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Issue: Vol. 5647 No. 1: 12 Jun. 2025
Type: Article
Published: 2025-06-12
DOI: 10.11646/zootaxa.5647.1.2
Page range: 27-50
Abstract views: 134
PDF downloaded: 5

Systematics of Ecuadorian anoles: Anolis bitectus Cope, 1864 and A. lemniscatus Boulenger, 1898, two junior synonyms of A. binotatus Peters, 1863 (Reptilia: Squamata: Anolidae)

Museo de Zoología; Departamento de Ciencias Biológicas; Pontificia Universidad Católica del Ecuador; Avenida 12 de Octubre 1076 y Roca; Apartado 17-01-2184; Quito; Ecuador
Department of Biology; University of New Mexico; 219 Yale Boulevard NE; Albuquerque; NM 87106; USA
Museo de Zoología; Departamento de Ciencias Biológicas; Pontificia Universidad Católica del Ecuador; Avenida 12 de Octubre 1076 y Roca; Apartado 17-01-2184; Quito; Ecuador; Unidad de Investigación; Instituto Nacional de Biodiversidad. Rumipamba 341 y Av. de los Shyris. Casilla postal: 17-07-8976. Quito; Ecuador; Integrative Biology Laboratory; Universidad Regional Amazónica Ikiam; Tena; Napo; Ecuador
Botanical Garden; Martin-Luther-University Halle-Wittenberg. Am Kirchtor 3; D-06108 Halle/Saale; Germany
Senckenberg Forschungsinstitut und Naturmuseum; Senckenberganlage 25; 60325 Frankfurt a.M.; Germany
Museo de Zoología; Departamento de Ciencias Biológicas; Pontificia Universidad Católica del Ecuador; Avenida 12 de Octubre 1076 y Roca; Apartado 17-01-2184; Quito; Ecuador
Reptilia Draconura Norops phylogeny synonymy types

Abstract

More than a century ago, three species of lizard with similar morphology were described from Ecuador: Anolis binotatus Peters 1863, A. bitectus Cope 1864, and A. lemniscatus Boulenger 1898. We studied the genetics, type material and morphological variation throughout the range of these forms to evaluate the taxonomic status of these names. Our phylogenetic analyses, based on mitochondrial genes such as COI and ND2, and a nuclear gene such as RAG1 revealed three geographically structured clades (northern, central and southern) similar in morphology and coloration, that did not correspond to the three species in question. Similarly, we found no justification for recognizing all three species based on examination of type material and comparisons of morphology across the geographic range of specimens assigned to the three species. Although there were significant differences in snout length and the number of scales between the interparietal and supraorbital semicircles among recovered molecular clades, we do not consider these differences as diagnostic at the species level. Therefore, we propose that A. binotatus is a senior synonym of both A. bitectus and A. lemniscatus. We provide a new description and variation of A. binotatus, including hemipenis, natural history, geographic distribution and conservation status.

 

