appendix ix

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Bathybiaster loripes Sladen, 1889. Leptychaster kerguelenensis Smith, 1876. Order Valvatidae. Asterinidae. Tremaster mirabilis Verrill, 1879. Odontasteridae.

APPENDIX IX

Marine benthic species of the Prince Edward Islands Evgeny A. Pakhomov

Thanks to the considerable oceanographic survey work undertaken at the Prince Edward Islands, and a substantial interest in the benthos by several taxonomists (especially M.L. Branch), the benthos has been reasonably well documented at the islands, at least in terms of the species that are present. Spatial surveys have typically not been undertaken. The reference list below provides an introduction to the literature. References Arnaud, F. & Branch, M.L., 1991. The Pycnogonida of subantarctic Marion and Prince Edward Islands: Illustrated keys to the species. South African Journal of Antarctic Research 21, 65-71.

Branch, M.L., Arnaud, P.M., Cantera, J. & Gianakouras, D., 1991. The benthic Mollusca and Brachiopoda of subantarctic Marion and Prince Edward Islands. South African Journal of Antarctic Research 21, 45-64.

Branch, M.L. 1994. The Polychaeta of subantarctic Marion and Prince Edward Islands: Illustrated keys to the species and results of the 1982-1989 University of Cape Town surveys. South African Journal of Antarctic Research 24, 3-52.

Branch, M.L., Griffiths, C.L., Kensley, B. & Sieg, J., 1991. The benthic Crustacea of subantarctic Marion and Prince Edward Islands: Illustrated keys to the species and results of the 1982-1989 University of Cape Town surveys. South African Journal of Antarctic Research 21, 3-44.

S.L. Chown & P.W. Froneman (eds.) The Prince Edward Islands • Land-Sea Interactions in a Changing Ecosystem, 416-430

Branch, M.L. & Hayward, P.J. 2005. New species of cheilostomatous Bryozoa from subantarctic Marion and Prince Edward Islands. Journal of Natural History 39, 2671-2704.

Rowe F.W.E. & Clark A.M., 1975. Notes on some echinoderms from Marion Island. Bulletin Britain Museum Natural History (Zoology) 28, 187-190.

Branch, M.L., Jangoux, M., Alva, V., Massin, C.I. & Stampanato, S., 1993. The Echinodermata of subantarctic Marion and Prince Edward Islands. South African Journal of Antarctic Research 23, 37-70.

Phylum Arthropoda Class Crustacea Subclass Cirripedia

Order Thoracica Suborder Lepadomorpha Scalpellum flavum Hoek, 1883 Subclass Malacostraca

Superorder Peracarida Order Amphipoda Suborder Gammaridea Acanthonotozomatidae Amphilochidae

Gnathiphimedia urodentata Bellan-Santini and Ledoyer, 1986 Gitanopsis marionis (Stebbing, 1888) Gitanopsis squamosa (Thomson, 1880) Gammaropsis longitarsus (Schellenberg, 1931)

Corophiidae

Gammaropsis sp. Haplocheira barbimana (Thomson, 1879) Lembos sp.

Didymocheliidae

Didymochelia edwardi Bellan-Santini and Ledoyer, 1986

Eophliantidae

Cylindryllioides mawsoni Nicholls, 1938 Atyloella magellanica (Stebbing, 1888) Atylopsis emarginatus Stebbing, 1888 Djerboa furcipes Chevreux, 1906 Eusiroides aberrantis Bellan-Santini and Ledoyer, 1986

Eusiridae

Eusiroides georgianus KH Barnard, 1932 Gondogeneia spinicoxa Bellan-Santini and Ledoyer, 1974 Harpinioides drepanocheir Stebbing, 1888 Oradarea edentata Thurston, 1974 Oradarea ocellata Thurston, 1974

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Ceradocopsis dufresni Bellan-Santini and Ledoyer, 1986 Ceradocopsis kergueleni Schellenberg, 1926 Gammaridae

Paramoera fissicauda (Dana, 1852) Pontogeneiella brevicornis (Chevreux, 1906) Schraderia gracilis Pfeffer, 1888 Carangolia cornuta Bellan-Santini and Ledoyer, 1986

Haustoriidae

Cardenio paurodactylus Stebbing, 1888 Urothoe marionis Bellan-Santini and Ledoyer, 1986 Urothoides lachneessa Stebbing, 1888 Cerapus oppositus KH Barnard, 1932 Ischyrocerus sp. A Ischyrocerus sp. B

Ischyroceridae

Jassa alonsoae Conlan, 1990 Pseuderichthonius gaussi Schellenberg, 1926 Pseudischyrocerus crenatipes Bellan-Santini and Ledoyer, 1986 Pseudischyrocerus distichon (KH Barnard, 1930) Ventojassa georgiana (Schellenberg, 1931)

Leucothoidae Liljeborgiidae

Leucothoe spinicarpa (Abildgraad, 1879) Leucothoe sp. Liljeborgia longicornis (Schellenberg, 1931) Liljeborgia pseudomacronyx Bellan-Santini and Ledoyer, 1986 Acontiostoma marionis Stebbing, 1888 Cheirimedon femoratus (Pfeffer, 1888) Hippomedon kergueleni (Miers, 1875) Kerguelenia antiborealis Bellan-Santini and Ledoyer, 1986

Lysianassidae

Lepidepecreella tridactyla Bellan-Santini, 1972 Parawaldeckia kidderi (Smith, 1876) Pseudorchomene coatsi (Chilton, 1912) Stomacontion acutibasalis (Bellan-Santini and Ledoyer, 1974) Stomacontion pepinii (Stebbing, 1888) ? Monoculodes antarcticus KH Barnard, 1932

