Range structure analysis: unveiling the internal structure of species’ ranges

Assessing risks of local extinction and shifts in species ranges are fundamental tasks in ecology and conservation. Most studies have focused either on the border of species’ range or on complex spatiotemporal dynamics of populations within the spatial distribution of species. The internal properties of species ranges, however, have received less attention due to a general lack of simple tools. We propose a novel approach within a metapopulation framework to study species ranges based on simple mathematical rules. We formulate and test a model of population fluctuations through space to identify key factors that regulate population density. We propose that spatial variability in species abundance reflects the interaction between temporal variability in population dynamics and the spatial variability of population parameters. This approach, that we call range structure analysis, integrates temporal and spatial properties to diagnose how each parameter contributes to species occupancy throughout its geographic range.     

Drivers of extinction risk in African mammals: the interplay of distribution state,human pressure,conservation response and species biology

Although conservation intervention has reversed the decline of some species, our success is outweighed by a much larger number of species moving towards extinction. Extinction risk modelling can identify correlates of risk and species not yet recognized to be threatened. Here, we use machine learning models to identify correlates of extinction risk in African terrestrial mammals using a set of variables belonging to four classes: species distribution state, human pressures, conservation response and species biology. We derived information on distribution state and human pressure from satellite-borne imagery. Variables in all four classes were identified as important predictors of extinction risk, and interactions were observed among variables in different classes (e.g. level of protection, human threats, species distribution ranges). Species biology had a key role in mediating the effect of external variables. The model was 90% accurate in classifying extinction risk status of species, but in a few cases the observed and modelled extinction risk mismatched. Species in this condition might suffer from an incorrect classification of extinction risk (hence require reassessment). An increased availability of satellite imagery combined with improved resolution and classification accuracy of the resulting maps will play a progressively greater role in conservation monitoring.     

Plant extinction risk under climate change: are forecast range shifts alone a good indicator of species vulnerability to global warming?

Models that couple habitat suitability with demographic processes offer a potentially improved approach for estimating spatial distributional shifts and extinction risk under climate change. Applying such an approach to five species of Australian plants with contrasting demographic traits, we show that: (i) predicted climate‐driven changes in range area are sensitive to the underlying habitat model, regardless of whether demographic traits and their interaction with habitat patch configuration are modeled explicitly; and (ii) caution should be exercised when using predicted changes in total habitat suitability or geographic extent to infer extinction risk, because the relationship between these metrics is often weak. Measures of extinction risk, which quantify threats to population persistence, are particularly sensitive to life‐history traits, such as recruitment response to fire, which explained approximately 60% of the deviance in expected minimum abundance. Dispersal dynamics and habitat patch structure have the strongest influence on the amount of movement of the trailing and leading edge of the range margin, explaining roughly 40% of modeled structural deviance. These results underscore the need to consider direct measures of extinction risk (population declines and other measures of stochastic viability), as well as measures of change in habitat area, when assessing climate change impacts on biodiversity. Furthermore, direct estimation of extinction risk incorporates important demographic and ecosystem processes, which potentially influence species’ vulnerability to extinction due to climate change.     

Do functional traits offset the effects of fragmentation? The case of large-bodied diurnal lemur species

Primates worldwide are faced with increasing threats making them more vulnerable to extinction. Anthropogenic disturbances, such as habitat degradation and fragmentation, are among the main concerns, and in Madagascar, these issues have become widespread. As this situation continues to worsen, we sought to understand how fragmentation affects primate distribution throughout the island. Further, because species may exhibit different sensitivity to fragmentation, we also aimed to estimate the role of functional traits in mitigating their response. We collated data from 32 large-bodied lemur species ranges, consisting of species from the families Lemuridae (five genera) and Indriidae (two genera). We fitted Generalized Linear Models to determine the role of habitat fragmentation characteristics, for example, forest cover, patch size, edge density, and landscape configuration, as well as the protected area (PA) network, on the species relative probability of presence. We then assessed how the influence of functional traits (dietary guild, home range size) mitigate the response of species to these habitat metrics. Habitat area had a strong positive effect for many species, and there were significantly negative effects of fragmentation on the distribution of many lemur species. In addition, there was a positive influence of PAs on many lemur species’ distribution. Functional trait classifications showed that lemurs of all dietary guilds are negatively affected by fragmentation; however, folivore-frugivores show greater flexibility/variability in terms of habitat area and landscape complexity compared to nearly exclusive folivores and frugivores. Furthermore, species of all home range sizes showed a negative response to fragmentation, while habitat area had an increasingly positive effect as home range increased in size. Overall, the general trends for the majority of lemur species are dire and point to the need for immediate actions on a multitude of fronts, most importantly landscape-level reforestation efforts.     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

