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Background: To what extent sex-related differences in cardiorespiratory fitness (CRF) impact postoperative patient mortality and corresponding implications for surgical risk stratification remains to be established. Methods: To examine this, we recruited 640 patients (366 males vs. 274 females) who underwent cardiopulmonary exercise testing prior to elective colorectal surgery. Patients were defined high risk if peak oxygen uptake was −1 min−1 and ventilatory equivalent for carbon dioxide at ‘anaerobic threshold’ >34. Between-sex CRF and mortality was assessed, and sex-specific CRF thresholds predictive of mortality was calculated. Results: Seventeen percent of deaths were attributed to sub-threshold CRF, which was higher than established risk factors for cardiovascular disease (CVD). The group (independent of sex) exhibited a 5-fold higher mortality (high vs. low risk patients hazard ratio = 4.80, 95% confidence interval 2.73–8.45, p < 0.001). Females exhibited 39% lower CRF (p < 0.001) with more classified high risk than males (36 vs. 23%, p = 0.001), yet mortality was not different (p = 0.544). Upon reformulation of sex-specific CRF thresholds, lower cut-offs for mortality were observed in females, and consequently, fewer (20%) were stratified with sub-threshold CRF compared to the original 36% (p < 0.001). Conclusions: Low CRF accounted for more deaths than traditional CVD risk factors, and when CRF was considered relative to sex, the disproportionate number of females stratified unfit was corrected. These findings support clinical consideration of ‘sex-specific’ CRF thresholds to better inform postoperative mortality and improve surgical risk stratification.

Background In the United Kingdom, the most common surgical approach for repair of open abdominal aortic aneurysms (AAAs) is transperitoneal (TP). However, retroperitoneal (RP) approach is favored in those with more complex vascular anatomy often requiring a cross-clamp on the aorta superior to the renal arteries. This study compared these approaches in patients matched on all major demographic, comorbid, anatomic, and physiological variables. Methods Fifty-seven patients (TP: n = 24; RP: n = 33) unsuitable for endovascular aneurysm repair underwent preoperative cardiopulmonary exercise testing prior to open AAA repair. The surgical approach undertaken was dictated by individual surgeon preference. Postoperative mortality, complications, and length of hospital stay (LoS) were recorded. Patients were further stratified according to infrarenal (IR) or suprarenal/supraceliac (SR/SC) surgical clamping. Systemic inflammation (C-reactive protein) and renal function (serum creatinine and estimated glomerular filtration rate) were recorded. Results Twenty-three (96%) of TP patients only required an IR clamp compared with 12 (36%) in the RP group. Postoperative systemic inflammation was lower in RP patients (p = 0.002 vs. TP) and fewer reported pulmonary/gastrointestinal complications whereas renal impairment was more marked in those receiving SR/SC clamps (p Conclusion Despite the demands of more complicated vascular anatomy, the clinical and economic benefits highlighted by these findings justify the more routine adoption of the RP approach for complex AAA repair.

Ecosystem functioning and services have provided a rationale for conservation over the past decades. Intertidal muddy sediments perform crucial functions, including erosion protection, nutrient cycling and carbon sequestration. Key mediators within this system include microphytobenthic biofilms, which bind sediment and influence nutrient biogeochemistry, and wading birds which feed extensively on sediment dwelling infauna, which in turn graze biofilms. Some wader species, specifically small sandpipers, have recently been found to interact directly with biofilms, e.g. by grazing, adding complexity to this food web. It has been suggested that infauna grazing by waders may lead to a reduction in biofilm grazing pressure by infauna. If this was the case it would likely have significant top-down, trophic cascade effects on key mudflat ecosystem functions. We carried out an exclusion experiment on the Colne Estuary, Essex, UK to examine whether wader presence significantly affects sediment erodibility, microphytobenthos fluorescence (Fo) and sediment-water nutrient fluxes (nitrate, nitrite, ammonia, phosphate and dissolved organic carbon (DOC)). Excluding wading birds caused significant changes in regulating and provisioning ecosystem functions, including mudflat erodibility and nutrient fluxes (with the exception of DOC). Fo was significantly greater, sediment critical shear strength increased, and the magnitude of nutrient fluxes increased where waders were absent. Excluding waders did not cause a significant difference in infauna density within the sediment, suggesting that a trophic cascade was not the cause of the measured effects. The dominant wading species at the site, dunlin Calidris alpina, is reported to consume biofilm and is considered to play a key role in maintenance of ecosystem stability on mudflats. After a period of 23 days, the arrival of the mud snail Peringia ulvae appeared to compensate for the absence of waders, consuming biofilm and causing a reduction in surface biofilm differences between presence/absence plots. This study introduces a previously unknown effect of wader foraging and bioturbation within this system, and highlights an area of wader science which could aid joint conservation and human provisioning action.

Metabarcoding has been used in a range of ecological applications such as taxonomic assignment, dietary analysis and the analysis of environmental DNA. However, after a decade of use in these applications there is little consensus on the extent to which proportions of reads generated corresponds to the original proportions of species in a community. To quantify our current understanding, we conducted a structured review and meta‐analysis. The analysis suggests that a weak quantitative relationship may exist between the biomass and sequences produced (slope = 0.52 ± 0.34, p

Environmental conditions play a major role in shaping the spatial distributions of pathogens, which in turn can drive local adaptation and divergence in host genetic diversity. Haemosporidians, such as Plasmodium (malaria), are a strong selective force, impacting survival and fitness of hosts, with geographic distributions largely determined by habitat suitability for their insect vectors. Here, we have tested whether patterns of fine‐scale local adaptation to malaria are replicated across discrete, ecologically differing island populations of Berthelot's pipits Anthus berthelotii. We sequenced TLR4, an innate immunity gene that is potentially under positive selection in Berthelot's pipits, and two SNPs previously identified as being associated with malaria infection in a genome‐wide association study (GWAS) in Berthelot's pipits in the Canary Islands. We determined the environmental predictors of malaria infection, using these to estimate variation in malaria risk on Porto Santo, and found some congruence with previously identified environmental risk factors on Tenerife. We also found a negative association between malaria infection and a TLR4 variant in Tenerife. In contrast, one of the GWAS SNPs showed an association with malaria risk in Porto Santo, but in the opposite direction to that found in the Canary Islands GWAS. Together, these findings suggest that disease‐driven local adaptation may be an important factor in shaping variation among island populations., We have investigated patterns of fine‐scale local adaptation to malaria between two ecologically divergent populations of an island bird. Our results found contrasting associations with malaria infection at candidate immune loci between the islands, suggesting that disease‐driven local adaptation may be an important factor in shaping variation among island populations.

