Buccinum Undatum (Linnaeus 1758) By
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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 Physiological thresholds through early ontogeny: the effects of temperature and hydrostatic pressure on the common whelk Buccinum undatum (Linnaeus 1758) by Kathryn Elizabeth Smith Thesis for the degree of Doctor of Philosophy January 2013 ‘It is not the strongest of the species that survives, nor the most intelligent. It is the one that is the most adaptable to change’ - Clarence Darrow (1857-1938) Physiological thresholds during development in Buccinum undatum Abstract UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES Ocean and Earth Sciences Doctor of Philosophy PHYSIOLOGICAL THRESHOLDS THROUGH EARLY ONTOGENY: THE EFFECTS OF TEMPERATURE AND HYDROSTATIC PRESSURE ON THE COMMON WHELK BUCCINUM UNDATUM (LINNAEUS 1758) By Kathryn Elizabeth Smith The eco-physiological thresholds controlling the distribution of marine invertebrates are of significance in understanding the evolution of marine diversity. This includes the direction of species radiation throughout the oceans. Range expansions occur as a result of evolutionary adaptations, or through environmentally or anthropogenically driven shifts in distribution. The success of such events is centred around a species ability to adapt; in order for a migration to be successful, all life history stages must tolerate the conditions of the new habitat. This thesis examines the thermal and hyperbaric thresholds affecting range extension in the marine environment. It focuses on the larval development of the shallow-water North Atlantic gastropod Buccinum undatum (Linnaeus 1758). The Buccinidae family consists of a wide range of shallow and deep- water species distributed globally. Improved knowledge on this topic will contribute to our understanding of the adaptations influencing both historical and modern shifts in the distribution of species. The thermal and hyperbaric ranges observed during development indicate B. undatum to have the capacity to develop at both temperatures and pressures outside its current distribution. Thermal acclimation to low temperature was also found to increase pressure tolerance during development. A shift in number of embryos developing and nurse egg partitioning per embryo indicate a decrease in developmental success at temperatures above those it is naturally exposed to. An increase in energetic expenditure, with both increasing temperature and pressure, relates to a rise in the metabolic cost associated with development under either condition. These results, combined with the known life history of B. undatum, suggest range expansion into deep water may be a plausible scenario, but tolerance of warmer conditions remains questionable due to the cold-induced spawning observed in this species. The results of this thesis support theories of high-latitude migrations into the deep sea via cold, isothermal waters, and indeed, suggest polar temperatures may promote the rate at which such range expansions occur. Additionally, the observed metabolic cost associated with development suggests hydrostatic pressure may induce bathymetric limits, which explain patterns observed in reproductive trends, eco-physiological adaptations, and faunal distribution in marine invertebrates throughout the oceans. Kathryn E. Smith 1 Physiological thresholds during development in Buccinum undatum Contents CONTENTS Abstract ........................................................................................................................... 1 Contents .......................................................................................................................... 2 List of tables .................................................................................................................... 5 List of figures .................................................................................................................. 6 Appendices ...................................................................................................................... 8 Author declaration ......................................................................................................... 9 Acknowledgements....................................................................................................... 10 CHAPTER 1. INTRODUCTION ............................................................................... 13 1.1. Evolution and colonization of the oceans: a historical perspective ....... 14 1.2. Species range limits and modern range expansions ................................ 20 1.3. Modern species distributions: significant variables and global patterns 23 1.4. Means of range expansion: the importance of larval development, maternal investment, and dispersal ........................................................... 28 1.5. The dominant factors affecting species range shifts: temperature and hydrostatic pressure .................................................................................. 34 1.6. Neogastropoda ........................................................................................ 39 1.7. Rationale, hypotheses and objectives ..................................................... 45 CHAPTER 2. EXPERIMENTAL DESIGN, METHODOLOGY AND EQUIPMENT ........................................................................................................ 47 2.1. Experimental design ............................................................................... 48 2.2. Sample collection ................................................................................... 51 2.3. Sample maintenance and egg mass use .................................................. 59 2.4. Egg mass variability ............................................................................... 60 2.5. Measurements and capsule dissection .................................................... 62 2.6. Temperature manipulation ..................................................................... 66 2.7. Hydrostatic pressure manipulation ......................................................... 66 2.8. Experimental protocols ........................................................................... 71 2.9. Data representation analysis and statistics ............................................. 75 2.10. Import legislations .................................................................................. 76 2 Kathryn E. Smith Physiological thresholds during development in Buccinum undatum Contents CHAPTER 3. INTRACAPSULAR DEVELOPMENT IN THE COMMON WHELK BUCCINUM UNDATUM (LINNAEUS 1758) .................................... 79 3.1. Abstract................................................................................................... 80 3.2. Introduction ............................................................................................ 80 3.3. Aims and objectives ............................................................................... 82 3.4. Materials and methods ............................................................................ 83 3.5. Results .................................................................................................... 86 3.6. Discussion............................................................................................... 94 CHAPTER 4. THERMAL TOLERANCE DURING EARLY ONTOGENY IN THE COMMON WHELK BUCCINUM UNDATUM (LINNAEUS 1758) .... 103 4.1. Abstract................................................................................................. 104 4.2. Introduction .......................................................................................... 104 4.3. Aims and objectives ............................................................................ 106 4.4. Materials and methods .......................................................................... 107 4.5. Results .................................................................................................. 109 4.6. Discussion............................................................................................. 117 CHAPTER 5. THE SUBTLE INTRACAPSULAR SURVIVAL OF THE FITTEST: MATERNAL INVESTMENT, SIBLING CONFLICT OR ENVIRONMENTAL EFFECTS? ..................................................................... 123 5.1. Abstract................................................................................................. 124 5.2. Introduction .......................................................................................... 124 5.3. Aims and objectives ............................................................................. 128 5.4. Materials and methods .........................................................................