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author:("jet, Walter")
1.  Unravelling the structure of species extinction risk for predictive conservation science 
Extinction risk varies across species and space owing to the combined and interactive effects of ecology/life history and geography. For predictive conservation science to be effective, large datasets and integrative models that quantify the relative importance of potential factors and separate rapidly changing from relatively static threat drivers are urgently required. Here, we integrate and map in space the relative and joint effects of key correlates of The International Union for Conservation of Nature-assessed extinction risk for 8700 living birds. Extinction risk varies significantly with species' broad-scale environmental niche, geographical range size, and life-history and ecological traits such as body size, developmental mode, primary diet and foraging height. Even at this broad scale, simple quantifications of past human encroachment across species' ranges emerge as key in predicting extinction risk, supporting the use of land-cover change projections for estimating future threat in an integrative setting. A final joint model explains much of the interspecific variation in extinction risk and provides a remarkably strong prediction of its observed global geography. Our approach unravels the species-level structure underlying geographical gradients in extinction risk and offers a means of disentangling static from changing components of current and future threat. This reconciliation of intrinsic and extrinsic, and of past and future extinction risk factors may offer a critical step towards a more continuous, forward-looking assessment of species' threat status based on geographically explicit environmental change projections, potentially advancing global predictive conservation science.
doi:10.1098/rspb.2010.1877
PMCID: PMC3061137  PMID: 20943690
birds; IUCN Red List; projected threat; future global change; structural equation models
2.  Global Gradients in Vertebrate Diversity Predicted by Historical Area-Productivity Dynamics and Contemporary Environment 
PLoS Biology  2012;10(3):e1001292.
A novel hierarchical framework integrates the effects of time, area, productivity, and temperature at their respective relevant scales and successfully predicts the latitudinal gradient in global vertebrate diversity.
Broad-scale geographic gradients in species richness have now been extensively documented, but their historical underpinning is still not well understood. While the importance of productivity, temperature, and a scale dependence of the determinants of diversity is broadly acknowledged, we argue here that limitation to a single analysis scale and data pseudo-replication have impeded an integrated evolutionary and ecological understanding of diversity gradients. We develop and apply a hierarchical analysis framework for global diversity gradients that incorporates an explicit accounting of past environmental variation and provides an appropriate measurement of richness. Due to environmental niche conservatism, organisms generally reside in climatically defined bioregions, or “evolutionary arenas,” characterized by in situ speciation and extinction. These bioregions differ in age and their total productivity and have varied over time in area and energy available for diversification. We show that, consistently across the four major terrestrial vertebrate groups, current-day species richness of the world's main 32 bioregions is best explained by a model that integrates area and productivity over geological time together with temperature. Adding finer scale variation in energy availability as an ecological predictor of within-bioregional patterns of richness explains much of the remaining global variation in richness at the 110 km grain. These results highlight the separate evolutionary and ecological effects of energy availability and provide a first conceptual and empirical integration of the key drivers of broad-scale richness gradients. Avoiding the pseudo-replication that hampers the evolutionary interpretation of non-hierarchical macroecological analyses, our findings integrate evolutionary and ecological mechanisms at their most relevant scales and offer a new synthesis regarding global diversity gradients.
Author Summary
Understanding what determines the distribution of biodiversity across the planet remains one of the critical challenges in biology and has gained particular urgency in the face of environmental change and accelerating species extinctions. Our study develops a novel analytical framework to jointly evaluate historical and contemporary environmental predictors of the latitudinal gradient in the diversity of terrestrial vertebrates. The number of vertebrate species is greater in warm, productive biomes, such as tropical forests, that have both a large size and a long evolutionary history. Using just a few key predictor variables—time, area, productivity, and temperature—we are now able to explain more than 80% of the variability in biodiversity among bioregions. By integrating each of these factors at both the regional and local scale in a hierarchical model, we are able to provide a consensus explanation for broad-scale diversity gradients that encompasses both ecological and evolutionary mechanisms.