Managing forest plantation stands in a way that retains productivity targets, but that also fosters biodiversity and stand resilience are key sustainable forest management goals. Current forestry policy advocates a diversification of forest stands to achieve these goals, favouring mixed age structures and polycultures over single-aged monocultures. Evidence is lacking, however, to support this management recommendation for biodiversity gains and related ecosystem service delivery. We used indices of taxonomic diversity and functional structure to compare ground vegetation communities in mixed and pure stands of Scots pine ( Pinus sylvestris ) and pedunculate oak ( Quercus robur ) in each of three study regions. We categorised the 91 vascular plant species identified into functional effect and response groups. We tested the hypotheses that ground vegetation communities (i) differ significantly in structure and composition between Scots pine and oak monocultures and (ii) show enhanced levels of taxonomic and functional diversity and functional redundancy in mixed stands of Scots pine and oak compared with monocultures. We explored the implications of any differences in the functional structure of ground vegetation communities in the different stand types on two ecosystem services: nutrient availability and levels of resource provisioning for herbivores. Nine functional response groups (RG) and seven functional effect groups (EG) were identified with considerable overlap in the RG and EG species grouping. Three RGs had traits characteristic of forests (spring flowering herbs, tree saplings and shrubs/climbers), one RG had traits characteristic of open habitats (annual ruderals) and the remaining RGs had more generalist traits (anemochorous perennials, graminoids and short perennials). No significant differences were found among stand types in terms of taxonomic diversity or richness of the different functional trait groups. Ground vegetation communities in the three study regions also had similar levels of functional redundancy across stand types. However, Scots pine and oak monocultures harboured significantly different abundances of species with distinct functional traits. In all three study regions, anemochorous perennials were significantly more abundant in Scots pine monocultures than oak monocultures, while two core forest groups (shrubs/climbers, spring flowering herbs) were significantly more abundant in oak monocultures. Mixed stands had intermediate abundances of these functional groups. These differences have implications for the comparative availability of food resources and shelter for wildlife, but also the mobilisation and temporal availability of nutrients in the two monocultures. Thus, mixtures of Scots pine and pedunculate oak can temper significant tree species identity influences on ground vegetation functional diversity.

Understanding the spatial scale at which selection acts upon adaptive genetic variation in natural populations is fundamental to our understanding of evolutionary ecology, and has important ramifications for conservation. The environmental factors to which individuals of a population are exposed can vary at fine spatial scales, potentially generating localized patterns of adaptation. Here, we compared patterns of neutral and major histocompatibility complex (MHC) variation within an island population of Berthelot's pipit (Anthus berthelotii) to assess whether landscape-level differences in pathogen-mediated selection generate fine-scale spatial structuring in these immune genes. Specifically, we tested for spatial associations between the distribution of avian malaria, and the factors previously shown to influence that distribution, and MHC variation within resident individuals. Although we found no overall genetic structure across the population for either neutral or MHC loci, we did find localized associations between environmental factors and MHC variation. One MHC class I allele (ANBE48) was directly associated with malaria infection risk, while the presence of the ANBE48 and ANBE38 alleles within individuals correlated (positively and negatively, respectively) with distance to the nearest poultry farm, an anthropogenic factor previously shown to be an important determinant of disease distribution in the study population. Our findings highlight the importance of considering small spatial scales when studying the patterns and processes involved in evolution at adaptive loci.

1. The ability to identify and quantify the constituent plant species that make up a mixed-species sample of pollen has important applications in ecology, conservation, and agriculture. Recently, metabarcoding protocols have been developed for pollen that can identify constituent plant species, but there are strong reasons to doubt that metabarcoding can accurately quantify their relative abundances. A PCR-free, shotgun metagenomics approach has greater potential for accurately quantifying species relative abundances, but applying metagenomics to eukaryotes is challenging due to low numbers of reference genomes.2. We have developed a pipeline, RevMet (Reverse Metagenomics), that allows reliable and semi-quantitative characterization of the species composition of mixed-species eukaryote samples, such as bee-collected pollen, without requiring reference genomes. Instead, reference species are represented only by ‘genome skims’: low-cost, low-coverage, short-read sequence datasets. The skims are mapped to individual long reads sequenced from mixed-species samples using the MinION, a portable nanopore sequencing device, and each long read is uniquely assigned to a plant species.3. We genome-skimmed 49 wild UK plant species, validated our pipeline with mock DNA mixtures of known composition, and then applied RevMet to pollen loads collected from wild bees. We demonstrate that RevMet can identify plant species present in mixed-species samples at proportions of DNA ≥1%, with few false positives and false negatives, and reliably differentiate species represented by high versus low amounts of DNA in a sample.4. The RevMet pipeline could readily be adapted to generate semi-quantitative datasets for a wide range of mixed eukaryote samples, which could include characterising diets, quantifying allergenic pollen from air samples, quantifying soil fauna, and identifying the compositions of algal and diatom communities. Our per-sample costs were £90 per genome skim and £60 per pollen sample, and new versions of sequencers available now will further reduce these costs.

