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Note to Users NOTE TO USERS This reproduction is the best copy available. UMI The utility of behaviour in rnacroevolutionary studles: analyses of caddény (Trichoptera) case building behaviour. Alison Elizabeth Stuart A thesis submitted in conforrnity with the requirements for the degree of Doctor of Philosophy Graduate Department of Zoology University of Toronto Q Copyright by Alison E. Stuart 2000 National Library Bibliothègue nationale 14 ~fcanah du Cana a Acquisitions and Acquisitions et Bibliographk Sewices seMces bibliographiques nie author has granted a non- L'auteur a accordé une Licence non exclusive licence aiiowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seîl reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/fih, de reproduction sur papier ou sur format électronique. The author retahs ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. The utility of behaviour in macroevolutionary studies: analyses of caddisfly (Trichoptera) case building behaviour. Doctor of Philosophy , 2000. Alison E. Stuart. Department of Zoology , University of Toronto. ABSTRACT Behavioural data have been largely ignored in macroevolutionary studies since the advent of modem phylogenetic systematics. The main purpose of this thesis is to contribute toward the revivai of behavioural data for macroevolutionary studies. 1 recognize four hinderances to the use of behavioural data. First, there is debate about what consti tu tes appropriate behavioural characters and behavioural units. Second, there remains concem about behavioural homology, lability and practicaiity. Third, poiential applications of behavioural data are not fully appreciated; and fourth. sharing and exchanging behavioural information is lirnited due to the lack of an adequate forum. 1 address each of these issues to various degrees. concentrating on the first and third. 1 believe that the key to reviving behavioural data in systematics is selecting the most applicable behavioural units. Locornotory components of behaviour (i.e., how an animal behaves) are the most applicable for determining genealogical relationships and thus should be the primary source of behavioural data for macroevolutionary studies. In this thesis 1 detail caddisfly case building and use this infornation to undertake ihree different macroevolutionary endeavoun: phylogeny reconstruction, assessrnent of behavioural evolution, and examination of broader macroevolutionary issues. 1 reconstruct four phylogenies within the Trichoptera using behavioural data: a subordinal-level analy sis, a family-1evel andysis and two generic-level analyses. Thcse analyses indicate that a complex behaviour shared by a group of organisms is likely to yield information about genealogical relationships. regardless of the relationship among the taxa or whether behavioural end-products appear markedly different. Using the behavioural information and a well-resolved phylogeny of the basal trichopteran relationships, 1 discuss a hypothesis of the evolution of case building in caddisflies. ii Building behaviour is anaiogous to development is because it is a hieruchical process that produces an end-product. I used the details of case-building behaviour to address the issue of homoplasy and to test for congruence between behavioural and end-product information. The variety of questions addressed in this thesis, al1 using locomotory components of case building in caddisflies. indicates the potentiai for this type of data in macroevolutionary studies. iii ACKNOWLEDGMENTS After what seems in some ways a very long four years, and in othen an unbelievably short period of time, 1 am feeling close enough to completion to begin writing my acknowledgments. This feels somewhat daunting because there are so many people to thank; 1 guess four years will do that to you. First 1 must Say that these are not cursory thank-you's; each person 1 mention, and probably some I will unwittingly forget, were an unbelievable source of academic ador personal support. I feel very pnviieged to have known each one of you and cannot imagine how 1 would have achieved this without you. To Doug Cume, rny supervisor, many thanks. Here we are. at the end of a pretty long and cool journey. 1 appreciate the knowledge that you passed dong, and I hope I can pass it dong to my own F1 someday! 