Does climate limit species richness by limiting individual species’ ranges?

Broad-scale geographical variation in species richness is strongly correlated with climate, yet the mechanisms underlying this correlation are still unclear. We test two broad classes of hypotheses to explain this pattern. Bottom-up hypotheses propose that the environment determines individual species’ ranges. Ranges then sum up to yield species richness patterns. Top-down hypotheses propose that the environment limits the number of species that occur in a region, but not which ones. We test these two classes of hypotheses using a natural experiment: seasonal changes in environmental variables and seasonal range shifts of 625 migratory birds in the Americas. We show that richness seasonally tracks the environment. By contrast, individual species’ geographical distributions do not. Rather, species occupy different sets of environmental conditions in two seasons. Our results are inconsistent with extant bottom-up hypotheses. Instead, a top-down mechanism appears to constrain the number of species that can occur in a given region.     

Geographical separation of two Ulex species at three spatial scales: does competition limit species' ranges?

It is a common assumption that species' ranges are limited by their physiological tolerances to climatic factors, Biotic factors, such as competition, are rarely considered. We investigated the distributions of Ulex minor and U. gallii at three spatial scales from geographic ranges to individual heaths - to examine whether the species are negatively associated, as predicted by the hypothesis that the ranges of the species are limited by competition with each other. Distribution maps for the British Isles and France (100 400 km² survey units) show the two species have largely separated, but slightly overlapping ranges. A region of range overlap on the heaths of Dorset, southern England was mapped using 4 ha survey squares. There was strong negative association between the species, and the heaths could be divided into zones where one species was dominant. There was some indication of edaphic differences between the U. minor -dominated zones and the U. gallii zones. The few heaths where the species co-occurred were surveyed using 4 m² quadrats placed along transects. Usually one species was widespread over the heath, while the other occurred in patches. The species were strongly negatively associated in all transects. Therefore, the two species showed strong negative associations at three mapping scales. Apparent co-occurrences detected at one spatial scale largely disappeared when species were mapped at finer scales, emphasising the fractal nature of distributions. This provides evidence that the distributions of the two species are not independent and that they cannot coexist, and therefore that their ranges are limited by competition. Over their ranges, competitive superiority is probably determined by the climate. At the range boundaries in the region of overlap, climate is not important, but other physical factors such as edaphic conditions may determine the outcome of competition.     

Can 'continuity indicator species' predict species richness or red-listed species of saproxylic beetles?

This study investigates the relationship between the abundance of wood-rotting fungus suggested as 'continuity indicator species' and environmental variables for the assemblage of saproxylic (wood-living) beetles associated with Fomitopsis pinicola fruiting bodies in a mature spruce forest in southeastern Norway. The presence of species thought to indicate continuity in old growth is one of the criteria used when finding and delineating small protected areas ('woodland key habitats') in Scandinavian forestry. Although it is clear that remnants of old-growth forest are important for many taxa, documentation as to which entities or species the indicator species indeed indicate is scarce. If stands with a continuous and unbroken input of dead wood have a unique assemblage of wood-rotting fungi, it seems relevant to ask if these stands also have a unique assemblage of rare saproxylic beetles. I find that the indicator species exhibit no significant correlations with beetle species richness or with the presence of red-listed saproxylic beetles as a group. The different characteristics of dead wood conditions are the most important environmental variables that explain both the species richness and the presence of red-listed beetles. Single-species analyses reveal contrasting relationships. The red-listed beetle Atomaria alpina shows a significant and positive association to the abundance of indicator species. Contrary, a group of three red-listed species with similar ecology in the family Cisidae exhibits a significant and negative association to indicator species abundance. This indicates that important patterns are concealed when considering general measures such as overall presence of red-listed beetles. Single-species studies are necessary in order to correctly understand how rare beetles respond to forestry activities and to develop a policy that can secure their continuing existence in the boreal forest.     

Does coevolution promote species richness in parasitic cuckoos?

Why some lineages have diversified into larger numbers of species than others is a fundamental but still relatively poorly understood aspect of the evolutionary process. Coevolution has been recognized as a potentially important engine of speciation, but has rarely been tested in a comparative framework. We use a comparative approach based on a complete phylogeny of all living cuckoos to test whether parasite–host coevolution is associated with patterns of cuckoo species richness. There are no clear differences between parental and parasitic cuckoos in the number of species per genus. However, a cladogenesis test shows that brood parasitism is associated with both significantly higher speciation and extinction rates. Furthermore, subspecies diversification rate estimates were over twice as high in parasitic cuckoos as in parental cuckoos. Among parasitic cuckoos, there is marked variation in the severity of the detrimental effects on host fitness; chicks of some cuckoo species are raised alongside the young of the host and others are more virulent, with the cuckoo chick ejecting or killing the eggs/young of the host. We show that cuckoos with a more virulent parasitic strategy have more recognized subspecies. In addition, cuckoo species with more recognized subspecies have more hosts. These results hold after controlling for confounding geographical effects such as range size and isolation in archipelagos. Although the power of our analyses is limited by the fact that brood parasitism evolved independently only three times in cuckoos, our results suggest that coevolutionary arms races with hosts have contributed to higher speciation and extinction rates in parasitic cuckoos.     

