Evolutionary Ecology

, Volume 28, Issue 2, pp 299–322 | Cite as

Ecology, song similarity and phylogeny predict natural hybridization in an avian family

  • Pamela M. Willis
  • Rebecca E. Symula
  • Irby J. Lovette
Original Paper


Hybridization is important in the evolution of many animal groups; however, broad scale patterns of natural hybridization are still poorly understood. Using phylogenetic comparative analyses, we tested for relationships between demographic, ecological, phenotypic and phylogenetic variables and the incidence of natural hybridization among 45 species of North American wood warbler. Since 1980, hybrids have been documented in 24 species (53 %). We detected negative relationships between the incidence of hybridization and both breeding range size and phylogenetic distance, and positive relationships between the incidence of hybridization and (1) song similarity, (2) the extent of breeding sympatry of species pairs and (3) the number of additional heterospecific warbler species that co-occur during breeding. Neither population size nor breeding habitat quality (as measured by threats to survival or reproduction in breeding areas) explained variation in the incidence of hybridization. Our results suggest a potential role of limited breeding habitat in wood warbler hybridization events and a net positive effect of breeding sympatry. They also support the hypothesis that mating signal similarity facilitates hybridization events and are consistent with an increase in reproductive isolation with increasing genetic divergence. Our results also suggest the presence of phylogenetic signal in wood warbler hybridization. By investigating natural hybridization at the taxonomic level of family, we have identified several previously undocumented patterns of natural hybridization. This study demonstrates the utility of examining the combined effects of ecology, demography, phenotype and phylogeny when studying variation in the expression of natural hybridization among taxa.


Hybridization Mate choice Breeding habitat Warblers Song Phylogenetic signal 



We thank Luke Harmon, Richard Glor, and Pierre Legendre for R code and programming assistance. Thanks to John Sullivan for phylogeny preparation and advice, and Wesley Hochachka for providing regional co-occurrence data and help with its interpretation. Darren Irwin and three anonymous reviewers provided very helpful feedback on earlier versions of the manuscript. P.M.W. was funded by a Zoology Scholarship Endowment for Excellence Fellowship.

Supplementary material

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Supplementary material 1 (PDF 3225 kb)


  1. Abbott R, Albach D, Ansell S, Arntzen J, Baird S, Bierne N, Boughman J, Brelsford A, Buerkle C, Buggs R (2013) Hybridization and speciation. J Evol Biol 26:229–246PubMedGoogle Scholar
  2. Agresti A (2007) An introduction to categorical data analysis, 2nd edn. Wiley, HobokenGoogle Scholar
  3. Alatalo RV, Lundberg A, Glynn C (1986) Female pied flycatchers choose territory quality and not male characteristics. Nature 323:152–153Google Scholar
  4. American Ornithologists’ Union (1998) Check-list of North American birds, 7th edn. American Ornithologist’s Union, Washington, DCGoogle Scholar
  5. Anich NM, Swelstad JA, Pierce B (2012) The first report of a Sutton’s warbler (Northern Parula x Yellow-throated warbler) in Wisconsin. Passeng Pigeon 74:149–155Google Scholar
  6. Arvidsson BL, Neergaard R (1991) Mate choice in the willow warbler—a field experiment. Behav Ecol Sociobiol 29(3):225–229Google Scholar
  7. Bain M (1996) A mystery warbler in southern Ontario. Birders J 5:134–135Google Scholar
  8. Baker MC, Boylan JT (1999) Singing behavior, mating associations and reproductive success in a population of hybridizing Lazuli and Indigo buntings. Condor 101:493–504Google Scholar
  9. Bank C, Hermisson J, Kirkpatrick M (2012) Can reinforcement complete speciation? Evolution 66:229–239PubMedGoogle Scholar
