Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Systematics, phylogeny and historical biogeography of the Pentagonaster clade (Asteroidea : Valvatida : Goniasteridae)

Christopher Mah
+ Author Affiliations
- Author Affiliations

Department of Invertebrate Zoology, National Museum of Natural History, MRC-163, PO Box 37012 Smithsonian Institution, Washington, DC 20560, USA. Email: mahch@si.edu

Invertebrate Systematics 21(4) 311-339 https://doi.org/10.1071/IS06049
Submitted: 11 November 2006  Accepted: 9 June 2007   Published: 20 September 2007

Abstract

Morphology-based phylogenetic hypotheses developed for living and fossil goniasterid asteroids have provided several unique opportunities to study bathymetric and biogeographic shifts for an ecologically important group of prominent, megafaunal invertebrates. A cladistic analysis of 18 ingroup taxa employing 65 morphological characters resulted in a single most parsimonious tree. The tree supports assignment of the Atlantic Tosia parva (Perrier, 1881) and the Pacific Tosia queenslandensis Livingstone, 1932 to new, separate genera. The phylogenetic tree supports offshore to onshore bathymetric shifts between basal and derived taxa. The phylogeny is also consistent with historical events surrounding the separation of Antarctica from Australia and South Africa. Buterminaster Blake & Zinsmeister, 1988 from the Eocene La Meseta Formation, Antarctic Peninsula, was included in the phylogenetic analysis and is now supported as the only fossil species in the genus Pentagonaster Gray, 1840. Pentagonaster stibarus H. L. Clark, 1914 is separated from synonymy with P. dubeni Gray, 1847 and resurrected as a valid species. The new genus, Akelbaster, gen. nov., shows unusual new structures that resemble cribiform organs, although their function has not been determined. One specific ingroup lineage, including Tosia and Pentagonaster, attains a much larger adult size than those of its sister-taxa, suggesting that Cope’s rule may apply to asteroids within this clade. Pentagonaster and related genera are revised. Descriptions of four new genera and three new species are presented, including: Akelbaster novaecaledoniae, gen. nov., sp. nov., Ryukuaster onnae, gen. nov., sp. nov., Eknomiaster beccae, sp. nov., Pawsonaster parvus, gen. nov., comb. nov. and Anchitosia queenslandensis, gen. nov., comb. nov.


Acknowledgements

It is with profound gratitude that I thank Nadia Ameziane, Philippe Bouchet, Bertrand Richer de Forges, Thomas Schlager, and the participants of the BATHUS 3, SMIB 3, 4, 5, and VAUBAN cruises for collection of material that was used in this paper. I am also grateful for the assistance of Cynthia Ahearn (NMNHIZ), Cathy Groves (LACM), and Bob Van Syoc (CAS) with specimen loans. Dan Blake, University of Illinois at Urbana-Champaign provided important discussion and a review of the paper. Thanks to Loisette Marsh, Western Australian Museum, Maria Byrne, University of Sydney, Tim O’Hara, Museum Victoria, and Mike Barker, University of Otago for discussions and answering my questions regarding topics in the paper. O’Hara and two anonymous reviewers provided useful comments. Kate Neil, New Zealand National Institute of Water & Atmospheric Research and Andrzej Gazdzicki, Polish Academy of Sciences provided me with reference images of asteroid specimens. This research was partly funded by NSF Polar Programs Postdoctoral Fellowship Grant # 0631245.


References


Atkinson D. (1994) Temperature and organism size. A biological law for ectotherms? Advances in Ecological Research 25, 1–58. open url image1

Ayres W. O. (1851) Remarks on new asteroids. Proceedings of the Boston Society of Natural History 4, 118. open url image1

Bell J. (1905) The Echinoderma found off the coast of South Africa. Asteroidea. Marine Investigations in South Africa 3, 241–253. open url image1

Benavides-Serrato M., Borrero-Perez G. H., Solano O. D., Navas G. R. (2005) Listado taxonónomico de los asteroideos (Echinodermata: Asteroidea) de la plataforma y el talud superior del Caribe colombiano. Revista de Biologia Tropical suppl. 3 53, 171–194. open url image1

