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Type: Article
Published: 2025-11-19
Page range: 151-183
Abstract views: 64
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Taxonomic revision of the polyphyletic Phyllodactylus bordai (Squamata: Phyllodactylidae), with the description of a new species

Facultad de Ciencias Forestales; Universidad Autónoma de Nuevo León; Carretera Nacional 85; Km. 145; 67700 Linares; Nuevo León; México.; Instituto de Biología; Departamento de Zoología; Colección Nacional de Insectos; Universidad Nacional Autónoma de México; Circuito Zona Deportiva S/N; C.U.; Coyoacán; 04510 Ciudad de México; México.
Universidad Autónoma de Yucatán; Campus de Ciencias Biológicas y Agropecuarias; Carretera Mérida-Xmatkuil Km. 15.5 Apdo.; Plan de Ayala ÌII; Itzimná; 97100 Mérida; Yucatán; México.
Totlok A.C.; Sinanche 174; C.P. 14240; Tlalpan; CDMX; Mexico.; Museo de Zoología “Alfonso L. Herrera” Facultad de Ciencias; Universidad Nacional Autónoma de México; Circuito exterior de CU; Ciudad Universitaria; 04510 Ciudad de México; México.
Museo de Zoología “Alfonso L. Herrera” Facultad de Ciencias; Universidad Nacional Autónoma de México; Circuito exterior de CU; Ciudad Universitaria; 04510 Ciudad de México; México.
Reptilia species delimitation gene flow introgression genomics reference genome Tehuacan-Cuicatlan Valley

Abstract

Through the re-evaluation of genomic data obtained from reduced genome representation sequencing method of the species belonging to Clade I within North American Phyllodactylus, we evidence the polyphyletic condition of the P. bordai taxon, which contains three independently evolving lineages. Gene flow tests demonstrated that the three lineages within P. bordai are genetically isolated and had no contact after their divergence. Furthermore, the divergence ages estimated for two of the lineages, from northern Guerrero and the Tehuacan-Cuicatlan Valley, are relatively deep, dating back to the mid-Oligocene and the mid-Eocene, respectively. Interestingly, we only obtained a gene flow signal between the most recently diverged lineage (from Huautla, Morelos) and its sister species (P. papenfussi), probably due to sporadic introgression pulses after their divergence in the early Pliocene. The evidence generated from the analysis of genomic, morphological, and climatic data allows us to propose taxonomic changes in accordance with the evolutionary history of the three main lineages, while avoiding the propagation of an artificial taxonomy within Clade I with respect to P. bordai. As such, in this study we describe Phyllodactylus ngiwa sp. nov., the earliest diverging lineage within the polyphyletic P. bordai taxon, from populations in the Tehuacan-Cuicatlan Valley. We also restrict P. bordai to populations from northern Guerrero. While the results suggest that the Morelos lineage is a distinct species from the rest of the North American Phyllodactylus species, more studies with increased sampling are necessary to assess this divergence in detail, including any interactions with its sister taxon, P. papenfussi.

 

References

  1. Blair, C., Méndez-de la Cruz, F.R., Ngo, A., Lindell, J., Lathrop, A. & Murphy, R.W. (2009) Molecular phylogenetics and taxonomy of leaf-toed geckos (Phyllodactylidae: Phyllodactylus) inhabiting the peninsula of Baja California. Zootaxa, 2027 (1), 28–42. https://doi.org/10.11646/zootaxa.2027.1.2
  2. Blair, C., Méndez-de la Cruz, F.R., Law, C. & Murphy, R. (2015) Molecular phylogenetics and species delimitation of leaf-toed geckos (Phyllodactylidae: Phyllodactylus) throughout the Mexican tropical dry forest. Molecular Phylogenetics and Evolution, 84, 254–265. https://doi.org/10.1016/j.ympev.2015.01.003
  3. Burriel-Carranza, B., Mochales-Riaño, G., Talavera, A., Els, J., Estarellas, M., Al-Saadi, S., Urriago-Suarez, J.D., Olsson, P.O., Matschiner, M. & Carranza, S. (2024) Clinging on the brink: Whole genomes reveal human-induced population declines and severe inbreeding in the Critically Endangered Emirati Leaf-toed Gecko (Asaccus caudivolvulus). Molecular Ecology, e17451. https://doi.org/10.1111/mec.17451
  4. Butler, B.O., Smith, L.L. & Flores-Villela, O. (2023) Phylogeography and taxonomy of Coleonyx elegans Gray 1845 (Squamata: Eublepharidae) in Mesoamerica: The Isthmus of Tehuantepec as an environmental barrier. Molecular Phylogenetics and Evolution, 178, 107632. https://doi.org/10.1016/j.ympev.2022.107632
  5. Canseco-Márquez, L. & Gutiérrrez-Mayén, M.G. (2010) Anfibios y reptiles del Valle de Tehuacán-Cuicatlán. Comisión Nacional para el conocimiento y uso de la biodiversidad, México, D.F., 302 pp.
