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«Luke Elden Dodd The Graduate School University of Kentucky 2010 FOREST DISTURBANCE AFFECTS INSECT PREY AND THE ACTIVITY OF BATS IN DECIDUOUS FORESTS ...»

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Abstract

OF DISSERTATION

Luke Elden Dodd

The Graduate School

University of Kentucky

2010

FOREST DISTURBANCE AFFECTS INSECT PREY AND THE ACTIVITY OF BATS

IN DECIDUOUS FORESTS

____________________________________

ABSTRACT OF DISSERTATION

_____________________________________

A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the College of Agriculture at the University of Kentucky By Luke Elden Dodd Lexington, Kentucky Director: Dr. Lynne K. Rieske-Kinney, Professor of Entomology Lexington, Kentucky 2010 Copyright © Luke Elden Dodd 2010

ABSTRACT OF DISSERTATION

FOREST DISTURBANCE AFFECTS INSECT PREY AND THE ACTIVITY OF BATS

IN DECIDUOUS FORESTS

The use of forest habitats by insectivorous bats and their prey is poorly understood. Further, while the linkage between insects and vegetation is recognized as a foundation for trophic interactions, the mechanisms that govern insect populations are still debated. I investigated the interrelationships between forest disturbance, the insect prey base, and bats in eastern North America.

I assessed predator and prey in Central Appalachia across a gradient of forest disturbance (Chapter Two). I conducted acoustic surveys of bat echolocation concurrent with insect surveys. Bat activity and insect occurrence varied regionally, seasonally, and across the disturbance gradient. Bat activity was positively related with disturbance, whereas insects demonstrated a mixed response. While Lepidopteran occurrence was negatively related with disturbance, Dipteran occurrence was positively related with disturbance. Shifts in Coleopteran occurrence were not observed. Myotine bat activity was most correlated with sub-canopy vegetation, whereas lasiurine bat activity was more correlated with canopy-level vegetation, suggesting differences in foraging behavior.

Lepidoptera were most correlated with variables describing understory vegetation, whereas Coleoptera and Diptera were more correlated with canopy-level vegetative structure, suggesting differences in host resource utilization.

I assessed the food habits of bats captured in mist nets. Morphological identification of prey suggested consumption of insect taxa varies across bat species and, at least for the most commonly-captured species, Myotis septentrionalis, across the region (Chapter Three). Trophic connections were further delineated between M.

septentrionalis and its prey by sequencing COI fragments of insect prey obtained from fecal samples. Prey identities were inferred for COI fragments using web-based searches (Chapter Four), as well as tree-building analyses (Chapter Five). Lepidoptera were detected most frequently in all prey identification procedures, though prey detection varied with procedure thus suggesting methodological bias. Prey species were identified using the Barcode of Life Database; the wingspan of prey consumed by M.

septentrionalis was smaller than that reported for other sympatric species.

My research demonstrates regional variation in bat activity, bat foraging, and prey occurrence across a gradient of forest disturbance. Conservation efforts should consider the importance of vegetation structure and plant species richness to sustain populations of both bats and their insect prey.

KEYWORDS: foraging ecology, predator-prey interactions, food habits, forest succession, Appalachia Unpublished dissertations submitted for the Doctor’s degree and deposited in the University of Kentucky Library are as a rule open for inspection, but are to be used only with due regard to the rights of the authors. Bibliographical references may be noted, but quotations or summaries of part may be published only with the permission of the author, and with the usual scholarly acknowledgments.

Extensive copying or publication of the dissertation in whole or in part also required the consent of the Dean of the Graduate School of the University of Kentucky.

A library that borrows this dissertation for use by its patrons is expected to secure the signature of each user.

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This project would not have been possible without the gracious assistance and cooperation of a number of groups. The National Council for Air and Stream Improvement, Inc., provided primary funding for this project, as well as field assistance.

