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MAGELLAN AND ANTARCTIC OPHIUROIDS 427
INTRODUCTION
World-wide there are about 1600 existing species
of ophiuroids; this class surpasses all other classes
of echinoderms in number of surviving species,
except asteroids. In many benthic communities of
the world oceans ophiuroids contribute significantly
to species diversity, abundance and biomass of the
fauna (see e.g. Hyman, 1955; Fell et al., 1969; Tyler,
1980; Barnes, 1987; Voß, 1988; Gerdes et al., 1992;
Dahm, 1996).
Echinoderms, and especially ophiuroids, are very
common on either side of the Drake Passage (Lar-
raín, 1995) and also in the high Antarctic (Voß,
1988; Dahm, 1996). This paper is a first attempt to
SCI. MAR., 63 (Supl. 1): 427-432 SCIENTIA MARINA 1999
MAGELLAN-ANTARCTIC: ECOSYSTEMS THAT DRIFTED APART. W.E. ARNTZ and C. RÍOS (eds.)
Ophiuroids (Echinodermata) of southern Chile and the
Antarctic: Taxonomy, biomass, diet and growth
of dominant species*
CORINNA DAHM
Alfred Wegener Institute for Polar and Marine Research, P.O. Box 120161, D-27515 Bremerhaven, Germany.
E-mail: cdahm@awi-bremerhaven.de
SUMMARY: This study aims on a first comparison of the shallow water (<550 m) ophiuroid fauna of the Magellan region
and the high-Antarctic Weddell Sea. Five species are common to both the Magellan region (22 species) and the Weddell
Sea (42 species). The most abundant Magellan species is Ophiuroglypha lymani, contributing 33% to total ophiuroid abun-
dance and 44% to total ophiuroid biomass. The diets of O. lymani and of three closely related (same sub-family Ophiuri-
nae) also dominant, Antarctic species are similar, indicate however slightly different feeding strategies. The Magellan
species tends more towards microphageous grazing, whereas the Weddell Sea species act more like scavengers. Within the
sub-family Ophiurinae growth performance of O. lymani is higher than in Antarctic species and in the range of boreal
species.
Key words: Ophiuroids, biomass, diet, growth, age.
RESUMEN: OFIUROIDEOS DEL SUR DE CHILE Y DE LA ANTÁRTIDA: TAXONOMÍA, BIOMASA, ALIMENTACIÓN Y CRECIMIENTO DE
LAS ESPECIES DOMINANTES.– El objetivo del presente estudio es realizar una primera comparación entre la fauna de ofiuroi-
deos de aguas someras (<550 m) de la región de Magallanes y del Mar de Weddell (alta Antártida). Cinco de las especies
estudiadas estan presentes tanto en la región de Magallanes (22 especies), como en el Mar de Weddell (42 especies). La
especie magallánica más abundante es Ophiuroglypha lymani, contribuyendo en un 33-100% a la abundancia numérica total
de ofiuroideos y en un 44-100% a la biomasa total de los mismos. La dieta de O. lymani y de tres especies taxonómicamente
próximas (pertenecientes a la misma subfamilia: Ophiurinae), también dominantes, en la Antártida son similares, aunque
presentan algunas diferencias en sus estrategias de alimentación. La especie magallánica tiende más hacia estrategias de tipo
ramoneador micrófago, mientras que las especies del Mar de Weddell actúan más como carroñeros. Dentro de la subfami-
lia Ophiurinae, el crecimiento de O. lymani es mayor en comparación con el de las especies antárticas y está dentro del rango
de las especies boreales.
Palabras clave: Ofiuroideos, biomasa, dieta, crecimiento, edad.
This is AWI publication no. 1481
*Accepted October 15, 1998.
compare the Magellan and high Antarctic ophiuroid
fauna. I will address the following questions:
1. How similar are the Magellan and the high
Antarctic ophiuroid fauna from a taxonomic point of
view?
2. How do ophiuroids contribute to total macro-
zoobenthos biomass in these regions?
3. Is the diet of the dominant species from the Mag-
ellan region comparable to the diet of closely relat-
ed as well as dominant species from the Antarctic?