Benham W. B. (1909) Scientific results of the New Zealand Government Trawling Expedition, 1907. Echinoderma. Records of the Canterbury Museum 1, 83–116. open url image1

Birkeland C. (1974) Interactions between a sea pen and seven of its predators. Ecological Monographs 44, 211–232.
Crossref | GoogleScholarGoogle Scholar | open url image1

Blake D. B. (1983) Some biological controls on the distribution of shallow water sea stars (Asteroidea: Echinodermata). Bulletin of Marine Science 33, 703–712. open url image1

Blake D. B. (1987) Classification and phylogeny of post-Paleozoic sea stars (Asteroidea: Echinodermata). Journal of Natural History 21, 481–528.
Crossref | GoogleScholarGoogle Scholar | open url image1

Blake D. B. (1990) Adaptive zones of the class Asteroidea (Echinodermata). Bulletin of Marine Science 46, 701–718. open url image1

Blake D. B., Aronson R. B. (1998) Eocene stelleroids (Echinodermata) at Seymour I., Antarctic Peninsula. Journal of Paleontology 72, 339–353. open url image1

Blake D. B., Jagt J. W. M. (2005) New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) from the Netherlands and Denmark and their palaeobiological significance. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique. Sciences de la Terre 75, 183–200. open url image1

Blake D. B., Zinsmeister W. J. (1988) Eocene asteroids (Echinodermata) from Seymour Island, Antarctic Peninsula. Geological Society of America Memoir 169, 489–498. open url image1

Bryan P. J., McClintock J. B., Hopkins T. S. (1997) Structural and chemical defenses of echinoderms from the northern Gulf of Mexico. Journal of Experimental Marine Biology and Ecology 210, 173–186.
Crossref | GoogleScholarGoogle Scholar | open url image1

Carrera-Rodriguez C. J., Tommasi L. R. (1977) Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil). Boletim Instituto de Oceanografia São Paulo 26, 51–130. open url image1

Chapelle G., Peck L. S. (1999) Polar gigantism dictated by oxygen availability. Nature 399, 114–115.
Crossref | GoogleScholarGoogle Scholar | open url image1

Clark A. M. (1952) On some echinoderms from the British Museum. Transactions of the Royal Society of South Africa 33, 193–221. open url image1

Clark A. M. (1953) Notes on asteroids in the British Museum (Natural History) IV. Tosia and Pentagonaster. Bulletin of the British Museum (Natural History) Zoology 1, 397–411. open url image1

Clark A. M. (1976) Asterozoa from Amsterdam and St. Paul Islands, Southern Indian Ocean. Bulletin of the British Museum (Natural History). Zoology 30, 247–261. open url image1

Clark A. M. (1993) An index of names of recent Asteroidea – Part 2: Valvatida. Echinoderm Studies 4, 187–366. open url image1

Clark A. M.,, Rowe F. W. E. (1971) Monograph of shallow-water Indo-West Pacific echinoderms. Trustees of the British Museum (Natural History) 690, 1–238. open url image1

Clark A. M., and Downey M. E. (1992). ‘Starfishes of the Atlantic.’ (Chapman and Hall: London.)

Clark A. M., and Courtman-Stock J. (1976). ‘The Echinoderms of Southern Africa.’ (British Museum of Natural History: London.)

Clark H. E. S. (1970) Sea-stars (Echinodermata: Asteroidea) from Eltanin Cruise 26 with a review of the New Zealand asteroid fauna. Zoology Publications from Victoria University of Wellington 52, 1–34. open url image1

Clark H. E. S., and McKnight D. G. (2001). The Marine Fauna of New Zealand: Echinodermata: Asteroidea (sea-stars), order Valvatida. NIWA Biodiversity Memoir 117.

Clark H. L. (1898) The echinoids and asteroids of Jamaica. Circular. Johns Hopkins University 18(137), 4–6. open url image1

Clark H. L. (1914) The echinoderms of the Western Australian Museum. Records of the Western Australian Museum 1, 132–173. open url image1

Clark H. L. (1916). report on the sea-lillies, starfishes, brittlestars, and sea-urchins obtained by the FIS Endeavor on the coasts of Queensland, New South Wales, Tasmania, Victoria, South Australia, and Western Australia. In ‘Biological Results of the Fishing Experiments carried on by the F.I.S. Endeavor, 1908–1914. Endeavour Research Vol. 4’. pp. 1–123.

