Continental speciation in the tropics: Contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar

Journal of Zoology

Volumn 275, Issue 4, 2008, Pages 423-440

  Raxworthy, C J ^a   Pearson, R G ^a   Zimkus, B M ^a,d   Reddy, S ^a,e   Deo, A J ^a,b,f   Nussbaum, R A ^c   Ingram, C M ^a  

^a DIVISION OF INVERTEBRATE ZOOLOGY   (United States)

^b NEW YORK UNIVERSITY   (United States)

^c UNIVERSITY OF MICHIGAN   (United States)

^d HARVARD UNIVERSITY   (United States)

^e FIELD MUSEUM OF NATURAL HISTORY   (United States)

^f UNIVERSITY OF PITTSBURGH SCHOOL OF MEDICINE   (United States)

Author keywords

Biogeography; Gekkonidae; Madagascar; Reptilia; Speciation; Systematics

Indexed keywords

ADAPTIVE RADIATION; BIOGEOGRAPHY; DISPERSAL; DIVERGENCE; LIZARD; MIOCENE; NEW SPECIES; PHYLOGENY; RELICT SPECIES; SPECIATION (BIOLOGY); TAXONOMY; TERTIARY;

INDIAN OCEAN; INDIAN OCEAN ISLANDS; MADAGASCAR;

GEKKONIDAE; PAROEDURA; REPTILIA; UROPLATUS;

EID: 48249126864     PISSN: 09528369     EISSN: 14697998     Source Type: Journal    
DOI: 10.1111/j.1469-7998.2008.00460.x     Document Type: Article

References (113)

