ResearchOnline@JCU This file is part of the following reference: Fromont, Jane (1990) A taxonomic study of tropical marine sponges (Porifera: Demospongiae: Haplosclerida and Petrosida) using morphological, chemical and reproductive character sets. PhD thesis, James Cook University. Access to this file is available from: http://eprints.jcu.edu.au/27402/ If you believe that this work constitutes a copyright infringement, please contact [email protected] and quote http://eprints.jcu.edu.au/27402/ A TAXONOMIC STUDY OF TROPICAL MARINE SPONGES (PORIFERA: DEMOSPONGIAE: HAPLOSCLERIDA AND PETROSIDA) USING MORPHOLOGICAL CHEMICAL AND REPRODUCTIVE CHARACTER SETS Thesis submitted by Jane Fromont MSc (Auckland, NZ) in July 1990 for the Degree of Doctor of Philosophy in the Department of Marine Biology at James Cook University of North Queensland Volume One of Two Volumes Statement on Access to Thesis. I, the undersigned, the author of this thesis, understand that James . Cook University of North Queensland will make it available for use within the University Library and, by microfilm or other photographic means, allow access to users in other approved libraries. All users consulting this thesis will have to sign the following statement: "In consulting this thesis I agree not to copy or closely paraphrase it in whole or in part without the written consent of the author; and to make proper written acknowledgement for any assistance which I have obtained from it." Beyond this, I do not wish to place any restriction on access to this thesis. ature) (date) DECLARATION. I declare that this thesis is my own work and has not been submitted in any form for another degree or diploma at any university or other institution of tertiary education. Information derived from the published or unpublished work of others has been acknowledged in the text and a list of references is given. J. Fromont 26 July 1990 1 ACKNOWLEDGEMENTS. I thank my supervising committee particularly Professor P. Bergquist (Auckland University, New Zealand) for suggesting the topic and for critical comment and advice for the duration of the study. I thank Associate Professor C. Alexander (Dept. Marine Biology, J.C.U.) for constant advice and support, and comments on drafts of the thesis, and Dr. P. Murphy (Australian Institute of Marine Science) for guidance and comment on chemical aspects of the thesis. Special thanks to Associate Professor D. Hopley (Sir George Fisher Centre for Tropical Marine Studies) for generous provision of desk space and administrative support throughout the study. I also thank Dr. J. Baker (AIMS) for the initial organisation which facilitated this study. Many scientists and fellow students provided assistance during this project. Staff and students of the Chemistry Department, Stanford University, California, gave help and friendship during my stay there. I thank Dr. C. Djerassi for providing funding for the visit. I particularly thank S. Kerr for expert guidance in the laboratory, for chemical analysis and sterol characterisations, and R. Kerr for critical comment and advice on sterol chemistry. I thank the many dive buddies and field assistants who volunteered their time for this study especially Katrina Barratt, Anne Jackson, Kerrie Licastro, Yvette MacNeil and David Sutton. I am grateful to Orpheus Island Research Station personnel for use of the facilities, Zollie Florian for provision of microscopes and technical advice, Leigh Winsor for histological advice, and the Bioactivity Unit (AIMS) for some of the photographs used in the thesis. Bill Williams, Brian McArdle and Martin Riddle provided advice and assistance with numerical analyses. David Sutton suggested many helpful comments on drafts of the thesis, Cathy Robinson advised on word processing and typed the tables, the proof reading team: John de Klerk, Ian Dight, Rundi Larsen, James Scandol, Jill St.John, David Sutton and Moya Tomlinson found the missing commas. Thankyou. Finally, many thanks to staff and students of the Sir George Fisher Centre for their friendship and support. This thesis is dedicated to Muriel Fromont. ABSTRACT. This study was undertaken to determine whether the separation of sponge species (Phylum: Porifera, Class: Demospongiae, Subclass: Ceractinomorpha) into two orders, the Haplosclerida and Petrosida, was valid. To this end selected species from each order were collected from the Great Barrier Reef, Queensland, Australia. Their taxonomic position was compared using a number of character sets: morphological, chemical and reproductive. The most useful morphological characters for taxonomic purposes were organisation of the internal sponge skeleton, its components and their quantities. These characters allowed separation of the sponge species examined into