Clark H. L. (1923a) The echinoderm fauna of South Africa. Annals of the South African Museum 13, 221–435. open url image1

Clark H. L. (1923b) Some echinoderms from West Australia. Journal of the Linnean Society. Zoology 35, 229–251. open url image1

Clark H. L. (1926) Echinoderms from the South African Fisheries and Marine Biological Survey. 2. Sea-stars. Report-Fisheries and Marine Biological Survey of the Union of South Africa 4(7), 1–33. open url image1

Clark H. L. (1938). Echinoderms from Australia, an account of collections made in 1929 and 1932. Memoirs of the Museum of Comparative Zoology 55.

Clark H. L. (1946). The echinoderm fauna of Australia. Its composition and its origin. Carnegie Institution of Washington Publication 566.

Clarke A., Crame J. A. (1992) The Southern Ocean benthic fauna and climate change: a historical perspective. Philosphical Transactions of the Royal Society of London B 338, 299–309.
Crossref | GoogleScholarGoogle Scholar | open url image1

Clarke A., Aronson R. B., Crame J. A., Gili J.-M., Blake D. B. (2004) Evolution and diversity of the benthic fauna of the Southern Ocean continental shelf. Antarctic Science 16, 559–568.
Crossref | GoogleScholarGoogle Scholar | open url image1

Coleman N. (1994). ‘Sea Stars of Australasia and their Relatives.’ (Underwater Geographic Pty Ltd: Brisbane.)

Crump R. G. (1968) The flight response in Struthiolaria papulosa gigas Sowerby. New Zealand Journal of Marine and Freshwater Research 2, 390–397. open url image1

Currie D. R., Parry G. D. (1999) Impacts and efficiency of scallop dredging on different soft substrates. Canadian Journal of Fisheries and Aquatic Sciences 56, 539–550.
Crossref | GoogleScholarGoogle Scholar | open url image1

Dakin W. J. (1960). ‘Australian Seashores.’ (Angus and Robertson: Sydney.)

Dartnall A. J. (1969) New Zealand sea stars in Tasmania. Papers and Proceedings of the Royal Society of Tasmania 103, 53–55. open url image1

Dartnall A. J. (1972). Sea stars of Tasmania. Tasmanian Year Book 1972. pp. 72–79.

Dartnall A. J. (1980) Tasmanian echinoderms. Fauna of Tasmania Handbook 3, 1–83. open url image1

Davie P. (1998). ‘Wild Guide to Moreton Bay: Wildlife and Habitats of a Beautiful Australian Coast—Noosa to the Tweed.’ (Queensland Museum Publication: Brisbane.)

Dingle R. V., Lavelle M. (2000) Antarctic Peninsula Late Cretaceous-Early Cenozoic palaeoenvironments and Gondwana palaeogeographies. Journal of African Earth Sciences 31, 91–105.
Crossref | GoogleScholarGoogle Scholar | open url image1

Downey M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions to Zoology 126, 1–158. open url image1

Edgar G. J. (1997). ‘Australian Marine Life: The Plants and Animals of Temperate Waters.’ (Reed Books: Melbourne.)

Endean R. (1957) The biogeography of Queensland’s shallow-water echinoderm fauna (excluding Crinoidea), with a rearrangement of the faunistic provinces of tropical Australia. Australian Journal of Marine and Freshwater Research 8, 233–273.
Crossref | GoogleScholarGoogle Scholar | open url image1

Farquhar H. (1895) Notes on New Zealand echinoderms. Transactions and Proceedings of the New Zealand Institute , 194–208. open url image1

Farquhar H. (1897) A contribution to the history of New Zealand echinoderms. Journal of the Linnean Society of London: Zoology 26, 186–198. open url image1

Farquhar H. (1898) Notes on New Zealand starfishes. Transactions of the New Zealand Institute 30, 187–191. open url image1

Farquhar H. (1927) Notes on New Zealand seastars. New Zealand Journal of Science and Technology 9, 237–240. open url image1

