Showing papers in "Global Ecology and Biogeography in 2011"

Status and distribution of mangrove forests of the world using earth observation satellite data

Abstract: Aim Our scientific understanding of the extent and distribution of mangrove forests of the world is inadequate. The available global mangrove databases, compiled using disparate geospatial data sources and national statistics, need to be improved.Here,we mapped the status and distributions of global mangroves using recently available Global Land Survey (GLS) data and the Landsat archive. Methods We interpreted approximately 1000 Landsat scenes using hybrid supervised and unsupervised digital image classification techniques. Each image was normalized for variation in solar angle and earth‐sun distance by converting the digital number values to the top-of-the-atmosphere reflectance. Ground truth data and existing maps and databases were used to select training samples and also for iterative labelling. Results were validated using existing GIS data and the published literature to map ‘true mangroves’. Results The total area of mangroves in the year 2000 was 137,760 km 2 in 118 countries and territories in the tropical and subtropical regions of the world. Approximately 75% of world’s mangroves are found in just 15 countries, and only 6.9% are protected under the existing protected areas network (IUCN I-IV). Our study confirms earlier findings that the biogeographic distribution of mangroves is generallyconfinedtothetropicalandsubtropicalregionsandthelargestpercentage of mangroves is found between 5° N and 5° S latitude. Main conclusions We report that the remaining area of mangrove forest in the world is less than previously thought. Our estimate is 12.3% smaller than the most recent estimate by the Food and Agriculture Organization (FAO) of the United Nations.We present the most comprehensive, globally consistent and highest resolution (30 m) global mangrove database ever created.We developed and used better mapping techniques and data sources and mapped mangroves with better spatial and thematic details than previous studies.

The HYDE 3.1 spatially explicit database of human‐induced global land‐use change over the past 12,000 years

Abstract: Aim This paper presents a tool for long-term global change studies; it is an update of the History Database of the Global Environment (HYDE) with estimates of some of the underlying demographic and agricultural driving factors. Methods Historical population, cropland and pasture statistics are combined with satellite information and specific allocation algorithms (which change over time) to create spatially explicit maps, which are fully consistent on a 5′ longitude/latitude grid resolution, and cover the period 10,000 bc to ad 2000. Results Cropland occupied roughly less than 1% of the global ice-free land area for a long time until ad 1000, similar to the area used for pasture. In the centuries that followed, the share of global cropland increased to 2% in ad 1700 (c. 3 million km2) and 11% in ad 2000 (15 million km2), while the share of pasture area grew from 2% in ad 1700 to 24% in ad 2000 (34 million km2) These profound land-use changes have had, and will continue to have, quite considerable consequences for global biogeochemical cycles, and subsequently global climate change. Main conclusions Some researchers suggest that humans have shifted from living in the Holocene (emergence of agriculture) into the Anthropocene (humans capable of changing the Earth's atmosphere) since the start of the Industrial Revolution. But in the light of the sheer size and magnitude of some historical land-use changes (e.g. as result of the depopulation of Europe due to the Black Death in the 14th century and the aftermath of the colonization of the Americas in the 16th century) we believe that this point might have occurred earlier in time. While there are still many uncertainties and gaps in our knowledge about the importance of land use (change) in the global biogeochemical cycle, we hope that this database can help global (climate) change modellers to close parts of this gap.

The effect of biodiversity on tree productivity: from temperate to boreal forests

Abstract: Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning.Experimental work has led to the proposal of mechanisms such as niche complementarity.However,few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions.(1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps.(1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables,as well as density and biodiversity indices,were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large-scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity‐ecosystem functioning (B-EF) research.

Increased water‐use efficiency during the 20th century did not translate into enhanced tree growth

Abstract: Aim The goals of this study are: (1) to determine whether increasing atmospheric CO2 concentrations and changing climate increased intrinsic water use efficiency (iWUE, as detected by changes in Δ13C) over the last four decades; and if it did increase iWUE, whether it led to increased tree growth (as measured by tree-ring growth); (2) to assess whether CO2 responses are biome dependent due to different environmental conditions, including availability of nutrients and water; and (3) to discuss how the findings of this study can better inform assumptions of CO2 fertilization and climate change effects in biospheric and climate models. Location A global range of sites covering all major forest biome types. Methods The analysis encompassed 47 study sites including boreal, wet temperate, mediterranean, semi-arid and tropical biomes for which measurements of tree ring Δ13C and growth are available over multiple decades. Results The iWUE inferred from the Δ13C analyses of comparable mature trees increased 20.5% over the last 40 years with no significant differences between biomes. This increase in iWUE did not translate into a significant overall increase in tree growth. Half of the sites showed a positive trend in growth while the other half had a negative or no trend. There were no significant trends within biomes or among biomes. Main conclusions These results show that despite an increase in atmospheric CO2 concentrations of over 50 p.p.m. and a 20.5% increase in iWUE during the last 40 years, tree growth has not increased as expected, suggesting that other factors have overridden the potential growth benefits of a CO2-rich world in many sites. Such factors could include climate change (particularly drought), nutrient limitation and/or physiological long-term acclimation to elevated CO2. Hence, the rate of biomass carbon sequestration in tropical, arid, mediterranean, wet temperate and boreal ecosystems may not increase with increasing atmospheric CO2 concentrations as is often implied by biospheric models and short-term elevated CO2 experiments.

