Ecological barriers mediate spatiotemporal shifts of bird communities at a continental scale.

Species' range shifts and local extinctions caused by climate change lead to community composition changes. At large spatial scales, ecological barriers, such as biome boundaries, coastlines, and elevation, can influence a community's ability to shift in response to climate change. Yet, ecological barriers are rarely considered in climate change studies, potentially hindering predictions of biodiversity shifts. We used data from two consecutive European breeding bird atlases to calculate the geographic distance and direction between communities in the 1980s and their compositional best match in the 2010s and modeled their response to barriers. The ecological barriers affected both the distance and direction of bird community composition shifts, with coastlines and elevation having the strongest influence. Our results underscore the relevance of combining ecological barriers and community shift projections for identifying the forces hindering community adjustments under global change. Notably, due to (macro)ecological barriers, communities are not able to track their climatic niches, which may lead to drastic changes, and potential losses, in community compositions in the future.

Drivers of the changing abundance of European birds at two spatial scales.

Detecting biodiversity change and identifying its causes is challenging because biodiversity is multifaceted and temporal data often contain bias. Here, we model temporal change in species' abundance and biomass by using extensive data describing the population sizes and trends of native breeding birds in the United Kingdom (UK) and the European Union (EU). In addition, we explore how species' population trends vary with species' traits. We demonstrate significant change in the bird assemblages of the UK and EU, with substantial reductions in overall bird abundance and losses concentrated in a relatively small number of abundant and smaller sized species. By contrast, rarer and larger birds had generally fared better. Simultaneously, overall avian biomass had increased very slightly in the UK and was stable in the EU, indicating a change in community structure. Abundance trends across species were positively correlated with species' body mass and with trends in climate suitability, and varied with species' abundance, migration strategy and niche associations linked to diet. Our work highlights how changes in biodiversity cannot be captured easily by a single number; care is required when measuring and interpreting biodiversity change given that different metrics can provide very different insights. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

Projected climate change impacts on the phylogenetic diversity of the world's terrestrial birds: more than species numbers.

Ongoing climate change is a major threat to biodiversity. As abiotic tolerances and dispersal abilities vary, species-specific responses have the potential to further amplify or ameliorate the ensuing impacts on species assemblages. Here, we investigate the effects of climate change on species distributions across non-marine birds, quantifying its projected impact on species richness (SR) as well as on different aspects of phylogenetic diversity globally. Going beyond previous work, we disentangle the potential impacts of species gains versus losses on assemblage-level phylogenetic diversity under climate change and compare the projected impacts to randomized assemblage changes. We show that beyond its effects on SR, climate change could have profound impacts on assemblage-level phylogenetic diversity and composition, which differ significantly from random changes and among regions. Though marked species losses are most frequent in tropical and subtropical areas in our projections, phylogenetic restructuring of species communities is likely to occur all across the globe. Furthermore, our results indicate that the most severe changes to the phylogenetic diversity of local assemblages are likely to be caused by species range shifts and local species gains rather than range reductions and extinctions. Our findings highlight the importance of considering diverse measures in climate impact assessments.

Global impacts of climate change on avian functional diversity.

Climate change is predicted to drive geographical range shifts, leading to fluctuations in species richness (SR) worldwide. However, the effect of these changes on functional diversity (FD) remains unclear, in part because comprehensive species-level trait data are generally lacking at global scales. Here, we use morphometric and ecological traits for 8268 bird species to estimate the impact of climate change on avian FD. We show that future bird assemblages are likely to undergo substantial shifts in trait structure, with a magnitude of change greater than predicted from SR alone, and a direction of change varying according to geographical location and trophic guild. For example, our models predict that FD of insect predators will increase at higher latitudes with concurrent losses at mid-latitudes, whereas FD of seed dispersing birds will fluctuate across the tropics. Our findings highlight the potential for climate change to drive continental-scale shifts in avian FD with implications for ecosystem function and resilience.

Phenological trends in the pre- and post-breeding migration of long-distance migratory birds.

