Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2019-12-19
Page range: 151–201
Abstract views: 174
PDF downloaded: 9

The sprightly little sphaerodactyl: Systematics and biogeography of
the Puerto Rican dwarf geckos Sphaerodactylus (Gekkota, Sphaerodactylidae)

Department of Biological Sciences, Sam Houston State University, Huntsville, TX 77341, USA
Department of Biological Sciences, Marquette University, Milwaukee, WI 53233, USA. Milwaukee Public Museum, Milwaukee, WI 53233, USA.
University of Puerto Rico Río Piedras, Río Piedras, PR, 003931, Puerto Rico.
Texas Invasive Species Institute, Sam Houston State University, Huntsville TX, 77341, USA.
Resonate, Reston VA 20190, USA.
University of Puerto Rico Bayamón, Bayamón, PR, 00959, Puerto Rico.
Department of Biological Sciences, Sam Houston State University, Huntsville, TX 77341, USA
Texas Tech University, Lubbock, TX 79409, USA.
Department of Biological Sciences, Marquette University, Milwaukee, WI 53233, USA. Milwaukee Public Museum, Milwaukee, WI 53233, USA. Bell Museum of Natural History, University of Minnesota, St Paul, MN 55113, USA.
Reptilia biogeography Caribbean Hispaniola lizard morphology Puerto Rico taxonomy species delimitation Squamata Virgin Islands

Abstract

Studies of the Caribbean herpetofauna (amphibians and reptiles) have made significant contributions to our knowledge of evolutionary patterns and processes. A prerequisite for these studies are accurate taxonomies and robust phylogenetic hypotheses. One notable Caribbean radiation lacking such data are dwarf geckos of the genus Sphaerodactylus. Systematics of the Puerto Rican Sphaerodactylus have been turbulent since the initial species descriptions and no molecular phylogenies exist that include complete or near-complete taxon sampling. Here, we combine a multi-locus molecular phylogeny with extensive morphological information to investigate the current diversity of Sphaerodactylus geckos from the Puerto Rican Bank, with a large number of species from Hispaniola as an outgroup. In particular, we focus our efforts on resolving the taxonomy of the Sphaerodactylus macrolepis Günther species complex. We find S. macrolepis sensu lato (currently two nominal species with nine subspecies) is made up of at least four diagnosable species within two clades: (1) the sister species Sphaerodactylus macrolepis sensu stricto from the Virgin Islands (including St. Croix) and Culebra, and S. parvus King from islands in the northern Lesser Antilles; and (2) all other Sphaerodactylus macrolepis subspecies from Puerto Rico, Vieques, and Culebra. We resurrect Sphaerodactylus grandisquamis Stejneger from synonymy to refer to all subspecies from Puerto Rico and elevate the subspecies Sphaerodactylus inigoi Thomas & Schwartz for geckos from Vieques and western Culebra. The resulting phylogeny and revised taxonomy will be a useful tool for subsequent research into Sphaerodactylus conservation and evolution.

 

References

  1. Barbour, T. (1921) Sphaerodactylus. Memoirs of the Museum of Comparative Zoology, 47, 217–278, plates 1–26.

    Barbour, T. (1937) Third list of Antillean reptiles and amphibians. Bulletin of the Museum of Comparative Zoology, 82, 77–166.

    Barker, B.S., Rodríguez-Robles, J.A., Aran, V.S., Montoya, A., Waide, R.B. & Cook, J.A. (2012) Sea level, topography and island diversity: Phylogeography of the Puerto Rican Red-eyed Coquí, Eleutherodactylus antillensis. Molecular Ecology, 21, 6033–6052.

    https://doi.org/10.1111/mec.12020

    Barley, A.J., Brown, J.M. & Thomson, R.C. (2018). Impact of model violations on the inference of species boundaries under the multispecies coalescent. Systematic Biology, 67, 269–284.

