Skip to main content Skip to main navigation menu Skip to site footer
Published: 2021-10-28

A new subspecies in the Turdus nudigenis complex (Aves: Turdidae) from the Cauca River valley of Colombia

Laboratorio de Biología Evolutiva de Vertebrados, Departamento de Ciencias Biológicas, Universidad de los Andes, Bogotá, Colombia.
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia.
Laboratorio de Biología Evolutiva de Vertebrados, Departamento de Ciencias Biológicas, Universidad de los Andes, Bogotá, Colombia. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia.
Facultad de Ciencias y Biotecnología, Universidad CES, Medellín, Colombia.
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia.
Aves Cauca canyon dry forest geographic variation incomplete lineage sorting paraphyly species limits


The taxonomy of the T. nudigenis complex has historically been challenging due to the uniformity in plumage among taxa, poorly known vocal variation, and allopatric distributions. Recent phylogenetic analyses have contributed to understanding relationships within the genus; however, much remains to be learned about the extent of phenotypic and genetic differentiation within taxa and its implications in species limits. Here, we analyze the taxonomic status of an enigmatic member of the T. nudigenis complex restricted to the Cauca River valley of Colombia. Our phylogenetic analyses suggest that the Cauca valley population is genetically distinctive from other members of the complex, although its sister relationships with respect to T. grayi and T. nudigenis could not be fully resolved due to paraphyly in the group. Based on similarity of plumage, calls, and biogeographic proximity we tentatively treat this taxon as a new subspecies of T. grayi. Populations from the northern Caribbean T. grayi casius/incomptus and the new taxon inhabit similar environmental conditions, being apparently isolated by the humid Nechí lowlands. A formal analysis of vocalizations coupled with a more robust genomic dataset are needed to resolve the systematic affinities and taxonomic status of several members of the T. nudigenis complex.



  1. Araya-Salas, M. & Smith-Vidaurre, G. (2017) warbleR: an r package to streamline analysis of animal acoustic signals. Methods in Ecology and Evolution, 8, 184–191.
    Arbeláez-Cortés, E. (2020) Defining the phylogeographic relationship between cis- and trans-Andean populations of Dendrocincla fuliginosa and Xenops minutus in Colombia. Revista Mexicana de Biodiversidad, 91, e912984.
    Avendaño, J.E., Arbeláez-Cortés, E. & Cadena, C.D. (2017) On the importance of geographic and taxonomic sampling in phylogeography: A reevaluation of diversification and species limits in a Neotropical thrush (Aves, Turdidae). Molecular Phylogenetics and Evolution, 111, 87–97.
    Avendaño, J.E., Cortés-Herrera, J.O., Briceño-L., E.R. & Rincón-Guarín, D.A. (2013) Crossing or bypassing the Andes: A commentary on recent range extensions of cis-Andean birds to the West of the Andes of Colombia. Orinoquia, 17, 207–214.
    Ayerbe-Quiñones, F. (2018) Guía Ilustrada de la Avifauna Colombiana. Wildlife Conservation Society, Bogotá, D.C., 440 pp.
    Bacon, C.D., Molnar, P., Antonelli, A., Crawford, A.J., Montes, C. & Vallejo-Pareja, M.C. (2016) Quaternary glaciation and the Great American Biotic Interchange. Geology, 44, 375–378.
    Bangs, O. (1898) On some birds from Santa Marta, Colombia. Proceedings of the Biological Society of Washington, 12, 131–144.
    Berlepsch, H.V. & Taczanowski, L. (1883) Liste des oiseaux recueillis par MM. Stolzmann et Siemiradzki dans l’Ecuadeur occidental. Proceedings of the Scientific Meetings of the Zoological Society of London, 1883, 536–577.
    Berrío, J.C., Hooghiemstra, H., Marchant, R. & Rangel, O. (2002) Late glacial and Holocene history of the dry forest area in the south Colombian Cauca Valley. Journal of Quaternary Science, 17, 667–682.
    Bonaparte, C.L.J.L. (1838) Birds collected by Colonel Velasquez de Leon during a fortnight’s scientific tour in Guatemala. Proceedings of the Zoological Society of London, 59, 114–119.
    Bonaparte, C.L.J.L. (1855) Catalogue des genres et sous-genres d’oiseaux contenues dans le museum Britannique. Compte Rendus Hebdomadaire des Séances de l’Académie des Sciences, 41, 649–661.
