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Recent Ostracoda From The Laptev Sea (Arctic Siberia) : Species Assemblages and Some Environmental Relationships
Recent Ostracoda From The Laptev Sea (Arctic Siberia) : Species Assemblages and Some Environmental Relationships
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Abstract
Ostracod assemblages from coretop sediments collected at 26 localities at different depths of the Laptev Sea shelf
and upper continental slope were investigated for assemblage studies. A total of 41 species belonging to 19 genera and
12 families have been identified. Three assemblages have been established that could be linked to environmental
factors such as water depth, bottom salinities, water mass circulation and sea-ice transportation. The species-rich and
abundant assemblages of the western and central Laptev Sea were related to the Atlantic waters occupying the upper
continental slope. These include relatively deep-water forms that show clear affinities to North Atlantic and Arctic
Ocean assemblages (Cytheropteron biconvexa, C. testudo, C. simplex, C. nodosoalatum, C. inflatum, C. porterae, Krithe
glacialis, K. minima, Pseudocythere caudata, Polycope punctata, P. orbicularis). In the eastern middle shelf region, the
assemblage is comprised of Acanthocythereis dunelmensis together with other normal marine species (Semicytherura
complanata, Elofsonella concinna, Cluthia cluthae). This assemblage seems connected to the winter flaw polynya which
is believed to be the main area of sediment entrainment into sea ice. The inner shelf assemblage of the southern
Laptev Sea is dominated by shallow-water euryhaline species (Paracyprideis pseudopunctillata and Heterocyprideis
sorbyana) with admixture of the brackish-water species Roundstonia macchesneyi. The unusual occurrence of a number
of shallow-water ostracod species on the upper continental slope may be explained by ice-rafting which these
ostracods are probably able to survive.
2 2002 Elsevier Science B.V. All rights reserved.
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PII: S 0 3 7 7 - 8 3 9 8 ( 0 2 ) 0 0 1 3 6 - 6
dence of variations in water depth, temperature^ (1995) identi¢ed 17 ostracod species representing
salinity characteristics, river runo¡ and bottom a typical inner shelf Arctic assemblage dominated
hydrodynamics. by Paracyprideis pseudopunctillata, Sarsicytheri-
Although ostracods could also play an impor- dea spp., Heterocyprideis sorbyana and Cluthia
tant role in palaeoenvironmental reconstructions cluthae. The authors conclude that a similar as-
of the shallow Siberian shelf seas, the data on semblage is also characteristic of the inner Alas-
Recent ostracods are limited, especially for the kan and Canadian shelf (e.g. Cronin, 1989).
Laptev Sea. Only ¢ve publications mentioning The present investigation was undertaken to
Recent ostracod species from this area are avail- provide a thorough assemblage study of the mod-
able to date (Akatova, 1946; Gorbunov, 1946; ern Ostracoda in the Laptev Sea. Using sediment
Cronin et al., 1991; Bauch et al., 1995; Erlenkeus- samples from the shallow inner shelf region down
er and von Grafenstein, 1999). Akatova (1946) to the upper continental slope, it is also attempted
gives a taxonomic description of seven ostracod to relate the assemblages to water depth, water
species found in the shallow waters around the mass properties and prevailing circulation pat-
New Siberian Islands (water depth range 14^68 terns. Finally, the modern ostracodal assemblages
m), while Gorbunov (1946) provides a total of of the Laptev Sea are compared with the available
26 Ostracoda species. Neale and Howe (1975) data from other Arctic areas.
used the data of Akatova (1946) in their review
of the high-latitude ostracod fauna in order to
assess the relationships between the Arctic ostra- 2. Environmental setting
cod faunas from di¡erent localities and the di-
verse fauna from the Russian Harbour (Novaya The Laptev Sea is an open marginal sea of the
Zemlya, Barents Sea). In general, they regard the Arctic Ocean occupying an area of 662U103 km2
ostracod fauna of the Arctic shelf waters as a between 70‡42’ to 81‡16’N and 95‡44’ to 143‡30’E
single circumpolar province with minor di¡eren- (Fig. 1). Its shelf has a vastly shallow topography,
ces between the eastern and western subprovinces. gently sloping northward down to water depths of
However, in their review the authors deal with 50^60 m, which is cut by several palaeoriver val-
data from the western Arctic. Seven ostracod leys. From October until mid-July the sea is cov-
samples from di¡erent parts of the Laptev Sea, ered with ice. A 1.5^2-m thick fast ice cover is
collected during the US Coast Guard Northwind characteristic of the shallow coastal zone. The
cruise in 1963, are listed in the Modern Arctic northern boundary of the fast ice runs approxi-
Podocopid Database (MAPD; Cronin et al., mately along the 20^25 m isobath, where an up to
1991). The total number of identi¢ed species is 100-km wide polynya can form in winter, separat-
nineteen. Three samples from the mid-depths, ing the fast ice from the region with drift ice.
one from the western shelf and two from the sub- Five large rivers empty into the sea, of which
marine Lena valley exhibit a high taxonomic di- the annual discharge from the Lena River consid-
versity (up to 15 species) with predominance of erably a¡ects the salinity in the eastern Laptev
Paracyprideis pseudopunctillata, Heterocyprideis Sea. In the shelf zone, the upper, freshened, water
fascis and Elofsonella neoconcinna. In contrast, layer is divided from the underlying saline water
all samples from the eastern Laptev Sea are tax- by a pycnocline, usually located in the depth
onomically rather poor, consisting of only three range from 5^7 to 20^25 m (Dmitrenko et al.,
species, Sarsicytheridea bradii, Heterocyprideis 1999).
sorbyana and Paracyprideis pseudopunctillata, the The temperature of the bottom water layer is
latter being the most abundant. Two other studies negative nearly all year round. At depths exceed-
(Bauch et al., 1995; Erlenkeuser and von Grafen- ing 80^100 m temperature rises up to 0.6^0.8‡C
stein, 1999) present merely lists of ostracod spe- due to the in£uence of the warm Atlantic waters.
