Are Toad-Headed Lizards Phrynocephalus przewalskii and P. frontalis (Family Agamidae) the Same Species? Defining Species Boundaries with Morphological and Molecular Data

Agnes Gozdzik, Jinzhong Fu

Abstract


Toad-headed lizards of the Phrynocephalus przewalskii complex provide a challenging case for delimiting species boundaries. We tested the species status of P. przewalskii and P. frontalis using mitochondrial DNA (mtDNA) sequence and morphological data. A phylogenetic analysis was applied to the mtDNA data and principal component analysis (PCA) was applied to the morphological data. Furthermore, Mantel tests were used to test congruence between the patristic distance matrix derived from the phylogenetic tree and Euclidean distance matrix derived from PCA. The phylogenetic tree presented deeply diverged discreet clades that largely correspond to the two putative species. Nevertheless, PCA revealed no distinct clustering of individuals. Unique maternal inheritance might explain the discrete mtDNA variations while nuclear gene based morphological variations were continuous. Mantal tests suggested the mtDNA and morphology diverged in concordance; both had evidence of a west to east clinal variation. We conclude that P. frontalis is a synonym of P. przewalskii. Furthermore, the Mantel test is a useful method to compare mtDNA data with morphological data, but insufficient to delimit species boundaries.

Keywords


species validity; Mantel test; mtDNA sequence; phylogeny; PCA; multivariate morphometrics; Phrynocephalus

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References


Arnold E. N. (1999), «Phylogenetic relationships of toad-headed lizards (Phynocephalus, Agamidae) based on morphology», Bull. Nat. Mus. Lond. Zool., 65, 1 – 13.

Avise J. C. and Ball R. J., Jr. (1990), «Principles of genealogical concordance in species concepts and biological taxonomy», Oxford Surveys Evol. Biol., 7, 45 – 67.

Barabanov A. V. and Ananjeva N. B. (2007), «Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae)», Zootaxa, 1399, 1 – 56.

Daltry J. C., Wuster W., and Thorpe R. S. (1996), «Diet and snake venom evolution», Nature, 379, 537 – 540.

Felsenstein J. (1985), «Confidence limits on phylogenies — An approach using the bootstrap», Evolution, 39, 783 – 791.

Fu J. and Zeng X. (2008), «How many species are in the genus Batrachuperus? A phylogeographical analysis of the stream salamanders (family Hynobiidae) from southwestern China», Mol. Ecol., 17, 1469 – 1488.

Funk D. J. and Omland K. E. (2003), «Species-level paraphyly and polyphyly: Frequency, causes, and consequences, with insights from animal mitochondrial DNA», Ann. Rev. Ecol. Evol. Syst., 34, 397 – 423.

Graybeal A. (1995), «Naming Species», Syst. Biol., 44, 237 – 250.

Hebert P. D. N., Penton E. H., Burns J. M., Janzen D. H., and Hallwachs W. (2004), «Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator», Proc. Natl. Acad. Sci. USA, 101, 14812 – 14817.

Knowles L. L. and Carstens B. (2007), «Delimiting species without monophyletic gene trees», Syst. Biol., 56, 887 – 895.

Maddison W. P. and Maddison D. R. (2003), MacClade: Analysis of Phylogeny and Character Evolution, Sinauer Associates, Sunderland.

Masters J. C. and Bragg N. P. (2000), «Morphological correlates of speciation in bush babies», Int. J. Primatol., 21, 793 – 813.

Montanucci R. R. (2004), «Geographic variation in Phrynosoma coronatum (Lacertilia, Phrynosomatidae): Further evidence for a Peninsular Archipelago», Herpetologica, 60, 117 – 139.

Nylander J. A. A. (2004), MrModeltest version 2.1 (Computer program distributed by the author), Uppsala University, Uppsala.

Olmstead R. G. (1995), «Species concepts and plesiomorphic species», Syst. Bot., 20, 623 – 630.

Pang J., Wang Y., Zhong Y., Hoelzel A. R., Papenfuss T. J., Zeng X., Ananjeva N. B., and Zhang Y. P. (2003), «A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences», Mol. Phylogen. Evol., 27, 398 – 409.

Pope C. H. (1935), The Reptiles of China, Am. Mus. Nat. Hist., New York.

Puorto G., Salomao M. D., Theakston R. D. G., Thorpe R. S., Warrell D. A., and Wüster W. (2001), «Combining mitochondrial DNA sequences and morphological data to infer species boundaries: Phylogeography of lanceheaded pitvipers in the Brazilian Atlantic forest, and the status of Bothrops pradoi (Squamata: Serpentes: Viperidae)», J. Evol. Biol., 14, 527 – 538.

Ronquist F. and Huelsenbeck J. P. (2003), «MRBAYES 3: Bayesian phylogenetic inference under mixed models», Bioinformatics, 19, 1572 – 1574.

Roberts M. A., Schwartz T. S., and Karl S. A. (2004), «Global population genetic structure and male-mediated gene flow in the green sea turtle (Chelonia mydas): Analysis of microsatellite loci», Genetics, 166, 1857 – 1870.

Sites J. W., Jr. and Marshall J. C. (2004), «Operational criteria for delimiting species», Ann. Rev. Ecol. Evol. Syst., 35, 199 – 227.

Strauch A. (1876), «Part III: Reptilia and Amphibia», in: Przehevalsky N. M. (ed.), Mongolia and the Rangut County. Vol. 2, Russian Imperial Geographic Society, St. Petersburg [in Russian].

Swofford D. L. (2002), PAUP*: Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4.0b10, Sinauer Associates, Sunderland.

Templeton A. R. (2001), «Using phylogeographic analyses of gene trees to test species status and processes», Mol. Ecol. 10, 779 – 791.

Wang Y. and Fu J. (2004), «Cladogenesis and vicariance patterns in the toad-headed lizard Phrynocepahlus versicolor species complex», Copeia, 2004, 199 – 206.

Wang Y. and Wang H. (1993), «Geographic variation and diversity in three species of Phrynocephalus in the Tengger Desert, Western China», Asiatic Herpetol. Res., 5, 65 – 73.

Wiens J. J. and Penkrot T. A. (2002), «Delimiting species using DNA and morphological variation and discordant species limits in spiny lizards (Sceloporus)», Syst. Biol., 51, 69 – 91.

Zhao E., Zhao K., and Zhou K. (1999), Fauna Sinica. Reptilia Vol. 2. Squamata, Lacertilia, Science Press, Beijing [in Chinese].




DOI: https://doi.org/10.30906/1026-2296-2009-16-2-107-118

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