Abstract
Recent DNA-based phylogenetic analyses of the family Thamnophilidae have shown that the genus Myrmotherula is polyphyletic. Traditional plumage-based taxonomy has been misleading in terms of identifying independently evolving lineages within the complex. Here, we integrate a molecular phylogeny with morphometric information and ancestral reconstruction of syringeal character states of the Musculi vocales ventrales, to investigate the taxonomic position of M. gularis, a species for which phylogenetic affinities have long been uncertain. We show that M. gularis represents a long branch in the tribe Thamnophilini that is not closely related to any other member of the Myrmotherula complex. Its relationships within the tribe remain uncertain because of the lack of phylogenetic resolution at the base of the tribe. M. gularis shares a derived character state of the M. vocalis ventralis with Taraba, Hypoedaleus, and Mackenziaena, which supports a close relationship between M. gularis and the large antshrikes. M. gularis can be diagnosed from Myrmotherula and Epinecrophylla by this condition of its M. vocalis ventralis, and from Isleria by plumage and other morphological traits. The phylogenetic and morphological distinctiveness of M. gularis does not warrant merging it into any other genus. We propose that this species be placed in a monotypic genus, for which the available name Rhopias applies.
References
Akaike, H. (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control, 19, 716–723.
Ames, P.L. (1971) The morphology of the syrinx in passerine birds. Peabody Museum Natural History Bulletin, 37, 1–194.
Barker, F.K., Barrowclough, G.F. & Groth, J.G. (2002) A phylogenetic hypothesis for passerine birds: taxonomic and biogeographic implications of an analysis of nuclear DNA sequence data. Proceedings of the Royal Society B-Biological Sciences, 269, 295–308.
Bornschein, M.R., Reinert, B.L. & Teixeira, D.M. (1995) Um novo Formicariidae do Sul do Brasil (Aves, Passeriformes). Série Técnico-Científica do Instituto Iguaçu de Pesquisa e Preservação Ambiental 1, Rio de Janeiro, 18pp.
Bravo, G.A., Chesser, R.T. & Brumfield, R.T. (2012) Isleria, a new genus of antwren (Aves: Passeriformes: Thamnophilidae). Zootaxa, 3195, 61–67.
Brumfield, R.T. & Edwards, S.V. (2007) Evolution into and out of the Andes: a Bayesian analysis of historical diversification in Thamnophilus antshrikes. Evolution, 61, 346–367.
Brumfield, R.T., Tello, J.G., Cheviron, Z.A., Carling, M.D., Crochet, N. & Rosenberg, K.V. (2007) Phylogenetic conservatism and antiquity of a tropical specialization: Army-ant-following in the typical antbirds (Thamnophilidae). Molecular Phylogenetics and Evolution, 45, 1–13.
Cabanis, J. & Heine, F. (1859–1860) Verzeichniss der ornithologischen Sammlung des Oberamtmann Ferdinand Heine auf Gut St. Burchard vor Halberstadt. Mit kritischen Anmerkungen und Beschreinbung der neuen arten systematisch bearbeitet. Museum Heineanum, 2, 1–175.
Cory, C.B. & Hellmayr, C.E. (1924) Catalogue of the birds of the Americas, part III: Pteroptochidae – Conopophagidae – Formicariidae. Field Museum of Natural History Publication Zoological Series, 223(XII), 369pp.
Develey, P.F. & Peres, C.A. (2000) Resource seasonality and the structure of mixed species bird flocks in a coastal Atlantic Forest of southeastern Brazil. Journal of Tropical Ecology, 16, 33–53.
Develey, P.F. & Stouffer, P.C. (2001) Effects of roads on movements by understory birds in mixed-species flocks in central Amazonian Brazil. Conservation Biology, 15, 1416–1422.
Galvão, A. & Gonzaga, L.P. (2011) Morphological support for placement of the Wing-banded Antbird Myrmornis torquata in the Thamnophilidae (Passeriformes: Furnariides). Zootaxa, 3122, 37–67.
Gómez, J.P., Bravo, G.A., Brumfield, R.T., Tello, J.G. & Cadena, C.D. (2010) A phylogenetic approach to disentangling the role of competition and habitat filtering in community assembly of Neotropical forest birds. Journal of Animal Ecology, 79, 1181–1192.
Gonzaga, L.A.P. (2001) Análise filogenética do gênero Formicivora Swainson, 1825 (Aves: Passeriformes: Thamnophilidae) baseada em caracteres morfológicos e vocais. PhD Dissertation. Universidade Federal do Rio de Janeiro, Rio de Janeiro, 215 pp.
Groth, J.G. & Barrowclough, G.F. (1999) Basal divergence in birds and phylogenetic utility of the nuclear RAG-1 gene. Molecular Phylogenetics and Evolution, 12, 115–123.
Hackett, S.J. & Rosenberg, K.V. (1990) Comparison of the phenotypic and genetic differentiation in South American antwrens (Formicariidae). The Auk, 107, 473–489.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
Irestedt, M., Fjeldså, J., Nylander, J.A.A. & Ericson, P.G.P. (2004) Phylogenetic relationships of typical antbirds (Thamnophilidae) and test of incongruence based on Bayes factors. BMC Evolutionary Biology, 4, 23.