Dung beetles (Scarabaeidae: Scarabaeinae) are sensitive to habitat perturbations and are easily studied, making them an ideal taxonomic group with which to evaluate the effects of low-intensity anthropogenic disturbances such as Reduced-Impact Logging. Here we examine the effect of a certified Reduced-Impact Logging operation on dung beetles, and demonstrate their suitability for use in rapid ecological impact studies. We sampled dung beetle assemblages, environmental variables and timber extraction rates across four treatment groups in closed canopy and canopy gaps in logged and unlogged forest in Guyana. Community analysis revealed that logged forest supported a more uniform dung beetle assemblage compared to unlogged forest. Differences in assemblage structure were driven by dissimilarity between closed canopy treatments, as plots in artificial and natural canopy gaps supported comparable assemblages. Indicator analyses were conducted across treatments, using a new approach (CLAM) and two well-established methods (IndVal, SIMPER). Two species respectively were classified as indicators of logged (Hansreia affinis and Eurysternus caribaeus) and unlogged forest (Canthidium aff. centrale and Deltochilum (Calhyboma) carinatum). BIO-ENV analysis demonstrated that tree extraction intensity, bare ground cover, and ground cover by leaf material were key factors influencing dung beetle assemblages. Despite the relatively low-impact of Reduced-Impact Logging reported by previous studies, we find that dung beetles are sensitive to even small changes in environmental conditions as a result of this form of anthropogenic disturbance. As dung beetles are a highly responsive taxonomic group, we illustrate that they represent a valuable taxon that can be used to critically evaluate best practice forestry operations and other disturbance activities, particularly in time constrained studies (e.g., rapid monitoring and environmental impact assessments). However, we recommend the use of multiple indicator analyses to monitor potential changes in assemblage composition, due to a lack of congruence between methods.

How the environment influences the transmission and prevalence of disease in a population of hosts is a key aspect of disease ecology. The role that environmental factors play in host-pathogen systems has been well studied at large scales, that is, differences in pathogen pressures among separate populations of hosts or across land masses. However, despite considerable understanding of how environmental conditions vary at fine spatial scales, the effect of these parameters on host-pathogen dynamics at such scales has been largely overlooked. Here, we used a combination of molecular screening and GIS-based analysis to investigate how environmental factors determine the distribution of malaria across the landscape in a population of Berthelot's pipit (Anthus berthelotii, Bolle 1862) on the island of Tenerife (Canary Islands, Spain) using spatially explicit models that account for spatial autocorrelation. Minimum temperature of the coldest month was found to be the most important predictor of malaria infection at the landscape scale across this population. Additionally, anthropogenic factors such as distance to artificial water reservoirs and distance to poultry farms were important predictors of malaria. A model including these factors, and the interaction between distance to artificial water reservoirs and minimum temperature, best explained the distribution of malaria infection in this system. These results suggest that levels of malaria infection in this endemic species may be artificially elevated by the impact of humans. Studies such as the one described here improve our understanding of how environmental factors, and their heterogeneity, affect the distribution of pathogens within wild populations. The results demonstrate the importance of measuring fine-scale variation - and not just regional effects - to understand how environmental variation can influence wildlife diseases. Such understanding is important for predicting the future spread and impact of disease and may help inform disease management programmes as well as the conservation of specific host species.

Commercial selective logging and the conversion of primary and degraded forests to agriculture are the biggest threats to tropical biodiversity. Our understanding of the impacts of these disturbances and the resulting local extinctions on the functional roles performed by the remaining species is limited. We address this issue by examining functional diversity (FD), which quantifies a range of traits that affect a species' ecological role in a community as a single continuous metric. We calculated FD for birds across a gradient of disturbance from primary forest through intensively logged forest to oil palm plantations on previously forested land in Borneo, Southeast Asia, a hotspot of imperilled biodiversity. Logged rainforest retained similar levels of FD to unlogged rainforest, even after two logging rotations, but the conversion of logged forest to oil palm resulted in dramatic reductions in FD. The few remaining species in oil palm filled a disproportionately wide range of functional roles but showed very little clustering in terms of functional traits, suggesting that any further extinctions from oil palm would reduce FD even further. Determining the extent to which the changes we recorded were due to under-utilization of resources within oil palm or a reduction in the resources present is an important next step. Nonetheless our study improves our understanding of the stability and resilience of functional diversity in these ecosystems and of the implications of land-use changes for ecosystem functioning.

Non-native species can be major drivers of biodiversity loss and cause economic damage. Predicting the potential distribution of a non-native species, and understanding the environmental factors that limit this distribution, is useful for informing their potential management. This is often carried out using species distribution models (SDMs) that attempt to classify grid cells as suitable or unsuitable for a species based on a set of environmental covariates. A key assumption of SDMs is that a species is in equilibrium with its environment. Spreading non-native species often violate this assumption due to dispersal limitation. We present a simple method for dealing with this problem: dispersal weighting (DW). This uses the probability that a species can disperse to a grid cell to weight a SDM. We use simulations to compare the ability of DW and unweighted models at parameterising the true species-environment relationship (SER) of a simulated species, and to test their ability at predicting the future distribution of this species. We investigate how varying the degree of spatial autocorrelation in explanatory variables affects the performance of the methods. Dispersal weighting models outperformed unweighted models at parameterising the SER, and at predicting the future distribution of the species when dispersal probabilities were incorporated into the model predictions. Unweighted models had a stronger tendency than DW models to overestimate the magnitude of relationships with spatially autocorrelated explanatory variables, but underestimate the magnitude of relationships with randomly distributed variables. We then applied our method to a real case study, using it to model the distribution of the non-native common waxbill Estrilda astrild in the Iberian Peninsula as a function of climate and land-use variables. The relative performance of DW and unweighted models reflected the results of the simulation. We conclude that DW models perform better than unweighted models at modelling the true SER of non-native species, and recommend using DW whenever enough data exist to create a dispersal model. © 2012 The Authors. Methods in Ecology and Evolution © 2012 British Ecological Society.

