Diversity of shell-bearing gastropods along
the western coast of the Arctic archipelago
Novaya Zemlya: an evaluation of modern
and historical data
Ivan O. Nekhaev & Ekaterina N. Krol
Polar Biology
ISSN 0722-4060
Polar Biol
DOI 10.1007/s00300-017-2140-1
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DOI 10.1007/s00300-017-2140-1
ORIGINAL PAPER
Diversity of shell-bearing gastropods along the western coast
of the Arctic archipelago Novaya Zemlya: an evaluation
of modern and historical data
Ivan O. Nekhaev1
•
Ekaterina N. Krol2
Received: 27 June 2016 / Revised: 15 May 2017 / Accepted: 6 June 2017
Ó Springer-Verlag GmbH Germany 2017
Abstract Accurate estimation of biodiversity is necessary
to provide a baseline for further ecosystem investigations
and protection of High Arctic regions. Novaya Zemlya is a
large Arctic archipelago located on the border between the
Barents and Kara Seas. Despite a long history of investigation, data on species composition along the coast of
Novaya Zemlya are presented only in sources from the
initial period of investigation in the late 19th to early 20th
century. We estimate herein the diversity of shell-bearing
Gastropoda found along the western coast of Novaya
Zemlya, based on both published sources and our original
identification of material collected between 1985 and 2008.
The total species list contains 86 species, 24 of which were
identified only from the recent material, while 28 were only
reported in literature. Our results demonstrate that modern
and historical data are significantly different. However,
these differences do not indicate any changes in species
composition in the region during the last century, but rather
arise for methodological reasons. Even combined, the two
types of data do not provide complete coverage of the
fauna. We also carried out a preliminary comparison
Electronic supplementary material The online version of this
article (doi:10.1007/s00300-017-2140-1) contains supplementary
material, which is available to authorized users.
& Ivan O. Nekhaev
inekhaev@gmail.com
Ekaterina N. Krol
krol.katerina@gmail.com
1
Laboratory of Macroecology and Biogeography of
Invertebrates, Saint Petersburg State University,
Universitetskaya emb. 7-9, Saint Petersburg, Russia 199034
2
Department of Biology, Murmansk State Technical
University, Sportivnaya str. 13, Murmansk, Russia 183010
between local coastal gastropod faunas from various parts
of the Barents Sea (including Novaya Zemlya).
Keywords Arctic Barents Sea Gastropoda Species
diversity Novaya Zemlya Historical data
Introduction
Arctic marine ecosystems are believed to have been
transformed recently due to the influence of multiple factors, including climatic changes, pollution, effects of fisheries, and invasive species (Britaev et al. 2010; Jørgensen
et al. 2015). One of the most important metrics of these
changes is alteration in taxonomical diversity over time, as
revealed by comparing modern and earlier investigations.
Such differences are often interpreted as substantial changes of the ecosystem structure influenced by local and/or
global factors (Britaev et al. 2010; Lyubina et al. 2014).
However, this approach has a significant drawback: Real
changes may be overestimated if the results of either of the
studies are incomplete (Nekhaev 2016). Given the limited
baseline knowledge regarding the Arctic fauna and
ecosystems in many regions, such incompleteness of biodiversity studies may be even harder to identify, and the
practical limitations of poor accessibility of many Arctic
regions make frequent and regular monitoring studies
difficult.
Only some large High Arctic marine shallow-water
ecosystems, especially coasts of Svalbard and Western
Greenland, undergo more or less regular hydrobiological
examination (Hop et al. 2002; Palerud et al. 2004; Beuchel
and Gulliksen 2008; Yesson et al. 2015). Both sites are
located in the Atlantic Ocean and are influenced by the tails
of the warm North Atlantic Current. By contrast,
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knowledge about bottom fauna of other Eurasian Arctic
archipelagos, such as Severnaya Zemlya, the New Siberian
Islands, Franz Josef Land, and Novaya Zemlya, is considerably poorer and limited to only occasionally collected
data (Gorbunov 1949; Golikov and Scarlato 1977; Denisenko et al. 1995). More detailed and accurate estimation
of biodiversity in these regions is necessary to provide a
baseline for further ecological research.
Gastropoda is one of the key groups of marine benthic
invertebrates. Aside from great taxonomical diversity and
abundance, gastropods demonstrate a wide spectrum of life
strategies and ecological preferences: from short-lived
algae eaters to long-lived predators or even parasites. Also,
some gastropod species are relatively easy to recognize
using simple shell characteristics. This makes gastropods
suitable as indicators of environmental conditions and
popular objects for monitoring studies in various environments (Golikov 1968; Zvonareva et al. 2015). Novaya
Zemlya is one of the largest but most poorly studied Arctic
archipelagos of Eurasia, located on the border between the
Barents and Kara Seas. Investigations of its environment
and fauna started in the early 18th century. The first report
on mollusks inhabiting the waters around the archipelago
was presented by Middendorff (1849) based on samples
collected during Baer’s expedition in 1837. Later, species
lists of mollusks from various parts of Novaya Zemlya
were published after the 1875–1876 and 1878–1879 cruises
by N.A.E. Nordenskiöld (Leche 1878; Aurivillius 1885).
Another survey of the archipelago was carried out in
1908; based on the material collected by this expedition, a
new species, Natica tenuistriata Dautzenberg et Fischer,
1911 (now Euspira tenuistriata), was described and an
annotated list of molluskan taxa published (Dautzenberg
and Fischer 1911). Dautzenberg also identified a small
collection of mollusks taken in a Russian survey in 1910
(Ivanov 1910). Substantial zoological material, including
marine invertebrates, was collected during the Norwegian
expedition to Novaya Zemlya in 1921, and a report on
mollusks was published by Grieg (1924).
The next stage of studies of Novaya Zemlya started in
the 1920s as part of investigations of the environment
along the Northeastern Passage, a shipping route between
the Atlantic and Pacific Oceans. Detailed species lists of
benthic animals and descriptions of the environmental
conditions were published for three studied areas: Moller
Bay, Chernaya Inlet, and Matochkin Shar Strait (Ushakov
1927; Gurjanova and Ushakov 1928; Ushakov 1931).