    https://doi.org/10.1093/sysbio/syx073

    Bauer, A.M. & Lamb, T. (2002) Phylogenetic relationships among members of the Pachydactylus capensis group of southern African geckos. African Zoology, 37, 209–220.

    https://doi.org/10.1080/15627020.2002.11657176

    Bennett, K.D. (2004) Continuing the debate on the role of Quaternary environmental change for macroevolution. Philosophical Transactions of the Royal Society B: Biological Sciences, 359, 295–303.

    https://doi.org/10.1098/rstb.2003.1395

    Botov, A., Phung, T.M., Ngyyen, T.Q., Bauer, A.M., Brennan, I.G. & Ziegler, T. (2015) A new species of Dixonius (Squamata: Gekkonidae) from Phu Quy Island, Vietnam. Zootaxa, 4040 (1), 048–058.

    https://doi.org/10.11646/zootaxa.4040.1.4

    Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.-H., Xie, D., Suchard, M.A, Rambaut, A. & Drummond, A.J. (2014) BEAST 2: A Software Platform for Bayesian Evolutionary Analysis. PLOS Computational Biology, 10, e1003537.

    https://doi.org/10.1371/journal.pcbi.1003537

    Bouckaert, R.R. & Heled, J. (2014) DensiTree 2: Seeing trees through the forest. bioRxiv. [posted online]

    https://doi.org/10.1101/012401

    Burbrink, F.T., Lawson, R. & Slowinski, J.B. (2000) Mitochondrial DNA phylogeography of the polytypic North American ratsnake (Elaphe obsoleta): A critique of the subspecies concept. Evolution, 54, 2107–2118.

    https://doi.org/10.1111/j.0014-3820.2000.tb01253.x

    Carlton, J.T., Chapman, J.W., Geller, J.B., Miller, J.A., Carlton, D.A., McCuller, M.I., Treneman, N.C., Steves, B.P. & Ruiz, G.M. (2017) Tsunami-driven rafting: Transoceanic species dispersal and implications for marine biogeography. Science, 357, 1402–1406.

    https://doi.org/10.1126/science.aao1498

    Darwin, C. (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. John Murray, London, 502 pp.

    https://doi.org/10.5962/bhl.title.68064

    Daza, J.D. & Bauer, A.M (2012) A new amber-embedded sphaerodactyl gecko from Hispaniola, with comments on morphological synapomorphies of the Sphaerodactylidae. Breviora, 2012, 1–28.

    https://doi.org/10.3099/529.1

    Daza, J.D., Bauer, A.M., Wagner, P. & Böhme, W. (2013) A reconsideration of Sphaerodactylus dommeli Böhme, 1984 (Squamata: Gekkota: Sphaerodactylidae), a Miocene lizard in amber. Journal of Zoological Systematics and Evolutionary Research, 51, 55–63.

    https://doi.org/10.1111/jzs.12001

    de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H., (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, pp. 57–75.

    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.

    https://doi.org/10.1080/10635150701701083

    Díaz-Lameiro, A.M., Oleksyk, T., Bird-Picó, F.J. & Martínez-Cruzado, J.C. (2013) Colonization of islands in the Mona Passage by endemic dwarf geckoes (genus Sphaerodactylus) reconstructed with mitochondrial phylogeny. Ecology and Evolution, 3, 4488–4500.

    https://doi.org/10.1002/ece3.770

    Dugo-Cota, Á., Vilà, C., Rodríguez, A. & Gonzalez-Voyer, A. (2019) Ecomorphological convergence in Eleutherodactylus frogs: A case of replicate radiations in the Caribbean. Ecology Letters, 22, 884–893.

    https://doi.org/10.1111/ele.13246

    Dunson, W.A. & Bramham, C.R. (1981) Evaporative water loss and oxygen consumption of three small lizards from the Florida Keys: Sphaerodactylus cinereus, S. notatus, and Anolis sagrei. Physiological Zoology, 54, 253–259.

    https://doi.org/10.1086/physzool.54.2.30155827

    Edgar, R.C. (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.