The U.S. Forest Service provided invaluable assistance in the field. I would like to further thank the U.S. Forest Service, the state of Tennessee, and Plum Creek Timber Company who gave permission for this research to be conducted on their lands. Additional funding sources for my research included Bat Conservation International, McIntire-Stennis Funding, Sigma Xi, and the University of Kentucky Graduate School. Thank you to all these organizations and agencies for making my research possible.

I would like to thank my graduate committee. Dr. Charles Fox and Dr. John Obrycki provided insight and critique regarding the design of this project. Dr. James Harwood was instrumental in my research and his involvement expanded my interests in molecular tools and applications. Dr. Lacki provided critical insight into study design and has continued to be strong mentor. I am particularly grateful to my advisor, Dr. Lynne Rieske-Kinney, whose guidance and academic support was invaluable to this student. I am truly in her debt and thank her for all she has taught me.





I would also like to thank Dr. Rodney Cooper, Dr. Eric Britzke, and Dr. Eric Chapman, all of whom contributed to the development of my research. I am grateful for their aid, patience, and tutelage.

I would like to thank all the technicians who made this project possible: Josh Adams, Melanie Antonik, Allison Barlows, Elizabeth Carlisle, Drew Chalkley, Spencer Gravitt, Jessica Rasmussen, Rebecca Smith, and Joe Wong. I would particularly like to

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leadership and efforts in the field; this project would not have been possible without the efforts of both of them.

I would finally like to thank those who provided moral support over the past four years. To my fellow lab mates, Josh Adkins, Melanie Antonik, Paul Ayayee, Dr. Michael Baker, Erin Barding, Josh Clark, Dr. Tom Coleman, Dr. Rodney Cooper, Dan Cox, Joe Johnson, Aerin Land, Rachael Mallis, and Heather Spaulding, I am grateful for your assistance and friendship. To my friends from Arkansas: the late-nighters from Tech and those from Hatfield, thank you for always being there. To all the other friends that I have made in Kentucky over the past few years: the beer club at Marikka’s, the D&D guys, and other grad students in the Entomology and Forestry Departments, thank you for the good times and keeping me sane. Finally, to my girlfriend Liz and my family: Mom, Ronda, Tonya, and my grandparents, thank you for your love and support!

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Acknowledgements

List of tables

List of figures

Chapter one: Introduction

Statement of issue

Objectives and hypotheses

Management implications

Chapter two: Bat activity and insect occurrence varies along a gradient of disturbance.... 7 Introduction

Methods

Study areas and disturbance

Bat activity

Insect occurrence

Vegetation assessment

Analyses

Results

Bat activity

Insect occurrence

Discussion

Chapter three: Regional variation in the food habits of bats in Central Appalachia......... 40

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Methods

Study areas and field collection

Dissection procedure and analysis

Results

Discussion

Chapter four: DNA-based techniques allow a high resolution analysis of prey selection by a forest-dwelling bat (Myotis septentrionalis)

Introduction

Materials and methods

Study areas and field collection

Prey identification and comparison across techniques

Results

Discussion

Chapter five: Development of a COI library of forest Lepidoptera and identification of the prey of Myotis septentrionalis using tree-based cladistic analyses

Introduction

Materials and methods

Study areas and field collection

Screening fecal samples

Development of sequence library

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Results

Discussion

Appendices

Appendix A: Description of study areas and land use history

Appendix B: Diagram and photograph of passive-monitoring acoustic detection system for bats

Appendix C: Photographs of trapping methods used for surveying nocturnal aerial insects

Appendix D: Canonical correspondence analysis of the relationship between bat activity and vegetation attributes in Central Appalachia, 2007-2008

Appendix E: Canonical correspondence analysis of the relationship between Lepidoptera and vegetation attributes in Central Appalachia, 2007-2008.................. 103 Appendix F: Canonical correspondence analysis of the relationship between Coleoptera and vegetation attributes in Central Appalachia, 2007-2008

Appendix G: Canonical correspondence analysis of the relationship between Diptera and vegetation attributes in Central Appalachia, 2007-2008

Appendix H: Species checklist of forest Lepidoptera captured across a gradient of silvicultural disturbance in Central Appalachia, 2007-2008

Appendix I: Observations of mating behavior in Lasiurus borealis

References

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Table 2.1.

