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BIOGEOGRAPHY of SOUTHERN POLAR BRYOZOANS D Barnes, S De Grave

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BIOGEOGRAPHY of SOUTHERN POLAR BRYOZOANS D Barnes, S De Grave BIOGEOGRAPHY OF SOUTHERN POLAR BRYOZOANS D Barnes, S de Grave To cite this version: D Barnes, S de Grave. BIOGEOGRAPHY OF SOUTHERN POLAR BRYOZOANS. Vie et Milieu / Life & Environment, Observatoire Océanologique - Laboratoire Arago, 2000, pp.261-273. hal- 03186927 HAL Id: hal-03186927 https://hal.sorbonne-universite.fr/hal-03186927 Submitted on 31 Mar 2021 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. VIE ET MILIEU, 2000, 50 (4) : 261-273 BIOGEOGRAPHY OF SOUTHERN POLAR BRYOZOANS D.K.A. BARNES* S. DE GRAVE** * Department of Zoology and Animal Ecology, University Collège Cork, Lee Mailings, Cork, Ireland ** The Oxford University Muséum of Natural History, Parks Road, Oxford, 0X1 3PW, United Kingdom ANTARCTICA ABSTRACT. - Despite the central rôle of Antarctica in the history and dynamics of BRYOZOANS oceanography in mid to high latitudes of the southern hémisphère there remains a POLAR FRONTAL ZONE paucity of biogeographic investigation. Substantial monographs over the last centu- ZOOQEOORAPHY ry coupled with major récent revisions have removed many obstacles to modem évaluation of many taxa. Bryozoans, being particularly abundant and speciose in the Southern Océan, have many useful traits for investigating biogeographic pat- terns in the southern polar région. They fossilise well, transitory morpho-stages between species are brief and most bryozoans have reduced dispersai opportunities due to possession of benthic larvae. Further to previously described broad faunistic patterns we demonstrate that bryozoans show sub-patterns within the Polar Frontal Zone. Four bryozoogeographical zones are recognised in Antarctica waters: East Antarctic seas, West Antarctic seas, the Antarctic Peninsula and the Scotia Arc. We discuss différences between species, genus and family level patterns in three ma- rine orders (of differing geological âge) and factors responsible for création of such distributions. ANTARCTIQUE RÉSUME. - En dépit du rôle central de l'Antarctique dans l'histoire et la dyna- BRYOZOAIRES mique de l'océanographie des latitudes moyennes à élevées de l'hémisphère sud, ZONE FRONTALE POLAIRE les investigations en biogéographie restent rares. Les monographies substancielles ZOOGÉOGRAPHIE des dernières décennies associées aux révisions majeures récentes ont levé divers obstacles dans l'évaluation moderne de nombreux taxons. Les Bryozoaires de l'Océan Sud, particulièrement abondants et riches en espèces, présentent de nom- breux traits significatifs pour les investigations sur la distribution biogéographique dans la région polaire du Sud. Ils se fossilisent bien, les morpho-stades ontogénéti- ques sont brefs et la plupart des Bryozoaires offrent peu d'opportunités à la disper- sion en raison de leurs larves benthiques. Au-delà des patterns à grande échelle déjà décrits, nous démontrons que les Bryozoaires présentent des sous-ensembles dans la Zone Frontale Polaire. Quatre zones zoogéographiques du groupe peuvent être reconnues dans les eaux antarctiques: les mers de l'Est et de l'Ouest antarctique, la Péninsule antarctique et l'Arc Scotia. Nous discutons des différences entre patterns au niveau des espèces, des genres et des familles appartenant à trois ordres marins (d'âge géologique différent) et des facteurs responsables d'une telle répartition. INTRODUCTION and fauna (Sara et al. 1992, Beu et al. 1997, Glasby & Alvarez 1999). Essentially, the results of thèse studies are in agreement with the biogeographic The continent of Antarctica and outlying scheme proposed by Hedgpeth (1969) and further subantarctic islands are isolated from most of the elaborated upon by Dell (1972). The basic feature land and continental shelves of the globe both of this scheme is three concentric zones denoting oceanographically (by the Antarctic Circumpolar the High Antarctic région around the continent it- Current) and bathymetry (by abyssal depths). His- self, the Antarctic Région extending to the Polar torically however, various plant and animal taxa Frontal Zone (PFZ) and the Subantarctic Région common to the austral landmasses bear testimony north of the PFZ. Within this tripartite division, a to a period when South America, Africa and the séries of districts is recognised, largely centred southern Australasian islands formed the around the various islands groups of the supercontinent Gondwana. Various attempts have Subantarctic Région. Bryozoans are potentially in- been made at biogeographic