Abstract
Megascops is the most species-rich owl genus in the New World, with 21 species currently recognized. Phylogenetic relationships within this genus are notoriously difficult to establish due to the considerable plumage similarity among species and polymorphism within species. Previous studies have suggested that the widespread lowland Amazonian M. watsonii might include more than one species, and that the Atlantic Forest endemic M. atricapilla is closely related to the M. watsonii complex, but these relationships are as yet poorly understood. A recently published phylogeny of Megascops demonstrated that M. watsonii is paraphyletic with respect to M. atricapilla and that genetic divergences among some populations of M. watsonii are equal to or surpass the degree of differentiation between some M. watsonii and M. atricapilla. To shed light on the taxonomic status of these species and populations within them, we conducted a multi-character study based on molecular, morphological, and vocal characters. We sequenced three mitochondrial (cytb, CO1 and ND2) and three nuclear genes (BF5, CHD and MUSK) for 49 specimens, covering most of the geographic ranges of M. watsonii and M. atricapilla, and used these sequences to estimate phylogenies under alternative Bayesian, Maximum Likelihood, and multilocus coalescent species tree approaches. We studied 252 specimens and vocal parameters from 83 recordings belonging to 65 individuals, distributed throughout the ranges of M. watsonii and M. atricapilla. We used Discriminant Function Analysis (DFA) to analyze both morphometric and vocal data, and a pairwise diagnostic test to evaluate the significance of vocal differences between distinct genetic lineages. Phylogenetic analyses consistently recovered six statistically well-supported clades whose relationships are not entirely in agreement with currently recognized species limits in M. watsonii and M. atricapilla. Morphometric analyses did not detect significant differences among clades. High plumage variation among individuals within clades was usually associated with the presence of two or more color morphs. By contrast, vocal analyses detected significant differentiation among some clades but considerable overlap among others, with some lineages (particularly the most widespread one) exhibiting significant regional variation. The combined results allow for a redefinition of species limits in both M. watsonii and M. atricapilla, with the recognition of four additional species, two of which we describe here as new. We estimated most cladogenesis in the Megascops atricapilla-M. watsonii complex as having taken place during the Plio-Pleistocene, with the development of the modern Amazonian and São Francisco drainages and the expansion and retraction of forest biomes during interglacial and glacial periods as likely events accounting for this relatively recent burst of diversification.
References
Alfaro, J.W.L., Boubli, J.P., Olson, L.E., Fiori, A.D., Wilson, B., Gutiérrez-Espeleta, G.A., Chiou, K.L., Schulte, M., Neitzel, S., Ross, V., Schwochow, D., Nguyen, M.T.T., Farias, I., Janson, C.H. & Alfaro, M.E. (2012) Explosive Pleistocene range expansion leads to widespread Amazonian sympatry between robust and gracile capuchin monkeys. Journal of Biogeography, 39, 272‒288.
https://doi.org/10.1111/j.1365-2699.2011.02609.x
American Ornithologists’ Union (1910) Check-list of North American Birds. 3rd Edition. American Ornithologists’ Union, New York, 430 pp.
Barrowclough, G.F., Groth, J.G., Mertz, L.A. & Gutiérrez, R.J. (2005) Genetic structure, introgression, and a narrow hybrid zone between northern and California spotted owls (Strix occidentalis). Molecular Ecology, 14, 1109–1120.
https://doi.org/10.1111/j.1365-294X.2005.02465.x
Batalha-Filho, H., Fjeldså, J., Fabre, P. & Miyaki, C.Y. (2013) Connections between the Atlantic and the Amazonian forest avifaunas represent distinct historical events. Journal of Ornithology, 154 (1), 41‒50.
https://doi.org/10.1007/s10336-012-0866-7
Batista, R., Aleixo, A., Vallinoto, M., Azevedo, L., Rego, P.S., Silveira, L.F., Sampaio, I. & Schneider, H. (2013) Molecular systematics and taxonomic revision of the Amazonian Barred Woodcreeper complex (Dendrocolaptes certhia: Dendrocolaptidae), with description of a new species from the Xingu-Tocantins interfluve. In: del Hoyo, J., Elliott, E., Sargatal, J., & Christie, D.A. (Eds.), Handbook of the Birds of the World. Special Volume: New Species and Global Index. First edition. Lynx Edicions, Barcelona, 800 pp.
Bergmann, C. (1847) Über die Verhältnisse der Wärmeökonomie der Thiere zu ihrer Grösse. Göttinger Studien, 3 (1), 595–708.
Bicudo, T.C., Sacek, V., Paes de Almeida, R., Bates, J.M. & Ribas, C.C. (2019) Andean tectonics and mantle dynamics as a pervasive influence on Amazonian ecosystem. Nature Scientific Reports, 9, 16879.
https://doi.org/10.1038/s41598-019-53465-y
Blotta, I., Prestinaci, F., Mirante, S. & Cantafora, A. (2005) Quantitative assay of total dsDNA with PicoGreen reagent and real-time fluorescent detection. Annali dell’Istituto Superiore di SanitaÌ, 41 (1), 119‒123.