Additionally, the new species Trichotropis herzensteini
Derjugin et Gurjanova, 1926 (now synonymized with Trichotropis borealis Broderip et Sowerby, 1829) was
described (Derjugin and Gurjanova 1926).
Publications based on later surveys of the region are
surprisingly scarce, and often present only brief ecological
123
summaries (e.g., Golikov and Averintcev 1977; Denisenko
et al. 1995; Pogrebov et al. 1997). Material collected during this period for the most part was not properly preserved
and is now not available for reexamination. However, some
portions of it are dispersed between taxonomic museum
collections or remain unidentified. Careful examination of
these samples may complement older data obtained during
the initial phase of investigation, although the latter still
remain the most useful source of faunistic information on
the Novaya Zemlya region.
The main objective of this study is to estimate the
species diversity of shell-bearing gastropod mollusks along
the western coast of Novaya Zemlya based on available
original material, taken during the cruises of past decades,
and published sources. We also compared the two datasets,
i.e., modern and historical, to determine whether either
dataset separately or both of them together can provide
reliable information regarding the fauna of the archipelago.
Materials and methods
Study area
The Arctic archipelago Novaya Zemlya (Hodaz Pevkz,
Novaja Semlja) is located between the Barents and Kara
Seas and consists of two major islands—Southern and
Northern—and several hundred smaller islands around
them. The Southern and Northern islands are divided by a
very narrow (1–2 km) strait called Matochkin Shar. The
shoreline of the archipelago is indented by numerous fjords
and inlets. Some of them, especially in the north, are influenced by tidewater glaciers. The typical depth along the
coast of Novaya Zemlya does not exceed 250 m. The
western coast in particular is influenced predominantly by
Arctic waters, although Atlantic ones also have some effect
in the extreme north of the region (Ozhigin et al. 2011).
Average water temperature at sea bottom, measured in
September, the warmest month, is about 0–1 °C (Loeng
1991; Ozhigin et al. 2011). The temperature range observed
during our recent investigation was from -1.83° to
?2.25 °C (Online Resource 1). The salinity of the coastal
waters was remarkably stable at about 34.8–34.9 psu with
nonsignificant variation (Ozhigin et al. 2011). At least two
regions of Novaya Zemlya (Matochkin Shar and Chernaya
Inlet) were used as nuclear weapon test sites during the
second part of the 20th century. However, dedicated study
revealed no significant effects of radioactive pollution on
macrobenthic communities (Pogrebov et al. 1997).
To describe the geographical distribution of gastropods
along Novaya Zemlya, we divided its coast into the following seven regions (Fig. 1): A—extreme north of the
archipelago southward to 75°500 N; B—coastal areas from
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Fig. 1 Map of study area, showing sample sites of material used
(black circles). Letters A–G indicate regions of previous and modern
investigations: A—extreme north of the archipelago southward to
75°500 N; B—coastal areas from 75°500 south to 74°200 N; C—central
part of the archipelago from 74°200 N to 71°500 N except Moller Bay
and Besimyannaya Bay; D—Moller Bay and Besimyannaya Bay; E—
Kostin Shar Strait and its vicinities; F—Chernaya Inlet; G—extreme
south of Novaya Zemlya southward to 70°500 N including Yugorskiy
Shar Strait
75°500 south to 74°200 N; C—central part of the archipelago
from 74°200 N to 71°500 N except Moller Bay (pakbd Mokkepa) and Besimyannaya Bay (<epsvzyyaz uy,a); D—
Moller Bay and Besimyannaya Bay; E—Kostin Shar Strait
(Rocnby iap) and its vicinities; F—Chernaya Inlet (X/
pyaz uy,a); G—extreme south of Novaya Zemlya southward to 70°500 N including Yugorskiy Shar Strait
(>uopcrbq iap). This division corresponds with sampling
areas of various surveys, both past and recent.
Matochkin Shar Strait was not included as a separate
research area in our study. A large amount of published and
original data on the distribution of seafloor fauna in the
strait was already summarized by Ushakov (1931). No
additional information has been reported since then, and
our recent samples also did not cover this area.
Sampling and sample processing
Material used for the present study had been collected in
the numerous cruises of RV Dalnie Zelentsy carried out
during the period of 1985–2008. Surveys covered the areas
along the western coast of Novaya Zemlya (A, B, C), inner
parts of Chernaya Inlet and Yugorskiy Shar Strait (F and G;
Online Resource 1), and some adjacent areas of the Barents
Sea (Fig. 1). Moller Bay and Kostin Shar (regions D and E
on Fig. 1) were not examined.
Material was taken from 4 to 14 sampling stations
within each region. Typically, seafloor sediments were
collected using grab samplers (0.1 m2 van Veen or 0.25 m2
Petersen) in three replicates, except for four stations in
Chernaya Inlet, where five replicates were taken with
0.1 m2 van Veen grab. Five additional samples were taken
with Sigsbee Trawl from regions B and C (Online
Resource 1).
Collected bottom sediments were sifted through 1-mmmesh sieve onboard and immediately fixed with 4 % formalin. After sorting in the laboratory under stereomicroscope, the material was transferred into 75 % ethanol.
Currently, samples are kept in the Laboratory of Macroecology and Biogeography of Invertebrates, Saint Petersburg State University (Saint Petersburg, Russia).
We found that the majority of the gastropods were live,
but occasional empty shells were also present. We did not
discard these data, and whenever information on empty
shells is used, it is specifically pointed out. Specimens in
such a poor state that they could not be identified to species
level were not considered.