  6. Card, D.C., Jennings, W.B. & Edwards, S.V. (2023) Genome evolution and the future of phylogenomics of non-avian reptiles. Animals, 13 (3), 1–33. https://doi.org/10.3390/ani13030471
  7. Casas-Andreu, G., Méndez-de la Cruz, F.R. & Camarillo, J.L. (1996) Anfibios y reptiles de Oaxaca. Lista, distribución y conservación. Acta Zoológica Mexicana, New Series, 69, 1–35. https://doi.org/10.21829/azm.1996.69691928
  8. Castro-Franco, R. & Uribe-Peña, Z. (1992) Dos subespecies nuevas de Phyllodactylus lanei (Sauria: Gekkonidae). Anales del Instituto de Biología UNAM Serie Zoología, 63 (1), 113–123.
  9. Chan, K.O. & Grismer, L.L. (2021) A standardized and statistically defensible framework for quantitative morphological analyses in taxonomic studies. Zootaxa, 5023 (2), 293–300. https://doi.org/10.11646/zootaxa.5023.2.9
  10. Chifman, J. & Kubatko, L. (2014) Quartet inference from SNP data under the coalescent model. Bioinformatics, 30 (23), 3317–3324. https://doi.org/10.1093/bioinformatics/btu530
  11. Danecek, P., Auton, A., Abecasis, G., Albers, C.A., Banks, E., DePristo, M.A., Handsaker, R.E., Lunter, G., Marth, G.T., Sherry, S.T., McVean, G., Durbin, R. & 1000 Genomes Project Analysis Group. (2011) The variant call format and VCFtools. Bioinformatics, 27 (15), 2156–2158. https://doi.org/10.1093/bioinformatics/btr330
  12. Dávalos-Álvarez, O.G., Nieto-Samaniego, Á.F., Alaniz-Álvarez, S.A., Martínez-Hernández, E. & Ramírez-Arriaga, E. (2007) Estratigrafía cenozoica de la región de Tehuacán y su relación con el sector norte de la falla de Oaxaca. Revista mexicana de ciencias geológicas, 24 (2), 197–215.
  13. Davis, H.R., Chan, K.O., Das, I., Brennan, I.G., Karin, B.R., Jackman, T.R., Brown, R.M., Iskandar, D.T., Nashriq, I., Grismer, L.L. & Bauer, A.M. (2020) Multilocus phylogeny of Bornean Bent-Toed geckos (Gekkonidae: Cyrtodactylus) reveals hidden diversity, taxonomic disarray, and novel biogeographic patterns. Molecular Phylogenetics and Evolution, 147, 106785. https://doi.org/10.1016/j.ympev.2020.106785
  14. Davis, H.R., Das, I., Leaché, A.D., Karin, B.R., Brennan, I.G., Jackman, T.R., Nashriq, I., Chan, K.O. & Bauer, A.M. (2021) Genetically diverse yet morphologically conserved: Hidden diversity revealed among Bornean geckos (Gekkonidae: Cyrtodactylus). Journal of Zoological Systematics and Evolutionary Research, 59 (5), 1113–1135. https://doi.org/10.1111/jzs.12470
  15. Davis, H.R., Nashriq, I., Woytek, K.S., Wikramanayake, S.A., Bauer, A.M., Karin, B.R., Brennan, I.G., Iskandar, D.T. & Das, I. (2023) Genomic analysis of Bornean geckos (Gekkonidae: Cyrtodactylus) reveals need for updated taxonomy. Zoologica Scripta, 52 (3), 249–263. https://doi.org/10.1111/zsc.12575
  16. Dixon, J.R. (1964) The systematics and distribution of lizards of the genus Phyllodactylus in North and Central America. New Mexico State University Research Center, Scientific Bulletin, 64 (1), 1–139.