Oedicerotidae

Monoculodes scrabriculosus KH Barnard, 1932 ? Oediceroides cinderella Stebbing, 1888

Pardaliscidae

Pardalisca marionis Stebbing, 1888

Phoxocephalidae

Harpinia obtusifrons Stebbing, 1888

Pleustidae

? Pleusymtes sp. Podocerus danae Stebbing, 1888

Podoceridae

Podocerus danae armatus Bellan-Santini and Ledoyer, 1986 Podocerus capillimanus Nicholls, 1938

Sebidae

418

Seba saundersii Stebbing, 1875

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Stegocephalidae

Andaniella integripes Bellan-Santini and Ledoyer, 1986 Probolisca ovata (Stebbing, 1888) Proboloides elliptica (Schellenberg, 1931) Proboloides sp. A

Stenothoidae

Proboloides sp. B Pseudothaumatelson cyproides Nicholls, 1938 Stenothoe sp. Thaumatelson herdmani Walker, 1906

Stilipedidae Talitridae

Alexandrella inermis Bellan-Santini and Ledoyer, 1986 Hyale grandicornis (Kroyer, 1945) Hyale hirtipalma (Dana, 1852) Suborder Caprellidea

Aeginellidae

? Eupariambus sp.

Order Isopoda Suborder Epicaridea Bopyridae

Bopyridae gen sp. Suborder Valvifera Antarcturus aculeatus Kussakin, 1967

Arcturidae

Microarcturus hirticornis (Monod, 1926) Neastacilla marionensis (Beddard, 1886)

Pseudidotheidae

Arcturides cornutus Studer, 1882 Suborder Anthuridea

Paranthuridae

Califanthura pingouin Kensley, 1980 Paranthura possessia Kensley, 1980 Suborder Flabellifera

Serolidae

Serolis septemcarinata Miers, 1847 Aega cf. crozetensis Kussakin and Vassina, 1982

Aegidae

Aega falklandica Kussakin, 1967 Aega semicarinata Miers, 1875 Gymodocella sp.

Sphaeromatidae

Dynamenella eatoni (Miers, 1875b) Euvallentina darwini (Cunningham, 1871) Exosphaeroma gigas (Leach, 1818) Suborder Gnathiidea

Gnathiidae

Gnathia antarctica (Studer, 1884)

Appendix IX

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Suborder Asellota Dendrotionidae Joeropsidae

Acanthomunna spinipes (Vanhöffen, 1914) Joeropsis curvicornis (Nicolet, 1849) Joeropsis marionis Beddard, 1886 Austrofilius furcatus Hodgson, 1910 Austroniscus ectiformis Vanhöffen, 1914

Janiridae

Iais pubescens (Dana, 1852) Ianisera trepidus Kensley, 1976 Notasellus sarsi Pfeffer, 1887 Munna instructa Cleret, 1973

Munnidae

Munna neglecta Monod, 1931 Munna neozelanica Chilton, 1892 Santia bicornis (Cleret, 1973)

Santiidae

Santia cf. hofsteni (Nordenstam, 1933) Santia cf. marmoratus (Vanhöffen, 1914)

Pleurogonidae

Munnogonium sp. Paramunna sp.

Order Tanaidacea Suborder Monokonophora Superfamily Apseudoidea Apseudidae

Apseudes spectabilis (Studer, 1884) Suborder Dikonophora Superfamily Paratanaoidea

Paratanaidae

Paratanais oculatus (Vanhöffen, 1914)

Leptocheliidae

Pseudonototanais werthi (Vanhöffen, 1914)

Nototanaidae

Nototanais antarcticus (Hodgson, 1902) Nototanais dimorphus (Beddard, 1886) Superfamily Tanaoidea Allotanais hirsustus (Beddard, 1886a) Tanais hirsustus, Stebbing, 1919 Pancoloides litoralis (Vanhöffen, 1914)

Tanaidae

Sinelobus stanfordi (Richardson, 1901) Zeuxo phytalensis Sieg, 1980 Zeuxoides helleri (Gerstaecker, 1888) Zeuxoides pseudolitoralis, Sieg 1980

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Superorder Eucarida Order Decapoda Suborder Natantia Campylonotidae Hippolytidae Nematocarcinidae

Campylonotus capensis Bate, 1888 Chorismus antarcticus (Pfeffer, 1887) Nauticaris marionis Bate, 1888 Nematocarcinus lanceopes Bate, 1888 Suborder Reptantia Section Branchyura

Hymenosomatidae

Halicarcinus planatus Fabricius, 1793 Section Anomura

Lithodidae

Lithodes murrayi Henderson, 1888 Paralomis aculeatus (Henderson, 1888)

Paguridae

Pagurodes inarmatus Henderson, 1888

Parapaguridae

Sympagurus dimorphus (Studer, 1883) = Parapagurus dimorphus (Studer, 1883)

Galatheidae

Munida spinosa Henderson, 1888

Chirostylidae

Uroptychus insignis Henderson, 1888

Phylum Brachiopoda Liothyrella sp. Magellania kerguelensis (Davidson, 1880) Platydia anomioides Scacchi, 1844

Phylum Mollusca Class Solenogastres Solenogastres gen. sp.

