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Evolutionary Patterns and Consequences of Developmental Mode in Cenozoic Gastropods from Southeastern Australia

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Evolutionary patterns and consequences of developmental mode in Cenozoic gastropods from southeastern Australia Thesis submitted in accordance with the requirements of the University of Liverpool for the degree of Doctor in Philosophy by Kirstie Rae Thomson September 2013 ABSTRACT Gastropods, like many other marine invertebrates undergo a two-stage life cycle. As the adult body plan results in narrow environmental tolerances and restricted mobility, the optimum opportunity for dispersal occurs during the initial larval phase. Dispersal is considered to be a major influence on the evolutionary trends of different larval strategies. Three larval strategies are recognised in this research: planktotrophy, lecithotrophy and direct development. Planktotrophic larvae are able to feed and swim in the plankton resulting in the greatest dispersal potential. Lecithotrophic larvae have a reduced planktic period and are considered to have more restricted dispersal. The planktic period is absent in direct developing larvae and therefore dispersal potential in these taxa is extremely limited. Each of these larval strategies can be confidently inferred from the shells of fossil gastropods and the evolutionary trends associated with modes of development can be examined using both phylogenetic and non-phylogenetic techniques. This research uses Cenozoic gastropods from southeastern Australia to examine evolutionary trends associated with larval mode. To ensure the species used in analyses are distinct and correctly assigned, a taxonomic review of the six families included in this study was undertaken. The families included in this study were the Volutidae, Nassariidae, Raphitomidae, Borsoniidae, Mangeliidae and Turridae. Phylogenetic analyses were used to examine the relationships between taxa and to determine the order and timing of changes in larval mode throughout the Cenozoic. Traditionally, planktotrophy has been considered the ancestral mode of development. However, using maximum parsimony and maximum-likelihood analysis, this research suggests that the ancestral developmental mode cannot be confidently determined in gastropods from southeastern Australia. Similarly, evidence that transitions between larval strategies might be reversible contradicts the general view that regaining the specialised structures associated with planktotrophy is so difficult that it is considered extremely unlikely to occur. When the timing of switches in larval mode was examined they were found to be scattered at different points in time rather than clustered to specific periods and therefore no inference can be made as to the likely factors driving transitions between larval modes. The correlation between mode of development and macroevolutionary trends was examined using non-phylogenetic techniques. The results do not concur with the hypothesis that species with planktotrophic larvae will exhibit wider geographic ranges, longer species durations and lower speciation rates then lecithotrophic or direct developing taxa. The iii analyses are thought to be hindered by a strong preservation bias and gaps within the fossil record. The quality of the fossil record and the congruence between phylogenies and stratigraphy is examined using the Stratigraphic Consistency Index, the Relative Completeness Index and the Gap Excess Ratio. iv ACKNOWLEDGEMENTS So many people have offered their support and guidance over the last four years and I hope that those people I have not had the chance to mention here know how incredibly grateful I am. Firstly, I thank my primary supervisor, Charlotte, for all her help, advice and encouragement. She has always provided me with guidance and reassurance when I needed it most and I simply couldn’t have finished this thesis without her. It has been great fun working with you Charlotte, and I will miss our meetings enormously! I would also like to thank my secondary supervisor, Jim, for always making time for me and for sharing his expertise. Thanks also go to the following people who made me so welcome during my visits to their institutions and who allowed me access to such wonderful collections: Mary-Anne Binnie and Ben McHenry at the South Australian Museum in Adelaide; Tom Darragh, Rolf Schmidt and David Holloway at the Museum Victoria in Melbourne; and Jon Todd, Consuelo Sendino and Martin Munt at the Natural History Museum, London. I am also very grateful to Chris Ah Yee and Janice Krause for arranging visits to less accessible or lesser known localities and for making my stay in Hamilton such a pleasant one and to Chris Goudey for allowing me to join him in the field and for showing me his truly amazing fossil collection, which was so very inspiring. My time in Liverpool has been so much fun and