Browning, M.R. (1989) The type specimens of Hekstra’s owls. Proceedings of the Biological Society of Washington, 102(2), 515‒519.
Burns K.J. & Naoki, K. (2004) Molecular phylogenetics and biogeography of Neotropical tanagers in the genus Tangara. Molecular Phylogenetics and Evolution, 32, 838–854.
https://doi.org/10.1016/j.ympev.2004.02.013
Cabanne, G.S., d’Horta, F.M., Sari, E.H.R., Santos, F.R. & Miyaki, C.Y. (2008) Nuclear and mitochondrial phylogeography of the Atlantic forest endemic Xiphorhynchus fuscus (Aves: Dendrocolaptidae): biogeography and systematics implications. Molecular Phylogenetics and Evolution, 49, 760–773.
https://doi.org/10.1016/j.ympev.2008.09.013
Camargo, A., Morando, M., Avila, L.J. & Sites Jr., J.W. (2012) Species delimitation with ABC and other coalescent-based methods: a test of accuracy with simulations and an empirical example with lizards of the Liolaemus darwinii complex (Squamata: Liolaemidae). Evolution, 66, 2834–2849.
https://doi.org/10.1111/j.1558-5646.2012.01640.x
Capurucho, J.M.G., Ashley, M.V., Ribas, C.C. & Bates, J.M. (2018) Connecting Amazonian, Cerrado, and Atlantic Forest histories: paraphyly, old divergences, and modern population dynamics in tyrant-manakins (Neopelma/Tyranneutes, Aves: Pipridae). Molecular Phylogenetics and Evolution, 127, 696‒705.
https://doi.org/10.1016/j.ympev.2018.06.015
Carnaval, A.C., Hickerson, M.J., Haddad, C.F.B., Rodrigues, M.T. & Moritz, C. (2009) Stability predicts genetic diversity in the Brazilian Atlantic forest hotspot. Science, 323, 785–789.
https://doi.org/10.1126/science.1166955
Carneiro, L.S., Gonzaga, L.P., Rêgo, P.S., Sampaio, I., Schneider, H. & Aleixo, A. (2012) Systematic revision of the Spotted Antpitta (Grallariidae: Hylopezus macularius), with description of a cryptic new species from Brazilian Amazonia. The Auk, 129 (2), 338‒351.
https://doi.org/10.1525/auk.2012.11157
Cassin, J. (1849) Descriptions of owls, presumed to be undescribed, specimens of which are in the collection of the Academy of Natural Sciences of Philadelphia. Proceedings of the Academy of Natural Sciences of Philadelphia, 4, 753‒774.
Chapman, F.M. (1928) Descriptions of new birds from eastern Ecuador and eastern Peru. American Museum Novitates, No. 332, 1‒12.
Chaves, J.C., Cuervo, A.M., Miller, M.J. & Cadena, C.D. (2010) Revising species limits in a group of Myrmeciza antbirds reveals a cryptic species within M. laemosticta (Thamnophilidae). The Condor, 112 (4), 718–730.
https://doi.org/10.1525/cond.2010.100098
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 16 February 2021)
Conn, J.E. & Mirabello, L. (2007) The biogeography and population genetics of Neotropical vector species. Heredity, 99, 245–256.
https://doi.org/10.1038/sj.hdy.6801002
Cory, C.B. (1918) Catalogue of Birds of the Americas. Part. II. Field Museum of Natural History Zoological Series XIII. Field Museum, Chicago, Illinois, 315 pp.
Dantas, S.M., Weckstein, J.D., Bates, J.M., Krabbe, N.K., Cadena, C.D., Robbins, M.B., Valderrama, E. & Aleixo, A. (2016) Molecular systematics of the new world screech-owls (Megascops: Aves, Strigidae): biogeographic and taxonomic implications. Molecular Phylogenetics and Evolution, 94, 626‒634.
https://doi.org/10.1016/j.ympev.2015.09.025
de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, Oxford, pp. 57‒75.
d´Horta, F.M., Cuervo, A.M., Ribas, C.C., Brumfield, R.T. & Miyaki, C.Y. (2013) Phylogeny and comparative phylogeography of Sclerurus (Aves: Furnariidae) reveal constant and cryptic diversification in an old radiation of rain forest understorey specialists. Journal of Biogeography, 40, 37–49.
https://doi.org/10.1111/j.1365-2699.2012.02760.x
Dickinson, E.C. & Remsen Jr., J.V. (Eds.), (2013) The Howard and Moore Complete Checklist of the Birds of the World. Vol. 1. 4th Edition. Non-Passerines, Aves Press, Eastbourne, 461 pp.
