Twenty‐five years of change in southern African passerine diversity: nonclimatic factors of change

We analysed more than 25 years of change in passerine bird distribution in South Africa, Swaziland and Lesotho, to show that species distributions can be influenced by processes that are at least in part independent of the local strength and direction of climate change: land use and ecological succession. We used occupancy models that separate species' detection from species' occupancy probability, fitted to citizen science data from both phases of the Southern African Bird Atlas Project (1987–1996 and 2007–2013). Temporal trends in species' occupancy probability were interpreted in terms of local extinction/colonization, and temporal trends in detection probability were interpreted in terms of change in abundance. We found for the first time at this scale that, as predicted in the context of bush encroachment, closed‐savannah specialists increased where open‐savannah specialists decreased. In addition, the trend in the abundance of species a priori thought to be favoured by agricultural conversion was negatively correlated with human population density, which is in line with hypotheses explaining the decline in farmland birds in the Northern Hemisphere. In addition to climate, vegetation cover and the intensity and time since agricultural conversion constitute important predictors of biodiversity changes in the region. Their inclusion will improve the reliability of predictive models of species distribution.     

Population dynamics can be more important than physiological limits for determining range shifts under climate change

Evidence is accumulating that species' responses to climate changes are best predicted by modelling the interaction of physiological limits, biotic processes and the effects of dispersal‐limitation. Using commercially harvested blacklip (Haliotis rubra) and greenlip abalone (Haliotis laevigata) as case studies, we determine the relative importance of accounting for interactions among physiology, metapopulation dynamics and exploitation in predictions of range (geographical occupancy) and abundance (spatially explicit density) under various climate change scenarios. Traditional correlative ecological niche models (ENM) predict that climate change will benefit the commercial exploitation of abalone by promoting increased abundances without any reduction in range size. However, models that account simultaneously for demographic processes and physiological responses to climate‐related factors result in future (and present) estimates of area of occupancy (AOO) and abundance that differ from those generated by ENMs alone. Range expansion and population growth are unlikely for blacklip abalone because of important interactions between climate‐dependent mortality and metapopulation processes; in contrast, greenlip abalone should increase in abundance despite a contraction in AOO. The strongly non‐linear relationship between abalone population size and AOO has important ramifications for the use of ENM predictions that rely on metrics describing change in habitat area as proxies for extinction risk. These results show that predicting species' responses to climate change often require physiological information to understand climatic range determinants, and a metapopulation model that can make full use of this data to more realistically account for processes such as local extirpation, demographic rescue, source‐sink dynamics and dispersal‐limitation.     

Is climate change causing the range contraction of Cape Rock-jumpers (Chaetops frenatus)?

Species distribution models often suggest strong links between climate and species' distribution boundaries and project large distribution shifts in response to climate change. However, attributing distribution shifts to climate change requires more than correlative models. One idea is to examine correlates of the processes that cause distribution shifts, namely colonization and local extinction, by using dynamic occupancy models. The Cape Rock-jumper (Chaetops frenatus) has disappeared over most of its distribution where temperatures are the highest. We used dynamic occupancy models to analyse Cape Rock-jumper distribution with respect to climate (mean temperature and precipitation over the warmest annual quarter), vegetation (proportion of natural vegetation, fynbos) and land-use type (protected areas). Detection/non-detection data were collected over two phases of the Southern African Bird Atlas Project (SABAP): 1987–1991 (SABAP1) and 2008–2014 (SABAP2). The model described the contraction of the Cape Rock-jumper's distribution between SABAP1 and SABAP2 well. Occupancy probability during SABAP1 increased with the proportion of fynbos and protected area per grid cell, and decreased with increases in mean temperature and precipitation over the warmest annual quarter. Mean extinction probability increased with mean temperature and precipitation over the warmest annual quarter, although the associated confidence intervals were wide. Nonetheless, our results showed a clear correlation between climate and the distribution boundaries of the Cape Rock-jumper, and in particular, the species' aversion for higher temperatures. The data were less conclusive on whether the observed range contraction was linked to climate change or not. Examining the processes underlying distribution shifts requires large datasets and should lead to a better understanding of the drivers of these shifts.     