Long-term shifts in abundance and distribution of a temperate fish fauna: a response to climate change and fishing practices

Abstract: Aim South-eastern Australia is a climate change hotspot with well-documented recent changes in its physical marine environment. The impact on and temporal responses of the biota to change are less well understood, but appear to be due to influences of climate,as well as the non-climate related past and continuing human impacts. We attempt to resolve the agents of change by examining major temporal and distributional shifts in the fish fauna and making a tentative attribution of causal factors. Location Temperate seas of south-eastern Australia. Methods Mixed data sources synthesized from published accounts, scientific surveys, spearfishing and angling competitions, commercial catches and underwater photographic records, from the ‘late 1800s’ to the ‘present’, were examined to determine shifts in coastal fish distributions. Results Forty-five species,representing 27 families (about 30% of the inshore fish families occurring in the region), exhibited major distributional shifts thought to be climate related. These are distributed across the following categories: species previously rare or unlisted (12), with expanded ranges (23) and/or abundance increases (30), expanded populations in south-eastern Tasmania (16) and extralimital vagrants (4).Another 9 species, representing 7 families, experienced longerterm changes (since the 1800s) probably due to anthropogenic factors, such as habitat alteration and fishing pressure: species now extinct locally (3), recovering (3), threatened (2) or with remnant populations (1). One species is a temporary resident periodically recruited from New Zealand. Of fishes exhibiting an obvious poleward movement, most are reef dwellers from three Australian biogeographic categories: widespread southern, western warm temperate (Flindersian) or eastern warm temperate (Peronian) species. Main conclusions Some of the region’s largest predatory reef fishes have become extinct in Tasmanian seas since the‘late 1800s’,most likely as a result of poorfishing practices. In more recent times, there have been major changes in the distribution patterns of Tasmanian fishes that correspond to dramatic warming observed in the local marine environment.

Pushing the limits in marine species distribution modelling: lessons from the land present challenges and opportunities

Abstract: Aim Species distribution models (SDMs) have been used to address a wide range of theoretical and applied questions in the terrestrial realm, but marine-based applications remain relatively scarce. In this review, we consider how conceptual and practical issues associated with terrestrial SDMs apply to a range of marine organisms and highlight the challenges relevant to improving marine SDMs. Location We include studies from both marine and terrestrial systems that encompass many geographic locations around the globe. Methods We first performed a literature search and analysis of marine and terrestrial SDMs in ISI Web of Science to assess trends and applications. Using knowledge from terrestrial applications, we critically evaluate the application of SDMs in marine systems in the context of ecological factors (dispersal, species interactions, aggregation and ontogenetic shifts) and practical considerations (data quality, alternative modelling approaches and model validation) that facilitate or create difficulties for model application. Results The relative importance of ecological factors to be considered when applying SDMs varies among terrestrial and marine organisms. Correctly incorporating dispersal is frequently considered an important issue for terrestrial models, but because there is greater potential for dispersal in the ocean, it is often less of a concern in marine SDMs. By contrast, ontogenetic shifts and feeding have received little attention in terrestrial SDM applications, but these factors are important to many marine SDMs. Opportunities also exist for applying more advanced SDM approaches in the marine realm, including mechanistic ecophysiological models, where water balance and heat transfer equations are simpler for some marine organisms relative to their terrestrial counterparts. Main conclusions SDMs have generally been under-utilized in the marine realm relative to terrestrial applications. Correlative SDM methods should be tested on a range of marine organisms, and we suggest further development of methods that address ontogenetic shifts and feeding interactions. We anticipate developments in, and cross-fertilization between, coupled correlative and process-based SDMs, mechanistic eco-physiological SDMs, and spatial population dynamic models for climate change and species invasion applications in particular. Comparisons of the outputs of different model types will provide insight that is useful for improved spatial management of marine species. © 2011 Blackwell Publishing Ltd.

When is a ‘forest’ a savanna, and why does it matter?

Abstract: Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high-rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C4 grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C4 grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life-history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire-resistant tree species in a C4 grass-dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.