  10. Bart J, Earnst SL (1999) Relative importance of male and territory quality in pairing success of male rock ptarmigan (Lagopus mutus). Behav Ecol Sociobiol 45:355–359Google Scholar
  11. Bayne EM, Hobson KA (2001) Effects of habitat fragmentation on pairing success of ovenbirds: importance of male age and floater behavior. Auk 118:380–388Google Scholar
  12. Blancher P, Rosenberg K, Panjabi A, Altman B, Bart J, Beardmore C, Butcher G, Demarest D, Dettmers R, Dunn E (2007) Guide to the Partners in Flight Population Estimates Database. Version: North American Landbird Conservation Plan 2004. Partners in Flight Technical Series No 5. Accessed Apr 2013
  13. Bolnick D (2009) Hybridization and speciation in centrarchids. In: Cooke SJ, Philipp DP (eds) Centrarchid fishes: diversity, biology and conservation. Wiley-Blackwell, Oxford, pp 39–69Google Scholar
  14. Bolnick DI, Near TJ (2005) Tempo of hybrid inviability in centrarchid fishes (Teleostei: Centrarchidae). Evolution 59:1754–1767PubMedGoogle Scholar
  15. Bonter D, Lovette IJ (2007) Name that warbler. Living Bird 26:32–35Google Scholar
  16. Boves TJ, Buehler DA, Massey PC (2010) Interspecific song imitation by a Cerulean Warbler. Wilson J Ornithol 122(3):583–587Google Scholar
  17. Brelsford A, Irwin DE (2009) Incipient speciation despite little assortative mating: the yellow-rumped warbler hybrid zone. Evolution 63:3050–3060PubMedGoogle Scholar
  18. Butlin R (1989) Reinforcement of premating isolation. In: Otte D, Endler JA (eds) Speciation and its consequences. Sinauer Associates, Sunderland, pp 158–179Google Scholar
  19. Byers BE (1995) Song types, repertoires and song variability in a population of chestnut-sided warblers. Condor 97:390–401Google Scholar
  20. Byers BE (1996) Geographic variation of song form within and among Chestnut-sided Warbler populations. Auk 113:288–299Google Scholar
  21. Byers BE (2007) Extrapair paternity in chestnut-sided warblers is correlated with consistent vocal performance. Behav Ecol 18:130–136Google Scholar
  22. Carlson CW (1981) The Sutton’s Warbler—a critical review and summation of current data. Atl Nat 34:1–11Google Scholar
  23. Chinn S (2000) A simple method for converting an odds ratio to effect size for use in meta-analysis. Stat Med 19:3127–3131PubMedGoogle Scholar
  24. Chiver I, Stutchbury BJ, Morton ES (2008) Do male plumage and song characteristics influence female off-territory forays and paternity in the hooded warbler? Behav Ecol Sociobiol 62:1981–1990Google Scholar
  25. Clark CW, Marler P, Beeman K (1987) Quantitative analysis of animal vocal phonology: an application to swamp sparrow song. Ethology 76:101–115Google Scholar
  26. Coleman SW, Patricelli GL, Coyle B, Siani J, Borgia G (2007) Female preferences drive the evolution of mimetic accuracy in male sexual displays. Biol Lett 3:463–466PubMedCentralPubMedGoogle Scholar
  27. Confer JL (2006) Secondary contact and introgression of Golden-winged warblers (Vermivora chrysoptera): documenting the mechanism. Auk 123:958–961Google Scholar
  28. Confer JL, Tupper SK (2000) A reassessment of the status of Golden-winged and Blue-winged warblers in the Hudson Highlands of southern New York. Wilson Bull 112:544–546Google Scholar
  29. Cortopassi KA, Bradbury JW (2000) The comparison of harmonically rich sounds using spectrographic cross-correlation and principal coordinates analysis. Bioacoustics 11:89–127Google Scholar
  30. Coyne JA, Orr HA (1989) Patterns of speciation in Drosophila. Evolution 43:362–381Google Scholar
  31. Coyne JA, Orr HA (2004) Speciation. Sunderland Associates Inc., SunderlandGoogle Scholar
  32. Coyne JA, Kim SY, Chang AS, Lachaise D, Elwyn S (2002) Sexual isolation between two sibling species with overlapping ranges: Drosophila santomea and Drosophila yakuba. Evolution 56:2424–2434PubMedGoogle Scholar
  33. Dale S (2001) Female-biased dispersal, low female recruitment, unpaired males, and the extinction of small and isolated bird populations. Oikos 92:344–356Google Scholar
