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Ormia Ochracea on the Cricket Gryllus Rubens

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Ormia Ochracea on the Cricket Gryllus Rubens Foraging theory as a model to examine parasitism by the Tachinid parasitoid fly Ormia ochracea on the cricket Gryllus rubens by Andrew J. Masson A thesis submitted in conformity with the requirements for the degree of Master of Science Department of Ecology and Evolutionary Biology University of Toronto © Copyright by Andrew Masson 2015 Foraging theory as a model to examine parasitism by the Tachinid parasitoid fly Ormia ochracea on the cricket Gryllus rubens Andrew Masson Master of Science Department of Ecology and Evolutionary Biology University of Toronto 2015 Abstract This paper tests the effects of auditory and chemical sensory input on the foraging and larviposition behaviour of an eavesdropping parasitoid fly. Ormia ochracea (Diptera: Tachinidae) use the dominant calling field cricket in their habitat as hosts for their larvae; females locate hosts by eavesdropping on the acoustic calls produced by male crickets. By manipulating directional cues in the auditory stimulus experienced by foraging females their ability to locate hosts and their ability to switch to a novel host following a switch in the location of the stimulus were examined. Success was dependent upon walking behaviour and proximity at stimulus change. After locating potential hosts females must deposit larva and differences in larviposition rates of females, after they had located an attractive sound source (cricket call), in the absence and presence of chemical cues associated with crickets indicate that females may use additional sensory modalities in host-foraging decisions. ii Table of Contents Contents Table of Contents ............................................................................................................... iii List of Tables ..................................................................................................................... iv List of Figures ..................................................................................................................... v Chapter 1 ............................................................................................................................. 1 1.1 Introduction ................................................................................................................... 1 1.1.1 Foraging theory and modeling .................................................................................. 1 1.1.2 Foraging parasitoid modeling .................................................................................... 2 1.1.3 Insect acoustic communication .................................................................................. 3 1.1.4 Ormia ochracea hearing ............................................................................................ 3 1.1.5 Phonotaxis in Ormia ochracea .................................................................................. 4 1.1.6 Chemical cue communication .................................................................................... 4 1.1.7 Life cycle .................................................................................................................. 5 1.1.8 Goals ......................................................................................................................... 6 1.2 Method .......................................................................................................................... 7 1.2.1 Study animal .............................................................................................................. 7 1.2.2 Room and arena set up ............................................................................................... 8 1.2.3 Experiment 1: Speaker turnoff ................................................................................... 8 1.2.4 Experiment 2: Speaker switch .................................................................................. 8 iii 1.2.5 Sound field ................................................................................................................. 9 1.2.6 Experiment 3: Chemical cue effect of larvaposit behaviour ...................................... 9 1.2.7 Recordings .............................................................................................................. 10 1.2.8 Statistics ................................................................................................................... 10 1.3 Results ........................................................................................................................ 11 1.3.1 Behavioural types .................................................................................................... 11 1.3.2 Speaker switch ........................................................................................................ 11 1.3.3 Speaker turnoff ........................................................................................................ 12 1.3.4 Larviposition ............................................................................................................ 12 1.4 Discussion .................................................................................................................. 13 References ........................................................................................................................ 19 Appendix ........................................................................................................................... 41 iv List of Tables (if any) Table 1: Speaker calibration ................................................................................................ 25 Table 2: Speaker turnoff modeling ...................................................................................... 26 Table 3: Speaker switch modeling ....................................................................................... 28 v List of Figures Figure 1: Calibration speaker 1 ............................................................................................. 30 Figure 2: Arena schematic ................................................................................................... 31 Figure 3: Sound calibration around speaker at turning distances ......................................... 32 Figure 4: Calibration of speaker 2 at turning points ............................................................ 33 Figure 5: Sound wave .......................................................................................................... 34 Figure 6: Walking behaviour speaker switch ........................................................................ 35 Figure 7: Walking behaviour speaker turnoff ...................................................................... 36 Figure 8: General linear model speaker turnoff: Distance vs. Success ................................. 37 Figure 9: General linear model speaker turnoff: Latency to respond post turnoff vs. Success ............................................................................................................................ 38 Figure 10: General linear model speaker switch: Distance vs. Success ............................... 39 Figure 11: Larviposition rates in 4 treatments ...................................................................... 40 vi 1 Chapter 1 1.1 Introduction 1.1.1 Foraging theory and modeling The process of searching for and consuming food, foraging, can be modeled under the assumption that individuals make decisions to maximize energy gain from foraging. In order to model an individual’s decision-making and predict behaviour, it is assumed that individuals will forage in an optimal manner. The basis of this assumption is that through natural selection a forager should behave in a manner that will maximize their energy intake while minimizing energy expenditure (Pyke et al. 1977; van Alpen et al. 2003). A mathematical model was developed by Charnov (1976), using the Marginal Value Theorem of Optimal Foraging, which is a restating and expansion of the work of MacArthur and Pianka (1966). In this model a predator is making decisions based on energy and time costs associated with the decision of either continuing to search in a patch (area of habitat with prey are available) or switch patches. There are three basic pieces of information that are required to generate an effective model for foraging: breadth of diet (specialist or generalist); strategies of movement (flight, walk, crawl); and the type of environment (patchy, uniform) (MacArthur & Pianka 1966; Charnov 1976). With this information the model is able to determine mathematically when the highest pay off for switching patches will occur given the quality of the current patch, and the status of other patches. The marginal value theory of optimal foraging determines theoretical values that an idealized optimal predator would use while foraging to exploit habitats. In this model a predator should continue searching in a patch as long as the resulting gain from time spent searching per unit food exceeds the loss (MacArthur & Pianka 1966; Charnov 1976). Prey items may be found while searching within a patch but will not be while switching patches. However, with increased time spent searching within a patch there is a decrease in prey availability - a phenomenon known as predator depression (Charnov 1976). This means that a patch should be exploited until the rate of fitness gain within the patch has decreased to a marginal value calculated in models as the mean fitness value of all patches (Wajnberg et al. 2000). In a theoretical
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