    https://doi.org/10.1093/nar/gkh340

    Ehlers, J. & Gibbard, P.L. (2007) The extent and chronology of Cenozoic global glaciation. Quaternary International, 164–165, 6–20.

    https://doi.org/10.1016/j.quaint.2006.10.008

    Erickson, E.E., Helmer, E.H., Brandeis, T.J. & Lugo, A.E. (2014) Controls on fallen leaf chemistry and forest floor element masses in native and novel forests across a tropical island. Ecosphere, 5, 1–48.

    https://doi.org/10.1890/ES13-00263.1

    Gamble, T., Bauer, A.M., Greenbaum, E. & Jackman, T.R. (2008a) Out of the blue: A novel, trans-Atlantic clade of geckos (Gekkota, Squamata). Zoologica Scripta, 37, 355–366.

    https://doi.org/10.1111/j.1463-6409.2008.00330.x

    Gamble, T., Simons, A.M., Colli, G.R. & Vitt, L.J. (2008b) Tertiary climate change and the diversification of the Amazonian gecko genus Gonatodes (Sphaerodactylidae, Squamata). Molecular Phylogenetics and Evolution, 46, 269–277.

    https://doi.org/10.1016/j.ympev.2007.08.013

    Gamble, T., Daza, J.D., Colli, G.R., Vitt, L.J. & Bauer, A.M. (2011) A new genus of miniaturized and pug-nosed gecko from South America (Sphaerodactylidae: Gekkota). Zoological Journal of the Linnean Society, 163, 1244–1266.

    https://doi.org/10.1111/j.1096-3642.2011.00741.x

    Gamble, T., Colli, G.R., Rodrigues, M.T., Werneck, F.P. & Simons, A.M. (2012) Phylogeny and cryptic diversity in geckos (Phyllopezus; Phyllodactylidae; Gekkota) from South America’s open biomes. Molecular Phylogenetics and Evolution, 62, 943–953.

    https://doi.org/10.1016/j.ympev.2011.11.033

    Gamble, T., Greenbaum, E., Jackman, T.R. & Bauer, A.M. (2015) Into the light: Diurnality has evolved multiple times in geckos. Biological Journal of the Linnean Society, 115, 896–910.

    https://doi.org/10.1111/bij.12536

    Geurgas, S.R. & Rodrigues, M.T. (2010) The hidden diversity of Coleodactylus amazonicus (Sphaerodactylinae, Gekkota) revealed by molecular data. Molecular Phylogenetics and Evolution, 54, 583–593.

    https://doi.org/10.1016/j.ympev.2009.10.004

    Gillespie, R. (2004) Community assembly through adaptive radiation in Hawaiian spiders. Science, 303, 356–359.

    https://doi.org/10.1126/science.1091875

    Grant, C. (1931) The Sphaerodactylus of Porto Rico, Culebra and Mona Islands. Journal of the Department of Agriculture of Porto Rico, 15, 199–213.

    Grant, C. (1932a) Sphaerodactylus grandisquamis, a valid species. Journal of the Department of Agriculture of Porto Rico, 16, 43–45.

    Grant, C. (1932b) Chart for determining the Sphaerodactylus of the Porto Rico region. Journal of the Department of Agriculture of Porto Rico, 16, 33–36.

    Grant, C. (1932c) The herpetology of Vieques Island. Journal of the Department of Agriculture of Porto Rico, 16, 37–39.

    Grant, C. & Beatty, H.A. (1944) Herpetological notes on St. Croix, Virgin Islands. Herpetologica, 2, 110–113.

    Grant, P.R. & Grant, B.R. (2011) How and Why Species Multiply: The Radiation of Darwin’s Finches. Princeton University Press, Princeton, , New Jersey, 272 pp.