4. Are there differences in growth and age between
closely related species from these regions?
MATERIAL AND METHODS
Investigation area, sampling and preservation
In the Magellan region (Fig. 1) 47 Agassiz
trawl samples (max. depth 350 m) were taken dur-
ing the Joint Chilean-German-Italian Magellan
Campaign in 1994 by RV “Victor Hensen” and
during the expedition ANT XIII/4 of RV
“Polarstern” in 1996. Stations, investigation area
and sample treatment are described in Arntz and
Gorny (1996) and Fahrbach and Gerdes (1997). In
the Weddell Sea (Fig. 1) 117 trawl samples were
taken during seven expeditions of RV “Polarstern”
between 1983 and 1992 (Dahm, 1996; max. depth
550 m). All samples were fixed in 4% formalde-
hyde buffered with hexamethylentetramine. In the
laboratory all individuals were identified to
species and then stored in 70% ethanol.
Estimation of biomass
Ophiuroid biomass was computed from multibox
corer samples taken by Gerdes et al. (1992) in the
Weddell Sea (see Dahm, 1996) and by Gerdes
(1999) in the Magellan region.
Stomach content analysis
The ophiuroid disc was cut around the perimeter
and the aboral surface removed to expose the stom-
ach. Contents were examined microscopically and
food items identified as exactly as possible. Ophi-
uroid diet was quantified using a points method
(Swynnerton and Worthington, 1940; Hynes, 1950)
modified by Brun (1972) and Dearborn et al. (1986)
which combines information on stomach fullness
and volumetric contribution of food items (for fur-
ther details see Fratt and Dearborn, 1984; Dearborn
et al., 1986; Dahm, 1996; Dahm and Brey, subm.).
Growth and age
Age of ophiuroids was determined by analyzing
the microstructure of annually formed growth rings
on the vertebral ossicles of the arms according to
Dahm (1993, 1996), Dahm and Brey (1998) and
Gage (1990a). I use the Richards function to model
individual growth:
St= S∞(1-D · e -K(t - to) )(1/D)
where S∞is asymptotic size, K (y-1) is the growth con-
stant, D is a shaping parameter and to(y) determines
the inflexion point of the curve. Growth performance
was computed according to Brey (in press):
ϕ= log(Mmax/Amax)
where Mmax is maximum body mass (kJ) and Amax is
maximum age (y).
RESULTS
Taxonomic similarity
In the Magellan region 22 species were found (15
of these identified to the species) belonging to 14
genera and 7 families in comparison to 42 species
belonging to 21 genera and 6 families in the Weddell
Sea. 5 species, 6 genera and 4 families are common
to both regions (Table 1).
428 C. DAHM
FIG. 1. – Sampling areas in the Magellan region and the Weddell Sea.
Biomass
According to the data of Gerdes (1999) and
Gerdes et al. (1992) ophiuroids contribute 1% (=0.07
gCorg m-2) to total macrozoobenthos biomass in the
Magellan region and about 4 % (= 0.44 gCorg m-2)in
the Weddell Sea. Taking into account a correction
necessary for underestimation of ophiuroid biomass
by corers (Dahm, 1996), true biomass is about 3% of
macrobenthic biomass in the Magellan region and
12% in the Weddell Sea, respectively.
Diet
Diet was compared within the sub-family Ophi-
urinae which includes the dominant ophiuroids of
both regions, Ophiuroglypha lymani in the Magellan
region and Ophionotus victoriae, Ophiurolepis geli-
da and Ophiurolepis brevirima in the Weddell Sea
(see Dahm, 1996). Figure 2 shows the diet composi-
tion of O. lymani after the first phytoplankton bloom
and subsequent sedimentation event of the year. The
species is omnivorous and feeds mainly on crus-
taceans (25%), sediment (25%) and phytodetritus
(23%). The three Antarctic species feed on the same
range of items shown in Figure 2, but their diet con-
tains distinctly more crustaceans (about 47%), less
phytodetritus (about 12%) and occasionally poly-
chaetes and sponge needles (Dahm, 1996).
