Abstract
Recent research on speciation has identified a central role for ecological divergence, which can initiate speciation when (i) subsets of a species or population evolve to specialize on different ecological resources and (ii) the resulting phenotypic modes become reproductively isolated. Empirical evidence for these two processes working in conjunction, particularly during the early stages of divergence, has been limited. We recently described a population of the medium ground finch, Geospiza fortis, that features large and small beak morphs with relatively few intermediates. As in other Darwin's finches of the Galápagos Islands, these morphs presumably diverged in response to variation in local food availability and inter- or intraspecific competition. We here demonstrate that the two morphs show strong positive assortative pairing, a pattern that holds over three breeding seasons and during both dry and wet conditions. We also document restrictions on gene flow between the morphs, as revealed by genetic variation at 10 microsatellite loci. Our results provide strong support for the central role of ecology during the early stages of adaptive radiation.
References
- Belkhir, K., Chikhi, L., Raufaste, N. & Bonhomme, F. 2004 Genetixlogiciel sous WindowsTM pour la génétique des populations, 4.05. Montpellier, France: Laboratoire Génome, Populations, Interactions CNRS UMR 5000, Université de Montpellier II. Google Scholar
Boag P.T& Grant P.R . 1981Intense natural selection in a population of Darwin's finches (Geospizinae) in the Galápagos. Science. 214, 82–85.doi:10.1126/science.214.4516.82. . Crossref, PubMed, ISI, Google ScholarGavrilets S Fitness landscapes and the origin of species. 2004Princeton, NJ:Princeton University Press. Google ScholarGislason D, Ferguson M.M, Skulason S& Snorrason S.S . 1999Rapid and coupled phenotypic and genetic divergence in Icelandic Arctic char (Salvelinus alpinus). Can. J. Fish. Aquat. Sci. 56, 2229–2234.doi:10.1139/cjfas-56-12-2229. . Crossref, ISI, Google ScholarGrant P.R Ecology and evolution of Darwin's finches. 1999Princeton, NJ:Princeton University Press. Google ScholarGrant B.R& Grant P.R . 1996High survival of Darwin's finch hybrids: effects of beak morphology and diets. Ecology. 77, 500–509.doi:10.2307/2265625. . Crossref, ISI, Google ScholarGrant P.R& Grant B.R . 1997Mating patterns of Darwin's finch hybrids determined by song and morphology. Biol. J. Linn. Soc. 60, 317–343.doi:10.1006/bijl.1996.0103. . Crossref, ISI, Google ScholarGrant P.R& Grant B.R . 2002Unpredictable evolution in a 30-year study of Darwin's finches. Science. 296, 707–711.doi:10.1126/science.1070315. . Crossref, PubMed, ISI, Google ScholarGrant P.R& Grant B.R . 2006Evolution of character displacement in Darwin's finches. Science. 313, 224–226.doi:10.1126/science.1128374. . Crossref, PubMed, ISI, Google ScholarGrant P.R, Abbot I, Schluter D, Curry R.L& Abbott L.K . 1985Variation in the size and shape of Darwin's finches. Biol. J. Linn. Soc. 25, 1–39. Crossref, ISI, Google ScholarGrant P.R, Grant B.R& Petren K . 2000The allopatric phase of speciation: the sharpbeaked ground finch (Geospiza difficilis) on the Galápagos islands. Biol. J. Linn. Soc. 69, 287–317.doi:10.1006/bijl.1999.0382. . Crossref, ISI, Google ScholarHendry A.P, Grant P.R, Grant B.R, Ford H.A, Brewer M.J& Podos J . 2006Possible human impacts on adaptive radiation: beak size bimodality in Darwin's finches. Proc. R. Soc. B. 273, 1887–1894.doi:10.1098/rspb.2006.3534. . Link, ISI, Google ScholarHerrel A, Podos J, Huber S.K& Hendry A.P . 2005Bite performance and morphology in a population of Darwin's finches: implications for the evolution of beak shape. Funct. Ecol. 19, 43–48.doi:10.1111/j.0269-8463.2005.00923.x. . Crossref, ISI, Google ScholarHigashi M, Takimoto G& Yamamura N . 1999Sympatric speciation by sexual selection. Nature. 402, 523–526.doi:10.1038/990087. . Crossref, PubMed, ISI, Google ScholarHuber S.K& Podos J . 