    Grismer, L.L., Wood Jr., P.L., Anuar, S., Riyanto, A., Ahmad, N., Muin, M.A., Sumontha, M., Grismer, J.L., Onn, C.K., Quah, E.S.H. & Pauwels, O.S.A. (2014) Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia,Thailand, and Indonesia. Zootaxa, 3880 (1), 1–147.

    https://doi.org/10.11646/zootaxa.3880.1.1

    Guarnizo, C.E., Werneck, F.P., Giugliano, L.G., Santos, M.G., Fenker, J., Sousa, L., D’Angiolella, A.B., Santos, A.R.d., Strüssmann, C., Rodrigues, M.T., Dorado-Rodrigues, T.F., Gamble, T. & Colli, G.R. (2016) Cryptic lineages and diversification of an endemic anole lizard (Squamata, Dactyloidae) of the Cerrado hotspot. Molecular Phylogenetics and Evolution, 94, 279–289.

    https://doi.org/10.1016/j.ympev.2015.09.005

    Günther, A. (1859) On the reptiles from St. Croix, West Indies, collected by Messrs, A. and B. Newton. Annals and Magazine of Natural History, Series 3, 4 (21), 209–217.

    https://doi.org/10.1080/00222935908697110

    Hass, C.A. (1991) Evolution and biogeography of West Indian Sphaerodactylus (Sauria: Gekkonidae): A molecular approach. Journal of the Zoological Society of London, 225, 525–561.

    https://doi.org/10.1111/j.1469-7998.1991.tb04323.x

    Hass, C.A. (1996) Relationships among West Indian geckos of the genus Sphaerodactylus: a preliminary analysis of mitochondrial 16S ribosomal RNA sequences. In: Powell, R. & Henderson, R.W. (Eds.), Contributions to West Indian Herpetology: A tribute to Albert Schwartz. Society for the Study of Amphibians and Reptiles, Ithaca, New York, pp. 175–194.

    Heatwole, H. & Levins, R. (1972) Biogeography of the Puerto Rican Bank: flotsam transport of terrestrial animals. Ecology, 53, 112–117.

    https://doi.org/10.2307/1935715

    Heatwole, H. & MacKenzie, F. (1967) Herpetogeography of Puerto Rico. IV. Paleogeography, faunal similarity and endemism. Evolution, 21, 429–438.

    https://doi.org/10.1111/j.1558-5646.1967.tb03400.x

    Hearty, P.J., Hollin, J.T., Neumann, A.C., O’Leary, M.J. & McCulloch, M. (2007) Global sea-level fluctuations during the Last Interglaciation (MIS5e). Quaternary Science Reviews, 26, 2090–2112.

    https://doi.org/10.1016/j.quascirev.2007.06.019

    Hedges, S.B. (1989) An island radiation: Allozyme evolution in Jamaican frogs of the genus Eleutherodactylus (Leptodactylidae). Caribbean Journal of Science, 25, 123–147.

    Hedges. S.B. & Conn, C.E. (2012) A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae). Zootaxa, 3288 (1), 1–244.

    https://doi.org/10.11646/zootaxa.3288.1.1

    Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct–developing frogs (Anura : Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737 (1), 1–182.

    https://doi.org/10.11646/zootaxa.1737.1.1

    Hedges, S.B., Marion, A.B., Lipp, K.M., Marin, J. & Vidal, N. (2014) A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49, 1–61.

    https://doi.org/10.31611/ch.49

    Hedges, S.B. & Thomas, R. (2001) At the lower size limit in amniote vertebrates: A new diminutive lizard from the West Indies. Caribbean Journal of Science, 37, 168–173.

