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The Evolution of Seed Dispersal Syndromes in Prunus

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The Evolution of Seed Dispersal Syndromes in Prunus University of Nevada, Reno The evolution of seed dispersal syndromes in Prunus A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Ecology, Evolution and Conservation Biology by Maurie J. Beck Stephen B. Vander Wall, Dissertation Advisor August, 2009 THE GRADUATE SCHOOL We recommend that the dissertation prepared under our supervision by MAURIE BECK entitled The Evolution Of Seed Dispersal Syndromes In Prunus be accepted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Stephen B. Vander Wall, Advisor Stephen H. Jenkins, Committee Member William S. Longland, Committee Member David W. Zeh, Committee Member Thomas J. Nickles , Graduate School Representative Marsha H. Read, Ph. D., Associate Dean, Graduate School August, 2009 i ABSTRACT There are two fruit types in the genus Prunus. The majority of species have fleshy- fruited drupes, which are considered the ancestral phenotype. In the deserts of Eurasia and North America there are also species that produce dry fruits and large nuts, suggesting this fruit type has originated independently on numerous occasions in response to dry conditions. Fleshy-fruited Prunus are dispersed by frugivorous animals, primarily birds and some mammalian carnivores. In this dissertation, by documenting complete seed fate pathways, I demonstrate that desert peach (Prunus andersonii), a dry, nut-producing species in the western Great Basin of North America, is only dispersed by scatter- hoarding rodents. Additionally, I demonstrate that western chokecherry (P. virginiana var. demissa, Rosaceae) is also dispersed by scatter-hoarding rodents, following primary dispersal by endozoochorous frugivores. This type of two-phased seed dispersal is a form of diplochory, a process that employs different modes of dispersal during sequential dispersal phases and usually offers unique benefits during each phase. Although not well documented, frugivory followed by scatter hoarding is believed to be more common than previously thought. In this case I show that phase I dispersal by frugivores transports seeds, often long distances, away from the parent plants. Scatter-hoarding rodents then harvest the seeds from feces and bury them in soil during phase II dispersal. Caching chokecherry seeds not only moves them away from the parent plant, but constitutes directed dispersal, a form of seed dispersal that disproportionately enhances seed and seedling survival. ii The transition in Prunus from primary dispersal by fruit-consuming animals to nut dispersal by scatter-hoarding rodents is difficult to envision. However, if the ancestor to desert peach or other dry-fruited species utilized diplochory, then the most parsimonious explanation is that the transformation from fleshy fruits to dry nuts was accompanied by the loss of frugivory and the reliance only on dispersal by scatter-hoarding rodents. In chapter 3 I review to similarities and differences between frugivory and dispersal by scatter-hoarding animals. iii ACKNOWLEDGEMENTS I am indebted to my major professor, Stephen B. Vander Wall, a consummate experimental ecologist and naturalist. I would also like to thank my graduate committee, Stephen H. Jenkins, William S. Longland, David W. Zeh, and Thomas J. Nickles for contributing to my dissertation and my development as a researcher. I would especially like to thank the Vander Wall lab, including Jenny Briggs, Julie Roth, Ted Thayer, Jennifer Armstrong, Jennifer Hollander, and Kellie Kuhn. They were like a family to me, helping tirelessly in the field and as supportive critics, separating the grain of ideas with merit from the chaff of those without. I thank Jenny Francis, Jessica Hay, and Chris Farrar for assistance in the field. Dr. Chris Ross provided me with one of my field sites. I received help with statistics from G. C. J. Fernandez, I. Aban, and D. Board for help. I received financial support from the Ecology, Evolution, and Conservation Biology (EECB) program and the Graduate Student Association at the University of Nevada, Reno and the Whittell Forest and Wildlife Area. I received additional financial assistance from C. Richard Tracy. Lucy Morris, Pauline Jasper, Lou Christiansen, and Cheri Briggs provided logistical support. I owe much to my family and friends for their companionship and love during this long journey, including my parents, Barbara Beck and Sam Sokolow, my godmother, Marion Kamins, my friends Dave and Betsy Harden, the Scarpaci family, Robby Weaver and Stacy Daghliesh, Jenseen Brons, Patti Doty, and the Jahn Family for providing a space conducive to finishing my dissertation, along with the fine food and well-timed threats. iv TABLE OF CONTENTS GENERAL INTRODUCTION................................................................................................... I REFERENCES.................................................................................................................... 3 Chapter 1. Desert peach (Prunus andersonii) and the transition in seed dispersal mode from frugivory to scatter-hoarding...................................................................................... 5 ABSTRACT........................................................................................................................ 6 INTRODUCTION................................................................................................................ 8 METHODS ...................................................................................................................... 10 Description of study sites............................................................................................ 10 Fruit development and production ............................................................................. 11 Foraging and removal of nuts .................................................................................... 12 Rodent community composition.................................................................................. 15 Caching behavior of rodents in field enclosures ........................................................ 15 Seed dispersal from source shrubs ............................................................................. 16 Seedling germination, emergence, and survival......................................................... 17 RESULTS ........................................................................................................................ 18 Fruit development and production ............................................................................. 18 Foraging and removal of nuts .................................................................................... 20 Rodent community composition.................................................................................. 24 Seed dispersal from source shrubs ............................................................................. 24 Seedling germination, emergence, and survival......................................................... 26 DISCUSSION ................................................................................................................... 29 ACKNOWLEDGEMENTS.................................................................................................. 39 REFERENCES.................................................................................................................. 39 TABLES .......................................................................................................................... 43 FIGURES......................................................................................................................... 51 Chapter 2. Diplochory in western chokecherry: you can’t judge a fruit by its mesocarp 61 ABSTRACT...................................................................................................................... 62 INTRODUCTION.............................................................................................................. 64 v METHODS ...................................................................................................................... 66 Study area and species................................................................................................ 66 Fruit development and crop size................................................................................. 67 Foraging on chokecherry............................................................................................ 67 Seed removal transects ............................................................................................... 70 Rodent community composition.................................................................................. 70 Caching behavior of rodents in field enclosures ........................................................ 71 Secondary dispersal by rodents .................................................................................. 71 Seedling emergence and survival ............................................................................... 72 Recruitment potential.................................................................................................. 74 RESULTS ........................................................................................................................ 74 Fruit development and crop size................................................................................. 74 Foraging on chokecherry...........................................................................................
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