  17. Dixon, J.R. (1966) Speciation and systematics of the gekkonid lizard genus Phyllodactylus of the islands of the Gulf of California. Proceedings of the California Academy of Science, 33, 415–452.
  18. Dixon, J.R. & Huey, R.B. (1970) Systematics of the lizards of the gekkonid genus Phyllodactylus of mainland South America. Natural History Museum of Los Angeles County Contribution in Science, 192, 1–78. https://doi.org/10.5962/p.241179
  19. dos Reis, M. & Yang, Z. (2019) Bayesian molecular clock dating using genome-scale datasets. In: Anisimova, M. (Ed.), Evolutionary genomics: Statistical and computational methods. Springer, New York, New York, pp. 309–330. https://doi.org/10.1007/978-1-4939-9074-0_10
  20. dos Reis, M. (2022) Dating microbial evolution with MCMCtree. In: Luo, H. (Ed.), Environmental microbial evolution methods and protocols. Humana, New York, New Yok, pp. 3–22. https://doi.org/10.1007/978-1-0716-2691-7_1
  21. Dubeux, M.J., Goncalves, U., Palmeira, C.N., Nunes, P.M., Cassimiro, J., Gamble, T., Werneck, F.P., Rodrigues, M.T. & Mott, T. (2022) Two new species of geckos of the genus Phyllopezus Peters, 1878 (Squamata: Gekkota: Phyllodactylidae) from northeastern Brazil. Zootaxa, 5120 (3), 345–372. https://doi.org/10.11646/zootaxa.5120.3.3
  22. Eaton, D.A.R. & Overcast, I. (2020) ipyrad: Interactive assembly and analysis of RADseq datasets. Bioinformatics, 36 (8), 2592–2594. https://doi.org/10.1093/bioinformatics/btz966
  23. Fitz-Gibbon, S., Hipp, A.L., Pham, K.K., Manos, P.S. & Sork, V.L. (2017) Phylogenomic inferences from reference-mapped and de novo assembled short-read sequence data using RADseq sequencing of California white oaks (Quercus section Quercus). Genome, 60 (9), 743–755. https://doi.org/10.1139/gen-2016-0202
  24. Flores-Tolentino, M., Ramírez-Rodríguez, J.R., Morales-Linares, J., Ibarra-Manríquez, G., Dorado, Ó. & Villaseñor, J.L. (2023) Delimitación geográfica y florística de la provincia fisiográfica de la Depresión del Balsas, México, con énfasis en el bosque tropical estacionalmente seco. Revista mexicana de biodiversidad, 94, e944985. https://doi.org/10.22201/ib.20078706e.2023.94.4985
  25. Flouri, T., Jiao, X., Rannala, B. & Yang, Z. (2018) Species Tree Inference with BPP using genomic sequences and the Multispecies Coalescent. Molecular Biology and Evolution, 35 (10), 2585–2593. https://doi.org/10.1093/molbev/msy147
  26. Flouri, T., Jiao, X., Rannala, B. & Yang, Z. (2020) A Bayesian implementation of the Multispecies Coalescent Model with introgression for phylogenomic analysis. Molecular Biology and Evolution, 37 (4), 1211–1223. https://doi.org/10.1093/molbev/msz296
  27. Flouri, T., Jiao, X., Huang, J., Rannala, B. & Yang, Z. (2023) Efficient Bayesian inference under the multispecies coalescent with migration. Proceedings of the National Academy of Sciences, 120 (44), 1–11. https://doi.org/10.1073/pnas.2310708120
