Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2026-04-13
Page range: 245-286
Abstract views: 205
PDF downloaded: 8

Facing a slithery dragon: systematics of the Petracola ventrimaculata (Boulenger, 1900) species complex (Squamata: Gymnophthalmidae) with description of five new species

Departamento de Herpetología; Museo de Historia Natural de San Marcos (MUSM); Av. Arenales 1256; Jesus Maria; Lima; Perú
Rainforest Partnership; 4005 Guadalupe St.; Austin; Texas 78751; USA; Instituto Peruano de Herpetología (IPH); Augusto Salazar Bondy 136; Urb. Higuereta; Surco; Lima; Perú
Rainforest Partnership; 4005 Guadalupe St.; Austin; Texas 78751; USA; Instituto Peruano de Herpetología (IPH); Augusto Salazar Bondy 136; Urb. Higuereta; Surco; Lima; Perú
Instituto Peruano de Herpetología (IPH); Augusto Salazar Bondy 136; Urb. Higuereta; Surco; Lima; Perú; Herbario KUELAP; Facultad de Ingeniería y Ciencias Agrarias; Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas (UNTRM); Chachapoyas 01001; Perú
Museo de Zoología; Centro de Investigaciones de la Biodiversidad CIBIO; Pontificia Universidad Católica del Ecuador; Avenida 12 de Octubre 1076 y Roca; Apartado 17–01–2184; Quito; Ecuador
Reptilia Andes Cercosaurinae conservation cryptic species Huancabamba Depression phylogeny taxonomy

Abstract

Nearly two decades ago, the gymnophthalmid lizard Petracola ventrimaculata was hypothesized to be a species complex based on dorsal and ventral color pattern differences among populations within a wide and disjunct geographical area in the northern Peruvian Andes. Here, we address this proposal using both molecular and morphological evidence. We analyzed a molecular matrix of Cercosaurinae, including 22 specimens and 78 DNA sequences of P. ventrimaculata sensu lato to study the phylogenetic relationships among its populations. We also conducted a detailed examination of the external morphology of 236 P. ventrimaculata specimens from several populations. Based on our analyses, we describe five new Petracola species, two of them previously assigned to P. ventrimaculata, from Cordillera Occidental and Cordillera Central in northern Peru. P. ventrimaculata sensu stricto is distributed across several localities in Cajamarca department in the northern part of the Cordillera Occidental in northwestern Peru. We hypothesize that the distribution patterns within the Petracola ventrimaculata species complex could be related to vicariant processes driven by the complex orogeny of the Huancabamba Depression and the Marañón River valley. Given its distinctive morphological traits and disjunct distribution, we recommend treating P. labioocularis as incertae sedis within Cercosaurinae, pending molecular data to resolve its phylogenetic placement. We present recommendations for future descriptions of new species of Petracola.

 