Tools for integrating range change,extinction risk and climate change information into conservation management

Ecological niche models (ENMs) are the primary tool used to describe and forecast the potential influence of climate change on biodiversity. However, ENMs do not directly account for important biological and landscape processes likely to affect range dynamics at a variety of spatial scales. Recent advances to link ENMs with population models have focused on the fundamental step of integrating dispersal and metapopulation dynamics into forecasts of species geographic ranges. Here we use a combination of novel analyses and a synthesis of findings from published plant and animal case studies to highlight three seldom recognised, yet important, advantages of linking ENMs with demographic modelling approaches: 1) they provide direct measures of extinction risk in addition to measures of vulnerability based on change in the potential range area or total habitat suitability. 2) They capture life‐history traits that permit population density to vary in different ways in response to key spatial drivers, conditioned by the processes of global change. 3) They can be used to explore and rank the cost effectiveness of regional conservation alternatives and demographically oriented management interventions. Given these advantages, we argue that coupled methods should be used preferentially where data permits and when conservation management decisions require intervention, prioritization, or direct estimates of extinction risk.     

Integrating species traits with extrinsic threats: closing the gap between predicting and preventing species declines

In studies of extinction risk, it is often insufficient to conclude that species with narrow ranges or small clutch sizes require prioritized protection. To improve conservation outcomes, we also need to know which threats interact with these traits to endanger some species but not others. In this study, we integrated the spatial patterns of key threats to Australian amphibians with species' ecological/life-history traits to both predict declining species and identify their likely threats. In addition to confirming the importance of previously identified traits (e.g. narrow range size), we find that extrinsic threats (primarily the disease chytridiomycosis and invasive mosquitofish) are equally important and interact with intrinsic traits (primarily ecological group) to create guild-specific pathways to decline in our model system. Integrating the spatial patterns of extrinsic threats in extinction risk analyses will improve our ability to detect and manage endangered species in the future, particularly where data deficiency is a problem.     

Predicting how populations decline to extinction

Global species extinction typically represents the endpoint in a long sequence of population declines and local extinctions. In comparative studies of extinction risk of contemporary mammalian species, there appear to be some universal traits that may predispose taxa to an elevated risk of extinction. In local population-level studies, there are limited insights into the process of population decline and extinction. Moreover, there is still little appreciation of how local processes scale up to global patterns. Advancing the understanding of factors which predispose populations to rapid declines will benefit proactive conservation and may allow us to target at-risk populations as well as at-risk species. Here, we take mammalian population trend data from the largest repository of population abundance trends, and combine it with the PanTHERIA database on mammal traits to answer the question: what factors can be used to predict decline in mammalian abundance? We find in general that environmental variables are better determinants of cross-species population-level decline than intrinsic biological traits. For effective conservation, we must not only describe which species are at risk and why, but also prescribe ways to counteract this.     

