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Of the Chukchi Sea Biogeographic Regionalization OceTHE OFFICIALa MAGAZINEn ogOF THE OCEANOGRAPHYra SOCIETYphy CITATION Denisenko, N.V., and J.M. Grebmeier. 2015. Spatial patterns of bryozoan fauna biodiversity and issues of biogeographic regionalization of the Chukchi Sea. Oceanography 28(3):134–145, http://dx.doi.org/10.5670/oceanog.2015.62. DOI http://dx.doi.org/10.5670/oceanog.2015.62 COPYRIGHT This article has been published in Oceanography, Volume 28, Number 3, a quarterly journal of The Oceanography Society. Copyright 2015 by The Oceanography Society. All rights reserved. USAGE Permission is granted to copy this article for use in teaching and research. Republication, systematic reproduction, or collective redistribution of any portion of this article by photocopy machine, reposting, or other means is permitted only with the approval of The Oceanography Society. Send all correspondence to: [email protected] or The Oceanography Society, PO Box 1931, Rockville, MD 20849-1931, USA. DOWNLOADED FROM HTTP://WWW.TOS.ORG/OCEANOGRAPHY RUSSIAN-AMERICAN LONG-TERM CENSUS OF THE ARCTIC Spatial Patterns of Bryozoan Fauna Biodiversity and Issues of Biogeographic Regionalization of the Chukchi Sea By Nina V. Denisenko and Jacqueline M. Grebmeier Bryozoan Parasmittina tatiani found in the Chukchi Sea. 134 Oceanography | Vol.28, No.3 ABSTRACT. Based on extensive data from a variety of sources, including Russian- biomass, estimating biomass for a major- American Long-term Census of the Arctic expeditions in 2009 and 2012, this paper ity of bryozoan species is difficult due to provides an update on the richness of bryozoans in the Chukchi Sea and evaluates the colonial and often encrusting nature the variation in bryozoan biodiversity along environmental gradients (e.g., depth, of the organism (N. Denisenko, 1983). bottom water temperature, and bottom sediment composition). Though bryozoans Moreover, old archived data do not con- have been studied in only about 77% of the Chukchi Sea, temperature gradients across tain information about bryozoan abun- geographical zones appear to control fauna richness. The inflow of Pacific water through dance. This fact may be one additional the Bering Strait into the Chukchi Sea strongly influences the distribution of the reason why there has been no statistical 204 registered bryozoan species, about 30% of which have a Pacific origin. The greatest estimation of bryozoan biodiversity for similarities between bryozoan fauna in the Chukchi Sea and those occurring further the Arctic. As a result, there has been no south in the Bering Sea are found in the southern and eastern parts of the Chukchi Sea. generalized report on bryozoan biodiver- The transitional zone between Arctic and Pacific high-boreal biogeographic regions sity or the relationship between faunal occurs in the western and northern areas of the Chukchi Sea. The dominance of boreal richness and environmental factors such bryozoan species over Arctic species in the southeastern and southern areas of the as depth, bottom water temperature, and Chukchi Sea identifies this area as part of a Pacific high-boreal region. The boundary bottom sediment composition. Taking between boreal and northern transitional zones corresponds to a 50:50 ratio of boreal to into account the intensity of zooben- Arctic species, with the transition extending from Cape Serdtse-Kamen’ in the western thos sampling in the Chukchi Sea (about Chukchi Sea northward to Point Franklin (68°30'N) in the eastern Chukchi Sea. 500 stations made by Russian expeditions and over a 1,000 stations by US expedi- INTRODUCTION (2008) increased the number of known tions alone), it could be assumed that the The study of Earth’s faunal biodiver- bryozoan species (see online supplemen- bryozoan records are sufficient to evalu- sity remains a pressing challenge within tary materials). Notably, most publica- ate the biodiversity of this faunal group; the field of biology. The number of pub- tions concerning bryozoans have been however, the quality of the available data lications devoted to this problem has mainly descriptive, with listings of spe- varies greatly and assumptions must be increased, although most studies are cies and their synonymy, and less focused made to utilize the different data sets. devoted to terrestrial ecosystems; only on their ecological importance. The Chukchi Sea is one of the world about 10% represent biodiversity stud- Understanding of Chukchi Sea bryo- ocean’s transitional areas, and there are ies in the marine environment (Hendriks zoan fauna has been enhanced by a syn- contrasting opinions regarding its bio- et al., 2006), and most of these have thesis using modern nomenclature of all geographic status. Some scientists con- been conducted in the temperate seas bryozoan faunal data