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Using microsatellite DNA variation, morphological measurements and sonagrams made from tape-recordings in the field, we examine the allopatric differentiation of six populations of the sharp-beaked ground finch, Geospiza difficilis , in the Galápagos archipelago. We ask how and why the populations became differentiated, and consider what the differences imply about speciation. An important factor is time G. difficilis is one of the phylogenetically oldest species. Populations became differentiated by adapting in beak morphology to different food supplies. Since beak size and shape also function in conspecific mate recognition and choice, the potential for reproductive isolation from sister and parental taxa arose as a correlated effect of natural selection on resource-exploiting traits. This conforms to a standard explanation for the origin of pre-mating isolation as a byproduct of adaptive changes in allopatry without being adaptive itself. However, this explanation is incomplete. Vocal characteristics also differentiated, partly as a result of natural and sexual selection independent of beak size change in environments with different sound transmitting properties. An additional element of chance is indicated by a comparison of closely-related populations on two islands, Wolf and Darwin, that are geographically close, and topographically and floristically similar. The populations have markedly different vocalizations. Morphological characters, vocalizations and genetic traits do not vary concordantly among the six populations. One population (Genovesa) is genetically more similar to other congeneric species. Phenotypic similarities with G. difficilis are the result of a uniquely long retention of shared ancestral traits. Arguments under the phylogenetic species concept justify recognizing this population as a separate species, Geospiza acutirostris. Under the biological species concept it should remain as currently classified, G. difficilis .

The allopatric phase of speciation: the sharp-beaked ground finch (Geospiza difficilis) on the Galápagos islands