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Pheno~Pic Plasticity in Marine Intertidal Gastropods DOCTOR OF Pheno~pic plasticity in marine intertidal gastropods By Ahmed Mohammed AI-Mazrouai A thesis submitted to the University of Plymouth in partial fulfilment for the degree of DOCTOR OF PHILOSOPHY School of Biological Sciences Faculty of Science November 2008 Ill Phenotypic plasticity in marine intertidal gastropods by Ahmed Mohammed AI- Mazrouai Abstract Phenotypic plasticity, the differential phenotypic expression of the same genotype in response to different environmental conditions, is a paradigm central of the study of evolution and ecology and is at the core of the "nature versus nurture" debate. Here, the marine gastropod Littorina littorea was used as a model to further our understanding of the potential role of phenotypic plasticity in intertidal systems. In the first study L. littorea was included in an investigation of induced defences across six species of intertidal marine gastropods in the families Littorinidae and Trochidae. Species differed in the magnitude and type of plastic response, which appeared to relate to their susceplihility Lo crab predation. Chapters three and four revealed that L. littorea was able to alter its degree of morphological plasticity depending on temporal variation in predation threat. Snails exposed to predation threat halfway through trials appeared to "catch up" snails continuously exposed to predator cues in, terms oftheir shell size, whereas snails experiencing a removal of predation cues showed a significant reduction in growth rate following this switch in predation. environment. A further investigation suggested that Littorina littorea demonstrated no significant difference in the morphological traits under variable predator threat versus a constant predator' threat environment. Finally, the interaction between biotic (predator) and abiotic (temperature) environmental effects revealed that snails maintained at 16 and 20° C demonstrated significant induced defences by growing larger and thicker shells, but there was no significant difference in induced defences between these two temperatures. However, the expression of induced defences was much lower at 24° C with only negative significant response in two of shell traits IV between control and predator cue treatments which may indicate that induced defences was inhibited at this temperature treatment. The implications of these results are discussed as is the potential applications of induced defences. V Contents Abstract ........................................................................................................................... iv Contents .......................................................................................................................... vi Figures ........................................................................................................................... viii Tables .............................................................................................................................. xi Acknowledgements ...................................................................................................... xlii Authors Declaration .................................................................................................... xiv Chapter 1 ......................................................................................................................... 1 Introduction ..................................................................................................................... l PHENOTYPIC PLASTICITY ....................................................................................... 2 General Background .................................................................................................... 2 Costs of and Limits to Plasticity ................................................................................. 3 Adaptive Value ofPhenotypic Plasticity .................................................................... 5 INDUCED DEFENCES ................................................................................................ 7 General Background .................................................................................................... 7 Ecological Consequences oflnduced Defences .......................................................... 8 Marine Gastropods as Models for Studying Induced Defences .................................. 9 THESIS AIMS ............................................................................................................. 11 Chapter 2 ................................................................................................................... 12 Chapter 3 ................................................................................................................... 13 Chapter 4 ................................................................................................................... 13 Chapter 5 ................................................................................................................... 13 Chapter 6 ................................................................................................................... 14 STUDY SPECIES AND FIELD COLLECTION SITES ............................................ 14 Common Periwinkle Littorina littorea (Linnaeus) ................................................... 14 Flat Periwinkle Littorina fabalis (Turton) ................................................................ 15 Flat Periwinkle Littorina obtusata (Linnaeus) .......................................................... 17 Thick Top Shell Osilinus linea la ( da Costa) ............................................................. 18 Grey Top Shell Gibbula cineraria (Linnaeus) .......................................................... 19 Flat Top Shell Gibbula umbilicalis (da Costa) ......................................................... 20 Predator Species ........................................................................................................... 21 Carcinus maenas (Linnaeus) .................................................................................... 21 Study Site ..................................................................................................................... 22 Chapter 2 ....................................................................................................................... 23 Predator-Induced Phenotypic Plasticity in Marine Gastropods: A Cross-Species Comparison ................................................................................................................... 23 SUMMARY ................................................................................................................. 24 lNTRODUCTION ....................................................................................................... 25 MATERIALS AND METHODS ................................................................................. 28 Study Organisms ....................................................................................................... 28 Experimental set up ................................................................................................... 29 Measuring Induced Defences .................................................................................... 29 Statistical Analysis ....................................................................................................... 31 RESULTS .................................................................................................................... 32 DISCUSSION .............................................................................................................. 36 Chapter 3 ..............................................•.........................................•........................•..... 41 Flexibility of Phenotypic Plasticity in Littorina littorea ............................................. 41 SUMMARY ................................................................................................................. 42 INTRODUCTION ....................................................................................................... 43 MATERIALS AND METHODS ................................................................................. 46 VI Study Organisms ....................................................................................................... 46 Experimental design and set up ................................................................................. 46 Measuring Induced Defences .................................................................................... 48 Statistical Analysis .................................................................................................... 48 RESULTS .................................................................................................................... 50 DISCUSSION .............................................................................................................. 57 Chapter 4 ....................................................................................................................... 61 The Effects of Variable Risk of Predation on Phenotypic Plasticity in the Marine Gastropod Littorina littorea .......................................................................................... 61 SUMMARY ................................................................................................................. 62 INTRODUCTION ......................................................................................................
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