Potential impacts of climate change on the environmental services of humid tropical alpine regions

Abstract: Aim Humid tropical alpine environments are crucial ecosystems that sustain biodiversity, biological processes, carbon storage and surface water provision. They are identified as one of the terrestrial ecosystems most vulnerable to global environmental change. Despite their vulnerability, and the importance for regional biodiversity conservation and socio-economic development, they are among the least studied and described ecosystems in the world. This paper reviews the state of knowledge about tropical alpine environments, and provides an integrated assessment of the potential threats of global climate change on the major ecosystem processes. Location Humid tropical alpine regions occur between the upper forest line and the perennial snow border in the upper regions of the Andes, the Afroalpine belt and Indonesia and Papua New Guinea. Results and main conclusions Climate change will displace ecosystem boundaries and strongly reduce the total area of tropical alpine regions. Displacement and increased isolation of the remaining patches will induce species extinction and biodiversity loss. Drier and warmer soil conditions will cause a faster organic carbon turnover, decreasing the below-ground organic carbon storage. Since most of the organic carbon is currently stored in the soils, it is unlikely that an increase in above-ground biomass will be able to offset soil carbon loss at an ecosystem level. Therefore a net release of carbon to the atmosphere is expected. Changes in precipitation patterns, increased evapotranspiration and alterations of the soil properties will have a major impact on water supply. Many regions are in danger of a significantly reduced or less reliable stream flow. The magnitude and even the trend of most of these effects depend strongly on local climatic, hydrological and ecological conditions. The extreme spatial gradients in these conditions put the sustainability of ecosystem management at risk.

ET come home: potential evapotranspiration in geographical ecology

Abstract: Aim Many macroecological analyses are based on analyses of climatological data, within which evapotranspiration estimates are of central importance. In this paper we evaluate and review the use of evapotranspiration models and data in studies of geographical ecology to test the likely sensitivity of the analyses to variation in the performance of different metrics of potential evapotranspiration. Location Analyses are based on: (1) a latitudinal transect of sites (FLUXNET) for 11 different land-cover types; and (2) globally gridded data.

Intra-specific variability and plasticity influence potential tree species distributions under climate change

Abstract: Aim To assess the effect of local adaptation and phenotypic plasticity on the potential distribution of species under future climate changes. Trees may be adapted to specific climatic conditions; however, species range predictions have classically been assessed by species distribution models (SDMs) that do not account for intra-specific genetic variability and phenotypic plasticity, because SDMs rely on the assumption that species respond homogeneously to climate change across their range, i.e. a species is equally adapted throughout its range, and all species are equally plastic. These assumptions could cause SDMs to exaggerate or underestimate species at risk under future climate change. Location The Iberian Peninsula. Methods Species distributions are predicted by integrating experimental data and modelling techniques. We incorporate plasticity and local adaptation into a SDM by calibrating models of tree survivorship with adaptive traits in provenance trials. Phenotypic plasticity was incorporated by calibrating our model with a climatic index that provides a measure of the differences between sites and provenances. Results We present a new modelling approach that is easy to implement and makes use of existing tree provenance trials to predict species distribution models under global warming. Our results indicate that the incorporation of intra-population genetic diversity and phenotypic plasticity in SDMs significantly altered their outcome. In comparing species range predictions, the decrease in area occupancy under global warming conditions is smaller when considering our survival–adaptation model than that predicted by a ‘classical SDM’ calibrated with presence–absence data. These differences in survivorship are due to both local adaptation and plasticity. Differences due to the use of experimental data in the model calibration are also expressed in our results: we incorporate a null model that uses survival data from all provenances together. This model always predicts less reduction in area occupancy for both species than the SDM calibrated with presence–absence. Main conclusions We reaffirm the importance of considering adaptive traits when predicting species distributions and avoiding the use of occurrence data as a predictive variable. In light of these recommendations, we advise that existing predictions of future species distributions and their component populations must be reconsidered.

Revisiting the historical distribution of Seasonally Dry Tropical Forests: new insights based on palaeodistribution modelling and palynological evidencegeb

Abstract: Aim To investigate the potential distribution of Seasonally Dry Tropical Forests (SDTFs) during the Quaternary climatic fluctuations; to reassess the formerly proposed ‘Pleistocenic arc hypothesis’ (PAH); and to identify historically stable and unstable areas of SDTF distributions in the light of palaeodistribution modelling. Location SDTFs in lowland cis-Andean eastern-central South America. Methods We first developed georeferenced maps depicting the current distributional extent of SDTFs under two distinct definitions (narrow and broad). We then generated occurrence datasets, which were used with current and past bioclimatic variables to predict SDTF occurrence by implementing the maximum entropy machine-learning algorithm. We obtained historical stability maps by overlapping the presence/absence projections of each of three climatic scenarios [current, 6 kyr bp during the Holocene, and 21 kyr bp during the Last Glacial Maximum (LGM)]. Finally, we checked the consistencies of the model prediction with qualitative comparisons of vegetation types inferred from available fossil pollen records. Results The present-day SDTF distribution is disjunct, but we provide evidence that it was even more disjunct during the LGM. Reconstructions support a progressive southward and eastward expansion of SDTFs on a continental scale since the LGM. No significant expansion of SDTFs into the Amazon Basin was detected. Areas of presumed long-term stability are identified and confirmed (the three nuclear regions, Caatinga, Misiones and Piedmont, plus the Chiquitano region), and these possibly acted as current and historical refugial areas. Main conclusions The LGM climate was probably too dry and cold to support large tracts of SDTF, which were restricted to climatically favourable areas relative to the present day (in contrast with the PAH, as it was originally conceived). Expansions of SDTFs are proposed to have occupied the southern portion of Caatinga nucleus more recently during the early–middle Holocene transition. We propose an alternative scenario amenable to further testing of an earlier SDTF expansion (either at the Lower Pleistocene or the Tertiary), followed by fragmentation in the LGM and secondary expansion in the Holocene. The stability maps were used to generate specific genetic predictions at both continental and regional scales (stable areas are expected to have higher genetic diversity and endemism levels than adjacent unstable areas) that can be used to direct field sampling to cover both stable (predicted refugia) and unstable (recently colonized) areas. Lastly, we discuss the possibility that SDTFs may experience future expansion under changing climate scenarios and that both stable and unstable areas should be prioritized by conservation initiatives.

