Polar Biol (2009) 32:611–622
DOI 10.1007/s00300-008-0561-6
ORIGINAL PAPER
Cumaceans (Crustacea) from the Bellingshausen Sea
and oV the western Antarctic Peninsula: a deep-water
link with fauna of the surrounding oceans
Jordi Corbera · Carles San Vicente · Jean-Claude Sorbe
Received: 21 August 2008 / Revised: 17 November 2008 / Accepted: 25 November 2008 / Published online: 17 December 2008
© Springer-Verlag 2008
Abstract During the austral summers of 2003 and 2006,
two cruise were carried out in the Bellingshausen Sea and
west oV Antarctic Peninsula on board of RV Hespérides.
Samples were collected at 26 stations with a multinet
Macer-GIROQ sled. A total of 557 cumaceans belonging to
36 species of Wve families were collected. Nannastacidae
was the most abundant and speciose family. Hemilamprops
pellucidus and Cyclaspis gigas were the most frequently
collected species (38.5% of sampling stations). Cumella
asutralis reached the highest density (514.7 individuals/
1,000 m2 at stn 7). Maximum species richness (S = 15) and
diversity (H⬘ = 3.53) was observed at one of the deepest
station. Positive correlations were found between the cumacean distribution and the organic content and percentage of
coarse sand of the sediments. Predominance of Nannastacidae in front of other cumaceans could be explained by their
type of feeding (i.e. predators or scavengers), which may be
more successful in the deep seaXoor of an oligotrophic sea
such as studied herein. The presence in the deepest sampling sites of species shared with faunas of surrounding
oceans suggests a link between these faunas and those of
deep Antarctic waters.
J. Corbera (&)
Carrer Gran 90, 08310 Argentona, Catalonia, Spain
e-mail: [email protected]
C. San Vicente
Carrer Nou 8, 43839 Creixell, Tarragona, Spain
J.-C. Sorbe
Station Marine, UMR 5805 (CNRS/UB1),
2 Rue Jolyet, 33120 Arcachon, France
Keywords Cumacea · Suprabenthos · Abundance ·
Environmental factors · Bellingshausen Sea ·
Antarctic Peninsula
Introduction
Antarctic has been isolated for a long time since the opening
of the Drake Passage (25–30 My BP) and the subsequent
establishment of the Circumantarctic Current (4.3–1.6 My
BP). This isolation has greatly aVected its marine fauna,
and diVerent processes (recolonization, evolution and speciation) have led to its current composition (Clarke and
Crame 1989; Gili et al. 2006).
However, these geological and biological processes
diVerently aVected the marine Antarctic fauna according to
taxa: species impoverishment in Wshes (Eastman and
Grande 1989), species enrichment in many other taxa such
as sponges, Lophophorata and pycnogonids (see Arntz
et al. 1997). In the case of crustaceans, reptant decapods are
nearly absent from Antarctic waters (Thatje and Arntz
2004), whereas peracaridans (amphipods, isopods, mysids,
cumaceans and tanaids) represent one of the most successful group in these areas (San Vicente et al. 1997, 2007;
Arntz et al. 2005). Amphipods and isopods not only show a
high species richness (De Broyer and Jazdzewski 1996;
Brandt et al. 2004) but also have colonized a wide range of
ecological niches. Although cumaceans are less speciose
than amphipods and isopods, they have a high level of
endemism (up to 91%; Corbera 2000).
The study of Antarctic cumaceans began with the Challenger expedition, and Sars (1887) described the Wrst four
species from this region. Later on, subsequent cruises
increased our knowledge on this taxonomical group, mainly
in the Antarctic Peninsula (Zimmer 1907a, b), the Ross Sea
123
612
(Jones 1971; Rehm et al. 2007), the Weddell Sea (Ledoyer
1993; Petrescu and Wittmann 2003) and the South Shetland
Islands (Corbera 2000). However, some Antarctic regions
remain poorly studied. This is the case of the Bellingshausen
Sea, from which only two specimens are so far known: a
preadult female of Cyclaspis gigas Zimmer, 1907 (=C. glacialis Hansen, 1908) and an adult female of Campylaspis
frigida Hansen, 1908 described from the material collected
by the RV Belgica.
One hundred years later, succeeding to a preliminary
report (Corbera and Ramos 2005), this study reports on
new data on the cumacean fauna from the Bellingshausen
Sea: species composition, taxocoenosis structure and biogeographic considerations.
Materials and methods
The cumaceans studied herein were collected during two
Spanish oceanographic cruises in the Bellingshausen Sea
and oV the western Antarctic Peninsula, BENTART-03
(January 30, 2003, to February 26, 2003) and BENTART06 (January 20, 2006, to February 11, 2006) on board of
RV Hespérides. After echo-sounding survey of each
selected sampling zone, a total of 35 stations were visited
during these two cruises (see San Vicente et al. 2009).
However, nine of them were apparently devoid of cumaceans (Stations 4, 11, 12, 20, 21, 22, 27–29) and were therefore excluded from this study. The 26 positive stations
(depth range 85–1,870 m) were sampled with a modiWed
Fig. 1 Position of the
BENTART-03 and BENTART06 sampling stations in the
Bellingshausen Sea and oV the
western Antarctic Peninsula
123
Polar Biol (2009) 32:611–622
version of the Macer-GIROQ suprabenthic sledge (Fig. 1;
Table 1). This gear [see full description in Cartes et al.
(1994)] is equipped with an opening–closing system and
with three superposed nets (0.5 mm mesh size) that simultaneously sample the 10–50 cm (N1), 55–95 cm (N2) and
100–140 cm (N3) water layers above the sea Xoor. During
BENTART cruises, it was towed over the bottom for
2–12 min at 1.5–2 knots.