Fell H. B. (1947) Key to the littoral asteroids of New Zealand. Tuatara 1, 20–23. open url image1

Fell H. B. (1952) Echinoderms from southern New Zealand. Zoology Publications from Victoria University College 18, 1–37. open url image1

Fell H. B. (1958) Starfishes of New Zealand. Tuatara 6, 127–142. open url image1

Fell H. B. (1960) Biological results of the Chatham Islands 1954 Chatham Islands 1954 Expedition. Part 2. Archibenthal and littoral echinoderms of the Chatham Islands. New Zealand Oceanographic Institute Memoirs 5, 55–75. open url image1

Fell H. B. ((1962a). ).’Nature in New Zealand: Native Sea Stars.’ (A. H. and A.W. Reed: Wellington.)

Fell H. B. (1962b) West-wind-drift dispersal of echinoderms in the Southern Hemisphere. Nature 193, 759–761.
Crossref | GoogleScholarGoogle Scholar | open url image1

Fenwick G. D., Horning D. S. (1980) Echinodermata of the Snares Islands, southern New Zealand. New Zealand Journal of Marine and Freshwater Research 14, 437–445. open url image1

Fisher W. K. (1911) Echinodermata of the Snares Islands, southern New Zealand.

Gemmill J. F. (1915) On the ciliation of asterids, and on the question of ciliary nutrition in certain species. Proceedings of the Zoological Society of London 1, 1–19. open url image1

Gill, J.B . (1998). ‘Powell’s Native Animals of New Zealand.’ 4th edn. (David Bateman: Auckland.)

Gosliner T. M., Behrens D. W., and Williams G. C. (1996). ‘Coral Reef Animals of the Indo-Pacific.’ (Sea Challengers Press: Monterey, CA.)

Gray J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias, Linnaeus). Annals and Magazine of Natural History 6, 175–184. open url image1

Gray J. E. (1847). Descriptions of some new genera and species of Asteriadae. Proceedings of the Zoological Society of London 1842, 72–82 (also in Ann. Mag. Nat. Hist. 20: 193–204).

Gray J. E. (1866). ‘Synopsis of the Species of Starfish in the British Museum.’ (British Museum: London.)

Halpern J. A. (1969) Biological investigations of the deep-sea. 50. The validity and generic position of Pentagonaster parvus Perrier (Echinodermata, Asteroidea). Proceedings. Biological Society of Washington 82, 403–506. open url image1

Halpern J. A. (1970) Goniasteridae (Echinodermata: Asteroidea) of the Straits of Florida. Bulletin of Marine Science 20, 193–286. open url image1

Hone D. W. E., Benton M. J. (2005) The evolution of large size: how does Cope’s Rule work? Trends in Ecology & Evolution 20, 4–6.
Crossref | GoogleScholarGoogle Scholar | open url image1

Hopkins T. S., Valentine J. F., McClintock J. B., Marion K., and Watts S. A. (1991). Community patterns of echinoderms associated with substrate and depth in the northern Gulf of Mexico. In ‘Biology of Echinodermata. Proceedings of the 7th International Echinoderm Conference, Atami, Japan, 9–14 September 1990’. (Eds T. Yanagisawa, I. Yasumasu, C. Oguro, N. Suzuki and T. Motokawa.) pp. 231–239. (Balkema: Rotterdam.)

Hrincevich A. D., Rocha-Olivares A., Foltz D. W. (2000) Phylogenetic analysis of molecular lineages in a species-rich subgenus of sea stars (Leptasterias subgenus Hexasterias). American Zoologist 40(3), 365–374. open url image1

Hunt G., Roy K. (2006) Climate change, body size evolution, and Cope’s Rule in deep-sea ostracodes. Proceedings of the National Academy of Sciences of the United States of America 103, 1347–1352.
Crossref | GoogleScholarGoogle Scholar | open url image1

Hutton F. W. (1872). ‘Catalogue of the Echinodermata of New Zealand, with Diagnoses of the Species.’ (James Hughes: Wellington.)

Jablonski D., and Bottjer D. J. (1988). Onshore-offshore evolutionary patterns in post-Paleozoic echinoderms: a preliminary analysis. In ‘Echinoderm Biology’. (Eds R. D. Burke and P. V. Mladenov.) pp. 81–90. (Balkema: Rotterdam.)

