University of Massachusetts - Amherst ScholarWorks@UMass Amherst Doctoral Dissertations 2014-current Dissertations and Theses Winter 2015 The effect of thermoregulation and roads on the movements and habitat selection of moose in Massachusetts David W. Wattles University of Massachusetts - Amherst, [email protected] Follow this and additional works at: http://scholarworks.umass.edu/dissertations_2 Recommended Citation Wattles, David W., "The effect of thermoregulation and roads on the movements and habitat selection of moose in Massachusetts" (2015). Doctoral Dissertations 2014-current. Paper 333. This Open Access Dissertation is brought to you for free and open access by the Dissertations and Theses at ScholarWorks@UMass Amherst. It has been accepted for inclusion in Doctoral Dissertations 2014-current by an authorized administrator of ScholarWorks@UMass Amherst. For more information, please contact [email protected]. THE EFFECT OF THERMOREGULATION AND ROADS ON THE MOVEMENTS AND HABITAT SELECTION OF MOOSE IN MASSACHUSETTS A Dissertation Presented by DAVID WILLIAM WATTLES Submitted to the Graduate School of the University of Massachusetts Amherst in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY February 2015 Department of Environmental Conservation Wildlife & Fisheries Conservation © Copyright by David W. Wattles 2015 All Rights Reserved THE EFFECT OF THERMOREGULATION AND ROADS ON THE MOVEMENTS AND HABITAT SELECTION OF MOOSE IN MASSACHUSETTS A Dissertation Presented by DAVID WILLIAM WATTLES Approved as to style and content by: _______________________________________ Stephen DeStefano, Chair _______________________________________ Todd K. Fuller, Member _______________________________________ Thomas Millette, Member ____________________________________ Curtice R. Griffin, Department Head Department of Environmental Conservation ACKNOWLEDGMENTS I would like to thank my advisor, Stephen DeStefano, for giving me the opportunity to work on this project, but mainly for the guidance, support, and friendship throughout the entire process. I cannot express my appreciation enough. I would like to thank my past and present committee members Todd K. Fuller, Thomas Millette, Robert D. Deblinger, and John E. McDonald, Jr., for guidance, suggestions, and constructive criticism that helped improve this project and dissertation. I want to recognize the hard work and friendship of Ken Berger in the difficult task of capturing moose and conducting fieldwork. I want to thank Kate for the love, support, and friendship through the happiness and frustrations of achieving my Ph.D. I thank my family for supporting me through the long process of completing this project. Thanks to all the friends and colleagues who made the work more enjoyable along the way. I want to recognize the individual state biologists and environmental police officers for helping on various aspects of the project, especially collaring and relocating problem animals. Thanks as well to the other field technicians and volunteers who assisted on the project. Special thanks to J. Finn for always having an open door and the generous giving of his time for help with statistical analysis. Thanks also to K. McCarthy, N. Rayl, B. Compton, M. Akresh, and B. Timm for statistical assistance on various aspects of the project. I want to thank the Massachusetts Division of Fisheries and Wildlife for providing long-term funding for moose research in Massachusetts through the Federal Aid in Wildlife Restoration program (W-35-R). The Massachusetts Department of Conservation and Recreation, U. S. Geological Survey, University of Massachusetts-Amherst, and Safari Club International have also provided funding and logistical support. iv ABSTRACT THE EFFECT OF THERMOREGULATION AND ROADS ON THE MOVEMENTS AND HABITAT SELECTION OF MOOSE IN MASSACHUSETTS FEBRUARY 2015 DAVID WILLIAM WATTLES, B.S., LAFAYETTE COLLEGE M.S., UNIVERSITY OF MASSACHUSETTS AMHERST Ph.D., UNIVERSITY OF MASSACHUSETTS AMHERST Directed by: Professor Stephen DeStefano Massachusetts, U.S.A. is located along the southern boundary of the geographic range of moose (Alces alces) in North America. This is an atypical environment for moose, because of its extremely high levels of human development and high year-round temperatures, which are possibly at the limits of moose physiological tolerances. I investigated the role of these two factors on moose movements and habitat selection to determine how human development of the landscape and temperature influence moose occupation of this extreme environment. In addition, the response of moose to these factors provides insights into the influence of development and temperature on individual fitness and population persistence. Thermal conditions in Massachusetts were consistently higher than the reported physiological tolerances of moose, and higher than parts of the range where high temperatures have negatively affected moose reproduction and survival. Moose greatly reduced their selection of open foraging habitat and increased selection for thermal shelters as temperatures increased above upper critical limits; this same pattern was apparent when comparing day (warmer) to night (cooler) activities. A strong transition in v the use of habitats occurred at dawn and dusk, which corresponded to peaks in hourly movement rates at these times, indicating a transition in state from foraging at night to bedding during the day. The ability of moose to adapt to this extreme thermal environment through thermoregulatory behaviors reveals the limitations of predicting species distributions based solely on theoretical temperature tolerances. Massachusetts has one of the highest rates of moose-vehicle collisions (MVCs) per-capita moose and resulting human fatalities. The majority (86%) of MVCs occurred on roads with high speed limits and traffic volumes, and occurred on these roads at much higher rates than would be predicted based on their availability on the landscape. The dense road network in Massachusetts had a strong negative effect on the movements and habitat selection patterns of moose. Moose avoided crossing roads and greatly reduced their use near roads. Road avoidance increased with increasing disturbance intensity associated with higher traffic volumes and busier times of day. Roadways reduced habitat availability and connectivity, were a major source of additive mortality for moose. vi TABLE OF CONTENTS Page ACKNOWLEDGMENTS ................................................................................................. iv ABSTRACT .........................................................................................................................v LIST OF TABLES ............................................................................................................. xi LIST OF FIGURES ......................................................................................................... xiv CHAPTER 1. THERMOREGULATORY BEHAVIOR OF MOOSE: A NORTHERN UNGULATE AT THE SOUTHERN EDGE OF ITS GEOGRAPHIC RANGE ..................................1 1.1 Abstract ..............................................................................................................1 1.2 Introduction ........................................................................................................2 1.3 Study Area .........................................................................................................5 1.4 Methods..............................................................................................................7 1.4.1 Assessing the Thermal Environment ..................................................7 1.4.1.1 Seasons ....................................................................................7 1.4.1.2 Ambient Temperatures versus Upper Critical Temperatures (UCTs) ...............................................................7 1.4.1.3 Heat Stress Index (HSI) ..........................................................8 1.4.1.4 Thermal Shelters .....................................................................9 1.4.2 Moose Habitat Use and Thermoregulatory Behavior .......................10 1.4.2.1 Capture and Marking ............................................................10 1.4.2.2 Temperature-Habitat Interactions .........................................10 1.4.3 Statistical Analysis ............................................................................12 1.4.3.1 Thermal Shelters ...................................................................12 1.4.3.2 Proportional Use of Cover Types .........................................13 1.4.3.3 Resource Selection Functions (RSF) ....................................13 1.5 Results ..............................................................................................................15 1.5.1 The Thermal Environment ................................................................15 vii 1.5.1.1 Ambeint Temperatures versus UCTs ....................................15 1.5.1.2 Heat Stress Index ..................................................................16 1.5.1.3 Thermal Shelters ...................................................................16 1.5.2 Moose Habitat Use and Thermoregulatory Behavior .......................17 1.5.2.1 Capture and Deployment of GPS Collars .............................17 1.5.2.2 Ambient Temperatures at GPS Locations.............................17 1.5.2.3 Proportional Habitat Use versus Ambient Temperatures............................................................................17