Ecological characteristics explain neutral genetic variation of three coastal sparrow species.

Eco-phylogeographic approaches to comparative population genetic analyses allow for the inclusion of intrinsic influences as drivers of intraspecific genetic structure. This insight into microevolutionary processes, including changes within a species or lineage, provides better mechanistic understanding of species-specific interactions and enables predictions of evolutionary responses to environmental change. In this study, we used single nucleotide polymorphisms (SNPs) identified from reduced representation sequencing to compare neutral population structure, isolation by distance (IBD), genetic diversity and effective population size (Ne) across three closely related and co-distributed saltmarsh sparrow species differing along a specialization gradient-Nelson's (Ammospiza nelsoni subvirgata), saltmarsh (A. caudacuta) and seaside sparrows (A. maritima maritima). Using an eco-phylogeographic lens within a conservation management context, we tested predictions about species' degree of evolutionary history and ecological specialization to tidal marshes, habitat, current distribution and population status on population genetic metrics. Population structure differed among the species consistent with their current distribution and habitat factors, rather than degree of ecological specialization: seaside sparrows were panmictic, saltmarsh sparrows showed hierarchical structure and Nelson's sparrows were differentiated into multiple, genetically distinct populations. Neutral population genetic theory and demographic/evolutionary history predicted patterns of genetic diversity and Ne rather than degree of ecological specialization. Patterns of population variation and evolutionary distinctiveness (Shapely metric) suggest different conservation measures for long-term persistence and evolutionary potential in each species. Our findings contribute to a broader understanding of the complex factors influencing genetic variation, beyond specialist-generalist status and support the role of an eco-phylogeographic approach in population and conservation genetics.

Mercury exposure of tidal marsh songbirds in the northeastern United States and its association with nest survival.

The biogeochemistry of tidal marsh sediments facilitates the transformation of mercury (Hg) into the biologically available form methylmercury (MeHg), resulting in elevated Hg exposures to tidal marsh wildlife. Saltmarsh and Acadian Nelson's sparrows (Ammospiza caudacutua and A. nelsoni subvirgatus, respectively) exclusively inhabit tidal marshes, potentially experiencing elevated risk to Hg exposure, and have experienced range-wide population declines. To characterize spatial and temporal variation of Hg exposure in these species, we sampled total mercury (THg) in blood collected from 9 populations spanning 560 km of coastline, including individuals resampled within and among years. Using concurrent nesting studies, we tested whether THg was correlated with nest survival probabilities, an index of fecundity. Blood THg ranged from 0.074-3.373 µg/g ww across 170 samples from 127 individuals. We detected high spatial variability in Hg exposure, observing differences of more than 45-fold across all individuals and 8-fold in mean blood THg among all study plots, including 4-fold between study plots within 4 km. Intraindividual changes in blood Hg exposure did not vary systematically in time but were considerable, varying by up to 2-fold within and among years. Controlling for both species differences and maximum water level, the dominant driver of fecundity in this system, nest survival probability decreased by 10% across the full range of female blood THg concentrations observed. We conclude that Hg has the potential to impair songbird reproduction, potentially exacerbating known climate-change driven population declines from sea-level rise in saltmarsh and Acadian Nelson's sparrows.

Mercury exposure in migrating songbirds: correlations with physical condition.

Many migratory songbirds are at high risk of methylmercury (MeHg) exposure due to their trophic position and foraging in and around wetland habitats. Methylmercury has the potential to alter migratory behaviors and physiology via neurological impairment or reduced flight performance and can be remobilized from songbird muscle tissue during migration, increasing the risk of acute MeHg exposure. To document MeHg exposure and its relationship with physical condition in migratory songbirds, we sampled passerine blood and feathers at a migration stopover site on Key Biscayne, FL during fall and spring from 2009 to 2012. We found evidence that spring blood total mercury (THg) concentrations decreased throughout the day and that fall feather THg concentrations changed over the migratory season. Total mercury exposure was marginally correlated with migratory fat stores and related to changes in pectoral muscle thickness by time of day. These patterns suggest that environmentally relevant levels of THg are related to, and may be influencing, the physical condition of free-living migrating songbirds. Further research and monitoring during the migratory period will be important to elucidate exposure risk across multiple species and assess the potential for effects during this complex period of the annual cycle.