Class Bivalvia Acesta sp. Adacnarca marionensis (Smith, 1885) Astarte longirostris d’Orbigny, 1846 Condylocardia sp. Cuspidaria kerguelenensis Smith, 1885 Cuspidaria cf. tenella, Smith 1907 Cyclopecten aviculoides Smith, 1885 Escalima goughensis (Melvill & Standen, 1907) Gaimardia adamsiorum Osorio & Arnaud, 1984 Gaimardia trapesina (Lamarck, 1819) Hiatella solida (Sowerby, 1834) Kellia nuculina Martens, 1881 Kidderia minuta Dall, 1876

Appendix IX

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Kidderia oblonga (Smith, 1898) Lasaea consanguinea Smith, 1877 Laternula elliptica King & Broderip, 1831 Limatula pygmaea (Philippi, 1845) Limatula simillima Thiele, 1912 Limopsis marionensis Smith, 1885 Lissarca miliaris (Philippi, 1845) Myonera fragilissima Smith, 1885 Mysella charcoti (Lamy, 1906) Neolepton umbonatum Smith, 1885 Nucula kerguelenensis Thiele, 1912 Palliolum clathratum Martens, 1881 Philobrya barbata Thiele, 1912 Philobrya kerguelenensis (Smith, 1885) Philobrya quadrata Thiele, 1912 Pteria sp. Thracia meridionalis Smith, 1885 Thyasira marionensis Smith, 1885 Yoldiella profundorum Melvill & Standen, 1912 Dosinia cf. pubescens Limopsis lilliei Smith, 1915 Lyonsiella cf. radiata Dall, 1889

Class Gastropoda Admete specularis (Watson, 1882) Amauropsis anderssoni Strebel, 1906 Amauropsis prasina (Watson, 1881) Anatoma euglypta (Pelseneer, 1903) Balcis amblia (Watson, 1883) Banzarecolpus austrina (Watson, 1881) Brookula cf. crassicostata (Strebel, 1908) Calliostoma delli McLean & Andrade, 1982 Cerithiella cf. werthi Thiele, 1912 Cerithiopsilla sp. Chlanidotella modesta (Martens, 1885) Diaphana kerguelenensis Thiele, 1912 Eatoniella kerguelenensis regularis (Smith, 1915) Eatoniella subrufescens (Smith, 1875) Eumetula macquariensis Tomlin, 1948 Eumetula sp. Falsilunatia soluta (Gould, 1847)

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Eumetula macquariensis Tomlin, 1948 Eumetula sp. Fusitriton magellanicus (Roeding, 1798) “Jeffreysia” edwardiensis (Watson, 1886) Kerguelenella lateralis (Gould, 1849) Laevilitorina caliginosa (Gould, 1848) Margarella expansa (Sowerby, 1838) Margarella porcellana Powell, 1951 Marseniopsis pacifica Bergh, 1886 Microdiscula subcanaliculata (Smith, 1875) Nacella delesserti (Philippi, 1849) Omalogyra cf. amotus (Philippi, 1841) Onoba transenna (Watson, 1886) Onoba paucicarinata (Ponder, 1983) Pareuthria regulus Watson, 1883 Powellisetia principis (Watson, 1886) Probuccinum edwardiensis (Watson, 1882) Puncturella conica (d’Orbigny, 1841) Retusa sp. Sinezona sp. Skenella edwardiensis (Watson, 1886) Solariella infundibulum (Watson, 1879) Spirotropis studeriana (Martens, 1878) Tectonatica impervia (Philippi, 1845) Toledonia limnaeaeformis (Smith, 1877) Torellia angulifera Waren, Arnaud & Cantera, 1986 Trophon declinans Watson, 1882 Trophon sepus Watson, 1882 Turbonilla lamyi Hedley, 1916 Typhlodaphne platamodes (Watson, 1881) Typhlodaphne translucida (Watson, 1881) Balcis sp. Bathydomus sp. Prosipho sp. Solariella sp. Toledonia elata Thiele, 1912

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Class Polyplacophora Hemiarthrum setulosum Dall, 1876 Lepidopleurus dorsuosus Haddon, 1886 Lepidopleurus kerguelenensis (Haddon, 1886) ? aff Placiphorella sp.

Class Scaphopoda Fissidentalium sp.

Class Cephalopoda Octopus magnificus Villaneuva et al. 1991

Phylum Pycnogonida Cilunculus kravcovi Pushkin, 1973 Tanystylum cavidorsum Stock, 1957 Tanystylum neorthetum Marcus, 1940

Ammotheidae

Tanystylum oedinotum Loman, 1923 Tanystylum ornatum Flynn, 1928 Pantopipetta australis (Hodgson, 1914) Austrodecus elegans Stock, 1957

Austrodecidae

Austrodecus goughense Stock, 1957 Austrodecus tristanense Stock, 1957

Callipallenidae

Pseudopallene glutus Pushkin, 1975

Pycnogonidae

Pycnogonum platylophum Loman, 1923

Colossendeidae

Colossendeis megalonyx megalonyx Hoek, 1881

Endeidae

Endeis viridis Pushkin, 1976

Rhynochothoracidae

Rhynchothorax australis Hodgson, 1907 Nymphon gracilipes Miers, 1875

Nymphonidae

Nymphon longicoxa Hoek, 1881 Nymphon sp.