References

  1. Aguilar, C., Wood, P.L., Jr., Cusi, J.C., Guzman, A., Huari, F., Lundberg, M., Mortensen, E., Ramirez, C., Robles, D., Suarez, J., Ticona, A., Vargas, V.J., Venegas, P.J. & Sites, J.W., Jr. (2013) Integrative taxonomy and preliminary assessment of species limits in the Liolaemus walkeri complex (Squamata, Liolaemidae) with descriptions of three new species from Peru. Zookeys, 364, 47–91. https://doi.org/10.3897/zookeys.364.6109
  2. Altamirano, M.A., Zaher, H., Lobo, L., Grazziotin, F.G., sales Nunes, P.M. & Trefaut Rodrigues, M. (2013) A new species of lizard genus Potamites from Ecuador (Squamata, Gymnophthalmidae). Zootaxa, 3717 (3), 345–358. https://doi.org/10.11646/zootaxa.3717.3.4
  3. Amézquita, A., Mazariegos-H, L.A., Cañaveral, S., Orejuela, C., Barragán-Contreras, L.A. & Daza, J.M. (2023) Species richness under a vertebral stripe: integrative taxonomy uncovers three additional species of Pholidobolus lizards (Sauria, Squamata, Gymnophthalmidae) from the north-western Colombian Andes. Zookeys, 1141, 119–148. https://doi.org/10.3897/zookeys.1141.94774
  4. Angulo, A. (2002) Anfibios y paradojas: perspectivas sobre la diversidad y las poblaciones de anfibios. Ecología Aplicada, 1, 105–109. https://doi.org/10.21704/rea.v1i1-2.237
  5. Angulo, A. & Icochea, J. (2010) Cryptic species complexes, widespread species and conservation: lessons from Amazonian frogs of the Leptodactylus marmoratus group (Anura: Leptodactylidae). Systematics and Biodiversity, 8 (3), 357–370. https://doi.org/10.1080/14772000.2010.507264
  6. Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117–126. https://doi.org/10.3897/zookeys.150.2109
  7. Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K., Meier, R., Winker, K., Ingram, K.K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology & Evolution, 22 (3), 148–155. https://doi.org/10.1016/j.tree.2006.11.004
  8. Cadle, J.E. (1991) Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Perú: New species and comments on relationships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadelphia, 143, 1–96.
  9. Chávez, G., Malqui, J. & Catenazzi, A. (2021) A new riparian Andean Potamites (Reptilia, Squamata, Gymnophtalmidae) from El Sira Mountains, central Peru, with comments on P. ecpleopus Cope 1875, and on the taxonomy and biogeography of Potamites. European Journal of Taxonomy, 760, 136–159. https://doi.org/10.5852/ejt.2021.760.1445
  10. Chernomor, O., Von Haeseler, A. & Minh, B.Q. (2016) Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology, 65, 997–1008. https://doi.org/10.1093/sysbio/syw037
  11. de Oliveira, D.P., de Carvalho, V.T. & Hrbek, T. (2016) Cryptic diversity in the lizard genus Plica (Squamata): phylogenetic diversity and Amazonian biogeography. Zoologica Scripta, 45, 630–641. https://doi.org/10.1111/zsc.12172
  12. de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless forms: species and speciation. Oxford University Press, New York, Oxford, pp. 57–75. https://doi.org/10.1093/oso/9780195109009.003.0005
  13. Doan, T.M. (2003) A south‐to‐north biogeographic hypothesis for Andean speciation: evidence from the lizard genus Proctoporus (Reptilia, Gymnophthalmidae). Journal of Biogeography, 30, 361–374. https://doi.org/10.1046/j.1365-2699.2003.00833.x
  14. Doan, T.M. & Castoe, T.A. (2005) Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus. Zoological Journal of the Linnean Society, 143, 405–416. https://doi.org/10.1111/j.1096-3642.2005.00145.x
  15. Duellman, W.E. (1979) The South American Herpetofauna: Its Origin, Evolution, and Dispersal. Monograph of the Museum of Natural History, University of Kansas. No. 7. Museum of Natural History, University of Kansas, Lawrence, Kansas, 485 pp. https://doi.org/10.5962/bhl.title.3207
