Species distribution modelling as a macroecological tool: a case study using New World amphibians

Although species distribution modelling (SDM) is widely accepted among the scientific community and is increasingly used in ecology, conservation biology and biogeography, methodological limitations generate potential problems for its application in macroecology. Using amphibian species richness in North and South America, we compare species richness patterns derived from SDM maps and ‘expert’ maps to evaluate if: 1) richness patterns derived from SDM are biased toward climate‐based explanations for diversity when compared to expert maps, since SDM methods are typically based on climatic variables; and 2) SDM is a reliable tool for generating richness maps in hyperrich regions where point occurrence data are limited for many species. We found that although three widely used SDM methods overestimated amphibian species richness in grid cells when compared to expert richness maps in both North and South America due to systematic overestimation of range sizes, diversity gradients were reasonably robust at broad scales. Further, climatic variables statistically explained patterns of richness at similar levels among the different richness sources, although climatic relationships were stronger in the much better known North America than in South America. We conclude that in the face of the high deforestation rates coupled with incomplete data on species distributions, especially in the tropics, SDM represents a useful macroecological tool for investigating broad‐scale richness patterns and the dynamics between species richness and climate.     

Type and spatial structure of distribution data and the perceived determinants of geographical gradients in ecology: the species richness of African birds

Aim Studies exploring the determinants of geographical gradients in the occurrence of species or their traits obtain data by: (1) overlaying species range maps; (2) mapping survey‐based species counts; or (3) superimposing models of individual species’ distributions. These data types have different spatial characteristics. We investigated whether these differences influence conclusions regarding postulated determinants of species richness patterns. Location Our study examined terrestrial bird diversity patterns in 13 nations of southern and eastern Africa, spanning temperate to tropical climates. Methods Four species richness maps were compiled based on range maps, field‐derived bird atlas data, logistic and autologistic distribution models. Ordinary and spatial regression models served to examine how well each of five hypotheses predicted patterns in each map. These hypotheses propose productivity, temperature, the heat–water balance, habitat heterogeneity and climatic stability as the predominant determinants of species richness. Results The four richness maps portrayed broadly similar geographical patterns but, due to the nature of underlying data types, exhibited marked differences in spatial autocorrelation structure. These differences in spatial structure emerged as important in determining which hypothesis appeared most capable of explaining each map's patterns. This was true even when regressions accounted for spurious effects of spatial autocorrelation. Each richness map, therefore, identified a different hypothesis as the most likely cause of broad‐scale gradients in species diversity. Main conclusions Because the ‘true’ spatial structure of species richness patterns remains elusive, firm conclusions regarding their underlying environmental drivers remain difficult. More broadly, our findings suggest that care should be taken to interpret putative determinants of large‐scale ecological gradients in light of the type and spatial characteristics of the underlying data. Indeed, closer scrutiny of these underlying data — here the distributions of individual species — and their environmental associations may offer important insights into the ultimate causes of observed broad‐scale patterns.     

Range maps and species richness patterns: errors of commission and estimates of uncertainty

Range maps are often combined into “range overlap maps” to estimate spatial variation in species richness. Range maps are, in most cases, designed to represent a species’ maximum geographical extent and not patterns of occupancy within the range. As a consequence, range maps overestimate occupancy by presenting false occupancy (errors of commission) within the interior of the range. To assess the implications of errors of commission when developing and applying range overlap maps, we used neutral landscapes to simulate range maps and patterns of occupancy within ranges. We explored several scenarios based on combinations of six parameters defining biogeographical and cartographic factors typically encountered by investigators. Our results suggest that, in general, uncertainty is lowest when map resolutions are moderately fine, the majority of species have geographically restricted ranges, species occur throughout their range, patterns of occupancy within the range are not correlated among species, and geographically local and widespread species tend to occupy different regions. Several of these outcomes are associated with broad geographical extents, the scale at which range overlap maps are typically applied. Thus, under most circumstances, reasonably accurate and precise representation of species richness patterns can be achieved. However, these representations can be improved by enhancing occupancy data for widespread species – a primary source of uncertainty – and selecting a map resolution that captures relevant biological and environmental heterogeneity. Hence, by determining where a study is situated within the scenarios examined in our simulations, uncertainty and its sources and implications can be ascertained. With this knowledge, research goals, methods, and data sources can be reassessed and refined and, in the end, conclusions and recommendations can be qualified. Alternatively, unique regional, taxonomic, or cartographic factors could be included in future simulations to provide direct assessments of uncertainty.     