Framework for quantifying population responses to disturbance reveals that coastal birds are highly resilient to hurricanes.

Changes in the frequency and severity of extreme weather may introduce new threats to species that are already under stress from gradual habitat loss and climate change. We provide a probabilistic framework that quantifies potential threats by applying concepts from ecological resilience to single populations. Our approach uses computation to compare disturbance-impacted projections to a population's normal range of variation, quantifying the full range of potential impacts. We illustrate this framework with projection models for coastal birds, which are commonly depicted as vulnerable to disturbances, especially hurricanes and oil spills. We found that populations of coastal specialists are resilient to extreme disturbances, with high resistance to the effects of short-term reductions in vital rates and recovery within 20 years. Applying the general framework presented here across disturbance-prone species and ecosystems would improve understanding of population resilience and generate specific projections of resilience that are needed for effective conservation planning.

Contrasting drivers of diversification rates on islands and continents across three passerine families.

Diversification rates vary greatly among taxa. Understanding how species-specific traits influence speciation rates will help elucidate mechanisms driving biodiversity over broad spatio-temporal scales. Ecological specialization and range size are two hypothesized drivers of speciation rates, yet each mechanism predicts both increases and decreases in speciation. We constructed a continuous index of specialization using avian bill morphology to determine the relative effect of specialization and range size and shape on speciation rates across 559 species within the Emberizoidea superfamily, a morphologically diverse New World clade. We found a significant positive correlation between specialization and speciation rate and a negative correlation with range size. Only the effect of specialization persisted after removing island endemics, however, suggesting that ecological specialization is an important driver of diversity across large macroevolutionary scales, and the relative importance of specific drivers may differ between islands and continents.

Subspecies delineation amid phenotypic, geographic and genetic discordance in a songbird.

Understanding the processes that drive divergence within and among species is a long-standing goal in evolutionary biology. Traditional approaches to assessing differentiation rely on phenotypes to identify intra- and interspecific variation, but many species express subtle morphological gradients in which boundaries among forms are unclear. This intraspecific variation may be driven by differential adaptation to local conditions and may thereby reflect the evolutionary potential within a species. Here, we combine genetic and morphological data to evaluate intraspecific variation within the Nelson's (Ammodramus nelsoni) and salt marsh (Ammodramus caudacutus) sparrow complex, a group with populations that span considerable geographic distributions and a habitat gradient. We evaluated genetic structure among and within five putative subspecies of A. nelsoni and A. caudacutus using a reduced-representation sequencing approach to generate a panel of 1929 SNPs among 69 individuals. Although we detected morphological differences among some groups, individuals sorted along a continuous phenotypic gradient. In contrast, the genetic data identified three distinct clusters corresponding to populations that inhabit coastal salt marsh, interior freshwater marsh and coastal brackish-water marsh habitats. These patterns support the current species-level recognition but do not match the subspecies-level taxonomy within each species-a finding which may have important conservation implications. We identified loci exhibiting patterns of elevated divergence among and within these species, indicating a role for local selective pressures in driving patterns of differentiation across the complex. We conclude that this evidence for adaptive variation among subspecies warrants the consideration of evolutionary potential and genetic novelty when identifying conservation units for this group.

Predictors of specialist avifaunal decline in coastal marshes.

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.

Seasonal fecundity is not related to geographic position across a species' global range despite a central peak in abundance.