based on both the sider the whole sea to be completely in the of the world ocean. collection of animals at the Zoological Arctic zone (Golikov, 1980; Andriyashev Biodiversity studies of fauna in Arctic Institute of the Russian Academy of and Shaposhnikova, 1985; Briggs, 1995), seas are steadily increasing, although Sciences and early data collected in the while others regard it as a transitional the number of publications on this issue framework of the Russian-American zone between the Arctic and boreo- Arctic remains limited. In recent years, a num- Long-term Census of the Arctic zones, depending on spatial location ber of reviews (e.g., Bluhm at al., 2011; (RUSALCA) program that began in 2004 (Grebmeier et al., 2006). Yet other scien- Kosobokova et al., 2011; Piepenburg (N. Denisenko, 2008; N. Denisenko and tists offer an intermediate interpretation, et al., 2011; Josefson et al., 2013) have Kuklinski, 2008). Additional collection where the southern part of the Chukchi been based on an increasing number of benthic samples during the RUSALCA Sea is a transitional zone, with the of field expeditions over the past two 2009 and 2012 expeditions further remainder belonging to the Arctic zone decades. However, the biodiversity and increased records of bryozoans and spe- (Mironov, 2013). Still others regard the spatial distribution of bryozoans, in par- cies counts beyond the 2008 publications. southern part of the Chukchi Sea as part ticular, still have not been estimated, We describe these findings here. of the Pacific boreal zone (Filatova, 1957; although this group is one of the most Magurran (2004) argued that spe- Scarlato, 1981; Petryashov at al., 2013). diverse fauna in the Arctic (Gontar and cies richness is the simplest way to esti- Based on our data compilations, first, Denisenko, 1989; Josefson et al., 2013). mate assemblage species diversity. we hypothesize that bryozoan diversity is Chukchi Sea bryozoans were one of However, estimation of species richness higher in the Chukchi Sea than previously the poorest known Arctic faunal types is quite difficult if information about recognized. Bryozoan species there are (around 140 species; Kluge, 1962; Gontar species composition is lacking, espe- distributed unequally and, in fact, form and Denisenko, 1989; Gontar, 2001) until cially for large regions. Although bio- local assemblages. The level of our under- recent compilations by N. Denisenko diversity is often estimated using quan- standing of bryozoan diversity depends (2008) and N. Denisenko and Kuklinski titative characteristics of abundance and on the intensity of bryozoan sampling Oceanography | September 2015 135 in different regions of the Chukchi Sea, were preserved in 4% formalin in sea- (the ratio between local and regional coincident with the known relationship water, buffered by hexamine. Within two diversity) was used to show similar- between the variability of environmental to three months, animals were sorted, ity or dissimilarity of species composi- factors (e.g., seawater temperature, sed- identified as to species, counted, and tion of regional faunas (Magurran, 2004). iment grain size) and the different water weighed, and subsequently archived in The data on species numbers were pro- masses in the region. Second, we hypoth- 70% ethanol. The results from these new cessed statistically using Microsoft Excel esize that bryozoans, as sessile animals, bryozoan samples were combined with and StatSoft Statistica. Station-based are suitable organisms to use to determine published data, including information rarefication curves were used to compare biogeographic zonations within the sea. from American waters in the eastern sec- species numbers from the whole Chukchi Our current analysis of spatial changes in tion of the Chukchi Sea. Both physical Sea and at different depth ranges. The the biogeographic composition of bryo- samples and electronic data are archived nonparametric Chao2 estimator was cal- zoans within the Chukchi Sea should help at the Zoological Institute of the Russian culated by means using the statistical reduce the disparity in interpretations of Academy of Sciences. PRIMER software (v.4) in order to pre- their biogeographic status in this region. In preparing the list of species, we dict the expected number of species that used the most recent information about would be observed in the Chukchi Sea MATERIALS AND METHODS species taxonomy (Dick and Ross, 1988; (Clarke and Warwick, 1994). Similarity This work is based on an analysis of Grischenko, 2002; Dick et al., 2005; of the bryozoan fauna within the Chukchi archived data and data available in the Kuklinski and Taylor, 2006; Kuklinski Sea and in comparison with nearby literature, faunal collections held at et al., 2007; N. Denisenko, 2008; Bock, areas (Bering
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