Relationships of climate,residence time,and biogeographical origin with the range sizes and species richness patterns of exotic plants in Great Britain

Based on atlas data with a 10-km cell resolution for 1,406 exotic plant species inhabiting Great Britain, we investigate the extent to which arrival time (residence time) and biogeographical origin (climate suitability) are associated with range sizes of exotic plants and how exotic plant richness is related to current climate and the human footprint. We grouped species according to four arrival periods (archaeophytes and three classes of neophytes), and three broad biogeographical origins, each reflecting a different macroclimate similarity with the study region (northern Holarctic > Mediterranean > and tropical–subtropical). While we found that mean range sizes increased with residence time, no strong effect of the region of origin on range size was detected. Also, across all groups, species richness was primarily and positively associated with temperature, whereas relationships with human footprint were much weaker, albeit also positive in all cases. The proportion of variance explained by environmental models of richness increased from groups comprising recently arrived species to those that arrived earlier, and from tropical–subtropical species to exotics coming from the Holarctic. Our data also illustrate how these trends translate into richness patterns and their association with climate, which become more similar to native richness patterns as residence time and macroclimatic matching increase. In contrast, broad-scale human alteration of ecosystems appeared to be less important for variation in exotic richness than climate, although we did not evaluate anthropogenic effects at finer scales.     

Rodent endemism,turnover and biogeographical transitions on elevation gradients in the northwestern Argentinian Andes

The aim of this research is to relate patterns of endemism and turnover along a local elevation gradient in northwest Argentina with continental biogeographical transitions. Specimen based records constituted the principal source of information to infer rodent distribution along the elevation gradient. I assessed elevational variation of richness, endemism and turnover by means of non-linear regression analysis. Then I identified five distributional patterns based on the overlap of species geographic range. Their frequency along elevation was used to validate biogeographical boundaries inferred by turnover rates. Eleven species out of 37 (30%) are endemic to the study area. Species richness and endemism were hump-shaped. The rate of endemism reached its maximum value at the upper limit of the forest (2500 m). By contrast, species turnover was U-shaped, with a small peak at 1500 m and a maximum at 3500 m. The species’ geographic range patterns were not randomly distributed along elevation but agglomerated at specific elevation. Species turnover and chorological analysis suggest two biogeographical boundaries, a weaker at 1500 m and a stronger at 3500 m. The 1500 m boundary marks the transition from assemblages dominated by Lowland-widespread fauna at lower elevation to Montane (Andean eastern slopes) species at middle elevation. This boundary is characterized by moderate species turnover and high species richness. The strong turnover rate at 3500 and the dominance of highland Andean and Andean-Patagonian species above this elevation suggest the occurrence of the transition between the Neotropical and Andean regions; which is characterised by an almost complete species replacement.     

The positive correlation between avian species richness and human population density in Britain is not attributable to sampling bias

Aim  To assess whether spatial variation in sampling effort drives positive correlations between human population density and species richness.
Location  British 10 × 10 km squares.
Methods  We calculated three measures of species richness from atlas data of breeding birds in Britain: total species richness, species richness standardised for sampling effort, and the number of species only recorded in supplementary casual records in a manner not standardised for survey effort. We then assessed the form of the relationship between these richness estimates and human population density, both with and without taking spatial autocorrelation into account.
Results  Both total and standardised species richness exhibit similar species richness–human population density relationships; species richness generally increases with human population density, but decreases at the very highest densities. Supplementary species richness is very weakly correlated with human population density.
Main conclusions  In this example, sampling effort only slightly influences the form of species richness–human population density relationships. The positive correlation between species richness and human population density and any resultant conservation conflicts are thus not artefactual patterns generated by confounding human density and sampling effort.     

Climate and landscape explain richness patterns depending on the type of species’ distribution data

