Geographical genetics: Conceptual foundations and empirical applications of spatial genetic data in wildlife management

Journal of Wildlife Management

Volumn 69, Issue 4, 2005, Pages 1434-1453

  Scribner, Kim T ^a   Blanchong, Julie A ^a,c   Bruggeman, Douglas J ^a   Epperson, Bryan K ^a   Lee, Cheng Yu ^a   Pan, Yu Wen ^a   Shorey, Rainy I ^a   Prince, Harold H ^a   Winterstein, Scott R ^a   Luukkonen, David R ^b  

^a MICHIGAN STATE UNIVERSITY   (United States)

^b MICHIGAN DEPARTMENT OF NATURAL RESOURCES   (United States)

^c UNIVERSITY OF WISCONSIN MADISON   (United States)

Author keywords

Autocorrelation; Canada geese; Disease ecology; Habitat equivalency analysis; Metapopulations; Population genetics; Spatial genetic structure; White tailed deer

Indexed keywords

CONSERVATION GENETICS; POPULATION GENETICS; WILDLIFE MANAGEMENT;

ANSER SP.; BRANTA CANADENSIS; ODOCOILEUS VIRGINIANUS;

EID: 33645659508     PISSN: 0022541X     EISSN: None     Source Type: Journal    
DOI: 10.2193/0022-541X(2005)69[1434:GGCFAE]2.0.CO;2     Document Type: Review

References (127)

