Abstract
Kingsnakes of the Lampropeltis getula complex range throughout much of North America. Using morphology and color pattern, Blaney made the last revision of this species complex nearly 30 years ago and recognized seven subspecies. Furthermore, Blaney hypothesized that populations in the eastern United States consist of two closely related taxa, L. g. getula & L. g. floridana, whichare morphologically divergent from all other subspecies. At the same time, Means hypothesized that an undescribed taxon existed in the Eastern Apalachicola Lowlands in the Florida panhandle. To test these hypotheses as well as help better understand phylogenetic relationships, we examine morphological characters and color pattern of L. getula throughout its range, particularly those populations in the eastern United States, and make comparisons to molecular data. We find that populations in the eastern United States represent a well-supported monophyletic group. Although some infraspecific clades (i.e., subspecies) within the L. getula complex may be weakly supported by homoplasious characters, at least one synapomorphy supports the monophyly of each group, including the two currently recognized subspecies in the eastern United States and the unnamed entity in the Eastern Apalachicola Lowlands, described herein as L. g. meansi. Justification for naming this natural clade at the infraspecific level (rather than species level) is provided. Furthermore, this panhandle clade is diagnosed by more synapomorphies than any other currently recognized taxon of L. getula, and overlaps in distribution with numerous other endemic plants and animals. All molecular analyses produced very similar tree topologies as our morphological dataset.References
Amos, B. & Hoelzel, A.R. (1991) Long-term preservation of whale skin for DNA analysis. Reports of the International Whaling Commission, Special Issue, 13, 99–103.
Arevalo, E., Davis, S.K. & Sites, J.W., Jr. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.
Armstrong, M.P., Frymire, D. & Zimmerer, E.J. (2001) Analysis of sympatric populations of Lampropeltis triangulum syspila and Lampropeltis triangulum elapsoides, in western Kentucky and adjacent Tennessee with relation to the taxonomic status of the scarlet kingsnake. Journal of Herpetology, 35, 688–693.
Avise, J.C. (1994) Molecular Markers, Natural History and Evolution. Chapman & Hall New York, New York, 511 pp.
Baum, D. (1992) Phylogenetic species concepts. Trends in Ecology and Evolution, 7, 1–2.
Baum, D.A. & Donoghue, M.J. (1995) Choosing among alternative “Phylogenetic” species concepts. Systematic Botany, 20, 560–573.
Baum, D.A., Donoghue, M.J. & Shaw, K.L. (1995) Genealogical perspectives on the species problem. In: Hoch, P.C. & Stephenson, A.G. (Eds.), Experimental and Molecular Approaches to Plant Biosystematics. Missouri Botanical Garden, St. Louis, Missouri, pp. 289–303.
Barbour, T. & Engels, W.L. (1942) Two interesting new snakes. Proceedings of the New England Zoological Club, 20, 101–104.
Bickham, J.W., Wood, C.C. & Patton, J.C. (1995) Biogeographic implications of cytochrome-b sequences and allozymes in sockeye (Oncorhynchus nerka). Journal of Heredity, 80, 140–144.
Blanchard, F.N. (1919) Two new snakes of the genus Lampropeltis. Occasional Papers of the Museum of Zoology, University of Michigan, 70, 1–11.
Blanchard, F.N. (1920) A synopsis of the king snakes: genus Lampropeltis Fitzinger. Occasional Papers of the Museum of Zoology, University of Michigan, 87, 1–8.
Blanchard, F.N. (1921) A revision of the king snakes: genus Lampropeltis. Bulletin of the U.S. National Museum, 114, 1–260.
Blaney, R.M. (1977) Systematics of the common kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47–103.
Blaney, R.M. (1979) The status of the Outer Banks kingsnake, Lampropeltis getulus sticticeps (Reptilia: Serpentes: Colubridae). Brimleyana, 1, 125–128.
Campbell, C.S. (1983) Systematics of the Andropogon virginicus complex (Gramineae). Journal of the Arnold Arboretum, 64, 171–254.
Campbell, C.S. (1986) Phylogenetic reconstructions and two new varieties in the Andropogon virginicus complex (Poaceae: Andropogoneae). Systematic Botany, 11, 280–292.
Clark, A. (1998) Reptile sheds yield high quality DNA. Herpetological Review, 29, 17–18.
Clewell, A.F. (1977) Geobotany of the Apalachicola River region. pp. 6–15. In: Livingston, R.J. & Joyce, E.A. (Eds.), Proceedings of the Conference on the Apalachicola Drainage System, Florida Marine Research Publication, 26.
Coile, N.C. (1996) Notes on Florida’s endangered and threatened plants. Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, Florida.
Conant, R. & Collins, J.T. (1998) A Field Guide to Reptiles and Amphibians of Eastern and Central North America. 4th ed. Houghton Mifflin Co., Boston, Massachusetts, 616 pp.
Davis, J.I. & Nixon, K.C. (1992) Populations, genetic variation, and the delimitation of phylogenetic species. Systematic Biology, 41, 421–435.
de Queiroz, K. & Donoghue, M.J. (1990) Phylogenetic systematics or Nelson’s version of cladistics? Cladistics, 6, 61–75.
