PHYLOGENETIC SYSTEMATIC ANALYSIS OF THE NEODERMATA (PLATYHELMINTHES) AND ASPIDOBOTHREA (TREMATODA, NEODERMATA) WITH INVESTIGATION OF THE EVOLUTION OF THE QUINONE TANNED EGGSBELL. David Zamparo A thesis submitted in codormity with the requirements for the degree of M. Sc. Graduate Department of Zodogy University of Toronto @Copyrightby David Zamparo 2ûû1 National Library Biblioth ue nationale 1*1 ,cm, du Cana% . .. et "4""""dBib iographic SeMms MIiographiques The author has granted a non- L'auteur a accordé une licence non exclusive licence aliowiag the exchsive permettant à la National Library of Canada to Bïbiiotheque nationale du Canada de reproduce, loan, distribute or sel1 reproduire, prêter, distribuer ou copies of this thesis in microforni, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur format dectronique. The author retains 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 otheIWise de celle-ci ne doivent être imprim6s reproduced without the author's ou autrement reproduits sans son permission. autorisation. Phylogenetic systematic analysis of the Neodermata (Platyhelminthes) and Aspidobothrea (Trematoda, Neodemata) with investigation of the evolution of the quinone tanned eggshell. Masters of Science, 2001. David Zamparo, Graduate Deputment of Zoology. University of Toronto. A phylogenetic analysis of the Neodermata and their closest relatives (the Rhabdocoela) was undertaken in order to provide a robust estimate of phylogeny. This phylogenetic analysis incorporates new character information and addresses a number of methodological issues raised by recent phylogenetic systematic analyses of the Platyhelminthes. A phylogenetic analysis of the Aspidobothna incorporates a new genus, Sychnocoryle Ferguson et al. 1999, and 16 new rnorphological characters. This analysis tests three previously proposed farnily-level hypotheses. The two phylogenetic systematic studies undertaken herein provides the basis for a study of the evolution of quinone- tanned eggshell arnong the parasitic platyhelminths. Its been hypothesized that quinone- tanned eggshell are a "pre-adaptation" (exaptation) to endoparasitism. 1evaluate this hypothesis by means of the comparative phylogenetic approach, which provides both 3 test and suggests future research. Such an endeavor takes over one's entk life and becomes an al1 consurning passion so that it is difficult to acknowledge ali those in one's life who deserve rightful recognition. To my loving wife, Shirley, who has more than anyone understood what this project has meant to me and who has sacrificed a great deal to afford me the opportunity to foîiow what is best described as a calling, 1 offer my most sincere thanks. To my supervisor, Dr. Deborah McLennan, 1 especially thank you for the opportunity which so many are unwilling to lend to young people. 1 also thank you for ail the support and encouragement 1 so desperately required throughout the gestation of this work. You have been most attentive to my needs, providing assistance whatever the situation, and never having to ask for it, 1 thank you. It has tnily been a great honor and privilege to have learned from such a distinguished and remarkable researcher. 1 can only hop that 1 have not been a disappointment. Special thanks to Dr. Daniel Brooks, for making advanced copies of his own and colleagues' material available to m.1 thank him for introducing me both to various researchea and to field work at the ACG in Costa Rica. I thank you for your mentoring in conducting field work and sharing your laboratory expertise with me. 1 thank you for patiently sitting on the side and allowed me to delve into a field of study you have invested so much in; you have shown by exarnple how one acts professionally in this occupation. Michelle Mattem who had. 1 am sure, the unbearable duty of sharing an offce with me for the past two years, I thank you for fruithl discussions on phylogenetic systematics and your indispensable technical assistance without which this work could not be possible. Al1 whose sojoums have taken them through the lab, Dr. Anindo Choudhuty, Dr. Fernando Marques and Bryan Rogers, you have al1 been inspuational to me. To the faculty and staff of the Department of Zoology, University of Toronto who took great care of me so that 1could apply myself fully to this project, 1 would like you to know that your work does not go unnoticed by graduate students. 1 would aiso like to thank Donna Stugyls and the whole staff at Gerstein Library Interlibrary Loans, University of Toronto for providing exceptional service. To my parents and extended farnily ,w ho suffered neglect at the very hands of this thesis, 1 dedicate this work to you in the hopes that it offers an explanation. iii .. Abstract ..........................................................................................JI *.. Acknowledgments.. ............................................................................lu . Table of Contents.. .............................................................................IV-VI CHAPTERONE- GENERALINTRODUCTION The Importance of Parasites.. .................................................................7-8 Intmducing the Neodemata.. .................................................................8- 12 Focus of the Thesis. .............................................................................12-14 CHA~Rm0- PHYLOGENETICANALYSIS OF THE RHABDOCOELA (PLATYHELMINTHES)WITH EMPHASIS ON THE NEODERMATAAND RELATIVES Introduction ......................................................................................15-16 Materials & Methods.. ......................................................................... 17-29 Results ............................................................................................29-3 1 Discussion ..................................................................................... ..314 Conclusions.................................................................................... ..40-4 1 CWR THREE- PHYLOGENETIC SYSTEMATICASSESSMENT OF THE ASPIWBOTHREA(PLATYHELMINTHES, NEODERMATA, TREMAToDA) Introduction. .................................................................................... S7-58 Materials & Methods.. ........................................................................ S9-63 Re~ulfs.......................................................................................... .63.H Discussion ........................................................................................64 .67 Conclusion...................................................................................... .6 7-68 CHAFTERFOUR- THE EVOLU~ONOF QUINONETANNED EGGs IN THE Introduction ......................................................................................7 6-79 Materials &Methods ............................................................................7 9-82 Results & Discussion ..........................................................................-83-9 1 LISTOF FIGURES Figure 2.1 .......................................................................................4243 Figure 2.2. ......................................................................................4445 Figure 2.3. ......................................................................................46-47 Figure 2.4 .......................................................................................4849 Figure 2.5. ....................................................................................$50-5 1 Figure 3.1...................................................................................... -69-70 Figure 3.2 ....................................................................................... 7 1-72 Figure 4.1 ....................................................................................... 92-93 Figure 4.2. ..................................................................................... .94-95 Figure 4.3. ..................................................................................... .96-97 Figure 4.4. ..................................................................................... .98-99 Figure 4.5.................................................................................... lWl01 Figure 4.6.................................................................................... 102-103 Figure 4.7 ....................................................................................104-105 Figure 4.8 ....................................................................................1M-107 Figure 4.9 .....................................................................................108-109 Table 2.1 ......................................................................................S2-56 Table 3.1. ......................................................................................7 3-75 Appendix 1................................................................................. 19% 197 Appendix 2................................................................................ .19 8.20 1 Appendix 3................................................................................. .202= 224 Chapter One GENERALINTRODUCTION THEIMPORTANCE OF PARASITES Parasitism is a ubiquitous and highly successful mode of life. Most avid naturalists