Extreme genetic similarity does not predict non‐breeding distribution of two closely related warblers

Detailed knowledge of migratory connectivity can facilitate effective conservation of Neotropical migrants by helping biologists understand where and when populations may be most limited. We studied the migratory behavior and non‐breeding distribution of two closely related species of conservation concern, the Golden‐winged Warbler (Vermivora chrysoptera) and Blue‐winged Warbler (Vermivora cyanoptera). Although both species have undergone dynamic range shifts and population changes attributed to habitat loss and social interactions promoting competition and hybridization, full life‐cycle conservation planning has been limited by a lack of information about their non‐breeding ecology. Because recent work has demonstrated that the two species are nearly identical genetically, we predicted that individuals from a single breeding population would have similar migratory timing and overwintering locations. In 2015, we placed light‐level geolocators on 25 males of both species and hybrids in an area of breeding sympatry at the Fort Drum Military Installation in Jefferson and Lewis counties, New York. Despite extreme genetic similarity, non‐breeding locations and duration of migration differed among genotypes. Golden‐winged Warblers (N = 2) overwintered > 1900 km southeast of the nearest Blue‐winged Warbler (N = 3) and spent nearly twice as many days in migration; hybrids (N = 2) had intermediate wintering distributions and migratory timing. Spring migration departure dates were staggered based on distance from the breeding area, and all birds arrived at the breeding site within 8 days of each other. Our results show that Golden‐winged Warblers and Blue‐winged Warblers in our study area retain species‐specific non‐breeding locations despite extreme genetic similarity, and suggest that non‐breeding locations and migratory timing vary along a genetic gradient. If the migratory period is limiting for these species, our results also suggest that Golden‐winged Warblers in our study population may be more vulnerable to population decline than Blue‐winged Warblers because they spend almost twice as many days migrating.     

An improved survey method for monitoring population trends of Golden‐winged Warblers and other patchily distributed birds

Conventional surveys designed to monitor common and widespread species may fail to adequately track population changes of rare or patchily distributed species that are often of high conservation concern. We evaluated the performance of a new monitoring approach that employs both a spatially balanced sampling design and a targeted survey protocol designed to estimate population trends of one such patchily distributed species, the Golden‐winged Warbler (Vermivora chrysoptera), in the Appalachian Mountains Bird Conservation Region (BCR 28), USA. Our spatially balanced survey consisted of 105 sample quads (one‐quarter Delorme Atlas pages) across the current range of Golden‐winged Warblers within BCR 28, each with five sample points located in early successional habitat. From 2009 to 2013, collaborators visited each sample point once per year during the peak breeding season and conducted a 17‐min survey consisting of passive observation and playback of conspecific songs and mobbing vocalizations. We used multi‐season, single‐species occupancy models to estimate probability of quad occupancy, detection probability, and occupancy dynamics for Golden‐winged Warblers and closely related Blue‐winged Warblers (Vermivora cyanoptera). Our survey protocol resulted in high estimates of detection probability for Golden‐winged (92%) and Blue‐winged (79%) warblers, with 47% and 56% of quads estimated to be initially occupied, respectively. Derived population trend estimates (λ) indicated an average decline in population of 6% for Golden‐winged Warblers and 7% for Blue‐winged Warblers, resulting in estimated 21% and 22% declines, respectively, in quad occupancy after 5 yr. Our results demonstrate that coupling a spatially balanced survey design in appropriate habitat with a playback protocol to increase detection rates is a viable strategy for tracking populations of Golden‐winged Warblers in the Appalachian Mountains BCR. Similar survey methods should be considered for other rare, declining, or patchily distributed bird species that require targeted monitoring.     