  28. Formenti, G., Theissinger, K., Fernandes, C., Bista, I., Bombarely, A., Bleidorn, C., Ciofi, C., Crottini, A., Godoy, J.A., Höglund, J., Malukiewicz, J., Mouton, A., Oomen, R.A., Paez, S., Palsbøll, P.J., Pampoulie, C., Ruiz-López, M.J., Svardal, H., Theofanopoulou, C., de Vries, J., Waldvogel, A.-M., Zhang, G., Mazzoni, C.J., Jarvis, E.D., Bálint, M., Formenti, G., Theissinger, K., Fernandes, C., Bista, I., Bombarely, A., Bleidorn, C., ˇCiampor, F., Ciofi, C., Crottini, A., Godoy, J.A., Hoglund, J., Malukiewicz, J., Mouton, A., Oomen, R.A., Paez, S., Palsbøll, P., Pampoulie, C., Ruiz-López, M.J., Svardal, H., Theofanopoulou, C., de Vries, J., Waldvogel, A.-M., Zhang, G., Mazzoni, C.J., Jarvis, E., Bálint, M., Aghayan, S.A., Alioto, T.S., Almudi, I., Alvarez, N., Alves, P.C., Amorim, I.R., Antunes, A., Arribas, P., Baldrian, P., Berg, P.R., Bertorelle, G., Böhne, A., Bonisoli-Alquati, A., Bostjancic, L.L., Boussau, B., Breton, C.M., Buzan, E., Campos, P.F., Carreras, C., Castro, L.F., Chueca, L.J., Conti, E., Cook-Deegan, R., Croll, D., Cunha, M.V., Delsuc, F., Dennis, A.B., Dimitrov, D., Faria, R., Favre, A., Fedrigo, O.D., Fernández, R., Ficetola, G.F., Flot, J.-F., Gabaldón, T., Galea Agius, D.R., Gallo, G.R., Giani, A.M., Gilbert, M.T.P., Grebenc, T., Guschanski, K., Guyot, R., Hausdorf, B., Hawlitschek, O., Heintzman, P.D., Heinze, B., Hiller, M., Husemann, M., Iannucci, A., Irisarri, I., Jakobsen, K.S., Jentoft, S., Klinga, P., Kloch, A., Kratochwil, C.F., Kusche, H., Layton, K.K.S., Leonard, J.A., Lerat, E., Liti, G., Manousaki, T., Marques-Bonet, T., Matos-Maraví, P., Matschiner, M., Maumus, F., Mc Cartney, A.M., Meiri, S., Melo-Ferreira, J., Mengual, X., Monaghan, M.T., Montagna, M., Mysłajek, R.W., Neiber, M.T., Nicolas, V., Novo, M., Ozretic, P., Palero, F., Parvulescu, L., Pascual, M., Paulo, O.S., Pavlek, M., Pegueroles, C., Pellissier, L., Pesole, G., Primmer, C.R., Riesgo, A., Rüber, L., Rubolini, D., Salvi, D., Seehausen, O., Seidel, M., Secomandi, S., Studer, B., Theodoridis, S., Thines, M., Urban, L., Vasemägi, A., Vella, A., Vella, N., Vernes, S.C., Vernesi, C., Vieites, D.R., Waterhouse, R.M., Wheat, C.W., Wörheide, G., Wurm, Y. & Zammit, G. (2022) The era of reference genomes in conservation genomics. Trends in Ecology and Evolution, 37 (3), 197–202. https://doi.org/10.1016/j.tree.2021.11.008
  29. Gable, S.M., Mendez, J.M., Bushroe, N.A., Wilson, A., Byars, M.I. & Tollis, M. (2023) The state of Squamate genomics: past, present, and future of genome research in the most speciose terrestrial vertebrate order. Genes, 14, 1387. https://doi.org/10.3390/genes14071387
  30. 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 (3), 943–953. https://doi.org/10.1016/j.ympev.2011.11.033
  31. GBIF.org. (2024) GBIF Occurrence Download. Available from: https://doi.org/10.15468/dl.7jb2ha (accessed 10 November 2024)
  32. Ghildiyal, K., Nayak, S.S., Rajawat, D., Sharma, A., Chhotaray, S., Bhushan, B., Dutt, T. & Panigrahi, M. (2023) Genomic insights into the conservation of wild and domestic animal diversity: A review. Gene, 886, 147719. https://doi.org/10.1016/j.gene.2023.147719
  33. Gippner, S., Travers, S.L., Scherz, M.D., Colston, T.J., Lyra, M.L., Mohan, A.V., Multzsch, M., Nielsen, S.V., Rancilhac, L., Glaw, F., Bauer, A.M. & Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https://doi.org/10.1016/j.ympev.2021.107311
  34. Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59 (3), 307–321. https://doi.org/10.1093/sysbio/syq010
  35. Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
