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Seasonal Life-History Adaptation in the Water

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Seasonal Life-History Adaptation in the Water SEASONAL LIFE -HISTORY ADAPTATION IN THE WATER STRIDER GERRIS LACUSTRIS DISSERTATION ZUR ERLANGUNG DES NATURWISSENSCHAFTLICHEN DOKTORGRADES DER BAYERISCHEN JULIUS -MAXIMILIANS -UNIVERSITÄT WÜRZBURG VORGELEGT VON BRENDA PFENNING AUS MARKTHEIDENFELD WÜRZBURG 2008 Eingereicht am: 11. Februar 2008 Mitglieder der Prüfungskommission: Vorsitzender: Prof. Dr. Martin J. Müller Erstgutachter: Prof. Dr. Hans Joachim Poethke Zweitgutachter: Prof. Dr. Jürgen Tautz Tag des Promotionskolloquiums: .............................................. Doktorurkunde ausgehändigt am: ............................................ Table of Contents Chapter 1 General Introduction 1 1.1 LIFE -HISTORY ADAPTATIONS TO VARIATIONS IN SEASON LENGTH ....... 4 1.2 ADAPTATIONS OF WATER STRIDERS TO SEASONALITY .............................. 9 1.3 OUTLINE OF THE THESIS .................................................................................. 12 Chapter 2 Species and Study Area 15 2.1 THE COMMON POND -SKATER GERRIS LACUSTRIS ...................................... 17 2.2 STUDY AREA ....................................................................................................... 20 Chapter 3 Variability in the life history of the water strider Gerris lacustris (Heteroptera: Gerridae) across small spatial scales 23 3.1 INTRODUCTION ................................................................................................. 26 3.2 METHODS ........................................................................................................... 28 Wing length pattern and life-cycles of reference populations and reared 3.2.1 29 individuals ............................................................................................................ 3.2.2 Wing length pattern of field- and forest-populations ........................................... 30 3.2.3 Population dynamics and voltinism of field- and forest- populations .................. 31 3.3 RESULTS .............................................................................................................. 31 Wing length pattern and life-cycles of reference populations and reared 3.3.1 32 individuals ............................................................................................................ 3.3.2 Wing length pattern of field- and forest-populations ........................................... 33 3.3.3 Population dynamics and voltinism of field- and forest- populations .................. 33 3.4 DISCUSSION ........................................................................................................ 35 Chapter 4 Alternative life histories in the water strider Gerris lacustris : 39 temporal constraint on wing morph and voltinism 4.1 INTRODUCTION ................................................................................................ 42 4.2 MATERIAL AND METHODS ................................................................ ............. 43 4.2.1 Effects of temperature on developmental time of eggs and larvae ......................... 43 4.2.2 Estimation of lower developmental threshold t 0 and thermal constant k .............. 45 Temporal constraint on voltinism and wing dimorphism on field- and forest- 4.2.3 46 ponds ................................................................................................ ..................... 4.3 RESULTS ................................................................................................ .............. 47 4.3.1 Developmental rate of eggs and larvae ................................................................ .. 47 4.3.2 Estimation of lower developmental threshold t 0 and thermal constant k .............. 48 Temporal constraint on voltinism and wing dimorphism on field- and forest- 4.3.3 49 ponds ................................................................................................ ..................... 4.4 DISCUSSION ................................................................................................ ........ 51 Chapter 5 Dealing with time constraints on development: the effect of 55 food availability 5.1 INTRODUCTION ................................................................................................ 58 5.2 MATERIAL AND METHODS ................................................................ ............. 59 5.2.1 Rearing experiment ................................................................ ............................... 59 5.2.2 Effect of feeding regime and hatching time on development ................................ 60 5.2.3 Mortality and wing phenology ................................................................ ............. 61 5.3 RESULTS ................................................................................................ .............. 61 5.3.1 Effect of feeding regime and hatching time on development ................................ 61 5.3.2 Mortality and wing phenology ................................................................ ............. 63 5.4 DISCUSSION ................................................................................................ ........ 64 CHAPTER 6 The annual reproductive pattern of the water strider Gerris lacustris in variable environments 67 6.1 INTRODUCTION ................................................................................................ 70 6.2 METHODS ................................................................................................ ............ 71 6.2.1 Diapause strategy of Gerris lacustris ................................................................ .... 71 6.2.2 Inter-individual variability of development ................................ .......................... 72 Fitness model: Calculating fitness for diapause and directly reproduction 6.2.3 strategies ................................................................................................ ............... 73 6.3 RESULTS ................................................................................................ .............. 74 6.4 DISCUSSION ................................................................................................ ........ 78 6.5 APPENDIX 1: FITNESS CALCULATION ................................ ............................ 80 6.6 APPENDIX 2: PARAMETER ESTIMATION ................................ ........................ 82 6.6.1 The environment ................................................................................................ ... 82 6.6.2 The species ................................................................................................ ............. 83 Summary 87 Zusammenfassung 93 References 99 Publications 111 Conferences & Workshops 112 Curriculum vitae 113 Danksagung 115 Erklärung 117 Chapter 1 General Introduction Chapter 1 – General Introduction 3 Chapter 1 General Introduction The components of an individual’s life that are closely related to fitness – primarily juvenile and adult survival, development rate, age and size at maturity, age-specific fecundity, offspring size and number, longevity and senescence – are called life-history traits. Due to their linkage to an individual’s fitness, selective pressure operates on these traits (Roff, 1992; Stearns, 1992). It is the purpose of life-history theory to explain how an individual’s life-history strategy, i.e. the combination and sequence of traits it shows during its life, are shaped by natural selection (Nylin & Gotthardt, 1998) and consequently why we find such a vast variation of patterns of growth, survival and reproduction in nature. Life-history theory suggests that an individual’s fitness is maximised in relation to the combination of extrinsic and intrinsic factors that act upon its life. The extrinsic factors are the ecological impacts that vary in space and time and force individuals to adjust their life-histories appropriately. The intrinsic factors are ‘trade-offs among live-history traits and lineage-specific constraints on the expression of genetic variation’ , which limit this adjustment (Stearns, 2000). Thus, to explain a life-history strategy observed in an organism, the critical environmental parameters that influence its traits and the major constraints and trade-offs that limit the variation of traits have to be identified. The environmental influences individuals in natural habitats are exposed to, as well as the theories on life-history adaptations to environmental factors are diverse (Roff, 2002); therefore it is necessary to narrow the subject matter down to the theories that are most meaningful for the type of organism under investigation. This thesis focuses on the life-history strategy adaptations of an annual insects species (with the potential to switch from uni- to multivoltine phenology) living in temperate latitudes. The major constraint these organisms are faced with is the problem of fitting their life-history appropriately to a seasonal environment (Levins, 1969; Tauber et al., 1986; Danks, 1994). The seasonal character of the environment forces poikilothermic organisms (like insects) to pass the climatically adverse part of the season in an appropriately adapted state of diapause. Reproduction, growth and development up to the species-specific diapause state thus have to be completed within the limited period of favourable conditions (Cohen, 1970; Dingle, 1978). The primary problems these organisms have to deal with are (i) to optimally adjust development time to the given season length with respect to the trade-offs connected with development (Roff, 1980)
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