References

  1. Amador, L., Ayala-Varela, F., Nárvaez, A.E., Cruz, K. & Torres-Carvajal, O. (2017) First record of the invasive Brown Anole, Anolis sagrei Duméril & Bibron, 1837 (Squamata: Iguanidae: Dactyloinae), in South America. Check List, 13 (2), 1–6. https://doi.org/10.15560/13.3.2083
  2. Andrews, R.M. (1971) Structural habitat and time budget of a tropical Anolis lizard. Ecology, 52 (2), 262–270. https://doi.org/10.2307/1934584
  3. Ayala-Varela, F., Caicedo, J.R., Calderón, M., Ines Hladki, A., Ramírez Pinilla, M., Renjifo, J., Rivas, G. & Urbina, N. (2020) Anolis binotatus. The IUCN Red List of Threatened Species, 2020, e.T44577183A44577188. https://doi.org/10.2305/IUCN.UK.2020-3.RLTS.T44577183A44577188.en
  4. Ayala-Varela, F., Valverde, S., Poe, S., Narváez, A.E., Yánez-Muñoz, M.H. & Torres-Carvajal, O. (2021) A new giant anole (Squamata: Iguanidae: Dactyloinae) from southwestern Ecuador. Zootaxa, 4991 (2), 295–317. https://doi.org/10.11646/zootaxa.4991.2.4
  5. Ayala-Varela, F.P., Troya-Rodríguez, D., Talero-Rodríguez, X. & Torres-Carvajal, O. (2014) A new Andean anole species of the Dactyloa clade (Squamata: Iguanidae) from western Ecuador. Amphibian & Reptile Conservation, Special Section, 8 (1), 8–24. [https://www.biodiversitylibrary.org/part/178775]
  6. Boulenger, G.A. (1898) An account of the reptiles and batrachians collected by Mr. W. F. H. Rosenberg in western Ecuador. Proceedings of the Zoological Society of London, 1898, 107–126. https://doi.org/10.1111/j.1096-3642.1898.tb03134.x
  7. Camacho Badani, T. (2013) Sistemática del complejo Pristimantis conspicillatus y Pristimantis peruvianus (Anura: Craugastoridae). Pontificia Universidad Católica del Ecuador, Quito. [unknown pagination]
  8. Castañeda, M.D.R. & Queiroz, K.D.E. (2013) Phylogeny of the Dactyloa clade of Anolis lizards: New insights from combining morphological and molecular data. Bulletin of the Museum of Comparative Zoology, 160 (7), 345–398. https://doi.org/10.3099/0027-4100-160.7.345
  9. Cope, E.D. (1864) Contributions to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences of Philadelphia, 16 (3), 166–176.
  10. Cruz-García, K., Barreno, M., Cuadrado, S., Moretta-Urdiales, M., Molina-Moreira, N., Salas, J.A., Torres-Domínguez, Á. & Narváez, A.E. (2023) Amphibians and reptiles of Isla Santay (Durán, Ecuador). Check List, 19 (3), 347–369. https://doi.org/10.15560/19.3.347
  11. D’Angiolella, A.B., Klaczko, J., Rodrigues, M.T. & Avila-Pires, T.C.S. (2016) Hemipenial morphology and diversity in South American anoles (Squamata: Dactyloidae). Canadian Journal of Zoology, 94 (4), 251–256. https://doi.org/10.1139/cjz-2015-0194
  12. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic acids research, 32 (5), 1792–1797. https://doi.org/10.1093/nar/gkh340
  13. Esri (2020) ArcMap. Version 10.8. Environmental Systems Research Institute, Redlans, California. [Software]
  14. Fitch, H.S., Echelle, A.F. & Echelle, A.A. (1976) Field observations on rare or little known mainland anoles. The University of Kansas science bulletin, 51, 91–128. https://doi.org/10.5962/bhl.part.24957
  15. Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular marine biology and biotechnology, 3 (5), 294–299.
  16. Garzón-Santomaro, C., Sánchez-Nivicela, J.C., Mena-Valenzuela, P., González-Romero, D. & Mena-Jaén, J. (2019) Anfibios, reptiles y aves de la provincia de El Oro. Una guía para la identificación de especies del páramo al manglar. Segunda Edición. Publicación Miscelánea No. 11. Serie de Publicaciones GADPEO-INABIO. GADPEO-INABIO, Machala, El Oro. [unknown pagination]
  17. Geneious Prime (2020) Geneious Prime. Version 2020.2.2. Biomatters Ltd. Auckland. [software]
