Aim Geographic variation in the species richness of birds has been shown to be strongly associated with annual water and energy levels (actual evapotranspiration, AET) at the global scale. However, the gradient in eastern North America appears to be anomalous, because richness is greatest around the Great Lakes, whereas AET is highest in the south-eastern US. Here I examine if birds may be responding to vegetation produced during the breeding season rather than to annual production. Location North America east of longitude 98degrees W. Methods The bird richness pattern was examined using climatic variables, remotely sensed estimates of annual and seasonal plant biomass, and time since areas were exposed by the retreating Laurentide ice sheet from 20,000 to 6000 yr BP. Results Average summer GVI (Global Vegetation Index, derived from NDVI) was found to be positively linearly associated with richness, explaining 82% of the variance, whereas the relationships between richness and annual measures of both AET and GVI were curvilinear. The pattern of retreat of the Laurentide ice sheet explained an additional 6% of the variance in richness, consistent with a previous analysis of Canadian birds. Main conclusions In eastern North America, a seasonal variable associated with plant production explains the diversity gradient rather than the annual measures, but it does not undermine a general conclusion that bird diversity is closely linked with plant biomass. Further, both contemporary and historical factors appear to influence the gradient, and an association between bird richness and the geographic pattern of glacial retreat is detectable in both climatic and plant-biomass models of bird diversity.

There is a well documented relationship between parasitoid species assemblage size and host feeding niche. Parasitoid assemblage size peaks on hosts thought to have intermediate levels of physical refuge. We examined the influence of refuges on parasitoid coexistence using pairs of specialist parasitoids in a controlled laboratory environment. Using physical barriers we excluded parasitoids from 0, 25, 50 or 75% of the hosts to simulate host refuge. We found no evidence that host refuges can promote parasitoid coexistence in a simplified laboratory environment. Results were similar whether pairs of parasitoid species were competitively disparate or competitively similar. Our results suggest that spatial heterogeneity in parasitoid attack rate was not sufficient to maintain parasitoid coexistence regardless of host refuge, and we argue that the level of spatial heterogeneity necessary to promote coexistence is rare in nature. We conclude that in most systems the coexistence of specialist parasitoids cannot be explained by a host refuge effect.

Models of host-parasitoid dynamics often assume constant levels of spatial heterogeneity in parasitoid attack rate, which tends to stabilize the interactions. Recently, authors have questioned this assumption and shown that outcomes of simple host-parasitoid models change if spatial heterogeneity is allowed to vary with parasitoid density. Here, we allow spatial heterogeneity to vary with either parasitoid density or percent parasitism in a model designed to explain specialist parasitoid coexistence on insect hosts with various levels of refuge. By examining this model we can evaluate the effect of varying spatial heterogeneity on a more complex model in which spatial heterogeneity is not considered the primary determinant of persistence. By modeling communities with one host and two parasitoid species, we show that the probability of species persistence for the competitively inferior parasitoid depends on the assumed relationship between spatial heterogeneity and both parasitoid density and percent parasitism. The probability of parasitoid coexistence is generally lower when spatial heterogeneity varies with parasitoid demographics. We conclude that the conditions for which host refuge promote specialist parasitoid coexistence are less common that proposed by the original model. Finally, we compared a model in which spatial heterogeneity varies with percent parasitism to data from laboratory trials and find a reasonable fit. We conclude that the change in spatial heterogeneity strongly influenced the outcome of the laboratory trials, and we suggest more research is necessary before researchers can assume constant spatial heterogeneity in future models.

Although it has long been held that plant diversity must influence animal diversity, the nature of this relationship remains poorly understood at large spatial scales. We compare the species richness patterns of vascular plants and mammals in north-eastern Spain using a 100-km(2) grain size to examine patterns of covariation. We found that the total mammal richness pattern, as well as those of herbivores and carnivores considered separately, only weakly corresponded to the pattern of plants. Rather, mammal richness was best described by climatic variables incorporating water inputs, and after adding these variables to multiple regression models, plant and mammal richness were virtually independent. We conclude that the observed association, although weak, is explained by shared responses of both groups to climate, and thus, plant richness has no influence on the richness pattern of Catalan mammals.

