Kornicker, Editors
ANTARCTIC OCEANOLOGY BIOLOGY OF THE ANTARCTIC SEAS IX
Joseph L. Reid, Editor Louis S. Kornicker, Editor
ANTARCTIC OCEANOLOGY II: THE AUSTRALIAN- BIOLOGY OF THE ANTARCTIC SEAS X
NEW ZEALAND SECTOR Louis S. Kornicker, Editor
Dennis E. Hayes, Editor BIOLOGY OF THE ANTARCTIC SEAS XI
Louis S. Kornicker, Editor
ANTARCTIC SNOW AND ICE STUDIES BIOLOGY OF THE ANTARCTIC SEAS XII
Malcolm Mellor, Editor David L. Pawson, Editor
ANTARCTIC SNOW AND ICE STUDIES II BIOLOGY OF THE ANTARCTIC SEAS XIII
A. P. Crary, Editor Louis S. Kornicker, Editor
BIOLOGY OF THE ANTARCTIC SEAS XIV
ANTARCTIC SOILS AND SOIL FORMING PROCESSES Louis S. Kornicker, Editor
J. C. F. Tedrow, Editor BIOLOGY OF THE ANTARCTIC SEAS XV
DRY VALLEY DRILLING PROJECT Louis S. Kornicker, Editor
L. D. McGinnis, Editor BIOLOGY OF THE ANTARCTIC SEAS XVI
GEOLOGY AND PALEONTOLOGY OF THE ANTARCTIC Louis S. Kornicker, Editor
Jarvis B. Hadley, Editor BIOLOGY OF THE ANTARCTIC SEAS XVII
GEOLOGY OF THE CENTRAL TRANSANTARCTIC MOUNTAINS Louis S. Kornicker, Editor
Mort D. Turner and John F. Splettstoesser, Editors
GEOMAGNETISM AND AERONOMY
ANTARCTIC TERRESTRIAL BIOLOGY
A. H. Waynick, Editor
George A. Llano, Editor
METEOROLOGICASLT UDIESA T PLATEAU STATION, TERRESTRIAL BIOLOGY II
ANTARCTICA
Bruce Parker, Editor
Joost A. Businger, Editor
TERRESTRIAL BIOLOGY III
OCEANOLOGY OF THE ANTARCTIC CONTINENTAL SHELF
Bruce Parker, Editor
Stanley S. Jacobs, Editor
STUDIES IN ANTARCTIC METEOROLOGY
Morton J. Rubin, Editor
UPPER ATMOSPHERE RESEARCH IN ANTARCTICA ANTARCTIC ASCIDIACEA
L. J. Lanzerotti and C. G. Park, Editors Patricia Kott
THE ROSS ICE SHELF: GLACIOLOGY AND GEOPHYSICS ANTARCTIC BIRD STUDIES
C. R. Bentley and D. E. Hayes, Editors Oliver L. Austin, Jr., Editor
ANTARCTIC PINNIPEDIA
Biological and Life Sciences William Henry Burt, Editor
ANTARCTIC CIRRIPEDIA
BIOLOGY OF THE ANTARCTIC SEAS William A. Newman and Arnold Ross
Milton O. Lee, Editor BIRDS OF THE ANTARCTIC AND SUB-ANTARCTIC
BIOLOGY OF THE ANTARCTIC SEAS I I George E. Watson
George A. Llano, Editor ENTOMOLOGY OF ANTARCTICA
BIOLOGY OF THE ANTARCTIC SEAS I II J. Linsley Gressitt, Editor
George A. Llano and Waldo L. Schmitt, Editors HUMAN ADAPTABILITY TO ANTARCTIC CONDITIONS
BIOLOGY OF THE ANTARCTIC SEAS IV E. K. Eric Gunderson, Editor
George A. Llano and I. Eugene Wallen, Editors POLYCHAETA ERRANTIA OF ANTARCTICA
BIOLOGY OF THE ANTARCTIC SEAS V Olga Hartman
David L. Pawson, Editor POLYcHAETA MYZOSTOMIDAE AND SEDENTARIA OF
BIOLOGY OF THE ANTARCTIC SEAS VI ANTARCTICA
David L. Pawson, Editor Olga Hartman
BIOLOGY OF THE ANTARCTIC SEAS VII RECENT ANTARCTIC AND SUBANTARCTIC BRACHIOPODS
David L. Pawson, Editor Merrill W. Foster
ANTARCTIC
Volume 44 RESEARCH
SERIES
Biology of the
Antarctic Seas XVII
Louis S. Kornicker, Editor
(cid:127) AmericaGne ophysiUcanli on
Washington, D.C.