cies found in coretop samples collected during Recent oceanographic investigations (Dmitrenko
two expeditions in the early 1990s. Bauch et al. et al., 2001a,b) have shown that at certain times
Fig. 1. Distribution of ostracods in the Laptev Sea surface sediments. Key: (1) location of stations of the TRANSDRIFT V ex-
pedition; (2) location of stations of the previous TRANSDRIFT expeditions (I-IK93, II-PM94, III-KD95); (3) abundance of os-
tracods in surface sediment samples (specimens per 100 g dry bulk sediment); (4) marine species; (5) euryhaline and brackish-
water species. Isobaths in metres.
there exist strong wind-induced reversal bottom bottom water salinity is lower in the eastern Lap-
currents that can cause a southward advection tev Sea (Dmitrenko et al., 2001a; Dmitrenko, un-
of these relatively warm Atlantic waters along publ. data). Following the distribution of the riv-
the palaeovalleys up to water depths of 20^ erine waters, the average summer surface water
40 m. Although the existing temperature variations salinity ranges from about 5 in the southeast to
are insigni¢cant for the benthic organisms in the 10^15 in the east and up to 30 in the west (Fig.
Laptev Sea, they appear to be good water mass 2a). In winter, due to reduced river runo¡ and
indicators. sea-ice formation, the average surface salinity is
Due to considerable river runo¡ and ice forma- higher, varying from 10 to 12 in the southeast to
tion in the Laptev Sea salinity is the most variable 33 in the west (Fig. 2b). Bottom salinity is less
feature that a¡ects the spatial distribution of dependent upon river runo¡. Both in summer
benthic organisms. In general, both surface and and in winter, it varies from 18^20 in the shallow
Fig. 2. (a) Average surface salinity in summer (Dmitrenko et al., 2001a). (b) Average surface salinity in winter (Dmitrenko et al., 2001a). (c) Average bottom salini-
ty in summer (Dmitrenko, Kirillov, unpubl. data). (d) Average bottom salinity in winter (Dmitrenko, Kirillov, unpubl. data). Isobaths in metres.
A. Stepanova et al. / Marine Micropaleontology 48 (2003) 23^48 27
Table 1
Location of stations, water depth, environmental data and percentage of live ostracod specimens
Station Water Latitude Longitude Temperature Salinity Sediment Live ostracod
depth (‡C) specimens
(m) (%)
PS-51/158-8 west 68 76‡57,49N 118‡35,37E 31.481 33.488 sandy silt 96
PS-51/159-8 west 61.6 76‡45,99N 116‡01,86E 31.617 33.687 clayey silt 95
PS-51/154-9 west 276.4 77‡16,61N 120‡36,03E 0.395 34.708 sandy silt 97
PM9402-3 west 47 75‡29,44N 115‡14,94E
PS-51/118-1 centre 121 77‡53,6N 132‡12,57E 30.7 sandy silt 80
PS-51/125-12 centre 127 77‡36,09N 130‡00,07E 31.54 silty sand 95
PS-51/114-13 centre 66 77‡35,52N 132‡15,82E 31.345 33.776 sandy^silty clay 99
PS-51/117-3 centre 76 77‡49,8N 132‡14,42E 30.949 34.105 sandy silt 92
PS-51/126-2 centre 85 77‡32,9N 130‡07,9E 31.54 silty sand 90
PS-51/135-2 east 51 76‡09,93N 133‡14,78E 31.6 sandy silt 100
PS-51/104-14 east 34 75‡57,83N 132‡09,06E 31.565 32.638 sandy silt 100
PS-51/106-1 east 33 75‡56,97N 132‡04,39E 31.5 silty sand 90
PS-51/105-3 east 33 75‡57,2N 132‡06,13E 31.590 32.821 sandy silt 94
PS-51/138-10 east 41 75‡09,18N 130‡49,75E 31.648 33.219 sandy silt 100
PS-51/094-3 south 31 74‡33,36N 130‡27,2E 31.592 32.673 silty sand ^
PS-51/093-1 south 33 74‡56,74N 130‡34,15E 31.608 32.992 sandy silt 100
PS-51/092-11 south 34 74‡35,5N 130‡08,4E 31.584 32.427 sandy^silty clay 100
PS-51/085-2 south 22 73‡33,9N 131‡16,3E 31.086 27.896 sandy^silty clay 98
PS-51/080-11 south 21 73‡27,83N 131‡39,0E 31.062 23.407 silty clay 85
IK9306-11 south 17.5 72‡00,63N 130‡59,23E clay
IK9307-3 south 20.7 72‡32,97N 131‡17,80E mud
IK93Z4-3 south 14 72‡01,90N 130‡07,55E silty clay
IK93Z5-4 south 11 71‡41,41N 137‡00,40E mud
PM9441-4 south 14 74‡00,00N 125‡59,29E
KD9541-13 south 22 73‡22,80N 129‡56,57E clayey silt
KD9565-11 south 21 73‡50,76N 120‡19,00E sandy silt
southeastern region to 30^34 at depths exceeding and from eight nearshore localities (water depths
30 m (Fig. 2c,d). down to 11 m) collected during the TRANS-
A general cyclonic circulation pattern is char- DRIFT I, II and III expeditions in 1993, 1994
acteristic of the surface and shelf bottom waters and 1995 (Table 1 ; Fig. 1). All samples from
(Dobrovol’skii and Zalogin, 1982; Pavlov and the TRANSDRIFT V expedition were obtained
Pavlov, 1999). The strong reversal currents (cur- using a giant box-corer. A total of 36 (two sam-
rent velocity up to 59 cm/s) recently recorded in ples per station) undisturbed surface samples (ap-
the submarine valleys make these troughs the proximately the uppermost centimetre) were taken
main area of interaction between the Arctic Ocean at 18 stations covering a water depth from 21 to
water and the Laptev Sea shelf water (Dmitrenko 276 m (Table 1). Coretop samples from the other
et al., 2001a,b). TRANSDRIFT expeditions were taken from both
box and kasten cores.
Both sets of samples were treated di¡erently.