  34. de Kort SR, den Hartog PM, ten Cate C (2002) Vocal signals, isolation and hybridization in the vinaceous dove (Streptopelia vinacea) and the ring-necked dove (S. capicola). Behav Ecol Sociobiol 51:378–385Google Scholar
  35. Detwiler KM, Burrell AS, Jolly CJ (2005) Conservation implications of hybridization in African cercopithecine monkeys. Int J Primatol 26:661–684Google Scholar
  36. Dobzhansky T (1937) Genetics and the origin of species, 1st edn. Columbia University Press, New YorkGoogle Scholar
  37. Dowling TE, Secor CL (1997) The role of hybridization and introgression in the diversification of animals. Annu Rev Ecol Syst 28:593–619Google Scholar
  38. Ducharme C, Lamontagne J (1992) Un cas d’hybridation chez les parulines? Dendroica caerulescens x Dendroica petechia. Quebec Oiseaux 4:6Google Scholar
  39. Dunn J, Garrett K (1997) A field guide to the warblers of North America. Houghton Mifflin, BostonGoogle Scholar
  40. Dunn PO, Whittingham LA, Freeman-Gallant CR, DeCoste J (2008) Geographic variation in the function of ornaments in the common yellowthroat Geothlypis trichas. J Avian Biol 39:66–72Google Scholar
  41. Eliason BC (1986) Female site fidelity and polygyny in the Blackpoll Warbler (Dendroica striata). Auk 103:782–790Google Scholar
  42. Erdfelder E (1984) Zur bedeutung und kontrolle des β-fehlers bei der inferenzstatistischen prüfung log-linearer modelle (The significance and control of the β-error during the inference-statistical examination of the log-linear models). Z Sozialpsychol 15:18–32Google Scholar
  43. Farnsworth A, Lovette IJ (2008) Phylogenetic and ecological effects on interspecific variation in structurally simple avian vocalizations. Biol J Linn Soc 94:155–173Google Scholar
  44. Faul F, Erdfelder E, Buchner A, Lang A-G (2009) Statistical power analyses using G* Power 3.1: tests for correlation and regression analyses. Behav Res Methods 41:1149–1160PubMedGoogle Scholar
  45. Ficken MS, Ficken RW (1966) Notes on mate and habitat selection in the Yellow Warbler. Wilson Bull 78:232–233Google Scholar
  46. Ficken MS, Ficken RW (1968) Reproductive isolating mechanisms in the blue-winged warbler-golden-winged warbler complex. Evolution 22:166–179Google Scholar
  47. Fitzpatrick BM (2004) Rates of evolution of hybrid inviability in birds and mammals. Evolution 58:1865–1870PubMedGoogle Scholar
  48. Freeman-Gallant CR, Taff CC, Morin DF, Dunn PO, Whittingham LA, Tsang SM (2010) Sexual selection, multiple male ornaments, and age- and condition-dependent signaling in the common yellowthroat. Evolution 64:1007–1017PubMedGoogle Scholar
  49. Fritz SA, Purvis A (2010) Selectivity in mammalian extinction risk and threat types: a new measure of phylogenetic signal strength in binary traits. Conserv Biol 24:1042–1051PubMedGoogle Scholar
  50. Garland T, Dickerman AW, Janis CM, Jones JA (1993) Phylogenetic analysis of covariance by computer simulation. Syst Biol 42:265–292Google Scholar
  51. Gee JM (2003) How a hybrid zone is maintained: behavioral mechanisms of interbreeding between California and Gambel’s quail (Callipepla californica and C. gambelii). Evolution 57:2407–2415PubMedGoogle Scholar
  52. Gerlai R (2007) Mate choice and hybridization in Lake Malawi cichlids, Sciaenochromis fryeri and Cynotilapia afra. Ethology 113:673–685Google Scholar
  53. Gholamhosseini A, Vardakis M, Aliabadian M, Nijman V, Vonk R (2013) Hybridization between sister taxa versus non-sister taxa: a case study in birds. Bird Study (ahead-of-print) 1–7. doi: 10.1080/00063657.2013.770815
  54. Gibbs JP, Faaborg J (1990) Estimating the viability of Ovenbird and Kentucky warbler populations in forest fragments. Conserv Biol 4:193–196Google Scholar
  55. Gill FB (2004) Blue-winged Warblers (Vermivora pinus) versus Golden-winged Warblers (V. chrysoptera). Auk 121:1014–1018Google Scholar
  56. Gill FB, Murray BG (1972a) Discrimination behavior and hybridization of the blue-winged and golden-winged warblers. Evolution 26:282–293Google Scholar