1. Andreone, F. Aprea, G. (2006). A new finding of Uroplatus alluaudi in north-eastern Madagascar widens considerably its distribution range (Reptilia, Gekkonidae). Acta Herpetol. 1, 121 125.
2. Angert, A.L. Schemske, D.W. (2005). The evolution of species' distributions: reciprocal transplants across the elevation ranges of Mimulus cardinalis and M. lewisii. Evolution 59, 1671 1684.
3. Austin, J.J., Arnold, E.N. Jones, C.G. (2004). Reconstructing an island radiation using ancient and recent DNA: the extinct and living day geckos (Phelsuma) of the Mascarene islands. Mol. Phyl. Evol. 31, 109 122.
4. Barraclough, T.G. Vogler, A.P. (2000). Detecting the geographical pattern of speciation from species-level phylogenies. Am. Nat. 155, 419 433.
5. Barraclough, T.G., Vogler, A.P. Harvey, P.H. (1998). Revealing the factors that promote speciation. Philos. Trans. Roy. Soc. Lond. Ser. B 353, 241 249.
6. Barraclough, T.G. Nee, S. (2001). Phylogenetics and speciation. TREE 16, 391 399.
7. Bauer, A.M. (1990). Phylogeny and biogeography of the geckos of southern Africa and the islands of the western Indian Ocean: a preliminary analysis. In Vertebrates in the tropics : 275 284. Peters, G. Hutterer, R. (Eds). Bonn : Museum Alexander Koenig.
8. Bauer, A.M., Jackman, T., Sadlier, R.A. Whitaker, A.H. (2006). A revision of the Bavayia validiclavis group (Squamata: Gekkota: Diplodactylidae), a clade of New Caledonian geckos exhibiting microendemism. Proc. Calif. Acad. Sci. 4th Ser. 57, 503 547.
9. Bauer, A.M. Russell, A.P. (1989). A systematic review of the genus Uroplatus (Reptilia: Gekkonidae), with comments on its biology. J. Nat. Hist. 23, 169 203.
10. Bermann, P.J. Russell, A. (2007). Systematics and biogeography of the widespread Neotropical gekkonid genus Thecadactylus (Squamata), with the description of a new cryptic species. Zool. J. Linn. Soc. 149, 339 370.
11. Besaire, H. (1972). Geologie de Madagascar I. Les terrains sédimentaires. Annales Géologiques de Madagascar 35, 1 463.
12. Böhme, A. Schönecker, P. (2003). Eine neue art der gattung Uroplatus Duméril, 1805 aus ost-Madagaskar (Reptilia: Squamata: Gekkonidae). Salamandra 31, 129 138.
13. Bouchard, P., Brooks, D.R. Yeates, D.K. (2005). Mosaic macroevolution in Australian wet tropic arthropods: community assemblage by taxon pulses. In Tropical rainforests. Past, present, and future : 425 469. Bermingham, E., Dick, C.W. Moritz, C. (Eds). Chicago : University of Chicago Press.
14. Brooks, D.R. McLennan, D.A. (1991). Phylogeny, ecology, and behavior. Chicago : University of Chicago Press.
15. Brooks, D.R., van Veller, M.G.P. McLennan, D.A. (2001). How to do BPA, really. J. Biogeog. 28, 345 358.
16. Brown, J.H. (2004). Concluding remarks. In Frontiers of biogeography. New directions in the geography of nature : 361 368. Lomolino, M.V. Heaney, L.R. (Eds). Sunderland, MA : Sinauer Associates.
17. Burney, D.A. (2003). Madagascar's prehistoric ecosystems. In The natural history of Madagascar : 47 51. Goodman, S.M. Bernstead, J.P. (Eds). Chicago : University of Chicago Press.
18. Butlin, R.K. (2004). Mystery of mysteries no longer? Evolution 58, 243 245.
19. Carranza, S., Arnold, E.N., Mateo, J.A. López-Juarado, L.F. (2000). Long-distance colonization and radiation in gekkonid lizards, Tarentola (Reptilia: Gekkonidae), revealed by mitochondrial DNA sequences. Proc. Roy. Soc. Lond. Ser. B 267, 637 649.
20. Chesser, R.T. Zink, R.M. (1994). Modes speciation in birds: a test of Lynch's method. Evolution 48, 490 497.
21. Coyne, J.A. Orr, H.R. (2004). Speciation. Sunderland, MA : Sinauer Associates.
22. Crisci, J.V., Katinas, L. Posadas, P. (2003). Historical biogeography. Cambridge, MA : Harvard University Press.
23. Cracraft, J. (1994). Species diversity, biogeography, and the evolution of biotas. Am. Nat. 34, 33 47.
24. Dimmick, W.W., Ghedotti, M.J., Grose, M.J., Maglia, A.M., Meinhardt, D.J. Pennock, D.S. (1999). The importance of systematic biology in defining units of conservation. Conserv. Biol. 13, 653 660.