that is all due to the wonderful staff and students. Thank you in particular to Tash, Paddy, Johnald, Jutley, Vickee and Gemma – you are all so amazing and the last four years wouldn’t have been nearly so good without you. A very special thank you goes to my former office-mate, Lis, who has been such an incredibly good friend to me. Coffee and yumyums will forever bring you to mind! A massive thank you goes to Gem who has supported me in so many ways over the last four years and whose friendship has reduced the likelihood of total insanity considerably. A massive thank you also goes to my friends from elsewhere and in particular to Rachael, Bekah and Ellie. Thanks for always being there for me and for your endless support. To my friends in Adelaide and Melbourne, thank you for making long visits so far away from home so much fun and thank you for all your hospitality. v My family have been so supportive over the last four years and I am so grateful for their unconditional love and their endless guidance. To my brother, Iain, thank you for being the best brother in the world! To Fiona, who is more like a sister than a cousin, I cannot thank you enough for all your support and to my Aunty Jaki, thank you for always believing in me. To Ed, I cannot thank you enough for all you have done for me throughout this journey. You have put up with a lot of crazy and you have made sure that when I come home every night it is normal and calm. That means so much to me. Thank you for all your love and support and your constant faith in me. I hope I can repay the favour one day. Finally, I would like to thank my parents. Mum and Dad, you are a constant source of inspiration to work hard, take (calculated) risks and to grab every opportunity that comes my way. Thank you for always encouraging me to attempt something new, for showing me the world and for all your love and support. Your belief in me and my abilities has kept me going even during the toughest moments. Thank you for everything. vi CONTENTS ABSTRACT iii ACKNOWLEDGEMENTS v LIST OF FIGURES xi LIST OF TABLES xv LIST OF PLATES xviii CHAPTER 1: 1 INTRODUCTION 1 1.1 LARVAL STRATEGIES 2 1.1.1 PLANKTOTROPHY 3 1.1.2 NONPLANKTOTROPHY 4 1.1.3 CLASSIFICATION SCHEMES 6 1.2 MACROEVOLUTIONARY CONSEQUENCES OF LARVAL STRATEGIES 9 1.2.1 GEOGRAPHIC DISTRIBUTION 9 1.2.2 SPECIES LONGEVITY 12 1.2.3 SPECIATION RATES 14 1.3 EVOLUTION OF LARVAL MODE 17 1.4 FACTORS DRIVING SWITCHES IN LARVAL STRATEGY 22 1.5 AIMS AND OBJECTIVES 25 CHAPTER 2: 2 GEOLOGICAL SETTING 29 2.1 SOUTH AUSTRALIA 31 2.1.1 ST VINCENT BASIN 31 2.1.2 MURRAY BASIN 36 2.2 VICTORIA 41 2.2.1 OTWAY BASIN 41 2.2.2 TORQUAY BASIN 50 2.2.3 PORT PHILLIP BASIN 53 2.2.4 GIPPSLAND BASIN 56 2.3 STRATIGRAPHIC CORRELATION 59 2.4 OTHER FORMATIONS 59 vii CHAPTER 3: 3 TAXONOMY OF CENOZOIC GASTROPODS FROM SOUTHEASTERN AUSTRALIA 63 3.1 FAMILY VOLUTIDAE 65 3.1.1 SYSTEMATIC PALAEONTOLOGY 66 3.2 FAMILY NASSARIIDAE 142 3.2.1 SYSTEMATIC PALAEONTOLOGY 143 3.3 FAMILY RAPHITOMIDAE 158 3.3.1 SYSTEMATIC PALAEONTOLOGY 158 3.4 FAMILY BORSONIIDAE 165 3.4.1 SYSTEMATIC PALAEONTOLOGY 165 3.5 FAMILY MANGELIIDAE 184 3.5.1 SYSTEMATIC PALAEONTOLOGY 184 3.6 FAMILY TURRIDAE 194 3.6.1 SYSTEMATIC PALAEONTOLOGY 195 CHAPTER 4: 4 PHYLOGENETIC ANALYSES OF CENOZOIC GASTROPODS FROM SOUTHEASTERN AUSTRALIA 207 4.1 METHODS 209 4.2 FAMILY VOLUTIDAE 211 4.2.1 SUBFAMILY ATHLETINAE 211 4.2.2 SUBFAMILY VOLUTINAE 214 4.2.3 SUBFAMILY AMORIINAE 216 4.2.4 SUBFAMILY ZIDONINAE 218 4.2.5 FAMILY VOLUTIDAE COMPOSITE TREE 220 4.3 FAMILY NASSARIIDAE 224 4.4 FAMILY RAPHITOMIDAE 227 4.5 FAMILY BORSONIIDAE 228 4.6 FAMILY MANGELIIDAE 230 4.7 FAMILY TURRIDAE 232 4.8 SUPERFAMILY CONOIDEA 234 4.9 NEOGASTROPOD COMPOSITE TREE 239 viii CHAPTER 5: 5 ORDER AND TIMING OF SWITCHES IN LARVAL MODE 243 5.1 INFERRING LARVAL MODE FROM FOSSIL GASTROPODS 244 5.2 RECONSTRUCTION OF ANCESTRAL DEVELOPMENTAL MODE 254 5.2.1 MAXIMUM PARSIMONY ANALYSIS 254 5.2.2 MAXIMUM-LIKELIHOOD ANALYSIS 261 5.3 NUMBER AND ORDER OF CHANGES IN DEVELOPMENTAL MODE 265 5.3.1 MAXIMUM PARSIMONY ANALYSIS 265 5.3.2 MAXIMUM PARSIMONY SENSITIVITY ANALYSIS 268 5.3.3 MAXIMUM-LIKELIHOOD ANALYSIS 270 5.4 CONGRUENCE OF PHYLOGENY AND STRATIGRAPHY 270 5.5 TIMING OF CHANGES IN DEVELOPMENTAL MODE 276 CHAPTER 6: 6 MACROEVOLUTIONARY CONSEQUENCES OF LARVAL STRATEGIES 279 6.1 MATERIALS AND METHODS 282 6.1.1 GEOGRAPHIC DISTRIBUTION 282 6.1.2 SPECIES LONGEVITY 283 6.1.3 SPECIATION RATES 283 6.1.4 SPECIATION EVENTS 283 6.1.5 COMPARATIVE ANALYSIS 284 6.2 RESULTS 285 6.2.1 GEOGRAPHIC DISTRIBUTION 285 6.2.2 SPECIES LONGEVITY 289 6.2.3 SPECIATION RATES 292 6.2.4 SPECIATION EVENTS 296 6.3 SUMMARY 303 CHAPTER 7: 7 CONCLUSIONS AND FUTURE WORK 305 7.1 TAXONOMY 305 7.2 PHYLOGENETIC ANALYSES 307 7.3 ORDER AND TIMING OF CHANGES IN LARVAL MODE 308 7.4 MACROEVOLUTIONARY CONSEQUENCES OF LARVAL STRATEGIES 310 ix REFERENCES 313 APPENDIX 1 – LOCALITY DATA 355 APPENDIX 2 – TIME SCALE 393 APPENDIX 3 – SPECIES DATA 397 x LIST OF FIGURES Figure 1.1 Idealised metazoan life cycle.