3. Materials and methods The ¢rst set (V400 cm2 ) was stained with Rose
Bengal, washed over a 63-Wm mesh-size sieve on-
Coretop sediment samples were collected from board, and later used for estimating the percent-
di¡erent parts of the Laptev Sea shelf and upper age of ostracod species collected alive (Table 1).
continental slope during the Russian^German We considered both strongly and slightly Rose
TRANSDRIFT V expedition in August 1998 Bengal coloured valves as living ones. Many of
the strongly coloured carapaces had their chiti- mate the riverine in£uence on the ostracod com-
nous soft body parts preserved. The number of munity (Fig. 3b), following the method applied by
less brightly Rose Bengal coloured valves and car- Polyak et al. (in press) who distinguish between
apaces was by far lower than that of strongly benthic foraminiferal species in the Kara Sea on
coloured ones. Thus, we combined all these speci- the basis of three environmental categories : river-
mens into the group of live ostracods, although proximal ( 6 15), river-intermediate (15^25), and
certain problems exist in distinguishing dead from river-distal ( s 25).
living specimens using this method (see discussion By applying cluster analysis, the studied ostra-
in Brouwers et al., 2000). The second set of sedi- cod samples were grouped in relation to their
ment samples (100 cm2 ) was freeze-dried, weighed geographical location and, hence, environmental
and also washed over a 63-Wm mesh-size sieve. conditions (water depth, salinity variations). The
After sieving, ostracods were picked, identi¢ed percentage data of 39 taxa from 20 samples (sam-
and counted. Ostracod abundance was calculated ples containing 1^2 species were excluded) were
per 100 grams of dry bulk sediment. Percentages entered into the Statistica 4.3 software package.
of di¡erent ecological groups, i.e. marine and We used the 1-Pearson r coe⁄cient and average
euryhaline together with brackish-water species, linkage cluster analysis to attain the cluster den-
were calculated for this set of samples (Fig. 1). drogram (Fig. 4).
Although all ostracods are well preserved, there
does exist a certain di¡erence between the two sets
of samples. The samples that were ¢rst freeze- 4. Results
dried and then washed contained mainly single
valves, whereas those that were stained and 4.1. The Laptev Sea Ostracoda fauna
washed already onboard during the expedition
had considerable numbers of complete carapaces. A total of 1122 valves and 595 carapaces were
Ecological characteristics of the species found collected from the entire set of 36 samples. In all,
in our samples are based on both our own data 35 species were identi¢ed to species level belong-
and published evidence on their occurrences in ing to 19 genera and 12 families. Four species
other Arctic and high-latitudinal seas (e.g. Cro- were identi¢ed to generic level and two species
nin, 1977, 1981; Cronin et al., 1991, 1994; remained unidenti¢ed (species A, B). For identi¢-
McDougall et al., 1986; Brouwers, 1990, 1994; cation we followed the taxonomy provided by Ni-
Brouwers et al., 2000; Jones et al., 1998; Lev, kolaeva (1989), with additions from Athersuch et
1983; Nikolaeva, 1989; Neale and Howe, 1975; al. (1989) and Brouwers (1990). The complete list
Reimnitz et al., 1992). In order to demonstrate of taxa with numbers of plates and ¢gures in this
some possible environmental preferences of the paper together with corresponding references
ostracods, average frequencies of the more abun- from the literature is given in Table 2 (see Plates
dant species were plotted vs. water depth and I^IV). The entire collection is stored at Moscow
average summer surface salinity (Fig. 3a,b). In State University, Geological Faculty, Chair of Pa-
terms of water depth, four groups were distin- laeontology (No. 292/1-196).
guished, i.e. ostracods preferring the shallow inner
shelf (11^30 m), the middle-outer shelf (30^60 m), 4.2. Composition and abundance of ostracods in
the upper continental slope (60^100 m) environ- surface samples
ments, and depths exceeding 100 m (in our case
100^276 m) (Fig. 3a). We used the average In general, ostracod abundance is high in the
summer surface salinity as a factor to approxi- southern nearshore zone and in the Khatanga
Fig. 3. (a) Average frequencies of ostracod species vs. water depth in discrete increments. (b) Average frequencies of ostracod
species vs. average summer surface salinity in discrete increments.
a) b)
Fig. 4. (a) Cluster analysis dendrogram of 20 Laptev Sea samples based on the relative abundance of 39 ostracod taxa. (b) De-
scription of the obtained assemblages (clusters).
palaeovalley, ranging from about 300 to 600 speci- those parts of the submarine river valleys where
mens per 100 g dry bulk sediment (Fig. 1). The the strongest bottom currents have been observed
abundance is also relatively high on the continen- (I. Dmitrenko and J.A. Ho«lemann, pers. com-
tal slope (160^350 specimens). Although samples mun.). Thus, it is probable that dead valves are
from the eastern middle shelf contain consider- being constantly removed from these sites. Such a
ably less valves (27^113 specimens), they have situation is quite common for hydrodynamically
the highest abundance of living ostracods (up to active environments (Frenzel and Boomer, in
100%; Table 1). These stations are restricted to press). Also, possibly due to the same reasons,
31
32 A. Stepanova et al. / Marine Micropaleontology 48 (2003) 23^48
1 2 3
4 5 6
7 8 9
10 11 12
13 14 15
no juvenile valves were found in the samples from mainly corresponding to the western^central and
shallow areas and only few juveniles were recog- southern^eastern regions. Most stations in the
nised in the deeper western and central regions western and central Laptev Sea are located on
(Tables 3 and 4). Surface sediments in the Laptev the upper continental slope (62^276 m water
Sea are mainly represented by olive-brown sandy depth). The southern and eastern stations are re-
silt (Table 1). The highest abundance of ostracods stricted to the shallow shelf areas (11^51 m). Sam-
is usually recorded in the more clayey sediments, ples from the western^central region represent a
as found in the submarine valleys (stations 80, single assemblage (cluster IV; Fig. 4), with two
159; Fig. 1), while the coarser sandy sediments subdivisions according to their water depths.
occur in the hydrodynamically more active zones The di¡erence between assemblages from the east-
and show lower ostracod abundances (stations ern (cluster I) and southern (clusters II, III) re-
from the eastern middle shelf). gions is well pronounced (Fig. 4).
