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ECOLOGICAL FACTORS INFLUENCING COMMUNITY STRUCTURE of PARASITES and THEIR HOSTS by Andrew William Bartlow a Dissertation Submitt

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ECOLOGICAL FACTORS INFLUENCING COMMUNITY STRUCTURE of PARASITES and THEIR HOSTS by Andrew William Bartlow a Dissertation Submitt ECOLOGICAL FACTORS INFLUENCING COMMUNITY STRUCTURE OF PARASITES AND THEIR HOSTS by Andrew William Bartlow A dissertation submitted to the faculty of The University of Utah in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology The University of Utah December 2017 Copyright © Andrew William Bartlow 2017 All Rights Reserved The University of Utah Graduate School STATEMENT OF DISSERTATION APPROVAL The dissertation of Andrew William Bartlow has been approved by the following supervisory committee members: Sarah E. Bush , Chair 08/22/2017 Date Approved Dale H. Clayton , Member 08/22/2017 Date Approved Frederick R. Adler , Member 08/22/2017 Date Approved M. Denise Dearing , Member 08/22/2017 Date Approved Eric A. Rickart , Member 08/22/2017 Date Approved and by M. Denise Dearing , Chair/Dean of the Department/College/School of Biology and by David B. Kieda, Dean of The Graduate School. ABSTRACT Understanding patterns and processes underlying the structure and assembly of ecological communities is a major goal of ecology. Community assembly involves a balance between dispersal events and interactions with biotic and abiotic elements that make up the local environment. Determining which processes dominate and under what circumstances remains unclear for parasite communities. Parasite communities of deer mice are examined to determine whether host traits, parasite interactions, and/or parasite dispersal is important in structuring within host parasite community structure. The results suggest that within host parasite communities accumulate in heavier, older hosts. Parasite dispersal plays a role in structuring parasite communities in deer mice. Helminths and ticks accumulated in older hosts, while fleas and lice did not. Helminths and lice were predicted to accumulate in older hosts because they are associated with their hosts for longer periods of time than ticks and fleas. In studies of parasite communities, it is important to document most, if not all, parasites of interest on or in a host. Using histopathology, I show that deer mice are not infected by parasites in other tissues that are not typically examined during standard parasitological surveys. I also survey rodents in the Great Basin for diseases that are caused by both infectious and non-infectious sources. Two pathogens were found in deer mice: the fungus Emmonsia crescens and the nematode Capillaria hepatica. The most common disease was extramedullary hematopoiesis (11%). I also investigated how habitat change influences species turnover. A major change in the environments of the Intermountain West is the expansion of pinyon-juniper woodlands (P-J). Rodent communities in the Great Basin are assessed to document species turnover following P-J conversion. I found that there was a diversity deficit (immigration credit) following P-J conversion because immigration of new species into P-J converted areas is slower than local species extinction. Diversity may increase if the immigrating species become more abundant. Parasites are also subject to environmental change, but little empirical data exist about how parasites will respond. Resampling historical sites following environmental change suggests that helminth diversity increased throughout the Great Basin region. This change was driven by a decrease in the prevalence and abundance of the pinworm Syphacia peromysci, making the relative abundances of helminth species more even. Complex life cycle parasites appear to be more stable to environmental change than those with direct life cycles because, in this system, complex life cycle parasites have lower host specificity. iv TABLE OF CONTENTS ABSTRACT...................................................................................................................... iii LIST OF TABLES............................................................................................................ vii LIST OF FIGURES............................................................................................................ x ACKNOWLEDGEMENTS............................................................................................. xiv Chapters 1. INTRODUCTION ...........................................................................................................1 Background ................................................................................................................ 1 Chapter Summaries .................................................................................................... 4 References .................................................................................................................. 7 2. THE IMPORTANCE OF HOST TRAITS AND PARASITE DISPERSAL ON THE STRUCTURE OF PARASITE COMMUNITITES IN DEER MICE ................................ 9 Abstract ...................................................................................................................... 9 Introduction ...............................................................................................................10 Methods ................................................................................................................... 14 Results...................................................................................................................... 19 Discussion .................................................................................................................22 Acknowledgements.................................................................................................. 26 References................................................................................................................ 27 3. HISTOLOGICAL SURVEY OF DISEASES AND PARASITES OF WILD RODENTS IN THE GREAT BASIN ............................................................................... 47 Abstract .................................................................................................................... 47 Introduction .............................................................................................................. 48 Methods ................................................................................................................... 50 Results ...................................................................................................................... 53 Discussion ................................................................................................................ 55 Acknowledgements .................................................................................................. 59 References ................................................................................................................ 59 4. SPECIES TURNOVER OF RODENTS FOLLOWING HABITAT CHANGE.. ........ 69 Abstract .................................................................................................................... 69 Introduction .............................................................................................................. 70 Methods ................................................................................................................... 74 Results...................................................................................................................... 79 Discussion ................................................................................................................ 82 Acknowledgements.................................................................................................. 87 References................................................................................................................ 87 5. IMPACT OF ENVIRONMENTAL CHANGE ON PARASITE COMMUNITIES ...106 Abstract .................................................................................................................. 106 Introduction............................................................................................................ 107 Methods ................................................................................................................. 112 Results .................................................................................................................... 119 Discussion .............................................................................................................. 122 Acknowledgements ................................................................................................ 128 References .............................................................................................................. 128 Appendices A. STABLE ISOTOPE ANALYSIS OF DEER MICE IN THE GREAT BASIN ........ 147 B. SPECIES OF LICE COLLECTED FROM RODENTS IN THE GREAT BASIN ... 148 C. ANIMALS INFESTED WITH BOTFLIES AT TIME OF CAPTURE .................... 149 D. HOST-PARASITE DATA FROM RODENTS IN THE GREAT BASIN ............... 150 E. HISTOPATHOLOGY RESULTS OF RODENTS .................................................... 167 F. SEQUENCES OF PARASITE SPECIES OF DEER MICE ...................................... 170 vi LIST OF TABLES 2.1. Life history traits of the seven taxonomic groups of parasites. In the column of time spent with host, short (S) refers to parasites that are associated with their hosts for up to several days or feed intermittently. Long (L) refers to parasites being associated with their hosts for long period of time (several weeks to months) .......................................... 33 2.2. The prevalence and
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