Jangoux M. (1984) Les asterides littoraux de Nouvelle-Caledonie. Bulletin du Museum National D’Histoire Naturelle (Zoologie) 6A(2), 279–293. open url image1

Jangoux M. (1986). Les astérides. In ‘Faune Tropicale. Vol. 25. Guide des Étoiles de Mer, Oursins et Autres Échinodermes du Lagon de Nouvelle-Caleodnie. A’. (Eds A. Guille et al.) pp. 111–153. (Office de la Recherche Scientifique et Technique Outre-Mer: Paris.)

Jangoux M., Aziz A. (1988) Les astérides (Echinodermata) récoltés autour de l’île de la Réunion par le N.O. Marion Dufresne en 1982. Bulletin du Museum National D’Histoire Naturelle (Zoologie) 10A(4), 631–650. open url image1

Jangoux M., deRidder C. (1987) Annotated catalogue of recent Echinoderm type specimens in the collection of the Rijksmuseum Van Natuurlijke Historie at Leiden. Zoologische Mededelingen 61, 79–96. open url image1

Johnson W. S. (1973) Respiration rates of some New Zealand echinoderms. New Zealand Journal of Marine and Freshwater Research 7(1&2), 165–169. open url image1

Keough M. J., Butler A. J. (1979) The role of asteroid predators in the organization of a sessile community on pier pilings. Marine Biology 51, 167–177.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kingsolver J. G., Pfennig D. W. (2004) Individual-level selection as a cause of Cope’s Rule of phyletic size increase. Evolution 58, 1608–1612. open url image1

Laguarda-Figueras A., Solis-Marin F. A., Duran-Gonzalez A., Ahearn C. G., Sanchez B. E., Torres-Vega J. (2005) Equinodermos (Echinodermataa) del Caribe Mexicano. Revista de Biologia Tropical suppl.3 53, 109–122. open url image1

Lawver L. A., Gahagan L. M. (2003) Evolution of Cenozoic seaways in the circum-Antarctic region.  Paleogeography Paleoclimatology Palaeoecology 198, 11–37.
Crossref | GoogleScholarGoogle Scholar | open url image1

Livingstone A. A. (1932) The Australian species of Tosia (Asteroidea). Records of the Australian Museum 18, 373–382. open url image1

Ludwig H. (1912) Über die J.E. Gray’schen Gattungen Pentagonaster und Tosia. Zoologischen Jahrbüchern 15, 1–44. open url image1

Lütken C. (1865) Kritiske Bemaerkniger om forsjellige Søstjerne (Asteriderne), med Beskrivelse af nogle nye Arter. Videnskabelige Meddelelsefra Dansk Naturhistorisk Forening 1864, 123–169. open url image1

Madsen F. J. (1961) The Porcellanasteridae: a monographic revision of an abyssal group of sea-stars. Galathea Report 4, 33–174. open url image1

Mah C. L. (2005a) A phylogeny of Iconaster and Glyphodiscus (Goniasteridae; Valvatida; Asteroidea) with descriptions of four new species. Zoosystema 27, 131–167. open url image1

Mah C. (2005b). Cladistic analysis of the Goniasteridae (Asteroidea: Valvatoidea): phylogeny, evolution and biodiversity. Ph.D. Thesis, University of Illinois, Chicago.

Mah C. L. (2006) Phylogeny and biogeography of the deep-sea goniasterid Circeaster (Echinodermata, Asteroidea, Goniasteridae) including descriptions of six new species. Zoosystema 28, 1–38. open url image1

Marsh L. M. (1976) Western Australian Asteroidea since H.L. Clark. Thalassia Jugoslavica 12, 213–225. open url image1

Marsh L. M. (1991). Shallow water echinoderms of the Albany region, south-western Australia. In ‘Proceedings of the Third International Marine Biological Workshop – The Marine Flora and Fauna of Albany, Western Australia, Vol. II’. (Eds F. E. Wells, D. I. Walker, H. Kirkman and R. Lethbridge.) pp. 439–482. (Western Australian Museum: Perth.)