Quantitative metrics of overlaps in Grinnellian niches: advances and possible drawbacks

Abstract: Aim Studies of environmental niche shift/niche conservatism that are based on species distribution modelling require a quantification of niche purity and potential overlap. Although various metrics have been proposed for this task, no comparisons of their performance are available yet that express the linearity of range shifts and error-proneness. Herein, we assess the performance of six niche overlap metrics using three sister pairs of plethodontid salamanders as well as artificial species to test for linearity of overlap curves, impacts of varying potential distribution sizes and study area sizes. Location North America, artificial environments. Methods Species distribution models for the salamanders were performed with Maxent, and artificial species were created in the R environment. Potential distributions of species with varying range sizes and extents of the study area were compared using the Bray–Curtis distance BC, Schoener's D, two different modifications of the Hellinger distance Imod, Icor, Pianka's O and Horn's R. Niche overlaps in ecological space were compared using linear discriminant analyses based on principal components. Results Simulations of niche overlaps revealed strong variations in the performance of the niche overlap metrics. In artificial species, BC and D performed best, followed by O, R and Icor, but the modified Hellinger distance Imod showed a nonlinear slope and a truncated range. Furthermore, the simulations suggest that, in proportionally small potential distributions on large grids, an inclusion of a high proportion of grid cells with low occurrence probabilities representing background noise may bias assessments of niche overlaps. Main conclusions Both the salamander examples and simulations suggest that Schoener's D and the Bray–Curtis distance BC are best suited to compute niche overlaps from potential distributions derived from species distribution models. However, like all analysed metrics, both D and BC are seriously affected by the inclusion of high numbers of grid cells where the species are probably absent, i.e. with low occurrence probabilities. Therefore, pre-processing to eliminate background noise in the potential distribution grids is highly recommended.

Treeline form – a potential key to understanding treeline dynamics

Abstract: Aim Treelines occur globally within a narrow range of mean growing season temperatures, suggesting that low-temperature growth limitation determines the position of the treeline. However, treelines also exhibit features that indicate that other mechanisms, such as biomass loss not resulting in mortality (dieback) and mortality, determine treeline position and dynamics. Debate regarding the mechanisms controlling treeline position and dynamics may be resolved by identifying the mechanisms controlling prominent treeline spatial patterns (or ‘form’) such as the spatial structure of the transition from closed forest to the tree limit. Recent treeline studies world-wide have confirmed a close link between form and dynamics. Location The concepts presented refer to alpine treelines globally. Methods In this review, we describe how varying dominance of three general ‘first-level’ mechanisms (tree performance: growth limitation, seedling mortality and dieback) result in different treeline forms, what ‘second-level’ mechanisms (stresses, e.g. freezing damage, photoinhibition) may underlie these general mechanisms, and how they are modulated by interactions with neighbours (‘third-level’ mechanisms). This hierarchy of mechanisms should facilitate discussions about treeline formation and dynamics. Results We distinguish four primary treeline forms: diffuse, abrupt, island and krummholz. Growth limitation is dominant only at the diffuse treeline, which is the form that has most frequently responded as expected to growing-season warming, whereas the other forms are controlled by dieback and seedling mortality and are relatively unresponsive. Main conclusions Treeline form provides a means for explaining the current variability in treeline position and dynamics and for exploring the general mechanisms controlling the responses of treelines to climatic change. Form indicates the relative dependence of tree performance on various aspects of the external climate (especially summer warmth versus winter stressors) and on internal feedbacks, thus allowing inferences on the type as well as strength of climate-change responses.

Variation in above-ground forest biomass across broad climatic gradients

Abstract: AimAn understanding of the relationship between forest biomass and climate is needed to predict the impacts of climate change on carbon stores.Biomass patterns have been characterized at geographically or climatically restricted scales,making it unclear if biomass is limited by climate in any general way at continental to global scales.Using a dataset spanning multiple climatic regions we evaluate the generality of published biomass‐climate correlations.We also combine metabolic theory and hydraulic limits to plant growth to first derive and then test predictions for how forest biomass should vary with maximum individual tree biomass and the ecosystem water deficit.