Does climate determine broad‐scale patterns of species richness? A test of the causal link by natural experiment

Aim Broad‐scale spatial patterns of species richness are very strongly correlated with climatic variables. If there is a causal link, i.e. if climate directly or indirectly determines patterns of richness, then when the climatic variables change, richness should change in the manner that spatial correlations between richness and climate would predict. The present study tests this prediction using seasonal changes in climatic variables and bird richness. Location We used a grid of equal area quadrats (37 000 km²) covering North and Central America as far south as Nicaragua. Methods Summer and winter bird distribution data were drawn from monographs and field guides. Climatic data came from published sources. We also used remotely sensed NDVI (normalized difference vegetation index — a measure of greenness). Results Bird species richness changes temporally (between summer and winter) in a manner that is close to, but statistically distinguishable from, the change one would predict from models relating the spatial variation in richness at a single time to climatic variables. If one further takes into account the seasonal changes in NDVI and within‐season variability of temperature and precipitation, then winter and summer richness follow congruent, statistically indistinguishable patterns. Main conclusions Our results are consistent with the hypothesis that climatic variables (temperature and precipitation) and vegetation cover directly or indirectly influence patterns of bird species richness.     

Does a facultative mutualism limit species range expansion?

The availability and quality of mutualists beyond a species’ range edge may limit range expansion. With the legume Chamaecrista fasciculata, we asked to what extent the availability and quality of rhizobia beyond the range edge limits host range expansion. We tested the effect of rhizobia availability on plant growth by transplanting seed from three locations into five sites spanning C. fasciculata’s range (interior, at the northern and western range edges, and beyond the range edges), and inoculating half the seeds with rhizobia. We recorded growth of all surviving plants, and, for the uninoculated plants, whether they had formed nodules or not. We isolated rhizobia from nodules collected on the uninoculated plants, and cross-inoculated seed from four populations (both range edge and interior populations) in the greenhouse to determine whether the quality of rhizobia differed between regions. We found that seeds transplanted beyond the range edge were less likely to be nodulated when they were not experimentally inoculated, and there was benefit to inoculation at all sites. In the greenhouse, the three inocula that formed nodules on plants, from the range interior, northern edge and beyond the northern edge, did not detectably differ in their effect on plant growth. These results suggest that low densities of suitable rhizobia beyond the range edge may limit range expansion of legume species.     

Does species diversity limit productivity in natural grassland communities?

Theoretical analyses and experimental studies of synthesized assemblages indicate that under particular circumstances species diversity can enhance community productivity through niche complementarity. It remains unclear whether this process has important effects in mature natural ecosystems where competitive feedbacks and complex environmental influences affect diversity–productivity relationships. In this study, we evaluated diversity–productivity relationships while statistically controlling for environmental influences in 12 natural grassland ecosystems. Because diversity–productivity relationships are conspicuously nonlinear, we developed a nonlinear structural equation modeling (SEM) methodology to separate the effects of diversity on productivity from the effects of productivity on diversity. Meta-analysis was used to summarize the SEM findings across studies. While competitive effects were readily detected, enhancement of production by diversity was not. These results suggest that the influence of small-scale diversity on productivity in mature natural systems is a weak force, both in absolute terms and relative to the effects of other controls on productivity.     

How to describe species richness patterns for bryophyte conservation?

A large amount of data for inconspicuous taxa is stored in natural history collections; however, this information is often neglected for biodiversity patterns studies. Here, we evaluate the performance of direct interpolation of museum collections data, equivalent to the traditional approach used in bryophyte conservation planning, and stacked species distribution models (S‐SDMs) to produce reliable reconstructions of species richness patterns, given that differences between these methods have been insufficiently evaluated for inconspicuous taxa. Our objective was to contrast if species distribution models produce better inferences of diversity richness than simply selecting areas with the higher species numbers. As model species, we selected Iberian species of the genus Grimmia (Bryophyta), and we used four well‐collected areas to compare and validate the following models: 1) four Maxent richness models, each generated without the data from one of the four areas, and a reference model created using all of the data and 2) four richness models obtained through direct spatial interpolation, each generated without the data from one area, and a reference model created with all of the data. The correlations between the partial and reference Maxent models were higher in all cases (0.45 to 0.99), whereas the correlations between the spatial interpolation models were negative and weak (−0.3 to −0.06). Our results demonstrate for the first time that S‐SDMs offer a useful tool for identifying detailed richness patterns for inconspicuous taxa such as bryophytes and improving incomplete distributions by assessing the potential richness of under‐surveyed areas, filling major gaps in the available data. In addition, the proposed strategy would enhance the value of the vast number of specimens housed in biological collections.     

How do climate warming and species richness affect CO2 fluxes in experimental grasslands?