The range of a species is determined by the balance of its demographic rates across space. Population growth rates are widely hypothesized to be greatest at the geographic center of the species range, but indirect empirical support for this pattern using abundance as a proxy has been mixed, and demographic rates are rarely quantified on a large spatial scale. Therefore, the texture of how demographic rates of a species vary over its range remains an open question. We quantified seasonal fecundity of populations spanning the majority of the global range of a single species, the saltmarsh sparrow (Ammodramus caudacutus), which demonstrates a peak of abundance at the geographic center of its range. We used a novel, population projection method to estimate seasonal fecundity inclusive of seasonal and spatial variation in life history traits that contribute to seasonal fecundity. We replicated our study over 3 years, and compared seasonal fecundity to latitude and distance among plots. We observed large-scale patterns in some life history traits that contribute to seasonal fecundity, such as an increase in clutch size with latitude. However, we observed no relationship between latitude and seasonal fecundity. Instead, fecundity varied greatly among plots separated by as little as 1 km. Our results do not support the hypothesis that demographic rates are highest at the geographic and abundance center of a species range, but rather they suggest that local drivers strongly influence saltmarsh sparrow fecundity across their global range.

Demographic analysis demonstrates systematic but independent spatial variation in abiotic and biotic stressors across 59 percent of a global species range.

The balance of abiotic and biotic stressors experienced by a species likely varies across its range, resulting in spatially heterogeneous limitations on the species' demographic rates. Support for spatial variation in stressors (often latitudinal gradients) has been found in many species, usually with physiological or correlative occupancy data, but it has rarely been estimated directly with demographic data. We collected demographic data from 23 sites spanning the majority of the Saltmarsh Sparrow (Ammodramus caudacutus) breeding range. Using data from 837 nests, we quantified the abiotic and biotic variables most important to nest survival, which is the dominant driver of both fecundity and population growth rate in this species. We separately estimated daily nest failure probability due to nest depredation (biotic stressor) and nest flooding (abiotic stressor), which collectively account for almost all nest failure in the species. Nest depredation decreased with latitude, whereas nest flooding was not related to latitude. Instead, nest flooding was best predicted by a combination of maximum high tide, extremity of rare flooding events, and date. For a single vital rate, we observed predictable variation in competing biotic and abiotic stressors across this species range. We observed that biotic and abiotic stressors were geographically independent, both on a large spatial scale and locally. Our results suggest that stressors on the fecundity of Saltmarsh Sparrow vary systematically across its range, but independently. The observed patterns of biotic and abiotic stress provide information for efforts to conserve the Saltmarsh Sparrow, which is considered threatened. Further, understanding the effects that different stressors, and their interactions, have on demographic rates is necessary to unravel the processes that govern species distributions and to effectively conserve biodiversity in the face of global change.

El balance de factores de estrés abióticos y bióticos para una especie probablemente varía a través de su rango, dando como resultado limitaciones espaciales heterogéneas en las tasas demográficas de la especie. Se ha verificado la existencia de variación espacial en los factores de estrés (usualmente gradientes latitudinales) para muchas especies, usualmente con datos fisiológicos o de ocupación correlativa, pero raramente se ha estimado directamente con datos demográficos. Colectamos datos demográficos de 23 sitios abarcando la mayoría del rango reproductivo de Ammodramus caudacutus. Usando datos de 837 nidos, cuantificamos las variables abióticas y bióticas más importantes para la supervivencia del nido, que es la variable que determina tanto la fecundidad como la tasa de crecimiento poblacional en esta especie. Por otra parte, estimamos la probabilidad de fracaso diario del nido debido a la depredación del nido (factor de estrés biótico) e inundación del nido (factor de estrés abiótico), que juntos representaron casi todos los fracasos del nido en esta especie. La depredación del nido disminuyó con la latitud mientras que la inundación del nido no se relacionó con la latitud. En cambio, la inundación del nido se predijo mejor por una combinación del máximo superior de la marea, la extremidad de los eventos de inundación raros y la fecha. Considerando una sola tasa vital, observamos variación predecible en los factores de estrés biótico y abiótico que compiten a través del rango de la especie. Observamos que los factores de estrés biótico y abiótico fueron geográficamente independientes tanto a una escala espacial grande como a la escala local. Nuestros resultados sugieren que los factores de estrés relacionados a la fecundidad de A. caudacutus varían sistemática pero independientemente a través de su rango. Los patrones observados de estrés biótico y abiótico brindan información para los esfuerzos de conservación de A. caudacutus, una especie considerada amenazada. Más aún, es necesario entender los efectos que los diferentes factores de estrés y sus interacciones tienen en las tasas demográficas para desenmarañar los procesos que gobiernan las distribuciones de las especies y para conservar la biodiversidad de manera eficiente en miras al cambio global.