Taxonomic diversity was found to decrease in
eastward direction, from on average 15^16 species 4.2.1. Western^central Laptev Sea
in the west to 5^7 species in the east and south The ostracod assemblage from the western^cen-
(Tables 5 and 6). The lowest diversity (1^2 spe- tral Laptev Sea exhibits the highest taxonomic
cies) was recorded in the nearshore zone close to diversity among the studied samples (Tables 3
the Lena delta and Yana estuary (Table 5) and in and 4). It is distinguished from the other assem-
the samples obtained in the Sannikov Strait near blages by the presence of relatively deep-living
New Siberian Islands and in the Yana valley species (Fig. 3b) such as Pseudocythere caudata,
(MAPD; Cronin et al., 1991). Krithe glacialis, K. minima, Cytheropteron in£a-
As the studied samples were obtained from geo- tum, C. biconvexa, C. nodosoalatum, C. simplex,
graphically di¡erent localities, they could be di- C. porterae, C. testudo, Polycope punctata, P. or-
vided into three groups: a western^central group, bicularis and P. bireticulata, all of which are typ-
a southern and an eastern group (Fig. 1 ; Tables ical of the Arctic seas and the North Atlantic
3^6). This geographical grouping is also sup- (Whatley et al., 1998; Cronin, 1989, 1996; Cronin
ported by cluster analysis (Fig. 4). The obtained et al., 1994, 1995; Joy and Clark, 1977; Jones et
dendrogram clearly de¢nes two major groupings al., 1998, 1999; Didie¤, 2001). However, their
Plate I.
1^3. Bythocythere constricta Sars, 1926; (1) MSU292/185, sample PS51/118-1, left valve, external view, U61; (2) MSU292/
169, sample PS51/117-3, right valve, external view, U74; (3) MSU292/189, sample PS51/158-8, left valve, internal
view, U61.
4. Pseudocythere caudata Sars, 1866; MSU292/177, sample PS51/154-9, carapace, U83.
5. Sclerochilus contortus (Norman, 1862); MSU292/188, sample PS51/158-8, left valve, external view, U62.
6. Jonesia acuminata (Sars, 1866); MSU292/163, sample PS51/105-3, carapace, U38
7, 8. Roundstonia macchesneyi (Brady and Crosskey, 1871); sample PS51/138-12, core section, 355^358 cm; (7) MSU292/
191, right valve, external view, U100; (8) MSU292/192, right valve, internal view, U103
9, 10. Cluthia cluthae (Brady, Crosskey and Robertson, 1874); sample 118-1; (9) MSU292/184, left valve, external view,
U125; (10) MSU292/74, right valve, internal view, U100.
11. Cytheropteron suzdalskyi Lev, 1972; MSU292/140, sample PS51/135-4, core section, 390^393 cm, left valve, external
view, U79.
12, 13. Cytheropteron sulense Lev, 1972; (12) MSU292/179, sample PS51/085-2, right valve, external view, U65; (13)
MSU292/135, sample PS51/135-4, core section, 490^493 cm, right valve, external view, U85
14. Cytheropteron elaeni Cronin, 1989; MSU292/144, sample PS51/135-4, core section, 390^393 cm, right valve, external
view, U102
15. Cytheropteron montrosiense (Brady, Crosskey and Robertson, 1874); MSU292/180, sample PS51/085-2, right valve,
external view, U94.
1 2 3
4 5 6
7 8 9
10 11 12
13 14 15
relative abundance in the samples as well as the Sea (33^51 m) form a single, well-de¢ned assem-
total species composition change with depth (Fig. blage (cluster I; Fig. 4) which is dominated by
4). Most samples combined into cluster IVa are four normal marine middle and outer neritic spe-
dominated by C. porterae, Cytheropteron arcua- cies (Table 5 ; Fig. 3a,b). Primarily, these are
tum and, with increasing depth, by Sarsicytheridea Acanthocythereis dunelmensis together with Semi-
punctillata and Polycope spp. (P. punctata, P. or- cytherura complanata, Elofsonella concinna and
bicularis, P. bireticulata and Polycope sp.). Three Cluthia cluthae. A single sample from the south-
samples, comprising cluster IVb, are dominated ern Laptev Sea (KD9565) is included into the
by shallow-water species Rabilimis septentrionalis same cluster, probably because it was obtained
and Heterocyprideis sorbyana, but also include from the shallow region to the west o¡ the Lena
many of the above-mentioned deep-living forms. delta where there is considerably less river runo¡
Increased abundance of shallow-water species that than in the area to the east and north o¡ it.
were found alive in these samples could result Therefore, the taxonomic composition of sample
from ice-rafting that is a well-known factor a¡ect- KD9565 also di¡ers from all other southern sam-
ing the distribution of ostracods in the Arctic ples.
(Jones et al., 1998, 1999; Cronin et al., 1994).
As seen in Fig. 3a, R. septentrionalis and H. sor- 4.2.3. Southern Laptev Sea
byana have in fact two abundance maxima ^ in Ostracod samples from the shallow southern
the shallow inner shelf zone and on the upper Laptev Sea are quite di¡erent from most other
continental slope within the water depth range samples, being dominated by inner shelf euryha-
of 60 to 100 m. Many specimens of these species line species (Table 6 ; Fig. 3a,b). Taxonomic diver-
are represented by juvenile valves and carapaces sity sharply decreases, down to 1^2 species per
(Tables 3 and 4). This all might point to the im- sample in the most shallow nearshore zone (sta-
portance of ice-rafting in distributing shallow- tions IK9307, IK93Z5, KD9541). The highest
water species across the Laptev Sea shelf. abundance of valves was recorded in the near-
shore area (600 specimens/100 g at station 80),
4.2.2. Eastern Laptev Sea whereas numbers sharply decrease northward
Samples from the eastern part of the Laptev (down to 27 at station 94). Samples from the
Plate II.
1. Cytheropteron in£atum (Brady, Crosskey and Robertson, 1874); MSU292/71, sample PS51/118-1, left valve, external
view, U70.