Data were analysed using PRIMER v5 software (Clark
and Gorley 2001). Univariate diversity indices (Shannon–
Wiener diversity H⬘-log2, evenness J) were calculated from
species abundances. Intersample similarities were calculated using Bray–Curtis coeYcients based on fourth-roottransformed species abundances, which downweights the
eVect of rare or abundant species. Intersample similarities
were ordinated using nonmetric multidimensional scaling
(MDS). The SIMPER routine was used to discriminate species that contributed greatly to distinction between station
groups detected by the cluster analysis. The BIOENV procedure was used to characterise the possible relationship
between the observed cumacean distributions in the investigated area and the measured environmental variables. SpeciWcally, depth (m) and the following abiotic variables were
considered in the analyses (data from Troncoso et al. 2007;
Troncoso and Aldea 2008; Saiz et al. 2008): potential
redox, organic content and all granulometric fractions of
surWcial sediments (gravel: >2 mm, coarse sand: >0.5 mm,
medium sand: >0.25 mm, Wne sand: >0.0625 mm, mud:
<0.0625 mm). Carbonates were not considered due to the
lack of data for some sampling stations. Stations 14, 18, 39
Station
17/2/07 69°21⬘16⬙
6
7
8
9
10
13
14
18
23
24
BS
BS
BS
BS
BS
BS
BS
WAP
WAP
WAP
30
31
33
34
35
36
37
38
39
41
42
43
BS
BS
BS
BS
BS
BS
BS
BS
WAP
WAP
WAP
WAP
68°50⬘08⬙
68°42⬘16⬙
68°49⬘57⬙
68°56⬘42⬙
68°07⬘40⬙
69°15⬘11⬙
69°25⬘56⬙
69°55⬘47⬙
69°55⬘37⬙
70°06⬘02⬙
12/2/10 63°21⬘37⬙
11/2/10 65°09⬘58⬙
11/2/10 65°27⬘20⬙
8/2/10
5/2/10
4/2/10
3/2/10
1/2/10
1/2/10
31/1/10 70°16⬘09⬙
29/1/10 69°57⬘46⬙
27/1/10 69°58⬘24⬙
21/1/10 70°14⬘15⬙
27/2/07 64°19⬘33⬙
26/2/07 64°55⬘56⬙
21/2/07 67°57⬘40⬙
12/2/07 70°44⬘16⬙
12/2/07 70°14⬘42⬙
7/2/07
7/2/07
6/2/07
5/2/07
Beginning
63°21⬘22⬙ 64°17⬘29⬙
65°09⬘54⬙ 68°56⬙01⬙
65°27⬘07⬙ 69°00⬘29⬙
64°17⬘06⬙
68°55⬙48⬙
69° 01⬘00⬙
69°35⬘12⬙
80°12⬘00⬙
69°15⬘19⬙ 80°12⬘11⬙
68°07⬘44⬙ 69°35⬘41⬙
80°50⬘52⬙
80°24⬘37⬙
85°08⬘25⬙
84°52⬘14⬙
69°25⬘55⬙ 80°50⬘30⬙
69°55⬘31⬙ 80°24⬘23⬙
69°55⬘31⬙ 85°09⬘13⬙
70°06⬘24⬙ 84°52⬘37⬙
84°11⬘20⬙
86°22⬘23⬙
69°57⬘49⬙ 86°22⬘08⬙
70°15⬘59⬙ 84°11⬘30⬙
87°26⬘37⬙
95°06⬘07⬙
61°58⬘43⬙
63°38⬘06⬙
69°58⬘20⬙ 87°26⬘54⬙
70°14⬘14⬙ 95°05⬘49⬙
64°19⬘38⬙ 61°58⬘50⬙
64°56⬘01⬙ 63°38⬘25⬙
71°03⬘46
78°04⬘26⬙
69°21⬘15⬙ 78°04⬘39⬙
67°57⬘37⬙ 71°03⬘37
77°44⬘18⬙
81°28⬘14⬙
81°46⬘56⬙
90°21⬘17⬙
69°49⬘42⬙ 77°49⬘48⬙
70°44⬘14⬙ 81°28⬘21⬙
70°14⬘37⬙ 81°46⬘06⬙
68°50⬘12⬙ 90°21⬘17⬙
90°40⬘49⬙
90°49⬘11⬙
68°49⬘53⬙ 90°49⬘18⬙
68°42⬘16⬙ 90°49⬘52⬙
90°35⬘16⬙
95°14⬘20⬙
95°13⬘40⬙
End
68°56⬘44⬙ 90°33⬘22⬙
70°29⬘37⬙ 95°14⬘29⬙
70°38⬘19⬙ 95°13⬘47⬙
End
Longitude W
249
1,272
358
154
1,339
513
563
1,136
620
435
1,395
1,799
1,870
1,052
657
356
492
608
498
540
87
363
192
124
743
524
Beginning
Depth (m)
246 580
1,275 215
358 560
146 356
1,343 124
516 235
559 523
1,137 535
612 726
431 332
1,395 199
1,798 209
1,869 191
1,059 172
646 301
356 139
491 148
618 332
494 104
539 182
85 130
385 34
196 152
123 87
743 100
524 77
End
Haul
length
(m)
NM
NM
NM
221.9
298.2
244.0
289.0
260.7
326.0
290.2
207.8
187.7
178.9
170.5
272.5
NM
NM
240.0
260.0
261.8
155.8
174.8
122.5
199.3
289.3
252.2
Eh
(mV)
NM
NM
NM
NM
5.98
5.70
8.51
7.36
1.80
4.02
5.31
7.01
1.99
8.32
6.75
NM
3.68
4.64
4.05
5.96
1.23
1.85
1.35
1.43
5.02
4.81
O.M.
(%)
NM
NM
NM
NM
0.83
0.64
0.47
2.40
1.27
1.38
2.54
2.97
5.87
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
NM
Carbonates
(%)
NM
NM
NM
NM
65.69
35.37
33.15
47.65
0.00
20.11
0.00
58.38
1.33
0.00
0.00
NM
34.7
10.2
15.8
3.9
0.1
0.00
0.00
0.14
81.4
14.3
Gravel
(%)
NM
NM
NM
NM
3.14
17.04
1.08
3.78
12.91
12.99
2.22
1.78
11.22
0.24
0.50
NM
5.10
3.60
5.20
6.10
0.80
0.20
0.10
0.14
1.80
7.90
Corase s.