  57. Gill FB, Murray BG (1972b) Song variation in sympatric Blue-winged and Golden-winged Warblers. Auk 89:625–643Google Scholar
  58. Gonzalez J, Düttmann H, Wink M (2009) Phylogenetic relationships based on two mitochondrial genes and hybridization patterns in Anatidae. J Zool 279:310–318Google Scholar
  59. Good P (2005) Permutation, parametric and bootstrap tests of hypotheses. Springer, LondonGoogle Scholar
  60. Grant PR, Grant BR (1997) Hybridization, sexual imprinting, and mate choice. Am Nat 149:1–28Google Scholar
  61. Grant BR, Grant PR (1998) Hybridization and speciation in Darwin’s finches: the role of sexual imprinting on a culturally transmitted trait. In: Howard DJ, Berlocher SH (eds) Endless forms: species and speciation. Oxford University Press, New York, pp 404–422Google Scholar
  62. Griffiths J, Crowley J, Messier W (2008) An intergeneric hybrid warbler (Setophaga x Geothlypis) captured in Big Sur, California. Abstracts of the AOU/COS/SCO 2008 Meeting, Portland, OR, 4–9 Aug 2008
  63. Gröning J, Hochkirch A (2008) Reproductive interference between animal species. Q Rev Biol 83:257–282PubMedGoogle Scholar
  64. Hadfield JD (2010) MCMC methods for multi-response generalized linear mixed models: the MCMCglmm R package. J Stat Softw 33:1–22Google Scholar
  65. Hadfield JD (2012) MCMCglmm course notes. Available at Accessed Apr 2013
  66. Hadfield JD, Nakagawa S (2010) General quantitative genetic methods for comparative biology: phylogenies, taxonomies and multi-trait models for continuous and categorical characters. J Evol Biol 23:494–508PubMedGoogle Scholar
  67. Harmon LJ, Glor RE (2010) Poor statistical performance of the Mantel test in phylogenetic comparative analyses. Evolution 64:2173–2178PubMedGoogle Scholar
  68. Harper SL, Vallender R, Robertson RJ (2010) Male song variation and female mate choice in the Golden-winged Warbler. Condor 112:105–114Google Scholar
  69. Harvey M, Pagel P (1991) The comparative method in evolutionary biology. Oxford University Press, OxfordGoogle Scholar
  70. Hill GE (2006) Female mate choice for ornamental coloration. In: Hill GE, McGraw KM (eds) Bird coloration, volume 2: function and evolution. Harvard University Press, Cambridge, pp 137–200Google Scholar
  71. Hill GE, McGraw KM (eds) (2006) Bird coloration, volume 2: function and evolution. Harvard University Press, CambridgeGoogle Scholar
  72. Howlett JS, Stutchbury BJ (2003) Determinants of between-season site, territory, and mate fidelity in Hooded Warblers (Wilsonia citrina). Auk 120:457–465Google Scholar
  73. Hsieh FY, Bloch DA, Larsen MD (1998) A simple method of sample size calculation for linear and logistic regression. Stat Med 17:1623–1634PubMedGoogle Scholar
  74. Hubbs CL (1955) Hybridization between fish species in nature. Syst Zool 4:1–20Google Scholar
  75. Huelsenbeck JP, Rannala B (2003) Detecting correlation between characters in a comparative analysis with uncertain phylogeny. Evolution 57:1237–1247PubMedGoogle Scholar
  76. Hurley G, Jones J II (1983) A presumed mixed Bay-breasted x Blackburnian Warbler nesting in West Virginia. Redstart 50:108–111Google Scholar
  77. Irwin DE, Brelsford A, Toews DP, MacDonald C, Phinney M (2009) Extensive hybridization in a contact zone between MacGillivray’s warblers Oporornis tolmiei and mourning warblers O. philadelphia detected using molecular and morphological analyses. J Avian Biol 40:539–552Google Scholar
  78. Janes SW, Ryker L (2013) Rapid change in a Type I song dialect of hermit warblers (Setophaga occidentalis). Auk 130:30–35Google Scholar
  79. Jouventin P, Bried J (2001) The effect of mate choice on speciation in snow petrels. Anim Behav 62:123–132Google Scholar
  80. Kappes PJ, Stutchbury BJ, Woolfenden BE (2009) The relationship between carotenoid-based coloration and pairing, within-and extra-pair mating success in the American Redstart. Condor 111:684–693Google Scholar