2. Cytheropteron champlainum Cronin, 1981; MSU292/59, sample PS51/117-3, right valve, external view, U86.
3, 4. Cytheropteron nodosoalatum Neale and Howe, 1973; (3) MSU292/81, sample PS51/158-8, left valve, external view,
U87; (4) MSU292/39, sample PS51/138-12, core section, 151^154 cm, left valve, external view, U85.
5, 6. Cytheropteron testudo Sars, 1869; sample PS51/118-1; (5) MSU292/69, right valve, external view, U96; (6) MSU292/
71, left valve, external view, U120.
7, 8. Cytheropteron biconvexa Whatley and Masson, 1979; (7) MSU292/76, sample PS51/118-1, left valve, external view,
U115; (8) MSU292/85, sample PS51/158-8, left valve, internal view, U100.
9. Cytheropteron inornatum Brady and Robertson, 1872; MSU292/84, sample PS51/158-8, right valve, external view,
U97.
10, 11. Cytheropteron porterae Whatley and Coles, 1987; (10) MSU292/61, sample PS51/117-3, right valve, external view,
U80; (11) MSU292/89, sample PS51/158-8, left valve, external view, U90.
12. Cytheropteron simplex Whatley and Masson, 1979; MSU292/60, sample PS51/117-3, right valve, external view, U96.
13, 14. Cytheropteron arcuatum (Brady, Crosskey and Robertson, 1874); sample PS51/118-1; (13) MSU292/78, left valve,
external view, U97; (14) MSU292/79, right valve, internal view, U115.
15. Semicytherura complanata (Brady, Crosskey and Robertson, 1874); MSU292/194, sample PS51/135-2, right valve,
external view, U70.
1 2 3
4 5 6
7 8 9
10 11 12
13 14 15
southern region comprised of more than two spe- water species Roundstonia macchesneyi were also
cies were arranged into clusters II and III (Fig. 4). identi¢ed.
Unlike the other regions, clustering of the taxo-
nomically poor ostracod samples from this area
with the most changeable environmental condi- 5. Discussion
tions gave rather controversial results. Cluster
III (stations 85 and IK9306) was included into 5.1. Distribution of recent ostracods in relation to
the major group of the western^central assem- environmental parameters
blage. This is only because these two samples con-
tain species of the Polycope genus (P. punctata, P. The data presented above show that there is
orbicularis, P. bireticulata and Polycope sp.) and signi¢cant variability in the distribution of ostra-
Bythocythere constricta, which are absent in the cod species over the Laptev Sea shelf and upper
remaining samples from the shallow eastern Lap- slope area, re£ecting varying environmental con-
tev Sea (Table 6). Also, cluster II includes one ditions (Fig. 3a,b). In terms of water depth, the
sample (PM9402) from the western Laptev Sea most pronounced di¡erences are found between
(47 m water depth) which is dominated by the the western^central assemblage (upper continental
shallow-water euryhaline species Paracyprideis slope, water depths s 60 m) and assemblages
pseudopunctillata and Heterocyprideis sorbyana. from the southern and eastern shelf regions (water
Paracyprideis pseudopunctillata predominates in depths less than 60 m). A comparison of the as-
the majority of samples from the southern region semblages on the basis of the Jaccard index (Jac-
(Fig. 3a,b). H. sorbyana and Semicytherura com- card, 1912) shows that there is a strong similarity
planata are subdominant species. All three species between the eastern and southern assemblages
are known to inhabit shallow inner shelf environ- (60%). On the contrary, the values of the Jaccard
ments (Nikolaeva, 1989; McDougall et al., 1986; index for the western^central assemblage, on the
Reimnitz et al., 1993; Brouwers et al., 1991; one hand, and the eastern^southern assemblages,
Brouwers et al., 2000). Two of them, except S. on the other hand, are less than 50%. This implies
complanata, are known to tolerate salinities as that the taxonomic composition of these two re-
low as 5^10 (Cronin, 1977; Neale and Howe, gions is signi¢cantly di¡erent. The relatively deep-
1975; Lev, 1983). Several valves of the brackish- living species occurring here are also known from
Plate III.
1. Semicytherura complanata (Brady, Crosskey and Robertson, 1874); MSU292/44, sample PS51/138-12, core section,
151^154 cm, right valve, internal view, U68.
2, 3. Palmenella limicola (Norman, 1865); sample PS51/135-2; (2) MSU292/153, right valve, external view, U100; (3)
MSU292/193, right valve, internal view, U84.
4, 5. Acanthocythereis dunelmensis (Norman, 1865); (4) MSU292/148, sample PS51/135-4, core section, 390^393 cm, right
valve, external view, U53; (5) MSU292/149, sample PS51/114-13, right valve, internal view, U43.
6. Rabilimis septentrionalis (Brady, 1868); MSU292/95, sample PS51/159-8, right valve, external view, U57.
7, 8. Heterocyprideis sorbyana (Jones, 1857); sample PS51/114-13; (7) MSU292/173, left valve, external view, U47; (8)
MSU292/174, right valve, internal view, U61.
9, 10. Heterocyprideis fascis (Brady and Norman, 1889); sample PM942-3, left valves, external view; (9) MSU292/205, U65;
(10) MSU292/206, U55.
11^13. Sarsicytheridea punctillata (Brady, 1864); (11) MSU292/171, sample PS51/114-13, right valve, external view, U63; (12)
MSU292/170, sample PS51/114-13, left valve, internal view, U56; (13) MSU292/101, sample PS51/154-9, left juvenile
valve, external view, U75.
14. Sarsicytheridea bradii (Norman, 1864); MSU292/96, sample PS51/159-8, right valve, external view, U57.
15. Krithe sp.; MSU292/62, sample PS51/117-3, left valve, external view, U75.
1 2 3
4 5 6
7 8 9
10 11 12
13 14 15
16 17
the North Atlantic and the Arctic Ocean. This is macchesneyi found in this assemblage tolerates a
not too surprising because the western^central low and £uctuating salinity and at times has been
Laptev Sea region is in£uenced by Atlantic waters found together with freshwater species (Schoning
penetrating the upper continental slope and outer and Wastegafird, 1999). This is the only assemblage
shelf (Dobrovol’skii and Zalogin, 1982). where the share of brackish-water and euryhaline
Salinity also seems to have an important impact species reaches 76^88%, thus exceeding that of
on the distribution of ostracods in the eastern marine species (Fig. 1).