25. Doebeli, M. Dieckmann, U. (2003). Speciation along environmental gradients. Nature 421, 259 264.
26. Doebeli, M., Dieckmann, U., Metz, J.A.J. Tautz, D. (2005). What we have also learned: adaptive speciation is theoretically plausible. Evolution 59, 691 695.
27. Drummond, A.J., Ho, S.Y.W., Phillips, M.J. Rambaut, A.J. (2006). Relaxed phylogenetics and dating with confidence. PloS Biol. 4, e88.
28. Drummond, A.J. Rambaut, A.J. (2007). 'BEAST': Bayesian evolutionary analysis by sampling trees. BMC Evol. Biol. 7, 214.
29. Duncan, R.A. Storey, M. (1992). The life cycle of Indian Ocean hotspots. Geophys. Monogr. 70, 91 103.
30. Elith, J., Graham, C., Anderson, R.P., Dudík, M., Ferrier, S., Guisan, A., Hijmans, R.J., Huettmann, F., Leathwick, J.R., Lehmann, A., Li, J., Lohmann, L.G., Loiselle, B.A., Manion, G., Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J.McC., Peterson, A.T., Phillips, S.J., Richardson, K., Scachetti-Pereira, R., Schapire, R.E., Soberón, J., Williams, S., Wisz, M.S. Zimmermann, N.E. (2006). Novel methods improve prediction of species' distributions from occurrence data. Ecography 29, 129 151.
31. Emerick, C.M. Duncan, R.A. (1982). Age progressive volcanism in the Comores Archipelago, Western Indian Ocean and implications for Somali plate tectonics. Earth Planet. Sci. Lett. 60, 415 428.
32. Endler, J.A. (1977). Geographic variation, speciation and clines. Princeton : Princeton University Press.
33. Fitzpatrick, B.M. Turelli, M. (2006). The geography of mammalian speciation: mixed signals from phylogenies and range maps. Evolution 60, 601 615.
34. Gavrilets, S. (2003). Perspective: models of speciation: what have we learned in 40 years? Evolution 57, 2197 2215.
35. Ghalambor, C.K., Huey, R.B., Marin, P.R., Tewksbury, J.J. Wang, G. (2006). Are mountain passes higher in the tropics? Janzen's hypothesis revisited. Int. Comp. Biol. 46, 5 17.
36. Giordano, A.R., Ridenhour, B.J. Storfer, A. (2007). The influence of altitude and topography on genetic structure in the long-toed salamander (Ambystoma macrodactylum). Mol. Ecol. 16, 1625 1637.
37. Gillot, P.-Y., Lefèvre, J.-C. Nativel, P.-E. (1994). Model for the structural evolution of the volcanoes of Réunion Island. Earth Planet. Sci. Lett. 122, 291 302.
38. Glaw, F., Kosuch, J., Henkel, F.-H., Sound, P. Böhme, W. (2006). Genetic and morphological variation of the leaf-tailed gecko Uroplatus fimbriatus from Madagascar, with description of a new species. Salamandra 42, 129 144.
39. Goodman, S.M. Bernstead, J.P. (Eds) 2003). The natural history of Madagascar. Chicago : University of Chicago Press.
40. Goodman, S.M., Raxworthy, C.J., Maminirina, C.P. Olson, L.E. (2006). A new species of Microgale (Family Tenrecidae) from the Tsaratanana Massif of northern Madagascar: description, phylogeny and biogeography. J. Zool. (Lond.) 270, 384 398.
41. Greenbaum, E., Bauer, A.M., Jackman, T.R.V. Glaw, F. (2007). A phylogeny of the enigmatic Madagascan geckos of the genus Uroplatus (Squamata: Gekkonidae). Zootaxa 1493, 41 51.
42. Graham, C.H., Ron, S.R., Santos, J.C., Schneider, C.J. Moritz, C. (2004). Integrating phylogenies and environmental niche models to explore speciation mechanisms in dendrobatid frogs. Evolution 58, 1781 1793.
43. Guisan, A. Thuiller, W. (2005). Predicting species distribution: offering more than simple habitat models. Ecol. Lett. 8, 993 1009.
44. Gübitz, T., Thorpe, R.S. Malhotra, A. (2005). The dynamics of genetic and morphological variation on volcanic islands. Proc. Roy. Soc. Lond. Ser. B 272, 751 757.
45. Hall, J.P.W. (2005). Montane speciation patterns in Ithomiola butterflies (Lepidoptera: Riodinidae): are they consistently moving up in the world? Proc. Roy. Soc. Lond. Ser. B 272, 2457 2466.
46. Hijmans, R.J., Cameron, S.E., Parra, J.L., Jones, P.G. Jarvis, A. (2005). Very high resolution interpolated climate surfaces for global land areas. Int. J. Climatol. 25, 1965 1978.
