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Type: Article
Published: 2021-10-21
Page range: 151-180
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A taxonomic assessment of Chlorospingus flavopectus phaeocephalus and Chlorospingus semifuscus (Passeriformes: Passerellidae), including the description of a new subspecies

Instituto Nacional de la Biodiversidad del Ecuador (INABIO), calle Rumipamba 341 y Av. de Los Shyris, Casilla 17-07-8976, Quito, Ecuador.
Laboratorio de Biología Evolutiva de Vertebrados, Departamento de Ciencias Biológicas, Universidad de los Andes, Bogotá, Colombia.
Instituto Nacional de la Biodiversidad del Ecuador (INABIO), calle Rumipamba 341 y Av. de Los Shyris, Casilla 17-07-8976, Quito, Ecuador. Universidad Nacional de Colombia, Facultad de Ciencias, Bogotá D.C., Colombia; Grupo de Investigación Evolución y Ecología de Fauna Neotropical. Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales COCIBA, Laboratorio de Zoología Terrestre y Museo de Zoología, Campus Cumbayá, Quito 170901, Ecuador.
Université Paris Est Créteil, LAB’URBA, F-94010 Créteil, France. Centre d'Ecologie et des Sciences de la Conservation (CESCO UMR 7204), MNHN, CNRS, Sorbonne Université, CP135, 57 rue Cuvier, 75005 Paris, France.
Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS, Sorbonne Université, EPHE, 57 rue Cuvier, CP51, 75005 Paris, France.
American Museum of Natural History, Department of Ornithology, Central Park West at 79th Street, New York, New York 10024-5192. Office: 212-769-5788
Universidad San Francisco de Quito, Laboratorio de Biología Evolutiva, Colegio de Ciencias Biológicas y Ambientales, Ecuador.
Aves bird systematics Andes Ecuador Tumbes-Chocó speciation hybridization speciation barriers population barriers Passerellidae

Abstract

Chlorospingus flavopectus, a widely distributed member of the New World sparrows and finches (Passerellidae), is among the most variable and complex Neotropical bird species. With up to 28 subspecies that inhabit montane forest from Mexico south to Argentina, it presents a recurring leapfrog pattern, with many genetically differentiated lineages, but even more morphologically distinguishable taxa. Chlorospingus flavopectus phaeocephalus is distributed along the eastern Andean slope (from southern Colombia to northern Peru) and in localized patches along the central and southwestern slopes of the Ecuadorian Andes, in the forest remnants of Chimborazo, El Oro, and adjacent Loja provinces. This allopatric distribution was based on plumage similarities, but no genetic or vocal analysis has tested if these populations share a common ancestry. Here, we compared data for populations of C. f. phaeocephalus, C. semifuscus semifuscus from northwest Ecuador, and C. s. livingstoni from west Colombia, and analyzed them in the context of other Chlorospingus. Our phylogenetic analyses revealed that the putative west Ecuadorian populations of C. f. phaeocephalus are nested within C. semifuscus. They also present diagnostic characters when compared to other populations of C. semifuscus and C. f. phaeocephalus. Based on these results, we recognize the population from El Oro and adjacent Loja Provinces as a new subspecies of C. semifuscus, and the population from Chimborazo Province as a morphological variation of C. s. semifuscus.

 

References

  1. Audacity Team (2019) Audacity(R): Free Audio Editor and Recorder. Windows 10 Pro. Version 2.3.3. Available from: https://audacityteam.org/ (accessed 20 January 2020)
    Avendaño, J.E., Stiles, F.G. & Cadena, C.D. (2013) A new subspecies of Common Bush-Tanager (Chlorospingus flavopectus, Emberizidae) from the east slope of the Andes of Colombia. Ornitología Colombiana, 13, 45–58.
    Ayerbe-Quiñones, F. (2018) Guía Ilustrada de la Avifauna Colombiana. Puntoaparte, Bogotá, 440 pp.