  36. Hogg, C.J. (2024) Translating genomic advances into biodiversity conservation. Nature Reviews Genetics, 25 (5), 362–373. https://doi.org/10.1038/s41576-023-00671-0
  37. INALI. (2020) Se revitalizará la lengua Ngiwa / Ngigua desde las propias comunidades, Instituto de Lenguas indígenas, Alcaldía Alvaro Obregón, Ciudad de México. Available from: https://www.inali.gob.mx/detalle/ (accessed 15 January 2025)
  38. Jombart, T. & Ahmed, I. (2011) adegenet 1.3-1: new tools for the analysis of genome-wide SNP data. Bioinformatics, 27 (21), 3070–3071. https://doi.org/10.1093/bioinformatics/btr521
  39. Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589. https://doi.org/10.1038/nmeth.4285
  40. Kamvar, Z.N., Brooks, J.C. & Grünwald, N.J. (2015) Novel R tools for analysis of genome-wide population genetic data with emphasis on clonality. Frontiers in Genetics, 6, 1–10. https://doi.org/10.3389/fgene.2015.00208
  41. Karger, D.N., Conrad, O., Böhner, J., Kawohl, T., Kreft, H., Soria-Auza, R.W., Zimmermann, N.E., Linder, P. & Kessler, M. (2017) Climatologies at high resolution for the Earth land surface areas. Scientific Data, 4. https://doi.org/10.1038/sdata.2017.122
  42. Kassambara, A. & Mundt, F. (2020) Factoextra: Extract and Visualize the Results of Multivariate Data Analyses. R Package Version 1.0.7. Available from: https://CRAN.R-project.org/package=factoextra (accessed 2 November 2024)
  43. Knaus, B.J. & Grünwald, N.J. (2017) VCFR: a package to manipulate and visualize variant call format data in R. Molecular Ecology Resources, 17 (1), 44–53. https://doi.org/10.1111/1755-0998.12549
  44. Kotsakiozi, P., Jablonski, D., Ilgaz, Ç., Kumlutaþ, Y., Avcý, A., Meiri, S., Itescu, Y., Kukushkin, O., Gvoždík, V., Scillitani, G., Roussos, S.A., Jandzik, D., Kasapidis, P., Lymberakis, P. & Poulakakis, N. (2018) Multilocus phylogeny and coalescent species delimitation in Kotschy’s gecko, Mediodactylus kotschyi: Hidden diversity and cryptic species. Molecular Phylogenetics and Evolution, 125, 177–187. https://doi.org/10.1016/j.ympev.2018.03.022
  45. Lara-Reséndiz, R.A., Arenas-Moreno, D.M. & Méndez-de la Cruz, F.R. (2013) Termorregulación diurna y nocturna de la lagartija Phyllodactylus bordai (Gekkota: Phyllodactylidae) en una región semiárida del centro de México. Revista chilena de historia natural, 86 (2), 127–135. https://doi.org/10.4067/S0716-078X2013000200002
  46. Lemos-Espinal, J.A. & Smith, G.R. (2020) A conservation checklist of the herpetofauna of Morelos, with comparisons with adjoining states. ZooKeys, 941, 121–144. https://doi.org/10.3897/zookeys.941.52011
  47. Lemos-Espinal, J.A. & Smith, G.R. (2024) The distribution, diversity and conservation of the Mexican herpetofauna among its biogeographic provinces. Journal for Nature Conservation, 82, 126714. https://doi.org/10.1016/j.jnc.2024.126714
  48. Lozano-Fernández, J. (2022) A practical guide to design and assess a phylogenomic study. Genome Biology and Evolution, 14 (9), evac129. https://doi.org/10.1093/gbe/evac129
  49. Mata-Silva, V., Johnson, J.D., Wilson, L.D. & García-Padilla, E. (2015) The herpetofauna of Oaxaca, Mexico: composition, physiographic distribution, and conservation status. Mesoamerican Herpetology, 2, 6–62.
