PATTERNS AND CONSEQUENCES OF DISPERSAL IN ARCTIC CHAR (PISCES: SALVELINUS ALPINUS) FROM THE CANADIAN ARCTIC by Jean-Sébastien Moore BSc., McGill University, 2004 MSc., McGill University, 2007 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (ZOOLOGY) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) October 2012 © Jean-Sébastien Moore, 2012 i !"#$%!&$' Dispersal can have a multitude of ecological and evolutionary consequences that can be either positive or negative for population fitness and persistence. In this thesis, I describe patterns of dispersal in Arctic Char (Salvelinus alpinus), and I explored some of its consequences. I first examined the consequences of post-glacial dispersal for the distribution of genetic variation across the Canadian range of the species. MtDNA sequences and microsatellite markers provided evidence that the populations of Arctic Char currently inhabiting the Arctic Archipelago probably recolonized from a small glacial refugium, most likely located in ice-free areas of the Archipelago itself. I also presented evidence that two glacial lineages of Char (an Arctic lineage and an Atlantic lineage) probably hybridized post-glacially in the eastern Arctic. Finally, the importance of contemporary dispersal in redistributing genetic variation was illustrated by the fact that anadromous populations have greater within-population genetic diversity, and are less genetically differentiated, than landlocked populations. Second, I used a genetic assignment approach to study patterns of dispersal among populations distributed around Cumberland Sound, Nunavut. Estimates of dispersal rates varied extensively depending on the analysis method used, but all were relatively high compared to other salmonid species. I also found evidence that overwintering individuals have a greater propensity to disperse to non-natal habitats than individuals destined to spawn that year. The consequences of this behaviour for local adaptation among populations was examined using a population genetic model parameterized with estimates of gene flow obtained from microsatellite data. Third, I tested alternative hypotheses for the co-existence of sympatric migratory ecotypes in three lakes of southeast Baffin Island. Microsatellite data showed that the resident and anadromous components of the population are not genetically differentiated, suggesting that migratory behavior is not a genetically fixed trait. Together, the three parts of my thesis provide a general understanding of the patterns and consequences of dispersal for Arctic Char. Since dispersal will be crucial for the response of Arctic Char to environmental change, I conclude by discussing how my work can serve as a foundation for future work on the role of dispersal in adaptation to a changing Arctic. ii (%)*!&)' Chapter 2 was a project I conceived and conducted in collaboration with R. Bajno, J.D. Reist, and E.B. Taylor. The samples were collected by various people working under J.D. Reist over the years. I collected all the microsatellite data and some of the mtDNA sequence data. I performed all data analyses, and wrote the original manuscript. All three other collaborators provided advice and contributed revisions. Chapter 3 was a project I conceived and conducted in collaboration with L.N. Harris, R.F. Tallman, and E.B. Taylor. Adult samples used in this study were collected by various people working under R.F. Tallman over the years, and I collected all samples of juveniles according to the sampling design I conceived. L.N. Harris and I conducted all the laboratory work. I performed all data analyses and wrote the original manuscript. All three collaborators provided advice and contributed revisions. A version of chapter 4 has been submitted as: “T.N. Loewen, L.N. Harris & R.F. Tallman (in review) Genetic analysis of sympatric migratory ecotypes of Baffin Island Arctic Char Salvelinus alpinus: alternative mating tactics or reproductively isolated strategies?” T.N. Loewen and I collected the samples, and L.N. Harris and I conducted the laboratory work. I conducted all data analyses and wrote the original version of the manuscript. All three collaborators provided advice and contributed revisions to the manuscript. Permits for Arctic Char collection were obtained from Fisheries and Oceans Canada (Licenses to fish for scientific purpose S-09/10-1007-NU and S-10/11-1030-NU). 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Q(D! ,?;?<A'>)<A@;)=:C'((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((('9K9! 8#$#"! U(*4*-06+0:!;('4*+/04!##############################################################################################!"="! 8#$#$! X+))!'./)(*+/0-12!-6-7*-*+/0!&')7!F13*+3!3&-1!3/7'!>+*&!3)+,-*'!3&-0:'Y ! "=D! Q(K! &I=C):U'<AMH<\C'(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((('9KP! ")VI)=U<HGEL'((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((('9KX! vi !GGA:>)4'!N'#?GGIAMA:;H<L'MH;A<)HIC'B=<'&EHG;A<'D'(((((((((((((((((((((((((((((((((((((((((((('9X9! !GGA:>)4'"N'#?GGIAMA:;H<L'MH;A<)HIC'B=<'&EHG;A<'K'(((((((((((((((((((((((((((((((((((((((((((('9YD! vii +.#$',*'$!"+)#' Table 1.1 Some definitions of dispersal from recent reviews on the topic. ..................... 27! Table 1.2 Estimates of dispersal rates from tagging studies performed on Arctic Char, Salvelinus alpinus, and Dolly Varden, S. malma. ..................................................... 28! Table 1.3 Summary of evidence for increased propensity to disperse when overwintering in Arctic Char, Salvelinus alpinus, and Dolly Varden, S. malma. ............................ 28! Table 1.4 Summary of studies examining the marine migrations of Arctic Char, Salvelinus alpinus, and Dolly Varden, S. malma. ..................................................... 29! Table 2.1. Results of the historical demography analyses using neutrality tests (Fu’s F s and Tajima’s D) and mismatch distribution analysis on mitochondrial DNA sequence variation (D-loop) in Arctic Char, Salvelinus alpinus. .............................. 