doi:10.1371/journal.pbio.1001292
PMCID: PMC3313913  PMID: 22479151
3.  Projected range contractions of montane biodiversity under global warming 
Mountains, especially in the tropics, harbour a unique and large portion of the world's biodiversity. Their geographical isolation, limited range size and unique environmental adaptations make montane species potentially the most threatened under impeding climate change. Here, we provide a global baseline assessment of geographical range contractions and extinction risk of high-elevation specialists in a future warmer world. We consider three dispersal scenarios for simulated species and for the world's 1009 montane bird species. Under constrained vertical dispersal (VD), species with narrow vertical distributions are strongly impacted; at least a third of montane bird diversity is severely threatened. In a scenario of unconstrained VD, the location and structure of mountain systems emerge as a strong driver of extinction risk. Even unconstrained lateral movements offer little improvement to the fate of montane species in the Afrotropics, Australasia and Nearctic. Our results demonstrate the particular roles that the geography of species richness, the spatial structure of lateral and particularly vertical range extents and the specific geography of mountain systems have in determining the vulnerability of montane biodiversity to climate change. Our findings confirm the outstanding levels of biotic perturbation and extinction risk that mountain systems are likely to experience under global warming and highlight the need for additional knowledge on species' vertical distributions, dispersal and adaptive capacities.
doi:10.1098/rspb.2010.0612
PMCID: PMC2982223  PMID: 20534610
climate change; dispersal; extinction risk; geographical range; global warming; mountain biodiversity
4.  Phylogenetic conservatism of environmental niches in mammals 
Phylogenetic niche conservatism is the pattern where close relatives occupy similar niches, whereas distant relatives are more dissimilar. We suggest that niche conservatism will vary across clades in relation to their characteristics. Specifically, we investigate how conservatism of environmental niches varies among mammals according to their latitude, range size, body size and specialization. We use the Brownian rate parameter, σ2, to measure the rate of evolution in key variables related to the ecological niche and define the more conserved group as the one with the slower rate of evolution. We find that tropical, small-ranged and specialized mammals have more conserved thermal niches than temperate, large-ranged or generalized mammals. Partitioning niche conservatism into its spatial and phylogenetic components, we find that spatial effects on niche variables are generally greater than phylogenetic effects. This suggests that recent evolution and dispersal have more influence on species' niches than more distant evolutionary events. These results have implications for our understanding of the role of niche conservatism in species richness patterns and for gauging the potential for species to adapt to global change.
doi:10.1098/rspb.2010.2207
PMCID: PMC3119006  PMID: 21208967
environmental niche; temperature; precipitation; Brownian rate parameter; space; phylogeny
5.  Disparities between observed and predicted impacts of climate change on winter bird assemblages 
Understanding how climate change affects the structure and function of communities is critical for gauging its full impact on biodiversity. To date, community-level changes have been poorly documented, owing, in part, to the paucity of long-term datasets. To circumvent this, the use of ‘space-for-time’ substitution—the forecasting of temporal trends from spatial climatic gradients—has increasingly been adopted, often with little empirical support. Here we examine changes from 1975 to 2001 in three community attributes (species richness, body mass and occupancy) for 404 assemblages of terrestrial winter avifauna in North America containing a total of 227 species. We examine the accuracy of space-for-time substitution and assess causal associations between community attributes and observed changes in annual temperature using a longitudinal study design. Annual temperature and all three community attributes increased over time. The trends for the three community attributes differed significantly from the spatially derived predictions, although richness showed broad congruence. Correlations with trends in temperature were found with richness and body mass. In the face of rapid climate change, applying space-for-time substitution as a predictive tool could be problematic with communities developing patterns not reflected by spatial ecological associations.
doi:10.1098/rspb.2009.0162
PMCID: PMC2817117  PMID: 19520804
avian communities; body-size distributions; climate change; longitudinal analysis; space-for-time substitution; species richness
6.  Projected impacts of climate change on regional capacities for global plant species richness 
Climate change represents a major challenge to the maintenance of global biodiversity. To date, the direction and magnitude of net changes in the global distribution of plant diversity remain elusive. We use the empirical multi-variate relationships between contemporary water-energy dynamics and other non-climatic predictor variables to model the regional capacity for plant species richness (CSR) and its projected future changes. We find that across all analysed Intergovernmental Panel on Climate Change emission scenarios, relative changes in CSR increase with increased projected temperature rise. Between now and 2100, global average CSR is projected to remain similar to today (+0.3%) under the optimistic B1/+1.8°C scenario, but to decrease significantly (−9.4%) under the ‘business as usual’ A1FI/+4.0°C scenario. Across all modelled scenarios, the magnitude and direction of CSR change are geographically highly non-uniform. While in most temperate and arctic regions, a CSR increase is expected, the projections indicate a strong decline in most tropical and subtropical regions. Countries least responsible for past and present greenhouse gas emissions are likely to incur disproportionately large future losses in CSR, whereas industrialized countries have projected moderate increases. Independent of direction, we infer that all changes in regional CSR will probably induce on-site species turnover and thereby be a threat to native floras.