    Barker, F.K., Burns, K.J., Klicka, J., Lanyon, S.M. & Lovette, I.J. (2013) Going to extremes: contrasting rates of diversification in a recent radiation of New World passerine birds. Systematic Biology, 62 (2), 298–320. https://doi.org/10.1093/sysbio/sys094
    Barreira, A.S. & García, N.C. (2019) Visual and acoustic communication in Neotropical birds: diversity and evolution of signals. In: Reboreda, J.C., Fiorini, V.D. & Tuero, D.T. (Eds.), Behavioral Ecology of Neotropical Birds. Springer, pp. 155–183. https://doi.org/10.1007/978-3-030-14280-3
    Berlepsch, H. & Taczanowski, L. (1884) Deuxième liste des oiseaux recueillis dans l’Ecuadeur occidental par MM. Stolzmann et Siemiradski. Proceedings of the Zoological Society of London, 52 (3), 281–313.
    Best, B.F., Clarke, C.T., Checker, M., Broom, A.L., Thewlis, R.M., Duckworth, W. & McNab, A. (1993) Distributional records, natural history notes, and conservation of some poorly known birds from southwestern Ecuador and northwestern Peru. Bulletin British Ornithologists’ Club, 113 (2), 108–119.
    BirdLife International (2020) Endemic Bird Areas factsheet: Costa Rica and Panama highlands. Available from: http://www.birdlife.org (accessed 14 October 2020)
    Bohórquez, C.I. & Stiles, F.G. (2002) The paradoxical social system of the Dusky Bush-Tanager (Chlorospingus semifuscus): Lekking in a nine-primaried oscine? Journal of Field Ornithology, 73 (3), 281–291. https://doi.org/10.1648/0273-8570-73.3.281
    Bonaccorso, E., Navarro-Sigüenza, A.G., Sánchez-González, L.A., Peterson, A.T. & García-Moreno, J. (2008) Genetic differentiation of the Chlorospingus ophthalmicus complex in Mexico and Central America. Journal of Avian Biology, 39 (3), 311–321. https://doi.org/10.1111/j.0908-8857.2008.04233.x
    Bond, J. & Meyer de Schauensee, R. (1940) On some birds from southern Colombia. Proceedings of the Academy of Natural Sciences of Philadelphia, 92, 153–169.
    Cabanis, J.L. (1851) Verzeichniss der ornithologischen Sammlung des Oberamtmann Ferdinand Heine auf Gut St. Burchard vor Halberstatdt. Museum Heineanum, 1, 1–234.
    Cadena, C.D., Córdoba-Córdoba, S., Londoño, G.A., Calderón, F.D., Martin, T.E. & Baptiste, M.P. (2007) Nesting and singing behavior of Common Bush-Tanagers (Chlorospingus ophthalmicus) in South America. Ornitología Colombiana, 5, 54–63.
    Cadena, C.D., Cheviron, Z.A. & Funk, W.C. (2011) Testing the molecular and evolutionary causes of a ‘leapfrog’ pattern of geographical variation in coloration. Journal of Evolutionary Biology, 24 (2), 402–414.
    Cadena, C.D., Pérez-Emán, J.L., Cuervo, A.M., Céspedes, L.N. Epperly, K.L. & Klicka, J.T. (2018) Extreme genetic structure and dynamic range evolution in a montane passerine bird: implications for tropical diversification. Biological Journal of the Linnean Society of London, 126 (3), 487–506.
    Cárdenas M.L., Gosling W.D., Pennington R.T., Poole, I., Sherlock, S.C. & Mothes, P. (2014) Forest of the tropical eastern Andean flank during the middle Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology, 393, 76–89.
    Caro, L.M., Caycedo-Rosales, P.C., Bowie, R.C.K., Slabbekoorn, H. & Cadena, C.D. (2013) Ecological speciation along an elevational gradient in a tropical passerine bird? Journal of Evolutionary Biology, 26 (2), 357–374. https://doi.org/10.1111/jeb.12055
    Carson, R.J. & Spicer, G.S. (2003) A phylogenetic analysis of the emberizid sparrows based on three mitochondrial genes. Molecular Phylogenetics and Evolution, 29 (1), 43–57. https://doi.org/10.1016/s1055-7903(03)00110-6
    Center for Conservation Bioacoustics (2014) Raven Pro: Interactive Sound Analysis Software. Version 1.5 Beta. Computer Software. The Cornell Lab of Ornithology, Ithaca, New York. Available from: http://ravensoundsoftware.com (accessed 20 October 2019)
    Chapman, F.M. (1925) Descriptions of one new genus and of species of birds from Peru and Ecuador. American Museum Novitates, 205, 1–11.