1. AMOS, W., AND A. BALMFORD. 2001. When does conservation matter? Heredity 87:257-265.
2. AVISE, J. C. 1991. Ten unorthodox perspectives on evolution prompted by comparative population genetic findings on mitochondrial DNA. Annual Review of Genetics 25:45-69.
3. BAKER, A. 2000. Molecular methods in ecology. Blackwell Scientific, London, United Kingdom.
4. BARBUJANI, G. 1987. Autocorrelation of gene frequencies under isolation-by-distance. Genetics 117:772-782.
5. BLANCHONG, J. A., K. T. SCRIBNER, AND S. R. WINTERSTEIN. 2002. Assignment of individuals to populations: Bayesian methods and multilocus genotypes. Journal of Wildlife Management 66:321-329.
6. _. 2003. Genealogical relationships influence the probability of infection with bovine tuberculosis and microgeographic genetic structure in free-ranging white-tailed deer. Dissertation, Michigan State University, East Lansing, USA.
7. _, K. T. SCRIBNER, B. K. EPPERSON, AND S. R. WINTERSTEIN. 2005. Changes in artificial feeding regulations impact white-tailed deer microgeographic genetic structure. Journal of Wildlife Management:in press.
8. BOWCOCK, A. M., A. RUIZ-LINARES, J. TOMFOHRED, E. MINCH, J. R. KIDD, AND L. L. CAVALLI-SFORZA. 1994. High resolution of human evolutionary trees with polymorphic microsatellites. Nature 368:455-457.
9. BOX, G. E. P., AND G. M. JENKINS. 1976. Time series analysis: forecasting and control. Holden-Day, San Francisco, California, USA.
10. BRUGGEMAN, D., M. JONES, F. LUPI, AND K. T. SCRIBNER. 2005. Calculating spatially-dependent biodiversity credits with landscape equivalency analysis to trade habitat patches within an endangered metapopulation. Environmental Management 36:518-534.
11. CADRIN, S. X., K. D. FREIDLAND, AND J. R. WALDMAN. 2005. Stock identification methods: applications in fishery science. Elsevier Academic Press, New York, USA.
12. CEGELSKI, C. C., L. P. WAITS, AND N. J. ANDERSON. 2003. Assessing population structure and gene flow in Montana wolverines (Gulo gulo). Molecular Ecology 12:2907-2918.
13. CHESSER R. K. 1991a. Gene diversity and female philopatry. Genetics 127:437-447.
14. _. 1991b. Influence of gene flow and breeding tactics on gene diversity within populations. Genetics 129:573-583.
15. _, AND R. J. BAKER. 1996. Effective sizes and dynamics of uniparentally and diparentally inherited genes. Genetics 144:1225-1235.
16. CLIFF, A. D., AND J. K. ORD. 1981. Spatial autocorrelation. Pion, London, United Kingdom.
17. COCKERHAM, C. C. 1969. Variance of gene frequencies. Evolution 23:72-84.
18. _. 1973. Analysis of gene frequencies. Genetics 74:679-700.
19. CORNUET, J. M., S. PIRY, G. LUIKART, A. ESTOUP, AND M. SOLIGNAC. 1999. New methods employing multilocus genotypes to select or exclude populations as origins of individuals. Genetics 153:1989-2000.
20. DALE, M. R., P. DIXON, M-J. FORTIN, P. LEGENDRE, D. E. MYERS, AND M. S. ROSENBERG. 2002. Conceptual and mathematical relationships among methods for spatial analysis. Ecography 25:558-577.
21. DAWSON, K. J., AND K. BELKHIR. 2001. A Bayesian approach to the identification of panmictic populations and assignment of individuals. Genetical Research 78:59-77.
22. DEBEVEC, E. M., R. B. GATES, M. MASUDA, J. PELLA, J. H. REYNOLDS, AND L. W. SEEB. 2000. SPAM (Version 3.2): statistics program for analyzing mixtures. Journal of Heredity 91:509-510.
23. DEGROOT, R. S., M. A. WILSON, AND R. M. J. BOUMANS. 2002. A typology for the classification, description and valuation of ecosystem functions, goods, and services. Ecological Economics 41:393-408.
24. DESALLE, R., AND G. AMATO. 2004. The expansion of conservation genetics. Nature Reviews Genetics 5:702-722.
25. DEWEY, S. E., AND J. S. HEYWOOD. 1988. Spatial genetic structure in a population of Psychotria nervosa, I: distribution of genotypes. Evolution 42:834-838.
26. DUNNING, JR., J. B., D. J. STEWART, B. J. DANIELSON, B. R. NOON, T. L. ROOT, R. H. LAMBERSON, AND E. E. STEVENS. 1995. Spatially explicit population models: current forms and future uses. Ecological Applications 5:3-11.
27. D'EON, R. G., S. M. GLAMM, E. PARFITT, AND M-J. FORTIN. 2002. Landscape connectivity as a function of scale and organism vagility in a real forested landscape. Conservation Ecology 6:10-24.
