Early stages of speciation with gene flow in the Amazilia Hummingbird (Amazilis amazilia) subspecies complex of Western South America.

Disentangling the factors underlying the diversification of geographically variable species with a wide geographical range is essential to understanding the initial stages and drivers of the speciation process. The Amazilia Hummingbird, Amazilis amazilia, is found along the Pacific coast from northern Ecuador down to the Nazca Valley of Peru, and is currently classified as six phenotypically differentiated subspecies. We aimed to resolve the evolutionary relationships of the six subspecies, to assess the geographical pattern and extent of evolutionary divergence, and to test for introgression using both a mtDNA marker and a genome-by-sequencing dataset from 86 individuals from across the species range. The consensus phylogenetic tree separated the six subspecies into three distinct clades, corresponding with the Ecuador lowlands (A. amazilia dumerilii), the Ecuador highlands (A. amazilia alticola and A. amazilia azuay), and the Peruvian coast (A. amazilia leucophoea, A. amazilia amazilia, and A. amazilia caeruleigularis). However, an unresolved mtDNA network suggests that the diversification of the subspecies was recent and rapid. We found evidence of gene flow among the subspecies A. amazilia dumerilii, A. amazilia alticola, and A. amazilia leucophoea, with strong genetic isolation of the subspecies A. amazilia azuay in the isolated Yunguilla Valley of Ecuador. Finally, environmental data from each subspecies' capture locations were concordant with the three distinct clades. Overall, our results suggest that both expansions into new habitats and geographic isolation shaped the present-day phylogeny and range of the A. amazilia subspecies, and that A. amazilia azuay may be genetically divergent enough to be considered a separate species.

Comparison of adult census size and effective population size support the need for continued protection of two Solomon Island endemics.

Because a population's ability to respond to rapid change is dictated by standing genetic variation, we can better predict a population's long-term viability by estimating and then comparing adult census size (N) and effective population size (N e ). However, most studies only measure N or N e , which can be misleading. Using a combination of field and genomic sequence data, we here estimate and compare N and N e in two range-restricted endemics of the Solomon Islands. Two Zosterops White-eye species inhabit the small island of Kolombangara, with a high elevation species endemic to the island (Z. murphyi) and a low elevation species endemic to the Solomon Islands (Z. kulambangrae). Field observations reveal large values of N for both species with Z. kulambangrae numbering at 114,781 ± 32,233 adults, and Z. murphyi numbering at 64,412 ± 15,324 adults. In contrast, genomic analyses reveal that N e was much lower than N, with Z. kulambangrae estimated at 694.5 and Z. murphyi at 796.1 individuals. Further, positive Tajima's D values for both species suggest that they have experienced a demographic contraction, providing a mechanism for low values of N e . Comparison of N and N e suggests that Z. kulambangrae and Z. murphyi are not at immediate threat of extinction but may be at genetic risk. Our results provide important baseline data for long-term monitoring of these island endemics, and argue for measuring both population size estimates to better gauge long-term population viability.

Rapid, complete reproductive isolation in two closely related Zosterops White-eye bird species despite broadly overlapping ranges.

Examining what happens when two closely related species come into secondary contact provides insight into the later stages of the speciation process. The Zosteropidae family of birds is one of the most rapidly speciating vertebrate lineages. Members of this family are highly vagile and geographically widespread, raising the question of how divergence can occur if populations can easily come into secondary contact. On the small island of Kolombangara, two closely related nonsister species of white-eyes, Zosterops kulambangrae and Zosterops murphyi, are distributed along an elevational gradient and come into secondary contact at mid-elevations. We captured 134 individuals of both species along two elevational transects. Using genotyping-by-sequencing data and a mitochondrial marker, we found no evidence of past hybridization events and strong persistence of species boundaries, even though the species have only been diverging for approximately 2 million years. We explore potential reproductive barriers that allow the two species to coexist in sympatry, including premating isolation based on divergence in plumage and song. We also conducted a literature review to determine the time it takes to evolve complete reproductive isolation in congeneric avian species/subspecies in secondary contact (restricted to cases where congeneric taxa are parapatric or have a hybrid zone), finding our study is one of the youngest examples of complete reproductive isolation studied in a genomic context reported in birds.