    Chapman, F.M. (1926) The distribution of birdlife in Ecuador. Bulletin American Museum of Natural History, 55, 1–784.
    Chavarría-Pizarro, T., Gutiérrez-Espeleta, G., Fuchs, E.J. & Barrantes, G. (2010) Genetic and morphological variation of the Sooty-capped Bush Tanager (Chlorospingus pileatus), a highland endemic species from Costa Rica and western Panama. Wilson Journal of Ornithology, 122 (2), 279–287. https://doi.org/10.1676/09-111.1
    Chavez-Paz, L.F., Rosero-Chates, E.B. & Muñoz-García, J. (2017) eBird Checklist: ebird: An online database of bird distribution and abundance. Web Application. eBird, Ithaca, New York. Available from: https://ebird.org/checklist/S38825296 (accessed 30 September 2019)
    Chesser, R.T., Burns, K.J., Cicero, C., Dunn, J.L., Kratter, A.W., Lovette, I.J., Rasmussen, P.C., Remsen, J.V. Jr., Rising, J.D., Stotz, D.F. & Winker, K. (2017) Fifty-eighth supplement to the American Ornithological Society's Check-list of North American Birds. Auk, 134 (3), 751–773. https://doi.org/10.1642/AUK-17-72.1
    Chitnis, S.S., Rajan, S. & Krishnan, A. (2020) Sympatric wren-warblers partition acoustic signal space and song perch height. Behavioral Ecology, 31 (2), 559–567. https://doi.org/10.1093/beheco/arz216
    Cicero, C., Mason, N.A., Benedict, L. & Rising, J.D. (2020) Behavioural, morphological, and ecological trait evolution in two clades of New World sparrows (Aimophila and Peucaea, Passerellidae). PeerJ, 8 (e9249), 1–30. [published online] https://doi.org/10.7717/peerj.9249
    Clements, J.F., Schulenberg, T.S., Iliff, M.J., Billerman, S.M., Fredericks, T.A., Sullivan, B.L. & Wood C.L. (2019) The eBird/Clements Checklist of Birds of the World. Version 2019. Available from: https://www.birds.cornell.edu/clementschecklist/download/ (accessed 1 September 2019)
    Cuesta, F., Peralvo, M., Baquero, F., Bustamante, M., Merino, A., Muriel, P., Freile, J. & Torres, O. (2015) Áreas Prioritarias para la Conservación del Ecuador Continental. Ministerio de Ambiente, CONDESAN, Pontificia Universidad Católica del Ecuador, GIZ, Quito, 111 pp.
    DaCosta, J.M., Spellman, G.M., Escalante, P. & Klicka, J. (2009) A molecular systematic revision of two historically problematic songbird clades: Aimophila and Pipilo. Journal of Avian Biology, 40 (2), 206–216. https://doi.org/10.1111/j.1600-048X.2009.04514.x
    Dodson, C.H. & Gentry, A.H. (1991) Biological extinction in western Ecuador. Annals of the Missouri Botanical Garden, 78, 273–295.
    Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29 (8), 1969–1973. https://doi.org/10.1093/molbev/mss0755
    Du Bus de Gisignies, M. (1855) Note sur quelques espéces inédites d'oiseaux. Bulletins de L'Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 22 (1), 150–157.