Phenological mismatch is often cited as a putative driver of population declines in long-distance migratory birds. The mechanisms and cues utilized to advance breeding ground arrival will impact the adaptability of species to further warming. Furthermore, timing of post-breeding migration potentially faces diverging selective pressures, with earlier onset of tropical dry seasons favouring migration advancement, while longer growing seasons in temperate areas could facilitate delayed departures. Despite this, few studies exist of migration phenology on the non-breeding grounds or on post-breeding passage. Here, we use first arrival and last departure dates of 20 species of trans-Saharan migratory birds from tropical non-breeding grounds (The Gambia), between 1964 and 2019. Additionally, we use first arrival and last departure dates, as well as median arrival and departure dates, at an entry/departure site to/from Europe (Gibraltar), between 1991 and 2018. We assess phenological trends in pre- and post-breeding migration, as well as individual species' durations of stay in breeding and non-breeding areas. Furthermore, we assess the extent to which inter-annual variation in these timings may be explained by meteorological and ecological variables. We find significant advances in pre-breeding migration at both locations, while post-breeding migration is delayed. At Gibraltar, these trends do not differ between first/last and median dates of migration. The combination of these trends suggests substantial changes in the temporal usage of the two continents by migratory birds. Duration of stay (of species, not individuals) within Europe increased by 16 days, on average, over the 27-year monitoring period. By contrast, duration of species' stays on the non-breeding range declined by 63 days, on average, over the 56-year monitoring period. Taken together these changes suggest substantial, previously unreported alterations to annual routines in Afro-Palaearctic migrants.

Global inequities and political borders challenge nature conservation under climate change.

Underlying sociopolitical factors have emerged as important determinants of wildlife population trends and the effectiveness of conservation action. Despite mounting research into the impacts of climate change on nature, there has been little consideration of the human context in which these impacts occur, particularly at the global scale. We investigate this in two ways. First, by modeling the climatic niches of terrestrial mammals and birds globally, we show that projected species loss under climate change is greatest in countries with weaker governance and lower Gross Domestic Product, with loss of mammal species projected to be greater in countries with lower CO2 emissions. Therefore, climate change impacts on species may be disproportionately significant in countries with lower capacity for effective conservation and lower greenhouse gas emissions, raising important questions of international justice. Second, we consider the redistribution of species in the context of political boundaries since the global importance of transboundary conservation under climate change is poorly understood. Under a high-emissions scenario, we find that 35% of mammals and 29% of birds are projected to have over half of their 2070 climatic niche in countries in which they are not currently found. We map these transboundary range shifts globally, identifying borders across which international coordination might most benefit conservation and where physical border barriers, such as walls and fences, may be an overlooked obstacle to climate adaptation. Our work highlights the importance of sociopolitical context and the utility of a supranational perspective for 21st century nature conservation.

Burning savanna for avian species richness and functional diversity.

Prescribed fire is used throughout fire-prone landscapes to conserve biodiversity. Current best practice in managing savanna systems advocates methods based on the assumption that increased fire-mediated landscape heterogeneity (pyrodiversity) will promote biodiversity. However, considerable knowledge gaps remain in our understanding of how savanna wildlife responds to the composition and configuration of pyrodiverse landscapes. The effects of pyrodiversity on functional diversity have rarely been quantified and assessing this relationship at a landscape scale that is commensurate with fire management is important for understanding mechanisms underlying ecosystem resilience. Here, we assess the impact of spatiotemporal variation in a long-term fire regime on avian diversity in North West Province, South Africa. We examined the relationship between (1) species richness, (2) three indices of functional diversity (i.e., functional richness, functional evenness, and functional dispersion) and four measures of pyrodiversity, the spatial extents of fire age classes, and habitat type at the landscape scale. We then used null models to assess differences between observed and expected functional diversity. We found that the proportion of newly burned (<1-yr post-fire), old, unburned (≥10 yr post-fire), and woodland habitat on the landscape predicted species and functional richness. Species richness also increased with the degree of edge contrast between patches of varying fire age, while functional dispersion increased with the degree of patch shape complexity. Lower than expected levels of functional richness suggest that habitat filtering is occurring, resulting in functional redundancy across our study sites. We demonstrate that evaluating functional diversity and redundancy is an important component of conservation planning as they may contribute to previously reported fire resilience. Our findings suggest that it is the type and configuration, rather than the diversity, of fire patches on the landscape that promote avian diversity and conserve ecological functions. A management approach is needed that includes significant coverage of adjacent newly burned and older, unburned savanna habitat; the latter, in particular, is inadequately represented under current burning practices.