Phylum Echinodermata Class Asteroidea Order Paxillosida Astropectinidae

Bathybiaster loripes Sladen, 1889 Leptychaster kerguelenensis Smith, 1876

Order Valvatidae Asterinidae

Tremaster mirabilis Verrill, 1879 Acodontaster elongatus (Sladen, 1889)

Odontasteridae

Odontaster meridionalis (Smith, 1876) Odontaster penicillatus (Philippi, 1870) Odontaster validus Koehler, 1906

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Ceramaster patagonicus (Sladen, 1889) Goniasteridae

Hippasteria hyadesi Perrier, 1891 Hippasteria falklandica Fisher, 1940 Pseudarchaster discus Sladen, 1889

Order Spinulosida Crossaster penicillatus Sladen, 1889 Solasteridae

Lophaster stellans Sladen, 1889 Solaster dianae Stampanato & Jangoux Solaster regularis Sladen, 1889

Pterasteridae

Diplopteraster semireticulatus (Sladen, 1882) Pteraster affinis Smith, 1876

Korethrasteridae

Peribolaster folliculatus Sladen, 1889

Poraniidae

Porania antarctica Smith, 1876

Ganeriidae

Perknaster densus Sladen, 1889 Henricia fisheri AM Clark, 1962 Henricia sp. aff. H. obesa (Sladen, 1889)

Echinasteridae

Henricia praetans (Sladen, 1889) Henricia sp. aff. H. simplex (Sladen, 1889) Henricia sp. aff. H. studeri (Perrier, 1891)

Order Forcipatulida Labidiasteridae

Labidiaster annulatus Sladen, 1889 Anasterias rupicola (Verrill, 1876) Anteliaster australis Fisher, 1940 Anteliaster scaber Smith, 1876

Asteriidae

Diplasterias meridionalis (Perrier, 1875) Pedicellaster hypernotius Sladen, 1889 Smilasterias scalprifera (Sladen, 1889) Smilasterias triremis (Sladen, 1889)

Class Ophiuroidea Suborder Euryalina Asteronichidae Gorgonocephalidae

Asteronyx loveni Müller & Troschel, 1842 Astrotoma agassizi Lyman, 1875 Gorgonocephalus chilensis (Philippi, 1858) Suborder Ophiomyxina

Ophiomyxidae

Ophioscolex (Ophiolycus) nutrix Mortensen, 1936

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Suborder Gnatophiurina Amphiuridae Amphiura algida Koehler, 1911 Amphiurinae

Amphiura angularis angularis Lyman, 1879 Amphiura antarctica Studer, 1876 Amphiura tomentosa Lyman, 1879

Amphilepidinae Amphilepis sp. aff. A. gymnopora Hertz, 1927 Suborder Laemophiurina Ophiacantha imago Lyman, 1878 Ophiacantha rosea Lyman, 1878 Ophiacanthidae

Ophiacantha vivipara Ljungman, 1870 Ophiocymbium cavernosum Lyman, 1880 Ophiolebes scorteus Lyman, 1878 Suborder Chilophiurina

Ophiodermatidae

Toporkovia antarctica (Lyman, 1882)

Ophiuridae Amphiophiura sp. Ophiocten amitinum Lyman, 1878 Ophiocten banzarei Madsen, 1964 Ophiurinae Ophionotus hexactis (Smith, 1876) Ophiophycis mirabilis Koehler, 1901 Ophiurolepis intorta (Lyman, 1878) Stegophiura elevata (Lyman, 1878)

Class Echinoidea Suborder Echinina Echinidae

Sterechinus agassizi Mortensen, 1936 Suborder Temnopleurina

Temnopleuridae

Pseudechinus marionis Mortensen, 1936

Class Holothuroidea Order Aspidochirotida Synallactidae Gephyrothuridae

Mesothuria edwardensis Massin, 1992 Synallactes challengeri (Théel, 1886) Pseudostichopus mollis (Théel, 1886)

Order Dendrochirotida Cucumaria kerguelensis (Théel, 1886) Cucumariidae

Pseudocnus laevigatus (Verrill, 1876) Cladodactyla crocea croceoides (Vaney, 1908)

Psolidae

426

Psolidium incertum (Théel, 1886) Psolus paradubiosus Carriol & Feral, 1985

The Prince Edward Islands • Land-Sea Interactions in a Changing Ecosystem

Order Apodida Chiridotidae

Taeniogyrus contortus (Ludwig, 1874) Paradota marionensis Massin, 1992

Class Crinoidea Order Comatulae Suborder Macrophyeata Antedonidae

Phrixometra exigua (Carpenter, 1888) Eumorphometra hirsuta (Carpenter, 1888)

Phylum Annelida Class Polychaeta Order Orbiniida Orbiniella dayi (Branch) Orbiniidae

Leitoscoloplos kerguelensis (McIntosh, 1885) Scoloplos (Leodamas) marginatus (Ehlers, 1897) Naineris sp.

Paraonidae

Cirrophorus lyra (Southern, 1914) Paraonella sp.

Order Spionida Boccardia polybranchia (Haswell, 1885) Polydora armata Langerhans, 1881 Spionidae

Rhynchospio glutaea (Ehlers, 1897) Scolelepis marionis Branch ? Spiophanes bombyx Claparéde, 1870 Spiophanes tcherniai Fauvel, 1951

Chaetopteridae

Chaetopterus variopedatus Reiner, 1804 Tharyx sp.

Cirratulidae

Cirratulus sp. Timarete antarcticus Monro, 1930 Cirriformia sp. recorded by Day 1971

Order Capitellida Capitella ‘capitata’ (Fabricius, 1780) Capitellidae

Notomastus latericeus Sars, 1851 Notomastus sp.