Driskell, A.C. & Christidis, L. (2004) Phylogeny and evolution of the Australo-Papuan honeyeaters (Passeriformes, Meliphagidae). Molecular Phylogenetics and Evolution, 31, 943–960.
https://doi.org/10.1016/j.ympev.2003.10.017
Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
https://doi.org/10.1186/1471-2148-7-214
Drummond, A.J., Ho, S.Y.W., Phillips, M.J. & Rambaut, A. (2006) Relaxed phylogenetics and dating with confidence. PLoS Biology, 4, e88.
https://doi.org/10.1371/journal.pbio.0040088
Eberhard, J.R. & Bermingham, E. (2005) Phylogeny and comparative biogeography of Pionopsitta parrots and Pteroglossus toucans. Molecular Phylogenetics and Evolution, 36, 288–304.
https://doi.org/10.1016/j.ympev.2005.01.022
Enríquez, P.L., Eisermann, K., Mikkola, H. & Motta-Junior, J.C. (2017) A review of the systematics of Neotropical owls (Strigiformes). In: Enriquez, P. (Ed.), Neotropical Owls. Springer, Cham, pp. 1‒670.
https://doi.org/10.1007/978-3-319-57108-9_2
Faircloth, B.C., McCormack, J.E., Crawford, N.G., Harvey, M.G., Brumfield, R.T. & Glenn, T.C. (2012) Ultraconserved elements anchor thousands of genetic markers spanning multiple evolutionary timescales. Systematic Biology, 6 (5), 717‒726.
https://doi.org/10.1093/sysbio/sys004
Fernandes, A.M., Wink, M. & Aleixo, A. (2012) Phylogeography of the Chestnut-tailed Antbird (Myrmeciza hemimelaena) clarifies the role of rivers in Amazonian biogeography. Journal of Biogeography, 39, 1524–1535.
https://doi.org/10.1111/j.1365-2699.2012.02712.x
Fernandes, A.M., Gonzalez, J., Wink, M. & Aleixo, A. (2013) Multilocus phylogeography of the Wedge-billed Woodcreeper Glyphorynchus spirurus (Aves, Furnariidae) in lowland Amazonia: Widespread cryptic diversity and paraphyly reveal a complex diversification pattern. Molecular Phylogenetics and Evolution, 66, 270–282.
https://doi.org/10.1016/j.ympev.2012.09.033
Fernandes, A.M., Wink, M., Sardelli, C.H. & Aleixo, A. (2014) Multiple speciation across the Andes and throughout Amazonia: the case of the Spot-backed Antbird species complex (Hylophylax naevius/Hylophylax naevioides). Journal of Biogeography, 41, 1094–1104.
https://doi.org/10.1111/jbi.12277
Fjeldså, J., Baiker, J., Engblom, G., Franke, I., Geale, D., Krabbe, N.K., Lane, D.E., Lezama, M., Schmitt, F., Williams, R.S.R., Ugarte-Núñez, J., Yábar, V. & Yábar, R. (2012) Reappraisal of Koepcke’s Screech Owl Megascops koepckeae and description of a new subspecies. Bulletin of the British Ornithologists’ Club, 132, 180‒193.
Fuchs, J., Pons, J., Pasquet, E., Raherilalao, M.J. & Goodman, S.M. (2007) Geographical structure of genetic variation in the Malagasy Scops-owl inferred from mitochondrial sequence data. The Condor, 109, 408–418.
https://doi.org/10.1093/condor/109.2.408
Fuchs, J., Pons, J., Goodman, S.M., Bretagnolle, V., Melo, M., Bowie, R.C.K., Currie, D., Safford, R., Virani, M.Z., Thomsett, S., Hija, A., Cruaud, C. & Pasquet, E. (2008) Tracing the colonization history of the Indian Ocean scops-owls (Strigiformes: Otus) with further insight into the spatio-temporal origin of the Malagasy avifauna. BMC Evolutionary Biology, 8, 197.
https://doi.org/10.1186/1471-2148-8-197
Gascon, C., Malcolm, J.R., Patton, J.L., Silva, M.N.F., Bogart, J.P., Lougheed, S.C., Peres, C.A., Neckel, S. & Boag, P.T. (2000) Riverine barriers and the geographic distribution of Amazonian species. Proceedings of National Academy of Sciences USA, 97, 13672–13677.
https://doi.org/10.1073/pnas.230136397
Gill, F.B. (2014) Species taxonomy of birds: which null hypothesis? The Auk, 131 (2), 150‒161.
https://doi.org/10.1642/AUK-13-206.1
Gill, F., Donsker, D. & Rasmussen, P. (2020) IOC World Bird List. Version 10.1. Available from: https://doi.org/10.14344/IOC.ML.10.1 (accessed 16 February 2021)
Gouy, M., Guindon, S. & Gascuel, O. (2010) SeaView Version 4: A Multiplatform Graphical User Interface for Sequence Alignment and Phylogenetic Tree Building. Molecular Biology and Evolution, 27 (2), 221‒224.
https://doi.org/10.1093/molbev/msp259
Hackett, S.J. (1996) Molecular phylogenetics and biogeography of tanagers in the genus Ramphocelus (Aves). Molecular Phylogenetics and Evolution, 5, 368–382.
https://doi.org/10.1006/mpev.1996.0032
Haffer, J. (1974) Avian Speciation in Tropical South America, with a Systematic Survey of the Toucans (Ramphastidae) and Jacamars (Galbulidae). Publications of the Nuttall Ornithological Club, Cambridge, Massachusetts, 390 pp.