doi:10.1098/rspb.2010.0120
PMCID: PMC2894901  PMID: 20335215
biodiversity patterns; global warming; water-energy dynamics; water-energy-richness hypothesis
7.  Global associations between terrestrial producer and vertebrate consumer diversity 
In both ecology and conservation, often a strong positive association is assumed between the diversity of plants as primary producers and that of animals, specifically primary consumers. Such a relationship has been observed at small spatial scales, and a begetting of diversity by diversity is expected under various scenarios of co-evolution and co-adaptation. But positive producer–consumer richness relationships may also arise from similar associations with past opportunities for diversification or contemporary environmental conditions, or from emerging properties of plant diversity such as vegetation complexity or productivity. Here we assess whether the producer–consumer richness relationship generalizes from plot to regional scale and provide a first global test of its strength for vascular plants and endothermic vertebrates. We find strong positive richness associations, but only limited congruence of the most diverse regions. The richness of both primary and higher-level consumers increases with plant richness at similar strength and rate. Environmental conditions emerge as much stronger predictors of consumer richness, and after accounting for environmental differences little variation is explained by plant diversity. We conclude that biotic interactions and strong local associations between plants and consumers only relatively weakly scale up to broad geographical scales and to functionally diverse taxa, for which environmental constraints on richness dominate.
doi:10.1098/rspb.2008.1005
PMCID: PMC2674345  PMID: 18832059
biodiversity; plants; vertebrates
8.  Space versus phylogeny: disentangling phylogenetic and spatial signals in comparative data 
Variation in traits across species or populations is the outcome of both environmental and historical factors. Trait variation is therefore a function of both the phylogenetic and spatial context of species. Here we introduce a method that, within a single framework, estimates the relative roles of spatial and phylogenetic variations in comparative data. The approach requires traits measured across phylogenetic units, e.g. species, the spatial occurrences of those units and a phylogeny connecting them. The method modifies the expected variance of phylogenetically independent contrasts to include both spatial and phylogenetic effects. We illustrate this approach by analysing cross-species variation in body mass, geographical range size and species-typical environmental temperature in three orders of mammals (carnivores, artiodactyls and primates). These species attributes contain highly disparate levels of phylogenetic and spatial signals, with the strongest phylogenetic autocorrelation in body size and spatial dependence in environmental temperatures and geographical range size showing mixed effects. The proposed method successfully captures these differences and in its simplest form estimates a single parameter that quantifies the relative effects of space and phylogeny. We discuss how the method may be extended to explore a range of models of evolution and spatial dependence.
doi:10.1098/rspb.2008.0905
PMCID: PMC2614254  PMID: 18796398
comparative method; spatial analysis; mammals
9.  Future battlegrounds for conservation under global change 
Global biodiversity is under significant threat from the combined effects of human-induced climate and land-use change. Covering 12% of the Earth's terrestrial surface, protected areas are crucial for conserving biodiversity and supporting ecological processes beneficial to human well-being, but their selection and design are usually uninformed about future global change. Here, we quantify the exposure of the global reserve network to projected climate and land-use change according to the Millennium Ecosystem Assessment and set these threats in relation to the conservation value and capacity of biogeographic and geopolitical regions. We find that geographical patterns of past human impact on the land cover only poorly predict those of forecasted change, thus revealing the inadequacy of existing global conservation prioritization templates. Projected conservation risk, measured as regional levels of land-cover change in relation to area protected, is the greatest at high latitudes (due to climate change) and tropics/subtropics (due to land-use change). Only some high-latitude nations prone to high conservation risk are also of high conservation value, but their high relative wealth may facilitate additional conservation efforts. In contrast, most low-latitude nations tend to be of high conservation value, but they often have limited capacity for conservation which may exacerbate the global biodiversity extinction crisis. While our approach will clearly benefit from improved land-cover projections and a thorough understanding of how species range will shift under climate change, our results provide a first global quantitative demonstration of the urgent need to consider future environmental change in reserve-based conservation planning. They further highlight the pressing need for new reserves in target regions and support a much extended ‘north–south’ transfer of conservation resources that maximizes biodiversity conservation while mitigating global climate change.
doi:10.1098/rspb.2007.1732
PMCID: PMC2602673  PMID: 18302999
conservation planning; conservation priorities; protected areas; future land-use and climate change; biodiversity
10.  The Worldwide Variation in Avian Clutch Size across Species and Space 
PLoS Biology  2008;6(12):e303.
