The evolution of critical thermal limits of life on Earth.

Understanding how species' thermal limits have evolved across the tree of life is central to predicting species' responses to climate change. Here, using experimentally-derived estimates of thermal tolerance limits for over 2000 terrestrial and aquatic species, we show that most of the variation in thermal tolerance can be attributed to a combination of adaptation to current climatic extremes, and the existence of evolutionary 'attractors' that reflect either boundaries or optima in thermal tolerance limits. Our results also reveal deep-time climate legacies in ectotherms, whereby orders that originated in cold paleoclimates have presently lower cold tolerance limits than those with warm thermal ancestry. Conversely, heat tolerance appears unrelated to climate ancestry. Cold tolerance has evolved more quickly than heat tolerance in endotherms and ectotherms. If the past tempo of evolution for upper thermal limits continues, adaptive responses in thermal limits will have limited potential to rescue the large majority of species given the unprecedented rate of contemporary climate change.

The geographical variation of network structure is scale dependent: understanding the biotic specialization of host-parasitoid networks.

Research on the structure of ecological networks suggests that a number of universal patterns exist. Historically, biotic specialization has been thought to increase towards the Equator. Yet, recent studies have challenged this view showing non-conclusive results. Most studies analysing the geographical variation in biotic specialization focus, however, only on the local scale. Little is known about how the geographical variation of network structure depends on the spatial scale of observation (i.e., from local to regional spatial scales). This should be remedied, as network structure changes as the spatial scale of observation changes, and the magnitude and shape of these changes can elucidate the mechanisms behind the geographical variation in biotic specialization. Here we analyse four facets of biotic specialization in host-parasitoid networks along gradients of climatic constancy, classifying the networks according to their spatial extension (local or regional). Namely, we analyse network connectance, consumer diet overlap, consumer diet breadth, and resource vulnerability at both local and regional scales along the gradients of both current climatic constancy and historical climatic change. While at the regional scale none of the climatic variables are associated to biotic specialization, at the local scale, network connectance, consumer diet overlap, and resource vulnerability decrease with current climatic constancy, whereas consumer generalism increases (i.e., broader diet breadths in tropical areas). Similar patterns are observed along the gradient of historical climatic change. We provide an explanation based on different beta-diversity for consumers and resources across the geographical gradients. Our results show that the geographical gradient of biotic specialization is not universal. It depends on both the facet of biotic specialization and the spatial scale of observation.

GlobTherm, a global database on thermal tolerances for aquatic and terrestrial organisms.

How climate affects species distributions is a longstanding question receiving renewed interest owing to the need to predict the impacts of global warming on biodiversity. Is climate change forcing species to live near their critical thermal limits? Are these limits likely to change through natural selection? These and other important questions can be addressed with models relating geographical distributions of species with climate data, but inferences made with these models are highly contingent on non-climatic factors such as biotic interactions. Improved understanding of climate change effects on species will require extensive analysis of thermal physiological traits, but such data are both scarce and scattered. To overcome current limitations, we created the GlobTherm database. The database contains experimentally derived species' thermal tolerance data currently comprising over 2,000 species of terrestrial, freshwater, intertidal and marine multicellular algae, plants, fungi, and animals. The GlobTherm database will be maintained and curated by iDiv with the aim to keep expanding it, and enable further investigations on the effects of climate on the distribution of life on Earth.

Stress from cold and drought as drivers of functional trait spectra in North American angiosperm tree assemblages.

Understanding how environmental change alters the composition of plant assemblages, and how this in turn affects ecosystem functioning is a major challenge in the face of global climate change. Assuming that values of plant traits express species adaptations to the environment, the trait-based approach is a promising way to achieve this goal. Nevertheless, how functional traits are related to species' environmental tolerances and how trait spectra respond to broad-scale environmental gradients remains largely unexplored. Here, we identify the main trait spectra for US angiosperm trees by testing hypotheses for the relationships between functional traits and species' environmental tolerances to environmental stresses, as well as quantifying the environmental drivers of assemblage means and variances of these traits. We analyzed >74,000 community assemblages from the US Forest Inventory and Analysis using 12 functional traits, five traits expressing species' environmental tolerances and 10 environmental variables. Results indicated that leaf traits, dispersal traits, and traits related to stem hydraulics were related to cold or drought tolerance, and their assemblage means were best explained by minimum temperatures. Assemblage means of traits related to shade tolerance (tree growth rate, leaf phosphorus content, and bark thickness) were best explained by aridity index. Surprisingly, aridity index, rather than minimum temperature, was the best predictors of assemblage variances of most traits, although these relationships were variable and weak overall. We conclude that temperature is likely to be the most important driver of functional community structure of North American angiosperm trees by selecting for optimum strategies along the cold and drought stress trade-off. In turn, water availability primarily affects traits related to shade tolerance through its effect on forest canopy structure and vegetation openness.