47. Hoffman, M.H. (2005). Evolution of the realized climatic niche in the genus Arabidopsis (Brassicaceae). Evolution 59, 1425 1436.
48. Hoskin, C.J., Higgie, M., McDonald, K.R. Moritz, C. (2005). Reinforcement drives rapid allopatric speciation. Nature 437, 1353 1356.
49. Huelsenbeck, J.P. Ronquist, F. (2001). MRBAYES. Bayesian inference of phylogeny. Bioinformatics 17, 754 755.
50. Hugall, A., Moritz, C., Moussalli, A. Stanisic, J. (2002). Reconciling paleodistribution models and comparative phylogeography in the Wet Tropics rainforest land snail Gnarosophia bellendenkerensis (Brazier 1875). Proc. Natl. Acad. Sci. USA 99, 6112 6117.
51. Humbert, H. (1955). Les térritoires phytogéographiques de Madagascar. Leur cartographie. Ann. Biol.3rd serie 31, 195 204.
52. Humphries, C.J. Parenti, L. (1999). Cladistic biogeography. 2nd edn: interpreting patterns of plant and animal distributions. Oxford : Oxford University Press.
53. Jenkins, M.D. (1987). Madagascar. An environmental profile. Gland, Switzerland : IUCN.
54. Jiggins, C.D., Mallarino, R., Willmott, K.R. Bermingham, E. (2006). The phylogenetic pattern of speciation and wing pattern change in neotropical Ithomia butterflies (Lepidoptera: Nymphalidae). Evolution 60, 1454 1466.
55. Johnson, N.K. Cicero, C. (2002). The role of ecological diversification in sibling speciation of Empidonax flycatchers (Tyrannidae): multigene evidence from mtDNA. Mol. Ecol. 11, 2065 2081.
56. Kluge, A.G. Nussbaum, R.A. (1995). A review of African-Madagascan gekkonid lizard phylogeny and biogeography (Squamata). Misc. Pubs. Mus. Zool. Univ. Mich. 183, 1 20.
57. Knouft, J.H., Losos, J.B., Glor, R.E. Kolbe, J.J. (2006). Phylogenetic analysis of the evolution of the niche in lizards of the Anolis sagrei group. Ecology 87 (Suppl.), 29 38.
58. Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. Wilson, A.C. (1989). Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proc. Natl. Acad. Sci. USA 86, 6196 6200.
59. Kozak, K.H., Graham, C.H. Wiens, J.J. (2008). Integrating GIS-based environmental data into evolutionary biology. TREE 23, 141 148.
60. Kozak, K.H. Wiens, J.J. (2006). Does niche conservatism promote speciation? A case study in North American salamanders. Evolution 60, 2604 2621.
61. Krause, D., Hartman, J.H. Wells, N.A. (1997). Late Cretaceous vertebrates from Madagascar: implications for biotic change in deep time. In Natural change and human impacts in Madagascar : 3 43. Goodman, S.M. Patterson, M. (Eds). Washington, DC : Smithsonian Press.
62. Kronauer, D.J.C., Bergmann, P.J., Mercer, J.M. Russell, A.P. (2005). A phylogeographically distinct and deep divergence in the widespread Neotropical turnip-tailed gecko, Thecadactylus rapicauda. Mol. Phyl. Evol. 34, 431 437.
63. Leaché, A.D. McGuire, J.A. (2006). Phylogenetic relationships of horned lizards (Phrynosoma) based on nuclear and mitochondrial data: evidence for a misleading mitochondrial gene tree. Mol Phyl. Evol. 39, 628 644.
64. Lessa, E.R., Cook, J.A. Patton, J.L. (2003). Genetic footprints of demographic expansion in North America, but not Amazonia, during the Late Quaternary. Proc. Natl. Acad. Sci. USA 100, 10331 10334.
65. Losos, J.B. Glor, R.E. (2003). Phylogenetic comparative methods and the geography of speciation. TREE 18, 220 227.
66. Losos, J.B., Leal, M., Glor, R.E., DeQueiroz, K. Hertz, P.E. (2003). Niche lability in the evolution of a Caribbean lizard community. Nature 424, 542 550.
67. Lukhtanov, V.A., Kandul, N.P., Plotkin, J.B., Dantchenko, A.V. Haig, D. (2005). Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies. Nature 436, 385 389.
68. Lynch, J.D. (1989). The gauge of speciation: on the frequencies of modes of speciation. In Speciation and its consequences : 527 553. Otte, D. Endler, J.A. (Eds). Sunderland, MA : Sinauer.
69. Martin, R.D. (1972). Adaptive radiation and behaviour of the Malagasy lemurs. Philos. Trans. Roy. Soc. (Lond.) B 264, 295 352.