Laptev Sea, especially, where there is a strong With increasing water depth and salinity, shal-
e¡ect of river runo¡. The average bottom water low normal-marine species become dominant
salinity ranges from 26 to 32 (Fig. 2), and the (Fig. 3b). These include mainly Acanthocythereis
bottom salinity measured in this area during the dunelmensis but also Semicytherura complanata,
expedition in 1998 varied from 23^27 at 21 m Cluthia cluthae and Elofsonella concinna which to-
water depth to 32 at 40^50 m water depth (Table gether sometimes comprise 100% of the assem-
1). It is known that taxonomic diversity usually blage (Fig. 1). This assemblage is also character-
decreases in lower salinities, while total abun- istic of the area where the £aw polynya forms
dance of ostracod valves is often greater (Frenzel during wintertime. A similar assemblage was
and Boomer, in press). Compared to the assem- found in sea-ice sediments sampled from an ice
blage from the western^central region with nor- £oe in the Beaufort Sea (Reimnitz et al., 1993).
mal marine salinity, the total number of species As shown by Reimnitz et al. (1992, 1993), the
found in the eastern and southern assemblages is inner and middle shelf (10 to 30^40 m water
considerably lower (20 vs. 33^34 species). How- depth) is the main area where benthic organisms
ever, the highest abundance of ostracod valves can be easily entrained into ice, mainly through
was recorded in the nearshore part of the Lena the mechanism of anchor ice formation. Some
valley (600 specimens/100 g). The ostracod assem- shallow-water species (Heterocyprideis sorbyana,
blage (Table 6) of the southern inner shelf zone S. complanata, C. cluthae, Roundstonia macches-
(11^33 m) with highly variable environmental neyi) have been reported even from central Arctic
conditions is enriched in euryhaline species Para- Ocean surface sediments as part of the ‘ice-rafted
cyprideis pseudopunctillata and Heterocyprideis assemblage’ (Jones et al., 1998, 1999; Cronin et
sorbyana. The brackish-water species Roundstonia al., 1994). Therefore, it may be concluded that
Plate IV.
1, 2. Krithe minima Coles, Whatley and Moguilevsky, 1994; sample PS51/158-8; (1) MSU292/82, right valve, external view,
U57; (2) MSU292/83, right valve, internal view, U62.
3, 4 Krithe glacialis (Brady, Crosskey and Robertson, 1874); sample PS51/118-1; (3) MSU292/65, left valve, external view,
U57; (4) MSU292/66, left valve, internal view, U62.
5^7. Paracyprideis pseudopunctillata (Swain, 1963); (5) MSU292/118, sample PS51/092-11, left valve, external view, U55;
(6) MSU292/116, sample PS51/092-11, right valve, external view, U52; (7) MSU292/8, sample PS51/138-12, core
section, 481^484 cm, left valve, internal view, U52.
8. Argilloecia cylindrica Sars, 1923; MSU292/100, sample PS51/126-2, right valve, external view, U94.
9, 10. Argilloecia sp.; (9) MSU292/93, sample PS51/114-6, right valve, external view, U96; (10) MSU292/64, sample PS51/
117-3, carapace, U72.
11, 12. Elofsonella concinna (Jones, 1857); sample PS51/114-6; (11) MSU292/90, right valve, external view, U49; (12)
MSU292/91, right valve, internal view, U47.
13. Elofsonella a¡. concinna (Jones, 1857); MSU292/94, sample PS51/159-8, right valve, external view, U92.
14, 15. Polycope bireticulata Joy and Clark, 1977; (14) MSU292/166, sample PS51/117-3, carapace, U75; (15) MSU292/195,
sample PS51/080-11, carapace, U90.
16. Polycope orbicularis Sars, 1866; MSU292/165, sample PS51/117-3, carapace, U92.
17. Polycope sp.; MSU292/175, sample PS51/125-12, carapace, U56.
Table 3
Occurrence of ostracods in coretop samples from the western Laptev Sea
Species Station 158-8 Station 159-8 Station 154-9 Station PM9402-3
stained unstained stained unstained stained unstained
Bythocythere constricta 4v3c
Pseudocythere caudata 2c 2c
Sclerochilus contortus 1v8c
Jonesia acuminata 4v2c
Cluthia cluthae 1v8c 3v 1v 9v 6c 10v
Cytheropteron sulense 2c 2v
C. arcuatum 2v6c 1v 1v 3v 1v2c
C. porterae 2v6c 15v 4v2c 4v1c 3v
C. testudo 2c 1v
C. simplex 3v5c 6v 5v13c
C. inornatum 2v
C. biconvexa 2v
C. in£atum 4v1jc 1v 1v
C. nodosoalatum 1v1c 4v
C. cf. nodosum 1v
Semicytherura complanata 2c 5v
Palmenella limicola 4v
Acanthocythereis dunelmensis 2v 2v
Rabilimis septentrionalis 1v8c 5v2j 34v13c 1v 2v1c 11v7j
Heterocyprideis sorbyana 2c 3v 16v4c 2v 8v 7v 10v
H. fascis 16v
Heterocyprideis sp. juv. 4j
Sarsicytheridea bradii 3v 6v
S. punctillata 2v 2v 18v1c14j 58v16c
Paracyprideis pseudopunctillata 1c 1v 23v25c 22v 15v
Krithe minima 1c 2v 2v
K. glacialis 8v1c 9v 1v 1c
Krithe sp. 1v
Argilloecia cylindrica 1c 1v2c
Elofsonella concinna 3v 3v
Elofsonella a¡. concinna 1v 1v 10v
Eucythere cf. argus 1v
Polycope punctata 1c 1c
Polycope spp. 2c 1v9c
Undetermined, species A 1v
Rose Bengal used for staining. Abbreviations: v, single valves; c, carapaces; j, juvenile valves; jc, juvenile carapaces.