Isler, M.L., Lacerda, D.R., Isler, P.R., Hackett, S.J., Rosenberg, K.V. & Brumfield, R.T. (2006) Epinecrophylla, a new genus of antwrens (Aves: Passeriformes: Thamnophilidae). Proceedings of the Biological Society of Washington, 119, 522–527.
Katoh, K., Misawa, K., Kuma, K-I. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066.
Kumar, S., Skjæveland, Å., Orr, R.J.S., Enger, P., Ruden, T., Mevik, B-H., Burki, F., Botnen, A. & Shalchian-Tabrizi, K. (2009) AIR: A batch-oriented web program package for construction of supermatrices ready for phylogenomic analyses. BMC Bioinformatics 2009, 10, 357.
Lewis, P.O. (2001) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology, 50, 913–925.
Maddison, W.P. & Maddison, D.R. (2000) MacClade 4.0. Sinauer Associates, Sunderland, 492p.
Maddison, W.P. & Maddison, D.R. (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75. Available from http://mesquiteproject.org (accessed 17 October 2011).
Meyer de Schauensee, R. (1966) The species of birds of South America and their distribution. Livingston Publishing Co. Narberth, Pennsylvania, 578 pp.
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Trees. In: Proceedings of the Gateway Computer Environments Workshop, New Orleans, LA, pp. 1–8.
Moyle, R.G., Chesser, R.T., Brumfield, R.T., Tello, J.G., Marchese, D.J. & Cracraft, J. (2009) Phylogeny and phylogenetic classification of the antbirds, ovenbirds, woodcreepers, and allies (Aves: Passeriformes: infraorder Furnariides). Cladistics, 25, 1–20.
Munn, C.A. & Terborgh, J.W. (1979) Multi-species territoriality in Neotropical foraging flocks. The Condor, 81, 338–347.
Nylander, J.A.A. (2004) MrModeltest version 2. Evolutionary Biology Center, Uppsala University, Uppsala, Sweden. Available from http://www.abc.se/~nylander/ (accessed 27 August 2011).
Nylander, J.A.A., Wilgenbusch, J.C., Warren, D.L. & Swofford, D.L. (2008) AWTY (are we there yet?): a system for graphical exploration of mcmc convergence in Bayesian phylogenetics. Bioinformatics, 24, 581–583.
Pearson, D.L. (1977) Ecological relationships of small antbirds in Amazonian bird communities. The Auk, 94, 283–292.
Peters, J.L. (1951) Check-list of the birds of the world: vol. VII. Museum of Comparative Zoology, Cambridge, 318 pp.
Prum, R.O. (1990) A test of the monophyly of the manakins (Pipridae) and of the cotingas (Cotingidae) based on morphology. Occasional Papers of the Museum of Zoology, University of Michigan, 723, 1–44.
R Development Core Team. (2011) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. Available from http://www.R-project.org (accessed 27 August 2011).
Ridgley, R.S. & Tudor, G. (1994) The birds of South America. Vol. II. The Suboscine Passerines. University of Texas Press, Austin, 814 pp.
Sclater, P.L. (1858) Synopsis of the American antbirds (Formicariidae). Part II., containing the Formicivorinae or ant-wrens. Proceedings of the Zoological Society of London, 26, 232–272.
Sibley, C. G., & Monroe, B.L. Jr. (1990) Distribution and taxonomy of birds of the World. Yale University Press. New Haven, Connecticut, 1117 pp.
Sick, H. (1997) Ornitologia brasileira. Editora Nova Fronteira, Rio de Janeiro, 862 pp.
Skutch, A.F. (1946) Life histories of two Panamanian antbirds. The Condor, 48, 16–28.
Skutch, A.F. (1969) Life histories of Central American birds, part III. Pacific Coast Avifauna, 35, 7–550.
Spix, J.B.V. (1825) Avium species novae, quas in itinere per Brasiliam annis MDCCCXVII- MDCCCXX jussu et auspiciis Maximiliani Josephi I Bavariae Regis suscepto collegit et descripsit. Tomo II. Typis Francisci Seraphi Hybschmanni, Munich, 85 pp.
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
Stotz, D.F. (1990) Foraging behavior and morphology in the avian genus Myrmotherula. PhD dissertation. University of Chicago, Illinois, 258 pp.
Stotz, D.F., Fitzpatrick, J.W., Parker III, T.A. & Moskovits, D.K. (1996) Neotropical birds: Ecology and conservation. University of Chicago Press, Chicago and London, 469 pp.
Swofford, D.L. (2003) PAUP*. Phylogenetic analysis using parsimony (*and other methods). Sunderland, Sinauer Associates, CD-ROM.
Swofford, D.L. & Maddison, W.P. (1987) Reconstructing ancestral character states under Wagner parsimony. Mathematical Biosciences, 87, 199–229.
Whitney, B.M. (1994) Behavior, vocalizations, and possible relationships of four Myrmotherula antwrens (Formicariidae) from eastern Ecuador. The Auk, 111, 469–475.
Wilgenbusch, J.C., Warren, D.L. & Swofford, D.L. (2004) AWTY: A system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. Available from http://ceb.csit.fsu.edu/awty (accessed 27 August 2011).
Zimmer, K.J. & Isler, M.L. (2003) Family Thamnophilidae (Typical Antbirds). In: Del Hoyo, J.; Elliot, A. & Christie, D.A. (Eds.), Handbook of the Birds of the World, Vol. 8. Broadbills to Tapaculos. Lynx Edicions, Barcelona, pp. 448–681.