Antarctic Intertidal Communities Huw James Grifﬁths1* and Catherine Louise Waller2

Journal of Biogeography (J. Biogeogr.) (2016) ORIGINAL The first comprehensive description of ARTICLE the biodiversity and biogeography of Antarctic and Sub-Antarctic intertidal communities Huw James Griffiths1* and Catherine Louise Waller2 1British Antarctic Survey, Cambridge CB30ET, ABSTRACT UK, 2Centre for Environmental and Marine Aim To describe the distribution of biodiversity and biogeographical patterns Sciences, University of Hull, Scarborough YO11 of intertidal organisms in southern temperate and polar waters. We hypothe- 3AZ, UK sized that there would be differences in community structure between the Antarctic, which is most affected by ice, and the sub-Antarctic and other neighbouring regions. We also hypothesized that rafting and West Wind Drift will be the significant drivers of biogeographical patterns. Additionally, the size, age, isolation, volcanic or glacial history of a region and the presence of large, beach dwelling, mammals and birds would all play a role in determining the level of biodiversity observed. Location South Atlantic, Indian and Pacific Oceans and the Southern Ocean. Methods We examined all available intertidal records from the Antarctic and sub-Antarctic with additional data from neighbouring regions for comparison and context. We compiled 3902 occurrences of 1416 species of high southern latitude intertidal organisms from 229 locations and used PRIMER 6 to per- form multivariate statistical analyses. Results The Antarctic and sub-Antarctic are shown to be distinct biogeographical regions, with patterns driven by a small number of widely distributed species. These wide-ranging molluscs and macroalgae dominate the biogeographical structure of the Southern Ocean intertidal, most likely as a result of rafting in the Antarctic Circumpolar Current. East Antarctic intertidal habitats are potentially isolated by the Ross and Weddell Sea ice shelves but represent a great unknown in this biogeographical scheme. Main Conclusions The view that the Antarctic intertidal is a lifeless desert does not hold true, with Antarctic Peninsula intertidal communities being richer and more diverse than those in southern South America and the sub- *Correspondence: Huw James Griffiths, British Antarctic islands. Changing conditions in the Antarctic and sub-Antarctic Antarctic Survey, High Cross, Madingley Road, intertidal mean that a representative baseline is needed (acquired through stan- Cambridge CB30ET, UK. dardized and quantitative sampling) to assess future changes and to detect any E-mail: [email protected]. This is an open access article under the terms invasive species. of the Creative Commons Attribution License, Keywords which permits use, distribution and reproduction in any medium, provided the climate change, island biogeography, macroalgae, mollusc, rafting, richness, original work is properly cited. Southern Ocean environment is undergoing dramatic changes. Glacial INTRODUCTION retreat has the potential to expose ever-increasing areas of With 87% of marine glacier fronts currently in retreat on rocky or sedimentary intertidal habitat (Clark et al., 2015). the Antarctic Peninsula (Cook et al., 2005; Scambos et al., In cool temperate regions the intertidal zone is often seen 2014; Wouters et al., 2015) the Antarctic intertidal as the marine ‘canary in the coal mine’ with changes in ª 2016 The Authors. Journal of Biogeography Published by http://wileyonlinelibrary.com/journal/jbi 1 John Wiley & Sons Ltd doi:10.1111/jbi.12708 H. J. Griffiths and C. L. Waller species ranges due to climate change (e.g. Helmuth et al., Aims 2006; Hawkins et al., 2008) and establishment of invasive species being observed first (e.g. Griffiths et al., 1992; The aims of this study are to draw together and synthesise Robinson et al., 2005). So it comes as little surprise that all available data on the Antarctic and sub-Antarctic inter- one of the 80 key questions raised by the first Scientific tidal biota and to determine if there are any commonalities Committee on Antarctic Research (SCAR) Antarctic and between sites and regions. Southern Ocean Science Horizon Scan was: ‘How will We hypothesize that there will be differences in commu- increases in the ice-free Antarctic intertidal zone impact bio- nity structure between the high Antarctic, which is most diversity and the likelihood of biological invasions?’ (Kenni- affected by ice, and the sub-Antarctic and other neigh- cutt et al., 2014). bouring regions. We also hypothesize that rafting and The Antarctic intertidal environment is considerably less West Wind Drift will be significant drivers of biogeograph- well sampled than either the surrounding deep sea (Brandt ical patterns. Additionally, the size, age, isolation, volcanic et al., 2014) or adjacent terrestrial habitats (Chown et al., or glacial history of a region and the presence of large, 1998; Shaw et al., 2010), the opposite situation of almost beach dwelling, mammals and birds will all play a role in anywhere else in the world. From initial observations, the determining the level of biodiversity observed in the Antarctic littoral environment would appear to be virtually intertidal. lifeless, with no visible zonation, one of the most obvious and defining features of rocky shores globally. It is character- METHODS ized by intense seasonal scouring by ice, winter ice encase- ment, high UV radiation and seasonally large variations in In this study we examine all available whole community temperature and salinity. However, despite the long-held intertidal records from the Antarctic and sub-Antarctic (fol- view that the Antarctic intertidal conditions are too extreme lowing the geographical definitions used by the Scientific to support obvious signs of macrofaunal life (Powell, 1965; Committee on Antarctic Research Biogeographic Atlas). All Clark et al., 2015), little known historic and recent studies islands south of the Polar Front are considered to be Antarc- show that intertidal communities can survive these extreme tic (Bouvet Island, South Georgia, South Orkney Islands, and harsh conditions. South Sandwich Islands, South Shetland Islands) as described The sub-Antarctic intertidal environment is often charac- in Linse et al. (2006). All other islands within the influence terized by highly exposed shores on extremely remote of the Circumpolar Current are regarded as sub-Antarctic islands. The sub-Antarctic Islands are situated in the path of (Fig. 1). The database was compiled from studies of the ecol- the Antarctic Circumpolar Current and are subjected to the ogy or whole communities of localities and avoiding studies force of the southern Westerly winds (Nikula et al., 2010). that only targeted single species or taxa. We compiled data These habitats are often rocky or boulder beaches comprised for comparison from southern South America, the Falkland of bedrock and ice-formed cobble pavements (Hansom, Islands, Tristan da Cunha, South Africa, Tasmania and 1983b). Ice-free coastlines are often dominated by dense beds southern New Zealand, although data from these comparison of the kelp Durvillaea antarctica (Frazer et al., 2009; Nikula regions are not exhaustive. et al., 2010) and other large macrophytes (Knox, 1960). The restriction of data to comparable community studies Unlike shores further south, there is often a distinct pattern and avoiding data from single taxon studies inevitably means of zonation from extreme high water to low water spring that some regions with existing published intertidal biodiver- tides (Simpson, 1976). sity knowledge, such as the New Zealand sub-Antarctic The earliest published record of Antarctic intertidal life Islands, are underrepresented in the database, results and dates back to the Belgian Antarctic Expedition of 1897-99 conclusions. This is illustrated in Figs 2 and 3 where distri- which collected two species of gastropod mollusc, Nacella butions for selected taxa are supplemented with additional polaris (formerly known as N. concinna) (Hombron & Jac- data from sources not used in our analyses to illustrate that quinot, 1841) and Laevilitorina caliginosa (Gould, 1849), and taxon-specific distributional data exist beyond the scope of a single species of bivalve, Mysella subquadrata (Pelseneer, our database. 1903) from the intertidal of Two Hummock Island off the We created a database of 3902 records of high southern West Antarctic Peninsula. During the Heroic Era of Antarctic latitude intertidal occurrences for 1416 species/taxa, verified Exploration, many national expeditions included biologists. using the World Register of Marine Species (WoRMS Edito- None of these early expeditions concentrated explicitly on rial Board 2015), from 229 locations (Fig. 1). Records date intertidal research; however, many opportunistically sampled back to 1897. The majority of data came from published the shores of Antarctica and the sub-Antarctic. Later manuscripts (see Appendix S1), with additional digital researchers and expeditions went on to undertake dedicated records from the Smithsonian Antarctic Invertebrates (http:// qualitative and quantitative research in the intertidal. The invertebrates.si.edu/antiz/). For South Africa, we used the history of Southern Ocean intertidal research and references Natural Geography in Shore Areas (NaGISA) data set from used to create the intertidal database are listed in the Global Biodiversity Information Facility (GBIF) (Rigby Appendix S1. et al., 2005). A record was considered to be intertidal if it 2 Journal of Biogeography ª 2016 The Authors. Journal of Biogeography

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