Traits such as clutch size vary markedly across species and environmental gradients but have usually been investigated from either a comparative or a geographic perspective, respectively. We analyzed the global variation in clutch size across 5,290 bird species, excluding brood parasites and pelagic species. We integrated intrinsic (morphological, behavioural), extrinsic (environmental), and phylogenetic effects in a combined model that predicts up to 68% of the interspecific variation in clutch size. We then applied the same species-level model to predict mean clutch size across 2,521 assemblages worldwide and found that it explains the observed eco-geographic pattern very well. Clutches are consistently largest in cavity nesters and in species occupying seasonal environments, highlighting the importance of offspring and adult mortality that is jointly expressed in intrinsic and extrinsic correlates. The findings offer a conceptual bridge between macroecology and comparative biology and provide a global and integrative understanding of the eco-geographic and cross-species variation in a core life-history trait.
Author Summary
Why do some bird species lay only one egg in their nest, and others ten? The clutch size of birds is one of the best-studied life-history traits of animals. Nevertheless, research has so far focused either on a comparative approach, relating clutch size to other biological traits of the species, such as body weight; or on a macroecological approach, testing how environmental factors, such as seasonality, influence clutch size. We used the most comprehensive dataset on clutch size ever compiled, including 5,290 species, and combined it with data on the biology and the environment of these species. This approach enabled us to merge comparative and macroecological methods and to test biological and environmental factors together in one analysis. With this approach, we are able to explain a major proportion of the global variation in clutch size and also to predict with high confidence the average clutch size of a bird assemblage on earth. For example, cavity nesters, such as woodpeckers, have larger clutches than open-nesting species; and species in seasonal environments, especially at northern latitudes, have larger clutches than tropical birds. The findings offer a bridge between macroecology and comparative biology, and provide a global and integrative understanding of a core life-history trait.
Why do some bird species lay only one egg in their nest, and others ten? An analysis combining comparative and macroecological approaches across more than half of all bird species explains the global variation in this trait with high confidence.
doi:10.1371/journal.pbio.0060303
PMCID: PMC2596859  PMID: 19071959
11.  Environment, Migratory Tendency, Phylogeny and Basal Metabolic Rate in Birds 
PLoS ONE  2008;3(9):e3261.
Basal metabolic rate (BMR) represents the minimum maintenance energy requirement of an endotherm and has far-reaching consequences for interactions between animals and their environments. Avian BMR exhibits considerable variation that is independent of body mass. Some long-distance migrants have been found to exhibit particularly high BMR, traditionally interpreted as being related to the energetic demands of long-distance migration. Here we use a global dataset to evaluate differences in BMR between migrants and non-migrants, and to examine the effects of environmental variables. The BMR of migrant species is significantly higher than that of non-migrants. Intriguingly, while the elevated BMR of migrants on their breeding grounds may reflect the metabolic machinery required for long-distance movements, an alternative (and statistically stronger) explanation is their occupation of predominantly cold high-latitude breeding areas. Among several environmental predictors, average annual temperature has the strongest effect on BMR, with a 50% reduction associated with a 20°C gradient. The negative effects of temperature variables on BMR hold separately for migrants and non-migrants and are not due their different climatic associations. BMR in migrants shows a much lower degree of phylogenetic inertia. Our findings indicate that migratory tendency need not necessarily be invoked to explain the higher BMR of migrants. A weaker phylogenetic signal observed in migrants supports the notion of strong phenotypic flexibility in this group which facilitates migration-related BMR adjustments that occur above and beyond environmental conditions. In contrast to the findings of previous analyses of mammalian BMR, primary productivity, aridity or precipitation variability do not appear to be important environmental correlates of avian BMR. The strong effects of temperature-related variables and varying phylogenetic effects reiterate the importance of addressing both broad-scale and individual-scale variation for understanding the determinants of BMR.
doi:10.1371/journal.pone.0003261
PMCID: PMC2533122  PMID: 18810267
12.  Environmental and historical constraints on global patterns of amphibian richness 
Our knowledge of the broad-scale ecology of vertebrate ectotherms remains very limited. Despite ongoing declines and sensitivity to environmental change, amphibian distributions are particularly poorly understood. We present a global analysis of contemporary environmental and historical constraints on amphibian richness, the first for an ectotherm clade at this scale. Amphibians are presumed to experience environmental constraints distinct from those of better studied endothermic taxa due to their stringent water requirements and the temperature dependence of their energetic costs and performance. Single environmental predictors set upper bounds on, but do not exclusively determine, amphibian richness. Accounting for differing regional histories of speciation and extinction helps resolve triangular or scattered relationships between core environmental predictors and amphibian richness, as the relationships' intercepts or slopes can vary regionally. While the magnitude of richness is strongly determined by regional history, within-region patterns are consistently jointly driven by water and temperature. This confirms that ecophysiological constraints extend to the broad scale. This coupling suggests that shifts in climatic regimes will probably have dramatic consequences for amphibians. Our results illustrate how the environmental and historical explanations of species richness gradients can be reconciled and how the perspectives are complements for understanding broad-scale patterns of diversity.