  50. Mata-Silva, V., García-Padilla, E., Rocha, A., Desantis, D.L., Johnson, J.D., Ramirez-Bautista, A. & Wilson, L.D. (2021) A reexamination of the herpetofauna of Oaxaca, Mexico: composition update, physiographic distribution, and conservation commentary. Zootaxa, 4996 (2), 201–252. https://doi.org/10.11646/zootaxa.4996.2.1
  51. Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37, 1530–1534. https://doi.org/10.1093/molbev/msaa015
  52. Moritz, C.C., Pratt, R.C., Bank, S., Bourke, G., Bragg, J.G., Doughty, P., Keogh, J.S., Laver, R.J., Potter, S., Teasdale, L.C., Tedeschi, L.G. & Oliver, P.M. (2018) Cryptic lineage diversity, body size divergence, and sympatry in a species complex of Australian lizards (Gehyra). Evolution, 72 (1), 54–66. https://doi.org/10.1111/evo.13380
  53. Morrone, J.J., Escalante, T. & Rodríguez-Tapia, G. (2017) Mexican biogeographic provinces: map and shapefiles. Zootaxa, 4277 (2), 277–279. https://doi.org/10.11646/zootaxa.4277.2.8
  54. Morrone, J.J. (2019) Regionalización biogeográfica y evolución biótica de México: encrucijada de la biodiversidad del Nuevo Mundo. Revista mexicana de biodiversidad, 90, 1–68. https://doi.org/10.22201/ib.20078706e.2019.90.2980
  55. Murphy, R.W., Blair, C. & Méndez-de la Cruz, F.R. (2009) A new species of leaf-toed gecko, genus Phyllodactylus (Squamata: Gekkota: Phyllodactylidae) from Guerrero, Mexico. South American Journal of Herpetology, 4 (1), 17–24. https://doi.org/10.2994/057.004.0103
  56. Ochoa-Ochoa, L., Flores-Villela, O., García-Vázquez, U., Correa-Cano, M. & Canseco-Márquez, L. (2006) Phyllodactylus bordai (Eslaboncillo). Área de distribución potencial, escala: 1:1000000. Museo de Zoología “Alfonso L. Herrera”, Facultad de Ciencias, Universidad Nacional Autónoma de México. Proyecto: DS009, Extraído del proyecto DS009: Áreas potenciales de distribución y GAP análisis de la herpetofauna de México. Available from: http://geoportal.conabio.gob.mx/metadatos/doc/html/phylld_bordgw.html (accessed 22 Novembre 2024)
  57. Paradis, E. (2010) pegas: an R package for population genetics with an integrated-modular approach. Bioinformatics, 26, 419–420. https://doi.org/10.1093/bioinformatics/btp696
  58. Paradis, E. & Schliep, K. (2019) ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics, 35, 526–528. https://doi.org/10.1093/bioinformatics/bty633
  59. Pembleton, L.W., Cogan, N.O.I. & Forster, J.W. (2013) StAMPP: An R package for calculation of genetic differentiation and structure of mixed‐ploidy level populations. Molecular Ecology Resources, 13 (5), 946–952. https://doi.org/10.1111/1755-0998.12129
  60. R Core Team (2023) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.R-project.org/ (accessed 23 Novembre 2024)
  61. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
  62. Ramírez-Reyes, T., Piñero, D., Flores-Villela, O. & Vázquez-Domínguez, E. (2017) Molecular systematics, species delimitation and diversification patterns of the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico. Molecular Phylogenetics and Evolution, 115, 82–94. https://doi.org/10.1016/j.ympev.2017.07.008
