The spectacular evolutionary radiation of hummingbirds (Trochilidae) has served as a model system for many biological studies. To begin to provide a historical context for these investigations, we generated a complete matrix of DNA hybridization distances among 26 hummingbirds and an outgroup swift (Chaeturu pelagica) to determine the principal hummingbird lineages. FITCH topologies estimated from symmetrized AT,H-C values and subjected to various validation methods (bootstrapping, weighted jackknifing, branch length significance) indicated a fundamental split between hermit (Eutoxeres uquilu, Threnetes ruckeri; Phaethornithinae) and nonhermit (Trochilinae) hummingbirds, and provided strong support for six principal nonhermit clades with the following branching order: (1) analyses on a matrix of unsymmetrized A values gave similar support for these relationships except that the branching order of the two Andean clades (2, 3 above) was unresolved. In general, subsidiary relationships were consistent and well supported by both matrices, sometimes revealing surprising associations between forms that differ dramatically in plumage and bill morphology. Our results also reveal some basic aspects of hummingbird ecologic and morphologic evolution. For example, most of the diverse endemic Andean assemblage apparently comprises two genetically divergent clades, whereas the majority of North American hummingbirds belong a single third clade. Genetic distances separating some morphologically distinct genera (Oreotrochilus, Agluiocercus, Lesbiu; Myrtis, Acestruru, Philodice) were no greater than among congeneric (Coeligenu) species, indicating that, in hummingbirds, morphological divergence does not necessarily reflect level of genetic divergence.

Ecological studies of hummingbird communities have emphasized the importance of local conditions and contemporary interactions in the development of these varied faunas. A timecalibrated, DNA hybridization-based phylogeny of the principal hummingbird lineages was used to examine historical aspects of hummingbird faunas in the species-rich tropical lowlands and Andes, and the relatively depauperate West Indies and temperate regions of Central and North America. Parsimony reconstructions of ancestral distributions indicate that these faunas are polyphyletic in origin, comprising several to many independent lineages. Based on the timing of geologic and cladogenic events, hummingbird faunas appear to have arisen more often by colonization than by large-scale vicariance, with multiple dispersals across water gaps, elevational gradients, and latitude. The extent to which particular lineages colonized different regions depended, however, on lineage ecology as well as on the habitat and age of the fauna. In general, the oldest extant trochilofauna, which today occupies the tropical lowlands, was the principal source of colonizing taxa. However, all regions except possibly the West Indies contributed taxa now found elsewhere, including in the tropical lowlands. The Andean fauna comprises several lineages with lowland origin (hermits, Mangoes, Brilliants, Coquettes, Emeralds) as well as at least one that arose in temperate regions outside South America (Bees). At least two lineages that colonized the West Indies gave rise to endemic genera (Mangoes to Eulampif, and Emeralds to Orthodynczu). Even groups that diversified in the highlands (Brilliants and Bees) gave rise to taxa that subsequently reinvaded the tropical lowlands. As the result of these varied histories, hummingbird communities cannot be arranged easily with respect to organizational complexity and coevolution with nectar sources. Although the physically insular faunas in the Andes and West Indies differ markedly in diversity, both were more strongly affected by colonization than the other faunas. A high potential for coevolution between hummingbirds and plants probably facilitated the successful establishment and radiation of the several Andeanassociated lineages. However, coexistence between the two most diverse Andean clades may have been favoured initially through different habitat preferences by their extra-Andean ancestors. In the tropical lowlands, by comparison, the basic separation between the forestdwelling hermits and canopy and edge-dwelling nonhermits appears to have evolved in situ. The low species and morphologic diversity of hummingbirds breeding north of Mexico reflects the predominance there of a single relatively recent lineage. The regional coexistence of numerous unrelated lineages implies that patterns of ancestry, colonization, and extinction contribute to the make-up of contemporary species-rich hummingbird faunas and serves to qualify the view that hummingbird communities are coadapted assemblages that resist change.

A survey of 166 hummingbird species reveals novel associations of bill-length sexual dimorphism (BLSD) with plumage and breeding behaviours. Across all species, female bills become proportionately longer than male bills (higher female-to-male BLSD ratio) as sexual dichromatism increases. However, male bills are proportionately longer (lower female-to-male BLSD ratio) in both lekkers (traditional group display) and clustered breeders (female harems or colonial nests) compared with dispersed breeders. The overall positive association of plumage with BLSD suggests that social status determines priority of access to nectar-providing £owers. Furthermore, the distinctive BLSD associated with breeding aggregations may arise from behaviours that impose constraints on the usual male priority at £owers: female dominance over males around nest colonies and male residence on lek-mating territories. These various factors appear to alter plumage and bill characters of both sexes to produce the range of dimorphisms within the various dispersed and aggregated breeding system categories. Feedback loops caused by ecological consequences of breeding behaviour may alter the evolutionary dynamics of breeding systems, bird^plant interactions, and competing pollinators, as well as help explain the lek paradox.

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Ecological studies of hummingbird communities have emphasized the importance of local conditions and contemporary interactions in the development of these varied faunas. A timecalibrated, DNA hybridization-based phylogeny of the principal hummingbird lineages was used to examine historical aspects of hummingbird faunas in the species-rich tropical lowlands and Andes, and the relatively depauperate West Indies and temperate regions of Central and North America. Parsimony reconstructions of ancestral distributions indicate that these faunas are polyphyletic in origin, comprising several to many independent lineages. Based on the timing of geologic and cladogenic events, hummingbird faunas appear to have arisen more often by colonization than by large-scale vicariance, with multiple dispersals across water gaps, elevational gradients, and latitude. The extent to which particular lineages colonized different regions depended, however, on lineage ecology as well as on the habitat and age of the fauna. In general, the oldest extant trochilofauna, which today occupies the tropical lowlands, was the principal source of colonizing taxa. However, all regions except possibly the West Indies contributed taxa now found elsewhere, including in the tropical lowlands. The Andean fauna comprises several lineages with lowland origin (hermits, Mangoes, Brilliants, Coquettes, Emeralds) as well as at least one that arose in temperate regions outside South America (Bees). At least two lineages that colonized the West Indies gave rise to endemic genera (Mangoes to Eulampif, and Emeralds to Orthodynczu). Even groups that diversified in the highlands (Brilliants and Bees) gave rise to taxa that subsequently reinvaded the tropical lowlands. As the result of these varied histories, hummingbird communities cannot be arranged easily with respect to organizational complexity and coevolution with nectar sources. Although the physically insular faunas in the Andes and West Indies differ markedly in diversity, both were more strongly affected by colonization than the other faunas. A high potential for coevolution between hummingbirds and plants probably facilitated the successful establishment and radiation of the several Andeanassociated lineages. However, coexistence between the two most diverse Andean clades may have been favoured initially through different habitat preferences by their extra-Andean ancestors. In the tropical lowlands, by comparison, the basic separation between the forestdwelling hermits and canopy and edge-dwelling nonhermits appears to have evolved in situ. The low species and morphologic diversity of hummingbirds breeding north of Mexico reflects the predominance there of a single relatively recent lineage. The regional coexistence of numerous unrelated lineages implies that patterns of ancestry, colonization, and extinction contribute to the make-up of contemporary species-rich hummingbird faunas and serves to qualify the view that hummingbird communities are coadapted assemblages that resist change.