Shallow-Water Mega-Epibenthic Communities in the High Antarctic (Weddell Sea) and the West Coast of the Antarctic Peninsula (Bellingshausen Sea)
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Shallow-water mega-epibenthic communities in the high Antarctic (Weddell Sea) and the west coast of the Antarctic Peninsula (Bellingshausen Sea) Juana Marianne Raguá Gil Universität Bremen, 2004 Stiftung Alfred-Wegener-Institut für Polar- und Meeresforschung in der Helmholtz-Gemeinschaft Stiftung Alfred-Wegener-Institut für Polar- und Meeresforschung, Bremerhaven Shallow-water mega-epibenthic communities in the high Antarctic (Weddell Sea) and the west coast of the Antarctic Peninsula (Bellingshausen Sea) Juana Marianne Raguá Gil Vorgelegt in der Universität Bremen (Fachbereich 2 – Biologie/Chemie) als Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) Bremen, März 2004 Advisory Committee: 1. Gutachter: Prof. Dr. Wolf E. Arntz (Alfred-Wegener-Institut für Polar- und Meeresforschung, Bremerhaven, Germany) 2. Gutachter: Prof. Dr. Andrew Clarke (British Antarctic Survey, Cambridge, United Kingdom) 1. Prüfer: Dr. habil. Julian Gutt (Alfred-Wegener-Institut für Polar- und Meeresforschung, Bremerhaven, Germany) 2. Prüfer. Prof. Dr. Matthias Wolff (Zentrum für Marine Tropenökologie Bremen, Germany) „Gedruckt mit Unterstützung des Deutschen Akademischen Austauschdienstes und des Alfred-Wegener-Instituts für Polar und Meeresforschung“ To my parents and my brother, for their unlimited love and support “PRIMERO ESTABA EL MAR TODO ESTABA OSCURO, NO HABIA SOL NI LUNA, NI GENTE, NI ANIMALES, NI PLANTAS EL MAR ERA LA MADRE, LA MADRE NO ERA GENTE NI NADA, NI COSA ALGUNA. ELLA ERA ESPIRITU DE LO QUE IBA A VENIR Y ELLA ERA PENSAMIENTO Y MEMORIA” Mitología Kogui - Kogui: grupo indígena colombiano - “FIRST THERE WAS THE SEA EVERYTHING WAS DARK, THERE WAS NEITHER SUN NOR MOON NO PEOPLE, NEITHER ANIMALS NOR PLANTS THE SEA WAS THE MOTHER, THE MOTHER WAS NO PEOPLE NOR ANYTHING, NOT EVEN SOMETHING. SHE WAS THE SPIRIT OF WHAT WAS GOING TO COME AND SHE WAS THOUGHT AND MEMORY” Kogui Mythology - Kogui: Colombian Indian group - Contents I Contents ABSTRACT .................................................................................................................. III ZUSAMMENFASSUNG............................................................................................... V RESUMEN ................................................................................................................. VIII 1 INTRODUCTION ...................................................................................................... 1 1.1 Origin of the Antarctic fauna................................................................................ 1 1.2 Biogeographic remarks: Antarctic Peninsula and high Antarctic ............................................................................................................... 2 1.3 Imaging techniques............................................................................................... 3 1.4 Objectives of this study ........................................................................................ 6 2 STUDY AREAS .......................................................................................................... 7 2.1 Weddell Sea.......................................................................................................... 7 2.1.1 The benthic fauna of the eastern Weddell Sea...................................................... 9 2.2 Bellingshausen Sea ............................................................................................. 10 2.2.1 The benthic fauna of the Bellingshausen Sea..................................................... 11 2.3 North-east Greenland.......................................................................................... 12 3 MATERIAL AND METHODS ................................................................................ 13 3.1 Video-Sampling.................................................................................................. 13 3.2 Identification....................................................................................................... 15 3.3 Data analyses ...................................................................................................... 15 4 RESULTS AND DISCUSSION................................................................................ 19 4.1 Distribution and community structure of shallow-water Antarctic mega-epibenthic communities............................................................................ 19 4.1.1 Hypotheses.......................................................................................................... 19 4.1.2 Antarctic shallow-water mega-epibenthos: shaped by circumpolar dispersion or local conditions?............................................................................................. 