    Felix, R. & Coral-Jaramillo, B. (2017) eBird Checklist: https://ebird.org/checklist/S41001626. eBird: An online database of bird distribution and abundance [web application]. eBird. Available from: http://www.ebird.org (accessed 30 September 2019)
    Ferrer-Paris, J.R., Zager, I., Keith, D.A., Oliveira‐Miranda, M.A., Rodríguez, J.P., Josse, C., González-Gil, M., Miller, R.M., Zambrana-Torrelio, C. & Barrow, E. (2019) An ecosystem risk assessment of temperate and tropical forests of the Americas with an outlook on future conservation strategies. Conservation Letters, 12 (2), e12623.
    Fjeldså, J. & Krabbe, N. (1990) Birds of the High Andes. Zoological Museum, University of Copenhagen & Svendborg, Apollo Books, Copenhagen, 876 pp.
    Freile, J.F. & Santander, T. (2005) Áreas Importantes para la Conservación de las Aves en Ecuador. Aves & Conservación (Corporación Ornitológica del Ecuador), BirdLife International, Conservación Internacional y Ministerio del Ambiente de Ecuador, Quito, 236 pp.
    Freile, J.F. & Restall, R. (2018) Birds of Ecuador. Bloomsbury Publishing, London, 656 pp.
    Freile, J.F., Brinkhuizen, D.M., Greenfield, P.J., Lysinger, M., Navarrete, L., Nilsson, J., Olmstead, S., Ridgely, R.S., Sánchez-Nivicela, M., Solano-Ugalde, A., Athanas, N., Ahlman, R. & Boyla, K.A. (2020) Lista de las Aves del Ecuador/Checklist of the Birds of Ecuador. Comité Ecuatoriano de Registros Ornitológicos. Available from: https://ceroecuador.wordpress.com/ (accessed 12 September 2020)
    García-Moreno, J., Ohlson, J. & Fjeldså, J. (2001) MtDNA sequences support monophyly of Hemispingus tanagers. Molecular Phylogenetics and Evolution, 21 (3), 424–435. https://doi.org/10.1006/mpev.2001.1027
    García-Moreno, J., Navarro-Sigüenza, A.G., Peterson, A.T. & Sánchez-González, L. A. (2004) Genetic variation coincides with geographic structure in the Common Bush-tanager (Chlorospingus ophthalmicus) complex from Mexico. Molecular Phylogenetic and Evolution, 33 (1), 186–196. https://doi.org/10.1016/j.ympev.2004.05.007
    Garzón-Santomaro, C., Pozo-Zamora, G., Sornoza-Molina, F. & Mena-Valenzuela, P. (2019) Aves del Páramo al Manglar. In: MECN-INB-GADPEO, 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 N° 11. GADPEO-INABIO, Quito, pp. 121–218.
    GBIF.org (2018a) GBIF Occurrence Download. 27 January 2018. Available from; https://doi.org/10.15468/dl.qqfgwk (accessed 11 February 2021)
    GBIF.org (2018b) GBIF Occurrence Download. 27 January 2018. Available from; https://doi.org/10.15468/dl.i0ewug (accessed 11 February 2021)
    GBIF.org (2018c) GBIF Occurrence Download. 27 January 2018. Available from: https://doi.org/10.15468/dl.p1x9pb (accessed 11 February 2021)
    GBIF.org (2018d) GBIF Occurrence Download. 27 January 2018. Available from; https://doi.org/10.15468/dl.zevcuq (accessed 11 February 2021)
    Gmelin, J.F. (1789) Savannarum. In: Linné, C.A., Systema naturae per regna tria naturae. Secundum Claffes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. AVES VI. PASSERES. 112 Fringilla. Editio Decima Tertia, Aucta, Reformata. Tomus Primus. PARS II. Apud J. B. Mollieré, Lugduni, pp. 921.
    Gregory-Wodzicki, K.M. (2000) Uplift history of the Central and Northern Andes: a review. Geological Society of America Bulletin, 112 (7), 1091–1105.
    Griscom, L. (1924) Descriptions of new birds from Panama and Costa Rica. American Museum Novitates, 141, 1–12.
    Hammer, Ø., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4 (1), 9.