This paper presents the results of 2 yr of CO(2) flux measurements on grassland communities of varying species richness, exposed to either the current or a warmer climate. We grew experimental plant communities containing one, three or nine grassland species in 12 sunlit, climate-controlled chambers. Half of these chambers were exposed to ambient air temperatures, while the other half were warmed by 3 degrees C. Equal amounts of water were added to heated and unheated communities, implying drier soils if warming increased evapotranspiration. Three main CO(2) fluxes (gross photosynthesis, above-ground and below-ground respiration) were measured multiple times per year and reconstructed hourly or half-hourly by relating them to their most important environmental driver. While CO(2) outputs through respiration were largely unchanged under warming, CO(2) inputs through photosynthesis were lowered, especially in summer, when heat and drought stress were higher. Above-ground CO(2) fluxes were significantly increased in multispecies communities, as more complementary resource use stimulated productivity. Finally, effects of warming appeared to be smallest in monocultures. This study shows that in a future warmer climate the CO(2) sink capacity of temperate grasslands could decline, and that such adverse effects are not likely to be mitigated by efforts to maintain or increase species richness.     

Assessments of species’ vulnerability to climate change: from pseudo to science

Climate change vulnerability assessments (CCVAs) are important tools to plan for and mitigate potential impacts of climate change. However, CCVAs often lack scientific rigor, which can ultimately lead to poor conservation prioritization and associated ecological and economic costs. We discuss the need to improve comparability and consistency of CCVAs and either validate their findings or improve assessment of CCVA uncertainty and sensitivity to methodological assumptions.     

How do climate warming and plant species richness affect water use in experimental grasslands?

Climate warming and plant species richness loss have been the subject of numerous experiments, but studies on their combined impact are lacking. Here we studied how both warming and species richness loss affect water use in grasslands, while identifying interactions between these global changes. Experimental ecosystems containing one, three or nine grassland species from three functional groups were grown in 12 sunlit, climate-controlled chambers (2.25 m² ground area) in Wilrijk, Belgium. Half of these chambers were exposed to ambient air temperatures (unheated), while the other half were warmed by 3°C (heated). Equal amounts of water were added to heated and unheated communities, so that warming would imply drier soils if evapotranspiration (ET) was higher. After an initial ET increase in response to warming, stomatal regulation and lower above-ground productivity resulted in ET values comparable with those recorded in the unheated communities. As a result of the decreased biomass production, water use efficiency (WUE) was reduced by warming. Higher complementarity and the improved competitive success of water-efficient species in mixtures led to an increased WUE in multi-species communities as compared to monocultures, regardless of the induced warming. However, since the WUE of individual species was affected in different ways by higher temperatures, compositional changes in mixtures seem likely under climatic change due to shifts in competitiveness. In conclusion, while increased complementarity and selection of water-efficient species ensured more efficient water use in mixtures than monocultures, global warming will likely decrease this WUE, and this may be most pronounced in species-rich communities.     

Identifying ‘climate keystone species’ as a tool for conserving ecological communities under climate change

Aim

Climate change affects ecological communities via impacts on species. The community's response to climate change can be represented as the temporal trend in a climate-related functional property that is quantified using a relevant functional trait. Noteworthy, some species influence this response in the community more strongly than others.

Innovation

Leveraging on the concept of keystone species, we propose that species with a strong effect on the community's functional response to climate change beyond their relative abundance can be considered as ‘climate keystone species’. We develop a stepwise tool to determine species' effects on a community's climate response and identify climate keystone species. We quantify the species-specific effect by measuring the difference in the community's climate response with and without the species. Next, we identify climate keystone species as those with a strong residual effect after weighting with their relative abundances in the community.

Main Conclusions

To illustrate the use of the stepwise tool with empirical data, we identify climate keystone species that have a strong effect on the change in the average temperature niche in North American bird communities over time and find the identification tool ecologically relevant. Identification of climate keystone species can serve as an additional conservation method to efficiently protect ecological communities and, in turn, the ecosystem functions they provide.     

Are species' responses to global change predicted by past niche evolution?

Predicting how and when adaptive evolution might rescue species from global change, and integrating this process into tools of biodiversity forecasting, has now become an urgent task. Here, we explored whether recent population trends of species can be explained by their past rate of niche evolution, which can be inferred from increasingly available phylogenetic and niche data. We examined the assemblage of 409 European bird species for which estimates of demographic trends between 1970 and 2000 are available, along with a species-level phylogeny and data on climatic, habitat and trophic niches. We found that species'' proneness to demographic decline is associated with slow evolution of the habitat niche in the past, in addition to certain current-day life-history and ecological traits. A similar result was found at a higher taxonomic level, where families prone to decline have had a history of slower evolution of climatic and habitat niches. Our results support the view that niche conservatism can prevent some species from coping with environmental change. Thus, linking patterns of past niche evolution and contemporary species dynamics for large species samples may provide insights into how niche evolution may rescue certain lineages in the face of global change.