20 4.2. Spatial patterns of the Antarctic benthos by examples....................................... 25 4.2.1 Horizontal seriation ............................................................................................ 26 4.2.2 Depth zonation.................................................................................................... 31 4.3 Biodiversity at different spatial scales: Parallelism with the Arctic?................. 35 4.3.1 Gamma and beta diversity .................................................................................. 37 4.3.2 Alpha diversity.................................................................................................... 41 II Contents 5 PUBLICATIONS....................................................................................................... 45 5.1 Publication I: Raguá-Gil JM, Gutt J, Clarke A, Arntz WE, Antarctic shallow- water mega-epibenthos: shaped by circumpolar dispersion or local conditions? 47 5.2 Publication II: Raguá-Gil JM, Gutt J, Arntz WE, Antarctic mega-epibenthos: horizontal seriation and depth zonation by examples......................................... 69 5.3 Publication III: Raguá-Gil JM, Gutt J, Starmans A, Arntz WE, Differences in mega-epibenthic diversity on polar shelves at different spatial scales ............... 98 6 ACKNOWLEDGEMENTS .................................................................................... 123 7 REFERENCES ........................................................................................................ 127 8 APPENDICES.......................................................................................................... 139 8.1 List of abbreviations ......................................................................................... 139 8.2 List of ROV stations ......................................................................................... 140 8.3 Tables of abundance of taxa ............................................................................. 141 8.4 List of taxa ........................................................................................................ 150 Abstract III ABSTRACT The topic of the present study is to compare shallow-water mega-epibenthic fauna of two biogeographic areas in the Antarctic, the Antarctic Peninsula (Bellingshausen Sea) and the high Antarctic (Weddell Sea). Continental shelves around Antarctica are unusual in being deep, and consequently shallow habitats at depths <150 m are scarce. In the Weddell Sea, for example, only two shallow sites with water depths between 55 and 160 m are known: the inner parts of Atka Bay and a recently discovered shallow bank off Four Seasons Inlet NE of Kapp Norvegia. In contrast, the Antarctic Peninsula has a true coast and thus, true littoral areas are common. This study (1) characterises and describes the distribution and spatial patterns of shallow-water (55-160 m) mega- epibenthic fauna in the Antarctic Peninsula (Marguerite Bay, Bellingshausen Sea) and at two sites in the high Antarctic (Weddell Sea), (2) analyses community structures and (3) compares benthic biodiversity in these areas at different spatial scales and with additional data from deep Antarctic shelf sites and corresponding shallow and deep areas in the Arctic, originated from previous studies. In order to achieve the composition of the Antarctic benthic community, all organisms (>1 cm) were identified and counted on a total of 17 sea bed video transects conducted at 14 stations using a Remotely Operated Vehicle (ROV) during four Antarctic expeditions with R.V. Polarstern and R.R.S. James Clark Ross (ANT XIII, XV, XVII; JR37). Multivariate analyses revealed significant faunal differences between Marguerite Bay (Bellingshausen Sea) and the Weddell Sea stations, Atka Bay and Four Seasons Bank. Echinoderms, especially ophiuroids, dominated Marguerite Bay, bryozoans and ascidians were abundant at Atka Bay, and hydroids and gorgonians were well represented at Four Seasons Bank. These differences can mainly be explained by the influence of local environmental conditions that are probably the primary factor shaping the Antarctic shallow-water epifauna, and not an intensive exchange with larger depths or limited dispersion due to scarce and isolated shallow areas. In addition, modes of reproduction and characteristics of the early life history (e.g. brooding, viviparity or budding) of key taxa may also shape patterns of species distribution in shallow benthic Antarctic communities. IV Abstract Spatial patterns of Antarctic benthic shelf communities were analysed. Stations with non-significant “seriation” values, indicating no sharp faunistic discontinuities, have the smallest range of species or faunal-heterogeneity but intermediate