    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 National Academy of Sciences, 115 (31), 7985–7990. https://doi.org/10.1073/pnas.1803908115
    Hellmayr, C.E. (1921) Beschreibt 12 neue Formen aus dem neotropischen Gebiet. In: Anzeiger der Ornithologischen Gesellschaft in Bayern, 4, 25–32.
    Hellmayr, C.E. (1936) Catalogue of birds of the Americas and the adjacent islands in Field Museum of Natural History. Tersinidae-Thraupidae. Field Museum of Natural History, Zoological Series, 18, 1–458.
    Hermes, C., Döpper, A., Schaefer, H.M. & Segelbacher, G. (2016) Effects of forest fragmentation on the morphological and genetic structure of a dispersal-limited, endangered bird species. Nature Conservation, 16, 39–59. https://doi.org/10.3897/natureconservation.16.10905
    Hermes, C., Jansen, J. & Schaefer, H.M. (2017) Habitat requirements and population estimate of the endangered Ecuadorian Tapaculo Scytalopus robbinsi. Bird Conservation International, 28 (2), 1–17. https://doi.org/10.1017/S095927091600054X
    Herzog, S.K., Terrill, R.S., Jahn, A.E., Remsen, J.V., Maillard-Z, O., García-Solíz, V.H., Macleod, R., Maccormick, A. & Vidoz, J.Q. (2016) Birds of Bolivia: Field Guide. Asociación Armonía, Santa Cruz de la Sierra, 489 pp.
    Hilty, S.L. & Brown, W.L. (1986) A Guide to the Birds of Colombia. Princeton University Press, Princeton, New Jersey, 836 pp.
    Hilty, S.L. (2003) Birds of Venezuela. Princeton University Press, Princeton, New Jersey, 878 pp.
    Hilty, S. (2020a) Common Bush-tanager (Chlorospingus flavopectus). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (Eds.), Handbook of the Birds of the World Alive. Lynx Editions, Barcelona. Available from: https://www.hbw.com/node/61812 (accessed 8 May 2020)
    Hilty, S. (2020b) Dusky Chlorospingus (Chlorospingus semifuscus), version 1.0. In: J. del Hoyo, Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (Eds.), Birds of the World. Cornell Lab of Ornithology, Ithaca. Available from: https://doi.org/10.2173/bow.dubtan1.01 (accessed 8 May 2020)
    Hooghiemstra, H. & Van der Hammen, T. (2004) Quaternary Ice-Age dynamics in the Colombian Andes: developing an understanding of our legacy. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 359 (1442), 173–181. https://doi.org/10.1098/rstb.2003.1420
    Hoorn, C., Wesselingh, F.P., Ter Steege, H., Bermudez, M.A., Mora, A., Sevink, J., Sanmartín, I., Sanchez-Meseguer, A., Anderson, L.C., Figueiredo J.P., Jaramillo, C., Riff, D., Negri, F.R., Hooghiemstra, H., Lundberg, J., Stadler, T, Särkinen, T. & Antonelli, A. (2010) Amazonia through time: Andean uplift, climate change, landscape evolution, and biodiversity. Science, 330 (6006), 927–931. https://doi.org/10.1126/science.1194585
    Isler, M.L. & Isler, P.R. (1999) The Tanagers: Natural History, Distribution, and Identification. Smithsonian Institution Press, Washington, D.C., 406 pp.
    Jobling, J.A. (2010) The Helm Dictionary of Scientific Bird Names from Aalge to Zusii. Helm, London, 432 pp.
    Johnson, N.K. & Brush, A.H. (1972) Analysis of polymorphism in the Sooty-capped Bush Tanager. Systematic Zoology, 21 (3), 245–262.
    Klicka, J., Burns, K. & Spellman, G.M. (2007) Defining a monophyletic Cardinalini: A molecular perspective. Molecular Phylogenetics and Evolution, 45 (3), 1014–1032. https://doi.org/10.1016/j.ympev.2007.07.006
    Klicka, J., Barker, F.K., Burns, K.J., Lanyon, S.M., Lovette, I.J., Chaves, J.A. & Bryson, R.W. Jr. (2014) A comprehensive multilocus assessment of sparrow (Aves: Passerellidae) relationships. Molecular Phylogenetics and Evolution, 77, 177–182.