For species identification, we examined shell characteristics and compared our data with modern taxonomical
literature on Northern Atlantic and Arctic gastropods
(Galkin 1955; Bouchet and Warén 1980, 1986, 1993;
Golikov 1980, 1987; Golikov and Sirenko 1988; Bogdanov
1990; Warén 1996; Chaban 1996; Kantor and Sysoev 2006;
Ohnhesiser and Malaquias 2013). Prior to this study,
original species descriptions and reference museum collections, including type series, stored in several European
museums [Swedish Museum of Natural History (Stockholm, Sweden), Natural History Museum of Denmark
(Copenhagen, Denmark), Zoological Institute of Russian
Academy of Sciences (Saint Petersburg, Russia), University Museum at the University of Oslo (Norway), and
University Museum of Bergen (Norway)] were examined
by the first author.
The taxonomical system generally follows that of
MolluscaBase (http://www.molluscabase.org) with some
remarks discussed in Nekhaev (2014). We did not apply a
definitive species identification for Anatoma Woodward,
1859, because this genus, which currently contains a single
arctic species, requires taxonomical revision (Høisæter and
Geiger 2011). In classification of cephalaspid genera
Cylichna and Retusa, we follow Chaban (2004) and Kantor
and Sysoev (2006).
All results obtained from these recent surveys were
combined to produce a single dataset of new information
on presence/absence of various species across the study
area.
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Analysis of published sources
Results
We created a combined dataset of presence/absence of all
species recorded by seven publications, each of them a
faunistic report from a single survey. Four expeditions
covered large areas (Leche 1878; Ivanov 1910; Dautzenberg and Fischer 1911; Grieg 1924), while the rest examined smaller locations: Yugorskiy Shar Strait (Aurivillius
1885), Chernaya Inlet (Gurjanova and Ushakov 1928), and
Moller Bay (Ushakov 1927). Only studies containing species lists with detailed information on material and collection sites were included.
To resolve synonymy, we predominantly used two
online taxonomical databases: MolluscaBase (http://www.
molluscabase.org) and CLEMAM (http://www.somali.
asso.fr). However, not all of the species names used by
previous authors are present in these databases. Moreover,
names are sometimes erroneously used in a single publication. In these cases, we adopted detailed lists of synonyms from the nomenclatural catalogs and taxonomical
revisions (Galkin 1955; Bogdanov 1990; Kantor and
Sysoev 2006). Associations of all names used by previous
authors with the valid species synonyms are given in
Online Resource 2.
On several occasions, species listed in the published
sources were not included in our analysis. First, since the
taxonomical status of some species of Mangeliidae reported by previous authors is often confused, we only included
species reported for this region of the Barents Sea by
Bogdanov (1990). We also omitted several species from
other families, if their occurrence in the study area was in
doubt (see ‘‘Results’’ section for further information).
Examination of recent material
Data analysis
Multidimensional scaling (MDS) together with clustering
were applied for comparison between known gastropod
fauna of Novaya Zemlya with other local gastropod faunas
of the Barents and Norwegian Seas. The unweighted pairgroup average algorithm was used for dendrogram
formation.
To estimate significance of differences between modern
and published data and between regions, analysis of similarities (ANOSIM) was carried out. ANOSIM, MDS, and
clustering were based on the Dice similarity index, which
corresponds to the Bray-Curtis index widely used for binary data in ecological studies. Expected species number for
modern data was calculated by using nonparametric
Chao 2 and Jacknife 1 estimators (Colwell and Coddington
1994). Only data on grab samples were used for such
estimation. All statistical analyses were carried out using
the PAST 2 statistical package (Hammer et al. 2001).
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We identified a total of 58 species of shell-bearing gastropod (Table 1), with 3 species (Trichotropis bicarinata,
Colus gracilis, and Retusa pertenuis) represented exclusively by empty shells. All recorded species but one are
already known from the Barents Sea and adjacent Arctic
waters (Sirenko 2001; Kantor and Sysoev 2006; Nekhaev
2014). The exception was a single specimen of Admete
Kröyer in Møller, 1842, which we provisionally identified
as Admete cf. solida. This individual had 26-mm-high shell
and corresponded well to the description and published
figure of the holotype of A. solida (Aurivillius 1885)
(Sysoev and Kantor 2002). However, in the Russian Arctic,
it has been reported only from the northern part of the
Bering Sea (Kantor and Sysoev 2006), a location significantly removed from our study area. It is also possible that
some undescribed species with similar shell morphology
may be present in the study area (Sirenko 2015).
The highest numbers of species were detected in
regions C and G (37 and 28, respectively) (Table 1). Both
areas are located in the south of Novaya Zemlya (Fig. 1).
The number of species in the northern regions A and B was
two times smaller: 13 and 15 species. Surprisingly, only 17
species were recorded in the Chernaya Inlet, although it is
located in the south of the archipelago. The most common
species were Cylichna alba (39 %), Frigidoalvania cruenta (28 %), and Solariella obscura (15 %).
The suggested number of species estimated by Chao 2
estimator was 77.2 with variance of 3.5, while the Jacknife
1 estimator value was lower at 67.5. These data as well as
the species accumulation curve (Fig. 2) show that the real
species diversity of the region was underestimated during
recent studies.
Analysis of published sources
A total of 62 species were detected during previous
investigations along the western coast of Novaya Zemlya
(Table 1). Nine more species [Onchidiopsis glacialis (Sars,
1851), Tachyrhynchus erosus (Couthouy, 1838), Neptunea
despecta (Linnaeus, 1758), Buccinum hydrophanum Hancock, 1846, Curtitoma novajasemljensis (Leche, 1878),
Frigidoalvania cruenta (Odhner, 1915), Eumetula arctica
(Mörch, 1857), Aartsenia candida (Møller, 1842), and
Cylichna densistriata (Leche, 1878)] were reported
exclusively from Matochkin Shar Strait (Ushakov, 1931)
and were not included in the total species count. Also, a
number of accounts were not added to the species list due
to their questionable reliability. We excluded three species
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Table 1 Species composition of shell-bearing Gastropoda along the western coast of Novaya Zemlya
Region
A
B
C
D
E
F
G
L
L
L, 1
6
1
L, 7
L
1
1, 4
7
1, 3
6
6
Patellogastropoda
Acmaeoidea Forbes, 1850
Lepetidae Gray, 1850
Lepeta caeca (Müller, 1776)
Lottiidae Gray, 1840
Tectura virginea (Müller, 1776)
Erginus rubellus (Fabricius, 1780)
Testudinalia testudinalis (Müller, 1776)
L
5
5
1
Vetigastropoda
Scissurelloidea Gray, 1817
Anatomidae McLean, 1989
Anatoma sp.
