Hybrid zones are often described as clines of genetic and phenotypic traits moving across species barriers through introgression. Yet, hybrid zones can also be spatially complex and shift over time, and dispersal and vicariance can isolate portions of a cline, potentially leading to hybrid lineage formation. We reassessed Sibley’s (1950) gradient between Collared Towhee (Pipilo ocai) and Spotted Towhee (P. maculatus) in Central Mexico to test whether it conformed to a typical tension-zone cline model. By comparing historical and modern data, we found that cline centers for genetic and phenotypic traits have not shifted over the course of 70 years. This equilibrium suggests that secondary contact between these species, which originally diverged over 2 million years ago, likely dates to the Pleistocene. Given the amount of mtDNA divergence, parental ends of the cline have very low autosomal nuclear differentiation (FST = 0.12). Dramatic and coincident cline shifts in mtDNA and throat color suggest the possibility of sexual selection as a factor in differential introgression, while a contrasting cline shift in green back color hints at a role for natural selection. Supporting the idea of a continuum between hybrid clines and hybrid lineage formation, the towhee gradient can be analyzed as one population under isolation-by-distance, as a two-population cline, and as three lineages experiencing divergence with gene flow. In the middle of the gradient, a hybrid lineage has become partly isolated, likely due both to forested habitat shrinking and fragmenting as it moved upslope after the last glacial maximum and a stark environmental transition. The towhee system offers a window into the potential outcomes of hybridization across a dynamic landscape including the creation of novel genomic and phenotypic combinations and incipient hybrid lineages.

Morphological data: traits measured from specimens with ruler and digital calipers

Genetic data: SNPs generated from sequence capture of ultraconserved elements and Illumina sequencing

Environmental data: extracted from environmental layers from localities associated with photo vouchers on iNaturalist