Donoghue. M.J. (1985) A critique of the biological species concept and recommendations for a phylogenetic alternative. Bryologist, 88, 172–181.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.
Fetzner, J.W. Jr. (1999) Extracting high-quality DNA from shed reptile skins: a simplified method. BioTechniques, 26, 1052–1054.
Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological applications. Herpetologica, 46, 87–104.
Gilbert, C.R. (1987) Zoogeography of the freshwater fish fauna of southern Georgia and peninsular Florida. Brimleyana, 13, 225–244.
Hammond, P.C. (1990[1991]) Patterns of geographic variation and evolution in polytypic butterflies. Journal of Research on the Lepidoptera, 29, 54–76.
Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42, 182–192.
Hillis, D.M., Bull, J.J., Larson, A., Davis, S.K. & Zimmer, E.A. (1990) Nucleic acids III: Sequencing. In: Hillis, D.M. & Moritz, S (Eds.), Molecular Systematics. Sinauer Associates, Sunderland, Massachusetts, pp. 318–370.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
International Code of Zoological Nomenclature. (1999) The International Code of Zoological Nomenclature. 4th ed., London.
Iudica, C.A., Whitten, W.M. & Williams, N.H. (2001) Small bones from dried mammal museum specimens as a reliable source of DNA. BioTechniques, 30, 732–736.
James, C.W. (1961) Endemism in Florida. Brittonia, 13, 225–244.
Judd, W.S. (1982) The taxonomic status of Oxypolis greenmanii (Apiaceae). Rhodora, 84, 265–279.
Judd, W.S., Campbell, C.S., Kellogg, E.A., Stevens, P.F. & Donoghue, M.J. (2002) Plant Systematics: A Phylogenetic Approach. 2nd ed. Sinauer Associates, Inc., Sunderland, Massachusetts, 576 pp.
Keogh, J.S. (1996) Evolution of the colubrid snake tribe Lampropeltini: a morphological perspective. Herpetologica, 52, 406–416.
Kessing, B., Croom, H., Martin, A., McIntosh, C., McMillan, W.O. & Palumbi, S. (1989) The simple fool’s guide to PCR. Ver. 1.0. Department of Zoology, University of Hawaii, Honolulu.
Krysko, K.L. (1995) Resolution of the controversy regarding the taxonomy of the kingsnake, Lampropeltis getula, in southern Florida. M.S. Thesis, Florida International University, Miami, Florida. 73 pp.
Krysko, K.L. (2001) Ecology, conservation, and morphological and molecular systematics of the kingsnake, Lampropeltis getula (Serpentes: Colubridae). Ph.D. Dissertation, University of Florida, Gainesville, Florida. 159 pp.
Krysko, K.L. (2002) Seasonal activity of the Florida kingsnake, Lampropeltis getula floridana (Serpentes: Colubridae), in southern Florida. American Midland Naturalist, 148, 102–114.
Krysko, K.L. & Franz, R. (2003) Systematics and conservation of the kingsnake (Lampropeltis getula) in Florida. Final Report. Florida Fish and Wildlife Conservation Commission, Tallahassee, Florida.
Krysko, K.L. & Smith, D.J. (2005) The decline and extirpation of kingsnakes, Lampropeltis getula, in Florida. In: Meshaka, W.E., Jr. & Babbitt, K.J. (Eds.), Status and Conservation of Florida Amphibians and Reptiles. Krieger Publ.,Malabar, Florida, pp. 132–141.
Maddison, W.P., Donoghue, M.J. & Maddison, D.R. (1984) Outgroup analysis and parsimony. Systematic Zoology, 33, 83–103.
Mayr, E. (1969) Principles of Systematic Zoology. McGraw-Hill, New York, New York, 428 pp.
McDade, L.A. (1992) Hybrids and phylogenetic systematics II. The impact of hybrids on cladistic analysis. Evolution, 46, 1329–1346.
Means, D.B. (1977) Aspects of the significance to terrestrial vertebrates of the Apalachicola River drainage basin, Florida. In: Livingston, R.J. & Joyce, E.A. (Eds.), Proceedings of the Conference on the Apalachicola Drainage System, Florida Marine Research Publication, 26, pp. 37–67.
Means, D.B. (1978) Rare: Apalachicola populations of the eastern common kingsnake including L. g. goini. In: McDiarmid, R.W. (Ed.), Rare and Endangered Biota of Florida, Volume III, Amphibians and Reptiles. University Presses of Florida, Gainesville, Florida. pp. 60–61.
Means, D.B. (1992) Rare: eastern common kingsnake, Apalachicola population. In: Moler, P.E. (Ed.), Rare and Endangered Biota of Florida, Volume III, Amphibians and Reptiles. University Presses of Florida, Gainesville, Florida. pp. 232–236.