    Krabbe, N., Skov, F., Fjeldså, J. & Petersen, I.K. (1998) Avian Diversity in the Ecuadorian Andes. DIVA Technical Report 4. Centre for Research on Cultural and Biological Diversity of Andean Rainforests, Rønde, 143 pp.
    Krabbe, N. (2008) Arid valleys as dispersal barriers to high-Andean forest birds in Ecuador. Cotinga, 29, 28–30.
    Lafresnaye, M.F. de (1840) Description de quelques nouvelles espèces d’oiseaux. Revue Zoologique par La Société Cuvierienne, 227–232.
    Lafresnaye, N.F.A.A. & D’Orbigny, A. (1838) Notice sur quelques oiseaux de Carthagène et de la partie du Mexique la plus voisine, rapportés par M. Ferdinand de Candé, officer de la marine royale. Revue Zoologique par La Societé Cuvierienne, 1838, 164–166.
    Lanfear, R., Calcott, B.S., Ho, Y. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyzes. Molecular Biology and Evolution, 29 (6), 1695–1701. https://doi.org/10.1093/molbev/mss020
    Lerner, H.R., Meyer, M., James, H.F., Hofreiter, M. & Fleischer, R.C. (2011) Multilocus resolution of phylogeny and timescale in the extant adaptive radiation of Hawaiian honeycreepers. Current Biology, 21 (21), 1838–1844. https://doi.org/10.1016/j.cub.2011.09.039
    MAE (Ministerio del Ambiente del Ecuador) (2012) Sistema de Clasificación de los Ecosistemas del Ecuador Continental. Subsecretaría de Patrimonio Natural, Quito, 232 pp.
    MAE (Ministerio del Ambiente del Ecuador) (2015) Estadísticas de Patrimonio Natural. Datos de Bosques, Ecosistemas, Especies, Carbono y Deforestación del Ecuador Continental. MAE, Quito, 19 pp.
    Maldonado-Sánchez, D., Gutiérrez-Rodríguez, C. & Ornelas, J.F. (2016) Genetic divergence in the Common Bush-tanager Chlorospingus ophthalmicus (Aves: Emberizidae) throughout Mexican cloud forests: The role of geography, ecology and Pleistocene climatic fluctuations. Molecular Phylogenetics and Evolution, 99, 76–88. https://doi.org/10.1016/j.ympev.2016.03.014
    McCarthy, E.M. (2006) Handbook of Avian Hybrids of the World. Oxford University Press, New York, 583 pp.
    Moore, J.V., Krabbe, N. & Jahn, O. (2013) Bird Sounds of Ecuador. A Comprehensive Collection. DVD. John V. Moore Nature Recordings, San Jose. [DVD-Rom]
    NCI [Naturaleza y Cultura Internacional] (2020) Se fortalece la conservación de zonas de alta diversidad biológica en El Oro. Available from: http://www.naturalezaycultura.org/spanish/htm/news/2020-01-Convenio.htm (accessed 8 August 2020)
    Nelson, E.W. (1912) Descriptions of new genera, species and subspecies of birds from Panama, Colombia and Ecuador. Smithsonian Miscellaneous Collections, 3, 1–25.
    Olson, S.L. (1983) Geographic variation in Chlorospingus ophthalmicus in Colombia and Venezuela (Aves: Thraupidae). Proceedings of the Biological Society of Washington, 96, 103–109.
    Olson, S.L. (1993) Intergradation between the bush-tanagers Chlorospingus punctulatus and C. ophthalmicus in western Panama (Aves: Thraupidae). Auk, 110 (1), 148–150.
    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. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
    O’Neill, J.P. & Parker III, T.A. (1981) New subspecies of Pipreola riefferii and Chlorospingus ophthalmicus from Peru. Bulletin of the British Ornithologists’ Club, 101, 294–299.
    Pons, J., Barraclough, T.G., Gomez-Zurita, J., Cardoso, A., Duran, D.P., Hazell, S., Kamoun, S., Sumlin, W.D. & Vogler, A.P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609.