doi:10.1098/rspb.2006.0436
PMCID: PMC2189569  PMID: 17327208
amphibians; biogeography; climate change; diversity gradient; historical contingency; water–temperature hypothesis
13.  Projected Impacts of Climate and Land-Use Change on the Global Diversity of Birds 
PLoS Biology  2007;5(6):e157.
Over the past few decades, land-use and climate change have led to substantial range contractions and species extinctions. Even more dramatic changes to global land cover are projected for this century. We used the Millennium Ecosystem Assessment scenarios to evaluate the exposure of all 8,750 land bird species to projected land-cover changes due to climate and land-use change. For this first baseline assessment, we assumed stationary geographic ranges that may overestimate actual losses in geographic range. Even under environmentally benign scenarios, at least 400 species are projected to suffer >50% range reductions by the year 2050 (over 900 by the year 2100). Although expected climate change effects at high latitudes are significant, species most at risk are predominantly narrow-ranged and endemic to the tropics, where projected range contractions are driven by anthropogenic land conversions. Most of these species are currently not recognized as imperiled. The causes, magnitude and geographic patterns of potential range loss vary across socioeconomic scenarios, but all scenarios (even the most environmentally benign ones) result in large declines of many species. Whereas climate change will severely affect biodiversity, in the near future, land-use change in tropical countries may lead to yet greater species loss. A vastly expanded reserve network in the tropics, coupled with more ambitious goals to reduce climate change, will be needed to minimize global extinctions.
Author Summary
Land conversion and climate change have already had significant impacts on biodiversity and associated ecosystem services. Using future land-cover projections from the recently completed Millennium Ecosystem Assessment, we found that 950–1,800 of the world's 8,750 species of land birds could be imperiled by climate change and land conversion by the year 2100. These projections are based on the assumption that birds will not dramatically shift their ranges in response to a changing climate, a process that would lessen the range contractions we predict. While climate change will be the principal driver of range contractions at higher latitudes, our projections reveal that land conversion (e.g., deforestation, conversion of grasslands to croplands, etc.) will have a much larger effect on species that inhabit the tropics. This is because birds in the tropics are especially diverse and tend to have small ranges, making them particularly vulnerable to extinction; in contrast, birds at higher latitudes are less diverse and tend to have large ranges. A vastly expanded reserve network in the tropics, coupled with more ambitious goals to reduce greenhouse gas emissions and monitor biodiversity impacts, will be needed to minimize global extinctions.
Evaluating changes in range size of land bird species using Millenium Ecosystem Assessment scenarios reveals that land conversion, as well as climate change, will lead to the decline of many species, particularly those in the tropics.
doi:10.1371/journal.pbio.0050157
PMCID: PMC1885834  PMID: 17550306
14.  Phenotypic plasticity in the scaling of avian basal metabolic rate 
Many birds exhibit short-term, reversible adjustments in basal metabolic rate (BMR), but the overall contribution of phenotypic plasticity to avian metabolic diversity remains unclear. The available BMR data include estimates from birds living in natural environments and captive-raised birds in more homogenous, artificial environments. All previous analyses of interspecific variation in BMR have pooled these data. We hypothesized that phenotypic plasticity is an important contributor to interspecific variation in avian BMR, and that captive-raised populations exhibit general differences in BMR compared to wild-caught populations. We tested this hypothesis by fitting general linear models to BMR data for 231 bird species, using the generalized least-squares approach to correct for phylogenetic relatedness when necessary. The scaling exponent relating BMR to body mass in captive-raised birds (0.670) was significantly shallower than in wild-caught birds (0.744). The differences in metabolic scaling between captive-raised and wild-caught birds persisted when migratory tendency and habitat aridity were controlled for. Our results reveal that phenotypic plasticity is a major contributor to avian interspecific metabolic variation. The finding that metabolic scaling in birds is partly determined by environmental factors provides further support for models that predict variation in scaling exponents, such as the allometric cascade model.
doi:10.1098/rspb.2005.3415
PMCID: PMC1560247  PMID: 16627278
metabolic scaling; captivity; allometry; basal metabolic rate

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