Marsh L. M., Fromont J., Salotti M. (1999) A catalogue of Recent echinoderm type specimens in the Western Australian Museum, Perth. Records of the Western Australian Museum 19, 391–411. open url image1

McClain C. R., Rex M. A. (2001) The relationship between dissolved oxygen concentration and maximum size in deep-sea turrid gastropods: an application of quantile regression. Marine Biology 139, 681–685.
Crossref | GoogleScholarGoogle Scholar | open url image1

McClintock J. B., Watts S. A., Marion K. R., Hopkins T. S. (1995) Gonadal cycle, gametogenesis and energy allocation in two sympatric mid-shelf sea stars with contrasting modes of reproduction. Bulletin of Marine Science 57, 442–452. open url image1

McKnight D. G. (1967) Additions to the echinoderm fauna of the Chatham Rise. New Zealand Journal of Marine and Freshwater Research 1, 291–313. open url image1

Möbius K. (1859). Neue Seesterne des Hamburger und Ieler Museums. Abhandlungen und Verhandlungen. Naturwissenschaftlicher Verein in Hamburg 4(2).

Mortensen T. (1925) Echinoderms of New Zealand and the Auckland-Campbell Islands. 3–5: Asteroidea, Holothuroidea and Crinoidea. Zoogeographical remarks on the echinoderm fauna of New Zealand and the Auckland-Campbell Islands. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening I Kjøbenhavn 79, 261–420. open url image1

Mortensen T. (1933) Papers from Dr. Th. Mortensen’s Pacific Expedition 1914–26. 65. Echinoderms of South Africa (Asteroidea and Ophiuroidea). Videnskabelige Meddelelser fra Dansk naturhistorisk Forening I Kjøbenhavn 93, 215–399. open url image1

Müller J., and Troschel F. H. (1842). ‘System der Asteriden. 1. Asteriae. 2. Ophiuridae.’ (F. Vieweg und Sohn: Braunschweig.)

O’Hara T. (1998) Origin of Macquarie Island echinoderms. Polar Biology 20, 143–151.
Crossref | GoogleScholarGoogle Scholar | open url image1

O’Loughlin P. M. (1991). Brooding and fission in shallow water echinoderms of southern Australia. In ‘Biology of Echinodermata’. (Eds T. Yanagisawa, I. Yasumasu, C. Oguro, N. Suzuki and T. Motokawa.) pp. 223–228. (Balkema: Rotterdam.)

Perrier E. (1875). ‘Révision de la Collection de Stellérides du Museum d’Histoire Naturelle de Paris.’ (Reinwald: Paris.)

Perrier E. (1881) Description sommaire des espèces nouvelle d’Astérides. Bulletin of the Museum of Comparative Zoology Harvard 9, 1–31. open url image1

Perrier E. (1884) Mémoire sur les étoiles de mer recueillis dans la Mer des Antilles et la Golfe de Mexique. Nouvelles Archives du Muséum d’Histoire naturelle Paris 6, 127–276. open url image1

Perrier E. (1894) Stellérides. Expédition Scientifique Travailleur-Talismann 3, 1–431. open url image1

Rowe F. W. E., and Gates J. (1995). Echinodermata. In ‘Zoological Catalogue of Australia 33.’ (Ed. A. Wells.) pp. 63–72. (CSIRO Publishing: Melbourne.)

Saba M., Irimura S., and Soyama I. (2002). ‘Sea Stars and Brittle Stars in Japanese Waters.’ (TBS, Britannica Co.: Tokyo.) [In Japanese]

Shepherd S. A. (1968) The shallow water echinoderm fauna of South Australia. I. The Asteroids. Records of the South Australian Museum 15, 729–756. open url image1

Shick J. M., Edwards K. C., Dearborn J. H. (1981) Physiological ecology of the deposit-feeding sea star Ctenodiscus crispatus: Ciliated surfaces and animal-sediment interactions. Marine Ecology Progress Series 5, 165–184. open url image1

Sladen W. P. (1889) Asteroidea. Report of the Scientific Results of H.M.S. Challenger 30, 1–893. open url image1

Smith A. B. (1984). Echinoid paleobiology. In ‘Special Topics in Palaeontology’. (Eds C. T. Scrutton and C. P. Hughes.) (Allen & Unwin: London.)

