Physiological factors are rarely proposed to account for variation in the morphology of feeding structures. Recently, bird bills have been demonstrated to be important convective and radiant heat sinks. Larger bills have greater surface area than smaller bills and could serve as more effective thermoregulatory organs under hot conditions. The heat radiating function of bills should be more important in open habitats with little shade and stronger convective winds. Furthermore, as a means of dumping heat without increasing water loss through evaporation, bills might play a particularly important thermoregulatory role in heat loss in windy habitat where fresh water is limited. North American salt marshes provide a latitudinal gradient of relatively homogeneous habitat that is windy, open, and fresh‐water limited. To examine the potential role of thermoregulation in determining bill size variation among ten species or subspecies of tidal marsh sparrows, we plotted bill size against maximum summer and minimum winter temperatures. Bill surface areas increases with summer temperature, which explained 82–89% of the variance (depending upon sex) when we controlled for genus membership. Latitude alone predicted bill surface area much more poorly than summer temperature, and winter temperatures explained < 10% of the variance in winter bill size. Tidal marsh sparrow bill morphology may, to a large degree, reflect the role of the bill in expelling excess body heat in these unbuffered, fresh‐water‐limited environments. This new example of Allen's rule reaffirms the importance of physiological constraints on the evolution of vertebrate morphologies, even in bird bills, which have conventionally been considered as products of adaptation to foraging niche.

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Coastal marshes are one of the world's most productive ecosystems. Consequently, they have been heavily used by humans for centuries, resulting in ecosystem loss. Direct human modifications such as road crossings and ditches and climatic stressors such as sea-level rise and extreme storm events have the potential to further degrade the quantity and quality of marsh along coastlines. We used an 18-year marsh-bird database to generate population trends for 5 avian species (Rallus crepitans, Tringa semipalmata semipalmata, Ammodramus nelsonii subvirgatus, Ammodramus caudacutus, and Ammodramus maritimus) that breed almost exclusively in tidal marshes, and are potentially vulnerable to marsh degradation and loss as a result of anthropogenic change. We generated community and species trends across 3 spatial scales and explored possible drivers of the changes we observed, including marsh ditching, tidal restriction through road crossings, local rates of sea-level rise, and potential for extreme flooding events. The specialist community showed negative trends in tidally restricted marshes (-2.4% annually from 1998 to 2012) but was stable in unrestricted marshes across the same period. At the species level, we found negative population trends in 3 of the 5 specialist species, ranging from -4.2% to 9.0% annually. We suggest that tidal restriction may accelerate degradation of tidal marsh resilience to sea-level rise by limiting sediment supply necessary for marsh accretion, resulting in specialist habitat loss in tidally restricted marshes. Based on our findings, we predict a collapse of the global population of Saltmarsh Sparrows (A. caudacutus) within the next 50 years and suggest that immediate conservation action is needed to prevent extinction of this species. We also suggest mitigation actions to restore sediment supply to coastal marshes to help sustain this ecosystem into the future.

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BackgroundEvolutionary processes, including selection and differential fitness, shape the introgression of genetic material across a hybrid zone, resulting in the exchange of some genes but not others. Differential introgression of molecular or phenotypic markers can thus provide insight into factors contributing to reproductive isolation. We characterized patterns of genetic variation across a hybrid zone between two tidal marsh birds, Saltmarsh (Ammodramus caudacutus) and Nelson’s (A. nelsoni) sparrows (n = 286), and compared patterns of introgression among multiple genetic markers and phenotypic traits.ResultsGeographic and genomic cline analyses revealed variable patterns of introgression among marker types. Most markers exhibited gradual clines and indicated that introgression exceeds the spatial extent of the previously documented hybrid zone. We found steeper clines, indicating strong selection for loci associated with traits related to tidal marsh adaptations, including for a marker linked to a gene region associated with metabolic functions, including an osmotic regulatory pathway, as well as for a marker related to melanin-based pigmentation, supporting an adaptive role of darker plumage (salt marsh melanism) in tidal marshes. Narrow clines at mitochondrial and sex-linked markers also offer support for Haldane’s rule. We detected patterns of asymmetrical introgression toward A. caudacutus, which may be driven by differences in mating strategy or differences in population density between the two species.ConclusionsOur findings offer insight into the dynamics of a hybrid zone traversing a unique environmental gradient and provide evidence for a role of ecological divergence in the maintenance of pure species boundaries despite ongoing gene flow.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0635-y) contains supplementary material, which is available to authorized users.

Conditions favoring population divergence in trophic features, such as the low levels of species richness and interspecific competition found on islands, can be similar to conditions that increase their sexual dimorphism or overall variance. Male emberizid sparrows of tidal marshes have undergone parallel evolution of large bills. We tested for parallel increases between dimorphism and overall variation in bill size by comparing three groups totaling 30 sparrow subspecies: tidal-marsh sparrows, nontidal relatives of tidal-marsh taxa, and representative sparrow taxa. Bill size (and not other features) showed the following patterns in tidal-marsh sparrows compared to nontidal relatives or sparrows at large: (1) an increase; (2) a greater increase in males than females; (3) an increase in sexual dimorphism; and (4) greater variation in females. A high degree of sexual dimorphism in bill size is consistent with the hypothesis that low levels of interspecific and high levels of intraspecific competition select for intraspecific niche divergence. Alternatively, increased sexual selection in tidal-marsh sparrows, vis-a-vis high densities and hence increased male-male competition, may account for the differentially large increase in bill size in males. Relaxed natural selection due to high ecosystem productivity and low interspecific competition may explain why, in tidal-marsh sparrows, female bills have diverged less than males and show higher levels of variability at larger sizes. Both the niche divergence and sexual selection hypotheses depend upon processes, particularly increases in population density, that are similar to those often reported for island passerines. However, the low species diversity and increased intraspecific competition of salt marsh faunas is probably a result of abiotic constraints on colonization (tides and salinity) rather than the isolating distances of island biotas. Thus, both a shift in bill size and increases in its dimorphism and variability may be favored by high productivity and abiotic constraints.

Theory suggests that different taxa having colonized a similar, challenging environment will show parallel or lineage‐specific adaptations to shared selection pressures, but empirical examples of parallel evolution in independent taxa are exceedingly rare. We employed comparative genomics to identify parallel and lineage‐specific responses to selection within and among four species of North American sparrows that represent four independent, post‐Pleistocene colonization events by an ancestral, upland subspecies and a derived salt marsh specialist. We identified multiple cases of parallel adaptation in these independent comparisons following salt marsh colonization, including selection of 12 candidate genes linked to osmoregulation. In addition to detecting shared genetic targets of selection across multiple comparisons, we found many novel, species‐specific signatures of selection, including evidence of selection of loci associated with both physiological and behavioral mechanisms of osmoregulation. Demographic reconstructions of all four species highlighted their recent divergence and small effective population sizes, as expected given their rapid radiation into saline environments. Our results highlight the interplay of both shared and lineage‐specific selection pressures in the colonization of a biotically and abiotically challenging habitat and confirm theoretical expectations that steep environmental clines can drive repeated and rapid evolutionary diversification in birds.