(%)
NM
NM
NM
NM
1.26
10.27
1.08
1.73
14.98
8.86
4.81
1.02
29.09
0.23
0.50
NM
3.80
4.10
7.90
4.40
4.90
0.20
0.10
0.32
1.10
7.50
Medium s.
(%)
Sledge haul length calculated from GPS-derived position of the research vessel at the beginning and at the end of each tow [according to Brandt and Barthel (1995)]
Eh Potential redox, O.M. organic matter, s sand, NM not measured
26
BS
BENTART-06
15/2/07 69°49⬘39⬙
5
BS
70°29⬘36⬙
2
1/2/07
1
BS
31/1/07 70°38⬘18⬙
Date
Latitude S
(d/m/y)
Beginning
BS
BENTART-03
Zone
NM
NM
NM
NM
2.72
16.15
3.96
9.13
59.89
26.32
20.74
8.88
49.43
1.53
7.10
NM
11.70
17.50
16.40
12.40
58.90
6.10
21.00
19.50
4.20
19.10
Fine s.
(%)
NM
NM
NM
NM
27.20
21.17
60.72
37.72
12.21
31.72
72.22
29.95
8.94
98.00
91.90
NM
44.70
64.60
54.70
73.20
35.30
93.50
78.80
79.90
11.50
51.20
Mud
(%)
Table 1 Date, geographical location, water depth and environmental bottom characteristics (from Troncoso et al. 2007; Saiz et al. 2008) of the suprabenthic stations sampled in the Bellingshausen
Sea (BS) and oV the western Antarctic Peninsula (WAP) during BENTART-03 and BENTART-06 cruises
Polar Biol (2009) 32:611–622
613
123
614
and 41–43 were also discarded for the same reason as well
as Station 23 (only one specimen sampled). All variables
except depth were previously transformed by log (x + 0.1).
Results
Faunal composition and diversity
A total of 557 cumacean specimens belonging to 35 species
and Wve families were censed from the 26 stations sampled
with the suprabenthic sled (Tables 2, 3). The Nannastacidae
was the most speciose and abundant family with 257 specimens (46.1% of the total number of cumaceans in the studied collection herein) of 19 species from six genera. Within
Nannastacidae, the genus Campylaspis was the most speciose (eight species) and abundant (9.7% of the total). The
Lampropidae, comprising three species of two genera, represented 14.5% of the total specimens collected. Although
more speciose than Lampropidae, the Diastylidae (seven
species of three genera) was less abundant (11.7% of the
total). Only three species of Bodotriidae and Leuconidae
were collected, which, respectively, represented 11.0 and
4.8% of the total specimens collected. Sixty-six specimens
of the present collection (11.8% of total) were not identiWed
due to their damaged condition.
The total densities of cumaceans ranged between 4.2
individuals/1,000 m2 at Station 23 (657–646 m, Antarctic
Peninsula) and 698.5 individuals/1,000 m2 at Station 7
(363–385 m, Bellingshausen Sea, near Peter I Island). The
highest values (>150 individuals/1,000 m2) were all
reported from the Bellingshausen Sea, where densities were
not correlated to depth on the contrary to the trend observed
oV the western Antarctic Peninsula (R = 0.877). Cumella
australis showed the highest observed species density at
Station 7 (514.7 individuals/1,000 m2), followed by Cumella meriodionalis (163.3 individuals/1,000 m2 at Station
31) and Vaunthompsonia inerme (134.6 individuals/
1,000 m2 at Station 8).
As shown in Table 3, Cyclaspis gigas and Hemilamprops pellucidus were the most frequently collected species
in the area investigated (ten stations), followed by Procampylaspis halei (eight stations) and Leptostylis crassicauda
(seven stations). Eight species were collected only at a
single station: Bathycuma capense, Campylaspis alisae,
C. breviramis, Styloptocuma sp. B, Hemilamprops merlini
and Diastylis sp. A from the Bellingshausen Sea; Campylaspis heterotuberculata and Cumella sp. A oV the western
Antarctic Peninsula. The maximum values of species richness (S = 15) and diversity (H⬘ = 3.53) were observed at
Station 30 (1,798.5 m) from the Bellingshausen Sea.
Cumaceans showed a clear vertical distribution gradient
in the near-bottom water layers sampled by the sedge. The
123
Polar Biol (2009) 32:611–622
total number of cumaceans collected in the 10–50, 55–95
and 100–140 cm water layers, respectively, were 92.8, 5.2
and 2.0%. A drastic abundance decrease mainly occurred
between the two lowermost water layers. Such a vertical
distributional trend was also observed for most cumacean
species. Twenty-two species were exclusively sampled in
the 10–50 cm water layer and 29 species were absent in the
100–140 cm water layer (Table 3).
Cumacean assemblages
Three main groups of stations are discriminated by the multivariate analysis of the 25 sampling sites (station 23 was
excluded from this analysis because only one unidentiWed
individual was sampled), based on the species density
(individuals/1,000 m2) (Fig. 2a). The MDS ordination plot
(Fig. 2b) shows similar results to those of the classiWcation,
with a good stress value (0.09). The main abiotic and biotic
features of these station groups are presented in Tables 4
and 5.
Group A includes six stations located around Peter I
Island and in Margarita Bay (depth range 85–385 m), on
muddy/Wne sand sediments (coarse sand fraction <1%) with
a low organic content (mean 1.5%; range 1.2–1.9%). The
cumacean taxocoenosis shows low values of species richness (mean S = 1.8 species/sample) and diversity (mean H⬘
0.80) but the highest abundance of individuals (mean 166.0
individuals/1,000 m2; range 24.7–698.5 individuals/
1,000 m2). Two species, Cumella australis and Vaunthompsonia inermis, account for more than 97% of the average
within group similarity.
