Name: Prof. Dr. Angelika Brandt
Institute: Zoological Institute and Zoological Museum
Address: Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
Phone: +40-40-4123-2278
Fax: +40-40-4123-3937
Email: abrandt@zoologie.uni-hamburg.de

Title of Proposal

Scientific Objectives and Pertinent Investigations already carried out
Objectives: To elucidate the potential origin of Antarctic benthic taxa (i.e. analyse the evolutionary biology and explain current community pattern), we must entirely focus on work on the ocean floor during one expedition, hence no time will be available for process studies in the water column.

It is supposed that in the Cenozoic the continental ice sheet never extended sufficiently down the shelf to completely eradicate the Antarctic shelf fauna (Brandt, 1991; Clarke & Crame, 1992). In the geological past, the breakup of Gondwana, and the subsequent isolation of Antarctica, the climatic changes with intermittent periods of global warming (influenced by the Milankovitch cyclicity) combined with global sea-level changes possibly determined faunal distributional ranges (migration processes in and out of the Antarctic) and limits. Extensions of the ice sheet may have enhanced speciation processes on the Antarctic continental shelf, suitably named the Antarctic "diversity pump" (Clarke & Crame, 1992).

The most appropriate tool for studies of evolutionary biology or zoogeographic origin of taxa is phylogenetic analysis. The fossil record greatly facilitates the recognition of ancient or derived character states (Brandt, 1998, in press), but unfortunately fossils are only available for some taxa. Rich fossil material is available from molluscs and decapod crustaceans, but none of the fossil species are still extant in the Antarctic nowadays. Other frequent Antarctic taxa, which are characterized by a high degree of endemism and which radiated in the Southern Ocean, like sponges, ascidians, ophiuroids, pycnogonids, and peracarid crustaceans are not well known in the fossil records. Besides the lack of fossils, systematic knowledge is too scarce for some of these taxa to be used for phylogenetic analyses of the recent Antarctic species. For this reason at present only few analyses of phylogenetic biogeography have been done for some peracarid taxa (e.g. Watling & Thurston, 1989; Brandt, 1991, 1992; Wägele & Brandt, 1992), and eunicemorph polychaetes (Orensanz, 1990), and evolution and origin of most Antarctic taxa remain unclear. However, first faunistic inventories of many Antarctic peracarid and other taxa have been completed since the 1960’s (e. g. Barthel & Tendal, 1994; Dell, 1972; 1990; Knox & Lowry, 1977; Andriashev, 1987; Hartman, 1964, 1966, 1978; Hartmann-Schröder & Rosenfeldt, 1988, 1989; Hain, 1990; Hartmann, 1997; Sieg, 1988; Brandt, 1991; Mühlenhardt-Siegel, 1988, in press; Brandt et al., submitted; Orensanz, 1990; Watling & Thurston, 1989; De Broyer & Jazdzewski, 1993, 1996; Warén & Hain, 1996; Herman & Dahms, 1992). These data provide the basis for faunistic comparisons of different regions of investigation and in combination with the data gathered during the planned cruise will help to find answers to questions of the potential origin of Antarctic benthic taxa as well as to the question of colonization of the deep sea from the Antarctic (submergence versus emergence of species):

• Are the present distributions of taxa the result of progressive retractions of the species from a former more cosmopolitic distribution, which was established during Jurassic or Cretaceous periods, when Gondwana was still clustered, or are these Gondwanian relicts?
• or have some of the taxa radiated in the Antarctic because of the extinction of potential competitors, i.e. has the emergence of new, adaptive zones (e.g. Peracarida because of the extinction of brachyurans) and occurrence of mass extinctions in the Antarctic in the Tertiary opened up previously occupied adaptive zones, and thus provided opportunities for spectacular adaptive radiations (Myers & Giller, 1988)?
• Is the Antarctic deep sea a reservoir of high species diversity within all taxa?
• Does the Antarctic deep-sea fauna differ from that of the deep sea of the other oceans?
• Is there still faunal exchange with the benthic fauna found in the area of the Antarctic Peninsula and the Magellan area or are there distinct topographical barriers to migration in and out of Antarctica via the deep sea?
• Is there a link between the Antarctic shelf and the deep-sea fauna of the Southern Ocean in present and past?
  