Characteristics of Golden‐winged Warbler territories in plant communities associated with regenerating forest and abandoned agricultural fields

In the Appalachian portion of their breeding range, Golden‐winged Warblers (Vermivora chrysoptera) nest in shrubland and regenerating forest communities created and maintained by disturbance. Because populations of Golden‐winged Warblers have exhibited precipitous declines in population throughout their Appalachian breeding range, management activities that create or maintain early successional habitat are a priority for many natural resource agencies and their conservation partners. Within these early successional habitats, however, additional information is still needed concerning the relative importance of different vegetation features in selection of breeding territories by Golden‐winged Warblers. Our objective, therefore, was to use logistic regression to estimate the probability of territory‐level occupancy by Golden‐winged Warblers in north‐central Pennsylvania at two sites, each with its own early successional community, based on vegetation characteristics. Our communities were composed of shrublands and regenerating forest sites resulting from two disturbances: agriculture and forest fire. Despite differences in vegetation structure, portions of both study areas (regenerating forest and old field) supported territorial Golden‐winged Warblers. Probability of territory occupancy by Golden‐winged Warblers increased with percent blackberry (Rubus) cover in the regenerating forest community, and decreased as basal area and distance to microedge increased (i.e., as vegetation patchiness decreased) in both communities. These habitat features have also been found to influence other aspects of Golden‐winged Warbler breeding ecology such as nest‐site selection, pairing success, and territory abundance. Vegetation features influencing Golden‐winged Warbler territory establishment can differ among shrubland and regenerating forest communities, and management decisions and outcomes may be affected by these differences. Our study provides a starting point for a more comprehensive hypothesis‐driven occupancy survey to investigate features of the territories of Golden‐winged Warblers across a broader geographic range and in different vegetation communities.     

Detection probability of Golden-winged Warblers during point counts with and without playback recordings

ABSTRACT Use of point‐count data to estimate population sizes of North American landbirds may be challenged by limitations on detection probability of particular species, thereby requiring correction factors to ensure accurate estimates. We estimated detection probability of Golden‐winged Warblers (Vermivora chrysoptera) during 3‐min point‐count surveys conducted both with and without use of playback recordings in a mixed shrubland‐forest habitat (clearcut area) and a 60‐m wide electric transmission line right‐of‐way (ROW) in central Pennsylvania from 20 May to 17 June 2002–2003. In addition, we assessed the value of playback with respect to response rates of warblers and distance within which warblers approached the observer. Without playback, detection probability was approximately 23% in the clearcut area and 61% in the ROW. Use of playback resulted in 7% and 19% net increases in probability at the clearcut area and the ROW, respectively; proportional increase was approximately 30% for both habitats. Warblers responded to playback 68% of the time, but response rate was greater within 100 m (72%) than beyond (53%). Most responses (85%) included approach of the warbler toward the observer, and most individuals approached within 10 m. We conclude that 3‐min point counts with playback do not yield detection probabilities sufficient to estimate population size of Golden‐winged Warblers without use of correction factors. Furthermore, detection probability in mixed shrubland‐forest habitats can be much lower than in linear habitats such as utility ROWs. Efforts to estimate population size of Golden‐winged Warblers from data of the North American Breeding Bird Survey should recognize that habitat structure has much influence on detection probability as it relates to distance at which an observer can hear (or see) warblers. Accordingly, we recommend that such efforts incorporate a maximum detection distance of 100–150 m in mixed shrubland‐forest habitats.     

Geolocators reveal migratory connectivity between wintering and breeding areas of Golden‐winged Warblers

The conservation of migratory songbirds is often impeded by a lack of understanding of how populations in breeding and wintering areas are geographically linked (migratory connectivity). In recent years, light‐level geolocators have improved our understanding of migratory connectivity. Such information is valuable for evaluating how conservation efforts align between the breeding and non‐breeding areas of at‐risk species, and help to more effectively prioritize the allocation of conservation funding. Golden‐winged Warblers (Vermivora chrysoptera) are imperiled migratory songbirds, but the extent to which conservation efforts in their breeding and non‐breeding areas coincide with patterns of migratory connectivity are not well known. We used light‐level geolocators to evaluate the extent to which conservation actions targeting Golden‐winged Warblers in Nicaragua and in their breeding range in North America align with patterns of migratory connectivity. We recovered six of 22 geolocators that had been deployed on male Golden‐winged Warblers at the El Jaguar Reserve during the winter of 2015–2016. All six males migrated to breeding areas in the western Great Lakes region that includes eastern Minnesota, northern Wisconsin, southwestern Ontario, and Michigan's Upper Peninsula. All six males also had similar migration routes, with spring stopovers in southern Mexico, Guatemala, and Belize, a trans‐Gulf flight, and a stopover in the region of Louisiana, Arkansas, eastern Oklahoma, and Texas. Our results, in combination with those of previous studies, demonstrate strong migratory connectivity between portions of the breeding and winter distributions of Golden‐winged Warblers currently targeted for conservation. However, additional studies are needed to improve our understanding of the stopover ecology of Golden‐winged Warblers, especially in areas where they remain for extended periods of time. Finally, patterns of migratory connectivity revealed in our study should be used in combination with existing demographic parameters for Golden‐winged Warblers in the western Great Lakes and Nicaragua to help inform full life cycle population models for this imperiled songbird.     

