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Life History Evolution in the Parasitoid Hymenoptera Ruth Elizabeth Traynor

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Life History Evolution in the Parasitoid Hymenoptera Ruth Elizabeth Traynor Life history evolution in the parasitoid Hymenoptera Ruth Elizabeth Traynor This thesis was submitted for the degree of Doctor of Philosophy University of York Department of Biology 2004 Abstract This thesis addresses life history evolution of the parasitoid Hymenoptera. It aims to identify assumptions that should be incorporated into parasitoid life history theory and the predictions that theory should aim to make. Both two species and multi-species comparative studies as well as up-to-date phylogenetic information are employed to investigate these issues. Anecdotal observations suggest that solitary parasitoids have narrower host ranges than closely related gregarious species. There are several possible reasons for this; for example gregarious species may be able to exploit larger bodied hosts because they can fully consume the host, which may be essential for successful pupation to occur. Comparative laboratory experiments between two closely related species of Aphaereta, one of which is solitary and the other gregarious, show no difference in the extent of host range. This study does, however, suggest that differences in the realized niche that each species occupies in the field may result from life history differences between the species. These differences may themselves have arisen due to solitary or gregarious development. The first multi-species study in the thesis uses a data set, compiled for the parasitic Hymenoptera by Blackburn (1990), to address factors that may influence body size and clutch size. This study builds on previous analyses of the data (see Blackburn 1990, 1991a/b, Mayhew & Blackburn 1999) through the use of up-to-date phylogenetic information. Evidence is found that the host stage attacked by a parasitoid is associated with both body and clutch size, due to the amount of resources available for the developing parasitoids. In addition, gregarious species found at high latitudes have a reduced clutch size relative to those found at low latitudes. Several cross-species associations, which are not evolutionarily correlated, are identified: larger wasps lay smaller clutches; when attacking the same host stage, koinobionts are larger than idiobionts; temperate species are larger than tropical species (Bergmann's rule). This study supports some theoretical models and hypotheses based on other empirical studies. A second multi-species study is carried out using a novel data set and up-to-date phylogenetic information for the Ichneumonoidea. Evidence supporting some aspects of the dichotomous hypothesis is found; for example, ectoparasitoids I idiobionts live longer than endoparasitoids I koinobionts and endoparasitoids are more fecund than ectoparasitoids. There is a trade-off between parasitoid body size and brood size, and also between fecundity and egg volume. Body size is positively correlated with development time, adult lifespan, and egg size. Host body size is positively correlated with parasitoid body size and brood size. Gregarious wasps are smaller, but attack larger hosts than solitary species and the former are more associated with external rather than internal pupation sites. Temperate parasitoids have larger geographic ranges, longer preadult lifespans and attack more host species than tropical parasitoids. Positive relationships are identified between parasitoid geographic range and a) host geographic range and b) the number of host species attacked. All of these results illustrate that several biological transitions are important regulators of life history variation within the Ichneumonoidea. The evolutionary lability of Ichneumonoidea traits is then investigated. Influential life history traits, such as ecto- I endoparasitism, idio- I koinobiosis, body size, and solitary I gregarious development, are all conserved traits. Less conserved traits include longevity, pre­ adult lifespan, geographic range, host niche and host stage attacked. The majority of variation 2 amongst traits was found at the family or subfamily level, suggesting that ancient evolutionary events are responsible for the majority of modern phenotypic diversity. 