Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

A new brooding species of the biscuit star Tosia (Echinodermata : Asteroidea : Goniasteridae), distinguished by molecular, morphological and larval characters

Kate M. Naughton A B C and Timothy D. O’Hara A

A Sciences Department, Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia.

B Department of Zoology, University of Melbourne, Vic. 3010, Australia.

C Corresponding author. Email:

Invertebrate Systematics 23(4) 348-366
Submitted: 8 May 2008  Accepted: 13 July 2009   Published: 30 October 2009


The biscuit star Tosia australis Gray, 1840 is a well known component of the shallow rocky reef fauna of south-eastern Australia. The putative T. australis species complex was subjected to reproductive, morphometric and molecular analyses. Molecular analyses of the data from three markers (mitochondrial COI and 16S rRNA and the nuclear non-coding region ITS2) confirmed the presence of a cryptic species, the morphology of which does not agree with any of the existing nominal species. Two separate reproductive modes were observed within the complex and documented via scanning electron microscopy. T. neossia, sp. nov., described herein from south-eastern Australia, is shown to release gametes from gonopores on the actinal surface. Embryos develop first into non-feeding, non-swimming brachiolaria, and then into tripod brachiolaria before metamorphosis. No surface cilia are present at any point throughout development of T. neossia. T. australis sensu stricto is shown to release gametes from the abactinal surface. Embryos develop into non-feeding, swimming brachiolaria before metamorphosis. Whereas T. australis var. astrologorum is confirmed as synonymous with T. australis, the status of the putative Western Australian taxon T. nobilis remains unresolved.

Additional keywords: Bayesian phylogenetic analysis, brooding, cilia, colouration, larval morphology, life-history evolution, mitochondrial DNA, nuclear DNA.


Atwood D. G. 1973 Larval development in the asteroid Echinaster echinophorus. Biological Bulletin 144 1 11 doi:10.2307/1540143

Bennett I. Pope E. C. 1960 Intertidal zonation of the exposed rocky shores of Tasmania and the relationship with the rest of Australia. Australian Journal of Marine and Freshwater Research 11 182 219 doi:10.1071/MF9600182

Birkeland C. Chia F.-S. Strathmann R. R. 1971 Development, substratum selection, delay of metamorphosis and growth in the seastar, Mediaster aequalis Stimpson. Biological Bulletin 141 99 108 doi:10.2307/1539994

Bosch I. 1989 Contrasting modes of reproduction in two Antarctic asteroids of the genus Porania, with a description of unusual feeding and non-feeding larval types. Biological Bulletin 177 77 82 doi:10.2307/1541836

Byrne M. 1992 Reproduction of sympatric populations of Patiriella gunnii, P. calcar and P. exigua in New South Wales, astrinid seastars with direct development. Marine Biology 114 297 316 doi:10.1007/BF00349533

Byrne M. 1995 Changes in larval morphology in the evolution of benthic development by Patiriella exigua (Asteroidea: Asterinidae), a comparison with the larvae of Patiriella species with planktonic development. Biological Bulletin 188 293 305 doi:10.2307/1542306

Byrne M. 2005 Viviparity in the sea star Cryptasterina hystera (Asterinidae) – conserved and modified features in reproduction and development. Biological Bulletin 208 81 91 doi:10.2307/3593116

Byrne M. 2006 Life history diversity and evolution in the Asterinidae. Integrative and Comparative Biology 46 243 254 doi:10.1093/icb/icj033

Byrne M. Cerra A. 2000 Lipid dynamics in the embryos of Patiriella species (Asteroidea) with divergent modes of development. Development, Growth & Differentiation 42 79 86 doi:10.1046/j.1440-169x.2000.00486.x

Byrne M. , Cerra A. , Hart M. W. , and Smith M. J. (1999). Life history diversity and molecular phylogeny in the Australian sea star genus Patiriella. In ‘The Other 99%. The Conservation and Biodiversity of Invertebrates’. (Eds W. Ponder and D. Lunney.) pp. 188–196. (Transactions of the Royal Zoological Society of New South Wales: Sydney.)

Byrne M. Hart M. W. Cerra A. Cisternas P. 2003 Reproduction and larval morphology of broadcasting and viviparous species in the Cryptasterina species complex. Biological Bulletin 205 285 294 doi:10.2307/1543292

Chia F.-S. 1968 The embryology of a brooding starfish, Leptasterias hexactis (Stimpson). Acta Zoologica 49 321 364

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 1 396 411 plates 42–46.

Clark H. L. 1928 The sea-lilies, sea-stars, brittle stars and sea-urchins of the South Australian Museum. Records of the South Australian Museum 3 361 482 figs 108–142.

Clarke K. N. , and Gorley R. N. (2006). ‘Primer-E v.6. User Manual/Tutorial.’ (Primer-E Ltd: Plymouth, UK.)