L
Fissurelloidea Fleming, 1822
Fissurellidae Fleming, 1822
Puncturella noachina (Linnaeus, 1771)
6
1
7
L, 1, 3
1
1, 4
L, 7
L
6
1, 4
7
Trochoidea Rafinesque, 1815
Margaritidae Thiele, 1924
Margarites costalis (Lovén in Gould, 1841)
L
5
Margarites groenlandicus (Gmelin, 1791)
Margarites helicinus (Phipps, 1774)
Margarites olivaceus (Brown, 1827)
5
Margarites striatus (Leach, 1819)
Solariellidae Powell, 1951
1, 3
1, 6
1, 4
7
L
6, 5
4
7
1, 6, 5
1
7
5
Solariella obscura (Couthouy, 1838)
5
Solariella varicosa (Mighels et Adams, 1842)
L, 1
1, 6, 5
L, 1
1
L
1
L, 2
L, 1
L, 1
Turbinoidea Rafinesque, 1815
Colloniidae Cossmann, 1917
Moelleria costulata (Møller, 1842)
L
L, 7
L
7
1
Caenogastropoda
Capuloidea Fleming, 1822
Capulidae Fleming, 1822
Trichotropis bicarinata (Sowerby, 1825)
S
Trichotropis borealis (Broderip et Sowerby, 1829)
L
Trichotropis conica Møller, 1842
L, 1
1, 6
1
L
Neoiphinoe kroeyeri (Philippi, 1849)
1
Littorinoidea Children, 1834
Littorinidae Children, 1834
Littorina obtusata (Linnaeus, 1758)
Littorina saxatilis (Olivi, 1792)
1
6
7
Naticoidea Guilding, 1834
Naticidae Guilding, 1834
Amauropsis islandica (Gmelin, 1791)
Cryptonatica affinis (Gmelin, 1791)
6
S, 5
L, 5
L, 1, 3
1, 6, 5
L, 2
1, 4, 5
7
Pseudopolinices nanus (Møller, 1842)
Euspira pallida (Broderip et Sowerby, 1829)
Euspira tenuistriata (Dautzenberg and Fischer 1911)
L, 1, 2
L
L, 1, 3
1, 6, 5
1
7
L, 1
4
Rissoidea Gray, 1847
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Table 1 continued
Region
A
B
C
D
E
F
G
Rissoidae Gray, 1847
Alvania scrobiculata (Møller, 1842)
L
Boreocingula castanea (Møller, 1842)
Frigidoalvania cruenta (Odhner, 1915)
Frigidoalvania janmayeni (Friele, 1878)
L
L
L
L
6
L
L
L
L
L
L
Pusillina tumidula (G.O. Sars, 1878)
L
L
Onoba improcera Warén, 1996
Velutinoidea Gray, 1840
Velutinidae Gray, 1840
Limneria undata (T. Brown, 1839)
5
Piliscus commodus (Middendorff, 1851)
6
7
1
L
Onchidiopsis groenlandica Bergh, 1853
1
Velutina velutina (Müller, 1776)
5
7
Muricoidea Rafinesque, 1815
Muricidae Rafinesque, 1815
Boreotrophon clathratus (Linnaeus, 1767)
S, 3
Boreotrophon truncatus (Strøm, 1768)
L, 1
5
7
Buccinoidea Rafinesque, 1815
Buccinidae Rafinesque, 1815
Buccinum polare Gray, 1839
Buccinum angulosum Gray, 1839
1
S
L
Buccinum ciliatum (Fabricius, 1780)
Buccinum cyaneum Bruguière, 1792
1
4
7
1
1
7
1, 2
6, 5
4
7
1, 2
1
L
1, 2
7
1, 2
1
5
5
3
5
5
L, 1
1
Buccinum maltzani Pfeffer, 1886
1
1
Buccinum scalariforme Møller, 1842
L, 1
Buccinum fragile Verkruzen, 1878
Buccinum glaciale (Linnaeus, 1761)
L
Buccinum undatum (Linnaeus, 1758)
Colus gracilis (da Costa, 1778)
Colus holboelli (Møller, 1842)
4
S
L
L
Colus islandicus (Mohr, 1876)
1
7
Colus sabini (Gray, 1824)
Neptunea communis (Middendorff, 1848)
1
6
7
S
L
Neptunea ventricosa (Gmelin, 1791)
4
Pyrulofusus deformis (Reeve, 1847)
Retifusus roseus (Dall, 1877)
1
L
Columbellidae Swainson, 1840
Astyris rosacea (Gould, 1840)
1
6, 5
4
7
L
L, 1
6
1
L, 7
L
L
1, 6
1, 4
7
2
Cancellariidae Forbes et Hanley, 1851
Admete cf. solida
Admete viridula (Fabricius, 1780)
L
L, 1
Conoidea Fleming, 1822
Mangeliidae P. Fischer, 1883
Curtitoma trevelliana (Turton, 1834)
Curtitoma violacea (Mighels et Adams, 1842)
1
Obesotoma simplex (Middendorff, 1849)
Obesotoma woodiana (Møller, 1842)
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L, 1, 2
L
L
L, 1
1
1
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Table 1 continued
Region
A
B
Oenopota harpa (Dall, 1884)
Oenopota impressa (Beck in Mwrch, 1869)
C
D
E
L, 1
1, 6
4
L, 1
1
1, 4
1
5
Oenopota pyramidalis (Strøm, 1788)
Propebela arctica (Adams, 1855)
Propebela exarata (Møller, 1842)
1
Propebela harpularia (Couthouy, 1838)
1
1
6
Propebela nobilis (Møller, 1842)
5
L
Propebela rugulata (Møller in Reeve, 1846)
L
L
Propebela scalaris (Møller, 1842)
1, 6, 5
G
7
L, 1
L
L, 7
1
7
2
1
7
4
1, 2
7
1
Propebela spitzbergensis (Friele, 1886)
F
1
6
7
Heterobranchia
Mathildoidea Dall, 1889
Mathildidae Dall, 1889
Turritellopsis stimpsoni Dall, 1919
6
L
Pyramidelloidea Gray, 1840
Pyramidellidae Gray, 1840
Menestho truncatula Odhner, 1915
L
L
Liostomia eburnea (Stimpson, 1851)
L, 1
1
7
6
7
L
Cephalaspidea
Diaphanoidea Odhner, 1914
Diaphanidae Odhner, 1914
Diaphana glacialis Odhner, 1907
L
Diaphana hiemalis (Couthouy, 1839)
L
L
Diaphana minuta Brown, 1827
6
Toledonia limnaeoides (Odhner, 1913)
7
L
L
L
Philinoidea Gray, 1850