Habitat shifts of endangered species under altered climate conditions: importance of biotic interactions

Predicting changes in potential habitat for endangered species as a result of global warming requires considering more than future climate conditions; it is also necessary to evaluate biotic associations. Most distribution models predicting species responses to climate change include climate variables and occasionally topographic and edaphic parameters, rarely are biotic interactions included. Here, we incorporate biotic interactions into niche models to predict suitable habitat for species under altered climates. We constructed and evaluated niche models for an endangered butterfly and a threatened bird species, both are habitat specialists restricted to semiarid shrublands of southern California. To incorporate their dependency on shrubs, we first developed climate‐based niche models for shrubland vegetation and individual shrub species. We also developed models for the butterfly's larval host plants. Outputs from these models were included in the environmental variable dataset used to create butterfly and bird niche models. For both animal species, abiotic–biotic models outperformed the climate‐only model, with climate‐only models over‐predicting suitable habitat under current climate conditions. We used the climate‐only and abiotic–biotic models to calculate amounts of suitable habitat under altered climates and to evaluate species' sensitivities to climate change. We varied temperature (+0.6, +1.7, and +2.8 °C) and precipitation (50%, 90%, 100%, 110%, and 150%) relative to current climate averages and within ranges predicted by global climate change models. Suitable habitat for each species was reduced at all levels of temperature increase. Both species were sensitive to precipitation changes, particularly increases. Under altered climates, including biotic variables reduced habitat by 68–100% relative to the climate‐only model. To design reserve systems conserving sensitive species under global warming, it is important to consider biotic interactions, particularly for habitat specialists and species with strong dependencies on other species.     

Evidence that climate sets the lower elevation range limit in a high‐elevation endemic salamander

A frequent assumption in ecology is that biotic interactions are more important than abiotic factors in determining lower elevational range limits (i.e., the “warm edge” of a species distribution). However, for species with narrow environmental tolerances, theory suggests the presence of a strong environmental gradient can lead to persistence, even in the presence of competition. The relative importance of biotic and abiotic factors is rarely considered together, although understanding when one exerts a dominant influence on controlling range limits may be crucial to predicting extinction risk under future climate conditions. We sampled multiple transects spanning the elevational range limit of Plethodon shenandoah and site and climate covariates were recorded. A two‐species conditional occupancy model, accommodating heterogeneity in detection probability, was used to relate variation in occupancy with environmental and habitat conditions. Regional climate data were combined with datalogger observations to estimate the cloud base heights and to project future climate change impacts on cloud elevations across the survey area. By simultaneously accounting for species’ interactions and habitat variables, we find that elevation, not competition, is strongly correlated with the lower elevation range boundary, which had been presumed to be restricted mainly as a result of competitive interactions with a congener. Because the lower elevational range limit is sensitive to climate variables, projected climate change across its high‐elevation habitats will directly affect the species’ distribution. Testing assumptions of factors that set species range limits should use models which accommodate detection biases.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Trophic interactions and abiotic factors drive functional and phylogenetic structure of vertebrate herbivore communities across the Arctic tundra biome

Communities are assembled from species that evolve or colonise a given geographic region, and persist in the face of abiotic conditions and interactions with other species. The evolutionary and colonisation histories of communities are characterised by phylogenetic diversity, while functional diversity is indicative of abiotic and biotic conditions. The relationship between functional and phylogenetic diversity infers whether species functional traits are divergent (differing between related species) or convergent (similar among distantly related species). Biotic interactions and abiotic conditions are known to influence macroecological patterns in species richness, but how functional and phylogenetic diversity of guilds vary with biotic factors, and the relative importance of biotic drivers in relation to geographic and abiotic drivers is unknown. In this study, we test whether geographic, abiotic or biotic factors drive biome‐scale spatial patterns of functional and phylogenetic diversity and functional convergence in vertebrate herbivores across the Arctic tundra biome. We found that functional and phylogenetic diversity both peaked in the western North American Arctic, and that spatial patterns in both were best predicted by trophic interactions, namely vegetation productivity and predator diversity, as well as climatic severity. Our results show that both bottom–up and top–down trophic interactions, as well as winter temperatures, drive the functional and phylogenetic structure of Arctic vertebrate herbivore assemblages. This has implications for changing Arctic ecosystems; under future warming and northward movement of predators potential increases in phylogenetic and functional diversity in vertebrate herbivores may occur. Our study thus demonstrates that trophic interactions can determine large‐scale functional and phylogenetic diversity just as strongly as abiotic conditions.     

