Geographic Protein Variation and Speciation in Salamanders of the Plethodon Jordani and Plethodon Glutinosus Complexes in the Southern Appalachian Mountains with the Description of Four New Species

  • Richard Highton
  • Robert B. Peabody


The purpose of this paper is to analyze geographic protein variation in two closely related complexes of southern Appalachian woodland salamanders. The Plethodon jordani complex is restricted to the southern Appalachian Mountains, and the P glutinosus complex is widely distributed in the eastern and central United States. When we began our allozyme studies on eastern Plethodon, both complexes were regarded as monophyletic and each was made the subject of a separate study. Each complex previously had been recognized as a single variable species, Plethodon glutinosus (Green,1818), and Plethodon jordani Blatchley (1901) (Hairston,1950; Highton,1962, Highton,1970, Highton,1972). Highton and MacGregor(1983) and Highton (1984, 1989), showed that there are 16 genetically divergent groups within the P. glutinosus complex and all were recognized taxonomically as separate species. Seven of these occur in the southern Appalachians and three, P. aureolus (Highton 1984); P. glutinosus, and P. teyahalee Hairston (1950), are sympatric at a site in Polk County, Tennessee, without evidence of hybridation (Highton 1984). Both complexes are geographically variable in coloration and size but most individuals are easily identified. Members of the P. glutinosus complex are characterized by the possession of dorsal white or brassy spotting (except in P. chattahoochee) and lateral white or yellow spotting. Salamanders of the P. jordani complex are characterized by usually lacking dorsal spotting, and, except for two groups, most populations lack lateral spotting.