Cylichnidae H. et A. Adams, 1854
Cylichna alba (Brown, 1827)
L
Cylichna occulta (Mighels et Adams, 1842)
L
Cylichna scalpta (Reeve, 1855)
L
L
L, 1
1, 6
1, 4
1
1
1
L
L, 7
L, 1, 2
L, 1
6
L, 7
L
6
L, 7
L
6
L
L
17
28
Philinidae Gray, 1850
Laona finmarchica (M. Sars, 1859)
L
Philine lima (Brown, 1827)
Philine quadrata (S. Wood, 1839)
L
L
Retusidae Thiele, 1925
Retusa pertenuis (Mighels, 1843)
S, 1
Retusa turrita Møller, 1842
Found in original material
13
15
37
Recorded in literature
5
12
30
43
30
30
13
Total
17
26
51
43
30
46
45
‘‘L’’ indicates presence of living specimens in the material seen, ‘‘S’’ indicates that only empty shells were found in our material; numbers
indicate records in literature sources: 1, Leche 1878; 2, Aurivillius 1885; 3, Ivanov 1910; 4, Dautzenberg and Fischer 1911; 5, Grieg 1924; 6,
Ushakov 1927; 7, Gurjanova and Ushakov 1928. Letters A–G indicate regions of previous and modern investigations: A, extreme north of the
archipelago southward to 75°500 N; B, coastal areas from 75°500 south to 74°200 N; C, central part of the archipelago from 74°200 N to 71°500 N
except Moller Bay and Besimyannaya Bay; D, Moller Bay and Besimyannaya Bay; E, Kostin Shar Strait and its vicinities; F, Chernaya Inlet; G,
extreme south of Novaya Zemlya southward to 70°500 N including Yugorskiy Shar Strait
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Analysis of species lists from recent and previous
surveys
Fig. 2 Species accumulation curve for grab samples
[Onoba aculeus (Gould, 1841), Peringia ulvae (Pennant,
1777), and Rissoella globularis (Jeffreys in Forbes et
Hanley, 1852] recorded by Ushakov (1927) and Gurjanova
and Ushakov (1928) because their known distribution in
the Barents Sea is limited to its southwestern part (Nekhaev
2014; Nekhaev et al. 2014), which is strongly influenced
by the warm North Atlantic Current. Their presence along
Novaya Zemlya requires confirmation. We reexamined all
available samples of Onoba aculeus, Peringia ulvae, and
Rissoella globularis taken by Gurjanova and Ushakov
during various expeditions and stored in the collections of
the Zoological Institute of RAS (Saint Petersburg), but
none originated from coastal waters of Novaya Zemlya.
Five mangelid species reported by previous authors [Curtitoma decussata (Couthouy, 1839), Obesotoma gigantea
(Mörch, 1869), Oenopota elegans (Møller, 1842), Propebela cancellata (Mighels et Adams, 1842), and P. angulosa (G.O. Sars, 1878)] but not listed for this part of the
Arctic by Bogdanov (1990) were also excluded.
The highest number of species (46) was registered by
Leche (1878) along the southern part of Novaya Zemlya.
Ushakov (1927) and Gurjanova and Ushakov (1928) presented extensive species lists containing 36 gastropod
species for Chernaya Inlet and 31 for Moller Bay. Species
lists given by other authors contained from 6 to 18
species.
Four species (Testudinalia testudinalis, Cryptonatica
affinis, Buccinum cyaneum, and B. glaciale) were recorded
in all geographical regions (Table 1). There was a moderate degree of repeatability of the species records across
seven previous surveys: B. cyaneum and C. affinis were
found in all studies, while 20 species were only mentioned
once. The highest species richness was also observed in the
southern part of the study area (region D—43 species,
regions E and G—30 species).
123
The combined total of species of shell-bearing gastropod,
identified in both our material and published sources,
reached 86. Only 35 species are shared between both
datasets (Dice similarity distance = 0.56). By contrast, 24
species were found exclusively in the recent material and
28 species were reported from previous surveys alone.
Analysis of similarities shows clear separation between
species lists obtained during our study and compiled from
previous investigations in all regions (ANOSIM, R = 0.57,
p = 0.02). By contrast, geographical regions along the
coast of Novaya Zemlya did not differ based on both
datasets (ANOSIM, R = -0.2, p = 0.82). Modern and
historical data demonstrated unequal proportions of species
numbers in different superfamilies (Fig. 3). While whelks
and top snails were frequently reported by previous
authors, representatives of the small-sized superfamilies,
e.g., Rissoidea and Philinoidea, were more common in the
recent material.