Life-history traits associated with fragmentation vulnerability of lizards in the Thousand Island Lake, China

Following habitat fragmentation, the remnant faunal community will undergo a period of species loss or 'relaxation.' Theory predicts that species with particular life-history traits, such as a small population size, small geographical range, low fecundity and large body size, should be more vulnerable to fragmentation. In this study, we investigated the relationships between the above life-history traits and the fragmentation vulnerability index (the number of islands occupied) of five lizard species inhabiting recently isolated land-bridge islands in the Thousand Island Lake, China. Data on life-history traits were collected from field surveys (population density) and from the literature (body size, clutch size and geographical range size). The species–area relationships for lizards sampled from the mainland versus on the islands differed significantly (i.e. the number of species inhabiting islands was decreased relative to similar-sized areas on the mainland), indicating that species extinction has occurred on all of the study islands following isolation. For the fragmentation vulnerability index, model selection based on Akaike's information criterion identified natural density at mainland sites as the best correlate of vulnerability to fragmentation, supporting the hypothesis that rare species are most vulnerable to local extinction and will be lost first from fragmented landscapes. In contrast, there was little evidence for an effect of lizards' snout–vent length, clutch size or geographical range size on fragmentation vulnerability. Identification of species traits that render some species more vulnerable to fragmentation than others has important implications for conservation and can be used to aid direct management efforts.     

Macroecology and extinction risk correlates of frogs

Aim  Our aim was to test whether extinction risk of frog species could be predicted from their body size, fecundity or geographical range size. Because small geographical range size is a correlate of extinction risk in many taxa, we also tested hypotheses about correlates of range size in frogs.
Location  Global.
Methods  Using a large comparative data set ( n  = 527 species) compiled from the literature, we performed bivariate and multiple regressions through the origin of independent contrasts to test proposed macroecological patterns and correlates of extinction risk in frogs. We also created minimum adequate models to predict snout–vent length, clutch size, geographical range size and IUCN Red List status in frogs. Parallel non-phylogenetic analyses were also conducted. We verified the results of the phylogenetic analyses using gridded data accounting for spatial autocorrelation.
Results  The most threatened frog species tend to have small geographical ranges, although the relationship between range and extinction risk is not linear. In addition, tropical frogs with small clutches have the smallest ranges. Clutch size was strongly positively correlated with geographical range size ( r ² = 0.22) and body size ( r ² = 0.28).
Main conclusions  Our results suggest that body size and fecundity only affect extinction risk indirectly through their effect on geographical range size. Thus, although large frogs with small clutches tend to be endangered, there is no comparative evidence that this relationship is direct. If correct, this inference has consequences for conservation strategy: it would be inefficient to allocate conservation resources on the basis of low fecundity or large body size; instead it would be better to protect areas that contain many frog species with small geographical ranges.     

Doomed to die? Predicting extinction risk in the true hawks Accipitridae

One of the most important tasks in conservation biology is identifying species at risk from extinction and establishing the most likely factors influencing this risk. Here, we consider an ecologically well-defined, monophyletic group of organisms, the true hawks of the family Accipitridae, which are not only among the most studied, but also contain some of the rarest bird species in the world. We investigate which intrinsic and extrinsic factors, covering morphology, life history and ecology, covary with International Union for the Conservation of Nature and Natural Resources threat status, as well as global population size and geographic range size. By decomposing threat status into population size and range size, we test whether any factors are generally important: we found that species with less habitat specialization, a larger clutch size and more plumage polymorphism were associated with lower extinction risk and larger population and range sizes. Species with special habitat requirements might be less capable of dealing with habitat transformation and fragmentation, while species with small clutch sizes might not be able to reverse population declines. Plumage polymorphism might indicate the size of the species' gene pool and could be a good marker of extinction risk. The analyses also emphasized that no single factor is likely to be sufficient when predicting the threat of extinction.     