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  1. Avise, J. C. 1975. Systematic value of electrophoretic data. Systematic Zoology 23:465–481.CrossRefGoogle Scholar
  2. Avise, J. C. 1994. Molecular Markers, Natural History and Evolution. Chapman and Hall, New York, New York, U.S.A.CrossRefGoogle Scholar
  3. Bailey, J. R. 1937. Notes on plethodont salamanders of the southeastern United States. Occasional Papers Museum of Zoology, University of Michigan (364):1–10.Google Scholar
  4. Berven, K. A. 1982. The genetic basis of altitudinal variation in the wood frog Rana sylvatica. I. An experimental analysis of life history traits. Evolution 36:962–983.CrossRefGoogle Scholar
  5. Berven, K. A., and D. E. Gill. 1983. Interpreting geographic variation in life-history traits. American Zoologist 23:85–97.Google Scholar
  6. Bishop, S. C. 1941. Notes on salamanders with descriptions of several new forms. Occasional Papers Museum of Zoology, University of Michigan (451):1–25.Google Scholar
  7. Blatchley, W. S. 1901. On a small collection of batrachians from Tennessee, with descriptions of two new species. 25th Annual Report, Indiana Department of Geology, 1900:759–763.Google Scholar
  8. Brewer, G. J. 1970. An Introduction to Isozyme Techniques. Academic Press, New York, New York U.S.A. Brimley, C. 5.1912. Notes on the salamanders of the North Carolina mountains with descriptions of two new forms. Proceedings of the Biological Society of Washington 25:135–140.Google Scholar
  9. Brimley, C. S. 1927. An apparently new salamander (Plethodson clemsonae) from S.C. Copeia 164:73–75.CrossRefGoogle Scholar
  10. Bruce, R. C. 1966. Occurrence of the salamander Plethodon jordani in the Piedmont of western South Carolina. Copeia 1966:888–889.CrossRefGoogle Scholar
  11. Carr, D. E. 1996. Morphological variation among species and populations of salamanders in the Plethodon glutinosus complex. Herpetologica 52:56–65.Google Scholar
  12. Cavalli-Sforza, L. L., and A. W. E Edwards. 1967. Phylogenetic analysis: models and estimation procedures. Evolution 21:550–570.CrossRefGoogle Scholar
  13. Coyne, J. A., and H. A. Orr. 1998. The evolutionary genetics of speciation. Philosophical Transactions, Royal Society of London B. 353:287–305.CrossRefGoogle Scholar
  14. Davis, B. J. 1964. Disc electrophoresis-II. Method and application to human serum proteins. Annals of the New York Academy of Science 121:404–427.CrossRefGoogle Scholar
  15. Dawley, E. 1984. Recognition of individual, sex and species odours by salamanders of the Plethodon glutinosus-Plethodon jordani complexes. Animal Behaviour 32:353–361.CrossRefGoogle Scholar
  16. Dawley, E. 1986. Behavioral isolating mechanisms in sympatric terrestrial salamanders. Herpetologica 42:156–164.Google Scholar
  17. Dawley, E. 1987. Species discrimination between hybridizing and non-hybridizing terrestrial salamanders. Copeia 1987:924–931.Google Scholar
  18. Duncan, R., and R. Highton. 1979. Genetic relationships of the eastern large Plethodon of the Ouachita Mountains. Copeia 1979:96–110.CrossRefGoogle Scholar
  19. Dunn, E. R. 1917. Reptile and amphibian collections from the North Carolina mountains, with especial reference to salamanders. Bulletin of the American Museum of Natural History 37:593–634.Google Scholar
  20. Dunn, E. R. 1926. The Salamanders of the Family Plethodontidae. Smith College, Northampton, Massachusetts, U.S.A.Google Scholar
  21. Dunn, E. R., and A. A. Heinze. 1933. A new salamander from the Ouachita Mountains. Copeia 1933:121–122.CrossRefGoogle Scholar
  22. Folkerts, G. W. 1971. Notes on South Carolina salamanders. Journal of the Elisha Mitchell Scientific Society 87:206–208.Google Scholar
  23. Fowler, H. W. 1917. Notes on salamanders. Proceedings of the Academy of Natural Sciences, Philadelphia 69:7–28.Google Scholar
  24. Green, J. 1818. Descriptions of several species of North American Amphibia, accompanied with observations. Journal of the Academy of Natural Sciences, Philadelphia 1:348–359.Google Scholar
  25. Good, D. A., and D. B. Wake. 1992. Geographic variation and speciation in the torrent salamanders of the genus Rhyacotriton (Caudata: Rhyacotritonidae). University of California Publications in Zoology 126:1–91.Google Scholar
  26. Grobman, A. B. 1944. The distribution of the salamanders of the genus Plethodon in eastern United States and Canada. Annals of the New York Academy of Science 45:261–316.CrossRefGoogle Scholar
  27. Hairston, N. G. 1949. The local distribution and ecology of the plethodontid salamanders of the southern Appalachians. Ecological Monographs 19:47–73.CrossRefGoogle Scholar
  28. Hairston, N. G. 1950. Intergradation in Appalachian salamanders of the genus Plethodon. Copeia 1950:262–273.CrossRefGoogle Scholar
  29. Hairston, N. G. 1951. Interspecies competition and its probable influence upon the vertical distribution of Appalachian salamanders of the genus Plethodon. Ecology 32:266–274.CrossRefGoogle Scholar
  30. Hairston, N. G. 1980a. Evolution under interspecific competition: field experiments on terrestrial salamanders. Evolution 34:409–420.CrossRefGoogle Scholar
  31. Hairston, N. G. 1980b. The experimental test of an analysis of field distributions: competition in terrestrial salamanders. Ecology 61:817–826.CrossRefGoogle Scholar
  32. Hairston, N. G. 1981. An experimental test of a guild: salamander competition. Ecology 62:65–72.CrossRefGoogle Scholar