58! Table 2.2. Differences in genetic diversity (allelic richness and expected heterozygocsity) and average pairwise F between landlocked and anadromous populations of Arctic ST Char, Salvelinus alpinus, assayed at nine microsatellite DNA loci. ......................... 58! Table 3.1. Information regarding the sampling locations used in this study, and measures of genetic diversity at each location for anadromous Arctic Char (Salvelinus alpinus) assayed at fourteen microsatellite DNA loci. ........................................................... 91! Table 3.2. Results of discriminant function analysis testing for trait differences between philopatric and dispersing individuals of anadromous Arctic Char (Salvelinus alpinus) sample from Cumberland Sound. ............................................................... 93! Table 4.2. Details of the primers and PCR reactions used for each of the four multiplexes used to assess genetic variation of resident and anadromous Arctic Char (Salvelinus alpinus). .................................................................................................................. 117! Table 4.3. Summary statistics of microsatellite variation for each ecotype of Arctic Char (Salvelinus alpinus) per sampling location.. ........................................................... 118! Table 4.4. Semi-matrix of pairwise F (Weir & Cockerham, 1984) values between all ST sampling locations and ecotypes of Arctic Char (Salvelinus alpinus). ................... 119! Table 4.5. Results of the analysis of molecular variance (AMOVA) examined among sampling locations and ecotypes of Arctic Char (Salvelinus alpinus). ................... 119! viii Table A.1. Name and geographic coordinates of sampling locations where specimens of Arctic Char (ARCH, Salvelinus alpinus) and Dolly Varden (DVCH, Salvelinus malma) used for mtDNA sequencing were collected. ............................................ 171! Table A.2. Sampling location information for the samples used in the microsatellite analysis of Arctic Char (Salvelinus alpinus) ........................................................... 174! Table A.3. Frequency of haplotypes of Arctic Char (Salvelinus alpinus) in each sampling locations. ................................................................................................................. 175! Table A.4. Summary population statistics for the microsatellite analysis across nine loci in Arctic Char (Salvelinus alpinus). ........................................................................ 179! Table B.1. Details of the primers and PCR reactions used for each of the 4 multiplexes. ................................................................................................................................. 183! Table B.2. Heterozygosity (observed and expected) and F (Weir & Cockerham 1984) IS per locus in samples of Arctic Char (Salvelinus alpinus) ....................................... 185! Table B.3. Pairwise F (Weir and Cockerham 1984) between each Arctic Char ST (Salvelinus alpinus) sample used in the study. ....................................................... 189! Table B.4. Summary of sibship reconstruction among samples of juvenile Arctic Char (Salvelinus alpinus) using the program COLONY ................................................. 192! ix +.#$',*'*./0%)#' Figure 1.1. Arctic Char, Salvelinus alpinus. The drawing is based on an anadromous specimen in spawning colours from Anderson Bay Lake in the Cambridge Bay Region of Nunavut, Canada. ..................................................................................... 30! Figure 1.2 Map of the northern hemisphere showing the Holarctic distribution of Arctic Char, Salvelinus alpinus, in dark shaded areas surrounded by dashed lines. ........... 31! Figure 2.1. Map of the Northern Hemisphere showing the distribution and names of the five major mitochondrial DNA lineages of Arctic Char (Salvelinus alpinus) identified in the Holarctic phylogeography study of Brunner et al. (2001). ............. 59! Figure 2.2. Maximum likelihood phylogenetic tree of Arctic Char (Salvelinus alpinus) mtDNA haplotypes generated using GARLI (Zwickl 2006). ................................... 60! Figure 2.3 (Next page) Map showing the geographical distribution of the two mtDNA lineages identified in North American Arctic Char (Salvelinus alpinus) and Dolly Varden (S. malma). ................................................................................................... 61! Figure 2.4. Results of the pairwise mismatch analysis done on the mtDNA data for two lineages of Arctic Char (Salvelinus alpinus). ........................................................... 63! Figure 2.5. Neighbour-joining tree of Cavalli-Sforza’s chord measure genetic distance from nine microsatellite loci assayed in samples of Arctic Char (Salvelinus alpinus). ................................................................................................................................... 64! Figure 2.6. Variation in expected heterozygosity as a function of waterway distance from a putative glacial refugium (Banks Island) in samples of Arctic Char (Salvelinus alpinus) assayed at nine microsatellite DNA loci. .................................................... 65! Figure 2.7. Results of the STRUCTURE analysis based on the most likely (sensu Evanno et al. 2005) number of genetic cluster: K = 4 in samples of Arctic Char (Salvelinus alpinus) assayed at nine microsatellite DNA loci. .................................................... 66! Figure 2.8. Relationship between pairwise genetic distance measured as linearized F ST (Rousset 1997) and geographical distance (km) in samples of Arctic Char (Salvelinus alpinus) assayed at nine microsatellite DNA loci. ................................. 67! x
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