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Copyright: Anna Syme, 2007 A Systematic Revision of the Cylindroleberididae (Crustacea: Ostracoda: Myodocopa) Anna Syme B.Sc. (Hons.) Submitted in total fulfilment of the requirements of the degree of Doctor of Philosophy, February 2007. Zoology Department The University of Melbourne Printed on archival quality paper i Copyright: Anna Syme, 2007 ABSTRACT Ostracods of the family Cylindroleberididae are speciose, widespread, and well- defined by morphological features including the possession of gills. Despite the family having been divided into three subfamilies, five tribes, 32 genera, and more than 200 species, the relationships between its species have never been analysed phylogenetically. Twenty-five of the 32 genera are not defined by unique features but rather combinations of characters, and may therefore be historical constructions rather than evolutionary groups. The classification is of limited functional and predictive use, and requires a systematic revision. This work resolves several outstanding taxonomic issues. A checklist of the 219 described species is presented, including comments on taxonomic uncertainty. One of the key points of uncertainty concerns the missing specimen upon which the type species Cypridina mariae and the type genus of the family (Cylindroleberis) is based. A neotype is therefore described which clarifies taxonomically important morphological characters, particularly of the first antenna and mandible. To facilitate identification of cylindroleberidid species in ecological surveys and biodiversity estimates, an interactive key is produced, applicable to species in all geographic areas. Three new cylindroleberidid species are described from coastal Australian waters. A cladistic analysis is undertaken in order to test current subfamily and generic concepts against phylogenetic criteria. Using Bayesian likelihood methods, a phylogeny is inferred from both morphological and molecular data. The morphological data come from 141 species and 66 characters. The molecular data are based on predominantly new sequences of both nuclear (28S) and mitochondrial (16S) DNA, for 22 species. The results of the phylogenetic analyses of both morphological and molecular data strongly support the monophyly of the Cylindroleberididae; however, the two phylogenies differ in the topology of subfamily relationships. Where the morphology- based phylogeny suggests that Asteropteroninae and Cyclasteropinae are sister taxa, and then sister to Cylindroleberidinae, the molecular-based phylogeny places Asteropteroninae within the paraphyletic Cylindroleberidinae. The position of the Cyclasteropinae is uncertain. There is much homoplasy in the morphological characters, iii Copyright: Anna Syme, 2007 and the molecular phylogeny is considered more accurate in its representation of relationships within the family. Character mapping on both the morphology- and molecular-based phylogenies indicates that most existing generic diagnostic character states do not uniquely define clades and are likely to be of limited use in any phylogenetically-based classification. However, one character of use is the posterior ridge on the inner carapace which defines two genus level clades. Two new characters are identified of taxonomic use: a lateral seta on the first antenna, and the shape of the posterior of the body. A known fossil cylindroleberidid is dated at 425 million years old; this date was used to calibrate divergence times between clades in the molecular phylogeny. A genus- level clade in the phylogeny is calibrated to 110 million years old. In the light of this temporal framework, it is not surprising that morphological characters show high levels of homoplasy, representing potential evolutionary convergence. Based on the resolution of taxonomic issues and phylogenetic results, six genera are revised. Cylindroleberis is re-diagnosed based on the neotype Cypridina mariae. The emended diagnosis results in the synonymy of the monotypic Polyleberis with Cylindroleberis. Synasterope is re-diagnosed based on the important morphological character of the posterior ridge, and Postasterope and Heptonema become synonymized with it. Although the monophyly of Parasterope is not demonstrated, it is retained as a broadly-defined genus and its diagnosis expanded to accommodate all former species of Synasterope that lack the posterior ridge. Sections of this thesis have been published in Syme and Poore (2006a; 2006b) and are incorporated in a manuscript in review (Syme and Poore in review). iv Copyright: Anna Syme, 2007 DECLARATION This is to certify that: i. the thesis comprises only my original work towards the PhD, ii. due acknowledgement has been made in the text to all other material used, iii. the thesis is less than 100,000 words in length, exclusive of tables, maps, bibliographies and appendices. Signed: v Copyright: Anna Syme, 2007 ACKNOWLEDGMENTS I thank my principal PhD supervisor, Dr Gary Poore (Museum Victoria), for support and advice, particularly regarding crustacean taxonomy, curation, and systematics. For assistance with applications and thesis revision I thank Dr David Macmillan (The University of Melbourne). Discussion and assistance was provided by staff, students and associates of Museum Victoria, particularly the Marine Invertebrates Department. In particular, I wish to thank Robin Wilson, Joanne Taylor, Paula Cisternas and Tim O’Hara. Part of this research was conducted at the laboratory of Todd Oakley (University of California, Santa Barbara), and I am grateful for the assistance, advice and use of laboratory equipment and samples, and help from other laboratory members. For discussion about the morphology and taxonomy of ostracods, I thank Louis Kornicker (Smithsonian Museum, Washington D.C.) and for assistance there I thank Elizabeth Harrison-Nelson. Andrew Parker (then at the Australian Museum, Sydney) provided advice regarding ostracod dissection. I am grateful to Linda and Andrew Syme who assisted with fieldwork. I thank Josh Mackie for discussion of molecular protocols and systematic methods, Ian Seymour for discussion regarding evolutionary theory and mathematics, and my family and friends for their support. This work was supported by an Australian Postgraduate Award (2003-2006). For participation in laboratory research in the USA in 2004 I am grateful for awards from The University of Melbourne (Postgraduate Overseas Research Experience Scholarship) and the Zoology Department (Alfred Nichols Fellowship). Attendance at the 2006 Workshop on Molecular Evolution (USA) was supported by the Australian Biological Resources Study (Student Travel Bursary), Museum Victoria (1854 Student Scholarship) and Australian Marine Sciences Association (Victorian Student Prize). I thank these organisations for their financial assistance. vii Copyright: Anna Syme, 2007 TABLE OF CONTENTS CHAPTER 1: RESEARCH OUTLINE........................................................ 1 1.1 Research problem.......................................................................... 1 Systematics............................................................................ 1 Ostracoda ............................................................................. 1 Cylindroleberididae.............................................................. 2 Current problems in systematics .......................................... 2 1.2 Research aim: systematic revision................................................. 3 Resolution of existing taxonomic issues ............................... 3 Phylogenetic analysis ........................................................... 3 1.3 Research approach......................................................................... 4 Critical review of current knowledge ................................... 4 Taxonomic and phylogenetic analysis.................................. 4 Discussion and revision........................................................ 4 CHAPTER 2: A REVIEW OF THE CYLINDROLEBERIDIDAE .......... 7 2.1 Introduction ................................................................................... 7 2.2 Biology .......................................................................................... 7 Morphology .......................................................................... 7 Genetics .............................................................................. 13 Ecology ............................................................................... 13 Phylogeny and evolution .................................................... 15 2.3 Classification ............................................................................... 17 Higher relationships........................................................... 17 History of the classification................................................ 19 Accepted classification ....................................................... 22 Available genera................................................................. 24 2.4 Current systematic problems ....................................................... 31 Outstanding taxonomic issues ............................................ 32 Limited investigation of phylogeny and evolution.............. 32 Problematic classification .................................................. 33 2.5 Conclusion................................................................................... 34 CHAPTER 3: A REVIEW OF SYSTEMATIC METHODOLOGY....... 35 3.1 Introduction ................................................................................
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