  81. Keller LF, Waller DM (2002) Inbreeding effects in wild populations. Trends Ecol Evol 17:230–241Google Scholar
  82. Kenyon HL, Toews DP, Irwin DE (2011) Can song discriminate between Macgillivray’s and Mourning Warblers in a narrow hybrid zone? Condor 113:655–663Google Scholar
  83. Khanna H, Gaunt SLL, McCallum DA (1997) Digital spectrographic cross-correlation: tests of sensitivity. Bioacoustics 7:209–234Google Scholar
  84. King DI, DeGraaf RM, Griffin CR (2001) Plumage coloration and reproductive success in male chestnut-sided warblers. Wilson Bull 113:239–242Google Scholar
  85. Kroodsma DE (1981) Geographical variation and functions of song types in warblers (Parulidae). Auk 98:743–751Google Scholar
  86. Kroodsma DE, Meservey WR, Pickert R (1983) Vocal learning in the Parulinae. Wilson Bull 95:138–140Google Scholar
  87. Lapointe F-J, Garland T Jr (2001) A generalized permutation model for the analysis of cross-species data. J Classif 18:109–127Google Scholar
  88. Latta SC, Parkes KC (2001) A possible Dendroica kirtlandii hybrid from Hispaniola. Wilson Bull 113:378–383Google Scholar
  89. Latta SC, Parkes KC, Wunderle JM (1998) A new intrageneric Dendroica hybrid from Hispaniola. Auk 115:533–537Google Scholar
  90. Leichty ER, Grier JW (2006) Importance of facial pattern to sexual selection in Golden-winged Warbler (Vermivora chrysoptera). Auk 123:962–966Google Scholar
  91. Lemaire F (1977) Mixed song, interspecific competition and hybridisation in the reed and marsh warblers (Acrocephalus scirpaceus and palustris). Behaviour 63:215–240Google Scholar
  92. Lemon R, Struger J, Lechowicz M (1983) Song features as species discriminants in American warblers (Parulidae). Condor 85:308–322Google Scholar
  93. Lengagne T, Grolet O, Joly P (2006) Male mating speed promote hybridization in the Rana lessonaeRana esculenta waterfrog system. Behav Ecol Sociobiol 60:123–130Google Scholar
  94. Lijtmaer DA, Mahler B, Tubaro PL (2003) Hybridization and postzygotic isolation patterns in pigeons and doves. Evolution 57:1411–1418PubMedGoogle Scholar
  95. Lovette IJ, Bermingham E (1999) Explosive speciation in the New World Dendroica warblers. Proc R Soc Lond B Biol 266:1629–1636Google Scholar
  96. Lovette IJ, Hochachka WM (2006) Simultaneous effects of phylogenetic niche conservatism and competition on avian community structure. Ecology 87(Suppl):S14–S28PubMedGoogle Scholar
  97. Lovette IJ, Pérez-Emán JL, Sullivan JP, Banks RC, Fiorentino I, Córdoba-Córdoba S, Echeverry-Galvis MA, Barker FK, Burns KJ, Klicka J (2010) A comprehensive multilocus phylogeny for the wood-warblers and a revised classification of the Parulidae (Aves). Mol Phylogenet Evol 57:753–770PubMedGoogle Scholar
  98. Mallet J (2005) Hybridization as an invasion of the genome. Trends Ecol Evol 20:229–237PubMedGoogle Scholar
  99. Mallet J (2007) Hybrid speciation. Nature 446(7133):279–283PubMedGoogle Scholar
  100. Mallet J, McMillan WO, Jiggins CD (1998) Mimicry and warning color at the boundary between races and species. In: Howard DJ, Berlocher SH (eds) Endless forms: species and speciation. Oxford University Press, New York, pp 390–403Google Scholar
  101. Marks B, Willard D (1996) Two new intrageneric warbler hybrids. Abstracts of the 114th meeting of the American Ornithologists’ Union, Boise State University, Boise, Idaho, 13–17 Aug 1996Google Scholar
  102. Martin PR, Martin TE (2001) Ecological and fitness consequences of species coexistence: a removal experiment with wood warblers. Ecology 82:189–206Google Scholar
  103. Martin PR, Fotheringham JR, Robertson RJ (1995) A Prairie Warbler with a conspecific and heterospecific song repertoire. Auk 112:770–774Google Scholar
  104. Mayr E (1963) Animal species and evolution. Belknap Press, CambridgeGoogle Scholar
  105. McCarthy E (2006) Handbook of avian hybrids of the world. Oxford University Press, New YorkGoogle Scholar
  106. McCaskie G (1984) Southern Pacific coast region. Am Birds 38:245–248Google Scholar