Aim: British estuarine ecosystems support large populations of protected migratory waders. Understanding how wader communities vary spatially and how they may be changing temporally can greatly improve the understanding of these dynamic ecosystems. Here, we explore the variation in functional diversity (using a range of morphological and ecological traits) in order to identify the processes shaping wader communities on British estuaries and how these processes may be changing. Location: England, Wales and Scotland. Methods: We use national survey data (Wetland Bird Survey) from 1980/1981 to 2006/2007 winter to calculate functional diversity (FD) – an index that measures trait dispersion – in wader communities on 100 estuaries. We test for evidence of non-random patterns of diversity and explore the relative importance of two key processes, environmental filtering and competition, in shaping these communities. Results: The observed FD was significantly and positively associated with species richness and to a lesser extent estuary area, followed by longitude. An increase in observed FD was observed since 1980, supported by a small but significant slope. In the majority of cases, changes in FD were mirrored by changes in species richness. Observed FD was on average lower than expected by chance, as indicated by a negative value of observed minus expected FD. However, this difference became less negative over time, with observed minus expected FD values increasing slightly, but significantly, over the study period. Main conclusions: Wader FD varies across British estuaries, and the relative influence of the processes by which communities are structured appears to be changing through time. We discuss the potential drivers underlying these patterns and the importance of identifying such drivers for the protection of wader communities.

Aim Niche partitioning within species assemblages is thought to influence species packing and/or total niche space occupied. The evolution of dung beetles (Scarabaeinae) is likely to have been strongly influenced by inter-specific competition, leading to niche partitioning. We consider whether local-scale processes leave a signature in regional patterns of functional diversity in dung beetle assemblages, and investigate the correlation between total exploited ecomorphological space and density of species packing with increased species richness. We test whether ecomorphological space occupied by local assemblages

Deforestation is a main driver of climate change and biodiversity loss. An incentive mechanism to reduce emissions from deforestation and forest degradation (REDD) is being negotiated under the United Nations Framework Convention on Climate Change. Here we use the best available global data sets on terrestrial biodiversity and carbon storage to map and investigate potential synergies between carbon and biodiversity-oriented conservation. A strong association (rS= 0.82) between carbon stocks and species richness suggests that such synergies would be high, but unevenly distributed. Many areas of high value for biodiversity could be protected by carbon-based conservation, while others could benefit from complementary funding arising from their carbon content. Some high-biodiversity regions, however, would not benefit from carbon-focused conservation, and could become under increased pressure if REDD is implemented. Our results suggest that additional gains for biodiversity conservation are possible, without compromising the effectiveness for climate change mitigation, if REDD takes biodiversity distribution into account.

Aims Tests of the energy hypothesis for the large-scale distribution of species richness have largely been concerned with the influence of two alternative forms of environmental energy, temperature and energy from primary productivity, both of which (at least in terrestrial systems) peak within the tropics. Taxa showing extra-tropical diversity peaks present a potential challenge to the generality of species–energy theory. One such group are pelagic seabirds of the order Procellariiformes that show not only an extra-tropical diversity peak but one confined to the Southern Ocean, hence a highly asymmetric one. They are distinct in being exceptionally adapted to take advantage of wind energy, which they may rely on for long-distance ocean foraging for the patchy resources needed to meet their energetic needs. Wind represents a readily available source of kinetic energy, shows a strong latitudinal gradient, and has been largely omitted from species–energy theory. Moreover, maximal benefits of wind are likely to be afforded in areas of greatest available contiguous ocean extent. We compare the relative importance of wind speed, ocean productivity (chlorophyll concentration), air temperature and available ocean extent (distance) in explaining large-scale global distribution of procellariiform species richness across the world's oceans. Location Global, oceanic. Methods Hierarchical partitioning, model selection, ordinary least squares (OLS) and spatial generalized least squares (GLS) regression. Results Hierarchical partitioning of non-spatial regression models indicates that ocean distance is the most important predictor of procellariiform species richness followed by wind speed and then temperature. In contrast, that of spatial regression models indicates the roughly equal importance of ocean distance and temperature, followed by wind speed. Although contributing additional model fit, ocean productivity is consistently the weakest predictor. Best-fit models include all four predictors and explain 67% of observed variation. The species–productivity relationship is negative overall, while the species–temperature relationship is hump-shaped. In contrast, ocean distance and wind speed are positively associated with species richness. Conclusions Large-scale procellariiform species richness distribution may represent a trade-off in the use of different energy forms, being highest in Southern Ocean areas where productive energy and temperature are relatively low, but where available ocean foraging extent and wind energy required to utilize it are near-maximal.

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal-area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower-taxon richness relative to higher-taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher-taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co-occurrence. Species and genus co-occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher-taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long-term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.

The growing proportion of human populations living in urban areas, and consequent trends of increasing urban expansion and densification fuel a need to understand how urban form and land use affect environmental quality, including the availability of urban green spaces. Here we use Sheffield as a case study of city-wide relationships between urban green space extent, quality (vegetation cover and tree-cover), and gradients in urban form and topography. The total area of buildings and length of the road network are equally strong negative predictors of extent of green space, while the former predictor is a more important negative influence upon green space quality. Elevation positively influences extent of green space but negatively influences tree-cover. In contrast, slope of terrain positively influences green space quality and is the best predictor of tree-cover. Overall housing density is a more important negative predictor of extent of green space and tree-cover than the densities of individual housing types. Nevertheless, the latter are more important influences upon levels of vegetation cover. Threshold effects of densities of different housing types suggest opportunities for optimising green space quality, with implications for housing policy. Variation in ecological quality of green space may partly reflect different historical intensities of industrial activity.

Green spaces play a crucial role in supporting urban ecological and social systems, a fact recognised in public policy commitments in both the UK and Europe. The amount of provision, the distribution of green space and the ease of access to such spaces are key contributors to social and ecological function in urban environments. We measured distance along the transport network to public green space available to households in Sheffield, and compared this with the distribution of private garden space. In addition, we used a geodemographic database, Mosaic UK, to examine how access to green space varies across different sectors of society. Public green spaces are chronically underprovided relative to recommended targets. For example, 64% of Sheffield households fail to meet the recommendation of the regulatory agency English Nature (EN), that people should live no further than 300 m from their nearest green space. Moreover, this figure rises to 72% if we restrict attention to municipal parks recognised by the local council. There is an overall reduction in coverage by green space when moving from neighbourhoods where green space is primarily publicly provided to those where it is privately provided. While access to public green space varies significantly across different social groups, those enjoying the greatest access include more deprived groups and older people. This study highlights the need for additional green space to be created and existing green space to be protected in light of increasing development pressure.