  16. Duellman, W.E. & Pramuk, J.B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of Northern Peru. Museum of Natural History, the University of Kansas, 13, 1–78. https://doi.org/10.5962/bhl.title.16169
  17. Echevarría, L.Y. & Venegas, P.J. (2013) Petracola ventrimaculatus (spotted lightbulb lizard) cannibalism. Herpetological Review, 44, 678–679.
  18. Echevarría, L.Y. & Venegas, P.J. (2015) A new elusive species of Petracola (Squamata: Gymnophthalmidae) from the Utcubamba basin in the Andes of northern Peru. Amphibian & Reptile Conservation, 9 (1), 26–33.
  19. Ely, C.V., de Loreto Bordignon, S.A., Trevisan, R. & Boldrini, I.I. (2017) Implications of poor taxonomy in conservation. Journal for Nature Conservation, 36, 10–13. https://doi.org/10.1016/j.jnc.2017.01.003
  20. Funk, W.C., Caminer, M. & Ron, S.R. (2011) High levels of cryptic species diversity uncovered in Amazonian frogs. Proceedings of the Royal Society B, Biological Sciences, 279, 1806–1814. https://doi.org/10.1098/rspb.2011.1653
  21. 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, 307–321. https://doi.org/10.1093/sysbio/syq010
  22. Hazzi, N.A., Moreno, J.S., Ortiz-Movliav, C. & Palacio, R.D. (2018) Biogeographic regions and events of isolation and diversification of the endemic biota of the tropical Andes. Proceedings of the Academy of Natural Sciences, 115 (3), 7985–7990. https://doi.org/10.1073/pnas.1803908115
  23. Hending, D. (2025) Cryptic species conservation: a review. Biological Reviews, 100, 258–274. https://doi.org/10.1111/brv.13139
  24. 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, 518–522. https://doi.org/10.1093/molbev/msx281
  25. IUCN (2024) Guidelines for Using the IUCN Red List Categories and Criteria. Version 16. IUCN Standards and Petitions Committee, Gland, Switzerland. [https://www.iucnredlist.org/documents/RedListGuidelines.pdf]
  26. 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
  27. Katoh, K., Rozewicki, J. & Yamada, K.D. (2019) MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics, 20, 1160–1166. https://doi.org/10.1093/bib/bbx108
  28. Kizirian, D., Bayefsky-Anand, S., Eriksson, A., Le, M. & Donnelly, M.A. (2008) A new Petracola and re-description of P. ventrimaculatus (Squamata: Gymnophthalmidae). Zootaxa, 1700 (1), 53–62. https://doi.org/10.11646/zootaxa.1700.1.4
  29. Kizirian, D.A. (1996) A review of Ecuatorian Proctoporus (Squamata: Gymnophthalmidae) with description of nine new species. Herpetological monographs, 1996, 85–155. https://doi.org/10.2307/1466981
  30. Köhler, G. & Lehr, E. (2004) Comments on Euspondylus and Proctoporus (Squamata: Gymnophthalmidae) from Peru, with the Description of Three New Species and a Key to the Peruvian Species. Herpetologica, 60 (4), 501–518. https://doi.org/10.1655/03-93
  31. Kok, P.J.R., Fouquet, A., Torres Carvajal, O., Means, D.B. & Rodrigues, M.T. (2025) New insights into the molecular phylogenetic relationships of lizards in the Neotropical genus Arthrosaura (Reptilia: Gymnophthalmidae) reveal rampant ‘cryptic’ speciation in the Guiana Shield. Zoological Journal of the Linnean Society, 204, zlaf045. https://doi.org/10.1093/zoolinnean/zlaf045
  32. Kuraku, S., Zmasek, C.M., Nishimura, O. & Katoh, K. (2013) aLeaves facilitates on-demand exploration of metazoan gene family trees on MAFFT sequence alignment server with enhanced interactivity. Nucleic Acids Research, 41, W22–W28. https://doi.org/10.1093/nar/gkt389
  33. Mace, G.M. (2004) The role of taxonomy in species conservation. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 359, 711–719. https://doi.org/10.1098/rstb.2003.1454
