The Behavioural Ecology of Social Associations in Fishes by Stacey Sui Yee Lee-Jenkins A thesis submitted to the Faculty of Graduate and Postdoctoral Affairs in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Carleton University Ottawa, Ontario ©2012 Stacey Sui Yee Lee-Jenkins Library and Archives Bibliotheque et Canada Archives Canada Published Heritage Direction du 1+1 Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-93686-3 Our file Notre reference ISBN: 978-0-494-93686-3 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distrbute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. Canada ABSTRACT My thesis focuses on the behavioural ecology of social associations using fishes as a study system. The first part of the thesis examined the influence of social familiarity on shoaling behaviour in the banded killifish (.Fundulus diaphanus) in the laboratory and the field. Killifish preferred familiar shoal mates and this preference occurred in the form of cliques in the social network of a wild population. These social cliques were assorted by body size and furthermore individuals with apparent key roles in the network were larger than others. The second part of the thesis examined how brood-mixing events shape and influence social associations based on familiarity and kinship during early ontogeny in the convict cichlid (Amatitlania siquia). First, I optimised microsatellite markers for this species to determine if brood mixing occurs in wild broods (in the Rio Cabuyo). I genotyped 50 wild broods and categorized them into sibsets. My data indicated that 78% of the broods were mixed (contained foreign young). Potentially 26% of broods contained young from extra-pair copulations. Given the mixed kinship in broods, I performed a four-way choice experiment where I disentangled the effects of social familiarity and kinship on social partner choice in cichlid young (= fry) at varying developmental stages. Smaller fry did not have a shoal preference, whereas larger fry preferred to associate with familiar kin. Lastly, given that fry become separated from their parents and brood mates (= family) during predation encounters in nature and that some fry have a preference for familiar kin, I performed a ‘homing’ experiment in both the field and laboratory to determine if fry that were experimentally displaced from their family would successfully return to it. Displacement distance influenced the likelihood and time displaced fry took to return to their family in both the field and laboratory, and in the laboratory trials, this relationship was influenced by a fry’s body length. The body length of the fish affected the outcome of the field trials; larger fry were more likely to refuge than return to the brood, and smaller fry were more likely to be eaten than to refuge. PREFACE Co-authorship Statement for Chapters 3, 7 and 9 Of the six data chapters included in this thesis, Chapter 3 and Chapter 7 are currently published in peer-reviewed articles and Chapter 9 is currently under review. For Chapter 3 and 9, Dr. Jean-Guy Godin and I collaboratively developed the experimental projects (the questions, logistics, experimental design and analyses). I performed all of the methods, data collections and analyses and wrote up the first draft of the manuscript for these chapters. Dr. Godin provided editorial suggestions and comments which improved these manuscripts. For Chapter 7, the initial development of this project was in collaboration between Dr. Godin, Dr. Myron Smith and I. The collection of convict cichlid fry in Costa Rica was performed by Dr. Godin and 1.1 performed all of the microsatellite optimisation procedures (DNA extraction, PCR, fragment analyses) and subsequent data analysis (allele binning and sibset identification). Adam Densmore was instrumental in the further refinement of the DNA extraction and PCR optimisation which was necessary for the future analysis of this large dataset. Dr. Smith provided me with the genetic technique training required to complete this project and provided lab space and equipment to do so. He also provided insights into the analysis of these data. I wrote up the first draft of this manuscript and all three co-authors provided editorial comments and suggestions for revision. I have received permission from all co-authors to include our collaborative manuscripts in my doctoral thesis (see Appendices). Moreover, I have permission from each publisher (Chapters 3 & 7) to reproduce these published manuscripts in my thesis. To cite this thesis as a whole document or any o f the unpublished chapters I recommend the following citation: Lee-Jenkins, S.S.Y. 2012. The Behavioural Ecology of Social Associations in Fishes. PhD Thesis, Carleton University, Ottawa, Ontario, Canada. The following citations are the published manuscripts (Chapters 3 & 7) reproduced herein and the submitted manuscript currently under peer-review (Chapter 9) at the time o f thesis submission: C hapter 3: Lee-Jenkins, S.S.Y. & Godin, J.-G.J. 2010. Social familiarity and shoal formation in juvenile fishes. Journal o f Fish Biology, 76, 580-590. Publisher: Wiley-Blackwell. C hapter 7: Lee-Jenkins, S.S.Y., Densmore, A.M., Godin, J.-G.J. & Smith, M.L. 2011. Polymorphic microsatellite loci optimised for studies on the convict cichlid fish (Amatitlania siquia). Environmental Biology o f Fishes, 92, 261-266. Publisher: Springer. C hapter 9: Lee-Jenkins, S.S.Y. & Godin, J.-G.J. Concurrent effects of familiarity and kinship on social affiliations in convict cichlid (Amatitlania siquia) young. Behaviour [under review: MS# BES-S-12-00095] v ACKNOWLEDGEMENTS This thesis is the result of an academic ‘journey’ and represents a few tangents from my original thesis research proposal. During my time as a graduate student, I have learned so much about adapting to current circumstances, both the good and the bad! This entire process has been both challenging and rewarding for me, and I have many people to thank. First, I would like to thank my thesis supervisor, Dr. Jean-Guy Godin. Thank you for your guidance, patience and support throughout my graduate career. You have provided me with so many amazing opportunities: field work in Costa Rica and Trinidad, presenting at a large variety of conferences, mentoring students, and moreover you have always encouraged me to expand my professional development (be involved!) and to further develop my skills as a researcher and teacher. I’ve enjoyed our chats about work/life balance and the many lab dinners at the Godins’ (Thank you Heather Godin for the delicious food!). I have learned so much about science during my time in the Godin Lab, thank you for a rewarding and unforgettable experience. My thesis advisory committee (Drs. Gabriel Blouin-Demers and Mark Forbes, and my supervisor) has provided many helpful suggestions during my committee meetings, for which I am grateful. Additionally, thank you Mark for adopting me into your lab when I first arrived and for the use of your microscope! Thank you to my friends (many of whom I met in grad school) who have been supportive throughout this process: Stacey Robinson, Colin Henein, Jen Schellinck, Kim Hair, Tonia Robb, Ryan Chlebak, Michelle Zanuttig, Amanda Goth, Katie Lucas, Jen McAllister, Andrea Main, Devon Metcalf, and Lara Annett. I have particularly enjoyed being involved in the Biology Department with Stacey Robinson; it has been fun being the ‘Staceys’ together! Moreover, I would also like to thank my past and present labmates, many of you have become my close friends: Adam, Alex, Sarah, Kim, Pat, Heather, Pierre-Luc, Kat, Rich, Rebecca, Alisa, Amelia, Steph, Shawn, Megan and Keegan. You have been the most amazing lab mates who have kept me laughing and sane! I’ve enjoyed my many discussions about science with Sarah, Pat and Heather. I would particularly like to thank Adam, Alex, Rebecca and Sarah who contributed directly to my thesis research. Dr. Brian Wisenden has been a very enthusiastic collaborator on the convict cichlid work. I would like to thank him for sharing his convict cichlid knowledge, introducing me to the Rio Cabuyo and Rio Potrero field sites in Costa Rica and for commenting on portions of my thesis research. I would also like to thank Drs. Grant Brown, Jen Snekser and Murray Itzkowitz who provided some breeding convict cichlids for the Godin Lab cichlid population, which made this thesis work possible.