Means, D.B.& Krysko, K.L. (2001) Biogeography and pattern variation of kingsnakes, Lampropeltis getula, in the Apalachicola region of Florida. Contemporary Herpetology, 5. Available from http://www.selu.edu/journals/ContemporaryHerpetology/ch/2001/5/index.htm.
Mishler, B.D. (1985) The morphological, developmental, and phylogenetic basis of species concepts in bryophytes. Bryologist, 88, 207–214.
Mishler, B.D. & Brandon, R.N. (1987) Individuality, pluralism, and the phylogenetic species concept. Biology and Philosophy, 2, 397–414.
Mishler, B.D. & Theriot, E. (2000) The Phylogenetic Species Concept sensu Mishler and Theriot: monophyly, apomorphy, and phylogenetic species concepts. In: Wheeler, Q.D. & Meier, R. (Eds.), Species Concepts and Phylogenetic Theory: A Debate. Columbia University Press, New York, New York, pp. 44–54.
Neill, W.T. (1954) Juveniles of Brooks’ kingsnake, Lampropeltis getulus brooksi. Copeia, 1954, 59.
Neill, W.T. & Allen, E.R. (1949) A new kingsnake (genus Lampropeltis) from Florida. Herpetologica, 5, 1–12.
Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University.
O’Hara, R.J. (1993) Systematic generalization, historical fate, and the species problem. Systematic Biology, 42, 231–246.
Proebstel, D.S., Evans, R.P., Shiozawa, D.K. & Williams, R.N. (1993) Preservation of nonfrozen tissue samples from a salmonine fish Brachymystax lenok (Pallas) for DNA analysis. Journal of Ichthyology, 9, 9–17.
Rodriguez-Robles, J.A. & De Jesus-Escobar, J.M. (1999) Molecular systematics of New World lampropeltinine snakes (Colubridae): Implications for biogeography and evolution of food habits. Biological Journal of the Linnean Society, 68, 355–385.
Rodriguez-Robles, J.A., De Jesus-Escobar, J.M., Denardo, D.F. & Staub, R.E. (1999) Phylogeography of the California mountain kingsnake, Lampropeltis zonata (Colubridae). Molecular Ecology, 8, 1923–1934.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Saiki, R.K., Gelfand, D.H., Stoffel, S., Scharf, S.J., Higuchi, R., Horn, G.T., Mullis, K.B. & Erlich, H.A. (1988) Primer-directed enzymatic amplification of DNA with thermostable DNA polymerases. Science, 239, 487–491.
Smith, H.M., Chiszar, D. & Montanucci, R.R. (1997) Subspecies and classification. Herpetological Review, 28, 13–16.
Swofford, D.L. (2000) PAUP*: phylogenetic analysis using parsimony. Ver. 4.0b8. Sinauer Associates, Inc., Sunderland, Massachusetts.
Tennant, A. (1997) Field guide to the snakes of Florida. Gulf Publishing Co., Houston, Texas, 257 pp.
Wagner, W.H., Jr. (1980) Origin and philosophy of the groundplan-divergence method of cladistics. Systematic Botany, 5, 173–193.
Ward, D.B. (Ed.) (1979) Rare and Endangered Biota of Florida, Volume V, Plants. University Presses of Florida, Gainesville, Florida, 175 pp.
Wheeler, Q.D. & Platnick, N.I. (2000) The Phylogenetic Species Concept sensu Wheeler and Platnick. In: Wheeler, Q.D. & Meier, R. (Eds.), Species Concepts and Phylogenetic Theory: A Debate. Columbia University Press, New York, New York, pp. 55–69.
White, P.S. & Densmore, L.D. (1992) Mitochondrial DNA isolation. In: A.R. Hoezel (ed.), Molecular Genetic Analysis of Populations: A Practical Approach. IRL Press, Oxford Univ. Press, New York, pp. 29–58.
Wiley, E.O. & Mayden, R.L. (2000) The Evolutionary Species Concept sensu Wiley and Mayden. In: Wheeler, Q.D. & Meier, R. (Eds.), Species Concepts and Phylogenetic Theory: A Debate. Columbia University Press, New York, New York, pp. 70–89.
Wilken, D. & Hartman, R.L. (1991) A revision of the Ipomopsis spicata complex (Polemoniaceae). Systematic Botany, 16, 143–161.
Wilson, E.O. & Brown, W.L., Jr. (1953) The subspecies concept and its taxonomic application. Systematic Zoology, 2, 97–111.
Yerger, R.W. (1977) Fishes of the Apalachicola River. In: Livingston, R.J. & Joyce, E.A. (Eds.), Proceedings of the Conference on the Apalachicola Drainage System, Florida Marine Research Publication, 26, pp. 22–33.
Zhang, D.X. & Hewitt, G.M. (1996) Nuclear integrations: challenges for mitochondrial DNA markers. Trends in Ecology and Evolution, 11, 247–251.
Zharkikh, A. & Li, W.H. (1992) Statistical properties of bootstrap estimation of phylogenetic variability from nucleotide sequences. I. Four taxa with a molecular clock. Molecular Biology and Evolution, 9, 1119–1147.