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U·M·I University Microfilms International a Bell & Howell Information Company 300 North Zeeb Road INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copysubmitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sectionswith small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. U·M·I University Microfilms International A Bell & Howell Information Company 300 North Zeeb Road. Ann Arbor. M148106-1346 USA 313:761-4700 800/521·0600 Order Number 9325033 Character release in the endangered Hawai'ian hoary bat, Lasiurus einereus semotus Jacobs, David Steve, Ph.D. University of Hawaii, 1993 Copyright @1993 by Jacobs, David Steve. All rights reserved. U·M-I 300 N. ZeebRd. Ann Arbor, MI48106 CHARACTER RELEASE IN THE ENDANGERED HAWAI'IAN HOARY BAT, LASlURUS CINEREUS SEMOTUS. A DISSERTATION SUBMITIED TO THE GRADUATE DMSION OF THE UNIVERSllY OF HAWAI'IIN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY MAY 1993 By David S. Jacobs Dissertation Committee: Leonard Freed, Chairperson Sheila Conant Michael Hadfield Stephen Palumbi Rebecca Cann © Copyright 1993 by David S. Jacobs iii To my Family and the Village that raised me. iv ACKNOWLEDGEMENTS I am indebted to F. Howarth and A Ziegler for providing unpublished manuscripts and information on work in progress, to C. Koehler and M. Kalcounis for providing wing tracings of the North American hoary bat, and to the Bernice P. Bishop Museum (Honolulu) and the Museum of Southwestern Biology (New Mexico) for providing bat skulls. This work was supported by grants from the Audubon Society, the state Division of Forestry and Wildlife, Hawai ' i, U. S. Fish and Wildlife, Hawai-Ian Telephone Co. and Alvin Y. Yoshinaga. v ABSTRACT The insectivorous Hawaiian hoary bat, Lasiurus cinereus semotus, is an endemic subspecies of the North American hoary bat, L. c. cinereus. This study investigated morphological and ecological divergence of the island population, with emphasis on the potential for character release. Divergence in the Hawai \ian hoary bat involved characters related to flight and feeding. The Hawai \ ian bat has undergone a 45% reduction in body size with allometric responses in size of its wings. These changes decreased wing loading (ratio of mass to wing area), without altering the high aspect ratio (wingsparr'jwing area) of the ancestral species, thereby permitting slower and more maneuverable flight near vegetation. High aspect ratio enables the bat to retain the fast and efficient flight of its ancestor in open areas. This increased flexibility in flight behavior has allowed the Hawai\ ian bat to expand its foraging habitat to include the open habitats of the North American hoary bat and the closed habitats of putative competitors of the North American hoary bat. There has been a relative increase in skull length of the Hawai \ ian bat, with non-allometric increases in the gape of the jaws, height of the coronoid process, and size of the masseter muscle. This gives the jaw more crushing power for more efficient processing of large and hard-bodied prey. In the open habitat the Hawai \ ian bat fed predominantly on moths as does the North American hoary bat. In the closed habitat the Hawai \ ian bat fed predominantly on beetles as do putative competitors of the North American hoary bat. The morphological and vi ecological changes are consistent with the assumptions and prediction of the character release hypothesis. However, uncertainties remain about the historical aspects of the assumptions. In addition, the interaction between flight speed, echolocation, and insect prey was investigated. Apparent prey selection by fast-flying bats in open habitats was the result of a decrease in detectability of smaller prey. Bats responded to changes in prey density by altering the pursuit component of handling time. vii TABLE OF CONTENTS Acknowledgements ~ Abstract. Vl List of Tables ~ ix List of Figures .x Chapter 1: Introduction 1 Chapter 2: Distribution and Abundance of the Endangered Hawai \ ian Hoary Bat, Lasiurus cinereus semotus, on the island of Hawai \i. 15 Introduction 15 Materials and Methods 16 Results 18 Discussion 24 Chapter 3: Morphological Divergence in the Hawai \ ian Hoary Bat, Lasiurus cinereus semotus 27 Introduction 27 Methods 32 Results 35 Discussion 41 Chapter 4: Foraging Strategies and Prey Selection in the Hawai \ ian Hoary Bat, Lasiurus cinereus semotus 49 Introduction 49 