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Reproductive Strategies in Parasitic Wasps Ian Charles Wrighton Hardy

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Reproductive Strategies in Parasitic Wasps Ian Charles Wrighton Hardy 1 Reproductive Strategies in Parasitic Wasps by Ian Charles Wrighton Hardy A thesis submitted for the degree of Doctor of Philosophy of the University of London and for the Diploma of Imperial College Department of Biology and Centre for Population Biology, Imperial College at Silwood Park, Ascot, Berkshire, SL5 7PY, U.K. 1991 (Submitted November 1990) 2 Abstract This thesis investigates the evolutionary ecology of reproduction by parasitoid wasps. In haplodiploid populations some females are constrained to produce sons only, theor­ etically, the optimal progeny sex ratio of unconstrained females may be influenced. Prevalences of constrained females are assessed in parasitoids of D ro so p h ila and from the literature. Constrained oviposition is generally rare, however, in some species constrained females are sufficiently common to affect unconstrained female’s sex ratios. Goniozus nephantidis females remain with their broods until the offspring pupate. G. nephantidis competes for hosts with conspecific and non-conspecific parasitoids. The costs of remaining seem at least partially offset by the prevention of oviposition by competing parasitoids. To predict clutch size, the relationship to the p e r c a p ita fitness of offspring must be known and also the parental trade-off between present and future reproduction. Since trade-offs are assumed unimportant in G. nephantidis clutch fitness should be maximised, this is achieved at the ’Lack clutch size’. Females adjust clutch size to host size. Manipulation of clutch size on standard hosts shows that developmental mortality is unaffected by clutch size, but larger females emerge from smaller clutches and have greater longevity and fecundity. Lack clutch sizes are calculated using longevity and fecundity as estimators of offspring fitness. Calculated clutch sizes are larger than those produced by the parasitoids. Disparities are attributed to incorrect assumptions about the importance of trade-offs or to inadequate measurement of offspring fitness (laboratory-based approaches are unable to assess all components of fitness). Precise sex allocation is theoretically advantageous over binomial sex allocation, since the number of mated females is maximised when brood sex ratio variances are reduced. G. nephantidis sex ratios are compared with those expected under binomial sex allocation. Precise sex ratios are produced at most brood sizes, although precision is limited, probably by factors such as pre-adult mortality. To Esther, to John and to the rest of my family. 4 Table of Contents Abstract ................................................................................................................................. 2 Table of Contents ................................................................................................................ 4 List of Figures ...................................................................................................................... 7 List of T ab les........................................................................................................................ 8 Acknowledgements ............................................................................................................ 9 Chapter One Introduction................................................................................................. 10 1.1 Behavioural ecology and reproductive strategies .......................................... 10 1.2 Parasitoids, parasitic wasp diversity and the Bethylidae ............................. 11 1.3 Parasitoids and evolutionary stu d ies ................................................................. 13 1.3.1 Maternal Care ............................................................................................... 13 1.3.2 Clutch size ..................................................................................................... 14 1.3.3 Sex ratio .......................................................................................................... 16 1.4 Statistical an alysis.................................................................................................. 21 Chapter Two Constrained Sex Allocation in Hymenoptera .................................... 23 2.1 Abstract ..................................................................................................................... 23 2.2 Introduction............................................................................................................. 24 2.3 Constrained Oviposition by D ro so p h ila Parasitoids...................................... 30 2.3.1 Biology of D r o s o p h ila parasitoids ........................................................... 30 2.3.2 Methods .......................................................................................................... 31 2.3.3 R esults............................................................................................................. 32 2.4 Literature R ev iew ................................................................................................... 34 2.5 Discussion ................................................................................................................ 39 2.5.1 D r o s o p h ila parasitoids ................................................................................ 39 2.5.2 General prevalence of constrained oviposition...................................... 39 Chapter 3 Brood guarding in the bethylid wasp Goniozus nephantidis ............... 42 3.1 Abstract ..................................................................................................................... 42 3.2 Introduction.............................................................................................................. 43 3.2.1 Biology of G. nephantidis........................................................................... 43 3.3 M ethods..................................................................................................................... 44 3.3.1 A nim als........................................................................................................... 44 5 3.3.2 Experimental procedures .................................................................. 45 3.4 Results ....................................................................................................... 46 3.4.1 Control .............................................................................................. 46 3.4.2 Brood survivorships ........................................................ .................. 47 3.4.3 Presence or absence of the initial female and intruder oviposition .. 48 3.4.4 Survival of the initial female's brood and intruder oviposition ........ 49 3.5 Discussion ...................... ........................................................................... 52 Chapter 4 Clutch size in Goniozus nephantidis .... ............ ........ .......... ................ 55 4.1 Abstract .................................... .......... .......................... ............................. 55 4.2 Introduction .................... .......... .............................. ................ ................... 56 4.2.1 Biology of G. nephantidis ................................................................. 57 4.3 Methods.......................... .......... .............................. ................................... 58 4.3.1 Animals ............................................................................................. 58 4.3.2 Clutch size and host size ................................................................... 59 4.3.3 Clutch size manipulations ................................................................. 59 4.3~4 The effects of the mother remaining with her clutch ........................ 59 4.3.5 Reproductive female's longevity and fecundity ............................... 60 4.3.6 Female longevity without reproduction ............................................ 60 4.3.7 Statistical analysis ............................................................................. 60 4.4 Results ....................................................................................................... 61 4.4.1 Clutch size and host size ...... ................ ............... .............................. 61 4.4.2 Clutch size manipulations ................................................................. 64 4.4.3 The effects of mother remaining with her clutch .......................... .... 67 4.4.4 Influence of adult size on fitness .................. .................................... 68 4.4.4.1 Female fecundity ...................................................................... 68 4.4.4.2 Reproductive female's longevity .......................... .................... 70 4.4.4.3 Female longevity without reproduction .......... ......................... 72 4.5 Discussion ................................................................................................. 73 Chapter 5 Precise Sex Ratios in Goniozus nephantidis ....................................... 82 5.1 Abstract ..................................................................................................... 82 5.2 Introduction ............................................................................................... 83 5.2.1 Evidence for precise sex ratios ........... ...... ........... ............................. 84 5.2.2 The biology of Goniozus nephantidis ............................................... 93 6 5.2.3 Primary
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