  34. Mamani, L., Vargas, V.J., Chaparro, J.C. & Catenazzi, A. (2023) Two new species of gymnophthalmid lizards of the genus Petracola (Squamata: Cercosaurinae) from the Andes of northeastern Peru, and their phylogenetic relationships. Amphibian & Reptile Conservation, 17, 161–173.
  35. Melo‐Sampaio, P.R., Passos, P., Prudente, A.L.C., Venegas, P.J. & Torres‐Carvajal, O. (2021) Systematic review of the polychromatic ground snakes Atractus snethlageae complex reveals four new species from threatened environments. Journal of Zoological Systematics and Evolutionary Research, 59, 718–747. https://doi.org/10.1111/jzs.12453
  36. MINAGRI. (2014) D.S. 004-2014. Decreto Supremo que aprueba la actualización de la lista de clasificación y categorización de las especies amenazadas de fauna silvestre legalmente protegidas. El Peruano, Normas Legales, Martes, 8, 520497–520504.
  37. Minh, B.Q., Nguyen, M.A.T. & Von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evololution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024
  38. More, A., Sánchez, I., Piana, R.P., Vallejos, J., Appleton, R.D. & Vallejos, J. (2022) Registros notables de mamíferos en una cordillera aislada, amenazada y no protegida en la depresión de Huancabamba, norte de Perú. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 46, 393–405. https://doi.org/10.18257/raccefyn.1624
  39. Murphy, J.C., Jowers, M.J., Lehtinen, R.M., Charles, S.P., Colli, G.R., Peres, A.K., Jr., Hendry, C.R. & Pyron, R.A. (2016) Cryptic, Sympatric Diversity in Tegu Lizards of the Tupinambis teguixin Group (Squamata, Sauria, Teiidae) and the Description of Three New Species. PLoS One, 11 (8), e0158542. https://doi.org/10.1371/journal.pone.0158542
  40. Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evololution, 32 (1), 268–274. https://doi.org/10.1093/molbev/msu300
  41. Nunes, P.M.S., Fouquet, A., Curcio, F.F., Kok, P.J.R. & Rodrigues, M.T. (2012) Cryptic species in Iphisa elegans Gray, 1851 (Squamata: Gymnophthalmidae) revealed by hemipenial morphology and molecular data. Zoological Journal of the Linnean Society, 166, 361–376. https://doi.org/10.1111/j.1096-3642.2012.00846.x
  42. Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. BioScience, 51 (11), 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
  43. Parra, V., Nunes, P.M.S. & Torres-Carvajal, O. (2020) Systematics of Pholidobolus lizards (Squamata, Gymnophthalmidae) from southern Ecuador, with descriptions of four new species. Zookeys, 954, 109–156. https://doi.org/10.3897/zookeys.954.50667
  44. Pfenninger, M. & Schwenk, K. (2007) Cryptic animal species are homogeneously distributed among taxa and biogeographical regions. BMC evolutionary biology, 7, 1–6. https://doi.org/10.1186/1471-2148-7-121
  45. Rodríguez, L.O. & Mamani, L. (2020) A new species of Petracola (Squamata: Gymnophthalmidae) from Río Abiseo National Park, San Martín, Peru. Amphibian & Reptile Conservation, 14 (1), 140–146.
  46. Simpson, G.G. (1951) The species concept. Evolution, 5, 285–298. https://doi.org/10.1111/j.1558-5646.1951.tb02788.x
  47. Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA11: molecular evolutionary genetics analysis version 11. Molecular biology and evolution, 38, 3022–3027. https://doi.org/10.1093/molbev/msab120
  48. Torres-Carvajal, O. (2007) Phylogeny and biogeography of a large radiation of andean lizards (Iguania, Stenocercus). Zoologica Scripta, 36, 311–326. https://doi.org/10.1111/j.1463-6409.2006.00284.x
  49. Torres-Carvajal, O., Echevarría, L.Y., Lobos, S.E., Venegas, P.J. & Kok, P.J.R. (2019) Phylogeny, diversity and biogeography of Neotropical sipo snakes (Serpentes: Colubrinae: Chironius). Molecular Phylogenetics and Evolution, 130, 315–329. https://doi.org/10.1016/j.ympev.2018.10.022