    Brinkhuizen, D.M. & Seimola, T. (2014) First record of Pacific Parrotlet Forpus coelestis in Colombia. Conservación Colombiana, 21, 30–32.
    Broennimann, O., Fitzpatrick, M.C., Pearman, P.B., Petitpierre, B., Pellissier, L., Yoccoz, N.G., Thuiller, W., Fortin, M.J., Randin, C., Zimmermann, N.E., Graham, C.H. & Guisan, A. (2012) Measuring ecological niche overlap from occurrence and spatial environmental data. Global Ecology and Biogeography, 21, 481–497.
    Broennimann, O., Petitpierre, B., Randin, C., Engler, R., Di Cola, V., Breiner, F., D`Amen, M., Pellissier, L., Pottier, J., Pio, D., Garcia Mateo, R., Hordijk, W., Dubuis, A., Scherrer, D., Salamin, N. & Guisan, A. (2015) ecospat: Spatial Ecology Miscellaneous Methods. R Package. Version 1. Available from: (accessed 3 April 2020)
    Cadena, C.D., Klicka, J. & Ricklefs, R.E. (2007) Evolutionary differentiation in the Neotropical montane region: Molecular phylogenetics and phylogeography of Buarremon brush-finches (Aves, Emberizidae). Molecular Phylogenetics and Evolution, 44, 993–1016.
    Catchpole, C.K. & Slater, P.J.B. (2008) Bird Song: Biological Themes and Variations. Second Edition. Cambridge University Press, Cambridge, 335 pp.
    Cerqueira, P.V., Santos, M.P.D. & Aleixo, A. (2016) Phylogeography, inter-specific limits and diversification of Turdus ignobilis (Aves: Turdidae). Molecular Phylogenetics and Evolution, 97, 177–186.
    Chapman, F.M. (1917) The distribution of bird-life in Colombia: a contribution to a biological survey of South America. Bulletin of the American Museum of Natural History, 36, 1–728.
    Charif, R.A., Waack, A.M. & Strickman, L.M. (2014) Raven Pro: Interactive sound analysis software. Version 1.5. The Cornell Lab of Ornithology. Ithaca, New York.
    Chesser, R.T., Burns, K.J., Cicero, C., Dunn, J.L., Kratter, A.W., Lovette, I.J., Rasmussen, P.C., Remsen, J.V., Stotz, D.F. & Winker, K. (2019) Sixtieth supplement to the American Ornithological Society’s check-list of North American birds. The Auk, 136, 1–23.
    Collar, N.J. (2015) Family Turdidae (thrushes). In: del Hoyo, J., Elliot, A. & Christie, D.A. (Eds.), Handbook of Birds of the World, Cuckoo-Shrikes to Thrushes. Vol. 10. Lynx Edicions, Barcelona, pp. 514–807.
    Dabelsteen, T. & Pedersen, S.B. (1990) Song and information about aggressive responses of blackbirds, Turdus merula: evidence from interactive playback experiments with territory owners. Animal Behaviour, 40, 1158–1168.
    Donegan, T.M., Salaman, P. & Caro, D. (2009) Revision of the status of various bird species occurring or reported in Colombia. Conservación Colombiana, 8, 80–86.
    Fick, S.E. & Hijmans, R.J. (2017) WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37, 4302–4315.
    Funk, D.J. & Omland, K.E. (2003) Species-level paraphyly and polyphyly: Frequency, causes, and consequences, with insights from animal mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics, 34, 397–423.
    Haffer, J. (1967) Zoogeographical notes on the” nonforest” lowland bird faunas of northwestern South America. El Hornero, 10, 315–333.
    van der Hammen, T. (1974) The Pleistocene changes of vegetation and climate in tropical South America. Journal of Biogeography, 1, 3–26.
    Hammer, Ø., Harper, D. & Ryan, P. (2001) Past: Paleontological statistics software package for education and data analysis. Paleontologia Electronica, 4, 1–9.
    Hardy, J.W. & Parker III, T.A. (1997) The nature and probable function of vocal copying in Lawrence’s Thrush, Turdus lawrencii. Ornithological Monographs, 48, 307–320.
    Hernández-Camacho, J.I., Hurtado, A., Ortiz, R. & Walschburger, T. (1992) Unidades biogeográficas de Colombia. In: Halffter, G. (Ed.), La Diversidad Biológica de Iberoamérica. Special Volume. Acta Zoológica Mexicana, Xalapa, pp. 105–152.