five families, and also distinguished sponges at the species level. Development of the ectosomal skeleton was found to differentiate genera, and microscleres were useful for distinguishing some species. As a result of the morphological study four new species of Haplosclerida and three new species of Petrosida were described. Analyses of sterol chemistry did not support clear separation of sponge species into two orders but were found to be useful at the species level. Sponge species were found to have both a qualitative and quantitative sterol fingerprint. Sterol complements of species remained remarkably constant with geographic locality and season. This character set provided evidence for a species complex within Xestospongia muta. Three specimens of this species contained significantly different sterol complements, and two of the specimens were more similar to other species in the data set (Xestospongia n.sp.1 and X. testudinaria) than they were to each other. Results of this chemotaxonomic study suggest that most species currently classified in the same genus are generic complexes, as few species of the same genus had similar sterol complements. No cyclopropene or cyclopropane ring structures, previously thought to characterise species of the Petrosida, were found in the side chains of the fifty four sterols found from all species examined. The reproductive biology of six species, three classified in the Haplosclerida and three in the Petrosida, was monitored for two to four years. The reproductive mode of these species clearly separated them into two orders. Species of the Haplosclerida were found to brood larvae which is the normal reproductive mode of the subclass Ceractinomorpha. Species of the Petrosida did not conform to the normal ceractinomorph pattern. The three species examined (Xestospongia exigua, X. testudinaria and Xestospongia n.sp.1) had separate sexes and broadcast eggs and sperm. iii Reproductive characters also distinguished sponges at the species level. One species complex, in the Petrosida, was differentiated into two species (X. testudinaria and Xestospongia n.sp.1) when temporal isolation of spawning events was discovered. The spawning events of the two species were consistently separated by at least 15 days and diel timing of these events in both species was from 0700 hours. The mechanism, and possible environmental cues, whereby these species consistently separate their spawning events was not determined although X. testudinaria may have a semi-lunar periodicity. No lunar pattern was observed for Xestospongia n.sp.1. Larva of Xestospongia n.sp.1 were seen in aquaria three days after the spawning event in 1989. This is the first reported observation of larva from a species of the Petrosida. Two haplosclerid species found to brood female reproductive products did so in brood chambers situated basally in sponge individuals. This location may be a protective mechanism to avoid damage from turbulence or dehydration. Females of a third haplosclerid species, Haliclona symbiotica, had reproductive products aligned along the central axis of branches of adults. This location of sexual products is most 4-Iy a spatial confinement related to the symbiotic association this sponge maintains with the macroalga, Ceratodictyon spongiosum. All species of both orders examined that were reproductively active released sexual products in spring and summer. As a result of this study, one character set (reproductive mode) clearly supported the division of species into two orders. Sterol chemistry and morphological characters did not, but were useful at lower taxonomic levels. The distrepancies between higher taxonomic groupings, such as genus and family, suggests the taxonomy of these sponges is not currently resolved. TABLE OF CONTENTS: VOLUME ONE. Page no. Acknowledgements, Abstract. ii CHAPTER ONE. GENERAL INTRODUCTION. 1 CHAPTER TWO, SPECIES DESCRIPTIONS OF THE HAPLOSCLERIDA. 6 Introduction. 6 Methods. 7 Species Descriptions. 8 Order Haplosclerida Topsent 1928. 8 Family Haliclonidae de Laubenfels 1932. 8 Genus Haliclona Grant 1835. 10 Haliclona amboinensis (Levi 1961). 11 Haliclona symbiotica (Bergquist & Tizard 1967). 12 Genus Cladocroce Topsent 1892. 13 Cladocroce aculeata Pulitzer-Finali 1982.. 13 Family Niphatidae van Soest 1980 14 Genus Niphates Duchassaing & Michelotti 1864. 16 Niphates n.sp. 17 Genus Amphimedon Duchassaing & Michelotti,1864 18 Amphimedon viridis Duchassaing & Michelotti 1864. 18 Amphimedon n.sp.1 20 Amphimedco n.sp 2 21 Amphimedon n.sp 3 22 Genus Gelliodes Ridley 1884. 23 Gelliodes fibulata. (Carter 1881). 23 Genus Siphanodictyon Bergquist 1965 24 Siphonodictyon' mucosa. Bergquist 1965. 26 Siphonodictyon coralllphagum Rfitzler 1971 (forma typica) 27 Family Callyspongiidae de