Cumulative nitrogen input drives species loss in terrestrial ecosystems

Abstract: Aim Elevated inputs of biologically reactive nitrogen (N) are considered to be one of the most substantial threats to biodiversity in terrestrial ecosystems. Several attempts have been made to scrutinize the factors driving species loss following excess N input, but generalizations across sites or vegetation types cannot yet be made. Here we focus on the relative importance of the vegetation type, the local environment (climate, soil pH, wet deposition load) and the experimentally applied (cumulative) N dose on the response of the vegetation to N addition. Location Mainly North America and Europe. Methods We conducted a large-scale meta-analysis of in situ N addition experiments in different vegetation types, focusing on the response of biomass and species richness. Results Whereas the biomass of grasslands and salt marshes significantly increased with N fertilization, forest understorey vegetation, heathlands, freshwater wetlands and bogs did not show any significant response. Graminoids significantly increased in biomass following N addition, whereas bryophytes significantly lost biomass; shrubs, forbs and lichens did not significantly respond. The yearly N fertilization dose significantly influenced the biomass response of grassland and salt marshes, while for the other vegetation types none of the collected predictor variables were of significant influence. Species richness significantly decreased with N addition in grasslands and heathlands [Correction added on 23 March 2011, after first online publication: ‘across all vegetation types’ changed to ‘in grasslands and heathlands’]. The relative change in species richness following N addition was significantly driven by the cumulative N dose. Main conclusions The decline in species richness with cumulative N input follows a negative exponential pathway. Species loss occurs faster at low levels of cumulative N input or at the beginning of the addition, followed by an increasingly slower species loss at higher cumulative N inputs. These findings lead us to stress the importance of including the cumulative effect of N additions in calculations of critical load values.

Choice of predictor variables as a source of uncertainty in continental‐scale species distribution modelling under climate change

Abstract: Aim: species distribution modelling is commonly used to guide future conservation policies in the light of potential climate change. However, arbitrary decisions during the model-building process can affect predictions and contribute to uncertainty about where suitable climate space will exist. For many species, the key climatic factors limiting distributions are unknown. This paper assesses the uncertainty generated by using different climate predictor variable sets for modelling the impacts of climate change. Location: Europe, 10° W to 50° E and 30° N to 60° N. Methods: using 1453 presence pixels at 30 arcsec resolution for the great bustard (Otis tarda), predictions of future distribution were made based on two emissions scenarios, three general climate models and 26 sets of predictor variables. Twenty-six current models were created, and 156 for both 2050 and 2080. Map comparison techniques were used to compare predictions in terms of the quantity and the location of presences (map comparison kappa, MCK) and using a range change index (RCI). Generalized linear models (GLMs) were used to partition explained deviance in MCK and RCI among sources of uncertainty. Results: the 26 different variable sets achieved high values of AUC (area under the receiver operating characteristic curve) and yet introduced substantial variation into maps of current distribution. Differences between maps were even greater when distributions were projected into the future. Some 64–78% of the variation between future maps was attributable to choice of predictor variable set alone. Choice of general climate model and emissions scenario contributed a maximum of 15% variation and their order of importance differed for MCK and RCI. Main conclusions: generalized variable sets produce an unmanageable level of uncertainty in species distribution models which cannot be ignored. The use of sound ecological theory and statistical methods to check predictor variables can reduce this uncertainty, but our knowledge of species may be too limited to make more than arbitrary choices. When all sources of modelling uncertainty are considered together, it is doubtful whether ensemble methods offer an adequate solution. Future studies should explicitly acknowledge uncertainty due to arbitrary choices in the model-building process and develop ways to convey the results to decision-makers

Beyond taxonomic diversity patterns: how do α, β and γ components of bird functional and phylogenetic diversity respond to environmental gradients across France?

Abstract: Aim To test how far can macroecological hypotheses relating diversity to environmental factors be extrapolated to functional and phylogenetic diversities, i.e. to the extent to which functional traits and evolutionary backgrounds vary among species in a community or region. We use a spatial partitioning of diversity where regional or γ-diversity is calculated by aggregating information on local communities, local or α-diversity corresponds to diversity in one locality, and turnover or β-diversity corresponds to the average turnover between localities and the region. Location France. Methods We used the Rao quadratic entropy decomposition of diversity to calculate local, regional and turnover diversity for each of three diversity facets (taxonomic, phylogenetic and functional) in breeding bird communities of France. Spatial autoregressive models and partial regression analyses were used to analyse the relationships between each diversity facet and environmental gradients (climate and land use). Results Changes in γ-diversity are driven by changes in both α- and β-diversity. Low levels of human impact generally favour all three facets of regional diversity and heterogeneous landscapes usually harbour higher β-diversity in the three facets of diversity, although functional and phylogenetic turnover show some relationships in the opposite direction. Spatial and environmental factors explain a large percentage of the variation in the three diversity facets (>60%), and this is especially true for phylogenetic diversity. In all cases, spatial structure plays a preponderant role in explaining diversity gradients, suggesting an important role for dispersal limitations in structuring diversity at different spatial scales. Main conclusions Our results generally support the idea that hypotheses that have previously been applied to taxonomic diversity, both at local and regional scales, can be extended to phylogenetic and functional diversity. Specifically, changes in regional diversity are the result of changes in both local and turnover diversity, some environmental conditions such as human development have a great impact on diversity levels, and heterogeneous landscapes tend to have higher diversity levels. Interestingly, differences between diversity facets could potentially provide further insights into how large- and small-scale ecological processes interact at the onset of macroecological patterns.