Group B includes two stations from the Bellingshausen
Sea and two others oV the western Antarctic Peninsula, at
depths between 246 and 1,059 m. Not sampled at Station
43, the substratum is represented by muddy sediments
(mud > 70%) at Stations 9 and 24 and by a mix of diVerent
granulometric fractions at Station 37 with a dominance of
the gravel fraction (35.4%). The organic content of surWcial
sediments is relatively high when compared to other station
groups (mean 6.7%; range 5.7–8.3%). The cumacean taxocoenosis shows medium values of species richness (mean
S = 3.3 species/sample) and diversity (mean H⬘ 1.48) and
the lowest abundance of individuals (mean 48.6 individuals/1,000 m2; range 13.8–123.6 individuals/1,000 m2). Two
species, Leptostylis crassicauda and Leucon antarcticus,
account for 100% of the average similarity in this group.
Group C includes most of the deepest stations from the
Bellingshausen Sea and two others oV the western Antarctic Peninsula (depth range 358–1,870 m). Not sampled at
Stations 41 and 42, in the remaining stations of this group,
the substratum composition is highly variable from place to
place: from sandy mud bottoms (i.e., Station 31 with mud
fraction >70%) to gravel bottoms (i.e., Station 2 with gravel
–
–
–
Bathycuma capense
–
–
–
–
–
–
Cumella meridionalis
Cumella australis
Cumella emergens
Cumella sp. A
Styloptocuma sp. A
Styloptocuma sp. B
–
Leucon antarcticus
–
–
Leucon assimilis
–
Eudorella gracilior
Family Leuconidae
–
–
–
–
–
25.0
–
–
–
–
–
–
–
–
–
–
–
Campylaspis sp.*
–
–
48.7 25.0
–
Campylaspis quadriplicata
Procampylaspis sp. B
–
Campylaspis maculata
–
–
–
Campylaspis breviramis
–
–
Campylaspis heterotuberculata
–
Procampylaspis compressa
–
Campylaspis antarctica
–
–
–
–
Campylaspis alisae
48.7
–
Campylaspis sp. A
–
–
Procampylaspis halei
–
Campylaspis excavata
–
–
–
–
2
Schizocuma molosa
–
Atlantocuma bidentatum
Family Nannastacidae
–
Vaunthompsonia inerme
1
Cyclaspis gigas
Family Bodotriidae
Station code
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
28.7
–
28.7
–
5
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
110.3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
38.5
–
–
–
–
–
–
–
–
38.5
–
–
–
–
–
–
–
–
–
–
–
73.5 134.6
24.7 625.0
–
–
–
–
–
–
–
–
8
73.5 134.6
7
24.7 514.7
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
6
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
9
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
8.4
–
–
–
–
–
–
–
–
8.4
–
8.4
–
–
–
–
–
–
8.4
–
–
8.4
14
–
–
–
–
–
–
–
–
7.5 25.3
–
–
7.5
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
3.8
–
–
3.8
10 13
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
4.2
–
–
–
–
–
–
–
–
–
9.0 4.2
–
–
–
–
–
–
–
–
9.0 –
–
–
–
–
–
–
–
–
–
–
–
9.0 –
109.0
14.5
–
94.5
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
23 24
9.0 –
–
18
77.8
23.9
17.9
–
–
–
–
7.5
–
–
7.5
–
–
6.3
–
–
–
–
–
29.9
12.0
–
23.9
–
–
–
–
–
–
18.8
–
6.3
–
–
–
–
–
–
–
11.3
7.5
–
–
–
–
3.4
–
–
–
–
–
–
–
–
–
–
–
–
–
36
–
–
–
–
–
–
–
–
–
2.4
–
9.6
–
2.3 –
7.0 –
–
–
–
9.3
–
–
7.0 –
–
–
37.9 –
10.3
–
–
–
–
3.8 –
–
–
–
–
–
2.3
–
–
2.3
35
20.7 –
–
–
–
43.0
10.3
–
–
10.3
34
3.8 –
–
–
–
3.8
89.7 163.3 15.1
–
–
–
–
6.3 –
12.6 –
–
–
–
–
33
–
–
–
–
17.9
–
–
17.9
–
–
–
–
–
–
–
–
–
–
–
–
7.0
–
7.0
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
37
7.2 10.6
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
2.2
2.2
10.1 –
–
–
10.1 –
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
34.8 15.1
–
–
–
–
–
–
2.2
–
5.8 –
–
–
4.3
–
4.3
–
4.3
11.6 –
–
17.4 –
–
–
11.2 –
–
–
–
–
–
40.4 52.7 42.4
–
–
–
–
–
–
2.2
8.6
43
23.3 10.8
–
–
23.3
42
2.2 –
–
–
–
–
–
–
49.2 24.6
–
10.1 –
–
–
–
–
–
17.9
–
11.2
6.7
41
3.5 –
–
–
–
–
–
39
10.1 –
–
20.2
–
–
–
–
–
–
–
–
–
–
38
2.4 10.6 –
4.8 –
–
39.3 209.3 213.6 45.3 115.3 25.6 12.0
–
–
–
–
–
–
–
–
–
–
13.1 –
–
–
–
–
–
–
12.0
13.1 –
–
–
31
41.9 –
–
35.9
30
13.1 –
6.5
–
–
6.5
26
Table 2 Density of the cumacean species (individuals/1,000 m2; 10–140 cm near-bottom water layer) at the 26 stations sampled with a suprabenthic sledge during BENTART-03 and BENTART-06 cruises
Polar Biol (2009) 32:611–622
615
123
123
–
–
16.2
Hemilamprops merlini
Lampropidae unid.