Discussion
Comparison of modern and historical data
We present herein the first assessment of the species
diversity along Novaya Zemlya based on both newly collected and previously published data. These datasets
overlap only partially and both provide incomplete faunistic information (as the theoretical estimates of species
richness attest). We believe that differences between the
two datasets do not indicate changes in species composition of the region during the last century, since neither
recently obtained material nor published sources alone can
fully describe the diversity of gastropods along the coast of
Novaya Zemlya. Previous studies, as well as our work, are
Fig. 3 Proportions of species number in main superfamilies of shellbearing Gastropoda found only during the recent investigations (1),
only known by historical data (3), and shared by both datasets (2)
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Polar Biol
based on a limited number of benthic sampling stations.
None of the surveys aimed specifically at detection of
taxonomical diversity along the coast of Novaya Zemlya.
Moreover, some data used here were obtained accidentally
during investigations of adjacent areas. For instance, Nordenskiöld’s expeditions (Leche 1878; Aurivillius 1885)
primarily explored Siberian seas. Similarly, the majority of
the recently collected material identified during this study
was taken incidentally in the course of ecological surveys
of neighboring regions of the Barents Sea. There were only
three dedicated expeditions examining the species composition of benthic invertebrates, including gastropods, and
each was focused on a relatively small area: Moller Bay,
Matochkin Shar Strait, and Chernaya Inlet (Ushakov 1927;
Gurjanova and Ushakov 1928; Ushakov 1931). Predictably, the highest species diversity was recorded in these
areas (Table 1). Nonetheless, we identified nine more
species from Chernaya Inlet found during the 2000 survey
and not mentioned by Gurjanova and Ushakov (1928).
Overall, the fauna of the northern part of the archipelago
was poorly sampled and still remains one of the least
understood areas of the Barents Sea.
Researchers from the late 19th to early 20th century
used trawls or dredges to sample bottom fauna. Samples
were usually sorted immediately after collection, so many
small mollusks were often overlooked, while larger animals were overrepresented. Conversely, during the last
decades, samples were collected almost exclusively by
grabs. This method underestimates the diversity of large
mollusks (e.g. Buccinidae, Mangeliidae, and Velutinidae),
although it allows accurate assessment of small animals. A
similar effect of sampling gear on the outcome of benthic
studies was reported from surveys of the Gorlo Strait,
White Sea (Solyanko et al. 2010). Another important reason for the differences between recent and previously
published data lies in unequal coverage of coastal versus
offshore areas during investigations. Most of the historical
material comes from near-shore sites: Chernaya Inlet,
Moller Bay, and numerous smaller bays and inlets. By
contrast, the bulk of the recent samples were collected
much farther away from the coastline (usually 3–20 km).
Since in the recent surveys there were no samples from
shallow waters, we were unable to find several typical
coastal species, such as Testudinalia testudinalis, Margarites helicinus, and Littorina spp.
Relationships with adjacent faunas
The fauna of the Barents Sea is heterogeneous and consists of
two main zoogeographical groups: boreal and Arctic. The
boreal communities generally occur in the southwestern part
of the sea, which is influenced by temperate Atlantic water
masses. Arctic fauna, associated with cold waters, is common in the north and east of the Barents Sea (Golikov 1989;
Jirkov 2013; Jørgensen et al. 2014). Some areas, e.g., the
Murman Coast and Svalbard, are considered to have mixed
fauna. Altogether, up to eight biogeographical units of the
Barents Sea have been defined, based on the local distribution of benthic species (e.g., Jirkov 2013).
The number of species of shell-bearing gastropod in the
Barents Sea is no less than 200 (Sirenko 2001; Kantor and
Sysoev 2006; Nekhaev 2014), therefore the 86 species
known from Novaya Zemlya amount to about 40 % of the
gastropod fauna in the whole sea. Similar species-level
diversity was reported in the entirely of the White Sea and
in the waters surrounding Franz Josef Land: 86 and 83
species, respectively (Sirenko 2001; Golikov and Scarlato
1977; Nekhaev 2008).
Figure 4 compares the species composition of shellbearing gastropods from Novaya Zemlya presented here
with those for Franz Josef Land (Golikov and Scarlato
1977; Nekhaev 2008), Svalbard (Palerud et al. 2004),
Northern Norway (Høisæter 2009), the Murman Coast
(Nekhaev 2014), and the White Sea (Sirenko 2001). The
Fig. 4 Dendrogram (a) and
MDS plot (b) comparing
taxonomical composition of
shell-bearing Gastropoda of
Franz Josef Land (FJL),
Murman Coast (Mu), Novaya
Zemlya (NZ), Northern Norway
(NN), Svalbard (Sv), and White
Sea (WS)
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fauna of Northern Norway was the richest, and most distant
from all others. Similarity distances between coastal
regions of the Barents Sea, including Novaya Zemlya, were
similar and formed no defined grouping within this cluster.
Our findings support the view that these regions are
occupied by somewhat differing faunistic complexes (Jirkov 2013; Jørgensen et al. 2014). Surprisingly, here we
found no differences between sites from boreal (Northern
Norway, Murman, Svalbard) and Arctic (Novaya Zemlya,
Franz Josef Land, and the White Sea) biogeographical
regions, although in previous studies these two regions
were distinguished by occurrence of species with different
types of distribution (Filatova 1957; Jirkov 2013). This
analysis represents only a preliminary comparison between
the most studied local gastropod faunas of the Barents Sea
and the adjacent areas. More detailed investigations are
still needed.
Acknowledgements We are grateful to Ann-Helen Rønning (Oslo,
Norway) for her valuable advice on early publications and help with
searching for literature and to Yuta Tamberg (Saint Petersburg,
Russia) for language editing. We also thank the staff of the Laboratory of Zoobenthos of Murmansk Marine Biological Institute (Murmansk, Russia) for kind assistance during our work with recent
material.
Funding The work was supported by the Russian Foundation for
Basic Research (grant no. 16-34-00372 mol_a).
