PHYLOGENETIC SIGNAL IN POLLINATION NETWORKS VARIES IN STRENGTH WITH POLLINATOR TYPE AND COMMUNITY PLANT SPECIES RICHNESS by Clea Margaret Moray B.Sc., University of Guelph, 2003 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE In the Biological Sciences Department © Clea Margaret Moray 2009 SIMON FRASER UNIVERSITY Fall 2009 All rights reserved. However, in accordance with the Copyright Act of Canada, this work may be reproduced, without authorization, under the conditions for Fair Dealing. Therefore, limited reproduction of this work for the purposes of private study, research, criticism, review and news reporting is likely to be in accordance with the law, particularly if cited appropriately. APPROVAL Name: Clea Moray Degree: Master of Science Title of Thesis: Phylogenetic signal in pollination networks varies in strength with pollinator type and community plant species richness. Examining Committee: Chair: Dr. Glen Tibbits Professor ________________________________________ Dr. Arne Mooers Senior Supervisor Associate Professor ________________________________________ Dr. Elizabeth Elle Supervisor Associate Professor ________________________________________ Dr. Diane Srivastava Supervisor Associate Professor University of British Columbia ________________________________________ Dr. Sarah Otto External Examiner Professor University of British Columbia Date Defended/Approved: December-17-2009 ii Declaration of Partial Copyright Licence The author, whose copyright is declared on the title page of this work, has granted to Simon Fraser University the right to lend this thesis, project or extended essay to users of the Simon Fraser University Library, and to make partial or single copies only for such users or in response to a request from the library of any other university, or other educational institution, on its own behalf or for one of its users. The author has further granted permission to Simon Fraser University to keep or make a digital copy for use in its circulating collection (currently available to the public at the “Institutional Repository” link of the SFU Library website <www.lib.sfu.ca> at: <http://ir.lib.sfu.ca/handle/1892/112>) and, without changing the content, to translate the thesis/project or extended essays, if technically possible, to any medium or format for the purpose of preservation of the digital work. The author has further agreed that permission for multiple copying of this work for scholarly purposes may be granted by either the author or the Dean of Graduate Studies. It is understood that copying or publication of this work for financial gain shall not be allowed without the author’s written permission. Permission for public performance, or limited permission for private scholarly use, of any multimedia materials forming part of this work, may have been granted by the author. This information may be found on the separately catalogued multimedia material and in the signed Partial Copyright Licence. While licensing SFU to permit the above uses, the author retains copyright in the thesis, project or extended essays, including the right to change the work for subsequent purposes, including editing and publishing the work in whole or in part, and licensing other parties, as the author may desire. The original Partial Copyright Licence attesting to these terms, and signed by this author, may be found in the original bound copy of this work, retained in the Simon Fraser University Archive. Simon Fraser University Library Burnaby, BC, Canada Last revision: Spring 09 ABSTRACT Phylogenetic constraints have been hypothesized to influence the complex network structure found in plant-pollinator communities. Here, I develop and test a conceptual model of factors that might modulate any signal of phylogenetic clustering of plants visited by individual pollinator species. Across 29 communities, pollinators visited weakly phylogenetically clustered plant species. Plant relatedness was positively correlated with community plant richness; and plants visited by versatile pollinators were phylogenetically random in small communities but became clustered as richness increased. However, these patterns were not explained by a simple dichotomous scoring of plants as restrictive or unrestrictive to unversatile pollinators. Collectively, these results suggest that constraints imposed by plant phylogeny on pollination networks are moderated by current ecological processes such as community assembly and pollinator foraging behaviour, but mechanisms are unclear. Future research should consider the opportunity for bidirectional interplay between ecological and phylogenetic effects. Keywords: plant-pollinator network; phylogenetic relatedness; phylogenetic signal; community structure; species richness; floral traits. iii ACKNOWLEDGEMENTS I thank Arne Mooers deeply for his guidance, enthusiasm and support in matters scientific and otherwise. I also greatly appreciate the helpful advice of Elizabeth Elle, Diane Srivastava, Jana Vamosi and Sally Otto. I thank the members of FAB*-lab and the Elle lab, especially Lisa Neame, Nicole Tunbridge, Laura Weir, Scott Pavey, Aki Mimoto and Emily Meuser, as well as Maxence Salomon, Samantha Vibert, and Tom Cowan for their assistance and encouragement. For their participation in the other half of my education at SFU, I thank Mike Letourneau, Ursula Arndt, Kathy McKay, Graham Lyons, Joel Blok, Derrick Harder, Amanda van Baarsen, and Haida Arsenault-Antolick. Finally, I am grateful to my parents, my grandmother Olga Glasser, my aunt Margaret Eggen, and Chris Dickert for their love and encouragement. iv TABLE OF CONTENTS Approval...........................................................................................................................ii Abstract...........................................................................................................................iii Acknowledgements.........................................................................................................iv Table of Contents............................................................................................................ v List of Figures ................................................................................................................vii List of Tables ................................................................................................................ viii 1: INTRODUCTION ........................................................................................................ 1 1.1 Overview: community ecology of pollination............................................................ 1 1.2 Phylogenetic constraints in pollination networks...................................................... 2 1.3 Effect of point-of-view on detection of phylogenetic constraints............................... 4 1.4 Between community variation in phylogenetic constraints....................................... 7 1.5 Within-community variation in phylogenetic constraints........................................... 9 1.6 Conceptual model ................................................................................................. 11 2: METHODS................................................................................................................ 19 2.1 Datasets ............................................................................................................... 19 2.2 Phylogenies .......................................................................................................... 20 2.3 Null model test of phylogenetic clustering of visited plants .................................... 22 2.4 Metric of phylogenetic clustering of visited plants.................................................. 23 2.5 Classification of floral morphology and pollinator type ........................................... 25 2.6 Mean node depth of community phylogenies ........................................................ 26 2.7 Phylogenetic signal ............................................................................................... 26 2.8 Statistical analysis................................................................................................. 27 2.8.1 Overall presence of phylogenetic clustering .............................................. 28 2.8.2 Relationship between plant species richness and phylogenetic signal of flower type............................................................................................. 28 2.8.3 Relationship between plant species richness and mean phylogeny node depth ................................................................................................ 28 2.8.4 Difference in phylogenetic clustering between pollinator types versus community plant species richness ............................................................. 29 2.8.5 Difference in pollinator preference for floral morphology versus community plant species richness ............................................................. 29 3: RESULTS ................................................................................................................. 30 4: DISCUSSION ........................................................................................................... 33 4.1 Summary of results ............................................................................................... 33 4.2 Overall presence of phylogenetic clustering (question 1) ...................................... 33