  50. Torres-Carvajal, O. & Hinojosa, K.C. (2020) Hidden diversity in two widespread snake species (Serpentes: Xenodontini: Erythrolamprus) from South America. Molecular Phylogenetics and Evolution, 146, 106772. https://doi.org/10.1016/j.ympev.2020.106772
  51. Torres-Carvajal, O., Lobos, S.E., Venegas, P.J., Chavez, G., Aguirre-Penafiel, V., Zurita, D. & Echevarría, L.Y. (2016) Phylogeny and biogeography of the most diverse clade of South American gymnophthalmid lizards (Squamata, Gymnophthalmidae, Cercosaurinae). Molecular Phylogenetics and Evolution, 99, 63–75. https://doi.org/10.1016/j.ympev.2016.03.006
  52. Torres-Carvajal, O. & Mafla-Endara, P. (2013) Evolutionary history of Andean Pholidobolus and Macropholidus (Squamata: Gymnophthalmidae) lizards. Molecular Phylogenetics and Evolution, 68, 212–217. https://doi.org/10.1016/j.ympev.2013.03.013
  53. Torres-Carvajal, O. & Teran, C. (2021) Molecular phylogeny of Neotropical Parrot Snakes (Serpentes: Colubrinae: Leptophis) supports underestimated species richness. Molecular Phylogenetics and Evolution, 164, 107267. https://doi.org/10.1016/j.ympev.2021.107267
  54. Torres-Carvajal, O., Venegas, P.J. & Sales Nunes, P.M. (2020) Description and Phylogeny of a New Species of Andean Lizard (Gymnophthalmidae: Cercosaurinae) from the Huancabamba Depression. South American Journal of Herpetology, 18 (1), 13–23. https://doi.org/10.2994/SAJH-D-18-00069.1
  55. Trifinopoulos, J., Nguyen, L.T., Von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE web server: fast and accurate phylogenetic trees under maximum likelihood. Nucleic Acids Res., 44, W232–W235. https://doi.org/10.1093/nar/gkw256
  56. Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics, 27, 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x
  57. Venegas, P., Torres, C., Suárez, J., Lehr, E. & Perez, J. (2019) Petracola ventrimaculatus. The IUCN Red List of Threatened Species, 2019, e.T48282388A48282419.
  58. Weigend, M. (2004) Four new species of Nasa ser. Alatae (Loasaceae) in the Amotape-Huancabamba zone of Peru. Novon, 14 (1) 134–146.
  59. Weigend, M., Rodríguez, E.F. & Arana, C. (2005) The relict forests of northwest Peru and southwest Ecuador. Revista Peruana de Biología, 12, 185–194. https://doi.org/10.15381/rpb.v12i2.2390
  60. Weir, J.T. (2009) Implications of genetic differentiation in neotropical montane forest birds. Annals of the Missouri Botanical Garden, 96 (3), 410–433. https://doi.org/10.3417/2008011
  61. Wiley, E.O. (1978) The evolutionary species concept reconsidered. Systematic Biology, 27, 17–26. https://doi.org/10.2307/2412809
  62. Wiley, E.O. & Lieberman, B.S. (2011) Phylogenetics: theory and practice of phylogenetic systematics. 2nd Edition. John Wiley & Sons, Hoboken, New Jersey, 440 pp. https://doi.org/10.1002/9781118017883
  63. Winger, B.M. & Bates, J.M. (2015) The tempo of trait divergence in geographic isolation: Avian speciation across the Marañon Valley of Peru. Evolution, 69, 772–787. https://doi.org/10.1111/evo.12607
  64. Yánez-Muñoz, M.H., Torres-Carvajal, O., Reyes-Puig, J.P., Urgiles-Merchán, M.A. & Koch, C. (2021) A new and very spiny lizard (Gymnophthalmidae: Echinosaura) from the Andes in northwestern Ecuador. PeerJ, 9, e12523. https://doi.org/10.7717/peerj.12523

How to Cite

Echevarría, L.Y., Venegas, P.J., García-Ayachi, L.A., Bravo, A.G.-. & Torres-Carvajal, O. (2026) Facing a slithery dragon: systematics of the Petracola ventrimaculata (Boulenger, 1900) species complex (Squamata: Gymnophthalmidae) with description of five new species. Zootaxa, 5792 (2), 245–286. https://doi.org/10.11646/zootaxa.5792.2.2