28. EPPERSON, B. K. 1993. Spatial and space-time correlations in systems of subpopulations with genetic drift and migration. Genetics 133:711-727.
29. _. 2003. Geographical genetics. Princeton University Press, Princeton, New Jersey, USA.
30. FAVRE, L., F. BALLOUX, J. GOUDET, AND N. PERRIN. 1997. Female-biased dispersal in a monogamous mammal Crocidura russula: evidence from field data and microsatellite patterns. Proceedings of the Royal Society of London Series B 264:127-132.
31. FELIX, A. B., H. CAMPA, K. F. MILLENBAH, S. R. WINTERSTEIN, AND W. E. MORITZ. 2004. Development of landscape-scale habitat-potential models for forest wildlife planning and management. Wildlife Society Bulletin 32:795-806.
32. FOREY, P. L., C. J. HUMPHRIES, AND R. A. VANE-WRIGHT. 1994. Systematics and conservation evaluation. Clarendon Press, Oxford, United Kingdom.
33. FROMONT, E., D. PONTIER, AND M. LANGLAIS. 1998. Dynamics of a feline retrovirus (FeLV) in host populations with variable spatial structure. Proceedings of the Royal Society of London B 265:1097-1104.
34. FULFORD, G. R., M. G. ROBERTS, AND J. A. P. HEESTERBEEK. 2002. The metapopulation dynamics of an infectious disease: tuberculosis in possums. Theoretical Population Biology 61:15-29.
35. GAGGIOTTI, O. E., S. P. BROOKS, W. AMOS, AND J. HARWOOD. 2004. Combining demographic, environmental and genetic data to test hypotheses about colonization events in metapopulations. Molecular Ecology 13:811-825.
36. GARNER, M. 2001. Movement patterns and behavior at winter feeding and fall baiting stations in a population of white-tailed deer infected with bovine tuberculosis in the northeastern lower peninsula of Michigan. Dissertation, Michigan State University, East Lansing, USA.
37. GRIFFITHS, R., M. C. DOUBLE, K. ORR, AND R. J. G. DAWSON. 1998. A DNA test to sex most birds. Molecular Ecology 7:1071-1076.
38. HANSEN, M. M., D. E. RUZZANTE, E. E. NIELSEN, AND K. L. D. MENSBERG. 2000. Microsatellite and mitochondrial DNR polymorphism reveals life-history-dependent interbreeding between hatchery and wild brown trout (Salmo trutta L.) Molecular Ecology 9:583-594.
39. HANSKI, I., AND M. E. GILPIN. 1997. Metapopulation biology: ecology, genetics, and evolution. Academic Press, San Diego, USA.
40. _, AND O. E. GAGGIOTTI. 2004. Ecology, genetics, and evolution of metapopulations. Academic Press, New York, USA.
41. HEDRICK, P. W. 2002. Application of molecular genetics to conservation: new issues and examples. Pages 367-387 in S. R. Beissinger, and D. R. McCullough, editors. Population viability analysis. Chicago Press, Chicago, Illinois, USA.
42. _, AND M. E. GILPIN. 1997. Genetic effective size of a metapopulation. Pages 165-181 in I. A. Hanski and M. E. Gilpin, editors. Metapopulation biology: ecology, genetics and evolution. Academic Press, New York, USA.
43. HOOPER, P. M., AND G. J. D. HEWINGS. 1981. Some properties of space-time processes. Geographical Analysis 13:203-223.
44. HUDSON, R. R. 1990. Gene genealogies and the coalescent process. Pages 1-44 in D. Futuyma and J. Antonovics, editors. Oxford surveys in evolutionary biology. Oxford University Press, Oxford, United Kingdom.
45. INMAN, R., K. T. SCRIBNER, J. WARRILLOW, H. H. PRINCE, AND D. LUUKKONEN. 2003. Canada goose harvest derivation from genetic analysis of tail feathers. Wildlife Society Bulletin 31:1126-1131.
46. JONES, C. A., AND K. A. PEASE. 1997. Restoration-based compensation measures in natural resource liability statutes. Contemporary Economic Policy 15:111-122.
47. KENDALL, W. L., AND J. D. NICHOLS. 2002. Estimating statetransition probabilities for unobservable states using capture-recapture/resighting data. Ecology 83:3276-3284.
48. KIMURA, M., AND WEISS, G. H. 1964. The stepping stone model of population structure and the decrease of genetic correlation with distance. Genetics 49:561-576.
49. LACY, R. C. 1987. Loss of genetic diversity from managed populations: interacting effects of drift, mutation, immigration, selection, and population subdivision. Conservation Biology 1:143-157.
50. _. 2000. Structure of the VORTEX simulation model for population viability analysis. Ecological Bulletin 48:191-203.
51. LANDE, R. 1998. Anthropogenic, ecological, and genetic factors in extinction and conservation. Research in Population Ecology 40:259-269.