Factors that influence vital rates of Seaside and Saltmarsh sparrows in coastal New Jersey, USA.

As saltmarsh habitat continues to disappear, understanding the factors that influence saltmarsh breeding bird population dynamics is an important step for the conservation of these declining species. Using five years (2011 - 2015) of demographic data, we evaluated and compared Seaside (Ammodramus maritimus) and Saltmarsh (A. caudacutus) sparrow apparent adult survival and nest survival at the Edwin B. Forsythe National Wildlife Refuge, New Jersey, USA. We determined the effect of site management history (unditched vs. ditched marsh) on adult and nest survival to aid in prioritizing future management or restoration actions. Seaside Sparrow apparent adult survival (61.6%, 95% CI: 52.5 - 70.0%) averaged >1.5 times greater than Saltmarsh Sparrow apparent adult survival (39.9%, 95% CI: 34.0 - 46.2%). Nest survival and predation and flooding rates did not differ between species, and predation was the primary cause of failure for both species. Apparent adult survival and nest survival did not differ between unditched and ditched marshes for either species, indicating that marsh ditching history may not affect breeding habitat quality for these species. With predation as the primary cause of nest failure for both species in New Jersey, we suggest that future research should focus on identification of predator communities in salt marshes and the potential for implementing predator-control programs to limit population declines.

Differential introgression and the maintenance of species boundaries in an advanced generation avian hybrid zone.

BACKGROUND: Evolutionary 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. RESULTS: Geographic 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. CONCLUSIONS: Our 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.

Plumage microorganism communities of tidal marsh sparrows.

Microorganism communities can shape host phenotype evolution but are often comprised of thousands of taxa with varied impact on hosts. Identification of taxa influencing host evolution relies on first describing microorganism communities and acquisition routes. Keratinolytic (keratin-degrading) microorganisms are hypothesized to be abundant in saltmarsh sediments and to contribute to plumage evolution in saltmarsh-adapted sparrows. Metabarcoding was used to describe plumage bacterial (16S rRNA) and fungal (ITS) communities in three sparrow species endemic to North America's Atlantic coast saltmarshes. Results describe limited within-species variability and moderate host species-level patterns in microorganism diversity and community composition. A small percentage of overall microorganism diversity was comprised of potentially keratinolytic microorganisms, warranting further functional studies. Distinctions between plumage and saltmarsh sediment bacteria, but not fungal, communities were detected, suggesting multiple bacterial acquisition routes and/or vertebrate host specialization. This research lays groundwork for future testing of causal links between microorganisms and avian host evolution.

Mercury in wing and tail feathers of hatch-year and adult tidal marsh sparrows.

We estimated mercury exposure and bioaccumulation in sparrow feathers to determine variation among age groups, between sparrow species, and between feather types. Results of feather mercury studies in piscivorous birds indicate that mercury concentrations tend to increase with age and differ between feather types; however, data for insectivorous birds are lacking. We estimated mercury exposure of two insectivorous and sympatric tidal marsh sparrows: coastal plain swamp sparrow (Melospiza georgiana nigrescens), and seaside sparrow (Ammodramous maritimus). Tidal marshes have favorable conditions for mercury methlyation, thus it is likely that tidal marsh sparrows are exposed to methylmercury. We found no difference in mercury concentrations between males and female birds of both species. Adult swamp sparrow feather mercury concentrations did not differ among adult age groups; therefore, mercury was not found to increase with age in sparrows at the site. Hatch-year birds had significantly greater feather mercury concentrations compared with adult birds for both species. Mercury concentrations in adult seaside sparrows were twice as high as those in adult swamp sparrows suggesting species-specific variation, although concentrations in hatch-year sparrow species did not differ. Mercury concentrations differed between feather types in adults of both species. The first primary feather of both species had at least three times greater mercury concentrations than the outer tail feather possibly reflecting varying depuration rates with feather type.

Patterns of introgression vary within an avian hybrid zone.