–
–
–
–
–
Diastylis corniculata
Diastylis mawsoni
Diastylis sp. A
Diastylis sp.*
Makrokylindrus
inscriptus
–
–
–
0.68
1.08
3
0.76
0.76
2
1
1
5
2
0.78 1.00 1.00
2.01 2.32 1.00
6
–
1
–
23.9
–
–
–
4
0.55 0.96
–
–
–
–
0.90
3.53
15
7
–
–
9
–
3.8
–
–
–
–
–
–
3.8
–
0.63 0.91
1.77 2.9
25.1
35.9 –
–
–
–
12.0 –
–
–
–
6.5 149.5
0.87 1.91
3
7.3
–
–
71.8 113.1 18.8
6.0 –
6.0 –
Total abundance, species richness, diversity and evenness of the corresponding taxocenoses. * Damaged specimens; – 0.0
1
–
7.3
–
–
–
–
–
–
–
6.5
6.5
–
–
59.8 113.1 18.8
–
33
–
–
7.0
9.3
35
–
–
–
–
–
–
8
–
–
–
–
–
–
–
–
5.3
–
–
–
5.3
37
6
2
–
4.8 –
2.4
4.7 –
–
–
–
–
–
–
–
2.4
–
–
–
–
–
36
0.91 0.95 0.89 0.92
3.02 2.87 2.32 0.92
10
29.3
58.6
–
37.9 –
–
–
–
–
20.7
–
27.6 16.3
–
–
20.7
6.9
34
–
–
–
–
–
–
–
–
–
–
39
7
2
–
–
8
2.2
–
–
–
–
–
–
–
–
–
2.2
–
–
2.2
–
41
0.96 0.35 0.81
2.69 0.35 2.45
30.2
20.2
20.2 –
–
–
–
–
–
–
–
50.4
–
–
30.2
20.2
38
43
–
–
–
–
–
–
–
4.3
9
7
4.3
0.92 0.94
2.94 2.66
34.9
23.2 10.8
–
5.8 –
–
–
5.8 –
5.8
5.8 –
6.5
17.4 –
–
–
–
17.4 –
42
0.86 0.91
1
15.1
–
–
–
–
–
–
–
–
7.3
–
–
–
–
–
31
Evenness (J)
2
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
30
1.37 0.91
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
26
3
–
–
–
–
–
–
–
–
–
8.4
–
–
–
–
23 24
Diversity (H⬘-log2)
–
–
–
13.8 12.0 15.1
–
–
–
–
3.8
–
–
8.4
18
Species richness (S)
–
–
–
–
–
12.0 11.3
6.9 –
–
–
–
–
37.7
–
–
3.8
33.9
14
81.1 37.5 28.7 24.7 698.5 173.1 13.8 12.0 79.2 42.1 18.0 4.2 123.6 58.8 562.2 351.8 75.4 241.1 55.9 24.0 15.9 151.3 52.7 64.7 133.6 41.0
–
–
–
–
–
–
6.9 –
–
–
–
–
–
13
Total D
(individuals/
1,000 m2)
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
10
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
9
Cumacea unid.
–
–
–
8
–
7
16.2 12.5
–
–
–
–
–
–
–
–
–
–
–
–
6
Makrokylindrus sp. A
–
–
–
–
–
–
–
–
–
–
–
–
5
–
–
–
–
–
16.2 12.5
–
–
–
–
–
–
2
Leptostylis antipus
Leptostylis crassicauda –
Family Diastylidae
–
16.2
Hemilamprops
pellucidus
1
Paralamprops asper
Family Lampropidae
Station code
Table 2 continued
616
Polar Biol (2009) 32:611–622
Polar Biol (2009) 32:611–622
617
Table 3 Abundance (N: number of individuals) of cumacean species
in the three near-bottom water layers (N1: 10–50 cm; N2: 55–95 cm;
N3: 100–140 cm; Nt: 10–140 cm) sampled by the sledge above the sea
Xoor and their frequency of occurrence (F) in the 26 sampling stations
Species
N2
N3
Nt
F (%)
Family Bodotriiidae
Cyclaspis gigas
28
1
0
29
38.5
Vaunthompsonia inerme
20
1
4
25
23.1
7
0
0
7
3.8
Bathycuma capense
20
Similarity
N1
0
A
40
60
80
Family Nannastacidae
Atlantocuma bidentatum
0
0
27
7.7
4
2
0
6
11.5
Campylaspis sp. A
7
0
0
7
11.5
Campylaspis alisae
1
0
0
1
3.8
Campylaspis antarctica
15
0
0
15
11.5
Campylaspis heterotuberculata
1
1
0
2
3.8
Campylaspis breviramis
1
0
0
1
3.8
Campylaspis maculata
8
0
0
8
7.7
10
0
0
10
19.2
Campylaspis quadriplicata
Campylaspis sp. *
2
2
0
4
7.7
Cumella meridionalis
52
0
0
52
19.2
Cumella australis
46
0
1
47
7.7
Cumella emergens
2
0
0
2
19.2
Cumella sp. A
1
0
0
1
23.1
Styloptocuma sp. A
8
0
0
8
7.7
Styloptocuma sp. B
1
0
0
1
3.8
Schizocuma molosa
6
0
0
6
11.5
Procampylaspis halei
24
3
1
28
30.8
Procampylaspis compressa
27
1
1
29
15.4
2
0
0
2
7.7
Eudorella gracilior
7
8
2
17
11.5
Leucon assimilis
5
0
0
5
7.7
Leucon antarcticus
3
2
0
8
7.7
Paralamprops asper
22
0
1
23
23.1
Hemilamprops pellucidus
53
2
0
55
38.5
Hemilamprops merlini
1
0
0
1
3.8
Lampropidae unid.