MAGELLAN AND ANTARCTIC OPHIUROIDS 429
TABLE 1. – Ophiuroid species found in the Weddell Sea (Antarcti-
ca; Dahm, 1996) and in the Magellan region. Note that seven fur-
ther Magellan species could not be identified to the species and are
hence not included in this table.
Species Weddell Magellan
Sea Region
Gorgonocephalidae:
Astrotoma agassizii Lyman, 1875 ••
Astrochlamys bruneus Koehler, 1912 •
Astrochlamys sol Mortensen, 1936 •
Astrohamma tuberculatum (Koehler, 1923) •
Gorgonocephalus chilensis (Philippi, 1858) •
Ophiacanthidae:
Ophiacantha pentactis Mortensen, 1936 •
Ophiacantha antarctica Koehler, 1901 ••
Ophiacantha vivipara Ljungman, 1870 ••
Ophiacantha frigida Koehler, 1908 •
Ophiacantha cosmica Lyman, 1878 •
Ophiosparte gigas Koehler, 1922 •
Ophiomitrella falklandica Mortensen, 1936 •
Ophiacamax gigas Koehler, 1901 •
Amphiuridae:
Amphiura proposita Koehler, 1922 •
Amphiura protecta Hertz, 1927 •
Amphiura lymani Studer, 1885 •
Amphiura algida Koehler, 1911 •
Amphiura belgicae Koehler, 1901 •
Amphiura eugeniae Ljungman, 1867 •
Amphiura deficiens Koehler, 1922 •
Amphiura microplax disjuncta
Mortensen, 1936 •
Amphiura dilatata gaussi Hertz, 1927 •
Ophioleucidae:
Ophioleuce regulare (Koehler, 1901) •
Ophiuridae:
Ophioceres incipiens Koehler, 1922 •
Glaciacantha dubium (Koehler, 1901) •
Glaciacantha döderleini (Hertz, 1927) •
Ophiocten dubium Koehler, 1901 •
Ophiocten megaloplax (Koehler, 1901) •
Ophiocten amitinum Lyman, 1878 •
Ophiuroglypha lymani (Ljungman, 1870) •
Ophionotus victoriae Bell, 1902 •
Ophioperla koehleri (Bell, 1908) •
Ophiosteira echinulata Koehler, 1922 •
Ophiosteira debitor Koehler, 1922 •
Ophiosteira rotundata Koehler, 1922 •
Ophiosteira senouqui Koehler, 1901 •
Ophiurolepis brevirima Mortensen, 1936 •
Ophiurolepis gelida (Koehler, 1901) •
Ophiurolepis martensi (Studer, 1885) •
Ophiurolepis tumescens Koehler,1922 •
Ophiogona döderleini (Koehler, 1901) ••
Ophiura rouchi (Koehler, 1912) •
Ophiura serrata Mortensen, 1936 •
Ophiura ambigua (Lyman, 1878) •
Homalophiura inornata (Lyman, 1882) •
Theodoria relegata (Koehler, 1922) •
Ophiozonella falklandica Mortensen, 1936 •
Hemieuryalidae:
Ophiochondrus stelliger Lyman, 1879 ••
Ophiactidae:
Ophiactis asperula (Philippi, 1858) •
Ophiomyxidae:
Ophioscolex nutrix Mortensen, 1936 •
Ophiomyxa vivipara Studer, 1876 •
Ophiodermatidae:
Toporkovia antarctica (Lyman, 1882) •
Total: 42 15 FIG. 2. – Diet composition of O. lymani in the Magellan region
(Beagle Channel, 200 - 350 m water depth, N = 63) in summer.
Age and growth
Figure 3 shows the growth curve of O. lymani in
the Beagle Channel fitted to size-at-age data
obtained from ossicle readings. The oldest speci-
mens of O. lymani were 20 years old with a disc
diameter of about 25 mm. For the three Antarctic
species, Dahm (1996) determined maximum age
and disc diameter to be 22 years and 32 mm for O.
victoriae, 33 years and 21 mm for Ophiurolepis gel-
ida and 25 years and 21 mm for Ophiurolepis bre-
virima. Growth performance of the Magellan
species O. lymani is higher than that of the three
Antarctic species but in the range of boreal species
(Fig. 4).