Species distributional or trait data based on range map (extent-of-occurrence) or atlas survey data often display spatial autocorrelation, i.e. locations close to each other exhibit more similar values than those further apart. If this pattern remains present in the residuals of a statistical model based on such data, one of the key assumptions of standard statistical analyses, that residuals are independent and identically distributed (i.i.d), is violated. The violation of the assumption of i.i.d. residuals may bias parameter estimates and can increase type I error rates (falsely rejecting the null hypothesis of no effect). While this is increasingly recognised by researchers analysing species distribution data, there is, to our knowledge, no comprehensive overview of the many available spatial statistical methods to take spatial autocorrelation into account in tests of statistical significance. Here, we describe six different statistical approaches to infer correlates of species? distributions, for both presence/absence (binary response) and species abundance data (poisson or normally distributed response), while accounting for spatial autocorrelation in model residuals: autocovariate regression; spatial eigenvector mapping; generalised least squares; (conditional and simultaneous) autoregressive models and generalised estimating equations. A comprehensive comparison of the relative merits of these methods is beyond the scope of this paper. To demonstrate each method?s implementation, however, we undertook preliminary tests based on simulated data. These preliminary tests verified that most of the spatial modeling techniques we examined showed good type I error control and precise parameter estimates, at least when confronted with simplistic simulated data containing spatial autocorrelation in the errors. However, we found that for presence/absence data the results and conclusions were very variable between the different methods. This is likely due to the low information content of binary maps. Also, in contrast with previous studies, we found that autocovariate methods consistently underestimated the effects of environmental controls of species distributions. Given their widespread use, in particular for the modelling of species presence/absence data (e.g. climate envelope models), we argue that this warrants further study and caution in their use. To aid other ecologists in making use of the methods described, code to implement them in freely available software is provided in an electronic appendix.

Increasing housing density is an important component of global land transformation, with major impacts on patterns of biodiversity. However, while there have been many studies of the changes in biodiversity across rural–urban gradients, which are influenced in large part by housing densities, how biodiversity changes across the full range of regional variation in housing density remains poorly understood. Here, we explore these relationships for the richness and abundance of breeding birds across Britain. Total richness, and that of 27 urban indicator species, increased from low to moderate household densities and then declined at greater household densities. The richness of all species increased initially faster with household density than did that of the urban indicator species, but nonurban indicator species richness declined consistently after peaking at a very low housing density. Avian abundance showed a rather different pattern. Total abundance and that summed across all urban indicator species increased over a wide range of household densities, and declined only at the highest household densities. The abundance of individual urban indicator species generally exhibited a hump-shaped relationship with housing density. While there was marked intraspecific variation in the form of such relationships, almost invariably avian abundance declined at housing densities below that at which the UK government requires new developments to be built. Our data highlight the difficulties of maintaining biodiversity while minimising land take for new development. High-density housing developments are associated with declines in many of those species otherwise best able to exploit urban environments, and those components of native biodiversity with which human populations are often most familiar.

Aim Spatial patterns of phylogenetic diversity (PD) aid our ability to discern diversification rate mechanisms underlying hypotheses for the large-scale distribution of biodiversity. We develop a predictive framework for the way in which spatial patterns of PD vary with those of species richness, depending on the balance between speciation and extinction rates. Within this framework, diversification processes thought to underlie the productive energy, ambient energy, topographic variability and habitat variety hypotheses predict that gradients of increase in species richness will be associated with: (1) decreasing extinction rates where driven by productive energy, hence increasing relative PD (i.e. PD controlling for species richness, or PDrel); (2) a similar positive relationship between ambient energy and PDrel; (3) increasing speciation rates where driven by topographic variability, hence decreasing PDrel; and (4) no consistent relationship between PDrel and habitat variety when driven by the latter. We test these predictions using distributional data on parrots. Location Neotropical, Afrotropical, Indo-Malayan and Australasian realms. Methods Spatial models were used to test the predictions. Results Globally, a positive association between productive energy and PDrel confirms prediction (1). However, within realms, hump-shaped relationships suggest the importance of decreasing extinction rates up to a threshold level of productive energy, and the increasing importance of speciation rates thereafter. Ambient energy is positively associated with PDrel in Australasia, Indo-Malaya, and globally, supporting prediction (2). However, this is driven by the coincidence of highest PDrel in areas of high ambient energy and intermediate productive energy (i.e. in seasonal tropical environments), which may be characterized by relatively low speciation and extinction rates. In the Neotropics, increasing topographic variability is associated with decreasing PDrel and increasing species richness, suggesting an increasing gradient of speciation, supporting prediction (3). Elsewhere, the signal of this mechanism may be obscured by collinearities with energy gradients. The lack of an overall relationship between habitat diversity and PDrel confirms prediction (4). Main conclusions Spatial patterns of PDrel in relation to environmental gradients may be sensitive to collinearities among those gradients. Nevertheless, patterns emerge which have implications for the relative importance of speciation and extinction processes in generating latitudinal diversity gradients.

Anthropogenic modification of habitats may reduce the resources available for native species, leading to population declines and extinction. These same habitats often have the highest richness of non-native species. This pattern may be explained if recently human-modified habitats provide novel resources that are more accessible to non-native species than native species. Using non-native birds in the Iberian Peninsula as a case study, we conduct a large-scale study to investigate whether non-native species are positively associated with human modified habitats, and to investigate whether this positive association may be driven by the presence of resources that are not fully exploited by native species. We do this by comparing the functional diversity and resource use of native and non-native bird communities in a recently human-modified habitat (rice fields) and in more traditional habitats in the Iberian Peninsula. The functional diversity of native bird communities was lower in rice fields, but non-native birds were positively associated with rice fields and plugged this gap. Differences in resource use between native and non-native species allowed non-native species to exploit resources that were plentiful in rice fields, supporting the role of underexploited resources in driving the positive association of non-native birds with rice fields. Our results provide a potential mechanism explaining the positive association of non-native species with anthropogenic habitats, and further work is needed to test if this applies more generally.