3 List of contents Chapter 1: Introduction to life history evolution 1.1 Introduction (page 13) 1.1.1 Life history evolution (page 13) 1.1.2 Multi-trait life history models (page 15) 1.2 Ecological niche evolution (page 16) 1.2.1 Defining a niche (page 16) 1.2.2 Niche evolution (page 17) 1.3 Empirical approaches in evolutionary biology (page 18) 1.3.1 General (page 18) 1.3.2 Cross-species studies (page 19) 1.4 Taxonomy and phylogeny (page 20) 1.5 Comparative analyses used in the thesis (page 22) 1.5.1 Cross-species analysis (page 22) 1.5.2 Independent contrast methods (page 22) 1.5.3 Measures of phylogenetic lability (page 23) 1.6 Introduction to parasitoids (page 25) 1.6.1 Taxonomy (page 25) 1.6.2 Biology (page 25) 1.7 Parasitoid life history evolution (page 26) 1.7.1 Introduction (page 26) 1.7.2 The 'dichotomous hypothesis' (page 26) 1.7.3 Other trait associations (page 30) 1.7.4 Evolution of the parasitoid niche (page 32) 1.8 Conclusion (page 34) Chapter 2: Host range in solitary and gregarious parasitoids: a laboratory experiment 2.1 Abstract (page 36) 2.2 Introduction (page 36) 2.3 Materials and Methods (page 37) 2.3.1 Cultures (page 37) 2.3.2 Host range experiments (page 39) 2.3.3 Statistical Analysis (page 40) 2.4 Results (page 40) 2.4.1 Proportion of pupae from which wasps emerged (page 40) 2.4.2 Number of parasitoid offspring 'produced (page 43) 2.5 Discussion (page 44) 2.5.1 Main findings (page 44) 2.5.2 Current findings (page 44) 2.5.3 Analytical issues (page 46) 2.5.4 Conclusions (page 47) 4 Chapter 3: A comparative study of body size and clutch size across the parasitoid Hymenoptera 3.1 Abstract (page 48) 3.2 Introduction (page 48) 3.3 Methods (page 50) 3.3.1 Data (page 50) 3.3.2 Analysis (page 51) 3.3.3 Phylogenetic assumptions (page 52) 3.4 Results (page 52) 3.4.1 Body size, clutch size and host stage attacked (page 52) 3.4.2 Body size, clutch size and development mode (page 56) 3.4.3 Body size, clutch size and latitude (page 58) 3.5 Discussion (page 59) 3.5.1 General findings (page 59) 3.5.2 Body size, clutch size and host stage attacked (page 60) 3.5.3 Body size, clutch size and development mode (page 61) 3.5.4 Body size, clutch size and latitude (page 62) 3.5.5 Analytical issues (page 62) 3.5.6 Conclusion (page 63) Chapter 4: A comparative analysis of life history evolution across the Ichneumonoidea (Hymenoptera) 4.1 Abstract (page 64) 4.2 Introduction (page 65) 4.3 Methods (page 67) 4.3.1 Data (page 67) 4.3.2 Analysis (page 70) 4.3.3 Phylogenetic assumptions (page 71) 4.4 Results (page 72) 4.4.1 General findings (page 72) 4.4.2 The dichotomous hypothesis (page 74) 4.4.3 Trade-offs and allometries (page 81) 4.4.4 Parasitoid body size and brood size, and host body size (page 85) 4.4.5 Parasitoid geographic range and geographic distribution (page 88) 4.4.6 Number of host species attacked (page 90) 4.5 Discussion (page 95) 4.5.1 Main findings (page 95) 4.5.2 Dichotomous hypothesis (page 95) 4.5.3 Trade-offs and allometries (page 97) 4.5.4 Parasitoid body size and brood size and host body size (page 98) 4.5.5 Parasitoid geographic range and geographic distribution (page 99) 4.5.6 Number of host species attacked (page 100) 4.5.7 Analytical issues (page 100) 5 4.5.8 Conclusions (page 102) Chapter 5: The evolutionary lability of life history traits in in the Ichneumonoidea (Hymenoptera) 5.1 Abstract (page 103) 5.2 Introduction (page 103) 5.3 Methods (page 106) 5.3.1 Data (page 106) 5.3.2 Analysis (page 109) 5.4 Results (page 111) 5.4.1 Comparing metrics (page 111) 5.4.2 Trait lability (page 114) 5.5 Discussion (page 117) 5.5.1 General findings (page 117) 5.5.2 Analytical issues (page 120) 5.5.3 Conclusions (page 121) Chapter 6: Conclusions and future prospects 6.1 Introduction (page 122) 6.2 Host use in solitary versus gregarious parasitoids (page 122) 6.3 The evolution of body size and clutch size across the parasitoid Hymenoptera (page 124) 6.4 Associations between life history traits across the Ichneumonoidea (page 127) 6.5 The lability and rate of evolution of traits in the Ichneumonoidea (page 130) 6.6 General conclusions (page 131) Appendices Appendix 1: Parasitic Hymenoptera life history data set (page 133) Appendix 2: Traditional taxonomy for the parasitoid Hymenoptera (page 146) Appendix 3: Part 1. Conservative c1adogram for the parasitic Hymenoptera (page 161) Appendix 3: Part 2. Ichneumonidae - conservative c1adogram for the parasitic Hymenoptera (page 162) Appendix 3: Part 3. Braconidae - conservative cladogram for the parasitic Hymenoptera (page 163) Appendix 3: Part 4a. Chalcidoidea - conservative cladogram for the parasitic Hymenoptera (page 164) Appendix 3: Part 4b. Chalcidoidea - conservative cladogram for the parasitic Hymenoptera (page 165) Appendix 4: Part 1. Highly resolved cladogram for the parasitic Hymenoptera (page 166) Appendix 4: Part 2. Ichneumonidae - highly resolved cladogram for the parasitic Hymenoptera (page 167) Appendix 4: Part 3. Braconidae - highly resolved cladogram for the parasitic Hymenoptera (page 168) 6 Appendix 4: Part 4a. Chalcidoidea - highly resolved cladogram for the parasitic Hymenoptera (page 169) Appendix 4: Part 4b. Chalcidoidea - highly resolved cladogram for the parasitic Hymenoptera (page 170) Appendix 5: Ichneumonoidea (Hymenoptera) life history data set (page 171) Appendix 6: Taxonomy for the Ichneumonoidea (page 232) Appendix 7: Part1. Braconidae - composite cladogram (page 244) Appendix 7: Part 2a. Ichneumonidae - composite cladogram (page 245) Appendix 7: Part 2b. Ichneumonidae continued - composite cladogram (page 246) Appendix 8: Reference list for the Ichneumonoidea data set (page 247) References (page 275) 7 List of figures and tables Chapter 1: Introduction to life history evolution Figure 1.1: Factors affecting life-history evolution.
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