Abstract
Haploblocks are regions of the genome that coalesce to an ancestor as a
single unit. Differentiated haplotypes in these regions can result from
the accumulation of mutational differences in low-recombination
chromosomal regions, especially when selective sweeps occur within
geographically structured populations. We introduce a method to identify
large well-differentiated haploblock regions (LHBRs), based on the
variance in standardized heterozygosity (ViSHet) of single nucleotide
polymorphism (SNP) genotypes among individuals, calculated across a
genomic region (500 SNPs in our case). We apply this method to the
greenish warbler (Phylloscopus trochiloides) ring species, using
a newly assembled reference genome and genotypes at more than 1 million
SNPs among 257 individuals. Most chromosomes carry a single distinctive
LHBR, containing 4-6 distinct haplotypes that are associated with
geography, enabling detection of hybridization events and transition
zones between taxa. LHBRs have exceptionally low within-haplotype
nucleotide variation and moderately low between-haplotype nucleotide
distance, suggesting their establishment through recurrent selective
sweeps at varying geographic scales. Meiotic drive is potentially a
powerful mechanism of producing such selective sweeps, and the LHBRs are
likely to often represent centromeric regions where recombination is
restricted. Links between populations enable introgression of favored
haplotypes and we identify one haploblock showing a highly discordant
distribution compared to the rest of the genome, being present in two
distantly separated geographic regions that are at similar latitudes in
both east and west Asia. Our results set the stage for detailed studies
of haploblocks, including their genomic location, gene content, and
contribution to reproductive isolation.