    Heinicke, M.P., Duellman, W.E. & Hedges, S.B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences of the United States of America, 104, 10092–10097.

    https://doi.org/10.1073/pnas.0611051104

    Hewitt, G.M. (2004) Genetic consequences of climatic oscillations in the Quaternary. Philosophical Transactions of the Royal Society B: Biological Sciences, 359, 183–195.

    https://doi.org/10.1098/rstb.2003.1388

    Huelsenbeck, J.P., Nielsen, R. & Bollback, J.P. (2003) Stochastic mapping of morphological characters. Systematic Biology, 52, 131–158.

    https://doi.org/10.1080/10635150390192780

    Iturralde-Vinent, M.A. (2006) Meso-Cenozoic Caribbean paleogeography: implications for the historical biogeography of the region. International Geology Review, 48, 791–827.

    https://doi.org/10.2747/0020-6814.48.9.791

    Iturralde-Vinent, M.A. & MacPhee, R.D.E. (1999) Paleogeography of the Caribbean region: Implications for Cenozoic biogeography. Bulletin of the American Museum of Natural History, 238, 1–95.

    Jones, G. (2017) Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. Journal of Mathematical Biology, 74, 447–467.

    https://doi.org/10.1007/s00285-016-1034-0

    Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.

    https://doi.org/10.1093/bioinformatics/bts199

    King, F.W. (1962) Systematics of Lesser Antillean lizards of the genus Sphaerodactylus. Bulletin of the Florida State Museum, Biological Sciences, 7, 1–52.

    Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874.

    https://doi.org/10.1093/molbev/msw054

    Librado, P. & Rozas, J. (2009) DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.

    https://doi.org/10.1093/bioinformatics/btp187

    Lin, L.-H., Xiang, J., Diong, C.-H., Du, Y. & Lin, C.-X. (2010) Phylogeography and population structure of the Reevese’s butterfly lizard (Leiolepis reevesii) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 56, 601–607.

    https://doi.org/10.1016/j.ympev.2010.04.032

    López-Ortiz, R. & Lewis, A.R. (2002) Seasonal abundance of hatchlings and gravid females of Sphaerodactylus nicholsi in Cabo Rojo, Puerto Rico. Journal of Herpetology, 36, 276–280.

    https://doi.org/10.1670/0022-1511(2002)036[0276:SAOHAG]2.0.CO;2

    Losos, J.B., Jackman, T.R., Larson, A., de Queiroz, K. & Rodriguez-Schettino, L. (1998) Contingency and determinism in replicated adaptive radiations of island lizards. Science, 279, 2115–2118.

    https://doi.org/10.1126/science.279.5359.2115

    Losos, J.B. (2009) Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles. University of California Press, Berkeley, California, 528 pp.

    https://doi.org/10.1525/california/9780520255913.001.0001

    Lovette, I.J. (2005) Glacial cycles and the tempo of avian speciation. Trends in Ecology and Evolution, 20, 57–59.

    https://doi.org/10.1016/j.tree.2004.11.011

    MacArthur, R.H. & Wilson, E.O. (1967) Theory of Island Biogeography. Princeton University Press, Princeton, New Jersey, 224 pp.

    MacLean, W. P. (1985). Water-loss rates of Sphaerodactylus parthenopion (Reptilia: Gekkonidae), the smallest amniote vertebrate. Comparative Biochemistry and Physiology A Physiology, 82, 759–761.

    https://doi.org/10.1016/0300-9629(85)90479-7

    MacLean, W.P. & Holt, R.D. (1979) Distributional patterns in St. Croix Sphaerodactylus lizards: The taxon cycle in action. Biotropica, 11, 189–195.

    https://doi.org/10.2307/2388038

    Mahler, D.L., Revell, L.J., Glor, R.E. & Losos, J.B. (2010) Ecological opportunity and the rate of morphological evolution in the diversification of Greater Antillean anoles. Evolution, 64, 2731–2745.

    https://doi.org/10.1111/j.1558-5646.2010.01026.x

    Mann, P., Draper, G. & Lewis, J.F. (1991) An overview of the geologic and tectonic development of Hispaniola. In: Mann, P., Draper, G. & Lewis, J.F. (Eds.), Geological and Tectonic Development of the North America-Caribbean Plate Boundary in Hispaniola. Geological Society of America Special Papers, 262, pp. 1–28.

    https://doi.org/10.1130/SPE262-p1

    Mayr, E. (1942) Systematics and the Origin of Species. Columbia University Press, New York, pp. 334.