Haffer, J. (1997) Contact zones between birds of southern Amazonia. In: Remsen Jr., J.V. (Ed.), Studies in Neotropical ornithology honoring Ted Parker. Ornithological Monographs, 48, pp. 281–306.
https://doi.org/10.2307/40157539
Hafner, M.S., Skudman, P.D., Villablanca, F.X., Spradling, T.A., Demastes, J.W. & Nadler, S.A. (1994) Disparate rates of molecular evolution in cospeciating hosts and parasites. Science, 265, 1087–1090.
https://doi.org/10.1126/science.8066445
Han, K., Robbins, M.B. & Braun, M.J. (2010) A multi-gene estimate of phylogeny in the nightjars and nighthawks (Caprimulgidae). Molecular Phylogenetics and Evolution, 55, 443–453.
https://doi.org/10.1016/j.ympev.2010.01.023
Hanna, Z.R., Henderson, J.B., Sellas, A.B., Fuchs, J., Bowie, R.C.K. & Dumbacher, J.P. (2017) Complete mitochondrial genome sequences of the Northern Spotted Owl (Strix occidentalis caurina) and the Barred Owl (Strix varia; Aves: Strigiformes: Strigidae) confirm the presence of a duplicated control region. PeerJ, 5, e3901.
https://doi.org/10.7717/peerj.3901
Harvey, M.G., Aleixo, A., Ribas, C.C. & Brumfield, R.T. (2017) Habitat association predicts genetic diversity and population divergence in Amazonian birds. American Naturalist, 190, 631‒648.
https://doi.org/10.1086/693856
Heidrich, P., König, C. & Wink, M. (1995) Molecular phylogeny of South American screech owls of the Otus atricapillus complex (Aves: Strigidae) inferred from nucleotide sequences of the mitochondrial cytochrome b gene. Zeitschrift für Naturforschung, 50c, 294‒302.
https://doi.org/10.1515/znc-1995-3-420
Hekstra, G.P. (1982) Description of twenty-four new subspecies of American Otus (Aves, Strigidae). Bulletin Zoologische Museum, Universiteit van Amsterdam, 9 (7), 49‒63.
Hellmayr, C.E. (1907) On a collection of birds made by Mr. W. Hoffmanns on the Rio Madeira, Brazil. Novitates Zoologicae, 14, 343‒412.
Hellmayr, C.E. (1910) The birds of the Rio Madeira. Novitates Zoologicae, 17, 257‒428.
https://doi.org/10.5962/bhl.title.875
Helm-Bychowski, K. & Cracraft, J. (1993) Recovering phylogenetic signal from DNA sequences: relationships within the corvine assemblage (Class Aves) as inferred from complete sequences of mitochondrial cytochrome-b gene. Molecular Biology and Evolution, 10, 1196–1214.
Herzog, S.K., Ewing, S.R., Evans, K.L., Maccormick, A., Valqui, T., Bryce, R. Kessler, M. & Macleod, R. (2009) Vocalizations, distribution, and ecology of the Cloud-forest Screech Owl (Megascops marshalli). The Wilson Journal of Ornithology, 121 (2), 240–252.
https://doi.org/10.1676/08-041.1
Hoek Ostende, L.W. van den, Dekker, R.W.R.J. & Keijl, G.O. 1997. Type-specimens of birds in the National Museum of Natural History, Leiden. Part 1. Non-passerines. Nationaal Natuurhistorisch Museum Technical Bulletin, 1, 1‒248.
InfoNatura: Animals and Ecosystems of Latin America [web application]. (2007) Version 5.0. NatureServe, Arlington, Virginia. Available from: http://www.natureserve.org/infonatura (accessed 18 March 2013)
Isler, M.L., Isler, P.R. & Whitney, B.M. (1998) Use of vocalizations to establish species limits in antbirds (Passeriformes: Thamnophilidae). The Auk, 115 (3), 577‒590.
https://doi.org/10.2307/4089407
Jacobsen, F., Friedman, N.R. & Omland, K.E. (2010) Congruence between nuclear and mitochondrial DNA: combination of multiple nuclear introns resolves a well-supported phylogeny of New World orioles (Icterus). Molecular Phylogenetics and Evolution, 56, 419–427.
https://doi.org/10.1016/j.ympev.2010.03.035
Kimball, R.T., Braun, E.L., Barker, F.K., Bowie, R.C.K., Braun, M.J., Chojnowski, J.L., Hackett, S.J., Han, K., Harshman, J., Heimer-Torres, V., Holznagel, W., Huddleston, C.J., Marks, B.D., Miglia, K.J., Moore, W.S., Reddy, S., Sheldon, F.H., Smith, J.V., Witt, C.C. & Yuri, T. (2009) A well-tested set of primers to amplify regions spread across the avian genome. Molecular Phylogenetics and Evolution, 50, 654‒660.
https://doi.org/10.1016/j.ympev.2008.11.018
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of mitochondrial DNA evolution in animals: Amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences, 86, 6196–6200.
https://doi.org/10.1073/pnas.86.16.6196
König, C. & Weick, F. (2008) Owls of the World. Christopher Helm/A&C Black Publishers Ltd., London, 528 pp.