The influence of different aspects of grouse moorland management on nontarget bird assemblages.

Conflict between stakeholders with opposing interests can hamper biodiversity conservation. When conflicts become entrenched, evidence from applied ecology can reveal new ways forward for their management. In particular, where disagreement exists over the efficacy or ethics of management actions, research clarifying the uncertain impacts of management on wildlife can move debates forwards to conciliation.Here, we explore a case-study of entrenched conflict where uncertainty exists over the impacts of multiple management actions: namely, moorlands managed for the shooting of red grouse (willow ptarmigan) Lagopus lagopus in the United Kingdom (UK). Debate over how UK moorlands should be managed is increasingly polarized. We evaluate, for the first time at a regional scale, the relative impacts of two major moorland management practices-predator control and heather burning-on nontarget bird species of conservation concern.Birds were surveyed on 18 estates across Northern England and Southeast Scotland. Sites ranged from intensively managed grouse moors to moorland sites with no management for grouse shooting. We hypothesised that both targeted predator control and burning regimes would enhance ground-nesting wader numbers and, as a consequence of this, and of increased grouse numbers, nontarget avian predators should also be more abundant on heavily managed sites.There were positive associations between predator control and the abundance of the three most widespread species of ground-nesting wader: strong effects for European golden plover Pluvialis apricaria and Eurasian curlew Numenius arquata and, less strongly, for common snipe Gallinago gallinago. These effects saturated at low levels of predator control. Evidence for effects of burning was much weaker. We found no evidence of enhanced numbers of nontarget predators on heavily managed sites.

The limits to population density in birds and mammals.

We address two fundamental ecological questions: what are the limits to animal population density and what determines those limits? We develop simple alternative models to predict population limits in relation to body mass. A model assuming that within-species area use increases with the square of daily travel distance broadly predicts the scaling of empirical extremes of minimum density across birds and mammals. Consistent with model predictions, the estimated density range for a given mass, 'population scope', is greater for birds than for mammals. However, unlike mammals and carnivorous birds, expected broad relationships between body mass and density extremes are not supported by data on herbivorous and omnivorous birds. Our results suggest that simple constraints on mobility and energy use/supply are major determinants of the scaling of density limits, but further understanding of interactions between dietary constraints and density limits are needed to predict future wildlife population responses to anthropogenic threats.

Bioenergy cropland expansion may offset positive effects of climate change mitigation for global vertebrate diversity.

Climate and land-use change interactively affect biodiversity. Large-scale expansions of bioenergy have been suggested as an important component for climate change mitigation. Here we use harmonized climate and land-use projections to investigate their potential combined impacts on global vertebrate diversity under a low- and a high-level emission scenario. We combine climate-based species distribution models for the world's amphibians, birds, and mammals with land-use change simulations and identify areas threatened by both climate and land-use change in the future. The combined projected effects of climate and land-use change on vertebrate diversity are similar under the two scenarios, with land-use change effects being stronger under the low- and climate change effects under the high-emission scenario. Under the low-emission scenario, increases in bioenergy cropland may cause severe impacts in biodiversity that are not compensated by lower climate change impacts. Under this low-emission scenario, larger proportions of species distributions and a higher number of small-range species may become impacted by the combination of land-use and climate change than under the high-emission scenario, largely a result of bioenergy cropland expansion. Our findings highlight the need to carefully consider both climate and land-use change when projecting biodiversity impacts. We show that biodiversity is likely to suffer severely if bioenergy cropland expansion remains a major component of climate change mitigation strategies. Our study calls for an immediate and significant reduction in energy consumption for the benefit of both biodiversity and to achieve the goals of the Paris Agreement.