Arenicolidae

Abarenicola ?assimilis Wells, 1963 Axiothella quadrimaculata Augener, 1914

Maldanidae

Lumbriclymenella robusta Arwidsson, 1911 Maldanella antarctica McIntosch, 1885 Nichomache lumbricalis Fabricius, 1780

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Order Opheliida Ammotrypane breviata Ehlers, 1913

Opheliidae

Travisia kerguelensis McIntosch, 1885 Scalibregma inflatum Rathke, 1843

Scalibregmidae

Order Flabelligerida ? Brada mammillata Grube, 1877

Flabelligeridae

Flabelligera induta Ehlers, 1897

Order Terebellida Petta assimilis McIntosch, 1885

Pectinariidae

Ampharete kerguelensis McIntosch, 1885

Ampharetidae

Anobothrus patagonicus (Kinberg, 1867)

Terebellidae Polycirrinae

Polycirrus hamiltoni Benham, 1921 Polycirrus kerguelensis (McIntosch, 1885)

Thelepinae Thelepus extensus Hutchings & Glasby, 1987 Lanice marionensis Branch Amphitritinae

Nicolea venustula (Montagu, 1818) Neoleprea streptochaeta (Ehlers, 1897) Axionice godfroyi (Gravier, 1911)

Trichobranchidae

Terebellides stroemii kerguelensis McIntosch, 1885

Sabellidae Fabriciinae Oriopsis limbata (Ehlers, 1897) Sabellinae Perkinsiana antarctica (Kinberg, 1867) Serpulidae Filograninae Protula tubularia tubularia (Montagu, 1803) Serpulinae Serpula vermicularis vermicularis Linnaeus, 1767 Helicosiphon platyspira Knight-Jones, 1978 Leodora perrieri (Caullery & Mesnil, 1897) Spirorbidae

Romanchella inventis (Harris, 1969) Paralaeospira levinseni (Caullery & Mesnil, 1897) Spirorbis patagonicus Day, 1971

Order Phyllodocida Suborder Phyllodociformia Genetyllis polyphylla (Ehlers, 1897) Phyllodocidae

Steggoa magalaensis (Kinberg, 1866) ? Steggoa hunteri (Benham, 1921) ? Eteone aurantiaca Smarda, 1861

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The Prince Edward Islands • Land-Sea Interactions in a Changing Ecosystem

Suborder Aphroditiformia Laetmonice producta Grube, 1877

Aphroditidae Polynoidae Lepidonotinae

Admetella longipedata McIntosch, 1885 Eulagisca corrientis McIntosch, 1885 Eucrantis mollis (McIntosch, 1876) Harmothoe crosetensis (McIntosch, 1885) Harmothoe magellanica (McIntosch, 1885)

Harmothoinae

Harmothoe spinosa Kinberg, 1855 Harmothoe ?kerguelensis Lagisca exanthema (Grube, 1856) Malmgreniella fimbria Branch Polyeunoa laevis McIntosch, 1885 Suborder Nereidiformia ? Hesione sp.

Hesionidae

Kefersteinia fauveli Averincev 1972 Autolytus sp. cf. A. simplex (Ehlers, 1900)

Syllidae Exogoninae

Exogone heterosetosa McIntosch, 1885 Brania rhopalophora (Ehlers, 1897) Pionosyllis nutrix Munro, 1936

Syllinae Pionosyllis cf ehlersiaeformis Augener, 1913 Typosyllis ‘variegata’ (Grube, 1860) Amblyosyllis granosa Ehlers, 1897 Typosyllis sp. Eusyllinae ? Pharyngeovalvata natalensis Day, 1951 Eusyllis blomstrandi Malmgren, 1867 Eusyllis kerguelensis McIntosch, 1885 ? Neanthes kerguelensis (McIntosch, 1885) Nereididae

Platynereis australis Smarda 1861 Pseudoneries anomalia Gravier, 1901 Suborder Glyceriformia Glycerella magellanica (McIntosch, 1885)

Glyceridae

Glycera kerguelensis McIntosch, 1885 Protodorvillea kefersteini (McIntosch, 1869)

Goniadidae

Goniada brunnea (Treadwell, 1906 revised Moore, 1911)

Nephtyidae

Aglaophamus ornatus Hartman, 1967

Order Amphinomida Eophrosinidae

Euphrosine cirrata Sars, 1862

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Order Eunicida Eunicidae Onuphidae Lumbrineridae

Eunice pennata (Muller, 1776) ? E. edwardsi (McIntosch, 1885) Nothria anoculata Orensanz, 1974 Lumbrinereis sp. aff L. fragilis ? Lumbrinereis heteropoda Marenzeller, 1879

Phylum Coelenterata Class Hydrozoa Tubulariidae

Tubularia sp.

Myriothelidae

Myriothela meridiana Briggs, 1939

Corynidae

?Coryne conferta Allman, 1877

Bongainvilliidae

Rhizorbagium antarcticum (Hickson & Gravely)

Hydractiniidae

Hydractinia parvispina Hartlaub, 1905 Campanularia subantarctica n. sp. Millard

Campanulariidae

Obelia geniculata (Linnaeus, 1758) Silicularia rosea Meyen, 1834 Sertularella picta (Meyen, 1834)

Sertulariidae

Symplectoscyphus marionensis n. sp. Symplectoscyphus sp.

