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University of Southampton Research Repository Eprints Soton University of Southampton Research Repository ePrints Soton Copyright © and Moral Rights for this thesis are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder/s. The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holders. When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given e.g. AUTHOR (year of submission) "Full thesis title", University of Southampton, name of the University School or Department, PhD Thesis, pagination http://eprints.soton.ac.uk UNIVERSITY OF SOUTHAMPTON FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES Ocean and Earth Science Volume 1 of 1 Life-history biology and biogeography of invertebrates in deep-sea chemosynthetic environments by Verity Nye Thesis for the degree of Doctor of Philosophy December 2013 UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES Ocean and Earth Science Thesis for the degree of Doctor of Philosophy LIFE-HISTORY BIOLOGY AND BIOGEOGRAPHY OF INVERTEBRATES IN DEEP-SEA CHEMOSYNTHETIC ENVIRONMENTS Verity Nye Globally-distributed, insular and ephemeral deep-sea hydrothermal vents with their endemic faunas provide ‘natural laboratories’ for studying the processes that shape global patterns of marine life. The continuing discovery of hydrothermal vents and their faunal assemblages has yielded hundreds of new species and revealed several biogeographic provinces, distinguished by differences in the taxonomic composition of their assemblages. However, the first-order question of how these provinces are separated remains unanswered. The recent discovery of the Beebe (~4600 m depth) and Von Damm (~2300 m depth) vent fields at the Mid-Cayman Spreading Centre and their faunas has provided a critical opportunity to further our understanding of the biodiversity, life-history biology and biogeography of vent species. Here I present Rimicaris hybisae sp. nov. (Caridea: Alvinocarididae), Iheyaspira bathycodon sp. nov. (Turbinidae: Skeneinae), and Lebbeus virentova sp. nov. (Caridea: Hippolytidae) from the Beebe and Von Damm vent fields. I elucidate the general reproductive features of the dominant species at Beebe and Von Damm (R. hybisae) and reveal a high degree of spatial variability in the population structure and reproductive features of this species at both these vent fields. I demonstrate inter-annual variation in the population structure and reproductive development of R. hybisae, superimposed upon the pattern of spatial variation, and hypothesise periodic or seasonal reproductive development in this species. Cluster analysis and the presence of Rimicaris-dominated faunal assemblages at the Beebe and Von Damm vent fields support higher-level taxonomic affinities with the fauna of Mid-Atlantic vents. These findings reveal previously unappreciated spatial variation in the reproductive development of a motile species at hydrothermal vents and expand our knowledge of the distribution of biodiversity. This work advances our understanding of biogeographic patterns of deep-sea chemosynthetic ecosystems, and provides a foundation to test the influence of various processes on the distribution of vent fauna at global scale. i Contents ABSTRACT......................................................................................................................i Contents...........................................................................................................................iii List of tables......................................................................................................................v List of figures..................................................................................................................vii Declaration of authorship.................................................................................................xi Acknowledgements.........................................................................................................xii 1. Introduction...................................................................................................................1 2. A new species of Rimicaris (Crustacea: Decapoda: Caridea: Alvinocarididae) from hydrothermal-vent fields on the Mid-Cayman Spreading Centre, Caribbean............43 3. Iheyaspira bathycodon new species (Vetigastropoda: Trochoidea: Turbinidae: Skeneinae) from the Von Damm Vent Field, Mid-Cayman Spreading Centre, Caribbean....................................................................................................................87 4. A new species of Lebbeus (Crustacea: Decapoda: Caridea: Hippolytidae) from the Von Damm Vent Field, Caribbean Sea.....................................................................109 5. Spatial variation in the population structure and reproductive biology of Rimicaris hybisae (Caridea: Alvinocarididae) at hydrothermal vents on the Mid-Cayman Spreading Centre.......................................................................................................135 6. Fine-scale spatial variation and time-series of reproductive development in Rimicaris hybisae (Caridea: Alvinocarididae) at hydrothermal vents on the Mid-Cayman Spreading Centre.......................................................................................................173 7. Summary and conclusions.........................................................................................213 Appendix 1: Phyletic overview of reproduction and developmental modes of invertebrates in deep-sea chemosynthetic environments..........................................237 iii iv List of tables 1.1 Estimated ages of groups of taxa living in contemporary chemosynthetic environments............................................................................................................14 2.1. Summary of known geographic distribution, bathymetric range and habitat of alvinocaridid shrimp species……………………………………………………....45 2.2. Morphological variation in Rimicaris hybisae sp. nov., Mid-Cayman Spreading Centre. ……………………………………………………………………………..57 3.1. Turbinid gastropods described from extant hydrothermal vents and cold seeps…..89 3.2. Morphological variation in Iheyaspira bathycodon sp. nov……………………… 91 4.1. Summary of geographical distribution and bathymetric range of Lebbeus species from hydrothermal vents………………………………………………………….111 5.1. Rimicaris hybisae. Sample and population data for 959 specimens used in this study………………………………………………………………………………139 5.2. Average minimum realised fecundity and embryo sizes of caridean shrimp from hydrothermal vents and cold seeps……………………………………………….143 5.3. Rimicaris hybisae. Spatial variation in population structure between the Beebe and Von Damm vent fields, January 2012…………………………………………….144 5.4. Rimicaris hybisae. Spatial variation in population structure within the Beebe Vent Field, January 2012……………………………………………………………….144 5.5. Rimicaris hybisae. Variation in body size (carapace length), January 2012……..144 6.1. Rimicaris hybisae. Sample and population data for 1012 specimens.....................178 6.2. Rimicaris hybisae. Variation in body size (carapace length), February 2013........182 6.3. Rimicaris hybisae. Spatial variation in population structure between the Beebe and Von Damm vent fields, February 2013..................................................................182 6.4. Rimicaris hybisae. Spatial variation in population structure within the Beebe Vent Field, February 2013...............................................................................................186 6.5. Rimicaris hybisae. Spatial variation in population structure within the Von Damm Vent Field, February 2013......................................................................................186 v vi List of figures 1.1. Global distribution of hydrothermal vent, cold seep and whale fall sites that have been studied with respect to their fauna..................................................................7 1.2. The geological, biological, chemical and physical history of the deep sea...........13 1.3. Location of the Mid-Cayman Spreading Centre (MCSC) and hydrothermal vent fields......................................................................................................................18 2.1. Rimicaris hybisae sp. nov., in situ at the Beebe and Von Damm vent fields........48 2.2. Rimicaris hybisae sp. nov., live specimen, from the Beebe Vent Field................54 2.3. Rimicaris hybisae sp. nov., holotype female from the Von Damm Vent Field: entire animal, carapace and cephalic appendages..................................................56 2.4. Rimicaris hybisae sp. nov., holotype female from the Von Damm Vent Field: abdominal somites, telson and uropods.................................................................58 2.5. Rimicaris hybisae sp. nov., holotype female from the Von Damm Vent Field: antennal appendages and mouthparts....................................................................60 2.6. Rimicaris hybisae sp. nov., male, from the Beebe Vent Field: SEM micrographs of maxilla and first pereopod.................................................................................62 2.7. Rimicaris hybisae
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