Gharrett et al.: Phylogeographic analysis of mitochondrial DNA variation in Oncorhynchus kisutch 
539 
Acknowledgments 
We appreciate the constructive and encouraging comments 
made by A.T. Beckenbach on an early draft. D. Churikov 
and B. Finney provided valuable discussions. We thank 
D. Churikov, J. Gharrett, and three anonymous reviewers 
for reviewing the manuscript. T. Chatto, S. Kelley, D. 
McBride, L. Schwartz, L. Shaul, C. Skaugstad, and S. Taylor 
kindly provided samples. This work is a result of research 
sponsored in part by the Saltonstall-Kennedy Fund NA36 
FD0179 and in part by the Alaska Sea Grant College Pro- 
gram, which is cooperatively sponsored by NOAA, Office of 
Sea Grant and Extramural Programs, Department of Com- 
merce, under grant NA90AA-D-SG066 (project R/02-16), 
and the University of Alaska with funds appropriated by 
the State. The manuscript was revised while AJG was at 
Hokkaido University and supported, in part, by the Japan 
Society for Promotion of Science. 
Literature cited 
Adams, N. S., W. J. Spearman, C. V. Burger, K. P. Currens, 
C. B. Schreck, and H. W. Li. 
1994. Variation in mitochondrial DNA and allozymes dis- 
criminates early and late forms of chinook salmon ( On- 
corhynchus tshawytscha) in Kenai and Kasilof rivers, 
Alaska. Can. J. Fish. Aquat. Sci. Shsuppl. 1 ): 172—181 . 
Anderson, S., A. T. Bankier, G. T. Barrell, M. H. DeBruijn, and 
A. R. Coulson. 
1981. Sequence and organization of the human mitochon- 
drial genome. Nature 290:457-465. 
Anderson, S., M. H. DeBruijn, A. R. Coulson, I. C. Eperon, and 
F. Sanger. 
1982. Complete sequence of bovine mitochondrial DNA. J. 
Mol. Biol. 156:683-717. 
Avise, J. C. 
1989. Gene trees and organismal histories: a phylogenetic 
approach to population biology. Evolution 43:1 192-1208. 
1992. Molecular population structure and the biogeographic 
history of a regional fauna: a case history with lessons for 
conservation biology. Oikos:63-76. 
Avise, J. C., J. Arnold, R. M. Ball, E. Bermingham, T. Lamb, 
J. E. Neigel, C. A. Reeb, and N. C. Saunders. 
1987. Intraspecific phylogeography: the mitochondrial DNA 
bridge between population genetics and systematics. Ann. 
Rev. Ecol. Syst. 18:489-522. 
Avise, J. C., R. M. Ball, and J. Arnold. 
1988. Current versus historical population sizes in verte- 
brate species with high gene flow: a comparison based 
on mitochondrial DNA lineages and inbreeding theory for 
neutral mutations. Mol. Biol. Evol. 5:331-344. 
Avise, J. C., B. W. Bowen, and T. Lamb. 
1989. DNA fingerprints from hypervariable mitochondrial 
genotypes. Mol. Biol. Evol. 6:258-269. 
Avise, J. C., B. W. Bowen, T. Lamb, A. B. Meylan, and 
E. Bermingham. 
1992. Mitochondrial DNA evolution at a turtle’s pace: evi- 
dence for low genetic variability and reduced microevo- 
lutionary rate in the Testudines. Mol. Biol. Evol. 9:457- 
473. 
Avise, J. C., G. S. Helfman, N. C. Saunders, and L. S. Hales. 
1986. Mitochondrial DNA differentiation in North Atlantic 
eels: population genetic consequences of an unusual life 
history pattern. Proc. Nat. Acad. Sci. US. 83:4350-4354. 
Bartlein, P. J., P. M. Anderson, M. E. Edwards, and 
P. F. McDowell. 
1991. A framework for interpreting paleoclimatic varia- 
tions in eastern Beringia. Quaternary International 12-1: 
73-83. 