Phylum Bryozoa Bitectiporidae

Parkermavella corrugata Branch & Hayward, 2005

Buffonellodidae

Ipsibuffonella hyalina Branch & Hayward, 2005 Cornucopina antlera Branch & Hayward, 2005

Bugullidae

Cornucopina bryonyae Branch & Hayward, 2005 Cornucopina elongata Branch & Hayward, 2005

Calloporidae Catenicellidae

Pyriporoides judyae Branch & Haward, 2005 Valdemunitella dianae Branch & Hayward, 2005 Catenicella glabrosa Branch & Hayward, 2005 Talivittaticella frigida (Waters, 1904)

Cellariidae

Paracellariae elizabethae Branch & Hayward, 2005

Cribrilinidae

Filaguria lithocrustata Branch & Hayward, 2005

Hippothoidae

Celleporella marionensis Branch & Hayward, 2005

Lacernidae

Phonicosia jousseaumei Jullen, 1888 Calloporina spinosa Branch & Hayward, 2005

Microporellidae

Fenestrulina pumicosa Branch & Hayward, 2005 Microporella mandibulata Branch & Hayward, 2005

Phidoloporidae Sclerodomidae

430

Reteporella gigantea (Busk. 1884) Reteporellina berylae Branch & Hayward, 2005 Sclerodomus myriozoides (Busk, 1884)

The Prince Edward Islands • Land-Sea Interactions in a Changing Ecosystem

The Prince Edward Islands Land-Sea Interactions in a Changing Ecosystem

Edited by Steven L Chown Stellenbosch University Pierre William Froneman Rhodes University

The Prince Edward Islands Land-Sea Interactions in a Changing Ecosystem Published by SUN PReSS, an imprint of AFRICAN SUN MeDIA (Pty) Ltd., Stellenbosch 7600 www.africansunmedia.co.za www.sun-e-shop.co.za All rights reserved. Copyright © 2008 Chown & Froneman No part of this book may be reproduced or transmitted in any form or by any electronic, photographic or mechanical means, including photocopying and recording on record, tape or laser disk, on microfilm, via the Internet, by e-mail, or by any other information storage and retrieval system, without prior written permission by the publisher. First edition 2008 ISBN 978-1-920109-85-1 Set in 11 Baskerville Cover design and typographic setting by Liezel Meintjes SUN PReSS is an imprint of AFRICAN SUN MeDIA (Pty) Ltd. Academic, professional and reference works are published under this imprint in print and electronic format. This publication may be ordered directly from www.sun-e-shop.co.za Printed and bound by ASM/USD, Ryneveld Street, Stellenbosch, 7600.

Contents Foreword.................................................................................................................. xi Preface ................................................................................................................... xiii Contributor Affiliations....................................................................................... xv 1. 1.1 1.2 1.3 1.3.1 1.3.2 1.3.3 1.4 1.5

The Prince Edward Islands in a global context................................. 1 Steven L. Chown and P. William Froneman Environmental change in the Antarctic region............................................ 2 The Prince Edward Islands..........................................................................4 Research at the Prince Edward Islands........................................................6 Early terrestrial work.................................................................................... 6 Marine investigations................................................................................... 7 Recent developments.................................................................................... 8 This book...................................................................................................... 8 References....................................................................................................9

2. 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 2.10

Oceanographic setting of the Prince Edward Islands...................17 Johann R. E. Lutjeharms and Isabelle J. Ansorge Conceptual developments.......................................................................... 20 Climate change...........................................................................................23 General current environment of the Prince Edward Islands . .................. 23 General hydrographic environment of the Prince Edward Islands ..........25 Frontal systems in the vicinity of the Prince Edward Islands.....................27 Water masses in the vicinity of the Prince Edward Islands........................30 Eddy behaviour in the vicinity of the Prince Edward Islands....................31 Conclusions................................................................................................33 Acknowledgements.....................................................................................33 References..................................................................................................34

3. 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8

Climate and climate change................................................................39 Peter C. le Roux The hyper-oceanic climate.........................................................................40 Atmospheric circulation and the frontal nature of the weather................. 44 Spatial variation in climate on Marion Island............................................49 Contemporary changes in climate.............................................................52 Mechanisms driving sub-Antarctic climate change.................................... 55 Future climate trends..................................................................................58 Acknowledgements.....................................................................................59 References..................................................................................................59

4. 4.1 4.2 4.3 4.3.1 4.3.2 4.3.2.1 4.3.2.2 4.3.2.3 4.3.3 4.3.4 4.4 4.5 4.6

Geology, geomorphology and climate change.................................65 Jan C. Boelhouwers, K. Ian Meiklejohn, Stephen D. Holness and David W. Hedding Geological evolution...................................................................................66 The glacial record...................................................................................... 67 Geomorphology......................................................................................... 74 Weathering ................................................................................................74 Present-day periglacial landforms, processes and environment ................ 75 Periglacial landforms..................................................................................75 Periglacial processes....................................................................................78 The periglacial environment......................................................................80 Other slope processes................................................................................. 82 Zoogeomorphology.................................................................................... 83 The record of Holocene climate change...................................................88 Discussion and conclusions........................................................................90 References..................................................................................................92

5. 5.1 5.2 5.3 5.4 5.5 5.6 5.7

Biology in theoceanographic environment....................................... 97 Christopher D. McQuaid and P. William Froneman Phytoplankton studies................................................................................. 98 Zooplankton studies.................................................................................102 Nekton studies.......................................................................................... 109 Biological interactions..............................................................................110 Impact of global climate change on biology of the oceanographic ............. environment............................................................................................. 113 Acknowledgements...................................................................................115 References................................................................................................116

6. 6.1 6.1.1 6.1.2 6.1.3 6.1.4 6.1.5 6.2 6.2.1 6.2.2 6.2.3 6.2.4 6.2.5 6.3 6.3.1 6.3.2 6.3.3