70. Mayr, E. (1982). Speciation and macroevolution. Evolution 36, 119 1132.
71. McDougall, I. (1971). The geochronology and evolution of the young oceanic island of Réunion, Indian Ocean. Geochim. Cosmochim. Acta 35, 261 270.
72. Mindell, D.P., Sorenson, M.D., Huddleston, C.J., Miranda, H. Jr., Knight, C.A., Sawchuk, S.J. Tamaki, Y. (1997). Phylogenetic relationships among and within select avian orders based on mitochondrial DNA. In Avian molecular evolution and systematics : 213 247. Mindell, D.P. (Ed.). London : Academic Press.
73. Moritz, C., Patton, J.L., Schneider, C.J. Smith, T.B. (2000). Diversification of rainforest faunas: an integrated molecular approach. Annu. Rev. Ecol. Syst. 31, 533 563.
74. Nussbaum, R.A., Raxworthy, C.J. Pronk, O. (1998). The ghost geckos of Madagascar: a further revision of the Madagascan leaf-toed geckos (Reptilia, Squamata, Gekkonidae). Misc. Pubs. Univ. Mich. 186, 1 26.
75. Ogden, R. Thorpe, R.S. (2002). Molecular evidence for ecological speciation in tropical habitats. Proc. Natl. Acad. Sci. USA 99, 13612 13615.
76. Olsen, L.K., Goodman, S.M. Yoder, A.D. (2004). Illumination of crptic species boundaries in long-tailed shrew tenrecs (Mammalia: Tenrecidae; Microgale), with new insights into geographic variation and distributional constraints. Biol. J. Linn. Soc. 83, 1 22.
77. Pastorini, J., Thalmann, U. Martin, R.D. (2005). A molecular approach to comparative phylogeography of extant Malagasy lemurs. Proc. Natl. Acad. Sci. USA 100, 5879 5884.
78. Pearson, R.G., Raxworthy, C.J., Nakamura, M. Peterson, A.T. (2007). Predicting species' distributions from small numbers of occurrence records: a test case using cryptic geckos in Madagascar. J. Biogeogr. 34, 102 117.
79. Peterson, A.T. Holt, R.D. (2003). Niche differentiation in Mexican birds: using point occurrences to detect ecological innovation. Ecol. Lett. 6, 774 782.
80. Peterson, A.T., Soberon, J. Sanchez-Cordero, V. (1999). Conservatism of ecological niches in evolutionary time. Science 285, 1265 1267.
81. Posada, D. Crandall, K.A. (1998). Modeltest: testing the model of DNA substitution. Bioinfomatics 14, 817 818.
82. Phillips, S.J., Anderson, R.P. Schapire, R.E. (2006). Maximum entropy modeling of species geographic distributions. Ecol. Model. 190, 231 259.
83. de Queiroz, A. (2005). The resurrection of oceanic dispersal in historical biogeography. TREE 20, 68 73.
84. Raxworthy, C.J. (2003). Introduction to the reptiles. In The natural history of Madagascar : 934 949. Goodman, S.M. Bernstead, J.P. (Eds). Chicago : University of Chicago Press.
85. Raxworthy, C.J., Ingram, C., Rabibisoa, N. Pearson, R. (2007). Applications of ecological niche modeling for species delimitation: a review and empirical evaluation using day Geckos (Phelsuma) from Madagascar. Syst. Biol. 56, 907 923.
86. Raxworthy, C.J. Nussbaum, R.A. (1997). Biogeographic patterns of reptiles in Eastern Madagascar. In Natural change and human impact in Madagascar : 124 141. Goodman, S.M. Patterson, B.D. (Eds). Washington : Smithsonian Press.
87. Rice, N.H., Martínez-Meyer, E. Peterson, A.T. (2003). Ecological niche differentiation in the Aphelocoma jays: a phylogenetic perspective. Biol. J. Linn. Soc. 80, 369 383.
88. Ronquist, F. (1996). DIVA, v. 1.1. Computer program for MacOS and Win32. Available from ()
89. Ronquist, F. (1997). Dispersal-vicariance analysis: a new approach to the quantification of historical biogeography. Syst. Biol. 46, 195 203.
90. Royer, J.Y., Sclater, J.G., Sandwell, D.T., Cande, S.C., Schlich, R., Munschy, M., Dyment, J., Fisher, R.L., Muller, R.D., Coffin, M.F., Patriat, P. Berg, H.G. (1992). Appendix 1 - Indian Ocean plate reconstructions since the late Jurassic. In Synthesis of results from scientific drilling in the Indian Ocean. Geophysical Monograph 70 : 471 475. Duncan, R.A., Rea, D.K., Kidd, R.B., von Radd, U. Weissel, J.K. (Eds). Washington, DC : American Geophysical Union.