No species is an island: testing the effects of biotic interactions on models of avian niche occupation

Traditionally, the niche of a species is described as a hypothetical 3D space, constituted by well‐known biotic interactions (e.g. predation, competition, trophic relationships, resource–consumer interactions, etc.) and various abiotic environmental factors. Species distribution models (SDMs), also called “niche models” and often used to predict wildlife distribution at landscape scale, are typically constructed using abiotic factors with biotic interactions generally been ignored. Here, we compared the goodness of fit of SDMs for red‐backed shrike Lanius collurio in farmlands of Western Poland, using both the classical approach (modeled only on environmental variables) and the approach which included also other potentially associated bird species. The potential associations among species were derived from the relevant ecological literature and by a correlation matrix of occurrences. Our findings highlight the importance of including heterospecific interactions in improving our understanding of niche occupation for bird species. We suggest that suite of measures currently used to quantify realized species niches could be improved by also considering the occurrence of certain associated species. Then, an hypothetical “species 1” can use the occurrence of a successfully established individual of “species 2” as indicator or “trace” of the location of available suitable habitat to breed. We hypothesize this kind of biotic interaction as the “heterospecific trace effect” (HTE): an interaction based on the availability and use of “public information” provided by individuals from different species. Finally, we discuss about the incomes of biotic interactions for enhancing the predictive capacities on species distribution models.     

Intermediate habitat associations by hybrids may facilitate genetic introgression in a songbird

Hybridization or the interbreeding of genetically discrete populations or species can occur where ranges of genetically distinct units overlap. Golden‐winged warblers Vermivora chrysoptera, a species that has been in steady decline for decades, highlight the potential population‐level consequences of hybridization. A major factor implicated in their decline is hybridization with their sister species, the blue‐winged warbler Vermivora cyanoptera, which has likely been exacerbated by historic and current land‐use practices. We examined habitat associations of golden‐winged and blue‐winged warblers, phenotypic hybrids, and cryptic hybrids (i.e. mismatch between plumage phenotype and genotype as identified by mitochondrial DNA) in an area of relatively recent range overlap and hybridization in northern New York, USA. To explore the robustness of these results, we then compared the patterns from New York with habitat associations from the central Pennsylvanian Appalachian Mountains where blue‐winged warblers either do not occur or are in very low abundance, yet cryptic golden‐winged warbler hybrids are present. From 2008 to 2011, we captured 122 birds in New York and 28 in Pennsylvania and collected blood samples, which we used to determine maternal ancestry. For each bird captured, we measured territory‐level (50‐m radius circles) habitat, and later used remote‐sensing data to quantify habitat on the territories and in surrounding areas (100‐, 250‐, and 500‐m radius circles). In New York, golden‐winged warblers occupied structurally heterogeneous territories surrounded by homogeneously structured, contiguous deciduous forest, far from urban areas. Blue‐winged warblers showed opposite associations, and hybrids’ habitat associations were typically intermediate. In Pennsylvania, the habitat associations of golden‐winged warblers and their cryptic hybrids were remarkably similar to those in New York. These findings suggest that patterns of habitat occupancy by hybrids may promote contact with golden‐winged warblers and thus likely facilitate genetic introgression, even in areas where the parental species are not sympatric.     