Tritrophic phenological match-mismatch in space and time.

Increasing temperatures associated with climate change may generate phenological mismatches that disrupt previously synchronous trophic interactions. Most work on mismatch has focused on temporal trends, whereas spatial variation in the degree of trophic synchrony has largely been neglected, even though the degree to which mismatch varies in space has implications for meso-scale population dynamics and evolution. Here we quantify latitudinal trends in phenological mismatch, using phenological data on an oak-caterpillar-bird system from across the UK. Increasing latitude delays phenology of all species, but more so for oak, resulting in a shorter interval between leaf emergence and peak caterpillar biomass at northern locations. Asynchrony found between peak caterpillar biomass and peak nestling demand of blue tits, great tits and pied flycatchers increases in earlier (warm) springs. There is no evidence of spatial variation in the timing of peak nestling demand relative to peak caterpillar biomass for any species. Phenological mismatch alone is thus unlikely to explain spatial variation in population trends. Given projections of continued spring warming, we predict that temperate forest birds will become increasingly mismatched with peak caterpillar timing. Latitudinal invariance in the direction of mismatch may act as a double-edged sword that presents no opportunities for spatial buffering from the effects of mismatch on population size, but generates spatially consistent directional selection on timing, which could facilitate rapid evolutionary change.

Flight range, fuel load and the impact of climate change on the journeys of migrant birds.

Climate change is predicted to increase migration distances for many migratory species, but the physiological and temporal implications of longer migratory journeys have not been explored. Here, we combine information about species' flight range potential and migratory refuelling requirements to simulate the number of stopovers required and the duration of current migratory journeys for 77 bird species breeding in Europe. Using tracking data, we show that our estimates accord with recorded journey times and stopovers for most species. We then combine projections of altered migratory distances under climate change with models of avian flight to predict future migratory journeys. We find that 37% of migratory journeys undertaken by long-distance migrants will necessitate an additional stopover in future. These greater distances and the increased number of stops will substantially increase overall journey durations of many long-distance migratory species, a factor not currently considered in climate impact studies.

Consistent response of bird populations to climate change on two continents.

Global climate change is a major threat to biodiversity. Large-scale analyses have generally focused on the impacts of climate change on the geographic ranges of species and on phenology, the timing of ecological phenomena. We used long-term monitoring of the abundance of breeding birds across Europe and the United States to produce, for both regions, composite population indices for two groups of species: those for which climate suitability has been either improving or declining since 1980. The ratio of these composite indices, the climate impact indicator (CII), reflects the divergent fates of species favored or disadvantaged by climate change. The trend in CII is positive and similar in the two regions. On both continents, interspecific and spatial variation in population abundance trends are well predicted by climate suitability trends.

Choice of baseline climate data impacts projected species' responses to climate change.

Climate data created from historic climate observations are integral to most assessments of potential climate change impacts, and frequently comprise the baseline period used to infer species-climate relationships. They are often also central to downscaling coarse resolution climate simulations from General Circulation Models (GCMs) to project future climate scenarios at ecologically relevant spatial scales. Uncertainty in these baseline data can be large, particularly where weather observations are sparse and climate dynamics are complex (e.g. over mountainous or coastal regions). Yet, importantly, this uncertainty is almost universally overlooked when assessing potential responses of species to climate change. Here, we assessed the importance of historic baseline climate uncertainty for projections of species' responses to future climate change. We built species distribution models (SDMs) for 895 African bird species of conservation concern, using six different climate baselines. We projected these models to two future periods (2040-2069, 2070-2099), using downscaled climate projections, and calculated species turnover and changes in species-specific climate suitability. We found that the choice of baseline climate data constituted an important source of uncertainty in projections of both species turnover and species-specific climate suitability, often comparable with, or more important than, uncertainty arising from the choice of GCM. Importantly, the relative contribution of these factors to projection uncertainty varied spatially. Moreover, when projecting SDMs to sites of biodiversity importance (Important Bird and Biodiversity Areas), these uncertainties altered site-level impacts, which could affect conservation prioritization. Our results highlight that projections of species' responses to climate change are sensitive to uncertainty in the baseline climatology. We recommend that this should be considered routinely in such analyses.

