Abstract
In the 1850s, two species of "Spotted" Nightingale-Thrush (Aves: Catharus) were independently described from montane rainforests of Guatemala, C. dryas (Gould, 1855) and Ecuador, C. maculatus (Sclater, 1858). However, due to similarities in plumage color, C. maculatus was reclassified as a subspecies of C. dryas in 1878, a decision that has been upheld for 137 years. We collected multiple lines of evidence including phylogenetic analysis of mitochondrial DNA sequences (ND2), discriminant and principal components analysis of morphometric and vocal data, and statistical modeling of ecological niches, that collectively indicate that C. d. dryas and C. d. maculatus are independent species. We recommend restoring species status to C. maculatus of South America and applying the common name Sclater’s Nightingale-Thrush to this species.
References
Armenta, J.K., Weckstein, J.D. & Lane, D.F. (2005) Geographic variation in mitochondrial DNA sequences of an Amazonian non-passerine: The Black-spotted Barbet complex. Condor, 107, 527–536.
https://doi.org/10.1650/0010-5422(2005)107[0527:GVIMDS]2.0.CO;2Auer, S.K., Bassar, R.D., Fontaine, J.J. & Martin, T.E. (2007) Breeding biology of passerines in a subtropical montane forest in northwestern Argentina. Condor, 109, 321–333.
https://doi.org/10.1650/0010-5422(2007)109[321:BBOPIA]2.0.CO;2Baird, S.F. (1872) Review of American Birds in the Museum of the Smithsonian Institution, Part 1. Smithsonian Miscellaneous Collections, 181, 1– 478.
Baldwin, R. (2009) Use of maximum entropy modeling in wildlife research. Entropy, 11, 854–866.
https://doi.org/10.3390/e11040854Bent, A.C. (1949) Life Histories of North American Thrushes, Kinglets, and Their Allies. United States National Museum Bulletin, 196, 1–452.
Bonaparte, C.L. (1850) Conspectus Generum Avium. E. J. Brill, Leiden, 543 pp.
https://doi.org/10.5962/bhl.title.70841Brodkorb, P. (1938) New birds from the district of Soconusco, Chiapas. Occasional Papers of the Museum of Zoology, 369, 1–7. [University of Michigan]
Cabanis, P.J.G. (1861) Uebersicht der im Berliner Museum befindlichen Vögel von Costa Rica. Journal für Ornithologie, 8, 323.
Cadena, C.D. & Cuervo, A.M. (2010) Molecules, ecology, morphology and songs in concert: how many species is Arremon torquatus (Aves: Emberizidae)? Biological Journal of the Linnaean Society, 99, 152–176.
https://doi.org/10.1111/j.1095-8312.2009.01333.xCarriker Jr., M.A. (1935) Descriptions of new birds from Peru and Ecuador, with critical notes on other little-known species. Proceedings of the Academy of Natural Sciences of Philadelphia, 87, 343–359.
Chapman, F.M. (1926) The distribution of bird-life in Ecuador. A contribution to a study of the origin of Andean bird-life. Bulletin of the American Museum of Natural History, 55, 1–783.
Clement, P. (2000) Thrushes. Princeton University Press, Princeton, New Jersey, 463 pp.
Clements, J.F. (2007) The Clements Checklist of Birds of the World. Cornell University Press, Ithaca, New York, 864 pp.
Cracraft, J. (1983) Species concepts and speciation analysis. In: Johnston, R.F. (Ed.), Current Ornithology. Vol. 1. Plenum Press, New York, pp. 159–187.
https://doi.org/10.1007/978-1-4615-6781-3_6
Deignan, H.G. (1961) Type specimens of birds in the United States National Museum. Smithsonian Bulletin, 221, 1–718.Dilger, W.C. (1956) Hostile behavior and reproductive isolating mechanisms in the avian genera Catharus and Hylocichla. Auk, 73, 313–353.
https://doi.org/10.2307/4082003Dobzhansky, T. (1937) Genetics and the Origin of Species. Columbia University Press, New York, 364 pp.
Domaniewski, J. (1918) Die geographischen Formen von Catharus dryas Gould. Bulletin International de l‘Academie Polonaise des Sciences et des Lettres (Cracovie), 1918, 134–136.