  33. Hairston, N. G. 1983a. Alpha selection in competing salamanders: experimental verification of an a priori hypothesis. American Naturalist 122:105–113.CrossRefGoogle Scholar
  34. Hairston, N. G. 1983b. Growth, survival, and reproduction of Plethodon jordani: trade-offs between selective pressures. Copeia 1983:1024–1035.CrossRefGoogle Scholar
  35. Hairston, N. G. 1993. On the validity of the name teyahalee as applied to a member of the Plethodon glutinosus complex (Caudata: Plethodontidae): a new name. Brimleyana 18:65–69.Google Scholar
  36. Hairston, N. G., Sr., R. H. Wiley, C. K. Smith, and K. A. Kneidel. 1992. The dynamics of two hybrid zones in Appalachian salamanders of the genus Plethodon. Evolution 46:930–938.CrossRefGoogle Scholar
  37. Harlan, R. 1825. Description of a variety of Coluber fulvius, Linn., a new species of Scincus, and two new species of Salamandra. Journal of the Academy of Natural Sciences, Philadelphia 5:226–242.Google Scholar
  38. Harrison, R. G. (Ed.). 1993. Hybrid Zones and the Evolutionary Process. Oxford University Press, New York, New York, U.S.A.Google Scholar
  39. Hass, C. A., R. Highton, and L. R. Maxson. 1992. Relationships among the eastern Plethodon: evidence from immunology. Journal of Herpetology 26:137–141.CrossRefGoogle Scholar
  40. Highton, R. 1962. Revision of North American salamanders of the genus Plethodon. Bulletin of the Florida State Museum 6:235–367.Google Scholar
  41. Highton, R. 1970. Evolutionary interactions between species of North American salamanders of the genus Plethodon. Part 1. Genetic and ecological relationships of Plethodon jordani and Plethodon glutinosus in the southern Appalachian Mountains. Evolutionary Biology 4:211–241.Google Scholar
  42. Highton, R. 1972. Distributional interactions among eastern North American salamanders of the genus Plethodon. Pp. 139–188. In P. C. Holt (Ed.), The Distributional History of the Biota of the Southern Appalachians. Part III: Vertebrates. Research Division Monograph, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, U.S.A.Google Scholar
  43. Highton, R. 1977. Comparison of microgeographic variation in morphological and electrophoretic traits. Evolutionary Biology 10:397–436.Google Scholar
  44. Highton, R. 1984. A new species of woodland salamander of the Plethodon glutinosus group from the southern Appalachian Mountains. Brimleyana 9:1–20.Google Scholar
  45. Highton, R. 1989. Biochemical evolution in the slimy salamanders of the Plethodon glutinosus complex in the eastern United States. Part I. Geographic protein variation. Illinois Biological Monographs 57:1–78.Google Scholar
  46. Highton, R. 1991. Molecular phylogeny of plethondonine salamanders and hylid frogs: statistical analysis of protein comparisons. Molecular Biology and Evolution 8:796–818.PubMedGoogle Scholar
  47. Highton, R. 1993. The relationship between the number of loci and the statistical support for the topology of UPGMA trees obtained from genetic distance data. Molecular Phylogenetics and Evolution 2:337–343.PubMedCrossRefGoogle Scholar
  48. Highton, R. 1995. Speciation in eastern North American salamanders of the genus Plethodon. Annual Review of Ecology and Systematics 26:579–600.CrossRefGoogle Scholar
  49. Highton, R. 1997. Geographic protein variation and speciation in the Plethodon dorsalis complex. Herpetologica 53:345–356.Google Scholar
  50. Highton, R. 1998a. Frequency of hybrids between introduced and native populations of the salamander Plethodon jordani in their first generation of sympatry. Herpetologica 54:143–153.Google Scholar
  51. Highton, R. 1998b. Is Ensatina eschscholtzii a ring-species? Herpetologica 54:254–278.Google Scholar
  52. Highton, R. 1999a. Geographic protein variation and speciation in the salamanders of the Plethodon cinereus group with the description of two new species. Herpetologica 55:43–90.Google Scholar
  53. Highton, R. 1999b. Hybridization in the contact zone between Plethodon richmondi and Plethodon electromorphus in northern Kentucky. Herpetologica 55:91–105.Google Scholar
  54. Highton, R. 2000. Detecting cryptic species using allozyme data. Pp. 215–241. In R. C. Bruce, R. G. Jaeger, and L. D. Houck (Eds.), The Biology of Plethodontid Salamanders. Kluwer Academic/Plenum Publishers, New York, New York, U.S.A.CrossRefGoogle Scholar
  55. Highton, R., and S. A. Henry. 1970. Evolutionary interactions between species of North American salamanders of the genus Plethodon. Part 2. Variation in the electrophoretic migration of plasma proteins of Plethodon jordani, P glutinosus, and their natural hybrids. Evolutionary Biology 4:241–256.Google Scholar
  56. Highton, R., and A. Larson. 1979. The genetic relationships of the salamanders of the genus Plethodon. Systematic Zoology 28:579–599.CrossRefGoogle Scholar
  57. Highton, R., and J. R. MacGregor. 1983. Plethodon kentucki Mittleman: a valid species of Cumberland Plateau woodland salamander. Herpetologica 39:189–200.Google Scholar
  58. Highton, R., and T. P. Webster. 1976. Geographic protein variation and divergence in populations of the salamander Plethodon cinereus. Evolution 30:33–45.CrossRefGoogle Scholar
  59. Key, K. H. L. 1981. Species, parapatry, and the morabine grasshoppers. Systematic Zoology 30:425–458.CrossRefGoogle Scholar