Understanding the patterns of species richness and their environmental drivers, remains a central theme in ecological research and especially in the continental scales where many conservation decisions are made. Here, we analyzed the patterns of species richness from amphibians, reptiles and mammals at the EU level. We used two different data sources for each taxon: expert-drawn species range maps, and presence/absence atlases. As environmental drivers, we considered climate and land cover. Land cover is increasingly the focus of research, but there still is no consensus on how to classify land cover to distinct habitat classes, so we analyzed the CORINE land cover data with three different levels of thematic resolution (resolution of classification scheme ˗ less to more detailed). We found that the two types of species richness data explored in this study yielded different richness maps. Although, we expected expert-drawn range based estimates of species richness to exceed those from atlas data (due to the assumption that species are present in all locations throughout their region), we found that in many cases the opposite is true (the extreme case is the reptiles where more than half of the atlas based estimates were greater than the expert-drawn range based estimates). Also, we detected contrasting information on the richness drivers of biodiversity patterns depending on the dataset used. For atlas based richness estimates, landscape attributes played more important role than climate while for expert-drawn range based richness estimates climatic variables were more important (for the ectothermic amphibians and reptiles). Finally we found that the thematic resolution of the land cover classification scheme, also played a role in quantifying the effect of land cover diversity, with more detailed thematic resolution increasing the relative contribution of landscape attributes in predicting species richness.     

Quantification of climatic niches in birds: adding the temporal dimension

Quantification of the climatic niche from geographic occurrences is an increasingly important tool for studying species’ relationships to their environment, for example to predict responses to climate change. However, as the geographic distributions of birds are seasonally dynamic, they pose a challenge to carrying out comparable and appropriate quantification of climatic niches. In this review, we first assess how relevant seasonal dynamics are across birds as a whole by compiling a database of migratory behaviour for 10 443 bird species. Second, we examine how studies have quantified climatic niches of birds. Finally, using Australia as a case study, we investigate how well existing distribution datasets represent temporal dynamics by comparing seasonal patterns of species richness obtained from point‐occurrence data with those from range maps and assess the consequences for niche quantification. We provide a consistent classification of migratory behaviour across all birds, and find that a huge variety exists between and within species that should be considered when quantifying climatic niches. Despite this, our review of the literature revealed that seasonal dynamics have often not been accounted for. For future studies, we provide a framework for selecting appropriate occurrence data depending on migratory behaviour and data availability. Our comparison of seasonal species richness patterns obtained from extent‐of‐occurrence range maps and point‐occurrence data suggests that range maps are less able to detect temporal dynamics of bird distributions than point‐occurrence data. We conclude that seasonally explicit range maps combined with climatic data for the corresponding time period can be used to adequately quantify climatic niches for resident birds, but are not adequate to quantify the climatic niches of migratory and nomadic species. Therefore, consistent quantification of climatic niches across all birds requires temporally explicit occurrence points. As such, increasing the availability of occurrence data and methods correcting biases should be a priority.     

Effects of historical and contemporary factors on global patterns in avian species richness

Aim  The aim of this study was to determine how regional and historical factors influence global patterns in avian species richness.
Location  Global.
Methods  Using a comprehensive data set including 710 World Wildlife Fund terrestrial ecoregions covering nearly all the land surface of the Earth, avian species richness was compared among six biogeographical regions after accounting for sample area, elevational range and climate. Analysis of variance and multiple regressions were used. Spatial autocorrelation in model residuals was accounted for.
Results  Significant effects of region on avian species richness were found in nearly all comparisons between biogeographical regions.
Main conclusions  Regional and historical processes have played a role in regulating large-scale avian species richness patterns across the globe. Avian species richness in different regions of the world cannot be accurately predicted by a single global model. Avian species richness in areas of similar environments may differ substantially between regions, and thus avian species richness in one biogeographical region cannot be predicted using the richness–environment relationship derived from the data of another biogeographical region, even one with similar environments.     

Biogeography and the structure of coral reef fish communities on isolated islands

Aim To determine the applicability of biogeographical and ecological theory to marine species at two remote island locations. This study examines how biogeography, isolation and species geographic range size influence patterns of species richness, endemism, species composition and the abundance of coral reef fishes. Location Christmas Island and the Cocos (Keeling) Islands in the tropical eastern Indian Ocean. Methods Published species lists and underwater visual surveys were used to determine species richness, endemism, species composition and abundance of reef fishes at the islands. These data were statistically compared with patterns of species composition and abundance from the neighbouring ‘mainland’ Indonesian region. Results The two isolated reef fish communities were species‐poor and contained a distinct taxonomic composition with an overrepresentation of species with high dispersal potential. Despite low species richness, we found no evidence of density compensation, with population densities on the islands similar to those of species‐rich mainland assemblages. The mix of Indian and Pacific Ocean species and the proportional representations of the various regional faunas in the assemblages were not influenced by the relative proximity of the islands to different biogeographical provinces. Moreover, species at the edge of their range did not have a lower abundance than species at the centre of their range, and endemic species had substantially higher abundances than widespread species. At both locations, endemism was low (less than 1.2% of the community); this may be because the locations are not sufficiently isolated or old enough to promote the evolution of endemic species. Main conclusions The patterns observed generally conform to terrestrial biogeographical theory, suggesting that similar processes may be influencing species richness and community composition in reef fish communities at these remote islands. However, species abundances differed from typical terrestrial patterns, and this may be because of the life history of reef fishes and the processes maintaining isolated populations.     

Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

The influence of spatial scale on cross-taxon congruence patterns and prediction accuracy of species richness

A comparison of species richness patterns of butterflies and birds was made using data from two grids of squares (small squares 137.5 km on a side and large squares 275 km on a side) covering western North America. Using geostatistical procedures, we found that the spatial patterns of species richness of these two taxa were related. The influence of grain size on the strength of this relationship was investigated by analysing the two data sets. For both data sets, the number of butterfly species in a square was a statistically significant predictor of the corresponding number of bird species. However, cross-validation techniques showed that the marginal improvement in prediction accuracy due to including butterflies as a predictor was greater in the large-square data. We explored the effect of areal extent on cross-taxon congruencies by investigating species richness patterns in four subsets of the small-square data. In regions with smaller areal extent, the cross-taxon congruence patterns were not substantially different from the pattern found in the full data set. Finally, using data-splitting techniques, we explored the relationships between prediction accuracy of species richness, sample size, areal extent of the sample, and grain size.     

Spatial congruence between biotic history and species richness of Muscidae (Diptera,Insecta) in the Andean and Neotropical regions

Considering that Earth and life evolve together, the present study aims to verify whether the species richness patterns are spatially congruent to biotic history. Niche conservatism was adopted as a background hypothesis to associate species richness with phylogenetic information. A parallel analysis between this procedure and cladistic biogeography was undertaken. Eleven Muscidae genera that were previously systematically reviewed for phylogenetic hypotheses were chosen for the analysis. The genera were split into ‘basal’ and ‘derived’ species, following terminal taxon root distance within each genus. Richness patterns were contrasted for the most basal and most derived 33% of species, and richness maps were constructed at 220 × 220 km grid size. A difference richness map was drawn by derived minus basal values (=derived?basal). For regions with difference values around zero, a component analysis was performed and compared with relationships established by other studies. Derived and basal species richness showed a very concise richness gradient in the Neotropical region and it was compatible with its known biogeographical history. In the Andean region, richness did not show any pattern. The area cladogram grouped Subantarctic subregion in a polytomy and Central Chile as a paraphyletic group. All hypotheses about area relationship were divergent and no vicariant pattern could be recognized in Andean region. In Neotropical region, Muscidae results corroborated a previous component relationship. The hypothesis that Paleogene climatic changes could drive the biotic component’s split was suggested. In the Andean region, recently ice sheet covering events had driven the species to disperse and/or extinct resulted in absence of pattern seen either in richness analysis or in component analysis. It is believed that species richness is linked to biotic history and this fact may be considered when evaluating hypotheses to explain broad‐scale richness gradients.     

Spatially-Explicit Estimation of Geographical Representation in Large-Scale Species Distribution Datasets

Much ecological research relies on existing multispecies distribution datasets. Such datasets, however, can vary considerably in quality, extent, resolution or taxonomic coverage. We provide a framework for a spatially-explicit evaluation of geographical representation within large-scale species distribution datasets, using the comparison of an occurrence atlas with a range atlas dataset as a working example. Specifically, we compared occurrence maps for 3773 taxa from the widely-used Atlas Florae Europaeae (AFE) with digitised range maps for 2049 taxa of the lesser-known Atlas of North European Vascular Plants. We calculated the level of agreement at a 50-km spatial resolution using average latitudinal and longitudinal species range, and area of occupancy. Agreement in species distribution was calculated and mapped using Jaccard similarity index and a reduced major axis (RMA) regression analysis of species richness between the entire atlases (5221 taxa in total) and between co-occurring species (601 taxa). We found no difference in distribution ranges or in the area of occupancy frequency distribution, indicating that atlases were sufficiently overlapping for a valid comparison. The similarity index map showed high levels of agreement for central, western, and northern Europe. The RMA regression confirmed that geographical representation of AFE was low in areas with a sparse data recording history (e.g., Russia, Belarus and the Ukraine). For co-occurring species in south-eastern Europe, however, the Atlas of North European Vascular Plants showed remarkably higher richness estimations. Geographical representation of atlas data can be much more heterogeneous than often assumed. Level of agreement between datasets can be used to evaluate geographical representation within datasets. Merging atlases into a single dataset is worthwhile in spite of methodological differences, and helps to fill gaps in our knowledge of species distribution ranges. Species distribution dataset mergers, such as the one exemplified here, can serve as a baseline towards comprehensive species distribution datasets.     