  18. 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
  19. 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 (6), 587–589. https://doi.org/10.1038/nmeth.4285
  20. Köhler, G. (2014) Characters of external morphology used in Anolis taxonomy. Definition of terms, advice on usage, and illustrated examples. Zootaxa, 3774 (3), 201–257. https://doi.org/10.11646/zootaxa.3774.3.1
  21. Kumazawa, Y. & Nishida, M. (1993) Sequence evolution of mitochondrial tRNA genes and deep-branch animal phylogenetics. Journal of Molecular Evolution, 37, 380–398. https://doi.org/10.1007/BF00178868
  22. Losos, J.B. (2009) Lizards in an evolutionary tree: Ecology and adaptive radiation of anoles. University of California Press, Berkeley, California, 507 pp. https://doi.org/10.1525/california/9780520255913.001.0001
  23. Macey, J.R., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular biology and evolution, 14 (1), 91–104. https://doi.org/10.1093/oxfordjournals.molbev.a025706
  24. Minh, B.Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30 (5), 1188–1195. https://doi.org/10.1093/molbev/mst024
  25. Nicholson, K.E. (2002) Phylogenetic analysis and a test of the current infrageneric classification of Norops (beta Anolis). Herpetological Monographs, 16, 93–120. https://doi.org/10.1655/0733-1347(2002)016[0093:paaato]2.0.co;2
  26. Nicholson, K.E., Crother, B.I., Guyer, C. & Savage, J.M. (2012) It is time for a new classification of anoles (Squamata: Dactyloidae). Zootaxa, 3477 (1), 1–108. https://doi.org/10.11646/zootaxa.3477.1.1
  27. Nicholson, K.E., Glor, R., Kolbe, J., Larson, A., Hedges, S.B. & Losos, J. (2005) Mainland colonization by island lizards. Journal of Biogeography, 32 (6), 929–938. https://doi.org/10.1111/j.1365-2699.2004.01222.x
  28. Ortega, J., Martínez, J.L. & Tirira, D.G. (2014) Historia de la mastozoología en Latinoamérica, las Guayanas y el Caribe. Editorial Murciélago Blanco y Asociación Ecuatoriana de Mastozoología, Quito and México, D.F., 328 pp.
  29. Paynter, R.A. (1993) Ornithological gazetteer of Ecuador. 2nd Edtion. Bird Dept., Museum of Comparative Zoology, Harvard University, Cambridge, Masschusetts, xi + 247 pp. [https://www.biodiversitylibrary.org/item/50689] https://doi.org/10.5962/bhl.title.14626
  30. Pesantes, O.S. (1994) A method for preparing the hemipenis of preserved snakes. Journal of Herpetology, 28 (1), 93–95. https://doi.org/10.2307/1564686
  31. Peters, W.C.H. (1863) Mittheilung über eine neue Arten der Saurier-Gattun Anolis. Monatsberichte der Preussischen Akademie der Wissenschaften zu Berlin, 1863, 135–149.
  32. Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18 (1), 37–89. https://doi.org/10.1655/0733-1347(2004)018[0037:POA]2.0.CO;2
  33. Poe, S., Nieto-Montes De Oca, A., Torres-Carvajal, O., De Queiroz, K., Velasco, J.A., Truett, B., Gray, L.N., Ryan, M.J., Köhler, G., Ayala-Varela, F., Ayala-Varela, F. & Ian Latella, A. (2017) A phylogenetic, biogeographic, and taxonomic study of all extant species of Anolis (Squamata; Iguanidae). Systematic Biology, 66 (5), 663–697. https://doi.org/10.1093/sysbio/syx029
  34. Prates, I., Melo-Sampaio, P.R., Queiroz, K.D., Carnaval, A.C., Rodrigues, M.T. & Drummond, L.D.O. (2020) Discovery of a new species of Anolis lizards from Brazil and its implications for the historical biogeography of montane Atlantic Forest endemics. Amphibia Reptilia, 41 (1), 87–103. https://doi.org/10.1163/15685381-20191179
  35. 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.), Forms: Species and Speciation. Oxford University Press, New York, New York, pp. 57–75.