2006Beak morphology and song features covary in a population of Darwin's finches (Geospiza fortis). Biol. J. Linn. Soc. 88, 489–498.doi:10.1111/j.1095-8312.2006.00638.x. . Crossref, ISI, Google ScholarKeller L.F, Grant P.R, Grant B.R& Petren K . 2001Heritability of morphological traits in Darwin's finches: misidentified paternity and maternal effects. Heredity. 87, 325–336.doi:10.1046/j.1365-2540.2001.00900.x. . Crossref, PubMed, ISI, Google ScholarKondrashov A.S& Kondrashov F.A . 1999Interactions among quantitative traits in the course of sympatric speciation. Nature. 400, 351–354.doi:10.1038/22514. . Crossref, PubMed, ISI, Google ScholarMallet J . 2007Hybrid speciation. Nature. 446, 279–283.doi:10.1038/nature05706. . Crossref, PubMed, ISI, Google ScholarOrr M.R& Smith T.B . 1998Ecology and speciation. Trends Ecol. Evol. 13, 502–506.doi:10.1016/S0169-5347(98)01511-0. . Crossref, PubMed, ISI, Google ScholarPetren K . 1998Microsatellite primers from Geospiza fortis and cross-species amplification in Darwin's finches. Mol. Ecol. 7, 1782–1784. Crossref, PubMed, ISI, Google ScholarPodos J . 2001Correlated evolution of morphology and vocal signal structure in Darwin's finches. Nature. 409, 185–188.doi:10.1038/35051570. . Crossref, PubMed, ISI, Google ScholarPodos J& Hendry A.P The biomechanics of ecological speciation. Ecology and biomechanics: a mechanical approach to the ecology of animals and plants, Herrel A, Speck T& Rowe N . 2006pp. 301–321. Eds. Boca Raton, FL:CRC Press. Crossref, Google ScholarPodos J& Nowicki S . 2004Beaks, adaptation, and vocal evolution in Darwin's finches. Bioscience. 54, 501–510.doi:10.1641/0006-3568(2004)054[0501:BAAVEI]2.0.CO;2. . Crossref, ISI, Google ScholarPodos J, Southall J.A& Rossi-Santos M . 2004Vocal mechanics in Darwin's finches: correlation of beak gape and song frequency. J. Exp. Biol. 207, 607–619.doi:10.1242/jeb.00770. . Crossref, PubMed, ISI, Google ScholarPrice T . 1987Diet variation in a population of Darwin's finches. Ecology. 68, 1015–1028.doi:10.2307/1938373. . Crossref, ISI, Google ScholarPritchard J.K, Stephens M& Donnelly P . 2000Inference of population structure using multilocus genotype data. Genetics. 155, 945–959. Crossref, PubMed, ISI, Google ScholarRatcliffe L.M& Grant P.R . 1983Species recognition in Darwin's finches (Geospiza, Gould). I. Discrimination by morphological cues. Anim. Behav. 31, 1139–1153.doi:10.1016/S0003-3472(83)80021-9. . Crossref, ISI, Google ScholarRaymond M& Rousset F . 1995Genepop version 1.2: population genetics software for exact tests and ecumenicism. J. Hered. 86, 248–249. Crossref, ISI, Google ScholarRueffler C, Van Dooren T.J.M, Leimar O& Abrams P.A . 2006Disruptive selection and then what?. Trends Ecol. Evol. 21, 238–245.doi:10.1016/j.tree.2006.03.003. . Crossref, PubMed, ISI, Google ScholarRundle H.D& Nosil P . 2005Ecological speciation. Ecol. Lett. 8, 336–352.doi:10.1111/j.1461-0248.2004.00715.x. . Crossref, ISI, Google ScholarRyan P.G, Bloomer P, Moloney C.L, Grant T.J& Delport W . 2007Ecological speciation in South Atlantic island finches. Science. 315, 1420–1423.doi:10.1126/science.1138829. . Crossref, PubMed, ISI, Google ScholarSambrook J, Fritsch E.F& Maniatis T Molecular cloning: a laboratory manual. 2nd edn.1989New York, NY:Cold Spring Harbor Laboratory Press. Google ScholarSeehausen O . 2004Hybridization and adaptive radiation. Trends Ecol. Evol. 19, 198–207.doi:10.1016/j.tree.2004.01.003. . Crossref, PubMed, ISI, Google ScholarSlatkin M . 1995A measure of population subdivision based on microsatellite allele frequencies. Genetics. 139, 457–462. Crossref, PubMed, ISI, Google ScholarSmith T.B . 1993Disruptive selection and the genetic basis of bill size polymorphism in the African finch Pyrenestes. Nature. 363, 618–620.doi:10.1038/363618a0. . Crossref, ISI, Google ScholarSmith T.B& Skulason S . 1996Evolutionary significance of resource polymorphisms in fishes, amphibians, and birds. Annu. Rev. Ecol. Syst. 27, 111–133.doi:10.1146/annurev.ecolsys.27.1.111. . Crossref, Google ScholarWeir B.S& Cockerham C.C . 1984Estimating F-statistics for the analysis of population structure. Evolution. 38, 1358–1370.doi:10.2307/2408641. . PubMed, ISI, Google Scholar