many of the shallow-water species found in our Abramova, pers. commun.). A survival of macro-
samples from the upper continental slope (Rabi- epibenthic organisms that were frozen into fast ice
limis septentrionalis, H. sorbyana, Paracyprideis is reported from the Gulf of St. Lawrence (Med-
pseudopunctillata) were also ice-rafted. It is inter- cof and Thomas, 1974). Therefore, it seems con-
esting in this respect that most ostracods (90^ ceivable that some shallow-water species are able
95%) in our samples from the TRANSDRIFT V to settle onto the sea£oor after having been ice-
expedition were found stained. It is unknown rafted.
whether ostracods can survive ice-rafting. How-
ever, it has been observed that some other crus- 5.2. Comparisons with other Arctic areas
taceans (4th and 5th copepodite stages of cyclops)
remained alive after being frozen into riverine and One of the ¢rst attempts to review Arctic shal-
lacustrine ice in the Lena River deltaic zone (E. low-water ostracod faunas was undertaken by
Table 4
Occurrence of ostracods in coretop samples from the central Laptev Sea
Species Station 118-1 Station 125-12 Station 114-3 Station 117-3 Station 126-2
stained unstained stained unstained stained unstained stained unstained stained unstained
Bythocythere constricta 1v2c 1v1c
Cluthia cluthae 8c 1c 4v 5v 12c 4v 5c 4v
Cytheropteron sulense 1v
C. arcuatum 9v 4v 1c 6c 2v 2v2c 12v
C. porterae 7v1c 5v 5v1c 1v 2v2c 5v 3v 2v
C. testudo 3v 4v 1v1c 1v3c 3v 4c
C. simplex 7v1c 2v1c 2v1c 1v 2v1c
C. biconvexa 2c 2c 1v
C. nodosoalatum 1v
C. champlainum 1v 2v
C. in£atum 7v 4v 1v
Semicytherura complanata 2v 2v 1v
Acanthocythereis dunelmensis 5v2c 4c 1v
Rabilimis septentrionalis 2v1c 9v16c 17vlj 1v1c 1v 3v5c 8v50j
Heterocyprideis sorbyana 5v 5v2c 3v 26v47c 25v1c 2v1c 4v 5v1c
Sarsicytheridea punctillata 6v1c 2v 19v5c 18v3c 4v1c 2v 8v3c 11v1c28j 4v 5v1c
S. bradii 14v 1v uv
Paracyprideis pseudopunctillata 3v 3v 8v 1v 3v2c 1v
Krithe minima 4v 2v1c 2v
K. glacialis 3v1c 6v 2v 5v4c 6v 1v 1v 8v4c 3v
Argilloecia cylindrica 1c 3c 2v 1v
Argilloecia sp. 2v
Elofsonella concinna 3v2c 6v
E. a¡. concinna 1c
Eucythere cf. argus 4v
Polycope spp. 10c 4v6c 3v13c 1c
Undetermined, species A 1v 1v
Rose Bengal used for staining. Abbreviation: v, single valves; c, carapaces; j, juvenile valves.
Table 5
Occurrence of ostracods in coretop samples from the eastern Laptev Sea
Species Station 104-14 Station 105-3 Station 106-1 Station 135-2 Station 138-10
stained unstained stained unstained stained unstained stained unstained stained unstained
Jonesia acuminata 1c 2v3c 1v
Roundstonia macchesneyi 1v 1c
Cluthia cluthae 2v 4v14c 3v 6c 10c
Cytheropteron arcuatum 2c 4v7c
C. elaeni 2v 3v7c 1v
C. suzdalskyi 1v 1v5c 3v1c
C. nodosoalatum 1v 1v 1v
Semicytherura complanata 9v 4c 4v 5c 3v11c 2c 6v1c
Palmenella limicola 1c 4v5c
Acanthocythereis dunelmensis 1v15c 14v 4v8c 6v 4v4c 3v 1v7c 2v
Sarsicytheridea bradii 3v
Paracyprideis pseudopunctillata 2c 1c 4v2c 1v3c 11v 1v1c
Argilloecia cylindrica 1v5c 1v 1v
Elofsonella concinna 1v2c 4v 4v4c 1v
Elofsonella a¡. concinna 4c 1v 2c
Polycope spp. 2v1c
Rose Bengal used for staining. Abbreviations: v, single valves; c, carapaces.
Bythocythere constricta 1c
Roundstonia macchesneyi 1v 1f 3v
Cluthia cluthae 1c 1v
Cytheropteron sulense 2v3c 1v 1c 3v 7v 1v
MARMIC 906 28-3-03
C. arcuatum 1c
C. montrosiense 1v3c 1v 2c 2v
C. elaeni 1v 1c 1v
C. suzdalskyi
Semicytherura complanata 10c 1v 2c 1v 1v
Palmenella limicola 2c 1v
Rabilimis septentrionalis 22v1c
Acanthocythereis dunelmensis 1c 3v
Heterocyprideis sorbyana 1c 9v2c 5v 1v 6v1c
Paracyprideis pseudopunctillata 3v3c 21v 17v17c 27v 6v1c 2v1c 23v 3v1c 41v10c 2v 2v
P. cf pseudopunctillata 1v
Elofsonella concinna 1v1c
E. a¡. concinna 1v 1v
Polycope spp. 7c 2v9c 1v
Undetermined, species B 1v
Rose Bengal used for staining. Abbreviations: v, single valves; c, carapaces; f, fragments.