DISCUSSION
Ophiuroids can tolerate a wide depth as well as
temperature range (e.g. Brey et al., 1996). Therefore
we should expect many Antarctic species to occur
outside the South Polar Sea (South America or New
Zealand). As shown here (Table 1), five high
Antarctic (Weddell Sea) species are present in South
America, whereas New Zealand and Antarctica have
only one species in common (Fell et al., 1969). The
range of many shelf and slope ophiuroid species is
controlled by depth, hence only the existence of
shallow water routes allows wider dispersal (Fell et
al., 1969). No such shallow water route existed in
the more recent history between New Zealand and
Antarctica, whereas there is a submerged ridge, the
Scotia Arc, between the Antarctic Peninsula and
South America (Fell et al., 1969). This is the most
likely migration route for eurybathic species and
may explain the higher number of common species.
However, there are no fossil records which might
indicate the direction or time scale of this migration
(Arntz et al., 1994).
Species numbers and biomass
In terms of species numbers and biomass, ophi-
uroids play a less significant role in the Magellan
region (22 species, 3% of biomass) than in the Wed-
dell Sea (42 species, 12% of biomass). The strong
presence of ophiuroids in environments with low
and highly seasonal food input such as polar seas or
many deep sea regions indicates special adaptations
and higher competitiveness of ophiuroids under
those conditions. Consequently, the lower biomass
of ophiuroids in the Magellan region may be related
430 C. DAHM
FIG. 3. – Growth of ossicle radius R and disk diameter S of O.
lymani in the Magellan region. Richards growth function fitted to
size-at-age data obtained from ossicle growth band readings (320
band readings from 42 individuals). Growth parameters:
R∞= 2.1 mm; K = 0.17; to= 8.58; D = 0.41
FIG. 4. – Growth performance (ϕ= log(Mmax/Amax)) of ophiuroids
from various regions. Dots: Species belonging to the sub-family
Ophiurinae. Circles: Species belonging to other taxa. Data sources:
Antarctic: Dahm (1996); sub-Antarctic: Morison (1979); Magellan:
this study; Boreal: Dahm (1993), Munday and Keegan (1992); War-
wick et al. (1978), Warwick and George (1980); George and War-
wick (1985); Deep Sea: Gage (1990b), Gage and Tyler (1981,
1982a, b).
to more favourable conditions for the benthic fauna.
Longer periods of food availability may enable other
taxa to compete better against ophiuroids. It should
be noted, however, that certain boreal benthic com-
munities may be dominated by ophiuroids, too, as
observed by George and Warwick (1985) or O’Con-
nor et al. (1986).
Diet, age and growth
It might have been more meaningful to compare
diet and growth of the same species in the Magellan
region and in the Weddell Sea, however, none of the
species encountered occurred in sufficient numbers
in both regions.
The diet of all four species of Ophiurinae
analysed here is basically similar (crustaceans, phy-
todetritus, sediment), but nonetheless indicates dif-
ferences in feeding strategy between the Magellan
and Antarctic species. The Magellan species O.
lymani contains comparatively more phytodetritus
and sediment, indicating some kind of micropha-
geous “grazing” strategy (Warner, 1982). The high-
er amount of crustaceans ingested by the Antarctic
species, 47% compared to 25% in O. lymani, indi-
cates that these species act more like scavengers,
which ingest phytodetritus and sediment either acci-
dentally or as a second choice food (Warner, 1982).
This difference may indicate that competition for the
principal food source of benthic animals, i.e. sedi-
mented matter, is higher in the Antarctic, forcing the
dominant ophiuroid species to vertically expand
their trophic niche (see Dahm, 1996).
With only one Magellan ophiuroid species ana-
lyzed so far (Fig. 3), I am not able to make solid
general statements about growth and growth perfor-
mance. My data, however, indicate that Antarctic
species show lower growth performance than close-
ly related species from the Magellan area or other
more northerly regions (Fig. 4). This is in accor-
dance with previous comparisons based on data
referring to various benthic taxa from Antarctic and
non-Antarctic regions (Arntz et al., 1994).
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