    QGIS Development Team (2020) QGIS Geographic Information System. Open Source Geospatial Foundation Project. Available from: http://qgis.osgeo.org (accessed 12 November 2019)
    Rambaut, A. & Drummond, A.J. (2007) TreeAnnotator. Program and documentation distributed by the author. Available from: http://beast.bio.ed.ac.uk/TreeAnnotator (accessed 10 December 2019)
    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
    Reid, N.M. & Carstens, B.C. (2012) Phylogenetic estimation error can decrease the accuracy of species delimitation: a Bayesian implementation of the general mixed Yule-coalescent model. BMC Evolutionary Biology, 12 (1), 196.
    Remsen, J.V. (1984) High incidence of “leapfrog” pattern of geographic variation in Andean birds: implications for the speciation process. Science, 224 (4645), 171–173.
    Remsen, J. V. Jr., Areta, J.I., Bonaccorso, E., Claramunt, S., Jaramillo, A., Lane, D.F., Pacheco, J.F., Robbins, M.B., Stiles, F.G. & Zimmer, K.J. (2021) A classification of the bird species of South America. American Ornithological Society. Proposal 802 by Mason, N.A. & Burns, K.J. (2017): Revise familial limits and the linear sequence of families within the nine-primaried oscines. Available from: https://www.museum.lsu.edu/~Remsen/SACCprop802.htm (accessed 13 April 2021)
    Ridgely, R.S. & Greenfield, P.J. (2001) The Birds of Ecuador. Status, Distribution and Taxonomy. Cornell University Press, Ithaca, New York, 848 pp.
    Ridgway, R. (1912) Color Standards and Nomenclature. Published by the Author, Washington, D.C., 43 pp.
    Ríos-Chelén, A.A., McDonald, A.N., Berger, A., Perry, A.C., Krakauer, A.H. & Patricelli, G.L. (2017) Do birds vocalize at higher pitch in noise, or is it a matter of measurement? Behavioral Ecology and Sociobiology, 71 (1), 29.
    Robbins, M.B. & Ridgely, R.S. (1990) The avifauna of an upper tropical cloud forest in southwestern Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia, 142, 59–71.
    Rocha-Méndez, A., Sánchez-González, L.A., Arbeláez-Cortés, E. & Navarro-Sigüenza, A.G. (2018) Phylogeography indicates incomplete genetic divergence among phenotypically differentiated montane forest populations of Atlapetes albinucha (Aves, Passerellidae). ZooKeys, 809, 125–148. https://doi.org/10.3897/zookeys.809.28743
    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–574. https://doi.org/10.1093/bioinformatics/btg180
    Salvin, O. (1864) Descriptions of seventeen new species of birds from Costa Rica. Proceedings of the Scientific Meetings of the Zoological Society of London, 32, 579–586.
    Sánchez-González, L.A., Navarro-Sigüenza, A.G., Peterson, A.T. & García-Moreno, J. (2007) Taxonomy of Chlorospingus ophthalmicus in Mexico and northern Central America. British Ornithologists’ Club, 127 (1), 34–49. https://doi.org/10.1111/zsc.12088
    Sánchez-González, L.A., Navarro-Sigüenza, A.G., Krabbe, N.K., Fjeldsa, J. & García-Moreno, J. (2015) Diversification in the Andes: the Atlapetes brush-finches. Zoologica Scripta, 44 (2), 135–152.
    Schulenberg, T.S., Stotz, D.F., Lane, D.F., O’Neill, J.P. & Parker, T.A. (2010) Birds of Peru. Revised and Updated Edition. Princeton University Press, Princeton, New Jersey, 664 pp.
    Sclater, P.L. (1859) List of the first collection of birds made by Mr. Louis Fraser at Pallatanga, Ecuador, with notes and descriptions of new species. Proceedings of the Zoological Society of London, 27, 135–147.
    Sclater, P.L. (1860) Lists of birds collected by Mr. Fraser in Ecuador, at Nanegal, Calacali, Perucho and Puellaro; with notes and descriptions of new species. Proceedings of the Zoological Society of London, 28, 83–97.