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests., Invertebrates are key components in the ecological functioning of tropical forests. Here, Ewers et al. show that, compared to primary forest, logging halves the contribution of invertebrate species to several key ecosystem processes, including litter decomposition.

Spatial patterns of species richness follow climatic and environmental variation, but could reflect random dynamics of species ranges (the mid-domain effect, MDE). Using data on the global distribution of birds, we compared predictions based on energy availability (actual evapotranspiration, AET, the best single correlate of avian richness) with those of range dynamics models. MDE operating within the global terrestrial area provides a poor prediction of richness variation, but if it operates separately within traditional biogeographic realms, it explains more global variation in richness than AET. The best predictions, however, are given by a model of global range dynamics modulated by AET, such that the probability of a range spreading into an area is proportional to its AET. This model also accurately predicts the latitudinal variation in species richness and variation of species richness both within and between realms, thus representing a compelling mechanism for the major trends in global biodiversity.

Aims To determine the shape of global species–range size distributions, the influence on these of island species, threatened species and patterns of latitudinal variation in range sizes, and the fit to logit-normal distributions. Location Global. Methods We take the spatial distributions of the raptors and owls of the world as exemplar data sets, document the shapes of their species–range size distributions, the influence of particular groups of species and of latitudinal variation in range sizes on these shapes, and the fit of these distributions to a variety of models. Results The global species–range size distributions of both raptors and owls are extremely right skewed on untransformed axes. They are not lognormally distributed, as has commonly been stated for species–range size distributions, nor logit-normally distributed as has been suggested might be the case. For raptors, departures from either a lognormal or a logit-normal are little mitigated by excluding groups of species that might be thought to distort the observed species–range size distribution, nor by the latitudinal gradient in geographical range size. For owls, the effects of excluding island and threatened species are more marked, with the fit of the species–range size distribution to a lognormal or a logit-normal becoming much closer. Conclusions A simple general description of the shape of species–range size distributions remains elusive. This constitutes a significant constraint on the development of theory as to how they are determined. Whilst in principle the fit of any given mechanistic model can be tested against one or more empirical data sets, whatever their form, a simple general mathematical description of species–range size distributions would make the process of rapidly testing the appropriateness of mechanistic models more straightforward.

Patterns of species composition in a neotropical termite assemblage were studied in relation to early effects of forest fragmentation as well as other sources of environmental and spatial heterogeneity. Termite diversity surveys were carried out at three mainland forest sites, and 10 islands of varying size, in an area of lowland tropical forest that had been flooded 4 y earlier, during the creation of the Petit Saut hydroelectric project in French Guiana. The ‘ghost forest’ of dead emergent trees in the flooded zone was also studied for its influence on island termite assemblages. Results suggested that the effects of forest fragmentation upon the total assemblage, and upon soil-feeders in particular, were subordinate to the influence of forest understorey palm density, and the closely associated gradients of soil humus depth and soil pH. Moreover, gradients for these three variables were uncorrelated with forest fragmentation and probably reflected spatial environmental heterogeneity pre-dating inundation events. Nevertheless, factors associated with forest fragmentation appeared to have had a significant effect on changes in termite species composition across the study site, primarily in structuring the wood- and leaf-litter-feeder assemblage. Effects upon the latter were not apparently a result of influx of species from the ghost forest. Purely spatial variation also influenced β-diversity changes in species composition across the site. In conclusion, the effects of forest fragmentation upon termites appear to have been relatively mild compared with other faunal groups, 4 y after flooding. Nevertheless, we predict that the effects of fragmentation on termite assemblages will ultimately be negative. This study also points to the importance of Amazonian understorey palms in structuring a tropical forest termite assemblage.

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to-date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty-seven standardized local termite diversity transects from twenty-nine locations across five tropical regions. Local-scale, intermediate-scale and large-scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south-east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large-scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate-scale geographical and site variables and, finally, local-scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long-term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter-regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.

Biomass collapse and its associated microclimatic stresses within recently isolated rain forest fragments may negatively affect species diversity of most resident taxa. However, for some decomposer organisms, increased resource availability via accompanying tree die-off may effect positive responses, at least for a time, with implications for rates of nutrient cycling and greenhouse gas release. This study investigates the early effects of forest fragmentation on a Neotropical termite assemblage. Numbers of encounters (surrogate for relative abundance) and species richness of wood and leaf-litter feeders, soil feeders, and the whole assemblage, were studied across true forest islands and mainland sites at a hydroelectric reservoir in French Guiana. Results showed no overall effect of fragmentation on either total termite encounters or species richness. However, numbers of encounters and species richness of wood and leaf-litter feeders showed positive responses to forest fragmentation. By contrast, soil feeders showed a negative response for numbers of encounters and no significant effect for species richness. Environmental data suggest that increased tree die-off, and other edge effects associated with biomass collapse, were underway at the time of sampling. Resulting increase in resource availability may therefore explain the positive influence on wood and leaf-litter feeders. A possible decrease in predation pressure from ants with decrease in island size was not tested for, but was a likely effect of the flooded matrix habitat. Fragmentation effects on soil feeder encounters may be due to the energetic and microclimatic constraints of feeding lower down the humification gradient of termite food substrates, but were not sufficient to affect species richness. The patterns revealed suggest that rates of wood decomposition following tree die-off, and of soil nutrient cycling, under different rain forest fragmentation scenarios, merit further study.

In south-east Asia, during Quaternary glaciations, increased seasonality and sea level decreases of ~120 m caused considerable fragmentation of the rainforest. It is believed that most of the region's core was covered with savannah vegetation, and rainforest was confined to a few refugia. We investigated the geographical extent of the savannah vegetation and the positions of the refugia, using a combination of palynonlogical, geological, fossil and termite data. Our results show that during the Last Glacial Maximum, most of Thailand, Peninsula Malaysia, western and southern Borneo, eastern and southern Sumatra, and Java were probably covered by savannah. Rainforest refugia were probably present in northern and eastern Borneo, northern and western Sumatra and the Mentawai islands. We also discuss a possible complete history of the region. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 453–466.