    Hilty, S.L. & Brown, W.L. (1986) A Guide to the Birds of Colombia. Princeton University Press, Princeton, New York, 996 pp.
    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.
    Klicka, J., Voelker, G. & Spellman, G.M. (2005) A molecular phylogenetic analysis of the “true thrushes” (Aves: Turdinae). Molecular Phylogenetics and Evolution, 34, 486–500.
    Lafresnaye, M.F. (1848) Description de quelques oiseaux nouveaux de Caracas (province de Venezuela) et de Bogota. Revue Zoologique, 11, 2–12.
    Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.
    Lara, C.E., Cuervo, A.M., Valderrama, S. V., Calderón-F, D. & Cadena, C.D. (2012) A new species of wren (Troglodytidae: Thryophilus) from the dry Cauca River Canyon, northwestern Colombia. Auk, 129, 537–550.
    Lawrence, G.N. (1862) Catalogue of a collection of birds, made in New Granada, by James McLeannan, Esq., of New York, with notes and descriptions of new species. Annals of the Lycaeum of Natural History of New York, 7, 461–479.
    Lawrence, G.N. (1869) Characters of some new South American birds, with notes on other rare or little known species. Annals of the Lycaeum of Natural History of New York, 9, 265–275.
    Leigh, J.W. & Bryant, D. (2015) PopART: Full-feature software for haplotype network construction. Methods in Ecology and Evolution, 6, 1110–1116.
    Lichtenstein, H. (1823) Verzeichnis der Doublettin des Zoologische Museum der Königl. Universitët zu Berlin: Beschreibung vieler bisher unbekannter Säugethieren, Vógeln, Anfibien und Fischen. Commission bei T. Trautwein, Berlin, 118 pp.
    Linnaeus, C. von (1758) Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. 10th ed. Holmiae, Laurentii Salvii, 823 pp.
    Marchant, R., Boom, A., Behling, H., Hooghiemstra, H., Melief, B., Geel, B. Van, Hammen, T. Van der & Wille, M. (2004) Colombian vegetation at the Last Glacial Maximum: a comparison of model- and pollen-based biome reconstructions. Journal of Quaternary Science, 19, 721–732.
    Mayr, E. (1963) Animal Species and Evolution. Harvard University Press, Cambridge, Massachusetts, 797 pp.
    McMullan, M., Donegan, T.M., Pantoja-Peña, G., Tuncer-Navarro, T., Bartels, A. & Ellery, T. (2018) Field Guide to the Birds of Colombia. Rey Naranajo Editores, Bogotá, D.C., 397 pp.
    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE). IEEE, New Orleans, Louisiana, pp. 1–8.
    Miller, M.J., Bermingham, E., Klicka, J., Escalante, P., do Amaral, F.S.R., Weir, J.T. & Winker, K. (2008) Out of Amazonia again and again: episodic crossing of the Andes promotes diversification in a lowland forest flycatcher. Proceedings of the Royal Society B: Biological Sciences, 275, 1133–1142.
    Miller, M.J., Lelevier, M.J., Bermingham, E., Klicka, J.T., Escalante, P. & Winker, K. (2011) Phylogeography of the Rufous-tailed Hummingbird (Amazilia tzacatl). The Condor, 113, 806–816.
    Miller, W.D.W. & Griscom, L. (1925) Notes on Central American birds, with descriptions of new forms. American Museum Novitates, 183, 1–14.
    Nylander, J.A.A., Olsson, U., Alström, P. & Sanmartin, I. (2008) Accounting for phylogenetic uncertainty in biogeography: A Bayesian approach to dispersal-vicariance analysis of the thrushes (Aves: Turdus). Systematic Biology, 57, 257–268.
    Palacios, C., García-R, S., Parra, J.L., Cuervo, A.M., Stiles, F.G., McCormack, J.E. & Cadena, C.D. (2019) Shallow genetic divergence and distinct phenotypic differences between two Andean hummingbirds: Speciation with gene flow? The Auk: Ornithological Advances, 136, ukz046.
    O’Neill, J.P., Lane, D.F. & Naka, L.N. (2011) A cryptic new species of thrush (Turdidae: Turdus) from western Amazonia. The Condor, 113, 869–880.
    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, 933–938.[0933:TEOTWA]2.0.CO;2
    Phillips, S.J., Dudík, M. & Schapire, R.E. (2018) Maxent software for modeling species niches and distributions. Version 3.4.1. Available from: (accessed 5 January 2020)
    Pinho, C. & Hey, J. (2010) Divergence with gene flow: Models and data. Annual Review of Ecology, Evolution, and Systematics, 41, 215–230.