Water–energy balance and the geographic pattern of species richness of western Palearctic butterflies

Abstract. 1. Using two sources of data to estimate butterfly species richness, the potential influences of 11 environmental variables on the richness gradient of butterflies in western/central Europe and northern Africa were examined with multiple regression and spatial autocorrelation analysis. A measure of water–energy balance, actual evapotranspiration, explained 79% of the variance in butterfly species richness using data derived from range maps, and 72% of the variance using data derived from grid‐based distribution maps. All other variables explained less than 4% of the variance in the regression models and differed depending on the data source. 2. The spatial analysis indicated that actual evapotranspiration successfully removed most of the spatial autocorrelation in both richness data sets at all spatial scales, confirming the ability of the model to account for the spatial pattern in butterfly richness. 3. Plant species richness, a rarely tested variable hypothesised to be an important determinant of herbivore diversity, was weakly associated with butterfly richness, suggesting that it has little or no direct influence on butterfly richness. 4. A historical variable, the length of time that areas have been exposed for recolonisation after the retreat of the ice sheet following the last ice age, was also not associated with richness patterns, indicating that butterfly richness is in equilibrium with contemporary climate. 5. It was not possible to confirm a result reported for Canadian butterflies that land cover diversity is a strong predictor of butterfly richness, possibly because of methodological differences in the studies, differences in the range of climates found in Canada and the western Palearctic, or because of the highly modified landscape characteristic of Europe. 6. Water–energy balance offers a parsimonious explanation for the butterfly richness gradient in this region, operating partially indirectly via effects on plant productivity and partially directly via physiological effects on butterflies, and this conclusion is robust to differences in the types of distribution maps used to estimate richness patterns.     

Determinants of regional species richness: an empirical analysis of the number of hawkmoth species (Lepidoptera: Sphingidae) on the Malesian archipelago

Aims Major patterns and determinants of the species richness of Sphingidae in the Malesian archipelago were investigated, including a distinction of richness patterns between subfamilies and range‐size classes. Location Southeast Asia, Malesia. Methods Using a compilation of specimen‐label data bases, geographic information system (GIS)‐supported estimates of distributional ranges for all Sphingidae species of Southeast Asia were used to assess the species richness of islands. Range maps for all species and checklists for 114 islands can be found at http://www.sphingidae‐sea.biozentrum.uni‐wuerzburg.de . Potential determinants of the species richness of islands were tested with general linear models. Results The estimated species richness of islands in the region is determined by biogeographical association, seasonality, availability of rain forest and island size. Species–area relationships are linear on a semi‐logarithmic representation, but not on a double‐logarithmic scale. Species richness of all sphingid subfamilies is influenced by biogeography. The presence of large rain‐forest areas affects mainly Smerinthinae, whereas distance from continental Asia is conspicuously irrelevant for this group. Widespread rather than geographically restricted species shape the overall distribution patterns of species richness. The altitudinal range of islands does not significantly affect species‐richness patterns, but its potential effects on geographically restricted species are discussed. Main conclusions As well as being affected by climatic and vegetation parameters, sphingid species richness is strongly influenced by a historical, directional dispersal process from continental Southeast Asia to the Pacific islands. This process did not apply equally to species of different taxonomic groups or range sizes. Widespread species decline in species richness towards the south‐east, whereas geographically restricted species exhibit an inverse pattern of species richness, probably because speciation becomes more important in this group within the more isolated island groups.     

Midpoint attractors and species richness: Modelling the interaction between environmental drivers and geometric constraints

We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor – a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon‐specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.     