Bartley, D. M., B. Bentley, P. G. Olin, and G. A. E. Gall. 
1992. Population genetic structure of coho salmon ( Oncor- 
hynchus kisutch). Calif. Fish. Game 78:88-104. 
Bartley, D. M„ and G. A. E. Gall. 
1990 . Genetic structure and gene flow in chinook salmon pop- 
ulations in California. Trans. Am. Fish. Soc. 119:55-71. 
Beacham, T. D., R. E. Withler, and A. P. Gould. 
1985. Biochemical genetic stock identification of pink salmon 
(Oncorhynchus gorbuscha) in southern British Columbia 
and Puget Sound. Can. J. Fish. Aquat. Sci. 42:1474-1483. 
Bentzen, P, G. C. Brown, and W. C. Leggett. 
1989. Mitochondrial DNA polymorphism, population struc- 
ture, and life history variation in American shad ( Alosa 
sapidissirna ). Can. J. Fish. Aquat. Sci. 46:1446-1454. 
Bentzen, P, W. C. Leggett, and G. G. Brown. 
1993. Genetic relationships among the shads (Alosa) 
revealed by mitochondrial DNA analysis. J. Fish. Biol. 
43:909-917. 
Bernatchez, L., and J. J. Dodson. 
1991. Phylogeographic structure in mitochondrial DNA of 
the lake whitefish (Coregonus clupeaformis ) and its rela- 
tion to Pleistocene glaciations. Evolution 45:1016-1035. 
Bibb, M. J., R. A. van Etten, C. T. Wright, M. W. Walberg, and 
D. A. Clayton. 
1981. Sequence and gene organization of mouse mitochon- 
drial DNA. Cell 26:167-180. 
Bickham, J. W., C. C. Wood, and J. C. Patton. 
1995. Biogeograplnc implications of cytochrome b sequences 
and allozymes in sockeye (Oncorhynchus nerka ). J. Hered. 
86:140-144. 
Brown, W. M., M. George, Jr., and A. C. Wilson. 
1979. Rapid evolution of animal mitochondrial DNA. Proc. 
Natl. Acad. Sci. USA 76:1967-1971. 
Brown, W. M., E. M. Prager, A. Wang, and A. C. Wilson. 
1982. Mitochondrial DNA sequences of primates: tempo and 
mode of evolution. J. Mol. Evol. 18:225-239. 
Brykov, V. A., N. Polyakova, L. A. Skurikhina, and 
A. D. Kukhlevsky. 
1996. Geographical and temporal mitochondrial DNA variabil- 
ity in populations of pink salmon. J. Fish Biol. 48:899-909. 
Burger, C. V., W. J. Spearman, and M. A. Cronin. 
1997. Genetic differentiation of sockeye salmon subpopu- 
lations from a geologically young Alaskan lake system. 
Trans. Am. Fish. Soc. 126:926-938. 
Carney, B. L., A. K. Gray, and A. J. Gharrett. 
1997. Mitochondrial DNA restriction site variation within and 
among five populations of Alaskan coho salmon (Oncorhyn- 
chus kisutch ). Can. J. Fish. Aquat. Sci. 54:940-949. 
Castelloe, J., and A. R. Templeton. 
1994. Root probabilities for intraspecific gene trees under 
neutral coalescent theory. Mol. Phylogenet. Evol. 3:102- 
113. 
Chang, Y-S., F-L. Huang, and T-B. Lo. 
1994. The complete nucleotide sequence and gene organiza- 
tion of carp (Cypnnus carpio) mitochondrial genome. J. 
Mol. Evol. 38:138-155. 
Cronin, M. A., W. J. Spearman, R. L. Wilmot, J. C. Patton, and 
J. W. Bickham. 
1993. Mitochondrial DNA variation in chinook (Oncorhyn- 