Variation in bat activity in Central Appalachia, 2007-2008

Table 2.2.

Canonical correspondence analyses of the relationships between bat activity and vegetation attributes and insect occurrence and vegetation attributes in Central Appalachia, 2007-2008

Table 2.3.

Variation in insects captured in blacklight traps in Central Appalachia, 2007Table 2.4. Variation in Diptera captured in malaise traps in Central Appalachia, 2007Table 3.1. Prey identified in fecal samples of bats in Central Appalachia, 2006-2008.... 50 Table 3.2. Percent volume (percent frequency) of Coleoptera and Lepidoptera consumed by Myotis septentrionalis across study sites in Central Appalachia, 2006-2008……….51 Table 4.1. Contribution of percent variation to overall χ2 scores among methods of identification

Table 4.2.

List of insect prey species identified by comparing COI sequences from the fecal samples of Myotis septentrionalis by comparison with BOLD

Table 4.3.

Comparison of total COI sequences from BOLD and GenBank databases..... 75 Table 5.1 Forest Lepidoptera collected across Central Appalachia and integrated into a COI sequence library

Table 5.2 Arthropods accessed from GenBank and integrated into a COI sequence library.

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Figure 2.1.

Synthesis of bat activity and insect occurrence across a gradient of forest disturbance in Central Appalachia, 2007-2008

Figure 3.1.

Variation in prey abundance across Central Appalachia, 2007-2008, as assessed by blacklight traps (Coleoptera and Lepidoptera) and malaise traps (Diptera).. 52 Figure 4.1. Prey volume identified in fecal samples of Myotis septentrionalis using morphological identification

Figure 4.2.

Frequency of occurrence of prey taxa in fecal pellets of Myotis septentrionalis across identification procedures.

Figure 4.3.

Lepidopteran prey abundance, as assessed by blacklight traps, compared with lepidopteran consumption, as assessed using BOLD

Figure 5.1.

Representative portion of the neighbor-joining tree constructed using potential prey taxa and fecal samples of Myotis septentrionalis collected in Central Appalachia, 2007-2008

Figure I.1. A copulating pair of Lasiurus borealis

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Beyond natural disturbance processes, human land use and resource extraction widely involves disturbance of forest vegetation and the broader ecosystem (Jones et al.

1999; Feldhake and Schumann 2005). In eastern North America, and Appalachia specifically, forests are fragmented and parceled; little remained unutilized by humans during the 19th and 20th centuries (Constanz 2000; Gragson and Bolstad 2006). Much of the land that was cleared for agriculture, and much of the land from which timber was harvested, has reverted to forestland (Jones et al. 1999; Gragson and Bolstad 2006). Of the human land use practices occurring in Appalachia, few are as prevalent as timber harvesting (Feldhake and Schumann 2005; Gragson and Bolstad 2006). Given this, an understanding of how silvicultural and other land-use practices impact forests is central to understanding the ecology and communities of forests in Appalachia and, more broadly, eastern North America.

A basic ecological understanding of vertebrate, invertebrate, and floral communities is fundamental to achieve goals for both ecological stewardship and for resource extraction (Guldin et al. 2007; Homyock and Haas 2009). Bats form an understudied but important assemblage of vertebrate predators in forests in North America (Fenton 2003; Brigham 2007). In recent years research on bat ecology has moved toward an investigation of how bats use their forest environments and how anthropogenic forces may affect them (Brigham 2007). Even so, relatively few studies have concurrently studied the land use and occurrence of bats and their insect prey base.

Consequently, the use of forest habitats by foraging bats, and how this habitat use is

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