Mutualism fails when climate response differs between interacting species

Successful species interactions require that both partners share a similar cue. For many species, spring warming acts as a shared signal to synchronize mutualist behaviors. Spring flowering plants and the ants that disperse their seeds respond to warming temperatures so that ants forage when plants drop seeds. However, where warm‐adapted ants replace cold‐adapted ants, changes in this timing might leave early seeds stranded without a disperser. We investigate plant seed dispersal south and north of a distinct boundary between warm‐ and cold‐adapted ants to determine if changes in the ant species influence local plant dispersal. The warm‐adapted ants forage much later than the cold‐adapted ants, and so we first assess natural populations of early and late blooming plants. We then transplant these plants south and north of the ant boundary to test whether distinct ant climate requirements disrupt the ant–plant mutualism. Whereas the early blooming plant's inability to synchronize with the warm‐adapted ant leaves its populations clumped and patchy and its seedlings clustered around the parents in natural populations, when transplanted into the range of the cold‐adapted ant, effective seed dispersal recovers. In contrast, the mutualism persists for the later blooming plant regardless of location because it sets seed later in spring when both warm‐ and cold‐adapted ant species forage, resulting in effective seed dispersal. These results indicate that the climate response of species interactions, not just the species themselves, is integral in understanding ecological responses to a changing climate. Data linking phenological synchrony and dispersal are rare, and these results suggest a viable mechanism by which a species' range is limited more by biotic than abiotic interactions – despite the general assumption that biotic influences are buried within larger climate drivers. These results show that biotic partner can be as fundamental a niche requirement as abiotic resources.     

Are large‐scale distributional shifts of the blue‐winged macaw (Primolius maracana) related to climate change?

Aim To evaluate whether observed geographical shifts in the distribution of the blue‐winged macaw (Primolius maracana) are related to ongoing processes of global climate change. This species is vulnerable to extinction and has shown striking range retractions in recent decades, withdrawing broadly from southern portions of its historical distribution. Its range reduction has generally been attributed to the effects of habitat loss; however, as this species has also disappeared from large forested areas, consideration of other factors that may act in concert is merited. Location Historical distribution of the blue‐winged macaw in Brazil, eastern Paraguay and northern Argentina. Methods We used a correlative approach to test a hypothesis of causation of observed shifts by reduction of habitable areas mediated by climate change. We developed models of the ecological niche requirements of the blue‐winged macaw, based on point‐occurrence data and climate scenarios for pre‐1950 and post‐1950 periods, and tested model predictivity for anticipating geographical distributions within time periods. Then we projected each model to the other time period and compared distributions predicted under both climate scenarios to assess shifts of habitable areas across decades and to evaluate an explanation for observed range retractions. Results Differences between predicted distributions of the blue‐winged macaw over the twentieth century were, in general, minor and no change in suitability of landscapes was predicted across large areas of the species’ original range in different time periods. No tendency towards range retraction in the south was predicted, rather conditions in the southern part of the species’ range tended to show improvement for the species. Main conclusions Our test permitted elimination of climate change as a likely explanation for the observed shifts in the distribution of the blue‐winged macaw, and points rather to other causal explanations (e.g. changing regional land use, emerging diseases).     

Local‐scale biotic interactions embedded in macroscale climate drivers suggest Eltonian noise hypothesis distribution patterns for an invasive grass

A hierarchical view of niche relations reconciles the scale‐dependent effects of abiotic and biotic processes on species distribution patterns and underlies most current approaches to distribution modeling. A key prediction of this framework is that the effects of biotic interactions will be averaged out at macroscales – an idea termed the Eltonian noise hypothesis (ENH). We test this prediction by quantifying regional variation in local abiotic and biotic niche relations and assess the role of macroclimate in structuring biotic interactions, using a non‐native invasive grass, Microstegium vimineum, in its introduced range. Consistent with hierarchical niche relations and the ENH, macroclimate structures local biotic interactions, while local abiotic relations are regionally conserved. Biotic interactions suppress M. vimineum in drier climates but have little effect in wetter climates. A similar approach could be used to identify the macroclimatic conditions under which biotic interactions affect the accuracy of local predictions of species distributions.     