The diversity and abundance of North American bird assemblages fail to track changing productivity.

Plant biomass or productivity and the species richness of birds are associated across a range of spatial scales. Species-energy theory is generally assumed to explain these correlations. If true, bird richness should also track productivity temporally, and there should be spatial and temporal relationships between productivity and both bird abundance and bird richness. Using the summer normalized difference vegetation index (NDVI) for 1982-2006 and the North American Breeding Bird Survey, we evaluated the response of avian richness and abundance to interannual changes in plant biomass or productivity. We found positive spatial relationships between richness and NDVI for all 25 years. Temporally, however, richness and NDVI were positively associated at 1579 survey sites and negatively associated at 1627 sites (mean r2 = 0.09). Further, total abundance and NDVI were unrelated spatially (r2 values spanning < 0.01 and 0.03) and weakly related temporally (mean r2 = 0.10). We found no evidence that productivity drives bird richness beyond the spatial correlations, and neither prediction arising from species-energy theory was confirmed. Spatial relationships between productivity and bird richness may thus be largely spurious, arising via covariance between plant biomass or productivity and vegetation structural complexity, and the latter may be driving bird communities. This is consistent with the MacArthurs' classic hypothesis that the vertical profile of foliage drives bird species diversity.

Evolutionary histories of soil fungi are reflected in their large-scale biogeography.

Although fungal communities are known to vary along latitudinal gradients, mechanisms underlying this pattern are not well-understood. We used high-throughput sequencing to examine the large-scale distributions of soil fungi and their relation to evolutionary history. We tested the Tropical Conservatism Hypothesis, which predicts that ancestral fungal groups should be more restricted to tropical latitudes and conditions than would more recently derived groups. We found support for this hypothesis in that older phyla preferred significantly lower latitudes and warmer, wetter conditions than did younger phyla. Moreover, preferences for higher latitudes and lower precipitation levels were significantly phylogenetically conserved among the six younger phyla, possibly because the older phyla possess a zoospore stage that is vulnerable to drought, whereas the younger phyla retain protective cell walls throughout their life cycle. Our study provides novel evidence that the Tropical Conservatism Hypothesis applies to microbes as well as plants and animals.

Community phylogenetics at the biogeographical scale: cold tolerance, niche conservatism and the structure of North American forests.

AimThe fossil record has led to a historical explanation for forest diversity gradients within the cool parts of the Northern Hemisphere, founded on a limited ability of woody angiosperm clades to adapt to mid-Tertiary cooling. We tested four predictions of how this should be manifested in the phylogenetic structure of 91,340 communities: (1) forests to the north should comprise species from younger clades (families) than forests to the south; (2) average cold tolerance at a local site should be associated with the mean family age (MFA) of species; (3) minimum temperature should account for MFA better than alternative environmental variables; and (4) traits associated with survival in cold climates should evolve under a niche conservatism constraint. LocationThe contiguous United States. MethodsWe extracted angiosperms from the US Forest Service's Forest Inventory and Analysis database. MFA was calculated by assigning age of the family to which each species belongs and averaging across the species in each community. We developed a phylogeny to identify phylogenetic signal in five traits: realized cold tolerance, seed size, seed dispersal mode, leaf phenology and height. Phylogenetic signal representation curves and phylogenetic generalized least squares were used to compare patterns of trait evolution against Brownian motion. Eleven predictors structured at broad or local scales were generated to explore relationships between environment and MFA using random forest and general linear models. ResultsConsistent with predictions, (1) southern communities comprise angiosperm species from older families than northern communities, (2) cold tolerance is the trait most strongly associated with local MFA, (3) minimum temperature in the coldest month is the environmental variable that best describes MFA, broad-scale variables being much stronger correlates than local-scale variables, and (4) the phylogenetic structures of cold tolerance and at least one other trait associated with survivorship in cold climates indicate niche conservatism. Main conclusionsTropical niche conservatism in the face of long-term climate change, probably initiated in the Late Cretaceous associated with the rise of the Rocky Mountains, is a strong driver of the phylogenetic structure of the angiosperm component of forest communities across the USA. However, local deterministic and/or stochastic processes account for perhaps a quarter of the variation in the MFA of local communities.

The imprint of Cenozoic migrations and evolutionary history on the biogeographic gradient of body size in New World mammals.

