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Type: Article
Published: 2023-05-10
Page range: 121-141
Abstract views: 505
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No longer in doubt: Discovery of a second specimen corroborates the validity of Anolis incredulus Garrido and Moreno 1998 (Reptilia, Iguania)

Department of Vertebrate Zoology; National Museum of Natural History; Smithsonian Institution; Washington DC 20560; USA
Department of Vertebrate Zoology; National Museum of Natural History; Smithsonian Institution; Washington DC 20560; USA; Department of Biological Sciences; The George Washington University; Washington; DC 20052; USA
Department of Vertebrate Zoology; National Museum of Natural History; Smithsonian Institution; Washington DC 20560; USA
Reptilia Anolis angusticeps Anolis isolepis Anolis guazuma Cuba ecological morphology species inquirenda phylogenetic relationships


The species Anolis incredulus was proposed based on a single, poorly preserved specimen from the Sierra Maestra (mountain range) of southeastern Cuba. As its name suggests, this species was considered likely to raise doubts when it was first proposed, and it has been explicitly treated by some recent authors as a species inquirenda (a species of doubtful identity). Here we report on a second specimen of Anolis incredulus discovered in the amphibian and reptile collection of the National Museum of Natural History (Smithsonian Institution) that was collected more than 100 years before the holotype. We describe this specimen in detail and compare it both with the description of the holotype of A. incredulus and with presumed closely related Cuban species, providing evidence that it matches closely with the former and is distinct from the latter, thus corroborating the status of A. incredulus as a valid species. We also score and measure the specimen for sets of morphological characters to make inferences about its phylogenetic relationships and ecology (structural habitat use). Our results indicate that Anolis incredulus is likely a member of a clade of mostly Cuban twig-anole species and that it is a member of the twig ecomorph category, although its reported green coloration suggests either an erroneous ecomorph assignment or a difference in color from that of most other species of Cuban twig anoles.