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.
https://doi.org/10.1093/nar/gkh340ESRI (2014) ArcMap 10. ESRI (Environmental Systems Resource Institute), Redlands, California. [software]
Farr, T.G., Rosen, P.A., Caro, E., Crippen, R., Duren, R., Hensley, S., Kobrick, M., Paller, M., Rodriguez, E., Roth, L., Seal, D., Shaffer, S., Shimada, J., Umland, J., Werner, M., Oskin, M., Burbank, D. & Alsdorf, D. (2007) The Shuttle Radar Topography Mission. Reviews of Geophysics, 45 (2), RG2004. [published online]
https://doi.org/10.1029/2005rg000183Garzione, C.N., Hoke, G.D., Libarkin, J.C., Withers, S., MacFadden, B., Eiler, J., Ghosh, P. & Mulch, A. (2008) Rise of the Andes. Science, 320, 1304–1307.
https://doi.org/10.1126/science.1148615Gmelin, J.F. (1789) Systema Naturae. Vol. 1. Part 2. Impensis Georg Emanuel Beer, Leipzig, 500 pp.
Gould, J. (1855) Description of a new bird from Guatemala, forming the type of a new genus. Proceedings of the Zoological Society, 22, 285.
https://doi.org/10.1111/j.1469-7998.1854.tb07276.xGreeney, H.F., Dyrcz, A., Mikusek, R. & Port, J. (2015) Cooperative breeding at a nest of Slaty-backed Nightingale-Thrushes (Catharus fuscater). Wilson Journal of Ornithology, 127, 323–325.
https://doi.org/10.1676/wils-127-02-323-325.1Halley, M.R., Heckscher, C.M. & Kalavacharla, V. (2016) Multi-generational kinship, multiple mating, and flexible modes of parental care in a breeding population of the Veery (Catharus fuscescens), a trans-hemispheric migratory songbird. PLOS ONE, 11 (6), e0157051.
https://doi.org/10.1371/journal.pone.0157051Hansen, M.C., Potapov, P.V., Moore, R., Hancher, M., Turubanova, S.A., Tyukavina, A., Thau, D., Stehman, S.V., Goetz, S.J., Loveland, T.R., Kommareddy, A., Egorov, A., Chini, L., Justice, C.O. & Townshend, J.R.G. (2013) High-resolution global maps of 21st-Century forest cover change. Science 342, 850–853. Data available on-line. Available from: http://earthenginepartners.appspot.com/science-2013-global-forest (accessed 10 May 2017)
Heckscher, C.M., Halley, M.R. & Stampul, P. (2015) Intratropical migration of the Veery (Catharus fuscescens) in South America with implications for migration theory. Journal of Tropical Ecology, 31, 285–289.
https://doi.org/10.1017/S0266467415000024Helbig, A.J., Knox, A.G., Parkin, D.T., Sangster, G. & Collinson, M. (2002) Guidelines for assigning species rank. Ibis, 144, 518–525.
https://doi.org/10.1046/j.1474-919X.2002.00091.xHellmayr, C.E. (1934) Catalogue of birds of the Americas and the adjacent islands in Field Museum of Natural History, including all species and subspecies known to occur in North America, Mexico, Central America, the West Indies, and islands of the Caribbean Sea, the Galapagos Archipelago and other islands which may be included on account of their faunal affinities. Field Museum of Natural History, Zoological Series, 13 (7), 1–531.
Hijmans, R.J., Cameron, S.E., Parra, J.L., Jones, P.G. & Jarvis, A. (2005) Very high-resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965–1978.
https://doi.org/10.1002/joc.1276Hilty, 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. 2nd Edition. Princeton University Press, Princeton, New Jersey, 878 pp.
Howell, S.N.G. & Webb, S. (1995) A Guide to the Birds of Mexico and Northern Central America. Oxford University Press, New York, 851 pp.
Huxley, J. (1942) Evolution: The Modern Synthesis. George Allen & Unwin, London, 609 pp.
Lafresnaye, F. de (1845) Description de quelques oiseaux nouveaux. Revue Zoologique, 8, 337–342.
Lafresnaye, F. de (1848) Description de quelques oiseaux nouveaux de Caracas (province de Venezuela) et de Bogota. Revue Zoologique, 11, 2–12.
Lanyon, W.E. (1979) Development of song in the Wood Thrush (Hylocichla mustelina), with notes on a technique for hand-rearing passerines from the egg. American Museum Novitates, 2666, 1–27.
Marshall, J.T. (2000) The Gray-cheeked Thrush, Catharus minimus, and its New England subspecies, Bicknell's Thrush, Catharus minimus bicknelli. Smithsonian Institution, Washington D.C., 136 pp.