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    Auckland Shell Club Auction Lot List - 24 October 2015 Albany Hall. Setup from 9am. Viewing from 10am. Auction starts at noon. Lot Type Reserve 1 WW Many SMALL CYPRAEIDAE including the rare Rosaria caputdraconis from Easter Is. Mauritian scurra from Somalia, Cypraea eburnea white from from, New Caledonia, Cypraea chinensis from Solomon Is Lyncina sulcidentata from Hawaii and heaps more. 2 WW Many CONIDAE including rare Conus queenslandis (not perfect!) Conus teramachii, beautiful Conus trigonis, Conus ammiralis, all from Australia, Conus aulicus, Conus circumcisus, Conus gubernator, Conus generalis, Conus bullatus, Conus distans, and many more. 3 WW BIVALVES: Many specials including Large Pearl Oyster Pinctada margaritifera, Chlamys sowerbyi, Glycymeris gigantea, Macrocallista nimbosa, Pecten glaber, Amusiium pleuronectes, Pecten pullium, Zygochlamys delicatula, and heaps more. 4 WW VOLUTIDAE: Rare Teramachia johnsoni, Rare Cymbiolacca thatcheri, Livonia roadnightae, Zidona dufresnei, Lyria kurodai, Cymbiola rutila, Cymbium olia, Pulchra woolacottae, Cymbiola pulchra peristicta, Athleta studeri, Amoria undulata, Cymbiola nivosa. 5 WW MIXTURE Rare Campanile symbolium, Livonia roadnightae, Chlamys australis, Distorsio anus, Bulluta bullata, Penion maximus, Matra incompta, Conus imperialis, Ancilla glabrata, Strombus aurisdianae, Fusinus brasiliensis, Columbarium harrisae, Mauritia mauritana, and heaps and heaps more! 6 WW CYPRAEIDAE: 12 stunning shells including Trona stercoraria, Cypraea cervus, Makuritia eglantrine f. grisouridens, Cypraea
  • Iberus : Revista De La Sociedad Española De Malacología

    Iberus : Revista De La Sociedad Española De Malacología

    © SociedadEspañola de Malacología Iberus,2A (1): 13-21,2006 Taxonomic notes on some Indo -Pacific and West African Drilliaspecies (Conoidea: Drilliidae) Notas taxonómicas sobre unas especies de Drillia del Indo -Pacifico y África occidental (Conoidea: Drilliidae) Donn L. TIPPETT* Recibido ell-XII-2003. Aceptado el' 7-III-2006 ABSTRACT Two uncommon turrid species, Drillia dunkeri (Weinkauff, 1 876) and Drillia enna (Dalí, 1918), have been reviewed on the basis of examination of type material, material discovered in the USNM collection or procured by the author, and study of the literature. The review permits clarification of their taxonomic status, localities, and ranges. Athird, common, species, Drillia regia (Habe and Murakami, 1970), confused originally with D. enna, is similarly clarified. Confused with Drillia dunkeri, Drillia umbilicata Gray, 1 838, type species of the genus, and a species resembling it closely, Drillia patriciae Bernard, 1 984 are also reviewed. RESUMEN Drillia dunkeri (Weinkauff, 1876) y Drillia enna (Dalí, 1918), dos especies de tórridos poco comunes, se revisan en base al material tipo, de colecciones en el USNM o procu- rado por el autor, y el estudio de la literatura. La revisión permite clarificar el estado taxo- nómico, localidades y de rango. Igualmente se aclara la taxonomía de una tercera espe- cie común, D. regia (Habe y Murakami, 1970), originalmente confunidida con D. enna. Debido a la confusión con D. dunkeri, también se revisa D. umbilicata Gray, 1838, la especie tipo del género, y una especie parecida, D. patriciae Bernard, 1984. KEY WORDS: Drillia, taxonomy. PALABRAS CLAVE: Drillia, taxonomía. INTRODUCTION Drillia dunkeri (Weinkauff, 1876), as the USNM collection reveáis 3 further Pleurotoma {Clavus) dunkeri, was de- specimens that are here identified as D.