Range expansion through fragmented landscapes under a variable climate

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high‐resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine‐resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine‐scale, short‐term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.     

Directionality of recent bird distribution shifts and climate change in Great Britain

There is good evidence that species' distributions are shifting poleward in response to climate change and wide interest in the magnitude of such responses for scientific and conservation purposes. It has been suggested from the directions of climatic changes that species' distribution shifts may not be simply poleward, but this has been rarely tested with observed data. Here, we apply a novel approach to measuring range shifts on axes ranging through 360°, to recent data on the distributions of 122 species of British breeding birds during 1988–1991 and 2008–2011. Although previously documented poleward range shifts have continued, with an average 13.5 km shift northward, our analysis indicates this is an underestimate because it ignores common and larger shifts that occurred along axes oriented to the north‐west and north‐east. Trailing edges contracted from a broad range of southerly directions. Importantly, these results are derived from systematically collected data so confounding observer‐effort biases can be discounted. Analyses of climate for the same period show that whilst temperature trends should drive species along a north–north‐westerly trajectory, directional responses to precipitation will depend on both the time of year that is important for determining a species' distribution, and the location of the range margin. Directions of species' range centroid shift were not correlated with spatial trends in any single climate variable. We conclude that range shifts of British birds are multidirectional, individualistic and probably determined by species‐specific interactions of multiple climate factors. Climate change is predicted to lead to changes in community composition through variation in the rates that species' ranges shift; our results suggest communities could change further owing to constituent species shifting along different trajectories. We recommend more studies consider directionality in climate and range dynamics to produce more appropriate measures of observed and expected responses to climate change.     

Global variation in thermal tolerances and vulnerability of endotherms to climate change

The relationships among species'' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species'' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals—a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species'' physiology and the geography of climate change will advance assessments of species'' vulnerability to climate change.     

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.     

Juçara palm ecological interactions threatened by climate and land-cover changes

Ongoing climate change has caused well-documented displacements of species' geographic distribution to newly climatically suitable areas. Ecological niche models (ENM) are widely used to project such climate-induced changes but typically ignore species' interspecific interactions that might facilitate or prevent its establishment in new areas. Here, we projected the change in the distribution of Juçara Palm (Euterpe edulis Mart., Arecaceae), a neotropical threatened palm, taking into consideration its ecological interactions. We run ENMs of E. edulis, plus its known seed dispersers (15 bird species) and predators (19 birds and mammals) under current and future climatic conditions. Additionally, for E. edulis, we removed deforested areas from the model. When considering only climate, climate change has a positive impact on E. edulis, with a predicted westward expansion and a modest southward contraction, with a 26% net gain in distribution by 2060. When removing deforested areas, however, climate change harms E. edulis, with a 66% predicted net distribution loss. Within the palm's distribution in this more realistic model, there is also a predicted reduction in the richness of its dispersers and predators. We conclude that the possible benefits of climate change to E. edulis' distribution are overshadowed by widespread habitat loss, and that global change is likely to disrupt some of its ecological interactions. The outcome of the interplay between the negative impact of the loss of dispersers, and the benefit of the loss of predators, is unclear, but the large contraction of E. edulis' range predicted here foresees a dim future for the species. Abstract in Portuguese is available with online material.     