  63. Ramírez-Reyes, T. & Flores-Villela, O. (2018) Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico. Zootaxa, 4407 (2), 151–190. https://doi.org/10.11646/zootaxa.4407.2.1
  64. Ramírez-Reyes, T., Blair, C., Flores-Villela, O., Piñero, D., Lathrop, A. & Murphy, R. (2020) Phylogenomics and molecular species delimitation reveals great cryptic diversity of leaf-toed geckos (Phyllodactylidae: Phyllodactylus), ancient origins, and diversification in Mexico. Molecular Phylogenetics and Evolution, 150, 106880. https://doi.org/10.1016/j.ympev.2020.106880
  65. Ramírez‐Reyes, T., Flores‐Villela, O., Piñero, D., Lathrop, A. & Murphy, R.W. (2021a) Genomic assessment of the Phyllodactylus tuberculosus complex (Reptilia: Phyllodactylidae) in America. Zoologica Scripta, 50 (5), 529–542. https://doi.org/10.1111/zsc.12492
  66. Ramírez-Reyes, T., Barraza-Soltero, I.K., Nolasco-Luna, J.R., Flores-Villela, O. & Escobedo-Galván, A.H. (2021b) A new species of leaf-toed gecko (Phyllodactylidae, Phyllodactylus) from María Cleofas Island, Nayarit, Mexico. ZooKeys, 1024, 117–136. https://doi.org/10.3897/zookeys.1024.60473
  67. Ramírez-Reyes, T., Velasco, J.A., Flores-Villela, O. & Piñero, D. (2022) Decoupling in diversification and body size rates during the radiation of Phyllodactylus: evidence suggests minor role of ecology in shaping phenotypes. Evolutionary Biology, 49 (3), 373–387. https://doi.org/10.1007/s11692-022-09575-z
  68. Rick, J.A., Brock, C.D., Lewanski, A.L., Golcher-Benavides, J. & Wagner, C.E. (2024) Reference genome choice and filtering thresholds jointly influence phylogenomic analyses. Systematic Biology, 73 (1), 76–101. https://doi.org/10.1093/sysbio/syad065
  69. SEMARNAT-CONANP. (2013) Programa de Manejo Reserva de la Biosfera Tehuacán-Cuicatlán. SEMARNAT Tlalpan CDMX, Tlalpan, Mexico City, 336 pp.
  70. Simpson, M.G. (2010) Phylogenetic systematics. In: Simpson, M.G. (Ed.), Plant Systematics. 2nd Edition. Academic Press, San Diego, California, pp. 17–52. https://doi.org/10.1016/B978-0-12-374380-0.50002-6
  71. Skipwith, P.L., Bi, K. & Oliver, P.M. (2019) Relicts and radiations: Phylogenomics of an Australasian lizard clade with east Gondwanan origins (Gekkota: Diplodactyloidea). Molecular Phylogenetics and Evolution, 140, 106589. https://doi.org/10.1016/j.ympev.2019.106589
  72. Šmíd, J., Mazuch, T., Nováková, L., Modrý, D., Malonza, P.K., Elmi, H.S.A., Carranza, S. & Moravec, J. (2019) Phylogeny and systematic revision of the gecko genus Hemidactylus from the Horn of Africa (Squamata: Gekkonidae). Herpetological Monographs, 33 (1), 26–47. https://doi.org/10.1655/HERPMONOGRAPHS-D-19-00010.1
  73. Smith, H.M. (1935) Miscellaneous notes on Mexican lizards. The University of Kansas Science Bulletin, 36, 119–155.
  74. Smith, H.M. & Taylor, E.H. (1950) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin United States Natlional Museum, 199, 1–253. https://doi.org/10.5479/si.03629236.199
  75. Soto-Trejo, F., Robles, F., Lira, R., Sánchez-González, L.A., Ortiz, E. & Dávila, P. (2024) The evolution of paleo-and neo-endemic species of Cactaceae in the isolated Valley of Tehuacán-Cuicatlán, Mexico. Plant Ecology and Evolution, 157 (1), 42–54. https://doi.org/10.5091/plecevo.110352