  36. de Queiroz, K. (2007) Species concepts and species delimitation. Systematic biology, 56 (6), 879–886. https://doi.org/10.1080/10635150701701083
  37. Rambaut, A. (2018) FigTree: Tree figure drawing tool. Available from: https://tree.bio.ed.ac.uk/software/figtree/ (accessed 10 May 2025)
  38. Reyes-Puig, C.P. & Ríos-Alvear, G.D. (2016) Ampliación del rango altitudinal de Anolis fasciatus (Squamata: Dactyloidae) en Ecuador. Avances en Ciencias e Ingenierías, 8 (1). [published online] https://doi.org/10.18272/aci.v8i14.448
  39. Ron, S.R. (2020) Regiones naturales del Ecuador. BIOWEB. Pontificia Universidad Católica del Ecuador. Available from: https://bioweb.bio/faunaweb/amphibiaweb/RegionesNaturales (acceseed 10 May 2025)
  40. RStudio Team (2020) RStudio: Integrated development for R. RStudio, PBC, Boston, Massachusetts. [program]
  41. Savage, J.M. (1997) On terminology for the description of the hemipenes of squamate reptiles. Herpetological Journal, 7, 23–25.
  42. Schulte, J.A. & Cartwright, E.M. (2009) Phylogenetic relationships among iguanian lizards using alternative partitioning methods and TSHZ1: a new phylogenetic marker for reptiles. Molecular Phylogenetics and Evolution, 50 (2), 391–396. https://doi.org/10.1016/j.ympev.2008.10.018
  43. Simpson, G.G. (1951) The species concept. Evolution, 5 (4), 285–298. https://doi.org/10.2307/2405675
  44. Simpson, G.G. (1961) Principles of animal taxonomy. Columbia University Press, New York City, New York, xii + 247 pp. https://doi.org/10.7312/simp92414
  45. Torres-Carvajal, O., Pazmiño-Otamendi, G., Ayala-Varela, F. & Salazar-Valenzuela, D. (2024) Reptiles del Ecuador. Version 2024.1. Museo de Zoología, Pontificia Universidad Católica del Ecuador, Quito. [program]
  46. Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44 (W1), 232–235. https://doi.org/10.1093/nar/gkw256
  47. Uetz, P., Freed, P., Aguilar, R., Reyes, F. & Hošek, J. (2023) The Reptile Database.
  48. VertNet (2020) VertNet Search Portal. Available from: http://portal.vertnet.org/search (accessed 10 May 2025)
  49. Vitt, L.J., Avila-Pires, T.C.S., Zani, P.A. & Espósito, M.C. (2002) Life in shade: The ecology of Anolis trachyderma (Squamata: Polychrotidae) in Amazonian Ecuador and Brazil, with comparisons to ecologically similar anoles. Copeia, 2, 275–286. https://doi.org/10.1643/0045-8511(2002)002[0275:LISTEO]2.0.CO;2
  50. Vitt, L.J., Avila-Pires, T.C.S., Zani, P.A., Sartorius, S.S. & Espósito, M.C. (2003) Life above ground: Ecology of Anolis fuscoauratus in the Amazon rain forest, and comparisons with its nearest relatives. Canadian Journal of Zoology, 81 (1), 142–156. https://doi.org/10.1139/z02-230
  51. Williams, E., Rand, H., Rand, A. & O’Hara, R. (1995a) A computer approach to the comparison and identification of species in difficult taxonomic groups. Breviora, 502, 1–47.
  52. Yánez-Muñoz, M.H., Morales, M., Reyes-Puig, M. & Meza-Ramos, P.A. (2013) Reserva Biológica Buenaventura: Entre la transición húmeda tropical y la influencia tumbesina. In: MECN, Jocotoco & Ecominga (Eds.), Herpetofauna en áreas prioriotarias para la conservación: El sistema de Reservas Jocotoco y Ecominga. Serie de Publicaciones del Museo Ecuatoriano de Ciencias Naturales (MECN), Fundación para la Conservación Jocotoco, Fundación Ecominga, Quito, pp. 62–76.

How to Cite

Ayala-Varela, F., Poe, S., Posse-Sarmiento, V., Fläschendräger, A., Köhler, G. & Torres-Carvajal, O. (2025) Systematics of Ecuadorian anoles: Anolis bitectus Cope, 1864 and A. lemniscatus Boulenger, 1898, two junior synonyms of A. binotatus Peters, 1863 (Reptilia: Squamata: Anolidae). Zootaxa, 5647 (1), 27–50. https://doi.org/10.11646/zootaxa.5647.1.2