1986
ANTARCTIC
Volume 44 RESEARCH
SERIES
BIOLOGY OF THE ANTARCTIC SEAS XVII
Louis S. KORNICKERE, ditor
Published under the aegis of the
Board of Associate Editors, Antarctic Research Series
Charles R. Bentley, Chairman
Samuel C. Colbeck, David H. Elliot,
Dennis E. Hayes, Louis S. Kornicker,
Heinz H. Lettau, John Meriwether,
and Bruce C. Parker
Library of Congress Catalog Card Number:
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1. Marine biology--Antarctic regions--Collected works. I.
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tion (National Research Council (U.S.)) IV. Series: Publication
(National Academy of Sciences (U.S.))
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Printed in the United States of America
CONTENTS
The Antarctic Research Series: Statement of Objectives
Board of Associate Editors vii
Preface ix
Can Pelagic Aggregations Cause Benthic Satiation? Feeding Biology
of the Antarctic Brittle Star Astrotoma agassizii (Echinodermata:
Ophiuroidea)
John H. Dearborn, Frank D. Ferrari, and Kelly C. Edwards
Pelagic Shrimps of the Family Oplophoridae (Crustacea: Decapoda)
From the Pacific Sector of the Southern Ocean: USNS Eltanin Cruises
10, 11, 14-16, 19-21, 24, and 25
Robert A. Wasmer 29
Observations of Marine Birds in the South Atlantic Ocean in the Late
Austral Autumn
Richard R. Eattin, John H. Dearborn, and William C. Townsend 69
THE ANTARCTIC RESEARCH SERIES:
STATEMENT OF OBJECTIVES
The Antarctic Research Series, an outgrowth of research done in the Antarctic
during the International Geophysical Year, was begun early in 1963 with a grant from
the National Science Foundation to AGU. It is a book series designed to serve
scientists and graduate students actively engaged in Antarctic or closely related
research and others versed in the biological or physical sciences. It provides a
continuing, authoritative medium for the presentation of extensive and detailed
scientific research results from Antarctica, particularly the results of the United
States Antarctic Research Program.
Most Antarctic research results are, and will continue to be, published in the
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trial biology, snow and ice, human adaptability, and geology.
Priorities for publication are set by the Board of Associate Editors. Preference is
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performing much the same function as the more formal expedition reports of most of
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The standards of scientific excellence expected for the series are maintained by the
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manuscripts within a short time can be guaranteed.
BOARD OF ASSOCIATE EDITORS
ANTARCTIC RESEARCH SERIES
vii
PREFACE
This volume includes three diverse studies on the fauna of the Antarctic
seas, The first concerns feeding preference of an echinoderm, the brittle
star Asterotoma Agassizii, collected on the continental shelf off South
Georgia and along the Antarctic Peninsula between 1975 and 1983, Stomach
contents revealed a diet consisting of, in order of their abundance, copepods,
mysids, chaetognaths, and euphausi[ds, and also unidentified crustacean and
organic remains, ostracodes, and amphipods, The presence of both herbiverous
and predatory copepods in the diet suggested to the authors that carbon fixed
in surface waters may be transferred to the benthos within a year, The second
st,(cid:127)dy concerns pelagic shrimps of the family Oplophoridae collected in the
Pacific sector mainly during USNS Eltanœn cruises between 1963 and 1966, The
material comprised five genera with 10 species, including two new species,
Keys are presented for the recognition of oplophorid genera and species, and
the patterns of distribution of species are related to major hydrographic
regions, The third study concerns seabirds observed in the South Atlantic
during a cruise of the ARA Islas Orcadas to South Georgia and the South Sand-
wich Islands in May and June 1975 and supplements the few records of bird
observations in that area made during the late austral autumn, The observa-
tions are presented as an annotated list and in tables, Some correlation was
observed between bird distribution and surface water temperature, but zonation
patterns were not pronounced,
Several anonymous reviewers contributed valuable criticisms for the
authors' cons [deration,
Louis S, Kornicker
ix
Biology of the Antarctic Seas XVII
Antarctic Research Series, Volume 44, Pages 1-28
CAN PELAGIC AGGREGATIONS CAUSE BENTHIC SATIATION?