Climatic associations of British species distributions show good transferability in time but low predictive accuracy for range change

Conservation planners often wish to predict how species distributions will change in response to environmental changes. Species distribution models (SDMs) are the primary tool for making such predictions. Many methods are widely used; however, they all make simplifying assumptions, and predictions can therefore be subject to high uncertainty. With global change well underway, field records of observed range shifts are increasingly being used for testing SDM transferability. We used an unprecedented distribution dataset documenting recent range changes of British vascular plants, birds, and butterflies to test whether correlative SDMs based on climate change provide useful approximations of potential distribution shifts. We modelled past species distributions from climate using nine single techniques and a consensus approach, and projected the geographical extent of these models to a more recent time period based on climate change; we then compared model predictions with recent observed distributions in order to estimate the temporal transferability and prediction accuracy of our models. We also evaluated the relative effect of methodological and taxonomic variation on the performance of SDMs. Models showed good transferability in time when assessed using widespread metrics of accuracy. However, models had low accuracy to predict where occupancy status changed between time periods, especially for declining species. Model performance varied greatly among species within major taxa, but there was also considerable variation among modelling frameworks. Past climatic associations of British species distributions retain a high explanatory power when transferred to recent time--due to their accuracy to predict large areas retained by species--but fail to capture relevant predictors of change. We strongly emphasize the need for caution when using SDMs to predict shifts in species distributions: high explanatory power on temporally-independent records--as assessed using widespread metrics--need not indicate a model's ability to predict the future.     

Expanding northward: influence of climate change,forest connectivity,and population processes on a threatened species' range shift

Species' ranges are dynamic, shifting in response to a large number of interrelated ecological and anthropogenic processes. Climate change is thought to be one of the most influential drivers of range shifts, but the effects of other confounded ecological processes are often ignored even though these processes may modify expected range responses to climate change. To determine the relative effects of climate, forest availability, connectivity, and biotic processes such as immigration and establishment, we examine range changes occurring in a species of bird, the Hooded Warbler (Wilsonia citrina). We focus predominantly on the periphery of the species' northern range in Canada but we also examine data from the entire species' range. Nesting records in southern Ontario were obtained from two breeding bird Atlases of Ontario separated by a period of 20 years (1981–1985 and 2001–2005), and the rate of range expansion was estimated by comparing the number of occupied areas in each Atlas. Twelve hypotheses of the relationship between the rate of range expansion and factors known to influence range change were examined using model‐selection techniques and a mixed modeling approach (zero‐inflated Poisson's regression). Cooler temperatures were positively related to a lack of range expansion indicating that climate constrained the species' distribution. Establishment probability (based on the number of occupied, neighboring Atlas squares) and immigration from populations to the south (estimated using independent data from the North American Breeding Bird Survey) were also important predictors of range expansion. These biotic process variables can mask the effects of forest availability and connectivity on range expansion. Expansion due to climate change may be slower in fragmented systems, but the rate of expansion will be influenced largely by biotic processes such as proximity to neighboring populations.     

Range-Wide Latitudinal and Elevational Temperature Gradients for the World's Terrestrial Birds: Implications under Global Climate Change

Species'' geographical distributions are tracking latitudinal and elevational surface temperature gradients under global climate change. To evaluate the opportunities to track these gradients across space, we provide a first baseline assessment of the steepness of these gradients for the world''s terrestrial birds. Within the breeding ranges of 9,014 bird species, we characterized the spatial gradients in temperature along latitude and elevation for all and a subset of bird species, respectively. We summarized these temperature gradients globally for threatened and non-threatened species and determined how their steepness varied based on species'' geography (range size, shape, and orientation) and projected changes in temperature under climate change. Elevational temperature gradients were steepest for species in Africa, western North and South America, and central Asia and shallowest in Australasia, insular IndoMalaya, and the Neotropical lowlands. Latitudinal temperature gradients were steepest for extratropical species, especially in the Northern Hemisphere. Threatened species had shallower elevational gradients whereas latitudinal gradients differed little between threatened and non-threatened species. The strength of elevational gradients was positively correlated with projected changes in temperature. For latitudinal gradients, this relationship only held for extratropical species. The strength of latitudinal gradients was better predicted by species'' geography, but primarily for extratropical species. Our findings suggest threatened species are associated with shallower elevational temperature gradients, whereas steep latitudinal gradients are most prevalent outside the tropics where fewer bird species occur year-round. Future modeling and mitigation efforts would benefit from the development of finer grain distributional data to ascertain how these gradients are structured within species'' ranges, how and why these gradients vary among species, and the capacity of species to utilize these gradients under climate change.     