Nationwide trophic cascades: changes in avian community structure driven by ungulates.

In recent decades, many ungulate populations have changed dramatically in abundance, resulting in cascading effects across ecosystems. However, studies of such effects are often limited in their spatial and temporal scope. Here, we contrast multi-species composite population trends of deer-sensitive and deer-tolerant woodland birds at a national scale, across Britain. We highlight the divergent fates of these two groups between 1994 and 2011, and show a striking association between the calculated divergence and a composite population trend of woodland deer. Our results demonstrate the link between changes in deer populations and changes in bird communities. In a period when composite population trends for deer increased by 46%, the community population trend across deer-sensitive birds (those dependent on understory vegetation) declined much more than the community trend for deer-tolerant birds. Our findings suggest that ongoing changes in the populations of herbivorous ungulates in many countries worldwide may help explain patterns of community restructuring at other trophic levels. Ungulate impacts on other taxa may require more consideration by conservation practitioners than they currently receive.

Predicting the Spatial Distribution of Wolf (Canis lupus) Breeding Areas in a Mountainous Region of Central Italy.

Wolves (Canis lupus) in Italy represent a relict west European population. They are classified as vulnerable by IUCN, though have increased in number and expanded their range in recent decades. Here we use 17 years of monitoring data (from 1993 to 2010) collected in a mountainous region of central Italy (Arezzo, Tuscany) in an ecological niche-based model (MaxEnt) to characterize breeding sites (i.e. the areas where pups were raised) within home ranges, as detected from play-back responses. From a suite of variables related to topography, habitat and human disturbance we found that elevation and distance to protected areas were most important in explaining the locality of wolf responses. Rendezvous sites (family play-back response sites) typically occurred between 800 and 1200 m a.s.l., inside protected areas, and were usually located along mountain chains distant from human settlements and roads. In these areas human disturbance is low and the densities of ungulates are typically high. Over recent years, rendezvous sites have occurred closer to urban areas as the wolf population has continued to expand, despite the consequent human disturbance. This suggests that undisturbed landscapes may be reaching their carrying capacity for wolves. This, in turn, may lead to the potential for increased human-wolf interactions in future. Applying our model, both within and beyond the species' current range, we identify sites both within the current range and also further afield, that the species could occupy in future. Our work underlines the importance of the present protected areas network in facilitating the recolonisation by wolves. Our projections of suitability of sites for future establishment as the population continues to expand could inform planning to minimize future wolf-human conflicts.

Predicting potential responses to future climate in an alpine ungulate: interspecific interactions exceed climate effects.

The altitudinal shifts of many montane populations are lagging behind climate change. Understanding habitual, daily behavioural rhythms, and their climatic and environmental influences, could shed light on the constraints on long-term upslope range-shifts. In addition, behavioural rhythms can be affected by interspecific interactions, which can ameliorate or exacerbate climate-driven effects on ecology. Here, we investigate the relative influences of ambient temperature and an interaction with domestic sheep (Ovis aries) on the altitude use and activity budgets of a mountain ungulate, the Alpine chamois (Rupicapra rupicapra). Chamois moved upslope when it was hotter but this effect was modest compared to that of the presence of sheep, to which they reacted by moving 89-103 m upslope, into an entirely novel altitudinal range. Across the European Alps, a range-shift of this magnitude corresponds to a 46% decrease in the availability of suitable foraging habitat. This highlights the importance of understanding how factors such as competition and disturbance shape a given species' realised niche when predicting potential future responses to change. Furthermore, it exposes the potential for manipulations of species interactions to ameliorate the impacts of climate change, in this case by the careful management of livestock. Such manipulations could be particularly appropriate for species where competition or disturbance already strongly restricts their available niche. Our results also reveal the potential role of behavioural flexibility in responses to climate change. Chamois reduced their activity when it was warmer, which could explain their modest altitudinal migrations. Considering this behavioural flexibility, our model predicts a small 15-30 m upslope shift by 2100 in response to climate change, less than 4% of the altitudinal shift that would be predicted using a traditional species distribution model-type approach (SDM), which assumes that species' behaviour remains unchanged as climate changes. Behavioural modifications could strongly affect how species respond to a changing climate.