1 think I was most excited when you told me you were challenging someone's opinion of behavioural plasticity by telling them about locomotory components of behaviour. There is a certain pride in knowing that you have been able to pass a little knowledge in the opposite direction. Special thanks to my comrnittee members, Dr. Chris Darling, Dr. Deborah McLennan and Dr. Locke Rowe. 1 appreciate the comments that you made on various drafts of my thesis, as well as the time you made for me generally over the years. 1 have leamed tons from al1 of you. Other faculty at the University that 1 would like to acknowledge specifically are: Dr. David Dunham, for his general interest in my ideas and his willingness to help in any way and Dr. Sherwin Desser, especidly for coming and chatting with me while 1 waited for the results of my proposal defense. 1 would Iike to that the caddisfly workers who answered my questions (Dr. Glenn Wiggins, Dr. Norman Anderson, Dr. Nancy Erman, Dr. Robert Wisseman, Dr. John Morse), took me collecting (Dr. Glenn Wiggins) and even did some collecting for me (Dr. Nancy Erman and Dr. Dave Larsen). Special thanks to my two field assistants, Lisa McLaughlin and Gerald Tetreault. It makes life so much easier when you have people helping you that you can aist without question. You were both incredible field assistants. iv Completing the thanks for the work side of my thesis, I would like to thank the Natural Sciences and Engineering Research Council of Canada for funding my Ph.D., the Environmental Youth Corps for funding my two field assistants, the Wildlife Research Station in Algonquin Park for some amazing years, and the H. J. Andrews field station in Oregon for an incredible summer in 1997. The personal and acadernic support I received dunng my PhD. was unparalleled. There were some tough times when I questioned my motivation, my desire and my abiiity, and many of the people mentioned above as well as al1 of those to follow were at one time or another called upon to help me out. Sometimes that involved taking me away from it dl. like my trips to New Zealand and Algonquin. not to mention the Woolly and the Albion, and sometimes it just rneant listening. Most of the four years were an exceptional expenence and I'm so glad that 1 had the support to make it through the few tough times and realize this dream of mine. 1 would like to first thank my family, my Mom and Dad and brothers. Rick, Cameron, Jed and Roger. It was great growing up in my family where 1 always knew that the sky was the lirnit. Roger is one of my greatest supporters and 1 one of his; my cousin Peter is so much like my brother that I cenainly consider him one of my immediate farnily, I especially thank both of you for al1 the times you have ken there for me over the years. Thanks so much to the ROM gang (Andy B., John S., Antonia G., Charmaine C., Raoul B.), the Wildlife Research Station gang (Mattman, Lisa E., Meg K., Seanna M.. Keith M., Albrecht S-H.,Kevin J.), the H. I. Andrews gang (Brooks S. Ryan U., Geraid T., Lauren A.), the kiwi gang (Charlie P., Tim W., Sara W., Rhys P.. Alison W.. Justin M.. Kane L, Andy G.), the Horsefly gang (Chene, *Darcy.Jennifer and Jason J.), rny rwrnmates past and present (Dave B., Dan H., Patrick K. Lee M. Daniel F., Julia D.) and my Guelph buddies (Dave J., Scott M., and the Mwneys). 1 want to thank the entire Barrett lab, you guys took me in like 1 was one of your own; excepting me even though I don? study plants! I appreciate al1 your heip, listening to my talk mn-throughs, reading parts of my thesis, giving me hints for my job interview, lening me crash at your houses and generally just being my friends. Thanks so much Linley J., Anne W., Andrea C., and Ange B. 1 want to give some extra special thanks to my very best friends, I am so lucky to have you dl. Meredith McKay, the New Zealand connection, ihanks for pushing me a little harder than 1 ever knew 1 could go; Joe Napolitano, it was so great you got back in touch, 1 can't imagine email without you; Sbeila MacLeod Potter and Derek Potter, thanks for lots of great times and arnazing conversations; Karen McCoy, my globetrotting buddy, 1 can't wait to meet up, catch up and hang out again. thanks for everything. So, that leaves Lisa McLaughlin and Fiona Hunter. It is hard to know where to start with either of hem, Lisa has been there for me though everything and almost every time. no matter what it was, we were laughing by the end. So, Lisa, it looks like we have both survived Our respective degrees I'm glad that we went through it together. 1 can't thank you enough, but 1 hope that after 70 more years of continued friendship, we'll be able to cal1 it even! And Fiona, well, I think I can honestly say that I wouldn't be here today if it weren't for her.
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