Quantifying ecological, morphological, and genetic variation to delimit species in the coast horned lizard species complex (Phrynosoma). Leaché, A. D., Koo, M. S., Spencer, C. L., Papenfuss, T. J., Fisher, R. N., & McGuire, J. A. Proceedings of the National Academy of Sciences, 106(30):12418–12423, July, 2009.
Quantifying ecological, morphological, and genetic variation to delimit species in the coast horned lizard species complex (Phrynosoma) [link]Paper  doi  abstract   bibtex   
Lineage separation and divergence form a temporally extended process whereby populations may diverge genetically, morphologically, or ecologically, and these contingent properties of species provide the operational criteria necessary for species delimitation. We inferred the historical process of lineage formation in the coast horned lizard () species complex by evaluating a diversity of operational species criteria, including divergence in mtDNA (98 specimens; 2,781 bp) and nuclear loci (−, 1,054 bp; 529 bp), ecological niches (11 bioclimatic variables; 285 unique localities), and cranial horn shapes (493 specimens; 16 landmarks). A phylogenetic analysis of mtDNA recovers 5 phylogeographic groups arranged latitudinally along the Baja California Peninsula and in California. The 2 southern phylogeographic groups exhibit concordance between genetic, morphological, and ecological divergence; however, differentiation is weak or absent at more recent levels defined by phylogeographic breaks in California. Interpreting these operational species criteria together suggests that there are 3 ecologically divergent and morphologically diagnosable species within the complex. Our 3-species taxonomic hypothesis invokes a deep coalescence event when fitting the mtDNA genealogy into the species tree, which is not unexpected for populations that have diverged recently. Although the hypothesis that the 3 phylogeographic groups distributed across California each represent distinctive species is not supported by all of the operational species criteria evaluated in this study, the conservation status of the imperiled populations represented by these genealogical units remains critical.
@article{leache_quantifying_2009,
	title = {Quantifying ecological, morphological, and genetic variation to delimit species in the coast horned lizard species complex ({Phrynosoma})},
	volume = {106},
	url = {http://www.pnas.org/content/106/30/12418.abstract},
	doi = {10.1073/pnas.0906380106},
	abstract = {Lineage separation and divergence form a temporally extended process whereby populations may diverge genetically, morphologically, or ecologically, and these contingent properties of species provide the operational criteria necessary for species delimitation. We inferred the historical process of lineage formation in the coast horned lizard () species complex by evaluating a diversity of operational species criteria, including divergence in mtDNA (98 specimens; 2,781 bp) and nuclear loci (−, 1,054 bp; 529 bp), ecological niches (11 bioclimatic variables; 285 unique localities), and cranial horn shapes (493 specimens; 16 landmarks). A phylogenetic analysis of mtDNA recovers 5 phylogeographic groups arranged latitudinally along the Baja California Peninsula and in California. The 2 southern phylogeographic groups exhibit concordance between genetic, morphological, and ecological divergence; however, differentiation is weak or absent at more recent levels defined by phylogeographic breaks in California. Interpreting these operational species criteria together suggests that there are 3 ecologically divergent and morphologically diagnosable species within the complex. Our 3-species taxonomic hypothesis invokes a deep coalescence event when fitting the mtDNA genealogy into the species tree, which is not unexpected for populations that have diverged recently. Although the hypothesis that the 3 phylogeographic groups distributed across California each represent distinctive species is not supported by all of the operational species criteria evaluated in this study, the conservation status of the imperiled populations represented by these genealogical units remains critical.},
	number = {30},
	urldate = {2009-08-04TZ},
	journal = {Proceedings of the National Academy of Sciences},
	author = {Leaché, Adam D. and Koo, Michelle S. and Spencer, Carol L. and Papenfuss, Theodore J. and Fisher, Robert N. and McGuire, Jimmy A.},
	month = jul,
	year = {2009},
	keywords = {Phrynosoma, Phylogeography, Species delimitation},
	pages = {12418--12423}
}

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