    Price, T. (2008) Speciation in Birds. Roberts and Company, Greenwood Village, Colorado, 480 pp.
    R Development Core Team (2016) R: A language and environment for statistical computing. Available from: (accessed 1 January 2016)
    Rambaut, A., Drummond, A.J. & Suchard, M. (2003) Tracer. Version 1.6. MCMC Trace Analysis Package. Institute of Evolutionary Biology, Department of Computer Science. Available from: (accessed 1 August 2014)
    Remsen, J. V., Areta, J.I., Bonaccorso, E., Claramunt, S., Jaramillo, A., Pacheco, J.F., Robbins, M.B., Stiles, F.G., Stotz, D.F. & Zimmer, K.J. (2021) A Classification of the Bird Species of South America. American Ornithologists’ Union. Available from: (accessed 20 February 2021)
    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
    Schultz, E.D., Pérez-Emán, J., Aleixo, A., Miyaki, C.Y., Brumfield, R.T., Cracraft, J. & Ribas, C.C. (2019) Diversification history in the Dendrocincla fuliginosa complex (Aves: Dendrocolaptidae): Insights from broad geographic sampling. Molecular Phylogenetics and Evolution, 140, 106581.
    Smith, B.T., McCormack, J.E., Cuervo, A.M., Hickerson, M.J., Aleixo, A., Cadena, C.D., Pérez-Emán, J., Burney, C.W., Xie, X., Harvey, M.G., Faircloth, B.C., Glenn, T.C., Derryberry, E.P., Prejean, J., Fields, S. & Brumfield, R.T. (2014) The drivers of tropical speciation. Nature, 515, 406–409.
    Smithe, F.B. (1975) Naturalists’ color guide. American Museum of Natural History Press, New York, NY, USA, 25 pp.
    Smithe, F.B. (1981) Naturalist’s Color Guide, Part III. American Museum of Natural History Press, New York, NY, USA, 37 pp.
    Snow, D. (1985) Systematics of the Turdus fumigatus/hauxwelli group of thrushes. Bulletin of the British Ornithologists’ Club, 105, 30–37.
    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
    Stiles, F.G. & Avendaño, J.E. (2019) Distribution and status of Turdus thrushes in white-sand areas of eastern Colombia, with a new subspecies of T. leucomelas. Zootaxa, 4567, 161–175.
    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolutione, 30, 2725–2729.
    Todd, W.E.C. (1931) Critical notes on Neotropical thrushes. Proceedings of the Biological Society of Washington, 44, 47–54.
    Vargas-Castro, L.E., Sánchez, N.V. & Barrantes, G. (2012) Repertoire size and syllable sharing in the song of the Clay-colored Thrush (Turdus grayi). The Wilson Journal of Ornithology, 124, 446–453.
    Vargas-Castro, L.E., Sánchez, N.V. & Barrantes, G. (2015) Song plasticity over time and vocal learning in Clay-colored Thrushes. Animal Cognition, 18, 1113–1123.
    Vieillot, L.J.P. (1818) Ornithologie. In: Nouveau Dictionaire d´Histoire Naturelle, appliquée aux arts, a l’Agriculture, à l´Économie rurale et domestique, à la Médicine, etc. par une Société de naturalistes et d´agriculteurs. Deterville, Paris. [unknown pagination]
    Voelker, G., Rohwer, S., Bowie, R.C.K. & Outlaw, D.C. (2007) Molecular systematics of a speciose, cosmopolitan songbird genus: Defining the limits of, and relationships among, the Turdus thrushes. Molecular Phylogenetics and Evolution, 42, 422–434.
    Werneck, F.P., Costa, G.C., Colli, G.R., Prado, D.E. & Sites, J.W. (2011) Revisiting the historical distribution of Seasonally Dry Tropical Forests: New insights based on palaeodistribution modelling and palynological evidence. Global Ecology and Biogeography, 20, 272–288.


How to Cite

AVENDAÑO, J. E. ., CUERVO, A. M. ., MONTOYA, P. ., PULGARÍN-R, P. C. ., & STILES, F. G. . (2021). A new subspecies in the <em>Turdus</em> <em>nudigenis</em> complex (Aves: Turdidae) from the Cauca River valley of Colombia. Zootaxa, 5060(1), 105–123.