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

Coleman N. (2007). ‘Sea-stars: Echinoderms of the Asia/Indo-Pacific.’ (Neville Coleman’s Underwater Geographic: Springwood, Qld.)

Drummond A. J. Rambaut A. 2007 BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology 7 214 221 doi:10.1186/1471-2148-7-214

Edgar G. J. (2000). ‘Australian Marine Life: The Plants and Animals of Temperate Waters.’ 1st edn, revised. (Reed New Holland: Sydney.)

Emlet R. B. 1994 Body form and patterns of ciliation in nonfeeding larvae of echinoderms: functional solutions to swimming in the plankton? American Zoologist 34 570 585

Gosselin P. Jangoux M. 1998 From competent larva to exotrophic juvenile: a morphofunctional study of the perimetamorphic period of Paracentrotus lividus (Echinodermata: Echinoidea). Zoomorphology 118 31 43 doi:10.1007/s004350050054

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

Grosberg R. K. , Levitan D. R. , and Cameron B. B. (1996). Simple extraction and RAPD–PCR protocols. In ‘Molecular Zoology: Advances, Strategies, and Protocols’. (Eds J. D. Ferraris and S. R. Palumbi.) pp. 470–473. (Wiley-Liss: New York.)

Haesaerts D. Jangoux M. Flammang P. 2006 Adaptations to benthic development: functional morphology of the attachment complex of the brachiolaria larva in the sea star Asterina gibbosa. Biological Bulletin 211 172 182 doi:10.2307/4134591

Hall T. 1998 BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41 95 98

Hamel J.-F. Mercier A. 1995 Prespawning behavior, spawning, and development of the brooding starfish, Leptasterias polaris. Biological Bulletin 188 32 45 doi:10.2307/1542065

Hart M. W. Byrne M. Smith M. J. 1997 Molecular phylogenetic analysis of life-history evolution in asterinid starfish. Evolution 51 1848 1861 doi:10.2307/2411007

Hart M. W. Byrne M. Johnson S. L. 2003 Patiriella pseudoexigua (Asteroidea: Asterinidae): a cryptic species complex revealed by molecular and embryological analyses. Journal of the Marine Biological Association of the United Kingdom 83 1109 1116 doi:10.1017/S002531540300835Xh

Hart M. W. Johnson S. L. Addison J. A. Byrne M. 2004 Strong character incongruence and character choice in phylogeny of sea stars of the Asterinidae. Invertebrate Biology 123 343 356

Hart M. W. Keever C. S. Dartnall A. J. Byrne M. 2006 Morphological and genetic variation indicate cryptic species within Lamarck’s little sea star, Parvulastra (=Patiriella) exigua. Biological Bulletin 210 158 167 doi:10.2307/4134604

Himmelman J. H. Lavergne Y. Cardinal A. Martel G. Jalbert P. 1982 Brooding behaviour of the Northern Sea Star Leptasterias polaris. Marine Biology 68 235 240 doi:10.1007/BF00409590

Hyman L. H. (1955). ‘Echinodermata: The Coelomate Bilateria.’ (McGraw-Hill: New York.)

Knowles L. L. Carstens B. C. 2007 Delimiting species without monophyletic gene trees. Systematic Biology 56 887 895 doi:10.1080/10635150701701091

Knowlton N. 1993 Sibling species in the sea. Annual Review of Ecology and Systematics 24 189 216 doi:10.1146/

Knowlton N. 2000 Molecular genetic analyses of species boundaries in the sea. Hydrobiologia 420 73 90 doi:10.1023/A:1003933603879

Komatsu M. Kano Y. T. Yoshizawa H. Akabane S. Oguro C. 1979 Reproduction and development of the hermaphroditic sea-star, Asterina minor Hayashi. Biological Bulletin 157 258 274 doi:10.2307/1541053

Kumar S. Tamura K. Jakobsen B. Nei M. 2001 MEGA2: molecular evolutionary genetics analysis software. Bioinformatics 17 1244 1245 doi:10.1093/bioinformatics/17.12.1244

Lane D. J. W. , and Hu J. M. L. (1994). Abbreviated development in Iconaster longimanus (Möbius): planktonic lecithotrophy in a tropical goniasterid sea star. In ‘Echinoderms through Time’. (Eds B. David, A. Guille, J.-P. Féral and M. Roux.) pp. 343–346. (A.A. Balkema: Rotterdam.)

Livingstone A. A. 1932 The Australian species of Tosia (Asteroidea). Records of the Australian Museum 18 373 382 plates 43–44.