The major steps of the breakup of Gondwana are summarized in more detail by Crame (1998 in press). In Jurassic times Laurasia and Gondwana were still connected (Kennett, 1977, Shackleton & Boersma, 1981). The breakup of Gondwana continued in the Cretaceous period with the isolation and northward movement of Africa, completed about 90 Ma ago. After the first rifting between Antarctica and Australia about 100 Ma ago (e.g. Lawver et al., 1992), full deep-water separation between Australia and Antarctica subsequently enhanced the establishment of a current around the eastern part of the Antarctic in the beginning of the Eocene until the Oligocene (» 55-35 Ma ago) (Kennett et al., 1975; Kennett & Stott, 1991). The subsequent establishment of the Circumpolar Current was interrupted at that time only by the existence of the land connection between the Antarctic Peninsula and South America (compare Lawver et al., 1992; Storey, 1996; Crame, 1998 in press). However, this current probably already supported the cooling of the ocean water and the subsequent isolation of Antarctica (compare Huber & Watkins, 1992). With the opening of the Drake Passage (beginning some 28 to 23 Ma ago) and the establishment of the Scotia Arc the Circumpolar Current became gradually effective and this led to another decline in the Antarctic surface water temperature (Crame, 1992; 1994; 1997; Clarke & Crame, 1992). The final formation of a deep-sea passage between South America and Antarctica (some 15 MA ago, compare Crame, 1998 in press) was probably a key event for the evolution of many taxa of these environments, as shown for the Antarctic krill (Euphausia superba and E. crystallorophias) and E. vallentinii from the Subantarctic (Patarnello et al., 1996).

The reconstruction of the development of the Scotia Arc illustrated that in the early Cenozoic (about 50 Ma ago), the islands of the Scotia Arc and the South Shetland Islands were situated much closer together, a fact which led to the hypothesis that Antarctica might have been recolonised by some taxa via the Scotia Arc before these islands were disassembled due to the faster movement of the Antarctic plate in clockwise direction (Lawver et al., 1992). A schematic illustration of the present condition shows that these islands are separated by deep-sea areas (cf. Hellmer & Bersch, 1985), and this separation will certainly make active immigration difficult for many shelf species. Even for the deep-sea species we do not know how and whether they might have colonized the Antarctic slopes or even shelf. Therefore, deep-sea data are necessary from different areas in the Scotia Arc region, in the deep Weddell Sea, and off the Antarctic Peninsula into the Drake Passage. It will also be interesting to see whether the Antarctic deep sea is as diverse as the shelf and whether the diversity differs from that of other deep-sea areas (compare Bluhm et al., 1995; Bluhm, 1993; Gray, 1993; 1994; Gray et al, 1997; Grassle, 1977; Grassle and Maciolek, 1992; Grassle and Morse-Porteous, 1987; Arntz et al., 1997).

• Connections with already existing national and international projects

Up to now, only very few deep-sea investigations have been carried out in the Southern Ocean, and none have been devoted exclusively to the deep sea. Some data were collected in the years between 1950 and 1960 as part of Russian and U.S. American expeditions (Eltanin, Glacier, Akademik Kurchatov, Akademik D. Mendeleiev) which mainly focussed on the shelf off the South Orkneys and South Sandwich Islands. The Beagle Channel was sampled in 1873-1876 (HMS Challenger), including several deep-sea stations.

More recent programmes, such as EPOS and EASIZ, also included collection of deep-sea data, but again only as an addition to sampling arrays concentrating on the shelf. GLOBEC, and JGOFS, projects carried out in the deep sea, both concentrate on process studies in the pelagic realm, and other large Antarctic projects, such as the Spanish Antarctic Programme (Programa Nacional de Investigación en la Antártida) are carried out exclusively in shallow waters.The only information on the benthic deep-sea fauna in the Weddell Sea available to date is based on a few stations sampled during EASIZ II in the austral summer of 1998, and because of the time necessary for sample analysis it is still preliminary (cruise report in preparation, Ber. Polarforsch). The EU projects BENGAL and OMEX focus more on process studies, however, the information obtained - even if in the North Atlantic - will be very useful for the planned Antarctic deep-sea cruise as well. Moreover, the faunistic data collected during those projects can be as useful as the aims of the programmes such as DIVERSITAS or the SYSTEMATICS AGENDA 2000 - Charting the Biosphere.

Reviews

• (not applicable)
• The project will be reviewed by:
• ? German proposals will be sent to the German Science foundation, we expect colleagues from other nations to submit their own proposals to their national science foundations
• (not applicable)

Geographic Location of Proposed Project

• study area, location of transects and/or stations with map (obligatory)

? The study areas for both legs (compare 6.) of the proposed deep-sea expedition with RV Polarstern shall be the following: After leaving Punta Arenas sampling shall start in the Drake passage along a sill, which is shallower than the surrounding waters. Subsequently, transect work will follow off the South Orkney Islands, east of the Antarctic Peninsula, some stations will then be taken in the deep Weddell Sea on the way to the Kapp Norvegia sampling area, where a transect is planned (see below). From there the expedition will proceed to an older island, situated slightly east of one of the South Sandwich Islands, and finally another transect into the deep sea is planned off South Georgia.