  1. Barbour, T. & Ramsden, C.T. (1919) The herpetology of Cuba. Memoirs of the Museum of Comparative Zoology at Harvard College, 47 (2), 69–213, 15 pls.
  2. Díaz, L.M., Cádiz, A., Velazco, K. & Kawata, M. (2022) A new dwarf green anole (Squamata: Dactyloidae) of the Anolis carolinensis species group, from western Cuba. Caribbean Herpetology, 84, 1–16.
  3. Estrada, A.R. & Hedges, S.B. (1995) A new species of Anolis (Sauria: Iguanidae) from eastern Cuba. Caribbean Journal of Science, 31 (1–2), 65–72.
  4. Etheridge, R. (1959) The Relationships of the Anoles (Reptilia, Sauria, Iguanidae) an Interpretation based on Skeletal Morphology. Ph.D. dissertation. University of Michigan, Ann Arbor, Michigan, 236 pp.
  5. Feiner, N., Jackson, I.S.C., Munch, K.L., Radersma, R. & Uller, T. (2020) Plasticity and evolutionary convergence in the locomotor skeleton of Greater Antillean Anolis lizards. eLife, 9, e57468.
  6. Feiner, N., Jackson, I.S.C., Stanley, E. & Uller, T. (2021a) Evolution of the locomotor skeleton in Anolis lizards reflects the interplay between ecological opportunity and phylogenetic inertia. Nature Communications, 12, 1525.
  7. Feiner, N., Jackson, I.S.C., Van der Cruyssen, E. & Uller, T. (2021b) A highly conserved ontogenetic limb allometry and its evolutionary significance in the adaptive radiation of Anolis lizards. Proceedings of the Royal Society of London B, 288, 20210226.
  8. Garrido, O.H. (1983) Nueva especie de Anolis (Lacertilia: Iguanidae) de la Sierra del Turquino, Cuba. Caribbean Journal of Science, 19 (3–4), 71–76.
  9. Garrido, O.H. (1985) Nueva subespecie de Anolis isolepis (Lacertilia: Iguanidae) para Cuba. Doñana, Acta Vertebrata, 12 (1), 41–49.
  10. Garrido, O.H. & Moreno, L.V. (1998) Nueva especie de Anolis (Lacertilia: Iguanidae) del Pico Turquino, Sierra Maestra, Cuba. Avicennia, 8/9, 35–40.
  11. Garrido, O.H. & Hedges, S.B. (2001) A new anole from the northern slope of the Sierra Maestra in eastern Cuba (Squamata: Iguanidae). Journal of Herpetology, 35 (3), 378–383.
  12. Glor, R.E, Losos, J.B. & Larson, A. (2005) Out of Cuba: overwater dispersal and speciation among lizards in the Anolis carolinensis subgroup. Molecular Ecology, 14 (8), 2419–2432.
  13. Harrison, A. & Poe, S. (2012) Evolution of an ornament, the dewlap, in females of the lizard genus Anolis. Biological Journal of the Linnean Society, 106 (1), 191–201.
  14. Howard, R.A. (1988) Charles Wright in Cuba, 1856–1867. Chadwyck-Healey, Alexandria, Virginia, 90 pp.
  15. Huie, J.M., Prates, I., Bell, R.C. & de Queiroz, K. (2021) Convergent patterns of adaptive radiation between island and mainland Anolis lizards. Biological Journal of the Linnean Society, 134, 85–110.
  16. International Commission on Zoological Nomenclature (ICZN) (1999) International Code of Zoological Nomenclature. International Trust for Zoological Nomenclature, London, 306 pp.
  17. Kikinis, R., Pieper, S.D. & Vosburgh, K.G. (2014) 3D Slicer: a platform for subject-specific image analysis, visualization, and clinical support. In: Jolesz, F.A. (Ed.), Intraoperative Imaging and Image-guided Therapy. Springer, New York, New York, pp. 277–289.
  18. 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.
  19. 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.
  20. Losos, J.B. (2009) Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles. University of California Press, Berkeley, California, 507 pp.
  21. Poe, S. (1998) Skull characters and the cladistic relationships of the Hispaniolan dwarf twig Anolis. Herpetological Monographs, 12, 192–236.
  22. Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37–89.[0037:POA]2.0.CO;2
  23. 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. & Latella, I. (2017) A phylogenetic, biogeographic, and taxonomic study of all extant species of Anolis (Squamata; Iguanidae). Systematic Biology, 66 (5), 663–697.
  24. Rodríguez Schettino, L. (1999) Systematic accounts of the species. In: Rodríguez Schettino, L. (Ed.), The Iguanid Lizards of Cuba. The University Press of Florida, Gainesville, Florida, pp. 104–380.
  25. Rodríguez Schettino, L., Losos, J.B., Hertz, P.E., de Queiroz, K., Chamizo, A.R., Leal, M. & Rivalta González, V. (2010) The anoles of Soroa: aspects of their ecological relationships. Breviora, 520, 1–22. [The supplementary material contains the previously unpublished manuscript Rand, A.S. & Williams, E.E. The anoles of Soroa.]
  26. Rolfe, S., Pieper, S., Porto, A., Diamond, K., Winchester, J., Shan, S., Kirveslahti, H., Boyer, D., Summers, A. & Maga, A.M. (2021) SlicerMorph: An open and extensible platform to retrieve, visualize and analyze 3D morphology. Methods in Ecology and Evolution, 12 (10), 1816–1825.
  27. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
  28. Ruibal, R. (1964) An annotated checklist and key to the anoline lizards of Cuba. Bulletin of the Museum of Comparative Zoology, Harvard University, 130 (8), 473–520. []
  29. Sabaj, M.H. (2020) Codes for natural history collections in ichthyology and herpetology. Copeia, 108 (3), 593–669.
  30. Sanger, T.J., Sherratt, E., McGlothlin, J.W., Brodie, E.D., Losos, J.B. & Abzhanov, A. (2013) Convergent evolution of sexual dimorphism in skull shape using distinct developmental strategies. Evolution, 67(8), 2180–2193.
  31. Savage, J.M. & Talbot, J.J. (1978) The giant anoline lizards of Costa Rica and western Panama. Copeia, 1978, 480–492.
  32. Schaad, E.W. & Poe, S. (2010) Patterns of ecomorphological convergence among mainland and island Anolis lizards, Biological Journal of the Linnean Society, 101 (4), 852–859.
  33. Underwood, L.M. (1905) A summary of Charles Wright’s explorations in Cuba. Bulletin of the Torrey Botanical Club, 32 (6), 291–300.
  34. Williams, E.E. (1969) The ecology of colonization as seen in the zoogeography of anoline lizards on small islands. The Quarterly Review of Biology, 44 (4), 345–389.
  35. Williams, E.E. (1995) IV. The morphological characters used in the ANOLEKEY described, defined, and illustrated. In: Williams, E.E., Rand, H., Rand, A.S. & O’Hara, R.J. (1995) A computer approach to the comparison and identification of species in difficult taxonomic groups. Breviora, 502, pp. 15–37. []
  36. Williams, E.E. & Rand, A.S. (1969) Anolis insolitus, a new dwarf anole of zoogeographic importance from the mountains of the Dominican Republic. Breviora, 326, 1–21. []