2
0
0
2
7.7
7
2
0
9
26.9
Procampylaspis sp. B
100
13
1
34
33
41
14
42
36
31
30
38
35
10
2
26
24
9
43
37
18
8
5
7
39
6
27
Campylaspis excavata
A
B
C
B
Stress: 0.09
39
7
6
43
9
8
18
26
31
30
5
38
41
33
14
1
A
B
C
34
13
24
35
42
37
36
2
10
Fig. 2 a Hierarchical cluster analysis of the 25 BENTART stations
based on fourth-root transformed abundances and Bray-Curtis similarities. b Non-metric multi-dimensional scaling ordination of the 25
BENTART stations based on fourth-root transformed abundances and
Bray-Curtis similarities
Family Leuconidae
Family Lampropidae
Family Diastylidae
Leptostylis crassicauda
Leptostylis antipus
21
0
0
21
19.2
Diastylis corniculata
3
0
0
3
7.7
Diastylis mawsoni
2
0
0
2
3.8
Diastylis sp. A
2
0
0
2
3.8
Diastylis sp. *
1
1
0
2
7.7
22
1
0
23
7.7
7.7
Makrokylindrus inscriptus
Makrokylindrus sp. A
Cumacea unid.
Total
* Damaged specimens
3
0
0
3
63
2
1
66
517
29
11
557
100
fraction >80%). At these stations, the coarse sand fraction
is never superior to 13% and the organic content ranges
between 1.8 and 8.5% (mean 4.9%). The cumacean taxocoenosis shows the highest values of species richness
(mean S = 6.7 species/sample) and diversty (mean H⬘ 2.36)
and intermediate abundance value (mean 131.4 individuals/
1,000 m2; range 12.0–562.2 individuals/1,000 m2). Eight
species, Hemilamprops pellucidus, Leptostylis antipus,
Cyclaspis gigas, Procampylaspis halei, Cumella meridionalis, Campylaspis quadriplicata, Paralamprops asper and
Procampylaspis compressa account for more than 92% of
the average within group similarity.
Relation between biotic and environmental variables
The BIOENV analysis (Table 6) shows that the highest correlation ( = 0.512) between the eight selected environmental variables and the biotic matrix of the 19 selected
sampling stations is given by a combination of four variables: depth, potential redox, organic content and coarse
sand fraction of surWcial sediments. Particularly, the
123
618
Polar Biol (2009) 32:611–622
Table 4 Mean values (§standard deviation) of abiotic and biotic variables in the three station groups A–C discriminated by the multivariate
analysis of the 25 selected sampling stations
Abiotic/biotic variables
Station groups
A
B
C
N
N
N
Depth (m)
213.9 § 122.3
6
589.3 § 337.8
4
910.1 § 511.8
15
Potential redox (mV)
174.9 § 38.5
5
225.4 § 48.4
3
256.7 § 46.0
12
Organic content (%)
1.5 § 0.3
4
6.7 § 1.4
3
4.9 § 2.0
13
0
0.6
1
2.2 § 1.7
8
Carbonates (%)
NM
Mud (%)
71.9 § 25.3
4
64.1 § 39.2
3
39.0 § 21.0
13
Fine sand (%)
26.4 § 22.7
4
10.0 § 7.6
3
19.2 § 17.4
13
Medium sand (%)
1.4 § 2.4
4
5.0 § 5.1
3
6.7 § 7.9
13
Coarse sand (%)
0.3 § 0.3
4
7.8 § 8.5
3
5.6 § 4.3
13
Gravel (%)
0.1 § 0.1
4
13.1 § 19.4
3
29.4 § 26.8
13
166.0 § 267.3
6
48.6 § 51.5
4
131.4 § 149.5
15
Density (individuals/1,000 m2)
Species richness (S)
1.8 § 0.8
6
3.3 § 2.6
4
6.7 § 3.5
15
Diversity (H⬘ ¡ log2)
0.80 § 0.33
4
1.48 § 1.02
3
2.36 § 0.71
14
Cum.%
N number of measurements, NM not measured
Table 5 Results of SIMPER
analysis: average similarity
within groups
Av.Abund
Av.Sim
Sim/SD
Contrib%
106.00
29.57
1.19
62.56
62.56
40.98
16.48
0.73
34.86
97.42
Group A (average similarity: 47.27%)
Cumella australis
Vaunthopmsonia inermis
Group B (average similarity: 20.20%)
Leptostylis crassicauda
5.15
14.20
0.85
70.31
70.31
Leucon antarcticus
6.29
6.00
0.41
29.69
100.00
Group C (average similarity: 24.76%)
Hemilamprops pellucidus
The main cumacean species are
ranked according to their average contribution to similarity
within the three station groups
A–C discriminated by the multivariate analysis of the 25 selected sampling stations
18.69
5.24
0.80
21.15
21.15
Leptostylis antipus
5.65
4.98
0.48
20.12
41.27
Cyclaspis gigas
6.99
3.81
0.68
15.37
56.63
Procampylaspis halei
8.88
3.68
0.56
14.84
71.48
Cumella meridionalis
1.42
0.39
5.73
77.21
Campylaspis quadriplicata
4.32
1.37
0.39
5.53
82.73
Paralamprops asper
5.85
1.21
0.31
4.87
87.61
Procampylaspis compressa
5.48
1.16
0.30
4.70
92.30
organic content of surWcial sediments is the variable matching the best result when each abiotic variable is considered
separately ( = 0.414).
Discussion
As shown in Table 7, the area investigated herein was more
speciose (35 species) than other Antarctic areas previously
studied, namely the South Shetlands Islands (25 species;
Corbera 2000), the Weddell Sea (26 species; Petrescu and
Wittmann 2003) and the Ross Sea (29 species; Rehm et al.
2007). Furthermore, it also showed an unexpected dominance
123
19.4
of the family Nannastacidae both in terms of number of
species (19 species) and abundance of individuals (54.3%
of total abundance), thus contrasting with the other areas
characterised by a lower representation of this family
(eight to nine species) and by the numerical dominance of
Bodotriidae (South Shetland Islands and Weddell Sea) or
Leuconidae (Ross Sea).