The exchange of CH4 between tropical forests and the atmosphere was determined by simultaneously measuring the net CH4 flux at the soil surface and assessing the flux contribution from soil-feeding termite biomass, both within the soil profile and in mounds. In Cameroon the flux of CH4 ranged from a net emission of 40.7 ng m–2 s–1 to a net CH4 oxidation of –53.0 ng m–2 s–1. Soil-inhabiting termite biomass was significantly correlated with CH4 flux. Termite mounds emitted up to 2000 ng s–1 mound–1. Termite-derived CH4 emission reduced the soil sink strength by up to 28%. Disturbance also had a strong effect on the soil sink strength, with the average rate of CH4 oxidation, at – 17.5 ng m–2 s–1, being significantly smaller (≈ 36%) at the secondary forest site than the –27.2 ng m–2 s–1, observed at the primary forest site. CH4 budgets calculated for each site indicated that both forests were net sinks for CH4 at – 6.1 kg ha–1 y–1 in the near-primary forest and – 3.1 kg ha–1 y–1 in the secondary forest. In Borneo, three forest sites representing a disturbance gradient were examined. CH4 oxidation rates ranged from 0 to – 32.1 ng m–2s–1 and a significant correlation between the net flux and termite biomass was observed only in an undisturbed primary forest, although the biomass was insufficient to cause net emission of CH4. Rates of CH4 oxidation were not significantly different across the disturbance gradient but were, however, larger in the primary forest (averaging – 15.4 ng m–2 s–1) than in an old-growth secondary forest (–13.9 ng m–2s–1) and a young secondary re-growth (– 10.8 ng m–2s–1). CH4 flux from termite mounds ranged from net oxidation in an abandoned mound to a maximum emission of 468 ng s–1 mound–1. CH4 budgets calculated for each site indicated that CH4 flux from termite mounds had an insignificant effect on the budget of CH4 at the regional scale at all three forest sites. Annual oxidation rates were – 4.8, – 4.2 and – 3.4 kg ha–1 y–1 in the primary, secondary and young secondary forests, respectively.

To characterize the structure and species density of termite assemblages in tropical forests, we developed a sampling protocol based on a single large quadrat (belt transect) that standardises sampling effort and area. Emphasizing the estimation of species richness, Roisin and Leponce (2004, Austral Ecology, 29: 637- 46) recommend sampling numerous small quadrats, and varying the number of quadrats depending on local species density. However, their method would result in the loss of standardisation and speed, taking up to four times longer to collect and process the samples. We show that even for a subtropical forest with low species density, our protocol produces robust results.

Many of the world's languages face serious risk of extinction. Efforts to prevent this cultural loss are severely constrained by a poor understanding of the geographical patterns and drivers of extinction risk. We quantify the global distribution of language extinction risk—represented by small range and speaker population sizes and rapid declines in the number of speakers—and identify the underlying environmental and socioeconomic drivers. We show that both small range and speaker population sizes are associated with rapid declines in speaker numbers, causing 25% of existing languages to be threatened based on criteria used for species. Language range and population sizes are small in tropical and arctic regions, particularly in areas with high rainfall, high topographic heterogeneity and/or rapidly growing human populations. By contrast, recent speaker declines have mainly occurred at high latitudes and are strongly linked to high economic growth. Threatened languages are numerous in the tropics, the Himalayas and northwestern North America. These results indicate that small-population languages remaining in economically developed regions are seriously threatened by continued speaker declines. However, risks of future language losses are especially high in the tropics and in the Himalayas, as these regions harbour many small-population languages and are undergoing rapid economic growth.

Fire is prevalent in the dry deciduous forests of south-east Asia, the majority of fires being man-made (Stott 1988). In Thailand, the 10-19% estimated extent of annual burning of dry deciduous dipterocarp forest (1984-1986 data on total estimated area burned each year; Settarak et al. 1987) is thought to be a severe under-estimate (Kanjanavanit 1992). The interaction of fire and termites is poorly known for Asian ecosystems, but African studies show they have complementary roles in the removal of wood and leaf litter (Collins 1981, Trapnell et al. 1976). Termites are the dominant arthropod decomposers in tropical forests (Collins 1983) and savannas (Collins 1981, Holt 1987), and are important mediators of carbon flux (Jones 1990, Lawton et al. 1996). Furthermore, these functions are dependent on the species assemblage structure of the termite community (Lawton et al. 1996). To date, the only study of the effects of fire on termite species richness and abundance is from Australia (Abensperg-Traun & Milewski 1995). Here, a preliminary survey of species richness of termites in fireprotected and fire-prone deciduous forest in northern Thailand is presented. The study suggests that the fungus-growing Macrotermitinae have a greater species richness, under certain seasonal fire regimes, compared to those in areas under complete fire protection. Species richness estimates were made for two dry deciduous dipterocarp forest sites: A, fire-protected and; B, unprotected, fire-prone, in Doi Suthep-Pui National Park, near Chiang Mai, northern Thailand. Site A covered a 10 ha

The effects of climate change on biodiversity should depend in part on climate displacement rate (climate-change velocity) and its interaction with species’ capacity to migrate. We estimated Late Quaternary glacial-interglacial climate-change velocity by integrating macroclimatic shifts since the Last Glacial Maximum with topoclimatic gradients. Globally, areas with high velocities were associated with marked absences of small-ranged amphibians, mammals, and birds. The association between endemism and velocity was weakest in the highly vagile birds and strongest in the weakly dispersing amphibians, linking dispersal ability to extinction risk due to climate change. High velocity was also associated with low endemism at regional scales, especially in wet and aseasonal regions. Overall, we show that low-velocity areas are essential refuges for Earth’s many small-ranged species.

Research over many decades has documented numerous environmental effects of urbanization, ranging from the loss and reconfiguration of green space to dramatic changes in ecosystems and biodiversity. Rather less is known about how urban form, in particular the density of urban development, alters environmental patterns and processes within cities. Investigation of the relationships between urban form and environmental structure and performance is an important issue in the urban sustainability debate and here we use that work to illustrate some of the key ideas in this newly emerging field. After outlining the general effects of urbanization on environment and biodiversity, we then consider in turn the relationships between urban form and patterns of green space, the degree to which urban environments can provide useful ecosystem services to human populations, and finally the responses of biodiversity to urban development.