Biogeographical patterns of soil molecular microbial biomass as influenced by soil characteristics and management

Abstract: Aim The spatial organization of soil microbial communities on large scales and the identification of environmental factors structuring their distribution have been little investigated. The overall objective of this study was to determine the spatial patterning of microbial biomass in soils over a wide extent and to rank the environmental filters most influencing this distribution.

Bergmann's Rule – what's in a name?

Abstract: Despite the great interest it generates, the definition of Bergmann's Rule is vague and often contested. Debate focuses on whether the rule should be described in terms of pattern or process, what taxa it should apply to and what taxonomic level it should be associated with. Here I review the historical development of studies of Bergmann's Rule. I suggest that Bergmann thought that his rule should be strongest at the intra-specific level, rather than between closely related species as is usually thought. I argue that the rule is a pattern that can be studied regardless of mechanism in any taxon and at any taxonomic level.

A global comparative analysis of elevational species richness patterns of ferns

Abstract: Aim: Elevational gradients offer an outstanding opportunity to assess factors determining patterns of species richness, but along single transects potential explanatory factors often covary, making it difficult to distinguish between competing hypotheses. Many previous studies on plants have interpreted their results as supporting the mid-domain effect (MDE) as a major determinant of species richness, even when climatic factors showed similarly high explanatory power. We compared fern species richness along 20 elevational transects to quantify the relative contribution of climate and MDE as drivers of elevational richness patterns. Location: Twenty transects world-wide. Methods: Ferns were sampled in 1039 plots of 400–2500 m2 each. Mean annual precipitation and temperature, epiphytic bryophyte cover (as a proxy for air humidity) and MDE predictions were included as independent variables. For each transect, we calculated multiple linear models and partitioned the variance to assess the relative contribution of the independent variables, selecting the most parsimonious models based on Akaike weights and multi-model inference. Results: Along most individual gradients, nearly all variance of fern species richness that could be attributed to either space or MDEs was collinear with climatic factors. Yet, the comparison across transects showed that elevational richness patterns are most parsimoniously accounted for by climatic conditions, especially by low water availability at low elevations and in dry regions in general, and by low temperatures at high elevations and in extra-tropical regions. Main conclusions: Fern species richness is most closely related to climatic factors, and while MDE, surface area and metapopulation processes may somewhat modify the patterns, their importance has been overstated in the past. Future research challenges include determining whether the richness–climate relationship reflects: (1) a direct relationship through the physiological tolerance of the plants, (2) an indirect influence of climate on ecosystem productivity, or (3) an evolutionary legacy of longer or faster diversification processes under certain climatic conditions.

Trophic level scales positively with body size in fishes

Abstract: Aim The existence of a body size hierarchy across trophic connections is widely accepted anecdotally and is a basic assumption of many food-web models. Despite a strong theoretical basis, empirical evidence has been equivocal, and in general the relationship between trophic level and body size is often found to be weak or non-existent. Location Global (aquatic). Methods Using a global dataset for fishes (http://www.fishbase.org), we explored the relationship between body size and trophic position for 8361 fishes in 57 orders. Results Across all species, trophic position was positively related to maximum length (r2= 0.194, b= 0.065, P < 0.0001), meaning that a one-level increase in trophic level was associated with an increase in maximum length by a factor of 183. On average, fishes in orders that showed significantly positive trophic level–body size relations [mean = 51.6 cm ± 11.8 (95% confidence interval, CI)] were 86 cm smaller than fishes in orders that showed no relation [mean = 137.1 cm ± 50.3 (95% CI), P < 0.01]. A separate slopes model ANCOVA revealed that maximum length and trophic level were positively correlated for 47% (27 of 57) of orders, with two more orders showing marginally non-significant positive relations; no significant negative correlations were observed. The full model (order × body size) explained 37% of the variation between body size and trophic position (P < 0.0001). Main conclusions Our results support recent models which suggest that trophic level and body size should be positively correlated, and indicate that morphological constraints associated with gape limitation may play a stronger role in determining body size in smaller fishes. Differences among orders suggest that the nature of the trophic level–body size relation may be contingent, in part, on evolutionary history.