    Meerwarth, H. (1901) Die Westindischen Reptilien und Batrachier des Naturhistorischen Museums in Hamburg. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 18, 1–41.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, 14 November 2010, pp. 1–8.

    https://doi.org/10.1109/GCE.2010.5676129

    Nava, S.S. (2006) Size does matter. Iguana, 13, 16–30.

    Nava, S.S., Lindsay, C.R., Powell, R. & Henderson, R.W. (2002) Sphaerodactylus parvus. Catalog of American Amphibians and Reptiles, 752, 1–2.

    Nei, M. & Li, W.H. (1979) Mathematical model for studying genetic variation in terms of restriction endonucleases. Proceedings of the National Academy of Sciences USA, 76, 5269–5273.

    https://doi.org/10.1073/pnas.76.10.5269

    Nielsen, R. (2002) Mapping mutations on phylogenies. Systematic Biology, 51, 729–739.

    https://doi.org/10.1080/10635150290102393

    Ogilvie, H.A., Bouckaert, R.R. & Drummond, A.J. (2017) StarBEAST2 brings faster species tree inference and accurate estimates of substitution rates. Molecular Biology and Evolution, 34, 2101–2114.

    https://doi.org/10.1093/molbev/msx126

    Oliver, P.M., Hutchinson, M.N. & Cooper, S.J.B. (2007) Phylogenetic relationships in the lizard genus Diplodactylus Gray and resurrection of Lucasium Wermuth (Gekkota, Diplodactylidae). Australian Journal of Zoology, 55, 197–210.

    https://doi.org/10.1071/ZO07008

    Padilla, L.F. (2006) Geographic variation in color pattern on Sphaerodactylus macrolepis Günther 1859, (Sauria: Gekkonidae). M. Sc. Thesis, University of Puerto Rico, San Juan. 153 pp.

    Page, R.D. & Lydeard, C. (1994) Towards a cladistic biogeography of the Caribbean. Cladistics, 10, 21–41.

    https://doi.org/10.1111/j.1096-0031.1994.tb00162.x

    Palumbi, S.R., Cipriano, F. & Hare, M.P. (2001) Predicting nuclear gene coalescence from mitochondrial data: The three-times rule. Evolution, 55, 859–868.

    https://doi.org/10.1554/0014-3820(2001)055[0859:PNGCFM]2.0.CO;2

    Papadopoulou, A. & Knowles, L.L. (2015) Genomic tests of the species-pump hypothesis: Recent island connectivity cycles drive population divergence but not speciation in Caribbean crickets across the Virgin Islands. Evolution, 69, 1501–1517.

    https://doi.org/10.1111/evo.12667

    Paradis, E. & Schilep, K. (2018) ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics, 35, 526–528.

    https://doi.org/10.1093/bioinformatics/bty633

    Patten, M.A. & Unitt, P. (2002) Diagnosability versus mean differences of sage sparrow subspecies. The Auk, 119, 26–35.

    https://doi.org/10.1093/auk/119.1.26

    Pepper, M., Doughty, P. & Keogh, J.S. (2006) Molecular phylogeny and phylogeography of the Australian Diplodactylus stenodactylus (Gekkota; Reptilia) species-group based on mitochondrial and nuclear genes reveals an ancient split between Pilbara and non-Pilbara D. stenodactylus. Molecular Phylogenetics and Evolution, 41, 539–555.

    https://doi.org/10.1016/j.ympev.2006.05.028

    Pinto, B.J., Titus-McQuillan, J., Daza, J.D. & Gamble, T. (2019a). Persistence of a geographically-stable hybrid zone in Puerto Rican dwarf geckos. Journal of Heredity, 110, 523–534.

    https://doi.org/10.1093/jhered/esz015

    Pinto, B.J., Colli, G.R., Higham, T.E., Russell, A.P., Scantlebury, D.P., Vitt, L.J. & Gamble, T. (2019b) Population genetic structure and species delimitation of a widespread, Neotropical dwarf gecko. Molecular Phylogenetics and Evolution, 133, 54–66.