Evaluating the effectiveness of conservation site networks under climate change: accounting for uncertainty.

We forecasted potential impacts of climate change on the ability of a network of key sites for bird conservation (Important Bird Areas; IBAs) to provide suitable climate for 370 bird species of current conservation concern in two Asian biodiversity hotspots: the Eastern Himalaya and Lower Mekong. Comparable studies have largely not accounted for uncertainty, which may lead to inappropriate conclusions. We quantified the contribution of four sources of variation (choice of general circulation models, emission scenarios and species distribution modelling methods and variation in species distribution data) to uncertainty in forecasts and tested if our projections were robust to these uncertainties. Declines in the availability of suitable climate within the IBA network by 2100 were forecast as 'extremely likely' for 45% of species, whereas increases were projected for only 2%. Thus, we predict almost 24 times as many 'losers' as 'winners'. However, for no species was suitable climate 'extremely likely' to be completely lost from the network. Considerable turnover (median = 43%, 95% CI = 35-69%) in species compositions of most IBAs were projected by 2100. Climatic conditions in 47% of IBAs were projected as 'extremely likely' to become suitable for fewer priority species. However, no IBA was forecast to become suitable for more species. Variation among General Circulation Models and Species Distribution Models contributed most to uncertainty among forecasts. This uncertainty precluded firm conclusions for 53% of species and IBAs because 95% confidence intervals included projections of no change. Considering this uncertainty, however, allows robust recommendations concerning the remaining species and IBAs. Overall, while the IBA network will continue to sustain bird conservation, climate change will modify which species each site will be suitable for. Thus, adaptive management of the network, including modified site conservation strategies and facilitating species' movement among sites, is critical to ensure effective future conservation.

Foraging ranges of immature African white-backed vultures (Gyps africanus) and their use of protected areas in southern Africa.

Vultures in the Gyps genus are declining globally. Multiple threats related to human activity have caused widespread declines of vulture populations in Africa, especially outside protected areas. Addressing such threats requires the estimation of foraging ranges yet such estimates are lacking, even for widespread (but declining) species such as the African white-backed vulture (Gyps africanus). We tracked six immature African white-backed vultures in South Africa using GPS-GSM units to study their movement patterns, their use of protected areas and the time they spent in the vicinity of supplementary feeding sites. All individuals foraged widely; their combined foraging ranges extended into six countries in southern Africa (mean (± SE) minimum convex polygon area =269,103±197,187 km(2)) and three of the vultures travelled more than 900 km from the capture site. All six vultures spent the majority of their tracking periods outside protected areas. South African protected areas were very rarely visited whereas protected areas in northern Botswana and Zimbabwe were used more frequently. Two of the vultures visited supplementary feeding sites regularly, with consequent reduced ranging behaviour, suggesting that individuals could alter their foraging behaviour in response to such sites. We show that immature African white-backed vultures are capable of travelling throughout southern Africa, yet use protected areas to only a limited extent, making them susceptible to the full range of threats in the region. The standard approach of designating protected areas to conserve species is unlikely to ensure the protection of such wide-ranging species against threats in the wider landscape.

Intraseasonal variation in reproductive effort: young males finish last.

Age-dependent reproductive timing has been observed in females of a number of species; older females often breed earlier in the season and experience higher reproductive success as a result. However, to date, evidence for within-season variation in reproductive effort (RE) for males has been relatively weak. Males are expected to time RE in light of intraseasonal variations in the availability of receptive females and competition with other males. Young males, which are typically smaller and less experienced, might benefit from breeding later in the season, when male-male competition is less intense. Using a long-term data set of Alpine chamois Rupicapra rupicapra, we sought to evaluate the hypothesis that younger males allocate highest RE late in the breeding season, at a time when older male RE has decreased substantially. Our results support this hypothesis, which suggests that intraseasonal variation in RE may be an adaptive life-history trait for males as well as females.