Life history traits and abundance can predict local colonisation and extinction rates of freshwater mussels

1. A critical need in conservation biology is to determine which species are most vulnerable to extinction. Freshwater mussels (Bivalvia: Unionacea) are one of the most imperilled faunal groups globally. Freshwater mussel larvae are ectoparasites on fish and depend on the movement of their hosts to maintain connectivity among local populations in a metapopulation. 2. I calculated local colonisation and extinction rates for 16 mussel species from 14 local populations in the Red River drainage of Oklahoma and Texas, U.S. I used general linear models and AIC comparisons to determine which mussel life history traits best predicted local colonisation and extinction rates. 3. Traits related to larval dispersal ability (host infection mode, whether a mussel species was a host generalist or specialist) were the best predictors of local colonisation. 4. Traits related to local population size (regional abundance, time spent brooding) were the best predictors of local extinction. The group of fish species used as hosts by mussels also predicted local extinction and was probably related to habitat fragmentation and host dispersal abilities. 5. Overall, local extinction rates exceeded local colonisation rates, indicating that local populations are becoming increasingly isolated and suffering an ‘extinction debt’. This study demonstrates that analysis of species traits can be used to predict local colonisation and extinction patterns and provide insight into the long‐term persistence of populations.     

The conservation status of African vertebrates is unrelated to environmental and spatial patterns in their geographic ranges

Statistical predictions of the impact of climate change on biodiversity assume that the environmental and spatial characteristics of contemporary species’ distributions reflect the conditions needed for their continued and prolonged existence. Here we explore this assumption by testing whether a species’ threatened status is associated with the amount of variation in its distribution range attributable to environmental and spatial patterns. Using a variation partitioning approach, we decomposed variation in the distribution ranges of 4423 vertebrate species in sub-Saharan Africa into components attributable exclusively to environmental variables (E|S), exclusively to spatial variables (S|E) or to the collinearity between environmental and spatial variables (E∩S). We found that species’ threatened status was unrelated to E|S, S|E or E∩S variation components, but that unexplained variation was higher for species threatened with extinction. This suggests that spatio-environmental patterns in species’ ranges likely underestimate the overall extinction threat caused by climate change. We also found clear geographic patterns in the strength of E|S, S|E or E∩S that differed amongst biogeographical regions, but no component was over- or underrepresented in the present-day protected area network. While there may be benefits to tailoring protected area expansion to differences between biogeographical regions, this should aim to incorporate species-specific information wherever possible.     

Reconstructing range dynamics and range fragmentation of European bison for the last 8000 years

Aim Understanding what constituted species’ ranges prior to large‐scale human influence, and how past climate and land use change have affected range dynamics, provides conservation planners with important insights into how species may respond to future environmental change. Our aim here was to reconstruct the Holocene range of European bison (Bison bonasus) by combining a time‐calibrated species distribution models (SDM) with a dynamic vegetation model. Location Europe. Method We used European bison occurrences from the Holocene in a maximum entropy model to assess bison range dynamics during the last 8000 years. As predictors, we used bioclimatic variables and vegetation reconstructions from the generalized dynamic vegetation model LPJ‐GUESS. We compared our range maps with maps of farmland and human population expansion to identify the main species range constraints. Results The Holocene distribution of European bison was mainly determined by vegetation patterns, with bison thriving in both broadleaved and coniferous forests, as well as by mean winter temperature. The heartland of European bison was in Central and Eastern Europe, whereas suitable habitat in Western Europe was scarce. While environmentally suitable regions were overall stable, the expansion of settlements and farming severely diminished available habitat. Main conclusions European bison habitat preferences may be wider than previously assumed, and our results suggest that the species had a more eastern and northern distribution than previously reported. Vegetation and climate transformation during the Holocene did not affect the bison’s range substantially. Conversely, human population growth and the spread of farming resulted in drastic bison habitat loss and fragmentation, likely reaching a tipping point during the last 1000 years. Combining SDM and dynamic vegetation models can improve range reconstructions and projections, and thus help to identify resilient conservation strategies for endangered species.     