König, C., Weick, F. & Becking, J.H. (1999) Owls: a Guide to the Owls of the World. Yale University Press, New Haven, Connecticut, 462 pp.
Krabbe, N. (2017) A new species of Megascops (Strigidae) from the Sierra Nevada de Santa Marta, Colombia, with notes on voices of New World screech-owls. Ornitología Colombiana, 16 (eA08), 1–27.
Leaché, A.D. & Fujita, M.K. (2010) Bayesian species delimitation in West African forest geckos (Hemidactylus fasciatus). Proceedings of the Royal Society B, 277, 3071–3077.
https://doi.org/10.1098/rspb.2010.0662
Leaché, A.D., Zhu, T., Rannala, B. & Yang, Z. (2019) The spectre of too many species. Systematic Biology, 68 (1), 168–181.
https://doi.org/10.1093/sysbio/syy051
Librado, P. & Rozas, J. (2009) DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.
https://doi.org/10.1093/bioinformatics/btp187
Malacco, G. (2013) [WA1004597, Megascops atricapilla (Temminck, 1822)]. Wiki Aves—A Enciclopédia das Aves do Brasil. Available from: http://www.wikiaves.com/1004597 (accessed 6 April 2016)
Marks, J.S., Cannings, R.J. & Mikkola, H. (1999) Family Strigidae. In: del Hoyo, J., Elliott, A., &
Sargatal, J. (Eds.), Handbook of the Birds of the World. Vol. 5. Barn-owls to Hummingbirds. Lynx Edicions, Barcelona, pp. 1‒759.
Marshall, J.T. (1967) Parallel variation in North and Middle American screech-owls. Monographs of the Western Foundation of Vertebrate Zoology, 1, 1‒72.
Marshall, J.T. (1978) Systematics of smaller Asian night birds based on voice. Ornithological Monographs, 25, 1‒58.
https://doi.org/10.2307/40166757
Marshall, J.T. & King, B. (1988) Genus Otus. In: Amadon, D. & Bull, J. (Eds.), Hawks and owls of the world: A distributional and taxonomic list. Proceedings of the Western Foundation of Vertebrate Zoology, 3, pp. 296–357.
Marshall J.T. (1991) The variable screech owl (Otus atricapillus) and its relatives. Wilson Bulletin, 103 (2), 314‒315.
Matzke, N.J. (2013) Probabilistic historical biogeography: new models for founder event speciation, imperfect detection, and fossils allow improved accuracy and modeltesting. Frontiers of Biogeography, 5, 242–248.
https://doi.org/10.21425/F55419694
McCormack, J.E., Harvey, M.G., Faircloth, B.C., Crawford, N.G., Glenn, T.C. & Brumfield, R.T. (2013) A phylogeny of birds based on over 1,500 loci collected by target enrichment and high throughput sequencing. PLoS One, 8, e54848.
https://doi.org/10.1371/journal.pone.0054848
McKay, B.D., Mays Jr., H.L., Wu, Y., Li, H., Yao, C.T., Nishiumi, I. & Zou, F. (2013) An empirical comparison of character-based and coalescent-based approaches to species delimitation in a young avian complex. Molecular Ecology, 22, 4943‒4957.
https://doi.org/10.1111/mec.12446
Miller, M.J., Bermingham, E., Klicka, J., Escalante, P., 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 Biological Sciences, 275 (1639), 1133‒1142.
https://doi.org/10.1098/rspb.2008.0015
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) “Creating the CIPRES Science Gateway for inference of large phylogenetic trees”. In: 2010 Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, pp. 1‒8.
https://doi.org/10.1109/GCE.2010.5676129
Naka, L.N. (2011) Avian distribution patterns in the Guiana Shield: implications for the delimitation of Amazonian areas of endemism. Journal of Biogeography, 38, 681–696.
https://doi.org/10.1111/j.1365-2699.2010.02443.x
Naka, L.N., Bechtoldt, C.L., Henriques, L.M.P. & Brumfield, R.T. (2012) The role of physical barriers in the location of avian suture zones in the Guiana Shield, Northern Amazonia. The American Naturalist, 179 (4), 115‒132.
