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Sexual Communication in Yellowjackets (Hymenoptera: Vespidae)

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Sexual Communication in Yellowjackets (Hymenoptera: Vespidae) Sexual Communication in Yellowjackets (Hymenoptera: Vespidae) by Nathan Derstine B.A., Eastern Mennonite University, 2010 Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in the Department of Biological Sciences Faculty of Science Nathan Derstine 2017 SIMON FRASER UNIVERSITY Spring 2017 Approval Name: Nathan Derstine Degree: Master of Science Title: Sexual Communication in Yellowjackets (Hymenoptera: Vespidae) Examining Committee: Chair: Harold Hutter Professor Gerhard Gries Senior Supervisor Professor Jenny Cory Supervisor Professor Peter Landolt Supervisor Research Entomologist US Department of Agriculture Sheila Fitzpatrick External Examiner Research Entomologist Agriculture and Agri-Food Canada Date Defended/Approved: April 11, 2017 ii Abstract To determine if and how pheromones mediate sexual communication of yellowjackets [Dolichovespula arenaria, D. maculata, Vespula alascensis, V. pensylvanica, V. squamosa], I took three approaches: (1) In field trapping experiments, I baited traps with a virgin queen (gyne) or a male and tested for their ability to attract prospective mates. I found that only gynes of D. arenaria attracted males. (2) In laboratory Y-tube olfactometer experiments with D. arenaria, D. maculata and V. pensylvanica, I used sibling or non- sibling gynes as a test stimulus, and found that only D. maculata gynes attracted conspecific males, provided they were non-siblings. These results imply an olfactory- based mechanism of nestmate recognition and inbreeding avoidance. (3) I tested the hypothesis that cuticular hydrocarbons (CHCs) differentiate sex, caste, and nest membership. I found that each caste had specific CHC profiles. My data demonstrate the diversity and complexity of sexual communication in yellowjacket wasps, and inspire follow-up studies to identify the sex pheromones. Keywords: Vespidae; Vespula; Dolichovespula; sex pheromones; cuticular hydrocarbons iii Dedication This thesis is dedicated to my parents, Kenton and Rhoda Derstine, who have always nurtured my curiosity, even when it took me far from home. To my brother and sister, Adam and Christina, who have endured more conversations about insect sex than they ever imagined possible. To my friend and mentor Matt Siderhurst who set me on this path. To my partner Allison Cornell, whose love and intelligence have been a pillar of support. And to the much maligned yellowjackets, through which I’ve learned a great deal, and who thrive beautifully despite the ignorant hatred often directed their way. iv Acknowledgements I am extremely grateful to Professor Gerhard Gries for his support, infectious optimism, willingness to dive into unknown territory, and unwillingness to give up. I’ve benefitted greatly from his example. I am equally grateful to Regine Gries, whose expertise and rationality turn ideas into results. Her work ethic, dedication, and kindness are remarkable, and this work would not have been possible without her. I would like to thank my committee members Dr. Peter Landolt and Dr. Jenny Cory for their constructive criticism and feedback. I thank the many undergraduate students who assisted me in data processing, nest collection, and wasp feeding: Nicole Szulc, Allison Canvin, Friday Chen, Lorna Tu, Polly Shao, Zeheng Guo, Hussein Hagehassan, Manpreet Jhutty, Michael Dong, Agnes Gui, and Saru Sandhu. I am very grateful to Sebastian Ibarra Jimenez, who with the advice and support of Alejandro Córdoba-Aguilar, Alejandro Daniel Camacho Vera, Alejandro Zaldívar-Riverón, and María Cristina Mayorga Martínez, as well as the Escuela Nacional de Ciencias Biológicas from the Instituto Politécnico Nacional and the Instituto de Biología and Instituto de Ecología from the Universidad Nacional Autónoma de México, acquired the permits (SGPA/DGVS/10989/14 dated 04 Nov 2014, SGPA/DGVS/12972/14 dated 09 Dec 2014, SAGARPA 4090687366 dated 05 Dec 2014) from the Mexican federal authorities SEMARNAT and SAGARPA, and who located our V. squamosa field site. I also thank the citizens of Santiago Apoala, Oaxaca, for their support in locating V. squamosa nests and for generously facilitating field research on their properties. This research was supported by Graduate Fellowships from Simon Fraser University to Nathan Derstine and by an NSERC – Industrial Research Chair to Professor Gerhard Gries, with former Contech Enterprises Inc. (now Scotts Canada Ltd.) as the industrial sponsor. v Table of Contents Approval ............................................................................................................................... ii Abstract ............................................................................................................................... iii Dedication ........................................................................................................................... iv Acknowledgements .............................................................................................................. v Table of Contents ................................................................................................................ vi List of Tables ..................................................................................................................... viii List of Figures....................................................................................................................... x Chapter 1. Introduction ..................................................................................................1 1.1. The role of pheromones in the reproductive biology of Hymenoptera .......................1 1.2. Recognition and Assessment of Mates ......................................................................2 1.3. Symphyta ....................................................................................................................4 1.4. Aprocrita: Aculeata: Apiformes ...................................................................................5 1.5. Aprocrita: Aculeata: Spheciformes .............................................................................6 1.6. Apocrita: Parasitica .....................................................................................................7 1.7. Future Directions ...................................................................................................... 10 1.8. References ............................................................................................................... 11 1.9. Tables and Figures .................................................................................................. 19 Chapter 2. Evidence for sex pheromones and inbreeding avoidance in select North American yellowjacket species (Hymenoptera: Vespidae) ................................................................................................... 25 2.1. Abstract .................................................................................................................... 25 2.2. Introduction .............................................................................................................. 26 2.3. Methods and Materials ............................................................................................ 28 2.3.1. Experimental Insects .................................................................................. 28 2.3.2. Field trapping experiments ......................................................................... 29 2.3.3. Y-tube olfactometer experiments ............................................................... 30 2.3.4. Statistics ..................................................................................................... 31 2.4. Results ..................................................................................................................... 31 2.4.1. Field trapping experiments ......................................................................... 31 2.4.2. Y-tube olfactometer experiments ............................................................... 31 2.5. Discussion ................................................................................................................ 32 2.5.1. Hypothesis 1: Gynes produce an airborne sex pheromone attractive to males ...................................................................................... 33 2.5.2. Hypothesis 2: Males prefer non-sibling gynes based on olfactory cues ............................................................................................................ 35 2.6. References ............................................................................................................... 36 2.7. Tables and Figures .................................................................................................. 42 Chapter 3. Cuticular hydrocarbons determine sex, caste, and nest membership in each of four species of yellowjackets (Hymenoptera: Vespidae) ........................................................................ 47 3.1. Abstract .................................................................................................................... 47 vi 3.2. Introduction .............................................................................................................. 47 3.3. Methods and Materials ............................................................................................ 49 3.3.1. Experimental Insects .................................................................................. 49 3.3.2. Body surface extracts
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