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Article
Published: 2021-03-26

Multi-character taxonomic review, systematics, and biogeography of the Black-capped/Tawny-bellied Screech Owl (Megascops atricapilla-M. watsonii) complex (Aves: Strigidae)

1Zoology Graduate Program, Universidade Federal do Pará/Museu Paraense Emilio Goeldi, Belém-PA, Brazil. Department of Zoology, Museu Paraense Emílio Goeldi, Belém-PA, Brazil.
Department of Ornithology, Academy of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA, USA 19096. Department of Biodiversity, Earth and Environmental Science, Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA, USA 19096.
Negaunee Integrative Research Center, Field Museum of Natural History, 1400 S Lake Shore Drive Chicago, IL, USA 60605.
Department of Zoology, Museu Paraense Emílio Goeldi, Belém-PA, Brazil. Graduate Program in Environmental Biology, Universidade Federal do Pará, Bragança-PA, Brazil.
Department of Biodiversity, Earth and Environmental Science, Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA, USA 19096.
Department of Zoology, Museu Paraense Emílio Goeldi, Belém-PA, Brazil. Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland.
Aves Bioacoustics cryptic speciation morphometry plumage variation population genetics

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

  1. 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

How to Cite

DANTAS, S. M., WECKSTEIN, J. D., BATES, J., OLIVEIRA, J. N., CATANACH, T. A., & ALEIXO, A. (2021). Multi-character taxonomic review, systematics, and biogeography of the Black-capped/Tawny-bellied Screech Owl (<em>Megascops</em> <em>atricapilla</em>-<em>M. watsonii</em>) complex (Aves: Strigidae). Zootaxa, 4949(3), 401–444. https://doi.org/10.11646/zootaxa.4949.3.1