Such an unusual dominance of Nannastacidae could be
referred to their peculiar feeding behaviour. Although
poorly known in cumaceans (see Foxon 1936; Dixon 1944;
Blazewicz-Paszkowicz and Ligowski 2002; Corbera 2006),
nannastacid species belonging to genera Campylaspis and
Procampylaspis have been considered as predators/
Polar Biol (2009) 32:611–622
619
Table 6 Best combinations of abiotic variables obtained through
BIOENV analysis according to Sperman’s rank correlation values,
which best matches the biotic matrix
Number of variables
Correlation
Selections
1
0.414
3
1
0.319
2
1
0.315
5
2
0.468
3, 5
2
0.450
2, 3
2
0.414
1, 5
3
0.498
1, 3, 5
3
0.480
2, 3, 5
3
0.474
1–3
4
0.512
1–3, 5
4
0.473
1, 3–5
4
0.471
1, 3, 5, 8
5
0.490
1–3, 5, 8
5
0.486
1–5
5
0.476
1–3, 5, 6
1: depth; 2: potential redox; 3: organic content; 4: gravel; 5: coarse
sand; 6: medium sand; 7: Wne sand; 8: mud
scavengers with respect to their piercing mouth appendages
(Jones 1976). Furthermore, in some of these species, fragments of foraminiferan test, polychaete setae and crustaceans remains were found in their gut or around their
feeding appendages (Kaestner 1967; Blazewicz-Paszkowicz
and Ligowski 2002). Most of the nannastacid species registered in the Bellingshausen Sea and oV the western Antarctic Peninsula during the BENTART expeditions actually
belong to Campylaspis (eight species) and Procampylaspis
(three species) and these putative predators/scavengers
represent 20.3% of the total cumacean abundance. Their
dominance is probably favoured by the oligotrophic regime
of the Bellingshausen Sea (Mouriño, personal communication), resulting in a ‘benthic desert’ (Saiz et al. 2008)
accentuated by episodic disturbances (iceberg scouring,
intense dropstone fall to bottom). Moreover, it was recently
shown that foraminiferans (putative preys of these nannastacids) constitute the dominant component of the infaunal
Table 7 Abundance (N number
of individuals) and species richness (S number of species) of
cumaceans in four Antarctic
regions (BS & WAP: Bellingshausen Sea and oV western
Antarctic Peninsula)
Bodotriidae
Nannastacidae
Leuconidae
Lampropidae
a
Corbera 2000, b Petrescu and
Wittmann 2003; cRehm et al.
2007; dthis study
Diastylidae
Total
benthos sampled in the study area (Saiz et al. 2008). The
concomitant low representation of cumacean suspensionand deposit-feeders is probably related to the low nutritive
input of pelagic origin to underlying benthic communities
(Gage and Tyler 1991).
In the same way, during BENTART investigations, the
numerical dominance of predators/scavengers was also
observed in other taxonomical groups. Considered as scavengers by De Broyer et al. (2001), the amphipods Lysianassidae represented the most abundant family of this group in
the suprabenthic material collected by the sledge (San
Vicente et al. 2009). According to Matallanas and Olaso
(2007), the benthic icthyofauna captured with baited traps
in the Bellingshausen Sea was dominated by Zoarcidae,
known for their predatory/necrophagous feeding habits.
Huge cumacean densities have been reported for some
localities such as the Gulf of Alaska, NE PaciWc (86,107
individuals/m2; Moore et al. 2007), oV the New Hampshire
Coast, NW Atlantic (38,674 individuals/m2; Gnewuch and
Croker 1973) or in the Okhotsk Sea, NW PaciWc (37,640
individuals/m2; Fadeev 2003). However, cumacean densities reported in literature are more often very much lower
than the preceding ones. In Antarctic waters, the highest
value was reported from shallow eutrophic waters of MacMurdo Sound (31,548 individuals/m2; Dayton and Oliver
1977). However, recent studies reported a mean density of
2,045.7 individuals/1,000 m2 for the continental shelf of
the Ross Sea (Rehm et al. 2007) and even lower values of
55.8 individuals/1,000 m2 and 183.4 individuals/1,000 m2
(ANDEEP II and III cruise, respectively) for the Weddell
Abyssal Plain (Brökeland et al. 2007). In the area investigated herein (samplings on the continental shelf and upper
slope), the mean density of cumaceans was 121.7 individuals/1,000 m2, of the same order of magnitude as that
reported from the Weddell Abyssal Plain. Although comparison of density data have to be done with care due to
diVerences in sampling methodology, the mean value
obtained during this study supports the aforementioned
poverty of the Bellingshausen Sea benthos.
In the area studied, the highest cumacean diversity values were registered between 1,000 and 2,000 m depth, in
concordance with observations reported from other areas,
South Shetland Is.a
Weddell Seab
N
N
S
Ross Seac
S
N
BS & WAPd
S
N
S
622
3
2,209
2
1,088
2
61
3
80
8
89
8
7,912
9
257
19
360
4
41
2
25,471
9
27
3
50
3
163
5
90
2
81
3
112
7
901
9
2,293
7
65
7
1224
25
3,403
26
36,854
29
491
35
123
620
Table 8 Known distribution of
the cumacean species mentioned
in this study
Polar Biol (2009) 32:611–622
Geographical zones
3
4
5
6
7
8
9
10
Bathycuma capense
+
+
Cumella meridionalis
+
+
Hemilamprops merlini
+
11
12
13
+
Campylaspis alisae
+
Atlantocuma bidentatum
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Campylaspis frigida
+
+
Leucon assimilis
+
Campylaspis maculata
+
Campylaspis quadriplicata
+
+
+
+
+
+
+
+
+
+
Vaunthompsonia inerme
+
+
+
+
+
+
+
+
Cumella australis
+
+
+
+
+
+
+
+
Eudorella gracilior
+
+
+
+
+
+
+
+
Cyclaspis gigas
+
+
+
+
+
+
+
Leptostylis antipus
+
+
+
+
+
+
+
+
+
+
+
Diastylis corniculata
+
Leucon antarcticus
+
+
+
+
+
+
Campylaspis antarctica
+
+
Campylaspis heterotuberculata
+
+
Campylaspis breviramis
+
Cumella emergens
+
+
+
Leptostylis crassicauda
+
+
+
Procampylaspis halei
+
+
+
Procampylaspis compressa
+
+
+
+
Paralamprops asper
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Makrokylindrus inscriptus
+
+
+
Campylaspis excavata
+
+
Schizocuma molosa
+
Cumella sp. A
+
Procampylaspis sp. B
+
+
Campylaspis sp. A
+
+
Styloptocuma sp. A
+
Styloptocuma sp. B
+
Diastylis sp. A
+
Makrokylindrus sp. A
+
+
+
Diastylis mawsoni
where cumacean diversity was higher at intermediate
depths on continental slopes (Rex 1981; Cartes and Sorbe
1997). Gage et al. (2004) suggested a poleward decline of
large-scale deep-sea cumacean biodiversity especially in
the northern hemisphere. However, such a pattern could be
diVerent in the southern hemisphere in view of recent studies on peracarid (Brandt et al. 2007) as well as on benthos
in general (Arntz and Gili 2001). Most studies on Antarctic
cumaceans were conducted on continental shelves (down to
600 m) and very little is known on this major taxonomical
group below 2,000 m depth. A greater sampling eVort at
123
2
Species
Hemilamprops pellucidus
Bold symbols (+) represent new
records for the Bellingshausen
Sea and Antarctic Peninsula
from the BENTART-03 and
BENTART-06 cruises according to Corbera and Ramos 2005,
Rehm et al. 2007, and this study.