  107. McNaught MK, Owens IP (2002) Interspecific variation in plumage colour among birds: species recognition or light environment? J Evol Biol 15:505–514Google Scholar
  108. Mendelson TC (2003) Sexual isolation evolves faster than hybrid inviability in a diverse and sexually dimorphic genus of fish (Percidae: Etheostoma). Evolution 57:317–327PubMedGoogle Scholar
  109. Moore JA (1957) An embryologist’s view of the species concept. In: Mayr E (ed) The species problem. American Association for the Advancement of Science, Washington, DC, pp 325–338Google Scholar
  110. Moravec ML, Striedter GF, Burley NT (2006) Assortative pairing based on contact call similarity in budgerigars, Melopsittacus undulatus. Ethology 112:1108–1116Google Scholar
  111. Morse D (1989) American warblers: an ecological and behavioral perspective. Harvard University Press, CambridgeGoogle Scholar
  112. Nagel L, Schluter D (1998) Body size, natural selection, and speciation in sticklebacks. Evolution 52:209–218Google Scholar
  113. Nelson DA (1989) The importance of invariant and distinctive features in species recognition of bird song. Condor 120–130Google Scholar
  114. Neville KJ, Vallender R, Robertson RJ (2008) Nestling sex ratio of golden-winged warblers Vermivora chrysoptera in an introgressed population. J Avian Biol 39:599–604Google Scholar
  115. Nuechterlein GL, Buitron D (1998) Interspecific mate choice by late-courting male western grebes. Behav Ecol 9:313–321Google Scholar
  116. Panhuis TM, Butlin R, Zuk M, Tregenza T (2001) Sexual selection and speciation. Trends Ecol Evol 16:364–371PubMedGoogle Scholar
  117. Panjabi A, Dunn E, Blancher P, Hunter W, Altman B, Bart J, Beardmore C, Berlanga H, Butcher G, Davis S, Demarest DW, Dettmers R, Easton W, Gomez de Silva Garza H, Iñigo-Elias E, Pashley N, Ralph CJ, Rich TD, Rosenberg K, Rustay CM, Ruth JM, Wendt JS, Will TC (2005) The Partners in Flight handbook on species assessment. Version 2005. Rocky Mountain Bird Observatory. Accessed Apr 2013
  118. Parkes KC (1978) Still another Parulid intergeneric hybrid (Mniotilta x Dendroica) and its taxonomic and evolutionary implications. Auk 95:682–690Google Scholar
  119. Parkes KC (1983) Three additional hybrid combinations in North American birds. In: Proceedings of the 101st meeting of the American Ornithologists’ Union, New York, 26–30 Sept 1983Google Scholar
  120. Pearson SF (2000) Behavioral asymmetries in a moving hybrid zone. Behav Ecol 11:84–92Google Scholar
  121. Pfennig KS (2007) Facultative mate choice drives adaptive hybridization. Science 318:965–967PubMedGoogle Scholar
  122. Pierotti R, Annett CA (1993) Hybridization and male parental investment in birds. Condor 95:670–679Google Scholar
  123. Pitocchelli J (1990) Plumage, morphometric, and song variation in Mourning (Oporornis philadelphia) and Macgillivray’s (O. tolmiei) warblers. Auk 107:161–171Google Scholar
  124. Presgraves DC (2002) Patterns of postzygotic isolation in Lepidoptera. Evolution 56:1168–1183PubMedGoogle Scholar
  125. Price T (2008) Speciation in birds. Roberts and Company, Greenwood VillageGoogle Scholar
  126. Price TD, Bouvier MM (2002) The evolution of F-1 postzygotic incompatibilities in birds. Evolution 56:2083–2089PubMedGoogle Scholar
  127. Price T, Lovette IJ, Bermingham E, Gibbs HL, Richman AD (2000) The imprint of history on communities of North American and Asian warblers. Am Nat 156:354–367Google Scholar
  128. Qvarnström A, Haavie J, Sæther SA, Eriksson D, Pärt T (2006) Song similarity predicts hybridization in flycatchers. J Evol Biol 19:1202–1209PubMedGoogle Scholar
  129. Raftery AE, Lewis S (1992) How many iterations in the Gibbs sampler. Bayesian Stat 4:763–773Google Scholar
  130. Randler C (2002) Avian hybridization, mixed pairing and female choice. Anim Behav 63:103–119Google Scholar
  131. Randler C (2004) Frequency of bird hybrids: does detectability make all the difference? J Ornithol 145:123–128Google Scholar