52. _, AND G. F. BARROWCLOUGH. 1987. Effective population size and genetic variation. Pages 87-123 in M. E. Soulé, editor. Viable populations for conservation. Cambridge University Press, London, United Kingdom.
53. LEBRETON, J-D., J. E. HINES, R. PRADEL, J. D. NICHOLS, AND J. A. SPENDELOW. 2003. Estimation by capture-recapture of recruitment and dispersal over several sites. Oikos 101:253-264.
54. LEGENDRE, P. 1993. Spatial autocorrelation: trouble or new paradigm? Ecology 74:1659-1673.
55. _, M. R. DALE, M-J. FORTIN, J. GUREVITCH, M. HOHN, AND D. MYERS. 2002. The consequences of spatial structure for the design and analysis of ecological field surveys. Ecography 25:601-615.
56. _, AND M-J. FORTIN. 1989. Spatial pattern and ecological analysis. Vegetation 80:107-138.
57. LOOMIS, J. B., AND D. S. WHITE. 1996. Economic benefits of rare and endangered species: summary and meta-analysis. Ecological Economics 18:197-206.
58. LYNCH, M. 1991. The genetic interpretation of inbreeding depression and outbreeding depression. Evolution 45:622-629.
59. MALÉCOT, G. 1948. Les mathématiques de l'héré dité. Masson, Paris, France. [In French.]
60. MANEL, S., M. K. SCHWARTZ, G. LUIKART, AND P. TABERLET. 2003. Landscape genetics: combining landscape ecology and population genetics. Trends in Ecology and Evolution 18:189-197.
61. MANLOVE, M. N., J. C. AVISE, H. O. HILLSTAD, P. R. RAMSEY, D. O. STRANEY, AND M. H. SMITH. 1975. Starch gel electrophoresis for the study of population genetics of white-tailed deer. Proceedings of the Southeastern Association of Game and Fish Commissioners 29:392-403.
62. MANTEL, N. 1967. The detection of disease clustering and a generalized regression approach. Cancer Research 27:209-220.
63. MATTHEWS, N. E., AND W. F. PORTER. 1993. Effect of social structure on genetic structure of free-ranging white-tailed deer in the Adirondack mountains. Journal of Mammalogy 74:33-43.
64. MCCAULEY, D. E. 1991. Genetic consequences of population extinction and recolonization. Trends in Ecology and Evolution 6:33-37.
65. MERKEL, D. M. 1988. Soil nutrients in glaciated Michigan landscapes: distribution of nutrients and relationships with stand productivity. Dissertation, Michigan State University, East Lansing, USA.
66. MILLAR, R. B. 1987. Maximum likelihood estimation of mixed stock fishery composition. Canadian Journal of Fisheries and Aquatic Sciences 44:583-590.
67. MILLS, L. S., AND F. W. ALLENDORF. 1996. The one-migrant-per-generation rule in conservation and management. Conservation Biology 10:1509-1518.
68. _, J. J. CITTA, K. P. LAIR, M. K. SCHWARTZ, AND D. A. TALLMON. 2000. Estimating animal abundance using noninvasive DNA sampling: promise and pitfalls. Ecological Applications 10:283-294.
69. MORITZ, C. 1994. Applications of mitochondrial DNA analysis in conservation: a critical review. Molecular Ecology 3:401-411.
70. NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION. 1999a. Discounting and the treatment of uncertainty in natural resource damage assessment. Technical paper 99-1. Damage Assessment and Restoration Program, Damage Assessment Center, Silver Springs, Colorado, USA.
71. _. 1999b. Habitat equivalency analysis: an overview. Policy and technical paper series, No. 95-1. Technical paper 99-1. Damage Assessment and Restoration Program, Damage Assessment Center, Silver Springs, Colorado USA.
72. NATIONAL RESEARCH COUNCIL. 1995. Science and the endangered species act. National Academy of Science, Washington, D.C., USA.
73. NEI, M. 1973. Analysis of gene diversity in subdivided populations. Proceedings of the National Academy of Sciences of the United States of America 70:3321-3323.
74. _. 1987. Molecular evolutionary genetics. Columbia University Press, New York, USA.
75. NEIGEL, J. E. 1997. A comparison of alternative strategies for estimating gene flow from genetic markers. Annual Review of Ecology and Systematics 28:105-128.
76. NICHOLS, J. D., J. E. HINES, J-D. LEBRETON, AND R. PRADEL. 2000. Estimation of contributions to population growth: a reverse-time capture-recapture approach. Ecology 81:3362-3376.
77. NOSS, R. F., M. A. O'CONNELL, AND D. D. MURPHY. 1997. The science of conservation planning. Island Press, Washington, D.C., USA.
