Evolution and Development of Wing Form, Body Size and Flight in Large- and Small-Bodied Fruit Bats (Artibeus Jamaicensis and Carollia Perspiciillata) Jason B
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University of Northern Colorado Scholarship & Creative Works @ Digital UNC Dissertations Student Research 8-1-2011 Evolution and development of wing form, body size and flight in large- and small-bodied fruit bats (Artibeus jamaicensis and Carollia perspiciillata) Jason B. Shaw Follow this and additional works at: http://digscholarship.unco.edu/dissertations Recommended Citation Shaw, Jason B., "Evolution and development of wing form, body size and flight in large- and small-bodied fruit bats (Artibeus jamaicensis and Carollia perspiciillata)" (2011). Dissertations. Paper 253. This Text is brought to you for free and open access by the Student Research at Scholarship & Creative Works @ Digital UNC. It has been accepted for inclusion in Dissertations by an authorized administrator of Scholarship & Creative Works @ Digital UNC. For more information, please contact [email protected]. UNIVERSITY OF NORTHERN COLORADO Greeley, CO The Graduate School THE EVOLUTION AND DEVELOPMENT OF WING FORM, BODY SIZE AND FLIGHT IN LARGE- AND SMALL-BODIED FRUIT BATS (ARTIBEUS JAMAICENSIS AND CAROLLIA PERSPICIILLATA) A Dissertation Submitted in Partial Fulfillment of the Requirements of the Degree of Doctor of Philosophy Jason B. Shaw College of Natural and Health Sciences School of Human Sciences Biological Education August, 2011 This Dissertation by: Jason B. Shaw Entitled: The Evolution and Development of Wing Form, Body Size, and Flight in Large- and Small-bodied Fruit Bats (Artibeus jamaicensis and Carollia perspicillata) has been approved as meeting the requirement for the Degree of Doctor of Philosophy in the College of Natural and Health Sciences in the School of Biological Sciences, Program of Biological Education Accepted by the Research Committee ____________________________________________________ Rick Adams, Ph.D., Chair ____________________________________________________ Stephen Mackessy, Ph.D., Committee Member ____________________________________________________ Mark Thomas, Ph.D., Committee Member ____________________________________________________ David Gilliam, Ph.D., Faculty Representative Date of Dissertation Defense . Accepted by the Graduate School ____________________________________________________________ Robbyn R. Wacker, Ph.D. Assistant Vice President of Research Dean of the Graduate School & International Admissions ABSTRACT Shaw, Jason B. The Evolution and Development of Wing Form, Body Size, and Flight in Large- and Small-bodied Fruit Bats (Artibeus jamaicensis and Carollia perspicillata). Published Doctor of Philosophy dissertation, University of Northern Colorado, 2011. Differences in developmental patterns important to diversification are produced through heritable variation of the onset/offset and timing of juvenile growth. As the size and shape of an organism changes during ontogeny, morphological, and behavioral components must adjust to accommodate proper function. This study explored the ontogenetic pathways of two closely related Phyllostomids differing in flight ability, body size, life history strategies, and developmental state at birth. We hypothesized that Artibeus jamaicensis and Carollia perspicillata will show ontogenetic differences that account for the diversification of morphological, body size, and behavioral patterns. Comparisons between the two species‟ flight development, growth rates, and morphometrics were made from day 1 to adult size (AJ n = 45, CP n = 25). Forearm length, mass, wing area, and wingspan were measured on a daily basis. Flight behavior was compared with juveniles being dropped from a 1 meter high roost from day 1 post- partum. Logistic growth equations were used to compare growth rates of all measured parameters and t-tests (p < 0.001) showed significant differences between the species of all measured variables. Muscle development in the pectoralis major was iii significantly different with A. jamaicensis having significantly more slow-twitch fibers. There were significant differences between the day of first flap (t-test, p = 0.01) and flight (t-test, p < 0.0001) with C. perspicillata achieving flight at 22 days and A. jamaicensis achieving flight 33 days post-partum. C. perspicillata was shown to be significantly more maneuverable than A. jamaicensis. Our data suggest that growth trends are significantly different with the more altricial A. jamaicensis developing faster than the more precocial C. perspicillata. Ontogenetic comparisons are important proxies when determining evolutionary diversification of closely related species. Data can be combined with phylogenetic information, providing possible mechanisms as to what factors could have influenced the divergence of closely related species. iv ACKNOWLEDGEMENTS I would like to thank the University of Northern Colorado and the School of Biological Sciences for giving me the opportunity to pursue my doctoral degree. Their financial support and teaching assistantships were crucial to me gaining experience as well as completing this experience and degree. I would like to thank my mentor Dr. Rick Adams for the continual support and guidance throughout my time here at UNC. His expertise and knowledge were invaluable, allowing for me to be successful in earning this degree. I would also like to thank my committee members Dr. Steve Mackessy, Dr. Mark Thomas, and Dr. Dave Gilliam for their support. Thanks to the members of the bat lab, Mark Hayes, Richard Carter, Emily Snode, Matt Willey, and Katelin Cravin, for their continual friendship and support. I would also like to thank Dr. John Hermanson for his guidance and ideas regarding the muscle work in this project. Without John I would have been lost. I would like to thank Dr. Tony Schountz for the financial support in keeping the bat colony. I would also like to thank the bats, without them this project would never have been accomplished. Lastly, I want to thank my family and friends whose support has been greatly appreciated. I would like to especially thank my wife Annette. Annette thanks for being patient and hanging in there for all these years, your support has been felt and I am indeed grateful for it. Sam, Joe, Tom, Phoebe, and Edmund, you have been awesome and I am thankful you were able to help me relax at the end of the day. v TABLE OF CONTENTS CHAPTER I. INTRODUCTION .......................................................................................1 Delimitations ..............................................................................................10 II. REVIEW OF LITERATURE ....................................................................11 Artibeus jamaicensis and Carollia perspicillata .......................................11 Ontogenetic Implications of Bat Ecology and Co-existence ....................14 The Evolution and Development of Wing Form and Body Size ..............19 Peramorphic Heterochrony ............................................................21 Paedomorphic Heterochrony .........................................................23 Allometric Comparisons ................................................................27 Evolution of Bats ...........................................................................28 Bat Development ...........................................................................32 Wing Morphology ..........................................................................34 Flight Behavior ..............................................................................37 Muscle Development .....................................................................39 III. METHODS AND MATERIALS ...............................................................43 Species and Housing ..................................................................................43 Ontogenetic Implications of Bat Ecology and Co-existence ....................45 Flight Development and Behavior .................................................45 Flight Maneuverability...................................................................47 The Evolution and Development of Wing Form and Body Size ..............49 Morphometrics ...............................................................................49 Bone Development.........................................................................51 Muscle Development .....................................................................53 Validity and Reliability ..............................................................................55 Statistical Analysis of Data ........................................................................55 vi CHAPTER IV. RESULTS ..................................................................................................58 Ontogenetic Implications of Bat ecology and Co-existence ......................59 Flight Development .......................................................................59 Fight Ability and Maneuverability.................................................62 The Evolution and Development of Wing Form and Body ......................73 Morphological Growth Patterns .....................................................73 Mass ...............................................................................................74 Forearm ..........................................................................................80 Wingspan .......................................................................................83 Wing Area and Length ...................................................................86 Digit Ossification .........................................................................104