Species vulnerability to climate change: impacts on spatial conservation priorities and species representation

Climate change may shrink and/or shift plant species ranges thereby increasing their vulnerability and requiring targeted conservation to facilitate adaptation. We quantified the vulnerability to climate change of plant species based on exposure, sensitivity and adaptive capacity and assessed the effects of including these components in complementarity‐based spatial conservation prioritisation. We modelled the vulnerability of 584 native plant species under three climate change scenarios in an 11.9 million hectare fragmented agricultural region in southern Australia. We represented exposure as species' geographical range under each climate change scenario as quantified using species distribution models. We calculated sensitivity as a function of the impact of climate change on species' geographical ranges. Using a dispersal kernel, we quantified adaptive capacity as species' ability to migrate to new geographical ranges under each climate change scenario. Using Zonation, we assessed the impact of individual components of vulnerability (exposure, sensitivity and adaptive capacity) on spatial conservation priorities and levels of species representation in priority areas under each climate change scenario. The full vulnerability framework proved an effective basis for identifying spatial conservation priorities under climate change. Including different dimensions of vulnerability had significant implications for spatial conservation priorities. Incorporating adaptive capacity increased the level of representation of most species. However, prioritising sensitive species reduced the representation of other species. We conclude that whilst taking an integrated approach to mitigating species vulnerability to climate change can ensure sensitive species are well‐represented in a conservation network, this can come at the cost of reduced representation of other species. Conservation planning decisions aimed at reducing species vulnerability to climate change need to be made in full cognisance of the sensitivity of spatial conservation priorities to individual components of vulnerability, and the trade‐offs associated with focussing on sensitive species.     

Traits help explain species' performance away from their climate niche centre

Aim

Climate change impacts on biota are variable across sites, among species and throughout individual species' ranges. Niche theory predicts that population performance should decline as site climate becomes increasingly different from the species' climate niche centre, though studies find significant variation from these predictions. Here, we propose that predictions about climate responses can be improved by incorporating species' trait information.

Location

Europe.

Methods

We used observations of plant species abundance change over time to assess variation in climate difference sensitivity (CDS), defined as how species performance (colonization, extinction and abundance change) relates to the difference of site climate from the mean temperature and precipitation of each species' range. We then investigated if leaf economics, plant size and seed mass traits were associated with the species' CDS.

Results

Species that performed better (e.g. increased in abundance) towards sites progressively cooler than their niche centre were shorter and had more resource-acquisitive leaves (i.e. lower leaf dry matter content or LDMC) relative to species with zero or the opposite pattern of temperature difference sensitivity. This result supports the hypothesis that if sites cooler than niche centres are more stressful for a species, then shorter stature is advantageous compared with taller species. The LDMC result suggests the environment selects for more resource-acquisitive leaf strategies towards relatively cooler climates with shorter growing seasons, counter to expectations that conservative strategies would be favoured in such environments. We found few consistent relationships between precipitation difference sensitivities and traits.

Main Conclusions

The results supported key a priori foundations on how trait-based plant strategies dictate species responses to climate variation away from their niche centre. Furthermore, plant height emerged as the most consistent trait that varied with species CDS, suggesting height will be key for theory development around species response to climate change.     