Spatial heterogeneity and habitat configuration overcome habitat composition influences on alpha and beta mammal diversity

The effects of habitat fragmentation on different taxa and ecosystems are subject to intense debate, and disentangling them is of utmost importance to support conservation and management strategies. We evaluated the importance of landscape composition and configuration, and spatial heterogeneity to explain α- and β-diversity of mammals across a gradient of percent woody cover and land use diversity. We expected species richness to be positively related to all predictive variables, with the strongest relationship with landscape composition and configuration, and spatial heterogeneity respectively. We also expected landscape to influence β-diversity in the same order of importance expected for species richness, with a stronger influence on nestedness due to deterministic loss of species more sensitive to habitat disturbance. We analyzed landscape structure using: (a) landscape metrics based on thematic maps and (b) image texture of a vegetation index. We compared a set of univariate explanatory models of species richness using AIC, and evaluated how dissimilarities in landscape composition and configuration and spatial heterogeneity affect β-diversity components using a Multiple Regression on distance Matrix. Contrary with our expectations, landscape configuration was the main driver of species richness, followed by spatial heterogeneity and last by landscape composition. Nestedness was explained, in order of importance, by spatial heterogeneity, landscape configuration, and landscape composition. Although conservation policies tend to focus mainly on habitat amount, we advocate that landscape management must include strategies to preserve and improve habitat quality and complexity in natural patches and the surrounding matrix, enabling landscapes to harbor high species diversity.     

Assessing Extinction Risk in Small Metapopulations of Golden‐headed Lion Tamarins (Leontopithecus chrysomelas) in Bahia State,Brazil

Golden‐headed lion tamarins (GHLTs; Leontopithecus chrysomelas) are endangered primates endemic to the Brazilian Atlantic Forest, where loss of forest and its connectivity threaten species survival. Understanding the role of habitat availability and configuration on population declines is critical for guiding proactive conservation for this, and other, endangered species. We conducted population viability analysis to assess vulnerability of ten GHLT metapopulations to habitat loss and small population size. Seven metapopulations had a low risk of extirpation (or local extinction) over the next 100 years assuming no further forest loss, and even small populations could persist with immediate protection. Three metapopulations had a moderate/high risk of extirpation, suggesting extinction debt may be evident in parts of the species’ range. When deforestation was assumed to continue at current rates, extirpation risk significantly increased while abundance and genetic diversity decreased for all metapopulations. Extirpation risk was significantly negatively correlated with the size of the largest patch available to metapopulations, underscoring the importance of large habitat patches for species persistence. Finally, we conducted sensitivity analysis using logistic regression, and our results showed that local extinction risk was sensitive to percentage of females breeding, adult female mortality, and dispersal rate and survival; conservation or research programs that target these aspects of the species’ biology/ecology could have a disproportionately important impact on species survival. We stress that efforts to protect populations and tracts of habitat of sufficient size throughout the species’ distribution will be important in the near‐term to protect the species from continuing decline and extinction.     

Global patterns of fragmentation and connectivity of mammalian carnivore habitat

Although mammalian carnivores are vulnerable to habitat fragmentation and require landscape connectivity, their global patterns of fragmentation and connectivity have not been examined. We use recently developed high-resolution habitat suitability models to conduct comparative analyses and to identify global hotspots of fragmentation and connectivity for the world's terrestrial carnivores. Species with less fragmentation (i.e. more interior high-quality habitat) had larger geographical ranges, a greater proportion of habitat within their range, greater habitat connectivity and a lower risk of extinction. Species with higher connectivity (i.e. less habitat isolation) also had a greater proportion of high-quality habitat, but had smaller, not larger, ranges, probably reflecting shorter distances between habitat patches for species with restricted distributions; such species were also more threatened, as would be expected given the negative relationship between range size and extinction risk. Fragmentation and connectivity did not differ among Carnivora families, and body mass was associated with connectivity but not fragmentation. On average, only 54.3 per cent of a species' geographical range comprised high-quality habitat, and more troubling, only 5.2 per cent of the range comprised such habitat within protected areas. Identification of global hotspots of fragmentation and connectivity will help guide strategic priorities for carnivore conservation.     