Mayr, E. (1942) Systematics and the Origin of Species, from the Viewpoint of a Biologist. Harvard University Press, Cambridge, Massachusetts, 372 pp.
Nuttall, T. (1840) A Manual of the Ornithology of the United States and Canada. 2nd Edition. Hilliard, Gray & Co., Boston, Massachusetts, 832 pp.
https://doi.org/10.5962/bhl.title.49241Olrog, C.C. (1973) Notas ornitologicas. IX. Sobre la collection del Instituto Miguel Lillo de Tucumán. Acta Zoologica Lilloana, 30, 7–11.
O'Neill, J.P., Lane, D.F. & Naka, L.N. (2011) A cryptic new species of thrush (Turdidae: Turdus) from western Amazonia. Condor, 113, 869–880.
https://doi.org/10.1525/cond.2011.100244Ouellet, H. (1993) Bicknell's Thrush: taxonomic status and distribution. Wilson Bulletin, 105, 545–754.
Ortiz-Ramírez, M.F., Andersen, M.J., Zaldívar-Riverón, A., Ornelas, J.F. & Navarro-Sigüenza, A.G. (2015) Geographic isolation drives divergence of uncorrelated genetic and song variation in the Ruddy-capped Nightingale-Thrush (Catharus frantzii; Aves: Turdidae). Molecular Phylogenetics and Evolution, 94, 74–86.
https://doi.org/10.1016/j.ympev.2015.08.017Outlaw, D.C., Voelker, G., Mila, B. & Girman, D.J. (2003) Evolution of long-distance migration in and historical biogeography of Catharus thrushes: a molecular phylogenetic approach. Auk, 120, 299–310.
https://doi.org/10.1642/0004-8038(2003)120[0299:EOLMIA]2.0.CO;2Pallas, P.S. (1811) Zoographia Rosso-asiatica. Petropoli, St. Petersburg, 568 pp.
Peterson, A.T. (2011) Ecological niche conservatism: a time-structured review of evidence. Journal of Biogeography, 38, 817–827.
https://doi.org/10.1111/j.1365-2699.2010.02456.xPhillips, A.R. (1969) An ornithological comedy of errors: Catharus occidentalis and C. frantzii. Auk, 86, 605–623.
https://doi.org/10.2307/4083450Phillips, A.R. & Rook, W. (1965) A new race of the Spotted Nightingale-Thrush from Oaxaca, Mexico. Condor, 67, 1–5.
https://doi.org/10.2307/1365376Phillips, A.R. (1991) Known Birds of North and Middle America. Part 2. Self-published, Denver, Colorado, 249 pp.
Price, T. (2008) Speciation in Birds. Roberts & Company Publishers, Greenwood Village, CO, 470 pp.
Raitt, R.J. & Hardy, J.W. (1970) Relationships between two partly sympatric species of thrushes (Catharus) in Mexico. Auk, 87, 20–57.
https://doi.org/10.2307/4083657R Core Team (2014) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: http://www.R-project.org/ (accessed 10 May 2017)
Ridgely, R.S. & Greenfield, P.J. (2001) The Birds of Ecuador. Vol. 2. Cornell University Press, Ithaca, New York, 740 pp.
Ridgely, R.S. & Tudor, G. (2009) Field Guide to the Songbirds of South America: The Passerines. University of Texas Press, Austin, Texas, 750 pp.
Ridgway, R. (1872) On the relation between color and geographical distribution in birds, as exhibited in melanism and hyperchromism. American Journal of Science, 3, 454–460.
https://doi.org/10.2475/ajs.s3-4.24.454Ridgway, R. (1882) Descriptions of two new thrushes from the United States. Proceedings of the United States National Museum, 4, 377–379.
https://doi.org/10.5479/si.00963801.240.374Ridgway, R. (1886) A Nomenclature of Colors and Compendium of Useful Knowledge for Ornithologists. Little Brown & Company, Boston, Massachusetts, 129 pp.
https://doi.org/10.5962/bhl.title.105202Ridgway, R. (1907) The birds of North and Middle America. In: United States National Museum Bulletin. Vol. 50. Part IV. Government Printing Office, Washington, D.C, 974 pp.
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.