A consistent occupancy–climate relationship across birds and mammals of the Americas

At broad spatial scales, species richness is strongly related to climate. Yet, few ecological studies attempt to identify regularities in the individual species distributions that make up this pattern. Models used to describe species distributions typically model very complex responses to climate. Here, we test whether the variability in the distributions of birds and mammals of the Americas relates to mean annual temperature and precipitation in a simple, consistent way. Specifically, we test if simple mathematical models can predict, as a first approximation, the geographical variation in individual species’ probability of occupancy for 3277 non‐migratory bird and 1659 mammal species. We find a Gaussian model, where the probability of occupancy of a 10⁴ km² quadrat decreases symmetrically and gradually around a species ‘optimal’ temperature and precipitation, was generally the best model, explaining an average of 35% of the deviance in probability of occupancy. The inclusion of additional terms had very small and idiosyncratic effects across species. The Gaussian occupancy–climate relationship appears general among species and taxa and explains nearly as much deviance as complex models including many more parameters. Therefore, we propose that hypotheses aiming to explain the broad‐scale distribution of species or species richness must also predict generally Gaussian occupancy–climate relationships. Synthesis Science aims to identify regularities in a complex natural world. General patterns should be identified before one searches for potential mechanisms and contingencies. However, species geographic distributions are often modelled as complex (sometimes black box), species‐specific, functions of their environment. We asked whether a simple model could account for as much of the geographic variation in a species' probability of occupancy, and be widely applicable across thousands of species. As a first approximation, we found that a simple Gaussian occupancy‐climate relationship is very common in Nature, whether it be causal or not.     

Phylogenetic fields through time: temporal dynamics of geographical co-occurrence and phylogenetic structure within species ranges

Species co-occur with different sets of other species across their geographical distribution, which can be either closely or distantly related. Such co-occurrence patterns and their phylogenetic structure within individual species ranges represent what we call the species phylogenetic fields (PFs). These PFs allow investigation of the role of historical processes—speciation, extinction and dispersal—in shaping species co-occurrence patterns, in both extinct and extant species. Here, we investigate PFs of large mammalian species during the last 3 Myr, and how these correlate with trends in diversification rates. Using the fossil record, we evaluate species'' distributional and co-occurrence patterns along with their phylogenetic structure. We apply a novel Bayesian framework on fossil occurrences to estimate diversification rates through time. Our findings highlight the effect of evolutionary processes and past climatic changes on species'' distributions and co-occurrences. From the Late Pliocene to the Recent, mammal species seem to have responded in an individualistic manner to climate changes and diversification dynamics, co-occurring with different sets of species from different lineages across their geographical ranges. These findings stress the difficulty of forecasting potential effects of future climate changes on biodiversity.     

Effect of Climate Change on Mediterranean Winter Ranges of Two Migratory Passerines

We studied the effect of climate change on the distribution of two insectivorous passerines (the meadow pipit Anthus pratensis and the chiffchaff Phylloscopus collybita) in wintering grounds of the Western Mediterranean basin. In this region, precipitation and temperature can affect the distribution of these birds through direct (thermoregulation costs) or indirect effects (primary productivity). Thus, it can be postulated that projected climate changes in the region will affect the extent and suitability of their wintering grounds. We studied pipit and chiffchaff abundance in several hundred localities along a belt crossing Spain and Morocco and assessed the effects of climate and other geographical and habitat predictors on bird distribution. Multivariate analyses reported a positive effect of temperature on the present distribution of the two species, with an additional effect of precipitation on the meadow pipit. These climate variables were used with Maxent to model the occurrence probabilities of species using ring recoveries as presence data. Abundance and occupancy of the two species in the study localities adjusted to the distribution models, with more birds in sectors of high climate suitability. After validation, these models were used to forecast the distribution of climate suitability according to climate projections for 2050–2070 (temperature increase and precipitation reduction). Results show an expansion of climatically suitable sectors into the highlands by the effect of warming on the two species, and a retreat of the meadow pipit from southern sectors related to rain reduction. The predicted patterns show a mean increase in climate suitability for the two species due to the warming of the large highland expanses typical of the western Mediterranean.     

