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.


Special thanks to Richard Emlet for providing specimens and larvae of Tosia australis for the study. Many thanks go also to Maria Byrne and Paula Cisternas (University of Sydney), Andrew Cabrinovic (British Museum of Natural History, London), Marc Eléaume (Muséum National d’Histoire Naturelle, Paris), Nick Kirby (Melbourne Aquarium), Carsten Lueter (Museum für Naturkunde, Berlin), David Macmillan and Joan Clark (University of Melbourne), Benjamin Ong (Museum Victoria) and Genefor Walker-Smyth (Tasmanian Museum and Art Gallery) for their invaluable contributions to this project. David Staples (Museum Victoria) provided the photographs used in Figs 1 and 8.


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Appendix 1. List of specimens sequenced for the current study

Haplotype codes are referred to on the phylogenetic trees (Figs 2, 3). Composite mitochondrial haplotype names shown on the mitochondrial tree consist of a numeral referring to the 16S haplotype and a letter referring to the COI haplotype, as shown in the table

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