Rowley, J.S. & Orr, R.T. (1964) The status of Frantzius' Nightingale Thrush. Auk, 81, 303–314.
https://doi.org/10.2307/4082686Ruegg, K.C. (2008) Genetic, morphological, and ecological characterization of a hybrid zone that spans a migratory divide. Evolution, 62, 452–466.
https://doi.org/10.1111/j.1558-5646.2007.00263.xSalvin, O. (1866) Descriptions of eight new species of birds from Veragua. Proceedings of the Zoological Society of London, 1, 67–74.
Salvin, O. & Godman, F. D. (1879) Biologia Centrali-Americana. Vol. 1. Aves. R.H. Porter, London, 512 pp.
Schulenberg, T.S., Stotz, D.F., Lane, D.F., O'Neill, J.P. & Parker, T.A. (2007) Birds of Peru. Princeton University Press, Princeton, New Jersey, 656 pp.
Sclater, P.L. (1858) Notes on a collection of birds received by M. Verreaux of Paris from the Rio Napo in the Republic of Ecuador. Proceedings of the Zoological Society of London, 26, 59–78.
https://doi.org/10.1111/j.1469-7998.1858.tb06346.xSclater, P.L. (1859) A synopsis of the thrushes (Turdidae) of the New World. Proceedings of the Zoological Society of London, 27, 321–347.
Seebohm, H. (1881) Catalogue of the Birds in the British Museum. Vol. 5. British Museum of Natural History, London, 426 pp.
Stein, R.C. (1956) A comparative study of "advertising song" in the Hylocichla thrushes. Auk, 73, 503–512.
https://doi.org/10.2307/4081948Stephens, J.F. (1817) General Zoology. Vol.10. Part 1. Privately published, London, 317 pp.
Stone, W. (1912) The phylogenetic value of color characters in birds. Journal of the Academy of Natural Sciences of Philadelphia, 15, 311–320.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), 4.0. Sinauer Associates, Sunderland, MA, 144 pp.
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: molecular evolutionary genetics analysis using Maximum Likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731–2739.
https://doi.org/10.1093/molbev/msr121Tobias, J.A., Seddon, N., Spottiswoode, C.N., Pilgrim, J.D., Fishpool, L.D.C. & Collar, N.J. (2010) Quantitative criteria for species delimitation. Ibis, 152, 724–746.
https://doi.org/10.1111/j.1474-919X.2010.01051.xTschudi, J.J. von (1845) Untersuchungen über die Fauna Peruana. Druck und Verlag von Scheitlin und Zollikofer, St. Gallen, 890 pp.
Voelker, G., Bowie, R.K.C. & Klicka, J. (2013) Gene trees, species trees, and Earth history combine to shed light on the evolution of migration in a model avian system. Molecular Ecology, 22, 3333–3344.
https://doi.org/10.1111/mec.12305Vuilleumier, F. (2003) Neotropical ornithology: Then and now. Auk, 120, 577–590.
https://doi.org/10.1642/0004-8038(2003)120[0577:PIONOT]2.0.CO;2Warren, D.L., Glor, R.E. & Turelli, M. (2008) Environmental niche equivalency versus conservatism: quantitative approaches to niche evolution. Evolution, 62, 2868–2883.
https://doi.org/10.1111/j.1558-5646.2008.00482.xWarren, D.L., Glor, R.E. & Turelli, M. (2010) ENMTools: A toolbox for comparative studies of environmental niche models. Ecography, 33, 607–611.
https://doi.org/10.1111/j.1600-0587.2009.06142.xWhitney, C.L. (1989) Geographical variation in Wood Thrush song: a comparison of samples recorded in New York and South Carolina. Behaviour, 111, 49–60.
https://doi.org/10.1163/156853989X00574Whitney, C.L. (1992) Temporal stability of song in a local population of Wood Thrushes. Wilson Bulletin, 104, 516–520.
Whitney, C.L. & Miller, J. (1987) Song learning in the wood thrush. Canadian Journal of Zoology, 65, 1038–1042.
https://doi.org/10.1139/z87-165Wiens, J.J. & Graham, C.H. (2005) Niche conservatism: integrating evolution, ecology, and conservation biology. Annual Review of Ecology, Evolution, and Systematics, 36, 519–539.
https://doi.org/10.1146/annurev.ecolsys.36.102803.095431Winker, K. (2000) Migration and speciation. Nature, 404, 36.
https://doi.org/10.1038/35003651Winker, K. & Pruett, C.L. (2006) Seasonal migration, speciation, and morphological convergence in the genus Catharus (Turdidae). Auk, 123, 1052–1068.
https://doi.org/10.1642/0004-8038(2006)123[1052:smsamc]2.0.co;2