Pelagic predators.................................................................................121 Peter G. Ryan & Marthán N. Bester The abundance, distribution and phenology of pelagic predators ashore........................................................................... 122 Pinnipeds..................................................................................................122 Penguins................................................................................................... 124 Albatrosses................................................................................................127 Petrels.......................................................................................................128 Other seabirds..........................................................................................129 Diets and foraging methods of pelagic predators....................................130 Pinnipeds..................................................................................................130 Penguins................................................................................................... 133 Albatrosses................................................................................................133 Petrels.......................................................................................................134 Other seabirds..........................................................................................135 Distribution of pelagic predators at sea................................................... 135 Pinnipeds..................................................................................................135 Penguins................................................................................................... 136 Albatrosses................................................................................................137

6.3.4 6.3.5 6.4 6.5 6.5.1 6.5.2 6.5.3 6.5.4 6.5.5 6.6 6.7 6.8 6.9 6.10

Petrels.......................................................................................................139 Other seabirds..........................................................................................140 Movements between islands..................................................................... 140 Recent population changes...................................................................... 141 Pinnipeds..................................................................................................141 Penguins................................................................................................... 143 Albatrosses................................................................................................144 Petrels.......................................................................................................145 Other seabirds..........................................................................................147 Summary of human impacts on pelagic predators.................................. 147 Predicting the effects of climate change................................................... 148 Other pelagic predators............................................................................ 149 Acknowledgements...................................................................................150 References................................................................................................151

7. 7.1 7.1.1 7.1.2 7.2 7.3 7.4

Nutrient dynamics in the vicinity of the Prince Edward Islands........................................................................165 Valdon R. Smith and P. William Froneman Oceanic inputs of nutrients to the Marion Island terrestrial system........ 166 Rainwater input.......................................................................................166 Bird and seal input................................................................................... 166 Terrestrial/pelagic coupling.....................................................................173 Impact of global climate change on nutrient dynamics of the Prince Edward Island ecosystem . ...........................................................175 References................................................................................................176

8. 8.1 8.1.1 8.1.1.1 8.1.1.2 8.1.1.3 8.1.2 8.1.2.1 8.1.2.2 8.1.2.3 8.1.2.4 8.1.3 8.1.3.1 8.1.3.2 8.1.3.3 8.1.3.4 8.2

Terrestrial and freshwater primary production and nutrient cycling . ..................................................................................181 Valdon R. Smith Terrestrial communities............................................................................ 182 Vegetation standing crop and primary production..................................182 Standing crop........................................................................................... 183 Primary production.................................................................................. 187 Growth season duration, primary productivity and production efficiency................................................................................190 Nutrient cycling........................................................................................191 Nutrient pool sizes....................................................................................191 Nutrient uptake, nutrient turnover times and nutrient costs of the ANP...............................................................................................193 Plant-available nutrient pools and their replenishment rates ..................195 Nutrient inputs......................................................................................... 196 Decomposition and nutrient mineralization............................................ 197 Decomposition and soil heterotrophic activity......................................... 197 Nutrient mineralisation and the role of soil fauna...................................198 Impact of invasive biota on nutrient cycling............................................ 200 Climate change and nutrient cycling........................................................201 Freshwater communities........................................................................... 203

8.2.1 8.2.2 8.2.3 8.3 8.4

The freshwater bodies..............................................................................203 Nutrient composition...............................................................................204 Primary production.................................................................................. 206 The trophic dynamics disparity between the island’s terrestrial and aquatic systems – how real is it?...............................................................207 References................................................................................................207

9. 9.1 9.2 9.2.1 9.2.2 9.2.3 9.2.4 9.2.5 9.2.6 9.2.7 9.3 9.3.1 9.3.2 9.3.3 9.3.4 9.3.5 9.4

Terrestrial vegetation and dynamics...............................................215 Niek J.M. Gremmen & Valdon R. Smith The Flora ................................................................................................ 216 Vegetation, plant communities and habitats............................................218 Mire Complex..........................................................................................219 Slope Complex.........................................................................................225 Fellfield Complex......................................................................................226 Polar Desert.............................................................................................. 227 Coastal Saltspray Complex...................................................................... 228 Biotic Complex . ......................................................................................229 Aquatic communities................................................................................230 Vegetation dynamics................................................................................. 231 Autogenic succession................................................................................231 Vegetation and climate change................................................................. 232 Changes in population sizes of indigenous animals.................................233 Impact of changing human activities....................................................... 236 Impact of invasions by alien plants.......................................................... 236 References................................................................................................241

10. 10.1 10.2 10.3 10.4 10.5 10.6 10.7 10.8

Spatial variation in the terrestrial biotic system.......................... 245 Melodie A. McGeoch, Peter C. le Roux, Elizabeth A. Hugo and Mawethu J. Nyakatya The model ............................................................................................... 247 The plant . ............................................................................................... 251 The landscape .........................................................................................254 The island . .............................................................................................. 261 Apparent mechanisms.............................................................................. 266 Implications of environmental change.....................................................267 Acknowledgements...................................................................................268 References................................................................................................270

11. 11.1 11.2 11.3 11.4 11.5 11.6

Regional membership: biogeography.............................................277 Steven L. Chown, David J. Marshall andEvgeny A. Pakhomov............................. 277 Historical biogeography: an ongoing conundrum...................................280 Ecological biogeography: determinants of richness................................. 285 New developments in phylogeography..................................................... 290 Conclusions..............................................................................................291 Acknowledgements...................................................................................292 References................................................................................................292

