Population Differentiation in the European Roe Deer by Non-Metric Skull Traits in Germany
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Population differentiation in the European roe deer (Capreolus capreolus) by non-metric skull traits in Germany D i s s e r t a t i o n zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. net) vorgelegt der Naturwissenschaftlichen Fakultät I Biowissenschaften der Martin-Luther-Universität Halle-Wittenberg von Frau Atefeh Rahbar geb. am: 16.09.1976 in Tehran, Iran Gutachter /in 1. Dr. habil. Wolf Rüdiger Große 2. Prof. Dr. Michael Stubbe 3. Prof. Dr. J. Zima Halle (Saale), 2009 Population differentiation in the European roe deer (Capreolus capreolus) by non-metric skull traits in Germany Abstract The non-metric skull divergence between populations of the roe deer, Capreolus capreolus (Linnaeus, 1758) were studied using 786 complete skulls (crania and mandibles) from three wildlife research areas in Germany which were from Hakel, Fallstein and Darss. A total of 292 male and 494 female were scored for 56 cranial non-metric traits. The dependence of incidence of characters on age, sex and correlation between variants was studied. Four traits were found to have sex dependence and eighteen traits displayed a dependence on age. Of 1540 calculated correlation coefficients between the variants only 140 (9.1%) were significantly different from zero at p < 0.05. But only 4 out of 140 (2.86% and 0.26% out of all) had a correlation value, equal or more than 0.3 that were excluded. After deleting all variants correlated with one another and dependence on sex and age, further analysis was based on the frequency of 34 traits. The Mean Measures of Divergence (MMD) was used to express the interpopulation differences. The MMD values were calculated between pair populations Fallstein and Darss; Fallstein and Hakel; and Hakel and Darss as 0.04671, 0.01572 and 0.04131 respectively and all were highly significant at P < 0.001. The measure of uniqueness (MU) was calculated for each sample as the sum of its epigenetic distance (MMD) and they were 0.08802, 0.06243 and 0.05703 for Darss, Fallstein and Hakel respectively. The cluster analysis the MU values proved existence two main clusters. The first one consists of two samples (Hakel and Fallstein) with low differentiation, contrary to a distinctly separated position of the sample Darss which from Baltic coast of Germany. i ACKNOWLEDGMENTS This project completion would not have been possible without the help and support of many people. First and foremost, I would like to thank Prof. Dr. Herman Ansorge for his assistance in helping me to create this project over last three years and revising my thesis and being a person to discuss methodical issues. I am also appreciative for giving his computer program to analyze epigenetic traits. I would also like to thank my advisor Dr. habil. Wolf Rüdiger Große, Prof. Dr. Michael Stubbe, Prof. Dr. Herman Ansorge. Their guidance helped me write my thesis and complete my studies. I would also like to thank Stubbe family (Prof. Dr. Hans Stubbe, Dr. Christoph Stubbe and Prof. Dr. Michael Stubbe) for allowing me to access their skull collections. I especially like to acknowledge again Prof. Dr. Michael Stubbe who offered this thesis and gave his papers about study areas to me and supported me whenever I needed advice. I would also like to thank Prof. Dr. J. Zima for doing kindness to me and to post his papers from Bruno University, Czech. This was a great help. The help of my husband, Mr. Faridoddin Rezazadeh, with statistic analysis was greatly valued. I am also grateful to Prof. Dr. Harwig Prange for invitation me to Martin-Luther university Hall- Wittenberg to pursue my graduate study (as a PhD student). Additionally, I am appreciative of Dr. Stephan Schäffer for help me to find equivalent names of some flora the areas under study. The assistance of Mr. Ronald Müller was also greatly appreciated. I want to thanks Ms. Heike Brünsdorf, librarian of the Institute, for her help and guide to search and to find needed literatures. Finally, I would also like to express my extreme gratitude to Dr. Joachim Wussow for all of his endless support and spiritual help in throughout the course of this study. Without his support and patience, I was not really able to do my research and it may never have been completed. ii List of Contents List of Contents........................................................................................................................ iii List of Figures ............................................................................................................................v List of Tables.............................................................................................................................vi Preface ..................................................................................................................................... vii 1. Introduction .....................................................................................................................1 1.1. Taxonomy, morphology, geographic range and habitat use of roe deer...........................1 1.1.1. Taxonomy ...................................................................................................................1 1.1.2. Morphology ................................................................................................................1 1.1.2.1. Characterization of the Genus Capreolus .............................................................1 1.1.2.2. The Siberian roe deer- Capreolus pygargus (Pallas, 1771) ..................................2 1.1.2.3. The European roe deer- Capreolus capreolus (Linnaeus, 1758) ..........................2 1.1.3. Geographic range........................................................................................................3 1.1.4. Habitat use ..................................................................................................................4 1.2. Study areas ........................................................................................................................4 1.2.1. Darss ...........................................................................................................................4 1.2.1.1. Location................................................................................................................4 1.2.1.2. Protective status....................................................................................................6 1.2.1.3. Coastal Geology ...................................................................................................6 1.2.1.4. Climate .................................................................................................................6 1.2.1.5. Flora......................................................................................................................6 1.2.2. Hakel...........................................................................................................................7 1.2.2.1. Location................................................................................................................7 1.2.2.2. Protective status....................................................................................................7 1.2.2.3. Climate .................................................................................................................7 1.2.2.4. Flora......................................................................................................................7 1.2.3. Fallstein.......................................................................................................................8 iii 1.2.3.1. Location................................................................................................................8 1.2.3.2. Protective status....................................................................................................8 1.2.3.3. Climate .................................................................................................................8 1.2.3.4. Flora......................................................................................................................8 1.3. Morphological study .........................................................................................................9 1.3.1. Genetics aspect of morphological study on roe deer ................................................11 1.3.2. Non- metric method as a morphological tool ...........................................................12 1.3.3. History of non-metric method...................................................................................14 1.3.4. Forms for the occurrence of non-metric character....................................................16 1.3.5. Variable heritability of non-metric characters ..........................................................17 1.3.6. Non-metric variability and population divergence ...................................................19 1.3.7. Population divergence in Roe deer ...........................................................................20 1.4. Study objectives ..............................................................................................................20 2. Material and Method ....................................................................................................21 2.1. The list of non-metric traits.............................................................................................22 2.2 Preliminary tests of non-metric traits...............................................................................27