Loss of adaptive variation during evolutionary responses to climate change

The changes in species' geographical distribution demanded by climate change are often critically limited by the availability of key interacting species. In such cases, species' persistence will depend on the rapid evolution of biotic interactions. Understanding evolutionary limits to such adaptation is therefore crucial for predicting biological responses to environmental change. The recent poleward range expansion of the UK brown argus butterfly has been associated with a shift in female preference from its main host plant, rockrose (Cistaceae), onto Geraniaceae host plants throughout its new distribution. Using reciprocal transplants onto natural host plants across the UK range, we demonstrate reduced fitness of females from recently colonised Geraniaceae‐dominated habitat when moved to ancestral rockrose habitats. By contrast, individuals from ancestral rockrose habitats show no reduction in fitness on Geraniaceae. Climate‐driven range expansion in this species is therefore associated with the rapid evolution of biotic interactions and a significant loss of adaptive variation.     

A Mechanistic Niche Model for Measuring Species' Distributional Responses to Seasonal Temperature Gradients

Niche theory is central to understanding how species respond geographically to climate change. It defines a species'' realized niche in a biological community, its fundamental niche as determined by physiology, and its potential niche—the fundamental niche in a given environment or geographic space. However, most predictions of the effects of climate change on species'' distributions are limited to correlative models of the realized niche, which assume that species are in distributional equilibrium with respect to the variables or gradients included in the model. Here, I present a mechanistic niche model that measures species'' responses to major seasonal temperature gradients that interact with the physiology of the organism. I then use lethal physiological temperatures to parameterize the model for bird species in North and South America and show that most focal bird species are not in direct physiological equilibrium with the gradients. Results also show that most focal bird species possess broad thermal tolerances encompassing novel climates that could become available with climate change. I conclude with discussion of how mechanistic niche models may be used to (i) gain insights into the processes that cause species to respond to climate change and (ii) build more accurate correlative distribution models in birds and other species.     

Living on the edge: British and Irish woodland birds in a European context

This paper reviews broad geographical patterns in the species composition of breeding woodland bird communities from Ireland to eastern Europe and outlines how processes affecting woodland birds in Britain and Ireland may differ from those operating in mainland Europe. Bird communities in British and Irish woodlands consist of a subset of the species occurring within European forests at similar latitudes. The occurrence of virtually all groups of forest birds is lower in Britain, and strikingly lower in Ireland, than in other temperate areas of mainland Europe. This phenomenon appears to form part of a west–east gradient in species diversity and is probably not just a consequence of insularity. Across this gradient there appears to be broad geographical constancy in the types (taxonomic, ecological and life-history groups) of species present. There is considerable spatial variation in habitat use by forest species within Europe. Some species in Britain probably use habitats in different ways to elsewhere for reasons related to competition, predation and historical adaptation to landscape change. Several species appear to reach the limits of their geographical ranges within Britain (i.e. in the absence of physical barriers). We suggest that range contraction of one of these species, the Common Nightingale Luscinia megarhynchos , in Britain may partially reflect redistribution into the highest quality areas in response to a wider population decline. It is argued that conclusions drawn from studies of forest birds in Britain do not necessarily apply in other regions and vice versa. There is a need for large-scale studies in Europe of the spatial variation in organization of forest bird assemblages, habitat use and the genetic structure of populations.     

Range shifting on a fragmented landscape

Projected responses of species' to climate change have so far included few of the factors that are important determinants of species' distributions within its range. In this paper we utilise a spatially explicit cellular lattice, colonisation–extinction model to investigate the effect of habitat loss, fragmentation and species characteristics on range shifting in response to climate change. Contrary to the predictions of patch occupancy in static climate models we show that fragmentation can have a positive effect on species survival when species have high colonisation rates. For species with low colonisation rates aggregative behaviours prevent success on fragmented landscapes at high levels of habitat loss, and range shifting is more successfully achieved where habitat is correlated. At levels of habitat loss near the extinction threshold, less fragmented landscapes can facilitate range shifting even for the best colonisers. We discuss how imposing a climate window may reduce percolation routes and have implications for the area of usable habitat at any given level of habitat availability. We demonstrate the importance of landscape structure for range shifting dynamics and argue that management of reserve networks needs to consider the requirements of species with different life history characteristics.