78. ODEN, N. L., AND R. R. SOKAL. 1978. Spatial autocorrelation in biology. 2. Some biological implications and four applications of evolutionary and ecological interest. Biological Journal of the Linnean Society 10:229-249.
79. _, AND _. 1986. Directional autocorrelation: an extension of spatial correlograms in two dimensions. Systematic Zoology 35:608-617.
80. PALSBOLL, P. J. 1999. Genetic tagging: contemporary molecular ecology. Biological Journal of the Linnean Society 68:3-22.
81. PAETKAU, D., W. CALVERT, I. STIRLING, AND C. STROBECK. 1995. Microsatellite analysis of population structure in Canadian polar bears. Molecular Ecology 4:347-354.
82. _, R. SLADE, M. BURDENS, AND A. ESTOUP. 2004. Genetic assignment methods for direct, real-time estimation of migration rate: a simulation-based exploration of accuracy and power. Molecular Ecology 13:55-65.
83. PEARCE, J. M., B. J. PIERSON, S. L. TALBOT, D. V. DERKSEN, D. KRAEGE, AND K. T. SCRIBNER. 2000. A genetic evaluation of morphology used to identify hunter-harvested Canada geese. Journal of Wildlife Management 64:863-874.
84. PELLA, J. J., AND G. B. MILNER. 1987. Use of genetic marks in stock composition analysis. Pages 247-276 in N. Ryman and F. Utter, editors. Population genetics and fishery management. University of Washington Press, Seattle, USA.
85. _, AND M. MASUDA. 2001. Bayesian methods for analysis of stock mixtures from genetic characters. Fisheries Bulletin 99:151-167.
86. PETIT, R. J., A. E. MOUSADIK, AND O. PONS. 1998. Identifying populations for conservation on the basis of genetic markers. Conservation Biology 12:844-855.
87. PFEIFFER, P. E., AND S. J. DEUTSCH. 1980. A three-stage iterative procedure of space-time modeling. Technometrics 22:35-47.
88. POLLOCK, K. H. 1982. A capture-recapture design robust to unequal probability of capture. Journal of Wildlife Management 46:757-760.
89. POSSINGHAM, H. P., S. J. ADELMAN, B. R. NOON, S. TROMBULAK, AND H. R. PULLIAM. 2001. Making smart conservation decisions. Pages 225-244 in M. E. Soulé and G. H. Orians, editors. Conservation biology: research priorities for the next decade. Island Press, Washington, D.C., USA.
90. PRIMMER, C. R., M. T. KOSKIEN, AND J. PIRONEN. 2000. The one that did not get away: individual assignment using microsatellite data detects a case of fishing competition fraud. Proceedings of the Royal Society of London Series B 267:1699-1704.
91. PRITCHARD, J. K., M. STEPHENS, AND P. DONNELLY. 2000. Inference of population structure using multilocus genotype data. Genetics 155:945-959.
92. QUELLER, D. C., AND K. F. GOODNIGHT. 1989. Estimating relatedness using genetic markers. Evolution 43:258-275.
93. RANNALA, B., AND J. L. MOUNTAIN. 1997. Detecting immigration using multilocus genotypes. Proceedings of the National Academy of Sciences of the United States of America 94:9197-9202.
94. RAY, N. 2005. Pathmatrix: a geographical information system tool to compute effective distance among samples. Molecular Ecology Notes 5:177-180.
95. REXTAD, E. C. 1992. Effect of hunting on annual survival of Canada geese in Utah. Journal of Wildlife Management 56:297-305.
96. ROSENBERG, M. S. 2000. PASSAGE. Pattern analysis, spatial statistics, and geographic exegesis. Version 1.0. Arizona State University, Tempe, USA.
97. SAITOU N., AND M. NEI. 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4:406-425.
98. SCHMITT, S. M., S. D. FITZGERALD, T. M. COOLEY, C. S. BRUNING-FANN, L. SULLIVAN, D. BERRY, T. CARLSON, R. B. MINNIS, J. B. PAYEUR, AND J. LIKARSKIE. 1997. Bovine tuberculosis in free-ranging white-tailed deer from Michigan. Journal of Wildlife Diseases 33:749-758.
99. SCRIBNER, K. T., AND R. K. CHESSER. 1993. Environmental and demographic components of spatial genetic structure in populations of the eastern cottontail rabbit. Journal of Mammalogy 74:1026-1044.
100. _, AND _. 2001. Group-structured models in analyses of the population and behavioral ecology of poikilothermic vertebrates. Journal of Heredity 98:201-210.
101. _, J. W. ARNTZEN, T. BURKE, N. CRUDDACE, AND R. S. OLDHAM. 2000. Environmental correlates of toad abundance and inter-population variation in measures of genetic diversity: estimates derived from single locus minisatellites. Biological Conservation 98:201-210.