Table 7
Laptev Sea Ostracoda compared with published materials from other Arctic regions
Laptev Sea Ostracoda (own data)a White Seab Barents Seac;d Kara East Chukchi Greenland Sea Western Greenland Beaufort
Sead Siberian Sead and coast and eastern Seac;d;f
Sead Scoresby Sunde Canadian Arcticc;d
Bythocythere constricta + + + + + +
Pseudocythere caudata + + + + +
Sclerochilus contortus + + + + + + + +
C. inornatum
C. montrosiense + + + +
C. nodosoalatum + + + + + +
C. porterae +
C. simplex + + +
C. sulense +
C. suzdalskyi +
C. testudo + +
Semicytherura complanata + +
Palmenella limicola + + + + + + +
Acanthocythereis dunelmensis + + + + + + + + + +
Rabilimis septentrionalis + + + + + + +
Heterocyprideis sorbyana + + + + + + + + + +
H. fascis + +
Sarsicytheridea bradii + + + + + + + + + +
S. punctillata + + + + + + + + + +
Paracyprideis pseudopunctillata + +
Krithe glacialis + +
K. minima +
Argilloecia cylindrica +
Elofsonella concinna + + + +
Eucythere argus +
Polycope bireticulata +
P. orbicularis + +
P. punctata +
Robertsonites tuberculatusa + + + + + +
Roundstonia globuliferaa + + + +
Elofsonella neoconcinnaa + +
Semicytherura a⁄nisa + + + +
43
44 Beaufort
A. Stepanova et al. / Marine Micropaleontology 48 (2003) 23^48
1^1000
Seac;d;f
V90
shown that, in general, the shallow-water fauna
31
34
+
forms a circumpolar province, although with cer-
Western Greenland
Canadian Arcticc;d
coast and eastern
2^3065
er Arctic regions reveals similar features in the
46
14
274^3355
tal species list of the Laptev Sea Ostracoda (our
data with additions from Cronin et al., 1991) and
the occurrence of these species in other Arctic
and
61
15
25
38
9
223^50021^37
Sead
10
60
+
40
+
20
22
+
Land
3^55
31
12
39
78
White Seab Barents Seac;d
31
15
+
25
6
Atlantic water.
Total number of species
d
a
e
f
long to the group of relatively deep-living species the Laptev Sea. Thus, nearly all species found in
found in the western^central Laptev Sea (Table the Laptev Sea are also present in the Beaufort
7). Sea, with the exception of some rare deep-living
Just as for the Kara Sea, the data on the East species of mainly North Atlantic origin (Pseudo-
Siberian Sea ostracods are given by Cronin et al. cythere caudata, Cytheropteron testudo, C. porte-
(1991). The data on ostracods are non-represen- rae).
tative, because most of the samples contain only Based on the comparisons above, it can be con-
one to three valves, and only two samples out of a cluded that the Laptev Sea ostracod assemblages
total of nine included more than 30 valves. form a link between the western Arctic assem-
From ¢ve samples recovered from the Chukchi blages, a¡ected by North Atlantic waters, and
Sea Cronin et al. (1991) identi¢ed 24 species of shallow-water assemblages also known from the
which nine are also found in the Laptev Sea (Ta- Alaskan coast. Although many species occur
ble 7). Most of them are typical representatives of throughout all Arctic seas, their abundance, com-
the ostracod assemblages from the eastern and bination of species in certain assemblages and
southern Laptev Sea. presence of indicator species allow to distinguish
Whatley et al. (1996, 1998) list 61 species from between two geographically di¡erent assemblage
the Greenland Sea and Scoresby Sund fjord sys- groups in the Laptev Sea. The most characteristic
tem. Some of these species occasionally also occur feature of the western and central Laptev Sea is
in samples from the deep western and central the presence of relatively deep-living species with
Laptev Sea area associated with Atlantic waters. clear a⁄nities to the North Atlantic and the Arc-
Neale and Howe (1975) describe Recent ostra- tic Ocean. On the other hand, due to similar en-
cods collected in fjords from the Western Green- vironmental conditions (considerable river dis-
land coast and Eastern Canadian Arctic. This charge, ice coverage, existence of polynya and
rather diverse fauna does not have much in com- ice-rafting), ostracod assemblages from the south-
mon with the assemblages from the Laptev Sea ern and eastern Laptev Sea have many species in
(Table 7). In the western Greenland coastal common with the inner and middle shelf assem-
area, all species that are also found in the Laptev blages of the Beaufort Sea.
Sea, are extremely rare and occur at shallow-
water locations.
Ostracods of the Beaufort Sea have been 6. Conclusions
studied in great detail. Neale and Howe (1975)
describe ostracods from the Colville River delta We have investigated Recent ostracods from
that represent a very typical shallow-water assem- surface samples of the Laptev Sea shelf and upper
blage. It is very similar to the southern Laptev slope in an attempt to provide a more complete
Sea inner shelf assemblage and consists of shal- list of species from an area which thusfar has
low-water euryhaline species (Table 7). Reimnitz remained practically unstudied in terms of its os-
et al. (1993) give a list of 13 ostracods entrained tracod fauna. Moreover, the spatial distribution
by sea ice from the shallow middle shelf of the patterns of ostracods in relation to di¡erent envi-
Beaufort Sea. Nine of these species were identi¢ed ronmental parameters were evaluated.
in the Laptev Sea (Table 7). As noted above, this The recognised ostracod fauna consists of a to-
assemblage is very similar to that from the eastern tal of 41 species, of which 35 were identi¢ed to
Laptev Sea shelf, where the polynya forms during species level. Three di¡erent assemblages were es-
wintertime. Cronin et al. (1991) analysed more tablished according to their water depth. The
than 300 samples from the Beaufort Sea obtained western^central assemblage from the upper conti-
from water depths ranging between 1 and more nental slope, related to Atlantic waters, is the
than 1000 m. Nearly all species ( s 90) listed in richest in specimens abundance and diversity. It
the MAPD (Cronin et al., 1991) occur in the includes deep-living species that are absent in
Beaufort Sea, 34 of them are in common with the shallow eastern and southern shelf regions.
The western^central assemblage exhibits the clos- seev (MSU) and N. Kupriyanova (VNIIOkean-
est a⁄nity to the North Atlantic and Arctic geologiya) for valuable discussions. I. Dmitrenko
Ocean faunas (Cytheropteron biconvexa, C. testu- and S. Kirillov (AARI) provided us with their
do, C. simplex, C. nodosoalatum, C. in£atum, C. oceanographic data. The manuscript greatly bene-
porterae, Krithe glacialis, K. minima, Pseudocy- ¢ted from review comments made by T. Cronin
there caudata, Polycope punctata, P. orbicularis). and C. Didie¤.
The eastern shelf region is occupied by the Acan-
thocythereis dunelmensis assemblage consisting of
several shallow, normal-marine species. This as- References
semblage occurs in the region where a £aw poly-
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Recent shallow marine ostracods from high latitudes: Im-
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