INFORMATION TO USERS This manuscript has been reproduced from the microfilm master- UMI films the text directly from the original or copy submitted. Thus some thesis and dissertation cupies are in typewriter face, while others may be from any type of computer printer. The quality al this I.produc(kn k dependent upon th. quality of the copy submW. Brdcen or bldistinct priM cokred or poor quality iltustralions and photographs, print Meedthrough, substandard margins, and improper alignment can adversely afkt reproduction. In the unlikely event that the author did not send UMI a cornpkte maiptand 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, charto) are reproduced by S8dl*oning the original, beginning at the upper left-hand comer and continuing from left b right in equal sections with small overlaps. Photographs included in the original manudpt hwe been mpoducad xerographically in this copy. Higher quality C x 9" black and white photographic prints are available for any photographs or illustratio~appearing in this cwfw an additional charge. Contact UMI directly to der. Bell & Howell Information and Learning 300 Norlh Zeeb Road, Ann Arbor, MI 48106-1346 USA NOTE TO USERS This reproduction is the best 'copy available National Library BibliitMque nationale 1+1 of,,, du Canada Acquisitions and Acquisitions et Bibliographic Services services bibliographiques 39s weuiiqtm street 395, rue Wellington Ottawa ON KIA ON4 OttawaON KIA ON4 Canada Canada The author has granted a non- L'auteur a accorde une licence non exclusive licence allowing the exclusive pernettant a la National Library of Canada to Bibliotheque nationale du Canada de reproduce, loan, distriiute or sell reproduke, preter, distrr-buer ou copies of this thesis in microform, vendre des copies de cette these sous paper or electronic formats. la forme de microfiche/fb, de reproduction sur papier ou sur format electronique. The author retains ownership of the L7auteurconserve la propriete du copyright in this thesis. Neither the droit d'auteur qui protege cette these. thesis nor substantial extracts &om it Ni la these ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent &e imprimes reproduced without the author7s ou autrement reproduits sans son permission. autorisation. INTEL AND INTRASPECIFIC PHYLOGEOGRAPHY OF NORTH AMERICAN ODOCOLLEUS DEER BASED ON MITOCHOND~DNA SEQUENCES by O Annette D. Greenslade, B.Sc. @onours) A thesis submitted to the School of Graduate Studies in partial llfillment of the requirements for the degree of Master of Science Department of Biology Memorial University of Newfoundand July 1998 St. John's Newfoundland ABSTRACT Phylogenetic analysis ofmitochondrid DNA (mtDNA) cytochrome b sequences identified four major mtDNA genotype assemblages among popuIations of Odocoileus in western North America. These assemblages correspond to northwestern black-tailed deer, southwestern mule deer, northwestern mule deer, and southwestern white-tailed deer. Approximate times of the divergence of these assemblages, calculated on the basis of pairwise nucleotide &quence divergence estimates, were used to construct a model of Odocoileus evolution in western North America. According to this model, black-tailed deer represent the ancestral OdocoiZ~mtDNA lineage. Mule deer and white-tailed deer diverged more recently in palaeontologicaI history. The southwestern white-tailed deer mtDNA genotype assemblage is more closely related to the mule deer mtDNA assemblages than it is to the southeastern white-tailed deer Lineage. This suggests that the southwestem white-tailed deer mtDNA lineage may have been effectively replaced by mule deer mtDNA through relatively recent hybridbation between the two species. Microgeographic analysis of deer fiom southern Alberta revealed a considerable degree of geographic structuring of mtDNA sequence genotypes, as did qualitative analysis of the geographic distribution of mule deer and white-tailed deer mtDNA genotypes in CalSornia and Alberta. The phylogeographic structure may be maintained by philopatry of the female (or family) social units of Odocoileus deer, despite their potentially high vagility . Gratitude is extended as appropriate to my supervisory committee (S. M. Carr, D. I. Innes, and E. H. Miller), my colleagues, and those researchers who provided deer samples: S. Lingle collected samples fkom the McIntyre Ranch near Mapth, Alberta, and the Waterton Lakes National Park, Alberta; M- Cronin (LGL Alaska) provided samples from southern California; C. Strobeck (University of Alberta, Edmonton) provided samples fiom British Cohunbia, Manitoba, Sasketchewan, and Ontario; and S. Carr &Iemorial University of Newfoundlaud, St. John's) provided