Meta‐analysis reveals the importance of matrix composition for animals in fragmented habitat

Abstract: Aim Connectivity is a key determinant of the distribution and abundance of organisms and is greatly influenced by anthropogenic landscape modification, yet we lack a synthetic perspective on the magnitude and extent of matrix effects on connectivity. We synthesize results from published studies to understand the importance of matrix effects on fragmented animal populations. Location Global. Methods We conduct a meta-analysis of 283 fragmented populations representing 184 terrestrial animal taxa to determine the strength of matrix composition effects on the occurrence and abundance of animals in fragmented habitat. Results Studies that use data on matrix composition report greater effects on abundance and occupancy of fragmented populations than studies that define connectivity without regard to the surrounding matrix (i.e. ‘binary’ studies that describe only characteristics of patch habitat). Main conclusions Our findings underscore that conservation strategies must consider the importance of matrix habitat, have important implications for metapopulation and metacommunity paradigms, and provide direct large-scale, multi-taxa evidence that matrix habitat is an important driver of ecological dynamics in heterogeneous landscapes.

How landscape dynamics link individual- to population-level movement patterns: A multispecies comparison of ungulate relocation data

Abstract: Aim To demonstrate how the interrelations of individual movements form large-scale population-level movement patterns and how these patterns are associated with the underlying landscape dynamics by comparing ungulate movements across species. Locations Arctic tundra in Alaska and Canada, temperate forests in Massachusetts, Patagonian Steppes in Argentina, Eastern Steppes in Mongolia. Methods We used relocation data from four ungulate species (barren-ground caribou, Mongolian gazelle, guanaco and moose) to examine individual movements and the interrelation of movements among individuals. We applied and developed a suite of spatial metrics that measure variation in movement among individuals as population dispersion, movement coordination and realized mobility. Taken together, these metrics allowed us to quantify and distinguish among different large-scale population-level movement patterns such as migration, range residency and nomadism. We then related the population-level movement patterns to the underlying landscape vegetation dynamics via long-term remote sensing measurements of the temporal variability, spatial variability and unpredictability of vegetation productivity. Results Moose, which remained in sedentary home ranges, and guanacos, which were partially migratory, exhibited relatively short annual movements associated with landscapes having very little broad-scale variability in vegetation. Caribou and gazelle performed extreme long-distance movements that were associated with broad-scale variability in vegetation productivity during the peak of the growing season. Caribou exhibited regular seasonal migration in which individuals were clustered for most of the year and exhibited coordinated movements. In contrast, gazelle were nomadic, as individuals were independently distributed and moved in an uncoordinated manner that relates to the comparatively unpredictable (yet broad-scale) vegetation dynamics of their landscape. Main conclusions We show how broad-scale landscape unpredictability may lead to nomadism, an understudied type of long-distance movement. In contrast to classical migration where landscapes may vary at broad scales but in a predictable manner, long-distance movements of nomadic individuals are uncoordinated and independent from other such individuals. Landscapes with little broad-scale variability in vegetation productivity feature smaller-scale movements and allow for range residency. Nomadism requires distinct integrative conservation strategies that facilitate long-distance movements across the entire landscape and are not limited to certain migration corridors.

Connectivity and the development of population genetic structure in Indo-West Pacific coral reef communities

Abstract: Aim: To identify connectivity patterns among coral reefs of the Indo-West Pacific. Projecting connectivity forward in time provides a framework for studying long-term source–sink dynamics in the region, and makes it possible to evaluate the manner in which migration shapes population genetic structure at regional scales. This information is essential for addressing critical gaps in knowledge for conservation planning efforts in one of the most biologically diverse regions on earth. Location: Coral reefs of the Indo-West Pacific, ranging from 15° S to 30° N and 95° E to 140° E. Methods: Individual-based biophysical dispersal models were used in conjunction with matrix projection to identify the expected patterns of exchange between coral reefs over time. Results: Present-day oceanographic conditions lead to the transport of larvae from the South China Sea into the Coral Triangle region via the Sulu Sea, and from northern Papua New Guinea and the Solomon Islands via Halmahera. The directionality of the system leads to the expected accumulation of organisms from outlying areas into the Coral Triangle region over time, particularly in the vicinity of the Maluku Islands and eastern Sulawesi. Coral reefs in Papua New Guinea, the Sulu Archipelago and areas within the Philippines are expected to be areas of high diversity as well. Main conclusions: Biophysical dispersal models, used in conjunction with matrix projection, provide an effective means of simulating connectivity structure across the Indo-West Pacific and thereby evaluating the directionality of genetic diversity. Migration appears to have a significant influence on population genetic structure in the region. Based on present-day ocean currents, coral reefs in the South China Sea, northern Papua New Guinea and the Solomon Islands are contributing to high levels of diversity in the Coral Triangle.

Geographic disparities and moral hazards in the predicted impacts of climate change on human populations

Abstract: Aim It has been qualitatively understood for a long time that climate change will have widely varying effects on human well-being in different regions of the world The spatial complexities underlying our relationship to climate and the geographical disparities in human demographic change have, however, precluded the development of global indices of the predicted regional impacts of climate change on humans Humans will be most negatively affected by climate change in regions where populations are strongly dependent on climate and favourable climatic conditions declineHere we use the relationship between the distribution of human population density and climate as a basis to develop the first global index of predicted impacts of climate change on human populations Location Global

Asymmetric boundary shifts of tropical montane Lepidoptera over four decades of climate warming

Abstract: Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered.Location Mount Kinabalu, Sabah, Malaysia.Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations.Results Species that reached their upper limits at 2500-2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (-300 m in elevational extent and -45 km2 in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m).Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.