Smith A. G., Smith D. G., and Funnell B. M. (1994). ‘Atlas of Mesozoic and Cenozoic Coastlines.’ (Cambridge University Press: Cambridge.)

Spencer W. K., and Wright C. W. (1966). Asterozoans, Part U: Echinodermata. In ‘Treatise on Invertebrate Paleontology, Vol. 3(1)’. (Ed. R. C. Moore.) pp. U4–U107. (University of Kansas Press: Lawrence.)

Spicer J. I., Gaston K. J. (1999) Amphipod gigantism dictated by oxygen availability? Ecology Letters [and reply] 2, 397–403.
Crossref | GoogleScholarGoogle Scholar | open url image1

Sumida P. Y. G., Tyler P. A., Billett D. S. M. (2001) Early juvenile development of deep-sea asteroids of the NE Atlantic Ocean, with notes on juvenile bathymetric distributions. Acta Zoologica 82, 11–40. open url image1

Swofford D. L. (2003). ‘PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods).’ Version 4. (Sinauer Associates: Sunderland, MA.)

Tenison-Woods J. E. (1878) A list of Australian starfishes. Transactions Proceedings and Report of the Philosophical Society of Adelaide South Australia 79, 89–93. open url image1

Tommasi L. R., de Oliveira E. (1976) Equinodermes do Brasil. V. Sobre algumas especies coletadas durante viagens do N/Oc. “Prof. W. Besnard Boletim Instituto de Oceanografia São Paulo 25, 77–100. open url image1

Town J. C. (1978) An annotated key to the echinoderms of the Kaikoura Peninsula. Mauri Ora 6, 47–55. open url image1

Verrill A. E. (1899) Revision of certain genera and species of starfishes, with descriptions of new forms. Transactions of the Connecticut Academy of Arts and Sciences 10, 145–234. open url image1

Villier L., and Kutscher M. (1998). How dissociated ossicles can further our understanding of the origins of the Neoasteroidea. An example from the Toarcian of Western Europe. In ‘Echinoderm Research’. (Eds C. Carnevali and M. Bonsaro.) pp. 417–422. (Balkema: Rotterdam.)

Villier L., David B., and Neraudeau D. (2001). Ontogenetic and morphological evolution of the ambulacral pores in Heteraster (early spatangoids). In ‘Echinoderms 2000’. (Ed. M. Barker.) pp. 563–567. (Swets & Zeitlinger: Lisse.)

Walenkamp J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen 147, 1–91. open url image1

Walenkamp J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana Shelf. Zoologische Verhandelingen 170, 1–97. open url image1

Walenkamp J. H. C. (1990) Systematics and zoogeography of Asteroidea (Echinodermata) from Inhaca I., Mozambique. Zoologische Verhandelingen 261, 1–86. open url image1

Waters J. M., Roy M. S. (2003a) Marine biogeography of southern Australia: phylogeographical structure in a temperate sea-star. Journal of Biogeography 30, 1787–1796.
Crossref | GoogleScholarGoogle Scholar | open url image1

Waters J. M., Roy M. S. (2003b) Global phylogeography of the fissiparous sea-star genus Coscinasterias. Marine Biology 142, 185–191. open url image1

Young M. W. (1929) Marine fauna of the Chatham Is. Transactions and Proceedings of the New Zealand Institute 60, 136–166. open url image1

Zeidler W., and Shepherd S. A. (1982). Sea-stars (class Asteroidea). In ‘Handbook of the Flora and Fauna of South Australia. Marine Invertebrates of Southern Australia. Pt. I’. (Eds S. A. Shepherd and I. M. Thomas.) pp. 400–418 (South Australian Government: Adelaide.)

Zitt J. (2005) The asteroid genus Haccourtaster (Echinodermata, Goniasteridae) in the Bohemian Cretaceous Basin, Czech Republic. Cretaceous Research 26, 225–237.
Crossref | GoogleScholarGoogle Scholar | open url image1










Appendix 1.  Data matrix
Click to zoom



Appendix 2.  Character list
Click to zoom