Richard G. Davies

Global hotspots of species richness are not congruent with endemism or threat

Biodiversity hotspots have a prominent role in conservation biology, but it remains controversial to what extent different types of hotspot are congruent. Previous studies were unable to provide a general answer because they used a single biodiversity index, were geographically restricted, compared areas of unequal size or did not quantitatively compare hotspot types. Here we use a new global database on the breeding distribution of all known extant bird species to test for congruence across three types of hotspot. We demonstrate that hotspots of species richness, threat and endemism do not show the same geographical distribution. Only 2.5% of hotspot areas are common to all three aspects of diversity, with over 80% of hotspots being idiosyncratic. More generally, there is a surprisingly low overall congruence of biodiversity indices, with any one index explaining less than 24% of variation in the other indices. These results suggest that, even within a single taxonomic class, different mechanisms are responsible for the origin and maintenance of different aspects of diversity. Consequently, the different types of hotspots also vary greatly in their utility as conservation tools.

Global distribution and conservation of rare and threatened vertebrates

Global conservation strategies commonly assume that different taxonomic groups show congruent geographical patterns of diversity, and that the distribution of extinction-prone species in one group can therefore act as a surrogate for vulnerable species in other groups when conservation decisions are being made. The validity of these assumptions remains unclear, however, because previous tests have been limited in both geographical and taxonomic extent. Here we use a database on the global distribution of 19,349 living bird, mammal and amphibian species to show that, although the distribution of overall species richness is very similar among these groups, congruence in the distribution of rare and threatened species is markedly lower. Congruence is especially low among the very rarest species. Cross-taxon congruence is also highly scale dependent, being particularly low at the finer spatial resolutions relevant to real protected areas. ‘Hotspots’ of rarity and threat are therefore largely non-overlapping across groups, as are areas chosen to maximize species complementarity. Overall, our results indicate that ‘silver-bullet’ conservation strategies alone will not deliver efficient conservation solutions. Instead, priority areas for biodiversity conservation must be based on high-resolution data from multiple taxa.

The Influence of Late Quaternary Climate-Change Velocity on Species Endemism

Regions with low glacial-interglacial climate-change velocity were essential refuges for many small-ranged species. 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.

Global Patterns of Geographic Range Size in Birds

Large-scale patterns of spatial variation in species geographic range size are central to many fundamental questions in macroecology and conservation biology. However, the global nature of these patterns has remained contentious, since previous studies have been geographically restricted and/or based on small taxonomic groups. Here, using a database on the breeding distributions of birds, we report the first (to our knowledge) global maps of variation in species range sizes for an entire taxonomic class. We show that range area does not follow a simple latitudinal pattern. Instead, the smallest range areas are attained on islands, in mountainous areas, and largely in the southern hemisphere. In contrast, bird species richness peaks around the equator, and towards higher latitudes. Despite these profoundly different latitudinal patterns, spatially explicit models reveal a weak tendency for areas with high species richness to house species with significantly smaller median range area. Taken together, these results show that for birds many spatial patterns in range size described in geographically restricted analyses do not reflect global rules. It remains to be discovered whether global patterns in geographic range size are best interpreted in terms of geographical variation in species assemblage packing, or in the rates of speciation, extinction, and dispersal that ultimately underlie biodiversity.

Spatial turnover in the global avifauna

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 Earths major land masses.

Human impacts and the global distribution of extinction risk

Understanding the global geographical distribution of extinction risk is a key challenge in conservation biology. It remains controversial, however, to what extent areas become threat hotspots simply because of high human impacts or due to predisposing ecological conditions. Limits to the taxonomic and geographical extent, resolution and quality of previously available data have precluded a full global assessment of the relative roles of these factors. Here, we use a new global database on the geographical distributions of birds on continents and continental islands to show that, after controlling for species richness, the best predictors of the global pattern of extinction risk are measures of human impact. Ecological gradients are of secondary importance at a global scale. The converse is true for individual biogeographic realms, within which variation in human impact is reduced and its influence on extinction risk globally is therefore underestimated. These results underline the importance of a global perspective on the mechanisms driving spatial patterns of extinction risk, and the key role of anthropogenic factors in driving the current extinction crisis.

Environmental and spatial influences upon species composition of a termite assemblage across neotropical forest islands

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.

City-wide relationships between green spaces, urban land use and topography

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.

Logging cuts the functional importance of invertebrates in tropical rainforest

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.

Feeding group responses of a Neotropical termite assemblage to rain forest fragmentation

Abstract. 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.