  132. Randler C (2006a) Behavioural and ecological correlates of natural hybridization in birds. Ibis 148:459–467Google Scholar
  133. Randler C (2006b) Extrapair paternity and hybridization in birds. J Avian Biol 37:1–5Google Scholar
  134. Reudink MW, Marra PP, Boag PT, Ratcliffe LM (2009) Plumage coloration predicts paternity and polygyny in the American redstart. Anim Behav 77:495–501Google Scholar
  135. Rhymer JM, Simberloff D (1996) Extinction by hybridization and introgression. Ann Rev Ecol Syst 27:83–109Google Scholar
  136. Rohwer S (1994) Two new hybrid Dendroica warblers and new methodology for inferring parental species. Auk 111:441–449Google Scholar
  137. Rohwer S, Martin PR (2007) Time since contact and gene flow may explain variation in hybrid frequencies among three Dendroica townsendi x D. occidentalis (Parulidae) hybrid zones. Auk 124:1347–1358Google Scholar
  138. Rohwer S, Wood C (1998) Three hybrid zones between Hermit and Townsend’s warblers in Washington and Oregon. Auk 115:284–310Google Scholar
  139. Rohwer S, Wood C, Bermingham E (2000) A new hybrid warbler (Dendroica nigrescens × D. occidentalis) and diagnosis of similar D. townsendi × D. occidentalis recombinants. Condor 102:713–718Google Scholar
  140. Rosenthal G (2013) Individual mating decisions and hybridization. J Evol Biol 26:252–255PubMedGoogle Scholar
  141. Ross L, Hardy NB, Okusu A, Normark BB (2013) Large population size predicts the distribution of asexuality in scale insects. Evolution 67:196–206PubMedGoogle Scholar
  142. Sauer J, Hines J, Fallon J (2008) The North American Breeding Bird Survey Results and Analysis 1966—2007. Version 5.15. USGS Patuxent Wildlife Research Center. Accessed Apr 2013
  143. Secondi J, Bretagnolle V, Compagnon C, Faivre B (2003) Species-specific song convergence in a moving hybrid zone between two passerines. Biol J Linn Soc 80:507–517Google Scholar
  144. Secondi J, Bordas P, Hipsley CA, Bensch S (2011) Bilateral song convergence in a passerine hybrid zone: genetics contribute in one species only. Evol Biol 38:441–452Google Scholar
  145. Seddon N (2005) Ecological adaptation and species recognition drives vocal evolution in neotropical suboscine birds. Evolution 59:200–215PubMedGoogle Scholar
  146. Seehausen O (2004) Hybridization and adaptive radiation. Trends Ecol Evol 19:198–207PubMedGoogle Scholar
  147. Seehausen O, Takimoto G, Roy D, Jokela J (2008) Speciation reversal and biodiversity dynamics with hybridization in changing environments. Mol Ecol 17:30–44PubMedGoogle Scholar
  148. Servedio MR, Noor MAF (2003) The role of reinforcement in speciation: theory and data. Ann Rev Ecol Evol Syst 34:339–364Google Scholar
  149. Sherry TW, Holmes RT (1989) Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment. Behav Ecol Sociobiol 25:327–333Google Scholar
  150. Short LL (1969) Taxonomic aspects of avian hybridization. Auk 86:84–105Google Scholar
  151. Sillett TS, Rodenhouse NL, Holmes RT (2004) Experimentally reducing neighbor density affects reproduction and behavior of a migratory songbird. Ecology 85:2467–2477Google Scholar
  152. Smith CE, Rohwer S (2000) A phenotypic test of Haldane’s rule in an avian hybrid zone. Auk 117:578–585Google Scholar
  153. Spector D (1992) Wood-warbler song systems: a review of paruline singing behaviors. Curr Ornithol 9:199–238Google Scholar
  154. Spiegelhalter DJ, Best NG, Carlin BP, Van Der Linde A (2002) Bayesian measures of model complexity and fit. J Roy Stat Soc B 64:583–639Google Scholar
  155. Stevenson H, Anderson B (1994) Birdlife of Florida. University Press of Florida, GainsvilleGoogle Scholar
  156. Taff CC, Steinberger D, Clark C, Belinsky K, Sacks H, Freeman-Gallant CR, Dunn PO, Whittingham LA (2012) Multimodal sexual selection in a warbler: plumage and song are related to different fitness components. Anim Behav 84:813–821Google Scholar
  157. Thomsen HM, Balsby TJ, Dabelsteen T (2013) Individual variation in the contact calls of the monomorphic Peach-fronted Conure, Aratinga aurea, and its potential role in communication. Bioacoustics (ahead-of-print) 1–13. doi: 10.1080/09524622.2013.779560