samples from western North America. DNA extractions of the samples from S. Lingle were carried out by J. Coffin et 01. of C. Strobeckrs research lab. S. Lingle also provided a UTM map showing the sampling location of most of the deer she collected; Figure 2 is based on this map. Many thanks to M. Dennison and A. Lynch (both at The Royal Ontario Museum) for help with preparing the maps, to 0. Haddrath (The Royal Ontario Museum) for advice with phylogenetic analyses, and to A. J. Baker me Royal Ontario Museum) for reading an earlier draft of my thesis and offering invaluable advice (thanks for the encouragement). Thank you to my supervisory committee for reading drafts of the thesis and for offering constructive criticism. Thanks also to my supervisor for help with using MEGA, REAP, and PAUP,and with developing a model of 0docoUeu.s evoIution. ALSO sincere gratitude is extended to G. Kemy, Graduate secretary of the Biology Department, for help with all the administrative details. TABLE OF CONTENTS Page - Abstract .................................................................................................................. n- Acknowledgements ................................................................................................. rn-.- Table of Contents.................................................................................................... v . List of Tables.-...................~................................................................................... .vu- List of Figures ........................................................................................................ wu--. Introduction.............................................................................................................. 1 Materials and Methods............................................................................................ 13 Results.,.,.......-......................................................................................................... 23 Microgeographic ana&sis ofdeer#om Magruth and Waterton Lakes National Park.. .................................................................. 23 New rntDNA sequence genowes identtperi:....................................................... 3 0 Comparison ofpreviously reported mtDNA sequence genotpes ond new &ta ..................................................................................... 32 Pattern of mcleotide substitution among Odocoileus mtDNA sequence genows..~..~~~..........................................~.............................. 33 Phylogenetic analyses of mtDNA sequence genotypes ........................................ 44 Discussion.-..,-......,....-....-..-... .................................................................................. 78 Palaeontological record of Odocoileus...................-........................................ .--78 Evohtionory history of Odocoileus bared on morphological and mtDNA sequence &ta ................. RR-R-RRRRRRRR--RRRRRRR~ .......................................... 79 Glacial refirgia, stochastic lineage sorting d introgressive hybridization of mtDNAA-............................................................. ........................86 Gene trees and species trees................................................................................ 91 Phylogeogap?iy of Odocoileus~..~~....................................................................... 93 References................................................................................................................ 97 LIST OF TABLES Table 1 Mitochondria1 DNA genotypes found in deer fiom the Mctntyre Ranch near Magrath, Alberta, and Waterton Lakes National Patk, Albem..............-........-.... d 2 Pairwise haplotype divergence estimates for the mtDNA genotypes identified in mule deer and white-tailed deer fkm the McIntyx Ranch near Magrath, Alberta, and Waterton Lakes National Park, Alberta.... ................................................................................................ 28 3 Indices of haplotype (h) and nucleotide (x) diversity within populations of mule deer and white-tailed deer fiom the McIntyre Ranch near Magrath, Alberta, and Waterton Lakes National Park, Alberta. ................................................ 29 4 Correspondence of 401-nucleotide sequence genotype with 307-nucleotide genotype (Carr and Hughes, 1993; Hughes, 1990; Hughes and Cam, 1990), and with RFLP- genotype (Cronin, 1991; Cronin and Bleich, 1995).............................. 3 1 5 Frequencies and distributions of variant nucleotides in 0docoileu.s within the 40 1-nucleotide cytochrome b sequence data. ....................... 43 6 Minimum and