In 1847, Karl Bergmann proposed that temperature gradients are the key to understanding geographic variation in the body sizes of warm-blooded animals. Yet both the geographic patterns of body-size variation and their underlying mechanisms remain controversial. Here, we conduct the first assemblage-level global examination of 'Bergmann's rule' within an entire animal class. We generate global maps of avian body size and demonstrate a general pattern of larger body sizes at high latitudes, conforming to Bergmann's rule. We also show, however, that median body size within assemblages is systematically large on islands and small in species-rich areas. Similarly, while spatial models show that temperature is the single strongest environmental correlate of body size, there are secondary correlations with resource availability and a strong pattern of decreasing body size with increasing species richness. Finally, our results suggest that geographic patterns of body size are caused both by adaptation within lineages, as invoked by Bergmann, and by taxonomic turnover among lineages. Taken together, these results indicate that while Bergmann's prediction based on physiological scaling is remarkably accurate, it is far from the full picture. Global patterns of body size in avian assemblages are driven by interactions between the physiological demands of the environment, resource availability, species richness and taxonomic turnover among lineages.

There can now be no doubt of the intellectual substance and cohesion of early Wyclifitism as expressed in the writings of educated clerks in immediate contact with the man himself. Most would accept, too, that this coherence was successfully transferred from Latin to English. However, although these Wyclifite scholars recognised the need for a corpus of literature to cater for a non-academic audience and provide the basis in ideas for a sustained movement, they had difficulty in supplying it. This might seem to offer easy comfort to those who are already doubtful whether people called Lollards could or did grasp Wyclifitism; whether they just amputated the bits they liked, debased or perverted them, or really did not take anything on board at all. Even some less cynical would agree that the Church itself played a large part in shaping Lollardy's ideas and characteristics: such is so often the way in the birth of protest movements. Indeed, to some hardliners ‘Lollardy’ seems little more than a scare-story invented by the Church in order to damn a large but motley crew of critics and dissenters. Or, if not the Church, then those historians in whom hope triumphs over experience.

Despite its wide implications for many ecological issues, the global pattern of spatial turnover in the occurrence of species has been little studied, unlike the global pattern of species richness. Here, using a database on the breeding distributions of birds, we present the first global maps of variation in spatial turnover for an entire taxonomic class, a pattern that has to date remained largely a matter of conjecture, based on theoretical expectations and extrapolation of inconsistent patterns from different biogeographic realms. We use these maps to test four predictions from niche theory as to the form that this variation should take, namely that turnover should increase with species richness, towards lower latitudes, and with the steepness of environmental gradients and that variation in turnover is determined principally by rare (restricted) species. Contrary to prediction, we show that turnover is high both in areas of extremely low and high species richness, does not increase strongly towards the tropics, and is related both to average environmental conditions and spatial variation in those conditions. These results are closely associated with a further important and novel finding, namely that global patterns of spatial turnover are driven principally by widespread species rather than the restricted ones. This complements recent demonstrations that spatial patterns of species richness are also driven principally by widespread species, and thus provides an important contribution towards a unified model of how terrestrial biodiversity varies both within and between the Earth's major land masses.

Theory predicts that biogeographic factors should play a central role in promoting population divergence and speciation. Previous empirical studies into biogeography and diversification have been relatively restricted in terms of the geographical area, phylogenetic scope, and the range of biogeographic factors considered. Here we present a global analysis of allopatric phenotypic divergence (measured as subspecies richness) across more than 9600 bird species. The main aim of this study was to examine the extent to which biogeographical factors can explain patterns of phenotypic divergence. Analysis of the taxonomic distribution of subspecies among species suggests that subspecies formation and extinction have occurred at a considerably faster rate than has species formation. However, the observed distribution departs from the expectation under a random birth-death model of diversification. Across 19 phylogenetic trees, we find no significant linear relationship between species age and subspecies richness, implying that species age is a poor predictor of subspecies richness. Both subspecies richness and subspecies diversification rate are found to exhibit low phylogenetic signal, meaning that closely related species do not tend to possess similar numbers of subspecies. As predicted by theory, high subspecies richness was associated with large breeding range size, island dwelling, inhabitation of montane regions, habitat heterogeneity, and low latitude. Of these factors, breeding range size was the variable that explained the most variation. Unravelling whether species that have invaded previously glacial areas have more or fewer subspecies than expected proves to be complicated due to a covariation between the postglacial colonization, latitude, geographic range size, and subspecies richness. However, the effect of postglacial colonization on subspecies richness appears to be small. Mapping the distribution of species' subspecies richness globally reveals geographical patterns that correspond to many of the predictions of the statistical models, but may also reflect geographical variation in taxonomic practice. Overall, we demonstrate that biogeographic models can explain about 30% of the global variation in subspecies richness in birds.

A major goal of ecology is to determine the causes of the latitudinal gradient in global distribution of species richness. Current evidence points to either energy availability or habitat heterogeneity as the most likely environmental drivers in terrestrial systems, but their relative importance is controversial in the absence of analyses of global (rather than continental or regional) extent. Here we use data on the global distribution of extant continental and continental island bird species to test the explanatory power of energy availability and habitat heterogeneity while simultaneously addressing issues of spatial resolution, spatial autocorrelation, geometric constraints upon species' range dynamics, and the impact of human populations and historical glacial ice-cover. At the finest resolution (1°), topographical variability and temperature are identified as the most important global predictors of avian species richness in multi-predictor models. Topographical variability is most important in single-predictor models, followed by productive energy. Adjusting for null expectations based on geometric constraints on species richness improves overall model fit but has negligible impact on tests of environmental predictors. Conclusions concerning the relative importance of environmental predictors of species richness cannot be extrapolated from one biogeographic realm to others or the globe. Rather a global perspective confirms the primary importance of mountain ranges in high-energy areas.