Mah C. 2005 A phylogeny of Iconaster and Glyphodiscus (Echinodermata, Asteroida, Valvata, Goniasteridae) with descriptions of four new species. Zoosystema 27 137 161

Mah C. 2007 Systematics, phylogeny and historical biogeography of the Pentagonaster clade (Asteroidea: Valvatida: Goniasteridae). Invertebrate Systematics 21 311 339 doi:10.1071/IS06049

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’. (Eds F. E. Wells, D. I. Walker, H. Kirkman and R. C. Lethbridge.) pp. 439–482. (Western Australian Museum: Perth.)

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

McEdward L. R. Janies D. A. 1997 Relationships among development, ecology and morphology in the evolution of echinoderm larvae and life cycles. Biological Journal of the Linnean Society. Linnean Society of London 60 381 400 doi:10.1111/j.1095-8312.1997.tb01502.x

McEdward L. R. Miner B. G. 2001 Larval and life-cycle patterns in echinoderms. Canadian Journal of Zoology 79 1125 1170 doi:10.1139/cjz-79-7-1125

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

Müller J. Troschel F. H. 1843 Neue Beitrage zur Kenntnis der Asteriden. Archiv für naturgeschichte 9 113 131

O’Hara T. D. , Byrne M. , and Cisternas P. (2004). The Ophiocoma erinaceus complex: another case of cryptic speciation in echinoderms. In ‘Echinoderms: München’. (Eds T. Heinzeller and J. H. Nebelsick.) pp. 537–542. (A.A. Balkema: Rotterdam.)

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. (A.A. Balkema: Rotterdam.)

O’Loughlin P. M. Waters J. M. 2004 A molecular and morphological revision of genera of Asterinidae (Echinodermata: Asteroidea). Memoirs of Museum Victoria 61 1 40

Palumbi S. R. , Martin A. , Romano S. , McMillan W. O. , Stice L. , and Grabowski G. (1996). ‘The Simple Fool’s Guide to PCR.’ (Kewalo Marine Laboratory, University of Hawaii: Honolulu, HI.)

Pernet B. 2003 Persistent ancestral feeding structures in nonfeeding annelid larvae. Biological Bulletin 205 295 307 doi:10.2307/1543293

Perrier E. 1869 Recherches sur les Pédicellaires et les Ambulacres des Astéries et des Oursins. Annales des Sciences Naturelles 12 197 304

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

Posada S. Crandall K. A. 1998 Modeltest: testing the model of DNA substitution. Bioinformatics 14 817 818 doi:10.1093/bioinformatics/14.9.817

Prowse T. A. A. Sewell M. A. Byrne M. 2007 Fuels for development: evolution of maternal provisioning in asterinid sea stars. Marine Biology 153 337 349 doi:10.1007/s00227-007-0809-7

Ronquist F. Huelsenbeck J. P. 2003 MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 1572 1574 doi:10.1093/bioinformatics/btg180

Rowe F. W. E. , and Gates J. (1995). ‘Echinodermata.’ (CSIRO: Melbourne.)

Scheibling R. E. Lawrence J. M. 1982 Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico. Marine Biology 70 51 62 doi:10.1007/BF00397296

Shepherd S. A. 1968 The shallow water echinoderm fauna of South Australia: Part 1. The asteroids. Records of the South Australian Museum 15 729 756

Strathmann R. R. (1988). Larvae, phylogeny and von Baer’s Law. In ‘Echinoderm Phylogeny and Evolutionary Biology’. (Eds C. R. C. Paul and A. B. Smith.) pp. 53–68. (Clarendon Press: Oxford, USA.)

Thompson J. D. Higgins D. G. Gibson T. J. 1994 CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22 4673 4680 doi:10.1093/nar/22.22.4673

Tominaga H. , Komatsu M. , and Oguro C. (1994). Aggregation for spawning in the breeding season of the sea-star, Asterina minor Hayashi. In ‘Echinoderms through Time’. (Eds B. David, A. Guille, J.-P. Féral and M. Roux.) pp. 369–373. (A.A. Balkema: Rotterdam.)

Tyler P. A. Pain S. L. 1982 Observations of gametogenesis in the deep-sea asteroids Paragonaster subtilis and Pseudarchaster parelii (Phanerozonia, Goniasteridae). International Journal of Invertebrate Reproduction 5 269 272

Tyler P. A. , Pain S. L. , and Gage J. D. (1982). Gametogenic cycles in deep-sea phanerozoan asteroida from the N.E. Atlantic. In ‘Echinoderms: Proceedings of the International Conference, Tampa Bay’. (Ed. J. M. Lawrence.) pp. 431–434. (A.A. Balkema: Rotterdam.)

Villinski J. T. Villinski J. T. Byrne M. Raff R. A. 2002 Convergent maternal provisioning and life history evolution in echinoderms. Evolution 56 1764 1775

Waters J. M. Roy M. S. 2003 Marine biogeography of southern Australia: phylogeographical structure in a temperate sea-star. Journal of Biogeography 30 1787 1796 doi:10.1046/j.0305-0270.2003.00978.x

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