Multiple environmental stressors increase the realised niche breadth of a forest‐dwelling fish

Understanding the determinants of species' niche breadth is important due to the negative relationship between niche breadth and extinction probability. Species tolerant to extremely harsh abiotic conditions (e.g. low pH or hypoxia) often have relatively small realised niches due to a trade‐off between abiotic and biotic stress tolerance, and are therefore particularly vulnerable to extinction. We hypothesised that tolerance to multiple extreme conditions (e.g. low pH, hypoxia and drought) in brown mudfish Neochanna apoda, would increase their realised niche breadth because each extreme would provide additive refuge against a dominant species, the banded kokopu Galaxias fasciatus. Fish distributions were surveyed in 65 peat‐swamp‐forest streams and pools in New Zealand, which varied in dissolved oxygen, acidity and hydroperiod. Mudfish distribution was extremely patchy, and almost completely allopatric with kokopu. Allopatry was driven mainly by pool hydroperiod; mudfish occupied 88 percent of temporary pools due to their tolerance of habitat drying, whereas kokopu were absent from temporary pools. Within permanent pools, mudfish occurrence was negatively related to pool oxygen and pH, whereas kokopu occurrence was positively related to these conditions. Pool conditions were independently distributed in the landscape, suggesting that each abiotic stressor offered additive refuge for mudfish from kokopu predation/competition. Consequently, the mudfish realised niche breadth depended on the number of abiotic factors driving their allopatry with kokopu. Given the widespread negative relationships between niche breadth and species extinction probabilities, our results indicate that tolerance to multiple stressors may play an important role in insuring species persistence against the multiple drivers of global environmental change.     

Microhabitat selection in the common lizard: implications of biotic interactions,age, sex,local processes,and model transferability among populations

Modeling species' habitat requirements are crucial to assess impacts of global change, for conservation efforts and to test mechanisms driving species presence. While the influence of abiotic factors has been widely examined, the importance of biotic factors and biotic interactions, and the potential implications of local processes are not well understood. Testing their importance requires additional knowledge and analyses at local habitat scale. Here, we recorded the locations of species presence at the microhabitat scale and measured abiotic and biotic parameters in three different common lizard (Zootoca vivipara) populations using a standardized sampling protocol. Thereafter, space use models and cross‐evaluations among populations were run to infer local processes and estimate the importance of biotic parameters, biotic interactions, sex, and age. Biotic parameters explained more variation than abiotic parameters, and intraspecific interactions significantly predicted the spatial distribution. Significant differences among populations in the relationship between abiotic parameters and lizard distribution, and the greater model transferability within populations than between populations are in line with effects predicted by local adaptation and/or phenotypic plasticity. These results underline the importance of including biotic parameters and biotic interactions in space use models at the population level. There were significant differences in space use between sexes, and between adults and yearlings, the latter showing no association with the measured parameters. Consequently, predictive habitat models at the population level taking into account different sexes and age classes are required to understand a specie's ecological requirements and to allow for precise conservation strategies. Our study therefore stresses that future predictive habitat models at the population level and their transferability should take these parameters into account.     

Experimental assessment of biotic and abiotic filters driving community composition

Species occurrence in a site can be limited by both the abiotic environment and biotic interactions. These two factors operate in concert, but their relative importance is often unclear. By experimentally introducing seeds or plants into competition‐free gaps or into the intact vegetation, we can disentangle the biotic and abiotic effects on plant establishment. We established a seed‐sowing/transplant experiment in three different meadows. Species were introduced, as seeds and pregrown transplants, into competition‐free gaps and the intact vegetation. They included 12 resident plants from the locality and 18 species typical for different habitats. Last two years, gaps were overgrown with vegetation from surrounding plants and we observed the competitive exclusion of our focal plants. We compared plant survival with the expected occurrence in target locality (Beals index). Many of the species with habitat preferences different from our localities were able to successfully establish from seeds and grow in the focal habitat if competition was removed. They included species typical for much drier conditions. These species were thus not limited by the abiotic conditions, but by competition. Pregrown transplants were less sensitive to competition, when compared to seedlings germinated from seeds. Beals index significantly predicted both species success in gaps and the ability to withstand competition. Survival in a community is dependent on the adaptation to both the abiotic environment and biotic interactions. Statistically significant correlation coefficients of the ratio of seedling survival in vegetation and gaps with Beals index suggest the importance of biotic interactions as a determinant of plant community composition. To disentangle the importance of abiotic and biotic effect on plant establishment, it is important to distinguish between species pool as a set of species typically found in given community type (determined by Beals index) and a set of species for which the abiotic conditions are suitable.     