  76. Swofford, D.L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts. [program]
  77. Taylor, E.H. (1942) Some geckos of the genus Phyllodactylus. University Kansas Science Bulletin, 28 (6), 91–112.
  78. Téllez-Valdés, O., Farías, V., Dávila-Aranda, P., Louis-Stein, J., Lira-Saade, R. & Botello, F.J. (2010) Diversidad de mamíferos en los dominios climáticos de la Reserva de la Biosfera Tehuacán-Cuicatlán, México. Revista Mexicana de Biodiversidad, 81 (3), 863–874. https://doi.org/10.22201/ib.20078706e.2010.003.656
  79. Theissinger, K., Fernandes, C., Formenti, G., Bista, I., Berg, P.R., Bleidorn, C., Bombarely, A., Crottini, A., Gallo, G.R., Godoy, J.A., Jentoft, S., Malukiewicz, J., Mouton, A., Oomen, R.A., Paez, S., Palsbøll, P.J., Pampoulie, C., Ruiz-López, M.J., Secomandi, S., Svardal, H., Theofanopoulou, C., de Vries, J., Waldvogel, A.-M., Zhang, G., Jarvis, E.D., Bálint, M., Ciofi, C., Waterhouse, R.M. & Mazzoni, C.J. (2023) How genomics can help biodiversity conservation. Trends in Genetics, 39 (7), 545–559. https://doi.org/10.1016/j.tig.2023.01.005
  80. Uetz, P., Slavenko, A., Meiri, S. & Heinicke, M. (2020) Gecko diversity: a history of global discovery. Israel Journal of Ecology and Evolution, 66 (3-4), 117–125. https://doi.org/10.1163/22244662-bja10003
  81. Uetz, P., Freed, P, Aguilar, R., Reyes, F., Kudera, J. & Hošek, J. (Eds.) (2024) The Reptile Database. Available from: http://www.reptile-database.org (accessed 31 October 2024)
  82. Valiente-Banuet, A., Casas, A., Alcántara, A., Dávila, P., Flores, H.N., Arizmendi, M.C., Villaseñor, J.L. & Ortega, J. (2000) La vegetación del Valle de Tehuacán-Cuicatlán. Boletín de la Sociedad Botánica de México, 67, 24–74. https://doi.org/10.17129/botsci.1625
  83. Valiente-Banuet, A., Solís, L., Dávila, P., Arizmendi, M.C., Silva-Pereyra, C., Ortega-Ramírez, J., Treviño-Carreón, J., Rangel-Landa, S. & Casas, A. (2009) Guía de la Vegetación del Valle de Tehuacán-Cuicatlán. CONABIO, Fundación para la Reserva de la Biósfera Cuicatlán, A. C., IE, CIECO, UBIPRO-FES-Iztacala, UNAM, Ciudad de Mexico, 211 pp.
  84. Villaseñor, J.L., Dávila, P. & Chiang, F. (1990) Fitogeografía del Valle de Tehuacán- Cuicatlán. Boletín de la Sociedad Botánica de México, 50, 135–149. https://doi.org/10.17129/botsci.1381
  85. Vitt, L.J. & Caldwell, J.P. (2014) Herpetology: An Introductory Biology of Amphibians and Reptiles. 4th Edition. Academic Press, San Diego, 757 pp. https://doi.org/10.1016/B978-0-12-386919-7.00002-2
  86. Vu, V.Q., Friendly, M. & Tavadyan, A. (2024) ggbiplot: A Grammar of Graphics Implementation of Biplots. Available from: https://github.com/friendly/ggbiplot and https://friendly.github.io/ggbiplot/ (accessed 2 November 2024) https://doi.org/10.32614/CRAN.package.ggbiplot
  87. Wickham, H., Hester, J., Chang, W. & Bryan, J. (2022) devtools: Tools to Make Developing R Packages Easier. Available from: https://devtools.r-lib.org/ and https://github.com/r-lib/devtools (accessed 2 November 2024)
  88. Wood Jr., P.L., Guo, X., Travers, S.L., Su, Y.C., Olson, K.V., Bauer, A.M., Grismer, L.L, Siler, C.D., Moyle, R.G., Andersen, M.J. & Brown, R.M. (2020) Parachute geckos free fall into synonymy: Gekko phylogeny, and a new subgeneric classification, inferred from thousands of ultraconserved elements. Molecular Phylogenetics and Evolution, 146, 106731. https://doi.org/10.1016/j.ympev.2020.106731
  89. Woolrich-Piña, G.A., García-Padilla, E., DeSantis, D.L., Johnson, J.D., Mata-Silva, V. & Wilson, L.D. (2017) The herpetofauna of Puebla, Mexico: composition, distribution, and conservation. Mesoamerican Herpetology, 4 (4), 794–884.
  90. Yang, Z. & Rannala, B. (2014) Unguided species delimitation using DNA sequence data from multiple loci. Molecular Biology and Evolution, 31 (12), 3125–3135. https://doi.org/10.1093/molbev/msu279

How to Cite

Ramírez-Reyes, T., Durán-Arceo, D.R., Palacios-Aguilar, R. & Flores-Villela, O. (2025) Taxonomic revision of the polyphyletic Phyllodactylus bordai (Squamata: Phyllodactylidae), with the description of a new species. Zootaxa, 5722 (2), 151–183. https://doi.org/10.11646/zootaxa.5722.2.1