Bird conservation in the Carpathian Ecoregion in light of long-term land use trends and conservation responsibility

Land use change alters species' abundance and distributions by affecting habitat availability and quality. The decline of bird populations worldwide is of major concern, and habitat protection and restoration are primary conservation actions. However, conservation decisions largely consider only short-term habitat changes and species’ population dynamics in a given area. Disregarding long term modifications in species’ available habitat, and the role of a given population for a species’ global population may lead to misdirected conservation action. Our goal here was to combine the assessment of conservation responsibility, with that of century-long available habitat dynamics, in order to inform better conservation practice. We compiled available habitat data for 170 bird species in the Carpathian Region from 1860 to 2010 from historic maps and satellite data. We analyzed these species’ range distributions, IUCN extinction risk and population trends, and we identified 29 species of high conservation responsibility, and all of them were forest or and grassland specialists. Furthermore, we found major land use trends including cropland abandonment and increase in forests and grasslands that resulted in increases in potential habitat for the species for which the Carpathians have high conservation responsibility. The loss of row-crop agriculture, on the other hand, reduced habitat for species for which the Carpathians do not have high responsibility, and thus subsidizing agriculture may not be warranted from a conservation perspective. More broadly, many regions worldwide are undergoing rapid land use changes, and we suggest that these should be analyzed relative to a given regions’ conservation responsibility to see if there are opportunities for conservation, i.e., cases similar to the Carpathians, where conservation efforts ‘only’ have to foster current land use trends, and make them permanent, rather than to try to revert the loss of habitat.     

Effects of species' ecology on the accuracy of distribution models

In the face of accelerating biodiversity loss and limited data, species distribution models – which statistically capture and predict species’ occurrences based on environmental correlates – are increasingly used to inform conservation strategies. Additionally, distribution models and their fit provide insights on the broad‐scale environmental niche of species. To investigate whether the performance of such models varies with species’ ecological characteristics, we examined distribution models for 1329 bird species in southern and eastern Africa. The models were constructed at two spatial resolutions with both logistic and autologistic regression. Satellite‐derived environmental indices served as predictors, and model accuracy was assessed with three metrics: sensitivity, specificity and the area under the curve (AUC) of receiver operating characteristics plots. We then determined the relationship between each measure of accuracy and ten ecological species characteristics using generalised linear models. Among the ecological traits tested, species’ range size, migratory status, affinity for wetlands and endemism proved most influential on the performance of distribution models. The number of habitat types frequented (habitat tolerance), trophic rank, body mass, preferred habitat structure and association with sub‐resolution habitats also showed some effect. In contrast, conservation status made no significant impact. These findings did not differ from one spatial resolution to the next. Our analyses thus provide conservation scientists and resource managers with a rule of thumb that helps distinguish, on the basis of ecological traits, between species whose occurrence is reliably or less reliably predicted by distribution models. Reasonably accurate distribution models should, however, be attainable for most species, because the influence ecological traits bore on model performance was only limited. These results suggest that none of the ecological traits tested provides an obvious correlate for environmental niche breadth or intra‐specific niche differentiation.     

Habitat specialization predicts genetic response to fragmentation in tropical birds

Habitat fragmentation is one of the most severe threats to biodiversity as it may lead to changes in population genetic structure, with ultimate modifications of species evolutionary potential and local extinctions. Nonetheless, fragmentation does not equally affect all species and identifying which ecological traits are related to species sensitivity to habitat fragmentation could help prioritization of conservation efforts. Despite the theoretical link between species ecology and extinction proneness, comparative studies explicitly testing the hypothesis that particular ecological traits underlies species‐specific population structure are rare. Here, we used a comparative approach on eight bird species, co‐occurring across the same fragmented landscape. For each species, we quantified relative levels of forest specialization and genetic differentiation among populations. To test the link between forest specialization and susceptibility to forest fragmentation, we assessed species responses to fragmentation by comparing levels of genetic differentiation between continuous and fragmented forest landscapes. Our results revealed a significant and substantial population structure at a very small spatial scale for mobile organisms such as birds. More importantly, we found that specialist species are more affected by forest fragmentation than generalist ones. Finally, our results suggest that even a simple habitat specialization index can be a satisfying predictor of genetic and demographic consequences of habitat fragmentation, providing a reliable practical and quantitative tool for conservation biology.