1: Deep Atlantic Ocean; 2: Deep
PaciWc Ocean; 3: Deep Indian
Ocean; 4: Magellan Region;
5: Kerguelen Islands; 6: South
Georgia; 7: South Orkney
Islands; 8: South Shetland
Islands; 9: Antarctic Peninsula;
10: Bellingshausen Sea; 11:
Ross Sea; 12: East Antarctica;
13: Weddell Sea
1
+
+
+
+
intermediate and deeper depths will probably reveal the
suspected high diversity of the Southern Ocean cumacean
fauna.
According to multivariate analysis, the main environmental factors structuring the cumacean taxocoenosis of the
area studied were the organic content and coarse sand fraction of surWcial sediments as well as water depth. Organic
content of sediments and depth were also pointed out as the
main factors structuring cumacean communities in a W.
Mediterranean coastal area aVected by waste water and
sludge discharges (Corbera and Cardell 1995). The
Polar Biol (2009) 32:611–622
structuring role of a peculiar sediment fraction on cumacean taxocenoses seems to be highly variable according to
geographical locations. In a Brazilian shelf community, Wne
sands and depth were mentioned as the main structuring
factors (Dos Santos and Pires-Vanin 1999). In the Persian
Gulf, it has been reported that there is an increasing density
of the dominant cumacean species together with the
increase of the gravel fraction of the substratum (D. Martin
et al., submitted). Such discrepancies according to substratum fractions are probably related to the feeding biology of
cumacean species, i.e., to their ability to exploit microXora
carried by sand particles that are size-suitable for mouth
appendage manipulations (see Dixon 1944).
Corbera (2000) censed 68 species from Antarctic waters
(south of the Antarctic Convergence), pointing out that
91% of them were endemic from these areas. After the
works of Blazewicz and Heard (1999), Blazewicz and
Heard (2001), Petrescu and Wittmann (2003) and Petrescu
(2006), the current number of Antarctic cumaceans now
reaches 75 species. The species collected during the BENTART-03 and BENTART-06 cruises represent more than a
third of the known Antarctic species (Table 8). Seven species are probably new to Science, belonging to genera
Campylaspis, Cumella, Styloptocuma, Procampylaspis,
Diastylis and Makrokylindrus, only known from the Bellingshausen Sea and oV the western Antarctic Peninsula.
Most of the other species show a wide geographical distribution at high latitudes, as demonstrated by records from
the Weddell Sea, the Ross Sea and the East Antarctic
waters. Among them, Vaunthompsonia inerme, Cumella
australis and Eudorella gracilior were also recorded from
the Scotia Arc (South Georgia and South Orkney Islands),
South Shetland Islands and the Antarctic Peninsula.
Diastylis mawsoni, Makrokylindrus inscriptus and Campylaspis excavata are only known from the High Antarctic
areas, whereas Schizocuma molosa was collected for the
Wrst time in the Bellinghausen Sea after its original Wnding
in the East Antarctic waters. Such a disjunct distribution is
probably due to the scarcity of sampling in deep Antarctic
waters and a wider distribution is suspected for this species.
Finally, four species previously known from deep waters
of the Atlantic (Bathycuma capense, Cumella meridionalis,
Hemilamprops merlini), the PaciWc Ocean (Campylaspis
alisae) and the Indian Ocean (Atlantocuma bidentatum)
were recorded for Wrst time from the deepest sampling stations of the Bellingshausen Sea, suggesting a link between
deep faunas of Antarctic waters and adjacent oceans. This
link is also supported by the Wnding, for Wrst time in Antarctic waters, of two species of the genus Styloptocuma,
which is widely distributed in deep waters of the Atlantic
and PaciWc Oceans.
In conclusion, the cumacean fauna of the area studied is
relatively poor in terms of abundance of individuals,
621
although highly diverse on bottoms deeper than 1,000 m
depth. The shelf fauna (down to 600 m depth) is mainly
represented by species endemic to Antarctic waters and
characterised by a circumantarctic distribution, whereas the
slope fauna shares many species with adjacent Atlantic,
Indian and PaciWc Oceans. Further studies are needed to
conWrm the high biodiversity of the bathyal Antarctic areas
and thus contribute to a better understanding of the origin
of the Antarctic cumacean fauna.
Acknowledgments The BENTART cruises were carried out under
the auspices of two Spanish Ministry of Science and Technology
(MCYT) Antarctic Programmes (REN2001-1074/ANT and
CGL2004-01856). We express our gratitude to the head of campaign
Ana Ramos, to the oYcers and crew of the RV Hesperides and to our
colleagues from the BENTART cruises in 2003 and 2006. B. Mouriño
(Universidad de Vigo) communicated useful comments on oceanographic data.
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