References
Aurivillius CWS (1885) Öfversigt öfver de af Vega-expeditionen
insamlade arktiska hafsmollusker. II. Placophora och Gastropoda. In: Vega-Expeditionens Velenskapliga Jakffagelser. F
& G Beijers Förlag, Stokholm, pp. 313–382
Beuchel F, Gulliksen B (2008) Temporal patterns of benthic
community development in an Arctic fjord (Kongsfjorden,
Svalbard): results of a 24 year manipulation study. Polar Biol
31:913–924. doi:10.1007/s00300-008-0429-9
Bogdanov IP (1990) Molluscs of the subfamily Oenopotinae (Gastropoda, Pectinibranchia, Turridae). Nauka, Leningrad (in
Russian)
Bouchet P, Warén A (1980) Revision of the North-East Atlantic
bathyal and abyssal Turridae (Mollusca, Gastropoda). J Mollus
Stud Suppl 8:1–119
Bouchet P, Warén A (1986) Revision of the North-East Atlantic
bathyal and abyssal Neogastropoda excluding Turridae (Mollusca, Gastropoda). Boll Malacol Suppl 1:123–296
Bouchet P, Warén A (1993) Revision of the Northeast Atlantic
bathyal and abyssal Mesogastropoda. Boll Malacol Suppl
3:579–840
Britaev TA, Rzhavsky AV, Pavlova LV, Dvoretsij AG (2010) Studies
on impact of Red King Crab (Paralithodes camtschaticus) on the
shallow-water benthic communities of the Barents Sea. J Appl
Ichthyol 26:66–73. doi:10.1111/j.1439-0426.2010.01494.x
Chaban EM (1996) Opisthobranch molluscs of the family Diaphanidae (Gastropoda: Opisthobranchia) of the Russian Seas.
Ruthenica 6:127–148 (In Russian)
123
Chaban EM (2004) The cephalaspidean mollusks (Mollusca, Opisthobranchia) of the Laptev Sea. Explor Fauna Seas 54:71–87 (In
Russian)
Colwell RK, Coddington AJ (1994) Estimating terrestrial biodiversity
through extrapolation. Philos Trans R Soc B 345:101–118
Dautzenberg P, Fischer H (1911) Mollusques et Brachiopodes
recueillis en 1908 par la Mission Bernard dans les mers du
Nord (Nouvelle-Zemble, mer de Barentz, mer Blanche ocean
Glacial, Norvege, mer du Nord). J de Cocnhyliologie 59:1–51
Denisenko SG, Luppova EN, Denisenko NV, Alekseev VA, Kasabov
AY, Frolova EA (1995) Quantitative distribution of benthos and
bottom communities structure on the shelf fauna on Novaya
Zemlya in the Barents Sea. In: Matishov GG (ed) Environment
and ecosystems of Novaya Zemlya (the archipelago and shelf).
KSC RAS Publishing, Apatity, pp 66–79 (In Russian)
Derjugin KM, Gurjanova EF (1926) New species of the molluscs
from the Russian Northern Seas. Proc Len Soc Nat 56:17–26 (In
Russian)
Filatova ZA (1957) Zoogeographical division of northern seas (by
distribution of Bivalvia). Trans Inst Oceanol 23:195–215 (in
Russian)
Galkin YuI (1955) Gastropod molluscs trochids of Far Eastern and
northern Seas of the USSR. Academy of Sciences of the USSR
publ, Moscow-Leningrad (In Russian)
Golikov AN (1968) Distribution and variability of long-lived benthic
animals as indicators of currents and hydrological conditions.
Sarsia 34:199–208. doi:10.1080/00364827.1968.10413382
Golikov AN (1980) Molluscs Buccininae of the World Ocean. Nauka,
Leningrad (in Russian)
Golikov AN (1987) Classis Gastropoda (part). In: Starobogatov Y,
Naumov AD (eds) Molluscs of the White Sea. Nauka, Leningrad, pp 41–149 (In Russian)
Golikov AN (1989) Arctic ocean Gastropod Prosobranchs. In:
Herman Y (ed) The arctic seas: climatology, oceanography,
geology and biology. Springer, New York, pp 325–340
Golikov AN, Averintcev VG (1977) Peculiarities of some bottom
ecosystems in the oil polluted part of the Kola Bay and near
Zhelaniye Cape (Novaya Zemlya). Biol Morya-Vlad 2:63–73 (In
Russian)
Golikov AN, Scarlato OA (1977) Composition, distribution and
ecology of gastropods and bivalves off Franz Joseph Land. In:
Scarlato OA (ed) Biocenoses of the Franz Josef Land and the
fauna of adjacent waters. Nauka, Leningrad, pp 313–390 (In
Russian)
Golikov AN, Sirenko BI (1988) The naticid gastropods in the boreal
waters of the western Pacific and Arctic Oceans. Malacol Rev
21:1–41
Gorbunov G (1949) Bottom life of the Novosiberian shoalwaters and
the central part of the Arctic Ocean. In: Gorbunov GP, Ushakov
PV (eds) Proceedings of the drifting expedition of Glavsevmorput on ice-breaker ‘‘G. Sedov’’. Biology Glavsevmorput Publ,
Moskva-Leningrad, pp 30–136 (In Russian)
Grieg JA (1924) Molluscs, Brachiopods and echinoderms from
Novaya Zemlya. Report of the scientific results of the Norwegian
expedition to Novaya Zemlya 1921. Videnskapsselskapet I
Kristiania, Kristiania
Gurjanova EF, Ushakov PV (1928) On the fauna of the Chernaya
Inlet on Novaya Zemlya. Explor Seas USSR 6:3–72
Hammer Ø, Harper DAT, Ryan PD (2001) Past: paleontological
statistics software package for education and data analysis.