investigations of the valuable for historical zoogeographic comparisons, southern Austral marine flora (e.g. Clayton 1994) in view of their limited dispersai through a pre- 262 BARNES DKA, DE GRAVE S dominance of benthic short duration larvae abun- Here we investigate the distributions and affini- dance. Furthermore the taxon is highly abundant in ties of bryozoans south of 47° in the southern hémi- Southern Océan waters and is easily accessible due sphère, including Magellanic Chile, Argentina, the to its pre-dominance in shallow waters. Bryozoans Falkland Islands, South Georgia, Bouvet Island, have some interesting biogeographical links be- Crozet Islands, Prince Edward Islands, Kerguelen tween the cool water margins of the southern At- Islands, Heard Island, Campbell Islands, South lantic, Pacific and Indian Océans (see Gordon Sandwhich Islands, South Orkney Islands, South 1989, Moyano 1996). Within Antarctic waters, Shetland Islands, Antarctic Peninsula, Weddell bryozoan collections have been made for more than Sea, Bellingshausen Sea, Ross Sea and East Ant- a century but from an early stage (Waters 1904) the arctic marine régions. diversity of the bryofauna was realised. Although major monographs were undertaken on some bryo- zoan groups, such as the Cyclostomatida (Borg 1926, 1944), others, such as the Cheilostomatida, METHOD were in a state of taxonomic flux. When combined with the level of Southern Océan exploration, high rate of species discovery and description of new The taxonomic référence material used in this study gênera and lack of tertiary fossil knowledge it is were the works of Busk (1884), Borg (1926, 1944), Has- unsurprising that comprehensive bryozoogeo- tings (1943), Brown (1952), Rogick (1965), Androsova graphical analyses have not been undertaken. The (1968), Lopez Gappa (1978), d'Hondt & Redier (1977), last few décades have seen a dramatic explosion of d'Hondt (1984), Moyano (1982, 1983, 1985, 1991, 1996, taxonomic and ecological work in southern polar 1999), Gordon (1984, 1986, 1989), the studies of Hay- waters for various reasons. Many new Antarctic ward and co-workers summarised in Hayward (1995) cheilostomatid bryozoan species and gênera have and Ostrovsky & Taylor (1996). In addition samples were collected over 8 years (from 1990-1998) from Tier- been described, whilst the taxonomic status of ra del Fuego, the Falkland Islands, South Georgia and many others has been elucidated (see Hayward Bird Island, the South Orkney Islands, the South She- 1995 and références therein). Early collections tland Islands, the Northern Antarctic Peninsula and Sou- from the subantarctic islands were studied by Busk thern Antarctic Peninsula. The main study localities and (1884) and further explored by d'Hondt & Redier islands are shown in Fig. 1. (1977) and d'Hondt (1984). In neighbouring ré- A species (présence-absence only) matrix by sites gions Moyano (1974) and Lopez Gappa (1978) de- was constructed for Cheilostomatida, Ctenostomatida scribed Chilean and Argentinian species, whilst and Cyclostomatida separately. This matrix was subjec- Hayward & Cook (1979, 1983) named South Afri- ted to Detrended Correspondence Analysis, as ordina- can species and Brown (1952), Powell (1967) and tion techniques are more appropriate if the underlying Gordon (1984, 1986, 1989) listed the rich fauna of structure in the data matrix consists of a gradient, rather New Zealand waters. than a more divisive structure. In common with ail ordi- nation techniques, DCA arranges the sites in a two-di- The Southern Océan has been found, at certain mensional (or higher) space, in which the points (i.e. depths, to be highly diverse (Brey et al. 1994) with sites) are arranged such that the points close together certain groups such as the polycheates and bryo- correspond to sites with similar species composition and zoans being particularly well represented (Clarke points that are far apart correspond to sites that are dissi- 1992) whilst other taxa most notably decapod and milar in species composition. DCA is considered a more balanamorph crustaceans largely being absent superior ordination technique, as it corrects for two of south of 54°. Some (but few) bryozoan taxa are the major faults of earlier techniques, i.e. the fact that common to Antarctica, South America, South Af- the ends of the axes are often compressed relative to the rica and Australasia (Moyano 1985). The extensive middle of the axes and the fact that the second axis often work on Magellanic species have demonstrated shows a systematic relation with the first axis. In the présent analysis, only the first two axes
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