In this respect the work should proceed as follows (compare Table 7 for sampling gear):

Leg 1 (@ five weeks minimum)

1. in the Drake Passage (on the shallower sill) in 1000 m depth at approximatelly 57.9°S 64°W, another 1000 m station should follow at about 59.30°S 59°W, then a 1500 m station should be taken at about 60°S 58°W, and finally a station with all gear will be taken in approximatelly 3500-4000 m at 61°S 57°W,
2. a transect of stations starting from 750 m, 1000 m, 1500 m, 2500 m, and 3500-4000 m shall be taken off the South Orkneys,
3. a similar transect with stations starting from 750 m, 1000 m, 1500 m, 2500 m, and 3500-4000 m shall be taken east of the Antarctic Peninsula.

Leg 2 (@ seven weeks minimum)

1. three deep stations in approximatelly 4000 m depth will be taken in the Weddell Sea on the way to Kapp Norvegia,
2. a transect of stations similar to leg 1 (750 m, 1000 m, 1500 m, 2500 m, and 3500-4000 m) shall be taken off Kapp Norvegia,
3. a similar transect shall be taken off an older island situated east of one of the South Sandwich Islands,
4. and finally a similar transect shall be taken off South Georgia.

From there it is planned to steam back to Punta Arenas directly.

? It is necessary to work from board of RV Polarstern, because sea ice will be encountered, even during the Antarctic summer in the area of the South Orkney Islands, in the area east of the Antarctic Peninsula, across the Weddell Sea and also off Kapp Norvegia.

Special Logistic Requirements ? 10.000 m of wire

• sufficient crew to work for 24 hours (three teams in shifts). Because of the long wire times we have to try to get enough crew members to work for 24 hours and to be able to employ even heavy gear during the nights. If necessary and the costs for the additional crew members cannot be paid by the BMBF, we have to apply for these extra costs to the German Science Foundation.

Proposed Time Frame

• Specific requirements concerning months and/or season and year
• ? Two legs of the planned deep-sea expedition (both starting and ending in Punta Arenas) shall be carried out between October and April of 2002, duration needed for twelve weeks including only steaming and station time, excluding supply tasks (for more detailed time frame see 7)
• Short justification of requested time frame, e.g., dependence on ice conditions, weather, etc.
• twelve weeks are the minimum of shiptime needed for both legs because of the long steaming time to the areas of investigation and also the time needed to take samples in the deep sea (see 6.-8.) (pay-out and haul time for deep sampling is time consuming).
• for this expedition we are depending on the Antarctic summer or early autumn season, because of the ice conditions, which would not allow working in some of the proposed areas in the Antarctic spring

Alternate time frames (not applicable)

Planned Gear and Instruments

• coring devices, grabs, nets, sondes, etc.

  - CTD
  - multiple-box corer (MG), includes video system
  - multiple corer (MUC)
  - epibenthic sledge (EBS)
  - Agassiz trawl (AGT)
  - REMOTS
  - possibly any camera system (FTS) (who can supply / help?)
  - traps (T)

Required Gear/ Instruments to be Provided by the Ship or the AWI

• winches, cranes, PDRs, computers (boats and helicopters not applicable)

  We would like to borrow the epibenthic sledge, Agassiz trawl and grab system from the AWI (could somebody help with the backup of one of these gears, we could supply an epibenthic sledge)

Required Laboratory Space and Special Accomodations

• laboratories, lab containers, cooling rooms or freezers

  - a minimum of two cool containers for keeping live animals (below deck)
  - cool room for chemicals and frozen material at -30°C, and -12°C
  - freezer for e.g. molecular biology at -80°C

• space for containers on deck, below deck, on top deck, etc.
  

  - collaborators, you are requested to bring your own container if you need one, e.g.for keeping animals or with your equipment
  - does anybody need an isotope container? If so, could that person function as a safety guard for that laboratory

Number of Participants

• essential
  (see list of interested potential participants) circular sent to:

• Wolf Arntz, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany.
• Dieter Gerdes, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Mike Thomson, British Antarctic Survey, High Cross, Madingley Road, Cambridge CB 3 0 ET, United Kingdom
• Alistair Crame, British Antarctic Survey, High Cross, Madingley Road, Cambridge CB 3 0 ET, United Kingdom
• Andy Clarke, British Antarctic Survey, High Cross, Madingley Road, Cambridge CB 3 0 ET, United Kingdom
• Horst-Kurt Schminke, Universität Oldenburg, Fachbereich 7, Biologie, Postfach 2503, Carl von Ossietzky Str., 26111 Oldenburg
• Wolfgang Wägele, Ruhr Universität Bochum, Geb. ND 05/755, 44780 Bochum
• Michael Türkay, Senckenberg Museum, Senckenberganlage 25, 60325 Frankfurt
• Hjalmar Thiel, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Peter Schalk, University of Amsterdam, Mauritskade 61, NL-1092 AD Amsterdam, Netherlands
• Magda Vinx, University of Gent, Zoological Institute, Marine Biology Section, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium
• Claude De Broyer, Institute Royal des Sciences Naturelles de Belgique, Rue Vautier 29, 1040 Bruxelles, Belgium
• John Gage, Scottish Association for Marine Science, Dunstaffnage Marine Laboratory, Oban, United Kingdom
• Paul Tyler, Department of Oceanography, University of Southampton, Southampton, United Kingdom
• Volker Storch, Ruprecht-Karls-Universität Heidelberg, Zoologisches Institut, Abt. Morphologie, Ökologie, Im Neuenheimer Felf 230, 69120 Heidelberg, Germany
• Mike Thurston, Southampton Oceanographic Centre Empress Dock, Southampton SO14 3ZH, United Kingdom
• Tony Rice, Institute of Oceanographic Sciences Deacon Laboratory, Wormley, Godalming, Surrey GU8 5UB, United Kingdom
• John Lambshead, Department of Zoology, The Natural History Museum, London SW7 5BD, United Kingdom
• Gordon Patterson, Department of Zoology, The Natural History Museum, London SW7 5BD, United Kingdom
• Maria Cristina Gambi, Stazione Zoologica "Anton Dohrn", Villa Comunale, Ischia, I-80121 Napoli, Italia
• Brigitte Hilbig, Zoological Institute and Zoological Museum, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
• Ute Mühlenhardt-Siegel, Zoological Institute and Zoological Museum, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
• Peter Emschermann, Fakultät für Biologie, Albert-Ludwigs-Universität, Schänzlestraße 1, 7800 Freiburg, Germany
• John Gray, Department of Marine Zoology and Marine Chemistry, School of Biology, PB 1064, University of Oslo, N-0316 Oslo, Norway
• Wim Vader, University of Tromsø, Tromsø Museum, Department of Zoology, Lars Thørings veg 10, N-9000 Tromsø, Norway
• Hartmut Bluhm, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Christian Borowski, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Anders Waren, Swedish Museum of Natural History, Department of Invertebrate Zoology, Box 50007, S-10405 Stockholm, Sweden
• Ole Tendal, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark
• Joerundur Svavarsson, University of Iceland, Department of Biology, Grensasvegur 12, 108 Reykjavik, Iceland
• Torleiv Brattegard, University of Bergen, Institut for Fisheries and Marine Biology Høyteknologisenteret, 5020 Bergen, Norway
• Jean-Claude Hureau, Museum National d’ Histoire Naturelle, 57, rue Cuvier, F-75005 Paris, France
• Heye Rumohr, University of Kiel, Institute for Marine Sciences, Düsternbrooker Weg 20, 24105 Kiel, Germany
• Don Rhoads, 22 Widgeon Road, Falmouth, MA 02540, USA
• James A. Blake, ENSR, 89 Water Street, Woods Hole, MA 02543, USA
• Margo Branch & husband, University of Cape Town, Zoological Institute
• German Pequeño, Zoological Department, Universidad Austral de Chile, Casilla 567, Valdivia, Chile
• Barbara Hecker, Hecker Consulting, 26 Mullen Way, Falmouth, MA 02540, USA
• Jörn Thiede, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Paul Dayton, Scripps Institution of Oceanography, A-001, La Jolla, California 92093, USA
• Oliver Coleman, Humboldt Museum, Invalidenstr. 43, 10115 Berlin, Germany
• Karin Riemann-Zürneck, Alfred-Wegener-Institut für Polar-und Meeresforschung, Postfach 120161, 27515 Bremerhaven, Germany
• Michael Rex, Department of Biology, University of Massachusetts, Boston, Massachusetts 02125, USA
• J. Frederick Grassle, Institute of Marine and Coastal Science, Cook College, P.O. Box 231, New Brunswick, NJ 08903, USA
• Michael Spindler, Institute for Polar Ecology, University of Kiel, Seefischmarkt, Geb. 12, Wischhofstr. 1-3, D-24148 Kiel, Germany
• DFG
  

Collaboration with the Following Groups

• essential (AWI, BAS, IRSNB,Scripps Institution of Oceanography, all scientists on board)
  

Financial Support

• funds will be/ are being raised (compare 4)
• financial support is guaranteed by (this information cannot be provided yet)
  

Remarks

Appendices