91. Schluter, D. (2001). Ecology and the origin of species. TREE 16, 372 380.
92. Schneider, C.J., Smith, T.B. Moritz, C. (1999). A test of alternative models of diversification in tropical rainforests: ecological gradients vs. rainforest refugia. Proc. Natl. Acad. Sci. USA 96, 13869 13873.
93. Shimodaira, H. Hasegawa, M. (1999). Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Mol. Biol. Evol. 16, 1114 1116.
94. Smith, A.G., Smith, D.G. Funnell, B.M. (1994). Atlas of Mesozoic and Cenzoic coastlines. Cambridge : Cambridge University Press.
95. Smith, T.B., Wayne, R.K., Girman, D. Bruford, M.W. (2005). Populations: the importance of ecotones in rainforest speciation. In Tropical rainforests. Past, present, and future : 148 165. Bermingham, E., Dick, C.W. Moritz, C. (Eds). Chicago : Chicago Press.
96. Sorenson, M.D., Ast, J.C., Dimcheff, D.E., Yuri, T. Mindell, D.P. (1999). Primers for a PCR-based approach to mitochondrial genome sequencing in birds & other vertebrates. Mol. Phyl. Evol. 12, 105 114.
97. Stiassney, M.L.J. dePinna, M.C.C. (1994). Basal taxa and the role of cladistic patterns in the evaluation of conservation priorities: a view from freshwater. In Systematics and conservation evaluation : 235 249. Forey, P.I., Humphries, C.J. Vane-Wright, I. (Eds). Oxford : Clarendon Press.
98. Storey, M., Mahoney, J.J., Saunders, A.D., Duncan, R.A., Kelly, S.P. Coffins, M.F. (1995). Timing of hot spot-related volcanism and the breakup of Madagascar and India. Science 267, 852 855.
99. *: Phylogenetic analysis using parsimony and other methods, version 4. Sunderland, MA : Sinauer.
100. Torsvik, T.H., Tucker, R.D. Ashwal L, D., et al 2000). Late Cretaceous India-Madagascar fit and timing of break-up related magmatism. Terra Nova 12, 220 225.
101. Turelli, M., Barton, N.H. Coyne, J.A. (2001). Theory and speciation. TREE 16, 330 343.
102. Via, S. (2001). Sympatric speciation in animals: the ugly duckling grows up. TREE 16, 381 390.
103. Wells, N.A. (2003). Some hypotheses on the Mesozoic and Cenzoic paleoenvironmental history of Madagascar. In The natural history of Madagascar : 16 34. Goodman, S.M. Bernstead, J.P. (Eds). Chicago : University of Chicago Press.
104. Wiens, J.J. Graham, C. (2005). Niche conservatism: integrating evolution, ecology, and conservation biology. Annu. Rev. Ecol. Syst. 36, 519 539.
105. Wiens, J.J., Reeder, T.W. Montes De Oca, A.N. (1999). Molecular phylogenetics and evolution of sexual dichromatism among populations of the Yarrow's spiny lizard (Sceloporus jarrovii). Evolution 53, 1884 1897.
106. Wiley, E.O. (1988). Parsimony analysis and vicariance biogeography. Syst. Zool. 37, 271 290.
107. Wilme, L., Goodman, S.M. Ganzhorn, J.U. (2006). Biogeographic evolution of Madagascar's microendemic biota. Science 312, 1063 1065.
108. Yang, Z. (1994). Maximum likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods. J. Mol. Evol. 39, 306 314.
109. Yesson, C. Culham, A. (2006). Phyloclimatic modeling: combining phylogenetics and bioclimatic modeling. Syst. Biol. 55, 785 802.
110. Yoder, A.D. Nowak, M.D. (2006). Has vicariance or dispersal been the predominant biogeographic force in Madagascar?: only time will tell. Annu. Rev. Ecol. Syst. 37, 405 431.
111. Yoder, A.D., Olson, L.E., Hanley, C., Heckman, K.L., Rasoloarison, R., Russell, A.L., Ranivo, J., Soarimalala, V., Karanth, K.P., Raselimananana, A.P. Goodman, S.M. (2005). A multidimensional approach for detecting species patterns in Malagasy vertebrates. Proc. Natl. Acad. Sci. USA 102, 6587 6594.
112. Yoder, A.D., Rasoloarison, R.M., Goodman, S.M., Irwin, J.A., Atsalis, S., Ravosa, M.J. Ganzhorn, J.U. (2000). Remarkable species diversity in Malagasy mouse lemurs (primates, Microcebus). Proc. Natl. Acad. Sci. USA 97, 11325 11330.
113. Zwickl, D.J. (2006). Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. PhD dissertation, The University of Texas at Austin.