FEEDING BIOLOGY OF THE ANTARCTIC BRITTLE STAR ASTROTOMA AGASSIZII
( ECHINODER(cid:127)MATAO: PHIUROIDEA)
John H. Dearborn
Department of Zoology, University of Maine, Orono, Maine 04469
Frank D. Ferrari
Smithson[an Oceanographic Sorting Center, Museumo f Natural History
$mithsonian Institution, Washington, D. C. 20560
Kelly C. Edwards
Department of Zoology, University of Maine, Orono, Maine 04469
Abstract. Information on the diet, feeding source to the brittle stars. These interac-
behavior, and surface morphology was obtained tions suggest that carbon fixed in surface
for Astrotoma agassizii, a large, simple-armed waters may be transferred to the benthos with-
member of the suborder Euryalina found on the in a year. In addition, frequent reports of
Antarctic shelf. Material was collected at E. antarctica to 1000 m suggest similar carbon
South Georgia and along the Antarctic Penin- transfer to mesopelagic depths. The extent to
sula between L975 and 1983. Extremely long, which this influx of epipelagic carbon affects
flexible arms which can be produced into sin- the predatory brittle star and euchaetid and
uous forms and tight coils, together with a influences benthic and mesopelagic biology
combination of girdle hooklets, hooked and remains to be determined.
paddle-shaped arm spines, and long podia are
employed to capture prey from the water col- Introduction
umn. Frequency-of-occurrence and points meth-
ods were used to examine the stomach contents Increasing uses of automatic underwater
of 115 specimens of Astrotoma agassiz(cid:127)i, of cameras, remote operating vehicles and manned
which 78 (67.8%) contained food. Mean number submersibles, and ever deeper scuba activities
of food types per feeding animal was 1.7, and are providing new opportunities for studies of
mean fullness value was 1.8, on a scale of the behavior of subtidal benthic invertebrates
16. The diet consisted of members of only two [e.g., Pawson, 1982]. Amongt he ecological
major taxa, Crustacea and Chaetognatha. Cope- relationships which can now be more effective-
pods occurred in 75.6% of brittle stars con- ly studied is the extent to which some benthic
taining food and were the dominant prey group, invertebrates obtain food from the water
followed by mysids (34.6%), chaetognaths rather than through various benthic processes.
(10.2%), and euphausiids (8.9%). Other prey The food and feeding mechanisms of brittle
included unidentified crustacean and organic stars (Echinodermata: Ophiuroidea) in particu-
remains, ostracodes, and amphipods. All cope- lar have received recent attention (reviewed
pods in the stomachs of Astrotoma were calan- by Warner [1982]). Brittle stars exhibit
olds belonging to 12 species generally consid- diverse food capture adaptations. The struct-
ered pelagic animals. Euc(cid:127)aeta antarctica and ural and functional morphology of members of
Calanoides acutus constituted about 80% of the the suborder Euryalina is especially intrigu-
stomach content copepods. This association of ing [Hendler and Miller, 1984]. Some genera
an herbivorous calanid and predatory euchaetid (e.g., Asteroporpa, Astrochlamys) have five
may result from the well-known seasonal migra- very long unbranched arms. Other genera show
tions of Calanoides. Upon completing their varying degrees of arm branching. In Astrocn-
epipelagic season, aggregations of lipid-rich, ida and Conocladus, for example, the arms are
late-stage copepodids of C. acutus might be branched only a few times, while in the most
expected to attract larger pelagic predators, complex Euryalina, the true basket stars (Got-
including the euchaetid. In shoal waters this gonocephalusa nd Astrophyton), the arms branch
association of downwardm igrating herbivores many times, resulting in a complex assemblage
and their predators may bring these calanolds of "twigs." The feeding branches of these
in close proximity to the substrate, thereby many-divided arms are raised in a fanlike
providing a rich, seasonally predictable food posture and oriented into the current to pro-
Copyright 1986
by the American Geophysical
2 BIOLOGY OF THE ANTARCTIC SEAS XVII
1966; Patent, 1970; Macurda, 1976; Meyer and
Lane, 1976; Wolfe, 1978; Hendler, 1982; LaBar-
beta, 1982; for general review see Warner,
1982]. These papers deal primarily with Gorg-
onocephalus, Astrophyton, or Astroboa.
