The Baltic Sea scale inventory of benthic faunal communities
ICES Journal of
Marine Science
ICES Journal of Marine Science (2016), 73(4), 1196– 1213. doi:10.1093/icesjms/fsv265
Original Article
The Baltic Sea scale inventory of benthic faunal communities
Mayya Gogina 1*, Henrik Nygård 2, Mats Blomqvist3, Darius Daunys4, Alf B. Josefson 5, Jonne Kotta6,
Alexey Maximov 7, Jan Warzocha 8, Vadim Yermakov 9, Ulf Gräwe1, and Michael L. Zettler 1
1
Leibniz Institute for Baltic Sea Research (IOW), Warnemünde, Seestr. 15, 18119 Rostock, Germany
Marine Research Centre, Finnish Environment Institute SYKE, PO Box 140, FI-00251 Helsinki, Finland
3
Hafok AB, 179 61 Stenhamra, Sweden
4
Coastal Research and Planning Institute, Klaipeda University, H. Manto Str. 84, LT-92294 Klaipėda, Lithuania
5
Department of Bioscience, Aarhus University, Frederiksborgvej 399, 4000 Roskilde, Denmark
6
Estonian Marine Institute, University of Tartu, Mäealuse 14, 12618 Tallinn, Estonia
7
Zoological Institute Russian Academy of Science, Universitetskaja nab. 1, 199034 St Petersburg, Russia
8
National Marine Fisheries Research Institute, ul. Kołła˛taja 1, 81-332 Gdynia, Poland
9
Latvian Institute of Aquatic Ecology, Daugavgrı̄vas Str. 8, LV-1048 Riga, Latvia
2
*Corresponding author: tel: +49 381 5197 393; fax: +49 381 5197 211; e-mail:
Gogina, M., Nygård, H., Blomqvist, M., Daunys, D., Josefson, A. B., Kotta, J., Maximov, A., Warzocha, J., Yermakov, V., Gräwe, U., and
Zettler, M. L. The Baltic Sea scale inventory of benthic faunal communities. – ICES Journal of Marine Science, 73: 1196 –1213.
Received 23 July 2015; revised 9 December 2015; accepted 14 December 2015; advance access publication 26 January 2016.
This study provides an inventory of the recent benthic macrofaunal communities in the entire Baltic Sea. The analyses of soft-bottom benthic invertebrate community data based on over 7000 locations in the Baltic Sea suggested the existence of 10 major communities based on species abundances
and 17 communities based on species biomasses, respectively. The low-saline northern Baltic, characterized by silty sediments, is dominated by
Monoporeia affinis, Marenzelleria spp., and Macoma balthica. Hydrobiidae, Pygospio elegans, and Cerastoderma glaucum dominate the community
in sandy habitats off the Estonian west coast and in the southeastern and southern Baltic Sea. Deep parts of the Gulf of Finland and central Baltic
Sea often experience hypoxia, and when oxygen levels in these regions recover, Bylgides sarsi was the first species to colonize. The southwestern
Baltic Sea, with high salinity, has higher macrofaunal diversity compared with the northern parts. To spatially interpolate the distribution of the
major communities, we used the Random Forest method. Substrate data, bathymetric maps, and modelled hydrographical fields were used as
predictors. Model predictions were in good agreement with observations, quantified by Cohen’s k of 0.90 for the abundance and 0.89 in the wet
weight-based model. Misclassifications were mainly associated with uncommon classes in regions with high spatial variability. Our analysis provides
a detailed baseline map of the distribution of benthic communities in the Baltic Sea to be used both in science and management.
Keywords: Baltic Sea, community analysis, large scale, macrozoobenthos, Random Forest, spatial distribution.
Introduction
In the recent decades, there has been a marked increase in benthic
habitat mapping, largely inspired by the development and implementation of various research and monitoring programmes,
management plans, and legislations, such as the 1992 EU Habitats
Directive. This has resulted in an exponential accumulation of
data hitherto stored as different non-standardized national datasets.
Here for the first time, we bring much of these datasets together and
harmonize them to address management issues on the large Baltic
Sea scale. Similar large-scale efforts were previously reported for
the North Sea in Rees et al. (2007), for the Barents Sea in
Anisimova et al. (2010).
The focus of our study was identification, description, and basin
scale mapping of major benthic macrofauna community distribution in the Baltic Sea. A number of works were previously published,
describing and mapping the benthic macrofauna communities in
different subbasins of the Baltic Sea (e.g. Warzocha, 1995; Laine,
2003; Glockzin and Zettler, 2008; Gogina et al., 2010). However,
# International Council for the Exploration of the Sea 2016.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/
by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
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The Baltic Sea scale inventory of benthic faunal communities
we identified a lack of joint data analyses covering the entire Baltic
Sea region based on new data collected since 2000.
At the beginning of the last century, Petersen (1913) introduced the
concept of infaunal communities for the description and quantification
of areas with similar fish food. He aggregated the infaunal species in
relation to broad geographic areas of relatively uniform sediments,
depth, and salinity. Petersen (1913) described various soft sediment
benthic macrofauna assemblages in different regions of the Baltic Sea.
The scale of the investigated area and its heterogeneity allowed for
high resolution of classification detail, and the definition of a number
of communities or assemblages. Summaries on the distribution of
fauna in the Baltic Sea can be found in works of early investigators
such as Hessle (1924), Välikangas (1933), Ekman (1933, 1935), and
Segerstråle (1957). Based on the findings outlined by Zenkevitch
(1963), Schiewer (2008) listed the most important Baltic Sea coenoses
or assemblages of species. These were (i) Abra alba-coenosis with
Corbula gibba, Arctica islandica, Lagis koreni, Nephtys spp., Diastylis
rathkei, and Ophiura albida which predominate in the western Baltic.
Towards the east (ii) Arctica-Astarte assemblages were increasingly
found. In the shallow part of the Baltic Proper (iii) Macoma baltica
coenosis dominated, whereas in the deeper part (iv) Macoma calcarea
coenosis (with Mya truncata and Astarte) occurred. Farther in the
eastern soft-bottom areas (v) Saduria-Monoporeia-Pontoporeia coenosis with euryhaline species were found, and Gotland Deep, strongly
affected by oxygen depletion, was colonized only by (vi) a species-poor
hypolimnic community. One of the deepest basins in the Baltic Sea,
the Landsort Deep, was characterized by long-term azoic, laminated
sediments (Schiewer, 2008). Elmgren (1978) gave a description of the
structure and dynamics of the Baltic Sea benthic communities from
the Bothnian Bay down to the Baltic Proper and summarized habitats
as “an almost uniquely simple benthic ecosystem”. Carman and
Cederwall (2001) provided an analysis of mean biomass data for the
main subbasins of the (...truncated)