SUMMARY A major challenge in biology is to understand the genetic basis of adaptation. One compelling idea is that groups of tightly linked genes (i.e. ‘supergenes’ [1, 2]) facilitate adaptation in suites of traits that determine fitness. Despite their likely importance, little is known about how alternate supergene alleles arise and become differentiated, nor their ultimate fate within species. Herein we address these questions by investigating the evolutionary history of a supergene in white-throated sparrows, Zonotrichia albicollis. This species comprises two morphs, tan and white, that differ in pigmentation and components of social behavior [3–5]. Morph is determined by alternative alleles at a balanced >100Mb inversion-based supergene, providing a unique system for studying gene-behavior relationships. Using over two decades of field data we document near-perfect disassortative mating among morphs, as well as the fitness consequences of rare assortative mating. We use de novo whole genome sequencing coupled with population- and phylo-genomic data, to show that alternate supergene alleles are highly divergent at over 1,000 genes, that these alleles originated prior to the split of Z. albicollis from its sister species, and may be polymorphic in Z. albicollis due to a past hybridization event. We provide evidence that the ‘white' allele may be degrading, similar to neo-Y/Wsex chromosomes. We further show that the ‘tan’ allele has surprisingly low levels of genetic diversity, yet does not show several canonical signatures of recurrent positive selection. We discuss these results in the context of the origin, molecular evolution, and possible fate of this remarkable polymorphism.

In territorial species, the reproductive success of a male is dependent on the quality of his territory. One important component of territory quality is spatial location. High‐quality territories not only should be located in areas of high food abundance and low predation, but also should be located in areas that offer optimal amounts of social interaction. Such optima might be different for individuals according to their sex, dominance, or genotype. We studied territory quality (size, vegetation structure, and placement) in a socially monogamous, polymorphic passerine, the White‐throated Sparrow (Zonotrichia albicollis), in order to determine how spatial attributes contribute to selection intensity on two genotypes. In this species, plumage (white and tan), behavior, and life‐history characteristics have a genetic basis and are correlated with the presence or absence of a chromosomal inversion. Using remote sensing and Geographic Information Systems (GIS), we found that the territories of white and tan males do not differ in size or vegetation structure, suggesting that these factors are not of primary importance to males when deciding where to establish a territory. Instead, we suggest that the placement of white and tan territories depends on the number of neighbors (particularly, white male neighbors). Tan males settle in low‐density, neighbor‐restrictive habitats where intruder pressure from white males seeking extra‐pair copulations is reduced. In contrast, white males tend to settle in high‐density areas where the probability of encountering neighboring fertile females is greatest. This segregation has led to intraspecific niche partitioning in the two disassortative pair types so that each male morph can best exploit its respective reproductive strategies. These factors may, in turn, contribute to the maintenance of this unusual mating system and, ultimately, the stability of the polymorphism in this species. Similar forces may be operating in other species without distinct morphological markers; we suggest that researchers keep social factors in mind when examining habitat selection.

Aim Ecological factors that vary along spatial scales can greatly influence the outcome of evolution. However, often it is difficult for researchers to identify significant ecological variables that might be associated with the geographical distribution of phenotypes and genotypes in nature. In this paper, we use remote sensing image texture analyses to investigate breeding territories of the white-throated sparrow ( Zonotrichia albicollis ), a polymorphic passerine species that has been shown to segregate spatially according to morph. Our aim was to examine how an integrative measure of landscape feature, such as image texture, could be used to distinguish subtle differences in habitat use and, presumably, habitat selection.Location A population of white-throated sparrows located 1.5 miles east of the Cranberry Lake Biological Station in the Adirondack Mountains of St Lawrence County, New York, was used in this study; the study site itself was approximately 1 km 2 .Methods We analysed 27 territories (16 white males; 11 tan males) to determine if image texture can be used to distinguish differences in vegetation and structure in territories of the two colour morphs. First (variance)-and second (homogeneity)-order image texture measurements with three different window sizes were derived using the green, red, and near infrared bands of Advanced Spaceborne Thermal Emission and Reflection Radiometer (ASTER) data. Texture was then placed into ttests to determine statistical differences between the territories of the two male morphs. ResultsSignificant differences between the territories of the two morphs were found in both variance and homogeneity texture measures in the red and green ASTER bands in all three window sizes. There were no differences with any near infrared texture measures.Main conclusions Examination of first-and second-order statistical measures indicated that the territories of tan males have significantly more spectral variance than the territories of white males, while white males' territories are significantly more homogeneous. These findings are consistent with the previous finding of the morphs settling in 'high' and 'low' neighbour density habitats that differ with respect to the amount of useable area on their borders. Previous study has also suggested that white and tan male territories differ according to overall landscape structure, not vegetation composition. Therefore, we suggest that red and green spectra may be more useful when examining differences in territory structure and possibly territory quality of similar edge-type species. Our results suggest that image texture is a useful tool for ecological and evolutionary modelling of the distributions of phenotypes/ genotypes across landscapes.

How reproductive strategies contribute to patterns of senescence in natural populations remains contentious. We studied reproductive senescence in the dimorphic white-throated sparrow, an excellent species for exploring this issue. Within both sexes the morphs use distinct reproductive strategies, and disassortative pairing by morph results in pair types with distinct parental systems. White morph birds are more colorful and aggressive than tan counterparts, and white males compete for extrapair matings, whereas tan males are more parental. Tan males and white females share parental care equally, whereas white males provide little parental support to tan females. We found morph-specific patterns of reproductive senescence in both sexes. White males exhibited greater reproductive senescence than tan males. This result likely reflects the difficulty of sustaining a highly competitive reproductive strategy as aging progresses rather than high physiological costs of competitiveness, since white males were also long-lived. Moreover, morph was not consistently related to reproductive senescence across the sexes, arguing against especially high costs of the traits associated with white morph identity. Rather, tan females exhibited earlier reproductive senescence than white females and were short-lived, perhaps reflecting the challenges of unsupported motherhood. Results underscore the importance of social dynamics in determining patterns of reproductive senescence.