The diets and feediL(cid:127)g behavior of Euryalina
with unbranched arms have received much less
attention. The most informative paper to date
is the account by Hendler and Miller [1984] on
Asteroporpa annulata (Gorgonocephalidae).
They report observations made in situ from the
submersible Johnson-Sea-Link I on a population
off central Florida in depths of 56-85 m. The
brittle stars were found perching on clumps of
Oculina varicosa, a scleractinian coral. The
majority of individuals assumed suspension
feeding postures only at night. Their diet
consisted primarily of pelagic organisms,
mostly copepods.
Astrotoma agassizii (Figures I and 2) is
the largest Antarctic member of the Gorgonoce-
phalidae with five unbranched arms. In large
specimens, with a disc diameter of 50-60 mm,
the arms may reach 600-700 mm in length. The
arms of such an individual would typically
have a basal diameter of about 12 mm and taper
gradually to about 1 mm at the tip. General-
ly, one or two ar(cid:127)ns are used to cling to the
substrate, with the remainder extended to feed
'._,.,. (Figure 2). This species is irregular in
occurrence on the Antarctic shelf, although it
may be locally abundant where conditions are
.. favorable. Information on the feeding biology
.,(cid:127)... . . of A. agassizii has been summarized by Dear-
. ,.
born [1977]. He reviewed comments by Motten-
sen [1936] and Fell [1961] on the planktonic
nature of the food of Astrotoma, suggested by
presence of copepods and hyperiid amphipods in
stomachs, and on the basis of bottom photo-
graphs presented some new information on the
extremely flexible arms and various feeding
postures of this species.
We report here on the diet, feeding behav-
ior, and morphology of 115 specimens of A.
agassizii taken at South Georgia and along the
Antarctic Peninsula between 1975 and 198J. We
suggest that the dominance of pelagic calanoid
copepods in stomachs of A. agassizii in aust-
ral autumn is correlated with seasonally epis-
odic behavior of the prey. Further, since
some prey copepods are pelagic herbivores, we
infer that carbon transferred directly to the
brittle star has been fixed in the water and
not simply recycled by benthic copepods.
Fig. 1o Astrotoma agassizii; dried Predator-prey interactions, the large body
specimens. (A) Entire, DD = 43 mm. (B) size of Astrotoma, and its local abundance on
Aboral disc surface, DD = 56 mm. (C) Oral the Antarctic shel. f make it .an important
disc surface, same specimen as that shown in macroinvertebrate for further investigation.
Figure lB.
Materials and Methods
duce an effective trap for zooplankton [Davis, Sample Collection
1966; Hendler, 1982]. Most reports on the
behavior of Euryalina have involved these A total of 115 specimens of A. agassizii
complexly branched basket stars [Davis, 1966; were obtained by trawl from eight stations
Fricke, 1966, 1968; Tsurnamal and Marder, around South Georgia and along the
DEARBORN ET AL.: FEEDING IN ASTROTOMA 3
Peninsula in 1975 and 1983 (Table 1). Samples the number of specimens feeding. Frequency of
were collected at five sites off South Georgia occurrence of individual food items was then
in May and June 1975 during ARA Islas Orcadas expressed as a percentage of feeding
cruise 0575 [DeWitt et alo, 1976]. Depths of animals. The first points method of analysis
capture rangedf rom 121 to 265 m. Both 1.5-m followed the general methodso f Brun [1972]
and 3-m beam trawls were used. In March and and Fratt and Dearborn [198(cid:127)]. In this meth-
April 1983, additional specimensw ere taken at od, upon initial dissection, the stomachw as
two sites along the Antarctic Peninsula during assigned an estimated overall fullness value
R/V Hero cruise 83-3 [Dearborn et al., of 0 (empty), 1 (trace), 2, 4, 8 (one-half
1984]. The southernmost location, in the full), 12, or 16 (maximumfu llness). Food
Lemaire Channel between Booth Island and the items, if any, in the gut were then catalogued
Danco Coast, was sampled twice with two types and assigned a point value from the same scale
of gear. Initial collections were madew ith a according to their estimated volumec ontribu-
1.5-m beam trawl. A 3-m otter trawl was used t(cid:127)on to the food bolus (see Fratt and Dearborn
[1(cid:127)84] for details). Results were expressed
at a second station in the same general loca-
tion. The otter trawl was more successful as the proportion of total points awarded to a
than the beam trawl in obtaining specimens of particular food item compared to the total
Astrotoma agassizii, but the extremely rough points assigned to all food items.