    https://doi.org/10.1016/j.ympev.2018.12.029

    Portik, D.M., Travers, S.L., Bauer, A.M. & Branch, W.R. (2013) A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ‘sky island’ of northern Mozambique. Zootaxa, 3710 (5), 415–435.

    https://doi.org/10.11646/zootaxa.3710.5.2

    Powell, R. & Henderson, R.W. (2001) On the taxonomic status of some Lesser Antillean Lizards. Caribbean Journal of Science, 37, 288–290.

    Questel, K. (2018) Discovery of geckos, Sphaerodactylus macrolepis Günther and Sphaerodactylus sputator Sparrman, on Prickly Pear Cays, Anguilla, British West Indies. Caribbean Herpetology, 61, 1–2.

    https://doi.org/10.31611/ch.61

    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology, 67, 901–904.

    https://doi.org/10.1093/sysbio/syy032

    Regalado, R. (2014) Does dichromatism variation affect sex recognition in dwarf geckos? Ethology, Ecology and Evolution, 27, 56–73.

    https://doi.org/10.1080/03949370.2014.885465

    Revell, L.J. (2012) phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3, 217–223.

    https://doi.org/10.1111/j.2041-210X.2011.00169.x

    Reynolds, R.G., Strickland, T.R., Kolbe, J.J., Falk, B.G., Perry, G., Revell, L.J. & Losos, J.B. (2017) Archipelagic genetics in a widespread Caribbean anole. Journal of Biogeography, 44, 2631–2647.

    https://doi.org/10.1111/jbi.13072

    Rivero, J.A. (1998) Los Anfibios y Reptiles de Puerto Rico (The Amphibians and Reptiles of Puerto Rico). Segunda Edición Revisada. Universidad de Puerto Rico, Editorial Universitaria, San Juan, Puerto Rico, 510 pp.

    Rocha, S., Vences, M., Glaw, F., Posada, D. & Harris, D.J. (2009) Multigene phylogeny of Malagasy day geckos of the genus Phelsuma. Molecular Phylogenetics and Evolution, 52, 530–537.

    https://doi.org/10.1016/j.ympev.2009.03.032

    Rodda, G.H., Perry, G.A.D., Rondeau, R.J. & Lazell, J. (2001). The densest terrestrial vertebrate. Journal of Tropical Ecology, 17, 331–338.

    https://doi.org/10.1017/S0266467401001225

    Rull, V. (2006) Quaternary speciation in the Neotropics. Molecular Ecology, 15, 4257–4259.

    https://doi.org/10.1111/j.1365-294X.2006.03052.x

    Scantlebury, D.P., Ng, J., Landestoy, M., Geneva, A. & Glor, R.E. (2011) Notes on activity patterns of five species of Sphaerodactylus (Squamata: Sphaerodactylidae) from the Dominican Republic. IRCF Reptiles and Amphibians, 18, 51–55.

    Scantlebury, D.P. (2014) Patterns of adaptive radiation in insular reptiles and amphibians. PhD dissertation, New York, 220 pp.

    Schmidt, K.P. (1920) Contributions to the herpetology of Porto Rico. Annals of the New York Academy of Sciences, 28, 167–200.

    https://doi.org/10.1111/j.1749-6632.1918.tb55351.x

    Schmidt, K.P. (1928) Amphibians and land reptiles of Porto Rico, with a list of those reported from the Virgin Islands. New York Academy of Sciences Scientific Survey of Porto Rico and the Virgin Islands, 10, 1–160.

    Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9, 671–675.

    https://doi.org/10.1038/nmeth.2089

    Singhal, S., Hoskin, C.J., Couper, P., Potter, S. & Moritz, C. (2018) A framework for resolving cryptic species: A case study from the lizards of the Australian wet tropics. Systematic Biology, 67, 1061–1075.

    https://doi.org/10.1093/sysbio/syy026

    Stamatakis, A. (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.

    https://doi.org/10.1093/bioinformatics/btu033

    Steinberg, D.S., Powell, S.D., Powell, R., Parmerlee, J.S. & Henderson, R.W. (2007) Population densities, water-loss rates, and diets of Sphaerodactylus vincenti on St. Vincent, West Indies. Journal of Herpetology, 41, 330–336.

    https://doi.org/10.1670/0022-1511(2007)41[330:PDWRAD]2.0.CO;2

    Stejneger, L. (1904) Herpetology of Porto Rico. Report of the United States National Museum, 1902, 549–724.

    https://doi.org/10.5962/bhl.title.11835

    Stephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68, 978–989.

    https://doi.org/10.1086/319501

    Sukumaran, J. & Knowles, L.L. (2017) Multispecies coalescent delimits structure, not species. Proceedings of the National Academy of Sciences, 114, 1607–1612.

    https://doi.org/10.1073/pnas.1607921114

    Thomas, R. (1965) A new gecko from the Virgin Islands. Quarterly Journal of the Florida Academy of Sciences, 28, 117–122.

    Thomas, R. & Schwartz, A. (1966) Sphaerodactylus (Gekkonidae) in the greater Puerto Rico region. Bulletin of the Florida State Museum, Biological Sciences, 10, 193–260.

    Thomas, R., Hedges, S.B. & Garrido, O.H. (1992) Two new species of Sphaerodactylus from Eastern Cuba (Squamata: Gekkonidae). Herpetologica, 48, 358–367.

    https://doi.org/10.2307/1564883

    Thorpe, R.S., Jones, A.G., Malhotra, A. & Surget-Groba, Y. (2008) Adaptive radiation in Lesser Antillean lizards: molecular phylogenetics and species recognition in the Lesser Antillean dwarf gecko complex, Sphaerodactylus fantasticus. Molecular Ecology, 17, 1489–1504.

    https://doi.org/10.1111/j.1365-294X.2007.03686.x

    Torstrom, S.M., Pangle, K.L. & Swanson, B.J. (2014) Shedding subspecies: The influence of genetics on reptile subspecies taxonomy. Molecular Phylogenetics and Evolution, 76, 134–143.

    https://doi.org/10.1016/j.ympev.2014.03.011.

    Werneck, F.P., Gamble, T., Colli, G.R., Rodrigues, M.T. & Sites, J.W. (2012) Deep diversification and long-term persistence in the South American ‘dry diagonal’: Integrating continent-wide phylogeography and distribution modeling of geckos. Evolution, 66, 3014–3034.

    https://doi.org/10.1111/j.1558-5646.2012.01682.x

    Williams, E.E. (1983) Ecomorphs, faunas, island size, and diverse end points in island radiations of Anolis. In: Huey, R.B., Pianka, E.R. & Schoener, T.W. (Eds.), Lizard Ecology: Studies of a Model Organism. Harvard University Press, Cambridge, Massachusetts, pp. 326–370.

    https://doi.org/10.4159/harvard.9780674183384.c18

    Wilson, E.O. & Brown Jr., W.L. (1953) The subspecies concept and its taxonomic application. Systematic Zoology, 2, 97–111.

    https://doi.org/10.2307/2411818

    Ziegler, T., Truong, N.Q., Schmitz, A., Stenke, R. & Rösler, H. (2008) A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae). Zootaxa, 1771 (1), 16–30.

    https://doi.org/10.11646/zootaxa.1771.1.2

    Zink, R.M. (2004) The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society B: Biological Sciences, 271, 561–564.

    https://doi.org/10.1098/rspb.2003.2617

    Zink, R.M. & Barrowclough, G.F. (2008) Mitochondrial DNA under siege in avian phylogeography. Molecular Ecology, 17, 2107–2121.

    https://doi.org/10.1111/j.1365-294X.2008.03737.x