Joint effects of species traits and environmental preferences on range edge shifts of British birds

Aim

Despite the strong evidence of species range shifts as a response to environmental change, attempts to identify species traits that modulate those shifts have been equivocal. We investigate the role of species traits and environmental preferences on birds' range shifts in Great Britain, an island where dispersal is limited by the English Channel and the North Sea.

Location

Great Britain (England, Scotland and Wales).

Taxa

Birds (Aves).

Time Period

1968–2011.

Methods

Using 404,949 occurrence records from two time periods, we investigated the potential drivers of leading and rear range edge shifts of breeding birds using phylogenetic linear mixed models. We hypothesized that shifts are influenced by species' trophic and morphological traits, dispersal abilities and environmental preferences, but also by the geographical boundaries of Great Britain.

Results

Geographical boundaries—the distance from the northern or southern boundaries of Britain—accounted for most of the variability in range edge shifts. Species traits and environmental preferences emerged as relevant drivers of range shifts only for northern and Passeriform species. Northern habitat specialist, those with more predators and those sensitive to precipitation were more likely to shift their rear edge poleward. For Passeriformes, habitat generalists, species with smaller dispersal capabilities, under higher predatory pressure or associated with forest and grassland were more likely to shift their rear edge poleward.

Main Conclusions

While geographical boundaries impose constraints on range shifts in British birds, the subtle effects of species traits and environmental preferences emerge as relevant predictors for Northern and passeriform species' rear edge shifts. This highlights the importance of accounting for geographical boundaries when predicting species responses to global change. Differential range shifts of species across different trophic levels could result in the reorganization of biotic interactions, with consequences for ecosystem structure and stability.     

Maladaptation beyond a geographic range limit driven by antagonistic and mutualistic biotic interactions across an abiotic gradient

Species’ geographic range limits often result from maladaptation to the novel environments beyond the range margin. However, we rarely know which aspects of the n‐dimensional environment are driving this maladaptation. Especially of interest is the influence of abiotic versus biotic factors in delimiting species’ distributions. We conducted a 2‐year reciprocal transplant experiment involving manipulations of the biotic environment to explore how spatiotemporal gradients in precipitation, fatal mammalian herbivory, and pollination affected lifetime fitness within and beyond the range of the California annual plant, Clarkia xantiana ssp. xantiana. In the first, drier year of the experiment, fitness outside the range edge was limited mainly by low precipitation, and there was some evidence for local adaptation within the range. In the second, wetter year, we did not observe abiotic limitations to plant fitness outside the range; instead biotic interactions, especially herbivory, limited fitness outside the range. Together, protection from herbivory and supplementation of pollen resulted in three‐ to sevenfold increases in lifetime fitness outside the range margin in the abiotically benign year. Overall, our work demonstrates the importance of biotic interactions, particularly as they interact with the abiotic environment, in determining fitness beyond geographic range boundaries.     

Sympatry leads to reduced body condition in chickadees that occasionally hybridize

Both abiotic and biotic drivers influence species distributions. Abiotic drivers such as climate have received considerable attention, even though biotic drivers such as hybridization often interact with abiotic drivers. We sought to explore the (1) costs of co‐occurrence for ecologically similar species that hybridize and (2) associations between ecological factors and condition to understand how abiotic and biotic factors influence species distributions. For two closely related and ecologically similar songbirds, black‐capped and mountain chickadees, we characterized body condition, as a proxy for fitness, using a 1358‐individual range‐wide dataset. We compared body condition in sympatry and allopatry with several abiotic and biotic factors using species‐specific generalized linear mixed models. We generated genomic data for a subset of 217 individuals to determine the extent of hybridization‐driven admixture in our dataset. Within this data subset, we found that ~11% of the chickadees had hybrid ancestry, and all hybrid individuals had typical black‐capped chickadee plumage. In the full dataset, we found that birds of both species, independent of demographic and abiotic factors, had significantly lower body condition when occurring in sympatry than birds in allopatry. This could be driven by either the inclusion of cryptic, likely poor condition, hybrids in our full dataset, competitive interactions in sympatry, or range edge effects. We are currently unable to discriminate between these mechanisms. Our findings have implications for mountain chickadees in particular, which will encounter more black‐capped chickadees as black‐capped chickadee ranges shift upslope and could lead to local declines in mountain chickadee populations.