102. _, R. FIELDS, S. TALBOT, J. PEARCE, AND M. PETERSEN. 2001. Sex-biased dispersal in threatened spectacled eiders: evaluation using molecular markers with contrasting modes of inheritance. Evolution 55:2105-2115.
103. _, J. A. WARRILLOW, J. O. LEAFLOOR, H. H. PRINCE, R. L. INMAN, D. R. LUUKKONEN, AND C. S. FLEGEL. 2003. Genetic methods for determining racial composition of Canada goose harvests. Journal of Wildlife Management 67:123-136.
104. _, M. C. WOOTEN, M. H. SMITH, AND P. E. JOHNS. 1984. Demographic and genetic characteristics of white-tailed deer subjected to still and dog hunting. Pages 197-212 in S. L. Beasom, editor. Symposium on game harvest. Caesar Kleberg Wildlife Management Institute, Kingsville, Texas, USA.
105. SHAO, J. 1993. Linear model selection by cross-validation. Journal of the American Statistical Association 88:486-494.
106. SLATKIN, M. 1985. Gene flow in natural populations. Annual Review of Ecology and Systematics 16:393-430.
107. SMITH, T. B., AND R. K. WAYNE. 1996. Molecular genetic approaches in conservation. Oxford University Press, New York, USA.
108. SMOUSE, P. E., J. C. LONG, AND R. R. SOKAL. 1986. Multiple regression and correlation extensions of the Mantel test of matrix correspondence. Systematic Zoology 35:627-632.
109. _, AND R. PEAKALL. 1999. Spatial autocorrelation analysis of individual multiallele and multilocus genetic structure. Heredity 82:561-573.
110. SOKAL, R. R., AND N. L. ODEN. 1978. Spatial autocorrelation in biology. 1. Methodology. Biological Journal of the Linnean Society 10:199-228.
111. _, _, AND B. A. THOMSON. 1997. A simulation study of microevolutionary inferences by spatial autocorrelation analysis. Biological Journal of the Linnean Society 60:73-93.
112. SORK, V. L., J. NASON, D. R. CAMPBELL, AND J. F. FERNANDEZ. 1999. Landscape approaches to historical and contemporary gene flow in plants. Trends in Ecology and Evolution 14:219-224.
113. SUGG, D. W., AND R. K. CHESSER. 1994. Effective population sizes with multiple paternity. Genetics 137:1147-1155.
114. _, _, F. S. DOBSON, AND J. L. HOOGLAND. 1996. Population genetics meets behavioral ecology. Trends in Ecology and Evolution 11:338-342.
115. TAYLOR, P. D., L. FAHRIG, K. HENEIN, AND G. MERRIAM. 1993. Connectivity is a vital element of landscape structure. Oikos 68:571-573.
116. TISHENDORF, L., AND L. FARHIG. 2000. How should we measure landscape connectivity? Landscape Ecology 15:633-641.
117. WASER, P. M., AND C. STROBECK. 1998. Genetic signatures of interpopulation dispersal. Trends in Ecology and Evolution 13:43-44.
118. WEIR B. S., AND C. C. COCKERHAM. 1984. Estimating F-statistics for the analysis of population structure. Evolution 38:1358-1370.
119. WILCOVE, D. S., D. ROTHSTEIN, J. DUBOW, A. PHILLIPS, AND E. LOSOS. 1998. Quantifying threats to imperiled species in the United States. Bioscience 48:607-615.
120. WILCOX, B. A., AND D. D. MURPHY. 1985. Conservation strategy: the effects of fragmentation on extinction. American Naturalist 125:879-887.
121. WILSON, G. A., AND B. RANNALA. 2003. Bayesian inference of recent migration rates using multilocus genotypes. Genetics 163:1177-1191.
122. WOOD, C. C., S. MCKINNELL, T. J. MULLIGAN, AND D. A. FOURNIER. 1987. Stock identification with the maximum-likelihood mixture model: sensitivity analysis and application to complex problems. Canadian Journal of Fisheries and Aquatic Sciences 44:866-881.
123. WOODS, J. G., D. PAETKAU, D. LEWIS, B. N. MCLELLAN, M. PROCTOR, AND C. STROBECK. 1999.Genetic tagging of free-ranging black and brown bears. Wildlife Society Bulletin 27:616-627.
124. WRIGHT, S. 1943. Isolation by distance. Genetics 28:114-138.
125. _. 1965. The interpretation of population structure by F-statistics with special regard to systems of mating. Evolution 19:395-420.
126. XU, S., C. J. KOBAK, AND P. E. SMOUSE. 1994. Constrained least squares estimation of mixed population stock composition from mtDNA haplotype frequency data. Canadian Journal of Fisheries and Aquatic Sciences 51:417-425.
127. YOUNG, A. G., AND G. M. CLARKE. 2000. Genetics, demography, and viability of fragmented populations. Cambridge University Press, Cambridge, United Kingdom.