12. 12.1 12.2 12.2.1 12.2.2 12.2.3 12.2.4 12.3 12.4 12.5

Conservation and management.......................................................301 Marienne S. de Villiers and John Cooper The evolving management of the islands................................................ 302 Major conservation concerns at the islands............................................. 307 Introductions of alien species................................................................... 307 Factors affecting marine resources........................................................... 313 Litter, pollution and waste management.................................................. 315 Disturbance.............................................................................................. 319 Conclusions..............................................................................................321 Acknowledgements...................................................................................323 References................................................................................................324

13. 13.1 13.2 13.3 13.4 13.5 13.6 13.7 13.7.1 13.7.2 13.8 13.9 13.10

Human history.....................................................................................331 John Cooper Who first saw the islands? The discovery era, 1663-1776........................332 Seal tongues for breakfast and seal skins for shoes: the period of exploitation, 1800-1930.......................................................................334 Gentleman scientists in suits: the exploration era, 1840-1940................. 336 South Africans arrive to stay: the annexation of 1947-1948...................337 South Africans living in the sub-Antarctic, 1948 to present..................... 338 Early aliens: how some introduced mammals became established, and others not....................................................................... 339 Gender and race in the sub-Antarctic and the policy of Apartheid........342 Early female pioneers: thwarting the “no doors on the toilets” syndrome...................................................................................................... 342 Racial issues: from servants to leaders...................................................... 344 Postscript: looking to the future................................................................ 346 Acknowledgements...................................................................................346 References................................................................................................347

14. 14. 14.2 14.2.1 14.2.2 14.2.3 14.2.4 14.3 14.4

Conclusion: change in terrestrial and marine systems..............351 Steven L. Chown and P. William Froneman...................................................... 351 Historic change and evolutionary responses............................................ 352 Modern change across the biological hierarchy.......................................354 Biological invasions..................................................................................355 Climate change and terrestrial systems ................................................... 358 Climate change and marine systems........................................................361 Exploitation..............................................................................................362 Land-sea interactions in a changing ecosystem........................................363 References................................................................................................364



Appendices

I

Terrestrial and freshwater diatoms of the Prince Edward Islands ..........373 Bart Van de Vijver and Niek J.M. Gremmen

II

Hepatics of the Prince Edward Islands....................................................379 Niek J.M. Gremmen

III

Mosses of the Prince Edward Islands...................................................... 383 Ryszard Ochyra

IV

Vascular plants of the Prince Edward Islands.......................................... 390 Niek J.M. Gremmen and Valdon R. Smith

V

Lichens of the Prince Edward Islands ....................................................394 Dag O. Øvstedal and Niek J.M. Gremmen

VI

Freshwater invertebrates of the Prince Edward Islands...........................399 Evgeny A. Pakhomov and Steven L. Chown

VII

The Terrestrial invertebrates of the Prince Edward Islands....................401 Steven L. Chown and David J. Marshall

VIII

Marine plankton of the Prince Edward Islands region............................410 Evgeny A. Pakhomov

IX

Marine benthic species of the Prince Edward Islands............................. 416 Evgeny A. Pakhomov

X

Marine fish of the Prince Edward Islands region....................................431 Evgeny A. Pakhomov

XI

Birds recorded at the Prince Edward Islands...........................................435 Peter G. Ryan

XII

Mammals recorded at thePrince Edward Islands....................................439 Marthán N. Bester

Plates . ................................................................................................................441 Index . ................................................................................................................458

Contributor Affiliations Isabelle J. Ansorge

P. William Froneman

Department of Oceanography, University of Cape Town, South Africa

Department of Zoology and Entomology, Rhodes University, South Africa

Marthán N. Bester

Niek J. M. Gremmen

Department of Zoology and Entomology, University of Pretoria, South Africa

Data Analyse Ekologie, The Netherlands

Jan C. Boelhouwers

David W. Hedding

Department of Earth Sciences, Uppsala University, Sweden

Department of Geography, Geoinformatics and Meteorology, University of Pretoria, South Africa

Steven L. Chown

Stephen D. Holness

Centre for Invasion Biology, Stellenbosch University, South Africa

Department of Geography, Geoinformatics and Meteorology, University of Pretoria, South Africa

John Cooper

Elizabeth A. Hugo

Avian Demography Unit, University of Cape Town, South Africa

Department of Conservation Ecology and Entomology, Stellenbosch University, South Africa

Marienne S. de Villiers

Johann R. E. Lutjeharms

Avian Demography Unit, University of Cape Town, South Africa

Department of Oceanography, University of Cape Town, South Africa

xv

xvi

David J. Marshall

Dag O. Øvstedal

Department of Biology, Universiti Brunei Darussalam, Brunei

Botanical Institute, Norway

Melodie A. McGeoch

Peter C. le Roux

Department of Conservation Ecology and Entomology, Centre for Invasion Biology, Stellenbosch University, South Africa

Department of Conservation Ecology and Entomology, Stellenbosch University, South Africa

Christopher P. McQuaid

Evgeny A. Pakhomov

Department of Zoology and Entomology, Rhodes University, South Africa

Department of Earth and Ocean Sciences, University of British Columbia, Canada and University of Forth Hare, South Africa

K. Ian Meiklejohn

Peter G. Ryan

Department of Geography, Geoinformatics and Meteorology, University of Pretoria, South Africa

Percy Fitzpatrick Institute for African Ornithology, University of Cape Town, South Africa

Mawethu J. Nyakatya

Valdon R. Smith

Department of Conservation Ecology and Entomology, Stellenbosch University, South Africa

Department of Botany and Zoology, Stellenbosch University, South Africa

Ryszard Ochyra

Bart Van de Vijver

Wladyslaw Szafer Institute of Botany, Polish Academy of Sciences

National Botanical Garden of Belgium, Belgium