BACKGROUND: Exploring hybrid zone dynamics at different spatial scales allows for better understanding of local factors that influence hybrid zone structure. In this study, we tested hypotheses about drivers of introgression at two spatial scales within the Saltmarsh Sparrow (Ammospiza caudacuta) and Nelson's Sparrow (A. nelsoni) hybrid zone. Specifically, we evaluated the influence of neutral demographic processes (relative species abundance), natural selection (exogenous environmental factors and genetic incompatibilities), and sexual selection (assortative mating) in this mosaic hybrid zone. By intensively sampling adults (n = 218) and chicks (n = 326) at two geographically proximate locations in the center of the hybrid zone, we determined patterns of introgression on a fine scale across sites of differing habitat. We made broadscale comparisons of patterns from the center with those of prior studies in the southern edge of the hybrid zone. RESULTS: A panel of fixed SNPs (135) identified from ddRAD sequencing was used to calculate a hybrid index and determine genotypic composition/admixture level of the populations. Another panel of polymorphic SNPs (589) was used to assign paternity and reconstruct mating pairs to test for sexual selection. On a broad-scale, patterns of introgression were not explained by random mating within marshes. We found high rates of back-crossing and similarly low rates of recent-generation (F1/F2) hybrids in the center and south of the zone. Offspring genotypic proportions did not meet those expected from random mating within the parental genotypic distribution. Additionally, we observed half as many F1/F2 hybrid female adults than nestlings, while respective male groups showed no difference, in support of Haldane's Rule. The observed proportion of interspecific mating was lower than expected when accounting for mate availability, indicating assortative mating was limiting widespread hybridization. On a fine spatial scale, we found variation in the relative influence of neutral and selective forces between inland and coastal habitats, with the smaller, inland marsh influenced primarily by neutral demographic processes, and the expansive, coastal marsh experiencing higher selective pressures in the form of natural (exogenous and endogenous) and sexual selection. CONCLUSIONS: Multiple drivers of introgression, including neutral and selective pressures (exogenous, endogenous, and sexual selection), are structuring this hybrid zone, and their relative influence is site and context-dependent.

Meta-analysis: Congruence of genomic and phenotypic differentiation across diverse natural study systems.

Linking genotype to phenotype is a primary goal for understanding the genomic underpinnings of evolution. However, little work has explored whether patterns of linked genomic and phenotypic differentiation are congruent across natural study systems and traits. Here, we investigate such patterns with a meta-analysis of studies examining population-level differentiation at subsets of loci and traits putatively responding to divergent selection. We show that across the 31 studies (88 natural population-level comparisons) we examined, there was a moderate (R 2 = 0.39) relationship between genomic differentiation (F ST ) and phenotypic differentiation (P ST ) for loci and traits putatively under selection. This quantitative relationship between P ST and F ST for loci under selection in diverse taxa provides broad context and cross-system predictions for genomic and phenotypic adaptation by natural selection in natural populations. This context may eventually allow for more precise ideas of what constitutes "strong" differentiation, predictions about the effect size of loci, comparisons of taxa evolving in nonparallel ways, and more. On the other hand, links between P ST and F ST within studies were very weak, suggesting that much work remains in linking genomic differentiation to phenotypic differentiation at specific phenotypes. We suggest that linking genotypes to specific phenotypes can be improved by correlating genomic and phenotypic differentiation across a spectrum of diverging populations within a taxon and including wide coverage of both genomes and phenomes.

Genomics of rapid ecological divergence and parallel adaptation in four tidal marsh sparrows.

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.

Bidirectional adaptive introgression between two ecologically divergent sparrow species.