Connecting species’ geographical distributions to environmental variables: range maps versus observed points of occurrence

Connecting the geographical occurrence of a species with underlying environmental variables is fundamental for many analyses of life history evolution and for modeling species distributions for both basic and practical ends. However, raw distributional information comes principally in two forms: points of occurrence (specific geographical coordinates where a species has been observed), and expert-prepared range maps. Each form has potential short-comings: range maps tend to overestimate the true occurrence of a species, whereas occurrence points (because of their frequent non-random spatial distribution) tend to underestimate it. Whereas previous comparisons of the two forms have focused on how they may differ when estimating species richness, less attention has been paid to the extent to which the two forms actually differ in their representation of a species’ environmental associations. We assess such differences using the globally distributed avian order Galliformes (294 species). For each species we overlaid range maps obtained from IUCN and point-of-occurrence data obtained from GBIF on global maps of four climate variables and elevation. Over all species, the median difference in distribution centroids was 234 km, and median values of all five environmental variables were highly correlated, although there were a few species outliers for each variable. We also acquired species’ elevational distribution mid-points (mid-point between minimum and maximum elevational extent) from the literature; median elevations from point occurrences and ranges were consistently lower (median −420 m) than mid-points. We concluded that in most cases occurrence points were likely to produce better estimates of underlying environmental variables than range maps, although differences were often slight. We also concluded that elevational range mid-points were biased high, and that elevation distributions based on either points or range maps provided better estimates.     

Bird dietary guild richness across latitudes,environments and biogeographic regions

Aim To integrate dietary knowledge and species distributions in order to examine the latitudinal, environmental, and biogeographical variation in the species richness of avian dietary guilds (herbivores, granivores, frugivores, nectarivores, aerial insectivores, terrestrial/arboreal insectivores, carnivores, scavengers, and omnivores). Location Global. Methods We used global breeding range maps and a comprehensive dietary database of all terrestrial bird species to calculate guild species richness for grid cells at 110 × 110 km resolution. We assessed congruence of guild species richness, quantified the steepness of latitudinal gradients and examined the covariation between species richness and climate, topography, habitat diversity and biogeographic history. We evaluated the potential of current environment and biogeographic history to explain global guild distribution and compare observed richness–environment relationships with those derived from random subsets of the global species pool. Results While most guilds (except herbivores and scavengers) showed strong congruence with overall bird richness, covariation in richness between guilds varied markedly. Guilds exhibited different peaks in species richness in geographical and multivariate environmental space, and observed richness–environment relationships mostly differed from random expectations. Latitudinal gradients in species richness were steepest for terrestrial/arboreal insectivores, intermediate for frugivores, granivores and carnivores, and shallower for all other guilds. Actual evapotranspiration emerged as the strongest climatic predictor for frugivores and insectivores, seasonality for nectarivores, and temperature for herbivores and scavengers (with opposite direction of temperature effect). Differences in species richness between biogeographic regions were strongest for frugivores and nectarivores and were evident for nectarivores, omnivores and scavengers when present‐day environment was statistically controlled for. Guild richness–environment relationships also varied between regions. Main conclusions Global associations of bird species richness with environmental and biogeographic variables show pronounced differences between guilds. Geographic patterns of bird diversity might thus result from several processes including evolutionary innovations in dietary preferences and environmental constraints on the distribution and diversification of food resources.     

Biogeographical and anthropogenic determinants of landscape-scale patterns of raptors in West African savannas

Strong raptor population declines have recently been reported in sub-Saharan West Africa, where the pressure on wildlife and their supporting habitats is particularly high. This makes it imperative to understand the role of land-use on landscape-scale patterns of raptors and to define priority areas for conservation. We examine landscape-scale community patterns of raptors in biogeographical zones with different degrees of anthropogenic land-use and assess the role of protected areas in maintaining such patterns. We recorded raptors along road transects in Cameroon’s savannas, covering four years and 7,340–7,700 km in wet and dry seasons, in three biogeographical zones: the relatively well-preserved Inundation and Guinea zones to the north and south of the heavily exploited Sudan zone. The Inundation zone had the largest species pool and Palearctic raptor richness and abundance. The Guinea zone had the largest Afrotropical raptor species pool, while raptor diversity and richness were higher there than in the Sudan zone. The abundance of only one species (Fox Kestrel) peaked in the Sudan zone and only one large-bodied raptor (Hooded Vulture) with a Sudan-centered distribution was more common there than in the other zones. Our results suggest that land-use as determined by protected areas and human exploitation may override the role of biogeographical zonation in shaping raptor assemblages. Comparable patterns of raptor richness and diversity inside and outside protected areas suggest that both protected areas and partly cultivated peripheral zones act as important foraging and source areas, ensuring the preservation of diverse raptor assemblages at the landscape scale. Finally, our data illustrate the comparatively high richness of Cameroon’s and West Africa’s savanna raptor communities on a continental and global scale, underlining their importance for raptor conservation.