  158. Thusius KJ, Peterson KA, Dunn PO, Whittingham LA (2001) Male mask size is correlated with mating success in the common yellowthroat. Anim Behav 62:435–446Google Scholar
  159. Toews DP, Brelsford A, Irwin DE (2011) Hybridization between Townsend’s Dendroica townsendi and black-throated green warblers D. virens in an avian suture zone. J Avian Biol 42:434–446Google Scholar
  160. Ulrich D, Ulrich S (1981) Observations of the 1980 Indiana Sutton’s warbler. Atl Nat 34:12–13Google Scholar
  161. Uy JAC, Moyle RG, Filardi CE (2009) Plumage and song differences mediate species recognition between incipient flycatcher species of the Solomon Islands. Evolution 63:53–164Google Scholar
  162. Vallender R, Friesen VL, Robertson RJ (2007) Paternity and performance of golden-winged warblers (Vermivora chrysoptera) and golden-winged x blue-winged warbler (V. pinus) hybrids at the leading edge of a hybrid zone. Behav Ecol Sociobiol 61:1797–1807Google Scholar
  163. Vallender R, Gagnon J-P, Lovette IJ (2009a) An intergeneric wood-warbler hybrid (Mniotilta varia x Dendroica coronata) and use of multilocus DNA analyses to diagnose avian hybrid origins. Wilson J Ornithol 121:298–305Google Scholar
  164. Vallender R, Van Wilgenburg SL, Bulluck LP, Roth A, Canterbury R, Larkin J, Fowlds RM, Lovette IJ (2009b) Extensive rangewide mitochondrial introgression indicates substantial cryptic hybridization in the Golden-winged Warbler (Vermivora chrysoptera). Avian Conserv Ecol 4:4Google Scholar
  165. Van Buskirk J (1997) Independent evolution of song structure and note structure in American wood warblers. Proc R Soc Lond B Biol 264:755–761Google Scholar
  166. Veen T, Borge T, Griffith SC, Saetre GP, Bures S, Gustafsson L, Sheldon BC (2001) Hybridization and adaptive mate choice in flycatchers. Nature 411:45–50PubMedGoogle Scholar
  167. Villemereuil P, Gimenez O, Doligez B (2012) Comparing parent-offspring regression with frequentist and Bayesian animal models to estimate heritability in wild populations: a simulation study for Gaussian and binary traits. Methods Ecol Evol 4:260–275Google Scholar
  168. Whitney KD, Ahern JR, Campbell LG, Albert LP, King MS (2010) Patterns of hybridization in plants. Perspect Plant Ecol 12:175–182Google Scholar
  169. Wiens JJ, Engstrom TN, Chippindale PT (2006) Rapid diversification, incomplete isolation, and the “speciation clock” in North American salamanders (genus Plethodon): testing the hybrid swarm hypothesis of rapid radiation. Evolution 60:2585–2603PubMedGoogle Scholar
  170. Wiley RH (1991) Associations of song properties with habitats for territorial oscine birds of eastern North America. Am Nat 138:973–993Google Scholar
  171. Wiley C, Fogelberg N, Sæther SA, Veen T, Svedin N, Kehlenbeck JV, Qvarnström A (2007) Direct benefits and costs for hybridizing Ficedula flycatchers. J Evol Biol 20:854–864PubMedGoogle Scholar
  172. Williams J (1996) Nashville warbler (Vermivora ruficapilla). In: Poole A (ed) The Birds of North America Online. Cornell Lab of Ornithology. Accessed Apr 2013
  173. Willis PM (2013) Why do animals hybridize? Acta Ethol 16:127–134Google Scholar
  174. Wirtz P (1999) Mother species-father species: unidirectional hybridization in animals with female choice. Anim Behav 58:1–12PubMedGoogle Scholar
  175. Wright R (2007) Sightings: early May to early June 2007. Birding July/August 30–31Google Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  • Pamela M. Willis
    • 1
    • 4
  • Rebecca E. Symula
    • 1
    • 3
  • Irby J. Lovette
    • 2
  1. 1.Section of Integrative BiologyUniversity of Texas at AustinAustinUSA
  2. 2.Fuller Evolutionary Biology Program, Cornell Lab of OrnithologyCornell UniversityIthacaUSA
  3. 3.Department of BiologyUniversity of MississippiOxfordUSA
  4. 4.Department of BiologyUniversity of VictoriaVictoriaCanada

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