Methods 54 Results 58 Discussion 62 Chapter 5: Character Release in the Hawai\ ian Hoary Bat, Lasiurus cinereus semotus 68 Introduction 68 Methods 72 Results 73 Discussion 75 Appendix: Museum Skull Specimen Numbers and Standard Errors of Skull Measurements 125 Literature Cited 126 viii LIST OF TABLES 2.1. Maximum Number of Bats at Each Foraging Site 81 2.2. Temporal Distribution of Bats 82 2.3. Vegetation Associated with Each Foraging Site 83 3.1 Body Mass and Forearm Data 84 3.2. Skull Dimensions 85 3.3. Allometric Exponents of Skull Measurements 86 3.4. Wing Parameter Data Wingspan, Wing Area, Wing Loading 87 3.5. Allometric Exponents of Wing Parameters 88 3.6. Wing Parameter Data Aspect Ratio and Tip Shape Index 89 4.1. Size of Dietary Items 90 4.2. Percentage of Artificial Prey Chased 91 4.3. Bat Acoustic Responses to Artificial Prey 92 4.4. Percentage of Capture Attempts on Artificial Prey 93 A.l. Museum Skull Specimens 125 A2. Standard Errors for Repeated Measures l25 ix LIST OF FIGURES Figure 2.1. Distribution of Bat Observations 94 2.2. Bat Activity and Insect Biomass 95 3.1. Wing Design for Closed and Open Habitats 96 3.2a. Skull Measurements 97 3.2b. Skull Measurements 98 3.2c. Skull Measurements 99 3.3. Definitions of Wing Parameters and Measurements Measurements 100 3.4. Plot of Mass against Skull Length 101 3.5. Plot of Mass Against Wingspan 102 3.6. Plot of Mass Against Wing Area 103 3.7. Plot of Mass Against Wing Loading 104 3.8. Plot of Mean Mass Against Mean Wing Loading 105 4.1a. Insect Orders in Bat Diet and Light Traps Closed Habitat (Ocean View), August 1991 106 4.1b. Insect Orders in Bat Diet and Light Traps Closed Habitat (Ocean View), September/October 1991.. 107 4.1c. Insect Orders in Bat Diet and Light Traps Closed Habitat (Ocean View), June 1992 108 4.1d. Insect Orders in Bat Diet and Light Traps Closed Habitat (Ocean View), August/September 1992 109 4.2. Insect Orders in Bat Diet and Light Traps Closed Habitat (Red Cinder Road) 110 x LIST OF FIGURES (contd.) Figure 4.3a. Insect Orders in Bat Diet and Light Traps Open Habitat (Pohakuloa), June 1992 111 4.3b. Insect Orders in Bat Diet and Light Traps Open Habitat (Pohakuloa), July 1992 112 4.3c. Insect Orders in Bat Diet and Light Traps Open Habitat (Pohakuloa), September 1992 113 4.4a. Proportion of Bats Eating each Insect Order Closed Habitat (Ocean View), August, 1991.. 114 4.4b. Proportion of Bats Eating each Insect Order Closed Habitat (Ocean View), September/October 1991.. 115 4.4c. Proportion of Bats Eating each Insect Order Closed Habitat (Ocean View), June 1992 116 4.4d. Proportion of Bats Eating each Insect Order Closed Habitat (Ocean View), August, 1991.. 117 4.5. Proportion of Bats Eating each Insect Order Closed Habitat (Red Cinder Road), August/September 1992 118 4.6. Proportion of Bats Eating each Insect Order Open Habitat (Pohakuloa) 119 4.7. Size Distribution of Insect Orders Closed Habitat (Ocean View) 120 4.8. Size Distribution of Insect Orders Closed Habitat (Red Cinder Road) 121 4.9. Size Distribution of Insect Orders Open Habitat (Pohakuloa) 122 5.1. Size Range of Prey Items 123 5.2. Proportions of Moths and Beetles in the Bat Diet. 124 xi CHAPTER 1 INTRODUCTION ''/ should like to know whether the case ofendemic bats in islands struck you: it has me especially; perhaps too strongly" (Charles Darwin in a letter to J. D. Hooker, dated March 15, 1859 -Burkhardt and Smith, 1991). DIVERGENT EVOLUTION IN INSULAR POPULATIONS Divergent evolution occurs when the descendants of an ancestral population develop different heritable phenotypes. Anagenesis is a type of divergent evolution that involves change within the same lineage over evolutionary time without speciation (Futuyma, 1986, pp.403). Such change may include geographical variation of a single character (e.g., beak depth) within the same species. Cladogenesis, on the other hand, involves change between an ancestral population and some of its genetic lines that become isolated. One population splits to give rise to two or more divergent populations that may become different species. Whereas anagenesis involves a gradual and directional change within a lineage, cladogenesis may involve rapid change as different gene pools encounter different environments. The distinction between anagenesis and cladogenesis does not imply that the two processes necessarily occur in isolation of each other.
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