bottom of the Lemaire Channel severely damaged The third method of analysis was a modifi-
the net. At Hope Bay at the eastern end of cation of the first points method by which we
the Trinity Peninsula, the second Antarctic attempted to mitigate the inflationary effect
Peninsula site, all trawling was done with a of stomachs containing only a trace of a sing-
1.5-m beam trawl. Hauls of short duration le food item on the totals for that food
were used to minimize damage to specimens and item. Under Brun's [1972] and Fratt and Dear-
contamination of stomach contents. The latter born's [1984] unmodified points methods a gut
was unlikely, however, because Astrotoma has a with a trace of a single item and a gut filled
relatively small mouth which does not gape in with the same item would both have contributed
the same manner as it does in some noneuryalid 16 points to the total for that prey species
brittle stars. because these earlier methods did not include
the overall fullness index in calculating the
Stomach Analysis point totals for each food item and thus did
not take into account the absolute amount of
Immediately upon capture the arms were each item present. Our modification involved
removed from specimens of Astrotoma agassizii multiplying the point value assigned to each
retained for stomach analyses, and the discs food item by the fullness index. For example,
were fixed in 70% ethanol. Large discs were if ophiuroid A had a nearly empty gut, with
injected with ethanol to ensure proper pres- only a trace of sediment present, it would
ervation of contents. No regurgitation of have been assigned a fullness index of ! and
stomach contents was evident upon contact with 16 points for sediment (having been the only
the fixative or during injection. Some live item present). In the final summation, earl-
specimens were retained in circulating sea- ier methods would have assigned this animal 16
water at ambient temperature in shipboard points for sediment, as would our new modifi-
holding tanks for behavioral observations in cation. If, on the other hand, ophiuroid B
laboratory aquaria at Palmer station. had a gut that was packed absolutely full of
Prior to dissection the disc diameter (DD) copepods (fullness index of 16 and a point
and width of arm base (AB) were measured to value for copepods of 16 because copepods were
the nearest 0.1 mm. Animals were dissected by the only food item present), the two methods
cutting around the disc perimeter and folding would have given very different results. Upon
back the aboral surface to expose the stomach summing, earlier methods would have assigned a
lining. Stomach contents were examined with a total of 16 points to copepods, whereas our
Wild M5 dissecting microscope, and the types modification assigned 256 (16 x 16) points to
of food and relative abundance of each were copepods. The earlier methods would have set
recorded. Photomicrographs were taken of equal values for sediment and copepods in the
represeJtative stomachs. After initial exam- above examples. By comparison, our modified
inat(cid:127)on, all food items were removed and iden- points method considered both the total volume
tified to the lowest possible taxonomic of the food bolus and the relative volume of
level. Copepods were identified by the second the particular prey items. This provided a
author (F.D.F.) and remaining prey by the more realistic evaluation of the relative
other authors (K.C.E. and J.H.D.). importance of the individual food items in the
Diets of A. agassizii were described by diet of the sample population.
three methods, one giving the frequency of A portion of the specimens from station 90
occurrence of food items and the other two, were inadvertently processed by the percent
both points methods, giving different volumet- frequency-of-occurrence analysis only. We
tic contributions of food items. All animals later designated material from that location
were analyzed by the first procedure. Pres- as coming from station 90 (all three diet
ence or absence of food was noted to determine analyses completed) or station 90a