https://doi.org/10.1086/664627
Nylander, J.A.A. (2004) MrModeltest 2.2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala. [program]
Patané, J.S.L., Weckstein, J.D., Aleixo, A. & Bates, J.M. (2009) Evolutionary history of Ramphastos toucans: molecular phylogenetics, temporal diversification, and biogeography. Molecular Phylogenetics and Evolution, 53, 923–934. https://doi.org/10.1016/j.ympev.2009.08.017
Patel, S., Weckstein, J.D., Patané, J.S.L., Bates, J.M. & Aleixo, A. (2011) Temporal and spatial diversification of Pteroglossus aracaris (Aves: Ramphastidae) in the Neotropics: Constant rate of diversification does not support an increase in radiation during the Pleistocene. Molecular Phylogenetics and Evolution, 58, 105–115.
https://doi.org/10.1016/j.ympev.2010.10.016
Paynter, R.A. (1982) Ornithological Gazetteer of Venezuela. Museum of Comparative Zoology, Cambridge, Massachusetts, 244 pp.
https://doi.org/10.5962/bhl.title.14633
Paynter, R.A. (1989) Ornithological Gazetteer of Paraguay. Museum of Comparative Zoology, Cambridge, Massachusetts, 59 pp.
https://doi.org/10.5962/bhl.title.14584
Paynter, R.A. & Traylor, M.A. (1991) Ornithological Gazetteer of Brazil. Museum of Comparative Zoology, Cambridge, Massachusetts, 788 pp.
Paynter, R.A. (1992) Ornithological Gazetteer of Bolivia. Museum of Comparative Zoology, Cambridge, Massachusetts, 185 pp.
https://doi.org/10.5962/bhl.title.14591
Paynter, R.A. (1993) Ornithological Gazetteer of Ecuador. Museum of Comparative Zoology, Cambridge, Massachusetts, 247 pp.
https://doi.org/10.5962/bhl.title.14626
Paynter, R.A. (1997) Ornithological Gazetteer of Colombia. Museum of Comparative Zoology, Cambridge, Massachusetts, 537 pp.
https://doi.org/10.5962/bhl.title.14638
Pereira, G.A., Dantas, S.M., Silveira, L.F., Roda, S.A., Albano, C., Sonntag, F.A., Leal, S., Periquito, M.C., Malacco, G.B. & Lees, A.C. (2014) Status of the globally threatened forest birds of northeast Brazil. Papéis Avulsos de Zoologia, 54 (14), 177‒194.
https://doi.org/10.1590/0031-1049.2014.54.14
Peters, J.L. (1940) Check-list of Birds of the World. Vol. 4. Museum of Comparative Zoology, Cambridge, Massachusetts, 291 pp.
Piacentini, V.Q., Aleixo, A., Agne, C.E., Mauricio, G.N., Pacheco, J.F., Bravo, G.A., Brito, G.R.R., Naka, L.N., Olmos, F., Posso, S., Silveira, L.F., Betini, G.S., Carrano, E., Franz, I., Lees, A.C., Lima, L.M., Pioli, D., Schunck, F., Amaral, F.R., Bencke, G.A., Cohn-Haft, M., Figueiredo, L.F.A., Straube, F.C. & Cesari, E. (2015) Annotated checklist of the birds of Brazil by the Brazilian Ornithological Records Committee/Lista comentada das aves do Brasil pelo Comitê Brasileiro de Registros Ornitológicos. Revista Brasileira de Ornitologia, 23, 91–298.
https://doi.org/10.1007/BF03544294
Proudfoot, G.A., Gehlbargh, F.R. & Honeycutt, R.L. (2007) Mitochondrial DNA variation and phylogeography of the Eastern and Western screech-owls. The Condor, 109, 617‒627.
https://doi.org/10.1093/condor/109.3.617
Pulido-Santacruz, P., Aleixo, A. & Weir, J.T. (2018) Morphologically cryptic Amazonian bird species pairs exhibit strong postzygotic reproductive isolation. Proceedings of the Royal Society B, 285 20172081.
https://doi.org/10.1098/rspb.2017.2081
Raposo do Amaral, F., Neves, L.G., Resende Jr., M.F.R., Mobili, F., Miyaki, C.Y., Pellegrino, K.C.M. & Biondo, C. (2015) Ultraconserved elements sequencing as a low-cost source of complete mitochondrial genomes and microsatellite markers in non-model amniotes. PLoS One, 10 (9), e0138446.
https://doi.org/10.1371/journal.pone.0138446
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Version 1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 16 February 2021)
Rannala, B. & Yang, Z. (2003) Bayes estimation of species divergence times and ancestral population sizes using DNA sequences from multiple loci. Genetics, 164 (4), 1645‒1656.
Rasmussen, P. C., Allen, D.N.S., Collar, N.J., Demeulemeester, B., Hutchinson, R.O., Jakosalem, P. G.C., Kennedy, R.S., Lambert, F.R. & Paguntalan, L.M. (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex. Forktail, 28, 1–20.
