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Molecular Genetic Status of Aleutian Canada Geese from Buldir and the Semidi Islands, Alaska ’

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Molecular Genetic Status of Aleutian Canada Geese from Buldir and the Semidi Islands, Alaska ’ Tlze Condor 102: 172-180 0 The Cooper Ornithological Society 2000 MOLECULAR GENETIC STATUS OF ALEUTIAN CANADA GEESE FROM BULDIR AND THE SEMIDI ISLANDS, ALASKA ’ BARBARAJ. PIERSON,JOHNM. PEARCEAND SANDRA L. TALBOT United StatesGeological SurveyBiological ResourcesDivision, Alaska Biological ScienceCenter, 1011 E. Tudor Rd., Anchorage,AK 99503, e-mail: [email protected] GERALD E SHIELDS* Institute of Arctic Biology, Universityof Alaska Fairbanks, Fairbanks, AK 99775.0180 KIM T. SCRIBNER Departmentof Fisheries and Wildlife, Michigan State University,East Lansing, MI 48824.1222 Abstract. We conductedgenetic analysesof Aleutian Canada Geese (Branta canadensis leucopareia) from Buldir Island in the western Aleutians and the Semidi Islands in the eastern portion of their breeding range. We compared data from seven microsatellite DNA loci and 143 basepairs of the control region of mitochondrialDNA from the two populations of Aleutian Canada Geese and anothersmall-bodied subspecies,the Cackling CanadaGoose (B. c. minima) which nests in western Alaska. The widely separatedisland-nesting Aleutian geese were genetically more closely related to each other than to mainland-nestingsmall- bodied geese. The populations of Aleutian geese were genetically differentiated from one another in terms of mitochondrial DNA haplotype and microsatellite allele frequencies, suggestinglimited contemporary gene flow and/or major shifts in gene frequency through genetic drift. The degree of population genetic differentiation suggeststhat Aleutian Canada Goose populations could be consideredseparate management units. There was someevi- dence of population bottlenecks,although we found no significant genetic evidence of non- random mating or inbreeding. Kev words: Aleutian Canada Geese,bottlenecks, Branta canadensisleucopareia, genet- ics, &rosatellites, mtDNA. INTRODUCTION been released due to the island’s remotenessand Three small-bodied subspeciesof Canada Geese difficult terrain (Byrd and Woolington 1983). (Branta canadensis) breed in Alaska: Taverner’s Restricted distribution and low remnant popu- Canada Geese (B. c. taverneri) nest predomi- lation numbers prompted the United States Fish nantly in the interior portion of the state and and Wildlife Service to list Aleutian Canada north of the Brooks Mountain range in northern Geese as endangered in 1967. Alaska; Cackling Canada Geese (B. c. minima) Hatch and Hatch (1983) subsequently de- nest predominantly in coastal regions of western scribed “Aleutian-like” Canada Geese from Kil- and southwestern Alaska; and Aleutian Canada iktagik of the Semidi Islands, and Bailey and Geese (B. c. leucopareia) nest only on a few Trapp (1984) discovered a population of “Aleu- small islands (Buldir, Nizki, Alaid, Agattu, Little tian-like” Canada Geese on Chagulak Island Kiska, Chagulak, Amukta) in the Aleutian chain (Fig. 1). Conventional morphometric studies and on the Semidi Islands (Kiliktagik and An- were inconclusive regarding the taxonomic and owik) southwestof Kodiak Island. reproductive status of these populations in rela- Aleutian Canada Geese were extirpated from tion to those on Buldir (S. Hatch, pers. comm.). most of their historical range (Fig. 1) after the However, Shields and Wilson (1987) determined introduction of non-native predatorsfor fur pro- that newly discovered “Aleutian-like” Canada duction between the 1830s and 1930s (USFWS Geese from Chagulak and Kiliktagik could not 1991). The only population known to survive be genetically differentiated from B. c. leuco- was on Buldir Island where predators had not pareia from Buldir based on 21 restriction en- zymes and examination of nearly 200 restriction fragments from mitochondrial DNA (mtDNA). I Received 13 May 1999. Accepted 20 October 1999. MtDNAs of the two other small-bodied Canada 2 Current address: Department of Biology and geese in Alaska (Cackling Canada Geese and Chemistry, Carroll College, Helena, MT 59601. Taverner’s Canada Geese) were clearly distin- 11721 GENETIC STATUS OF ALEUTIAN CANADA GEESE 173 FIGURE 1. Historical distribution (polygons) and current U.S. (black circles) breeding populationsof Aleutian Canada Geese (USFWS 1991). Locations sampled in this study were Aleutian Canada Geese from Buldir Island and the Semidi Islands and Cackling Canada Geese from the Yukon-Kuskokwim Delta. guishable from those of the three island popu- geese winter in the same areas as the Buldir lations of Aleutian Canada Geese. The discovery geese (USFWS 1991). of the two previously unknown breeding popu- Previous genetic studies of Aleutian Canada lations of Aleutian Canada Geese on Chagulak Geese (allozymes, Morgan et