Geographic Variation in the Bahamian Brown Racer, Alsophis Vudii A
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Geographic Variation in the Bahamian Brown Racer, Alsophis Vudii A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science Christina Wieg June 2009 © June 2009 Christina Wieg. All Rights Reserved. This thesis entitled Geographic Variation in the Bahamian Brown Racer, Alsophis Vudii by CHRISTINA WIEG has been approved for the Program of Environmental Studies and the College of Arts and Sciences by Matthew W. White Associate Professor of Biological Science Benjamin M. Ogles Dean, College of Arts and Sciences Abstract WIEG, CHRISTINA, M.S., June 2009, Environmental Studies GEOGRAPHIC VARIATION IN THE BAHAMIAN BROWN RACER, ALSOPHIS VUDII (48 pp.) Director of Thesis: Matthew M. White Nine meristic and fourteen morphometric characters were examined to determine geographic variation in the five subspecies of the insular snake species, Alsophis vudii. In addition, museum specimens from the closely derived species Alsophis cantherigerus were examined. Canonical discriminate analysis determined that, based on the morphological data, significant differences were observed among subspecies. However, significant differences were also observed between island populations. My results suggest that the current subspecies designations oversimplify the variation within the species and are not an adequate reflection of variation present among the island populations. Additional research is required in order to determine the causes of the observed variation within the species. Possible explanations for observed variation may include multiple colonization events or environmental variation among islands. Approved: Matthew M. White Associate Professor of Biological Sciences Dedication I dedicate this work to my husband, Rick Perkins. Without his constant support my goals would have never become a reality. I would also like to dedicate this work to my lovely daughter, Raven Perkins. I never knew unconditional love until I looked into her eyes. 5 Acknowledgments I would like to thank my committee members, Matthew W. White, Donald B. Miles, and James Dyer for their advice and assistance on this project. I would especially like to thank advisor Dr. White for his support, patience, and understanding during the development of this project. I would also like to thank committee member Dr. Miles for his assistance in the statistical analysis of the data and the use of his laboratory and Scott Moody for his guidance in the development of the methods. Special thanks to Gene Mapes and Cheryl Hanzel for their assistance. Funding for the project was provided by Masters in Environmental Studies Program, and a Graduate Student Senate Grant. Specimens were provided by the Natural History Museum at the University of Kansas and the Museum of Comparative Zoology at Harvard University. 6 Table of Contents Abstract ............................................................................................................................... 3 Dedication .......................................................................................................................... 4 Acknowledgments.............................................................................................................. 5 List of Tables ..................................................................................................................... 7 List of Figures .................................................................................................................... 8 Introduction ........................................................................................................................ 9 Systematics of Alsophis vudii ...................................................................................... 11 Materials and Methods ..................................................................................................... 14 Statistical Analysis ....................................................................................................... 15 Results .............................................................................................................................. 17 Morphological Variables ................................................................................................. 17 Meristic Variation ........................................................................................................ 23 Phylogenetic Analysis .................................................................................................. 27 Discussion ........................................................................................................................ 31 References ........................................................................................................................ 36 Appendix A: Institutional Abbreviations ......................................................................... 39 Appendix B. Sample size (N), means, and standard deviations (SD) for 14 morphological characters by subspecies, island, and sex. ............................................... 41 Appendix C: Sample size (N), means, and standard deviations (SD) for 5 meristic characters by subspecies, island, and sex ......................................................................... 45 7 List of Tables Table Page 1. Description of and codes used for meristic and morphometric variables. .................. 14 2. Mahalanobis distance for squared distance to subspecies for morpholmetric data .... 18 3. Results from the CDA using subspecies as a grouping variable. ............................... 18 4. The five canonical correlation structures of the morphometric characteristics .......... 20 5. Mahalanobis distance for squared distance to subspecies of A. vudii for meristic characters.. ....................................................................................................................... 23 6. The canonical correlation structure of the meristic characteristics from subspecies designations of A. vudii .................................................................................................... 24 7. The canonical structures of the meristic characteristics ............................................. 26 8 List of Figures Figure Page 1. Distribution of the five subspecies of Alsophis vudii: 1) A.v. vudii 2) A. v. aterrimus 3) A. v. picticeps 4) A. v. raineyi 5) A. v. utowanae.. ......................................................12 2. CDA by of morphometric data by island populations ...............................................18 3. Cluster analysis based on class means from the Canonical Variates Analysis of morphological variables of the island populations. ........................................................21 4. Cluster Analysis based on shape variation from the Canonical Variates Analysis of morphological variables of the island populations .........................................................22 5. CDA by of meristic data by island populations. .........................................................25 6. Cluster analysis based on class means from the Canonical Variates Analysis of meristic variables of the island populations……………………………………………..27 7. Phylogenetic analysis using the continuous maximum likelihood based on means for the morphological variables of the island populations of A. vudii………………………………………………….………………………………….28 8. Phylogenetic Analyisis of the continous maximum likelihood using the means of the Meristic data from the island popultations………………………………………………30 9 Introduction Geographic variation within species has been the focus of many studies (Endler, 1977; Pearson et al., 2002). Patterns of variation associated with environmental heterogeneity can allow the grouping of local populations into systematic categories. More importantly, studies of geographic variation can provide insight into the process of differentiation and speciation. The presence of geographic variation can be a potential isolating mechanism which may further the separation between the two populations (Mayr, 1999). The isolation of insular species results in greater limitation in gene flow and is often the result of founder effect (MacArthur and Wilson, 1967). As a result, insular species often express a greater degree of geographic variation among populations and between island populations and mainland populations. As a result, biologists have long had an interest in studying the geographic variation among populations of island species. A wide variety of geographic variation has been observed among insular reptile species. Much of the observed variations have resulted in subspecies designations. However, the usefulness of subspecies designation in describing the actual variation observed among a species has been questioned (Barrowclough, 1982). The designation of a subspecies portrays a phenotypic uniformity across an area of distribution. However, a frequently observed pattern of variation consists of a lack of concordance of clines in different characters (independent geographic variation), reoccurrence of characters in several geographic areas (polytopic subspecies), and every population differing in some character (microgeographic races) (Wilson and Brown, 1953). Therefore, subspecies 10 designations do not tend to accurately reflect the geographic variation present within a species. In addition, subspecies designations were often based on observed variation in a single specimen or variation as a result