Primary productivity and animal species richness are associated across spatial scales and extents. The productivity hypothesis had been credited for explaining these correlations by asserting that places with more energy fixed in the system by plants should support more individuals, whereby the community accumulates more species. Many have interpreted the spatial association between richness and productivity as support for the hypothesis. If true, bird richness should track productivity temporally, and there should be spatial and temporal relationships between productivity and both bird abundance and richness.

We tested these predictions in the breeding season and winter across space and time. Using a remotely-sensed primary productivity proxy (NDVI) and avian data from two large North American citizen science surveys, the Breeding Bird Survey (BBS) and Christmas Bird Count (CBC), we evaluated the response of avian richness and abundance to interannual changes in plant productivity across 25 years. In the breeding season we found positive spatial relationships between richness and NDVI each year, but when evaluated temporally no support for the hypothesis was found. Richness and NDVI were positively associated at only half (49.3%) of BBS sites (mean r2 = 0.09). Further, total abundance and productivity were unrelated.

Despite summer findings, it is possible that richness depends more on productivity in winter when birds are stressed by harsh conditions. Using CBC data we found that across several thousand communities the number of individuals was not spatially associated with productivity, providing no support for the productivity hypothesis.

In these and earlier tests of the hypothesis, trait-neutral partitioning of resources across individuals is assumed, though it may be more realistic that partitioning is in unequal shares. To test for energetic trait differences, we regressed total avian biomass at sites against NDVI and found that avian biomass is independent of productivity.

We conclude that primary productivity is unlikely the driver of bird diversity. Spatial relationships between productivity and bird richness may be spuriously arising via covariance between productivity and vegetation structural complexity, and the latter may be driving bird communities, consistent with the MacArthurs’ classic hypothesis that the vertical profile of foliage drives bird species diversity.

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.

Aim To investigate the relative contributions of current vs. historical factors in explaining broad-scale diversity gradients using a combination of contemporary factors and a quantitative estimate of the temporal accessibility of areas for recolonization created by glacial retreat following the most recent Ice Age. Location The part of the Nearctic region of North America that was covered by ice sheets during the glacial maximum 20 000 BP. Methods We used range maps to estimate the species richness of mammals and terrestrial birds in 48 400 km(2) cells. Current conditions in each cell were quantified using seven climatic and topographical variables. Historical conditions were estimated using the number of years before present when an area became exposed as the ice sheets retreated during the post-Pleistocene climate warming. We attempted to tease apart contemporary and historical effects using multiple regression, partial regression and spatial autocorrelation analysis. Results A measure of current energy inputs, potential evapotranspiration, explained 76-82% of the variance in species richness, but time since deglaciation explained an additional 8-13% of the variance, primarily due to effects operating at large spatial scales. Because of spatial covariation between the historical climates influencing the melting of the ice sheet and current climates, it was not possible to partition their effects fully, but of the independent effects that could be identified, current climate explained two to seven times more variance in richness patterns than age. Main Conclusions Factors acting in the present appear to have the strongest influence on the diversity gradient, but an historical signal persisting at least 13 000 years is still detectable. This has implications for modelling changes in diversity patterns in response to future global warming.

Studies mostly focused on communities of primary producers have shown that species richness provides and promotes fundamental ecosystem services. However, we know very little about the factors influencing ecosystem services provided by higher trophic levels in natural food webs. Here we present evidence that differences in food web structure and the richness of herbivores in 19 plant-herbivore-parasitoid food webs influence the service supplied by natural enemies, namely, the parasitism rates on hosts. Specifically, we find that parasitoids function better in simple food webs than in complex ones, a result relevant to biological control practice. More generally, we show that species richness per se only contributes partially to the understanding of higher-level ecosystem services in multitrophic communities, and that changes in food web complexity should also be taken into account when predicting the effects of human-driven disturbances in natural communities.