GENETIC SUBDIVISION IN SCOTTISH RED DEER AND
NORTH AMERICAN WAPITI
by
PETER A. DRATCH
A thesis submitted in fulfilment of the requirement
for the degree of Doctor of Philosophy to the
University of Edinburgh
1983
E D I
0 ABSTRACT
To better understand the genetic structure of wildlife populations, an isozyme study of Scottish red deer and North American wapiti was conducted. Tissue samples from 943 individuals representing 28 localities were analysed by starch gel electrophoresis. The average amount of variation detected in a random selection of 28 to 34 enzyme loci did not significantly differ from that found in other mammals. There were significant differences between localities in both average heterozygosity and gene frequency.
The results suggest that population subdivision characterizes both
European red deer and North American wapiti. The polymorphism observed could be used in discriminating both populations and individuals. Nonetheless, the absolute genetic diversity was less than that measured between conspecific deer in Europe and North
America. By this electrophoretic measure, red deer and North American wapiti could be considered a single species, (Cervus elaphus L.) though not a panmictic population.
ii Declaration
This thesis has been composed by myself and it has not been submitted in any previous application for a degree. The work reported within was executed by myself, unless otherwise stated.
August 1983
iii ACKNOWLEDGEMENTS
Foremost, this project would not have been possible without the
help of stalkers and rangers in Scotland, as well as hunters and
biologists in the United States who aided in the collection of red
deer and wapiti samples. The co-operation of the Nature Conservancy
Council, the Forestry Commission, the Red Deer Commission and the
private estate-owners who permitted samples to be collected on their
land was also vital to the project.
I would like to thank Alan Archibald and John Bowman at the Animal
Breeding Research Organization in Edinburgh; Ulf Gyllensten and
Christina Reuterwall at the University of Stockholm; Mike Smith at the
Savannah River Ecology Laboratory and Fred Utter at the National
Marine Fisheries Service for providing not only laboratory facilities and materials, but also technical advice.
Financial assistance for the field work in the United States was supported by a grant from Sigma Xi, the Scientific Research Society.
Thanks to Susan Brotherstone and Jonathan Rasbash for advice in using the analysis of variance program (Harvey), and particular thanks to Andrew Sandford for advice and assistance in computing throughout the data analysis.
This study would never have been possible without the encouragement from afar of my parents, Claire and Joe Dratch.
Finally, I am indebted to my supervisors. To Dr. Trudy Mackay for helping me to see some of the complexities in the genetics of populations, and to Dr. W.E.S. Mutch for sound advice and warm support throughout my years in Edinburgh.
iv CONTENTS
TITLE PAGE j
ABSTRACT
DECLARATION
ACKNOWLEDGEMENTS iv
CONTENTS v, vi
CHAPTER 1 INTRODUCTION 1
1.1 Evolution and Classification 1 1.2 Eriropean red deer and North American wapiti 7 1.3 Applying population genetics 10 1.4 Electrophoretjc studies of deer 19 CHAPTER 2 MATERIALS and METHODS 24 2.1 Sampling sites, collection and storage 24 2.2 Electrophoretic procedure 27 2.3 Analysis of data 30 CHAPTER 3 DESCRIPTION OF POLYMORPHIC ENZYMES 33 3.1 Interpretation and notation 33 3.2 Isocitrate dehydrogenase 34 3.3 Superoxide dismutase 3.4 Glucose phosphate isomerase 38 3.5 Phosphoglucomutase 40 3.6 Malic enzyme 41 3.7 Mannose phosphate isomerase 43 3.8 Peptidase 43 CHAPTER 4 ELECTROPHORETIC VARIATION IN RED DEER 46 4.1 Scottish populations, 1979 46 4.2 Comparison with northern European populations 51 4.3 Retesting red deer from Scotland, 1980 60 CHAPTER 5 ENZYME POLYMORPHISM SAND QUANTITATIVE TRAITS 64 5.1 Neutrality vs. selection hypotheses 64 5.2 Local adaptation 65 5.3 Correlation with quantitative characters 67 CHAPTER 6 ELECTROPHORETIC VARIATION IN WAPITI 73 6.1 Why wapiti were tested 73 6.2 Heterogeneity in Northwest populations, 1981 75 6.3 Comparison with red deer 80 CHAPTER 7 DISCUSSION 90 CHAPTER 8 CONCLUSION 106 REFERENCES 109 APPENDICES 124
v CONTENTS (cont'd)
Appendix 1 Literature review of deer electrophoresis. 124 Appenxix 2 Buffer recipes and enzyme loci analysed. 135 Appendix 4a Heterogeneity tests. 137 Appendix 4b Collection sites, northern European red deer. 14 Appendix 4c Comparison of dendrogram and phylogenetic tree. 142 Appendix 4d Phylogenetic tree of Scottish red deer, 1980. 144 Appendix 5 Examples of analysis of variance, red deer 1980.146 Appendix 7 Poacher detection probability 149