  60. Larson, A. 1984. Neontological inferences of evolutionary pattern and process in the salamander family Plethodontidae. Evolutionary Biology 17:119–217.CrossRefGoogle Scholar
  61. Larson, A., and R. Highton. 1978. Geographic protein variation and divergence in the salamanders of the Plethodon welleri group (Amphibia: Plethodontidae). Systematic Zoology 27:431–448.CrossRefGoogle Scholar
  62. Lazell, J. 1998. New salamander of the genus Plethodon from Mississippi. Copeia 1998:967–970.CrossRefGoogle Scholar
  63. Manzo, P. A. 1988. A Morphometric Analysis of Plethodon jordani Blatchley and Its Hybrids. M. S. Thesis, University of Maryland, College Park, Maryland, U.S.A.Google Scholar
  64. Maxson, L. R., R. Highton, and D. B. Wake. 1979. Albumin evolution and its phylogenetic implications in the plethodontid salamander genera Plethodon and Ensatina. Copeia 1979:502–508.CrossRefGoogle Scholar
  65. Maxson, L. R., and R. D. Maxson. 1979. Comparative albumin and biochemical evolution in plethodontid salamanders. Evolution 33:1057–1062.CrossRefGoogle Scholar
  66. Mittleman, M. B. 1948. American Caudata, V: notes on certain Appalachian salamanders of the genus Plethodon. Journal of the Washington Academy of Science 38:416–419.Google Scholar
  67. Mittleman, M. B. 1951. American Caudata. VII. Two new salamanders of the genus Plethodon. Herpetologica 7:105–112.Google Scholar
  68. Nei, M. 1972. Genetic distance between populations. American Naturalist 106:283–292.CrossRefGoogle Scholar
  69. Orr, H. A. 1996. Dobzhansky, Bateson, and the genetics of speciation. Genetics 144:1331–1335.PubMedGoogle Scholar
  70. Peabody, R. B. 1978. Electrophoretic Analysis of Geographic Variation of Two Appalachian Salamanders, Plethodon jordani and Plethodon glutinosus. Ph.D. Dissertation, University of Maryland, College Park, Maryland, U.S.A.Google Scholar
  71. Petranka, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, D.C., U.S.A.Google Scholar
  72. Pope, C. H., and N. G. Hairston. 1948. Two new subspecies of the salamander Plethodon shermani. Copeia 1948:106–107.CrossRefGoogle Scholar
  73. Pope, C. H., and S. H. Pope. 1951. A study of the salamander Plethodon ouachitae and the description of an allied form. Bulletin of the Chicago Academy of Science 9:129–152.Google Scholar
  74. Reagan, N. L. 1992. Evolution of Sexual Isolation in Salamanders of the Genus Plethodon. Ph.D. Dissertation, University of Chicago, Chicago, Illinois, U.S.A.Google Scholar
  75. Saitou, N., and M. Nei. 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4:406–425.PubMedGoogle Scholar
  76. Selander, R. R., M. H. Smith, S. Y. Yang, W. E. Johnson, and G. B. Gentry. 1971. Biochemical polymorphism and systematics in the genus Peromyscus. I. Variation in the old-field mouse (Peromyscus polionotus). Studies in Genetics VI, University of Texas Publications 7103:49–90.Google Scholar
  77. Shaw, C. R., and R. Prasad. 1970. Starch gel electrophoresis of enzymes-a compilation of recipes. Biochemical Genetics 4:297–320.PubMedCrossRefGoogle Scholar
  78. Smith-Gill, S. J. 1983. Developmental plasticity: developmental conversion versus phenotypic modulation. American Zoologist 23:47–55.Google Scholar
  79. Smith-Gill, S. J., and K. A. Berven. 1979. Predicting amphibian metamorphosis. American Naturalist 113:563–585.CrossRefGoogle Scholar
  80. Stejneger, L. 1906. A new salamander from North Carolina. Proceedings of the United States National Museum 30:559–562.CrossRefGoogle Scholar
  81. Swofford, D. L., and R. B. Selander. 1981. BIOSYS-1: a FORTRAN program for the comprehensive analysis of electrophoretic data in population genetics and systematics. Journal of Heredity 72:281–283.Google Scholar
  82. Templeton, A. R. 1989. The meaning of species and speciation: a genetic perspective. Pp. 2–38. In D. Otte and J. A. Endler (Eds.), Speciation and Its Consequences. Sinauer, Sunderland, Massachusetts, U.S.A.Google Scholar
  83. Thorpe, J. P. 1982. The molecular clock hypothesis: biochemical evolution, genetic differentiation, and systematics. Annual Review of Ecology and Systematics 13:139–168.CrossRefGoogle Scholar
  84. Tilley, S. G., and M. J. Mahoney. 1996. Patterns of genetic differentiation in salamanders of the Desmognathus ochrophaeus complex (Amphibia: Plethodontidae). Herpetological Monographs 10:1–42.CrossRefGoogle Scholar
  85. Wake, D. B. 1981. The application of allozyme evidence to problems in the evolution of morphology. Proceedings of the Second International Congress of Systematics and Evolutionary Biology:257–270.Google Scholar
  86. Wake, D. B., and C. J. Schneider. 1998. Taxonomy of the plethodontid salamander genus Ensatina. Herpetologica 54:279–298.Google Scholar
  87. Wynn, A. H. 1986. Linkage disequilibrium and a contact zone in Plethodon cinereus on the Del-Mar-Va peninsula. Evolution 40:44–54.CrossRefGoogle Scholar
  88. Wynn, A. H., R. Highton, and J. F. Jacobs. 1988. A new species of rock-crevice dwelling Plethodon from Pigeon Mountain, Georgia. Herpetologica 44:135–143.Google Scholar

Copyright information

© Springer Science+Business Media New York 2000

Authors and Affiliations

  • Richard Highton
    • 1
  • Robert B. Peabody
    • 1
  1. 1.Department of BiologyUniversity of MarylandCollege ParkUSA
  2. 2.Department of BiologyStonehill CollegeNorth EastonUSA

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