al. 1977; mtDNA, and Kiliktagik and an increase from approxi- Shields and Wilson 1987) were based on small mately 800 birds in the mid-1970s to 32,000 sample sizes and were not sufficiently polymor- birds in 1999, led the U.S. Fish and Wildlife phic to provide data with which to establishpop- Service to propose delisting Aleutian Canada ulation relationships and comparative population Geese as threatened (DeGange 1999). levels of variation. Recently, techniques includ- Despite the lack of genetic evidence of dis- ing analysis of biparentally-inherited nuclear mi- tinctions, banding data suggestthere are at least crosatellite loci have proven to be highly infor- two demographically distinct sub-populations of mative regarding questions about dispersal, gene Aleutian Canada Geese. Buldir birds stage and flow, and relationships of isolated populations winter in northern and central California, re- (McDonald and Potts 1997). In this study we spectively, whereas Semidi birds winter in used seven recently developed biparentally-in- Oregon (USFWS 1991). The large distance be- herited microsatellite loci (Buchholz et al. 1998, tween breeding sites on Buldir and those on the Cathey et al. 1998) and DNA sequenceanalysis Semidi Islands, the time interval since the breed- of 143 base pairs of domain I of the matemally- ing distribution was more continuous (ca. 1750, inherited avian mtDNA control region to study DeGange 1999), and the two populations’ cur- Aleutian Canada Geese from Buldir and the rent use of different wintering areas suggestthe Semidi Islands. These two non-coding genetic two populations may be reproductively isolated. systems are subject to higher rates of mutation Migration patterns and wintering locations for than coding regions of the genome (Brown et al. geese of Amukta and Chagulak of the central 1982, Goldstein and Pollock 1997) and conse- portion of the Aleutian Island chain are not well quently are thought to be more informative at defined, but some banding data indicate these the population level. Our objective was to use 174 BARBARA J. PIERSON ET AL. molecular genetic markers to assessthe extent primer, and non-labeled primers at locus-specific of gene flow among populations, the potential concentrations [0.8 FM (TTUCG-4) or 0.36 uM genetic effects of population bottlenecks, and (TTUCG-5, TTUCG-1, Bcap 1, Bcap. 3) non- genetic relationships of Aleutian Canada Geese labeled forward primer, 1.2 pM (TTUCG-4) or from Buldir and the Semidi Islands. For com- 0.4 pM (TTUCG-5, ‘ITUCG-1, Bcap. 1, Bcap. parison, we used Cackling Canada Geese from 3) or 0.12 pM (Beak 9) or 0.04 PM (Beak 11) the Yukon-Kuskokwim (Y-K) Delta, which also reverse primer]. Thermocycler conditions in- have experienced reductions in population num- cluded initial denaturation at 94°C for 2 min, bers, although not as drastic as Aleutian Canada followed by 25-35 cycles of denaturing at 94°C Geese have. for 1 min, annealing at locus-specific tempera- tures 56°C (TTUCG-4, Beak 1, Bcap 3, Beau METHODS 9, Bca(* 11) or 60°C (TTUCG-5, TTUCG-1) for SAMPLE COLLECTION AND PREPARATION 1 min, and extending at 72°C for 1 min. During 1991, whole blood samples were col- Genotypes at each microsatellite locus were lected from geese at Buldir Island (n = 29), the assigned relative to an Ml3 sequence standard Semidi Islands (n = S), and from captive geese and internal controls (individuals of known ge- raised at the University of California, Davis notype run on each gel) after standardpolyacryl- from eggs taken at Buldir (n = 5). In addition, amide gel electrophoresis and autoradiography. tissue samples (n = 12) identified as being from Microsatellite allele frequencies, estimates of Semidi Islands birds based on morphology and observed and expected heterozygosity, and ge- their discrete wintering location were collected netic distance (chord distance, Cavalli-Sforza from carcassesfound on wintering grounds in and Edwards 1967) were derived using BIOSYS 1993 and 1994 in Tillamook County, Oregon. (version 1.7, Swofford and Selander 1981). The Samples were placed in tissue preservation buff- chord distance has been shown to be robust with er (4 M Urea, 0.2 M NaCl, 100 mM Tris-HCl respect to tree topology for microsatellite loci pH 8.0, 0.5% n-Lauryl sarkosine, 10 mM surveyed for i&a-specific populations (Takezaki EDTA) and shipped at ambient temperatures. and Nei 1996). Genetic distances were used to Blood quills from molting Cackling Canada generate a multi-locus neighbor-joining tree in Geese (n = 41) were collected in 1995 on the MEGA (Kumar et al. 1993). Bootstrap values Y-K Delta and were placed in the tissue pres- could not be generated for the tree due to an ervation buffer described above. All samples insufficient number of populations. Estimates of were stored at -20°C in the laboratory prior to Wright’s (1969) inbreeding coefficient (F) were analysis. Genomic
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