Cross-scale variation in species richness–environment associations

Abstract: Aim The scale dependence of many ecological patterns and processes implies that general inference is reliant on obtaining scale-response curves over a large range of grains. Although environmental correlates of richness have been widely studied, comparisons among groups have usually been applied at single grains. Moreover, the relevance of environment–richness associations to fine-grain assemblages has remained surprisingly unclear. We present a first global cross-scale assessment of environment–richness associations for birds, mammals and amphibians from 2000 km down to c. 20 km. Location World-wide. Methods We performed an extensive survey of the literature for well-sampled terrestrial vertebrate inventories over clearly defined small extents. Coarser grain richness was estimated from the intersection of extent-of-occurrence range maps with concentric equal-distance circles around fine-grain assemblage location centroids. General linear and simultaneous autoregressive models were used to relate richness at the different grains to environmental correlates. Results The ability of environmental variables to explain species richness decreases markedly toward finer grains and is lowest for fine-grained assemblages. A prominent transition in importance occurs between productivity and temperature at increased grains, which is consistent with the role of energy affecting regional, but not local, richness. Variation in fine-grained predictability across groups is associated with their purported grain of space use, i.e. highest for amphibians and narrow-ranged and small-bodied species. Main conclusions We extend the global documentation of environment–richness associations to fine-grained assemblages. The relationship between fine-grained predictability of a group and its ecological characteristics lends empirical support to the idea that variation in species fine-grained space use may scale up to explain coarse-grained diversity patterns. Our study exposes a dramatic and taxonomically variable scale dependence of environment–richness associations and suggests that environmental correlates of richness may hold limited information at the level of communities.

Carbon : nitrogen stoichiometry in forest ecosystems during stand development

Abstract: Aim Carbon (C) and nitrogen (N) stoichiometry is a critical indicator of biogeochemical coupling in terrestrial ecosystems. However, our current understanding of C : N stoichiometry is mainly derived from observations across space, and little is known about its dynamics through time. Location Global secondary forests. Methods We examined temporal variations in C : N ratios and scaling relationships between N and C for various ecosystem components (i.e. plant tissue, litter, forest floor and mineral soil) using data extracted from 39 chronosequences in forest ecosystems around the world. Results The C : N ratio in plant tissue, litter, forest floor and mineral soil exhibited large variation across various sequences, with an average of 145.8 ± 9.4 (mean ± SE), 49.9 ± 3.0, 38.2 ± 3.1 and 18.5 ± 0.9, respectively. In most sequences, the plant tissue C : N ratio increased significantly with stand age, while the C : N ratio in litter, forest floor and mineral soil remained relatively constant over the age sequence. N and C scaled isometrically (i.e. the slope of the relationship between log-transformed N and C is not significantly different from 1.0) in litter, forest floor and mineral soil both within and across sequences, but not in plant tissue either within or across sequences. The C : N ratio was larger in coniferous forests than in broadleaf forests and in temperate forests than in tropical forests. In contrast, the N–C scaling slope did not reveal significant differences either between coniferous and broadleaf forests or between temperate and tropical forests. Main conclusions These results suggest that C and N become decoupled in plants but remain coupled in other ecosystem components during stand development.

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Abstract: Aim? The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. ...

Plant traits and extinction in urban areas: a meta-analysis of 11 cities

Abstract: Aim Urban environments around the world share many features in common, including the local extinction of native plant species. We tested the hypothesis that similarity in environmental conditions among urban areas should select for plant species with a particular suite of traits suited to those conditions, and lead to the selective extinction of species lacking those traits. Location Eleven cities with data on the plant species that persisted and those that went locally extinct within at least the last 100 years following urbanization. Methods We compiled data on 11 plant traits for 8269 native species in the 11 cities and used hierarchical logistic regression models to identify the degree to which traits could distinguish species that persisted from those that went locally extinct in each city. The trait effects from each city were then combined in a meta-analysis. Results The cities fell into two groups: those with relatively low rates of extinction (less than 0.05% species per year – Adelaide, Hong Kong, Los Angeles, San Diego and San Francisco), for which no traits reliably predicted the pattern of extinction, and those with higher rates of extinction (> 0.08% species per year – Auckland, Chicago, Melbourne, New York, Singapore and Worcester, MA), where short-statured, small-seeded plants were more likely to go extinct. Main conclusions Our analysis reveals patterns in trait selectivity consistent with local studies, suggesting some consistency in trait selection by urbanization. Overall, however, few traits reliably predicted the pattern of plant extinction across cities, making it difficult to identify a priori the extinction-prone species most likely to be affected by urban expansion.