Ree, R.H. & Sanmartín, I. (2018) Conceptual and statistical problems with the DEC+J model of founder-event speciation and its comparison with DEC via model selection. Journal of Biogeography, 45 (4), 741‒749.
https://doi.org/10.1111/jbi.13173
Remsen Jr., J.V. (2005) Pattern, process, and rigor meet classification. The Auk, 122, 403–413.
https://doi.org/10.1093/auk/122.2.403
Remsen Jr., J.V., Areta, J.I., Cadena, C.D., Claramunt, S., Jaramillo, A., Pacheco, J.F., Robbins, M.B., Stiles, F.G., Stotz, D.F. & Zimmer, K.J. (2020) A Classification of the Bird Species of South America. Version 8 June 2020. American Ornithological Society. Available from: http://www.museum.lsu.edu/~Remsen/SACCBaseline.htm (accessed 16 February 2021)
Ribas, C.C., Miyaki, C.Y. & Cracraft, J. (2009) Phylogenetic relationships, diversification and biogeography in Neotropical Brotogeris parakeets. Journal of Biogeography 36, 1712–1729.
https://doi.org/10.1111/j.1365-2699.2009.02131.x
Ribas, C.C., Moyle, R.G., Miyaki, C.Y. & Cracraft, J. (2007) The assembly of montane biotas: linking Andean tectonics and climatic oscillations to independent regimes of diversification in Pionus parrots. Proceedings of the Royal Society B, 274, 2399–2408.
https://doi.org/10.1098/rspb.2007.0613
Ribas, C.C., Aleixo, A., Nogueira, A.C.R., Miyaki, C.Y. & Cracraft, J. (2012) A palaeobiogeographic model for biotic diversification within Amazonia over the past three million years. Proceedings of the Royal Society B, 279, 1712‒1729.
https://doi.org/10.1098/rspb.2011.1120
Rocha, T.C., Sequeira, F., Aleixo, A., Rêgo, P.S., Sampaio, I., Schneider, H. & Vallinoto, M. (2015) Molecular phylogeny and diversification of a widespread Neotropical rainforest bird group: The Buff-throated Woodcreeper complex, Xiphorhynchus guttatus/susurrans (Aves: Dendrocolaptidae). Molecular Phylogenetics and Evolution, 85, 131‒140.
https://doi.org/10.1016/j.ympev.2015.02.004
Roda, S.A. & Pereira, G.A. (2006) Distribuição recente e conservação das aves de rapina florestais do Centro Pernambuco. Revista Brasileira de Ornitologia, 14 (4), 331‒344.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572‒1574.
https://doi.org/10.1093/bioinformatics/btg180
Roulin, A., Burri, R. & Antoniazza, S. (2011) Owl melanin-based plumage redness is more frequent near than away from the equator: implications on the effect of climate change on biodiversity. Biological Journal of the Linnean Society, 102, 573–582.
https://doi.org/10.1111/j.1095-8312.2010.01614.x
Sambrook, J. & Russel, D.W. (2001) Molecular Cloning: A Laboratory Manual. 3rd Edition. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, 999 pp.
Salter, J.F., Oliveros, C.H., Hosner, P.A., Manthey, J.D., Robbins, M.B., Moyle, R.G., Brumfield, R.T. & Faircloth, B.C. (2020) Extensive paraphyly in the typical owl family (Strigidae). The Auk, 137 (1), 1‒15.
https://doi.org/10.1093/auk/ukz070
Salzburger, W., Ewing, G.B. & Von Haeseler, A. (2011) The performance of phylogenetic algorithms in estimating haplotype genealogies with migration. Molecular Ecology, 20, 1952–1963.
https://doi.org/10.1111/j.1365-294X.2011.05066.x
Sangster, G., King, B.F., Verbelen, P. & Trainor, C.R. (2013) A new owl species of the genus Otus (Aves: Strigidae) from Lombok, Indonesia. PLoS ONE, 8 (2), e53712.
https://doi.org/10.1371/journal.pone.0053712
Schultz, E.D., Perez-Eman, 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.
https://doi.org/10.1016/j.ympev.2019.106581
Sclater, P.L. (1858) On some new or little-known species of Accipitres, in the collection of the Norwich Museum. Proceedings of the Zoological Society of London, 1858, 128‒133.
https://doi.org/10.1111/j.1469-7998.1858.tb06354.x
Silva, J.M.C., Rylands, A.B. & Fonseca, G.A.B. (2005) O destino das áreas de endemismo da Amazônia. Megadiversidade, 1 (1), 124‒131.
Silva, S.M., Peterson, A.T., Carneiro, L., Burlamaqui, T.C.T., Ribas, C.C., Sousa-Neves, T., Miranda, L.S., Fernandes, A.M., d’Horta, F.M., Araújo-Silva, L.E., Batista, R., Bandeira, C.H.M.M., Dantas, S.M., Ferreira, M., Martins, D.M., Oliveira, J., Rocha, T.C., Sardelli, C.H., Thom, G., Rêgo, P.S., Santos, M.P., Sequeira, F., Vallinoto, M. & Aleixo, A. (2019) A dynamic continental moisture gradient drove Amazonian bird diversification. Science Advances, 5 (7), eaat5752.
https://doi.org/10.1126/sciadv.aat5752
Smith, B.T., Ribas, C.C., Whitney, B.M., Hernández-Banõs, B.E. & Klicka, J. (2013) Identifying biases at different spatial and temporal scales of diversification: A case study in the Neotropical parrotlet genus Forpus. Molecular Ecology, 22, 483‒494.
https://doi.org/10.1111/mec.12118
Smithe, F.B. (1975) Naturalist’s Color Guide. American Museum of Natural History, New York, New York, xiii + 229 pp.