    Sclater, P.L. & Salvin, O. (1873) Nomenclator Avium Neotropicalium. J. W. Elliot, London, 163 pp.
    Sclater, P.L. & Salvin, O. (1877) Descriptions of six new species of South-American birds. Proceedings of the Zoological Society of London, 1887 (3), 521–522.
    Sclater, P.L. & Salvin, O. (1879a) Descriptions of some new tanagers of the genus Buarremon. Ibis, 3, 425–427.
    Sclater, P.L. & Salvin, O. (1879b) On the birds collected by the late Mr. T. K. Salmon in the State of Antioquia, United States of Colombia. Proceedings of the Zoological Society of London, 47 (1), 486–550.
    Smithe, F. (1975) Naturalist’s Color Guide. Part I. Museum of Natural History Press, New York.
    Smith, B.T., McCormack, J.E., Cuervo, A.M., Hickerson, M.J., Aleixo, A., Cadena, C.D., Perez-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 (7527), 406–409. https://doi.org/10.1038/nature13687
    Sosa-López, J.R., González, C. & Navarro-Sigüenza, A.G. (2013) Vocal geographic variation in Mesoamerican Common Bush-Tanagers (Chlorospingus ophthalmicus). Wilson Journal of Ornithology, 125 (1), 24–33. https://doi.org/10.1676/12-051.1
    Stephenson, T. & Whittle, S. (2013) The Warbler Guide. Princeton University Press, Princeton and Oxford, 560 pp.
    Stotz, D.F., Fitzpatrick, J.W., Parker, T.A. & Moskovits, D.K. (1996) Neotropical Birds: Ecology and Conservation. University of Chicago Press, Chicago, Illinois, 478 pp.
    Strewe, R. (2001) Notes on nests and breeding activity of fourteen bird species from southwestern Colombia. Ornitología Neotropical, 12, 265–269.
    Sullivan, B.L., Wood, C.L., Iliff, M.J., Bonney, R.E., Fink, D. & Kelling, S. (2009) eBird: a citizen-based bird observation network in the biological sciences. Biological Conservation, 142, 2282–2292.
    Taczanowski, L. (1874) Description des oiseaux nouveaux du Pérou central. Proceedings of the Zoological Society of London, 42 (1), 129–140.
    The Document Foundation (2020) LibreOffice Writer. (version 6.4.3.2). Available from: https://www.libreoffice.org/discover/writer/ (accessed 12 September 2020)
    Underdown, C.E. (1932) A study of Chlorospingus ophthalmicus (Dubus) and its allies. Ibis, 74, 638–655. https://doi.org/10.1111/j.1474-919X.1932.tb00355.x
    von Berlepsch, H.G. (1901) Mitteilungen Über die von den Gebrüdern G. und O. Garlepp in Bolivia gesammelten Vögel und Beschreibungen neuer Arten. Journal für Ornithologie, 49, 81–89. https://doi.org/10.1007/BF02206275
    Weir, J.T., Bermingham, E., Miller, M.J., Klicka, J. & González, M.A. (2008) Phylogeography of a morphologically diverse Neotropical montane species, the Common Bush-Tanager (Chlorospingus ophthalmicus). Molecular Phylogenetics and Evolution, 47, 650–664.
    Yánez-Múñoz, M., Sánchez, J.C. & Medina, G. (2015) Cap. I. Características Relevantes de la Provincia de El Oro. In: MECN-INB-GADPEO, Aves, Anfibios y Reptiles de la Provincia de El Oro. Una Guía para Ecosistemas Andino-Costeros. Publicación Miscelánea Nº 7. MECN-INB-GADPEO, Quito, pp. 1–12.
    Zimmer, J.T. (1947) Studies of Peruvian birds no. 52. The genera Sericossypha, Chlorospingus, Cnemoscopus, Hemispingus, Conothraupis, Chlorornis, Lamprospiza, Cissopis and Schistochlamys. American Museum Novitates, 1367, 4–5.

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