Palaeontol Electron 41:1–9
Høisæter T (2009) Distribution of marine, benthic shell bearing
gastropods along the Norwegian coast. Fauna Norvegica
28:5–106
Author's personal copy
Polar Biol
Høisæter T, Geiger D (2011) Species of Anatoma (Gastropoda:
Anatomidae) in Norwegian and adjacent waters, with description
of two new species. Nautilus 125:89–112
Hop H, Pearson T, Hegseth EN, Kovacs KM, Wiencke C,
Kwasniewski S, Eiane K, Mehlum F, Gulliksen B, WlodarskaKowalczuk M, Lydersen C (2002) The marine ecosystem of
Kongsfjorden, Svalbard. Polar Res 21:167–208
Ivanov SS (1910) Preliminary report of zoological work during the
cruise of ‘‘Dmitry Solunskiy’’ in the summer 1910. Mat Investig
Nov Zemlya 2:95–110 (In Russian)
Jirkov IA (2013) Biogeography of the Barents Sea benthos. Invertebr
Zool 10:69–88
Jørgensen LL, Ljubin P, Skjoldal HR, Ingvaldsen RB, Anisimova N,
Manushin I (2014) Distribution of benthic megafauna in the
Barents Sea: baseline for an ecosystem approach to management.
ICES J Mar Sci 72:595–613. doi:10.1093/icesjms/fsu106
Jørgensen LL, Planque B, Thangstad TH, Certain G (2015) Vulnerability of megabenthic species to trawling in the Barents Sea.
ICES J Mar Sci 73:i84–i97. doi:10.1093/icesjms/fsv107
Kantor YuI, Sysoev AV (2006) Marine and brackish water Gastropoda of Russia and adjacent countries: an illustrated catalogue. KMK Scientific, Moscow
Leche W (1878) Ofversight ofver de af svenska expeditionerna till
Novaja Semlja och Jenissej 1875 och 1876 issamlade HafsMollusker. Handlingar Svenska Vetenskaps-Akademiens
16:1–86
Loeng H (1991) Features of the physical oceanographic conditions of
the Barents Sea. Polar Res 10:5–18
Lyubina OS, Bryazgin VF, Raznovskaya SV (2014) The composition
and distribution of benthic amphipods (Crustacea: Amphipoda)
in the Southern Barents Sea. Russ J Mar Biol 40:241–254
Middendorff AT (1849) Beiträge zu einer Malacozoologia Rossica. II.
Aufzählund und Beschreinbung zur Meeresfauna Russlands
gehörigen Einscharler. Mémoires de l’Académie Impériale des
Sciences, ser. 6. Classe Mathématique, Physique et Naturelles
8:329–516
Nekhaev IO (2008) Species composition and the structure of
gastropod mollusk fauna around Franz Joseph Land. Nat Shelf
Archipel Eur Arct 8:263–269 (In Russian)
Nekhaev IO (2014) Marine shell-bearing Gastropoda of Murman
(Barents Sea): an annotated check-list. Ruthenica 24:75–121
Nekhaev IO (2016) Newly arrived or previously overlooked: is there
evidence for climate-driven changes in the distribution of
molluscs in the Barents Sea? Biodivers Conserv 25:807–825.
doi:10.1007/s10531-016-1104-z
Nekhaev IO, Deart YuV, Lubin PA (2014) Molluscs of the genus
Onoba (H. et A. Adams 1852) from the Barents Sea and adjacent
waters (Gastropoda: Rissoidae). Proceedings of the Zoological
Institute of RAS 318:268–279
Ohnhesiser LT, Malaquias M (2013) Systematic revision of the
gastropod family Philinidae (Mollusca: Cephalaspidea) in the
north-east Atlantic Ocean with emphasis on the Scandinavian
Peninsula. Zool J Linn Soc 167:273–326. doi:10.1111/zoj.12000
Ozhigin VK, Ingvaldsen RB, Loeng H, Boitsov VD, Karsakov AL
(2011) Introduction to the Barents Sea. In: Jakobsen T, Ozhigin
VK (eds) The Barents Sea Ecosystem, resources, management
Half a century of Russian-Norwegian cooperation. Tapir Academic, Trondheim, pp 39–76
Palerud R, Gulliksen B, Brattegard T, Sneli J-A, Vader W (2004) The
marine macro-organisms in Svalbard waters. In: Prestrud P,
Strøm H, Goldman HV (eds) A catalogue of the terrestrial and
marine animals of Svalbard. Norwegian Polar Institute, Tromso,
pp 5–56
Pogrebov VB, Fokin SI, Galtsova VV, Ivanov GI (1997) Benthic
communities as influenced by nuclear testing and radioactive
waste disposal off Novaya Zemlya in the Russian Arctic. Mar
Pollut Bull 35:333–339
Sirenko BI (ed) (2001) List of species of free-living invertebrates of
Eurasian Arctic Seas and adjacent deep waters. ZIN RAS publ,
Saint Petersburg
Sirenko BI (2015) Two new species of gastropods of the subfamily
Admetinae (Gastropoda: Cancellariidae) from the Sea of
Okhotsk. Biol Morya-Vlad 41:174–178 (In Russian)
Solyanko K, Spiridonov V, Naumov A (2010) Benthic fauna of the
Gorlo Strait, White Sea: a first species inventory based on data
from three different decades from the 1920s to 2000s. Mar
Biodiv 41:441–453. doi:10.1007/s12526-010-0065-9
Sysoev AV, Kantor YuI (2002) On some species of prosobranchiate
gastropods from Russian waters described by C.W.S. Aurivillius
in 1885. Ruthenica 12:113–118
Ushakov PV (1927) On zoogeography of the coastal zones of the
Moller Bay. Expl Seas USSR 4:18–80
Ushakov PV (1931) Benthic communities of the Matochkin Shar
(Novaya Zemlya). Expl Seas USSR 12:5–128
Warén A (1996) New and little known Mollusca from Iceland and
Scandinavia. Part. 3. Sarsia 81:197–245
Yesson C, Simon P, Chemshirova I, Gorham T, Turner CJ, Arboe
NH, Blicher ME, Kemp KM (2015) Community composition of
epibenthic megafauna on the West Greenland Shelf. Polar Biol
38:2085–2096. doi:10.1007/s00300-015-1768-y
Zvonareva S, Kantor Yu, Xinzheng L, Britayev T (2015) Long-term
monitoring of Gastropoda (Mollusca) fauna in planted mangroves in central Vietnam. Zool Stud 54:39. doi:10.1186/s40555015-0120-0
123
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