Snethlage, E. (1914) Catálogo das aves amazônicas. Boletim do Museu Paraense Emílio Goeldi de Historia Natural e Etnografia, 8, 1‒530.
Soares, L.M.D.S., Bates, J., Carneiro, L.S., Santos, M.P.D. & Aleixo, A. (2019) Molecular systematics, biogeography and taxonomy of forest-falcons in the Micrastur ruficollis species complex (Aves: Falconidae). Journal of Avian Biology, 50 (4), e01943.
https://doi.org/10.1111/jav.01943
Sorenson, M.D., Ast, J.C., Dimcheff, D.E., Yuri, T. & Mindell, D.P. (1999) Primers for a PCR-based approach to mitochondrial genome sequencing in birds and other vertebrates. Molecular Phylogenetics and Evolution, 12, 105–114.
https://doi.org/10.1006/mpev.1998.0602
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446
Stephens, L. & Traylor, M.A. (1983) Ornithological Gazetteer of Peru. Museum of Comparative Zoology, Cambridge, Massachusetts, 271 pp.
Stephens, L. & Traylor, M.A. (1985) Ornithological Gazetteer of the Guianas. Museum of Comparative Zoology, Cambridge, Massachusetts, 121 pp.
Stephens, M. & Donnelly, P. (2003) A comparison of Bayesian methods for haplotype reconstruction from population genotype data. American Journal of Human Genetics, 73, 1162–1169.
https://doi.org/10.1086/379378
Stephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68, 978–989.
https://doi.org/10.1086/319501
Swofford, D.L. (2002) PAUP* Phylogenetic Analysis Using Parsimony (*and other methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]
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 (10), 2731–2739.
https://doi.org/10.1093/molbev/msr121
Temminck, C.J. (1822) Nouveau recueil des Planches Coloriées d’oiseaux. Vol. 2. Livraison 25. Pl. 145. F. G. Levrault, Libraire-editeur, Paris, 278 pp.
Thom, G. & Aleixo, A. (2015) Cryptic speciation in the White-shouldered Antshrike (Thamnophilus aethiops, Aves-Thamnophilidae): the tale of a transcontinental radiation across rivers in lowland Amazonia and the northeastern Atlantic Forest. Molecular Phylogenetics and Evolution, 82, 95‒110.
https://doi.org/10.1016/j.ympev.2014.09.023
van der Weyden, W.J. (1975) Scops and screech owls: vocal evidence for a basic subdivision in the genus Otus (Strigidae). Ardea, 63, 65‒77.
Weir, J.T. & Schluter, D. (2008) Calibrating the avian molecular clock. Molecular Ecology, 17, 2321‒2328.
https://doi.org/10.1111/j.1365-294X.2008.03742.x
Weir, J.T. & Price, M. (2011) Andean uplift promotes lowland speciation through vicariance and dispersal in Dendrocincla woodcreepers. Molecular Ecology, 20, 4550‒4563.
https://doi.org/10.1111/j.1365-294X.2011.05294.x
Winger, B.M. & Bates, J.M. (2015) The tempo of trait divergence in geographic isolation: avian speciation across the Marañon Valley of Peru. Evolution, 69 (3), 772‒787.
https://doi.org/10.1111/evo.12607
Wink, M., Heidrich, P., Sauer-Gürth, H., El-Sayed, A.A. & Gonzalez, J.M. (2008) Molecular phylogeny and systematics of owls (Strigiformes). In: König, C. & Weick, F. (Eds.), Owls of the World. 2nd Edition. Christopher Helm, London, pp. 42–63.
Wink, M. & Heidrich, P. (1999) Molecular evolution and systematics of owls (Strigiformes). In: König, C., Weick, F.E. & Becking, J.H. (Eds.), Owls: A Guide to the Owls of the World. Yale University Press, New Haven, pp. 39‒57.
Wink, M. & Heidrich, P. (2000) Molecular systematics of owls (Strigiformes) based on DNA-sequences of the mitochondrial cytochrome b gene. In: Chancelor, R.D. & Meyburg, B.U. (Eds.), Raptors at Risk. WWGBP, Hancock House, Surrey, British Columbia, pp. 819‒828.
Yang, Z. & Rannala, B. (2010) Bayesian species delimitation using multilocus sequence data. Proceedings of National Academy of Sciences USA, 107, 9264–9269.
https://doi.org/10.1073/pnas.0913022107