Natural hybrid zones can be used to dissect the mechanisms driving key evolutionary processes by allowing us to identify genomic regions important for establishing reproductive isolation and that allow for transfer of adaptive variation. We leverage whole-genome data in a system where two bird species, the saltmarsh (Ammospiza caudacuta) and Nelson's (A. nelsoni) sparrow, hybridize despite their relatively high background genetic differentiation and past ecological divergence. Adaptive introgression is plausible in this system because Nelson's sparrows are recent colonists of saltwater marshes, compared to the specialized saltmarsh sparrow that has a longer history of saltmarsh adaptation. Comparisons among whole-genome sequences of 34 individuals from allopatric and sympatric populations show that ongoing gene flow is shaping the genomic landscape, with allopatric populations exhibiting genome-wide FST estimates close to double of that observed in sympatry. We characterized patterns of introgression across the genome and identify regions that exhibit biased introgression into hybrids from one parental species. These regions offer compelling candidates for genes related to tidal marsh adaptations suggesting that adaptive introgression may be an important consequence of hybridization. These findings highlight the value of considering the landscapes of both genome-wide introgression and divergence when characterizing the evolutionary forces that drive speciation.

Wind patterns as a potential driver in the evolution and maintenance of a North American migratory suture zone.

Suture zones are areas where range contact zones and hybrid zones of multiple taxa are clustered. Migratory divides, contact zones between divergent populations that breed adjacent to one another but use different migratory routes, are a particular case of suture zones. Although multiple hypotheses for both the formation and maintenance of migratory divides have been suggested, quantitative tests are scarce. Here, we tested whether a novel factor, prevailing winds, was sufficient to explain both the evolution and maintenance of the Cordilleran migratory divide using individual-based models. Empirical observations of eastern birds suggest a circuitous migratory route across Canada before heading south. Western breeders, however, travel south along the Pacific coast to their wintering grounds. We modeled the effect of wind on bird migratory flights by allowing them to float at elevation using spatially explicit modeled wind data. Modeled eastern birds had easterly mean trajectories, whereas western breeders showed significantly more southern trajectories. We also determined that a mean airspeed of 18.5 m s(-1) would be necessary to eliminate this difference in trajectory, a speed that is achieved by waterfowl and shorebirds, but is faster than songbird flight speeds. These results lend support for the potential importance of wind in shaping the phylogeographic history of North American songbirds.

Genotype-environment associations support a mosaic hybrid zone between two tidal marsh birds.

Local environmental features can shape hybrid zone dynamics when hybrids are bounded by ecotones or when patchily distributed habitat types lead to a corresponding mosaic of genotypes. We investigated the role of marsh-level characteristics in shaping a hybrid zone between two recently diverged avian taxa - Saltmarsh (Ammodramus caudacutus) and Nelson's (A. nelsoni) sparrows. These species occupy different niches where allopatric, with caudacutus restricted to coastal marshes and nelsoni found in a broader array of wetland and grassland habitats and co-occur in tidal marshes in sympatry. We determined the influence of habitat types on the distribution of pure and hybrid sparrows and assessed the degree of overlap in the ecological niche of each taxon. To do this, we sampled and genotyped 305 sparrows from 34 marshes across the hybrid zone and from adjacent regions. We used linear regression to test for associations between marsh characteristics and the distribution of pure and admixed sparrows. We found a positive correlation between genotype and environmental variables with a patchy distribution of genotypes and habitats across the hybrid zone. Ecological niche models suggest that the hybrid niche was more similar to that of A. nelsoni and habitat suitability was influenced strongly by distance from coastline. Our results support a mosaic model of hybrid zone maintenance, suggesting a role for local environmental features in shaping the distribution and frequency of pure species and hybrids across space.

Helping struggling students in introductory biology: a peer-tutoring approach that improves performance, perception, and retention.

The high attrition rate among science, technology, engineering, and mathematics (STEM) majors has long been an area of concern for institutions and educational researchers. The transition from introductory to advanced courses has been identified as a particularly "leaky" point along the STEM pipeline, and students who struggle early in an introductory STEM course are predominantly at risk. Peer-tutoring programs offered to all students in a course have been widely found to help STEM students during this critical transition, but hiring a sufficient number of tutors may not be an option for some institutions. As an alternative, this study examines the viability of an optional peer-tutoring program offered to students who are struggling in a large-enrollment, introductory biology course. Struggling students who regularly attended peer tutoring increased exam performance, expert-like perceptions of biology, and course persistence relative to their struggling peers who were not attending the peer-tutoring sessions. The results of this study provide information to instructors who want to design targeted academic assistance for students who are struggling in introductory courses.