Ecology, evolution, and historical events all contribute to biogeographic patterns, but studies that integrate them are scarce. Here we focus on how biotic exchanges of mammals during the Late Cenozoic have contributed to current geographic body size patterns. We explore differences in the environmental correlates and phylogenetic patterning of body size between groups of mammals participating and not participating in past biotic exchanges. Both the association of body size with environmental predictors and its phylogenetic signal were stronger for groups that immigrated into North or South America than for indigenous groups. This pattern, which held when extinct clades were included in the analyses, can be interpreted on the basis of the length of time that clades have had to diversify and occupy niche space. Moreover, we identify a role for historical events, such as Cenozoic migrations, in configuring contemporary mammal body size patterns and illustrate where these influences have been strongest for New World mammals.

Global climate change is confounding species conservation strategies.

Most organisms face similar problems with respect to their conservation in the face of global climate change. Here, we examine probable effects of climate change on the hyperdiverse plant family Orchidaceae. In the 20th century, the major concerns for orchid conservation revolved around unsustainable harvest for the orchid trade and, more importantly, land conversion from natural ecosystems to those unable to support wild orchid populations. Land conversion included logging, fire regimes and forest conversions to agricultural systems. Although those forms of degradation continue, an additional suite of threats has emerged, fueled by global climate change. Global climate change involves more than responses of orchid populations to increases in ambient temperature. Increasing temperature induces secondary effects that can be more significant than simple changes in temperature. Among these new threats are extended and prolonged fire seasons, rising sea levels, increases in cyclonic storms, seasonal climate shifts, changes in orthographic wind dew point and increased drought. The long-term outlook for orchid biodiversity in the wild is dismal, as it is for many animal groups, and we need to start rethinking strategies for conservation in a rapidly changing world.

Ice age climate, evolutionary constraints and diversity patterns of European dung beetles.

Current climate and Pleistocene climatic changes are both known to be associated with geographical patterns of diversity. We assess their associations with the European Scarabaeinae dung beetles, a group with high dispersal ability and well-known adaptations to warm environments. By assessing spatial stationarity in climate variability since the last glacial maximum (LGM), we find that current scarab richness is related to the location of their limits of thermal tolerance during the LGM. These limits mark a strong change in their current species richness-environment relationships. Furthermore, northern scarab assemblages are nested and composed of a phylogenetically clustered subset of large-range sized generalist species, whereas southern ones are diverse and variable in composition. Our results show that species responses to current climate are limited by the evolution of assemblages that occupied relatively climatically stable areas during the Pleistocene, and by post-glacial dispersal in those that were strongly affected by glaciations.

Niche conservatism as an emerging principle in ecology and conservation biology.

The diversity of life is ultimately generated by evolution, and much attention has focused on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to instead remain similar over time [niche conservatism (NC)] has many consequences for the fundamental patterns and processes studied in ecology and conservation biology. Here, we describe the mounting evidence for the importance of NC to major topics in ecology (e.g. species richness, ecosystem function) and conservation (e.g. climate change, invasive species). We also review other areas where it may be important but has generally been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic interactions) and conservation (e.g. habitat modification). We summarize methods for testing for NC, and suggest that a commonly used and advocated method (involving a test for phylogenetic signal) is potentially problematic, and describe alternative approaches. We suggest that considering NC: (1) focuses attention on the within-species processes that cause traits to be conserved over time, (2) emphasizes connections between questions and research areas that are not obviously related (e.g. invasives, global warming, tropical richness), and (3) suggests new areas for research (e.g. why are some clades largely nocturnal? why do related species share diseases?).

Phylogeny, niche conservatism and the latitudinal diversity gradient in mammals.

Biologists have long searched for mechanisms responsible for the increase in species richness with decreasing latitude. The strong correlation between species richness and climate is frequently interpreted as reflecting a causal link via processes linked to energy or evolutionary rates. Here, we investigate how the aggregation of clades, as dictated by phylogeny, can give rise to significant climate-richness gradients without gradients in diversification or environmental carrying capacity. The relationship between climate and species richness varies considerably between clades, regions and time periods in a global-scale phylogenetically informed analysis of all terrestrial mammal species. Many young clades show negative richness-temperature slopes (more species at cooler temperatures), with the ages of these clades coinciding with the expansion of temperate climate zones in the late Eocene. In carnivores, we find steeply positive richness-temperature slopes in clades with restricted distributions and tropical origins (e.g. cat clade), whereas widespread, temperate clades exhibit shallow, negative slopes (e.g. dog-bear clade). We show that the slope of the global climate-richness gradient in mammals is driven by aggregating Chiroptera (bats) with their Eutherian sister group. Our findings indicate that the evolutionary history should be accounted for as part of any search for causal links between environment and species richness.

Climate, niche conservatism, and the global bird diversity gradient.

We tested the proposition that there are more species in the tropics because basal clades adapted to warm paleoclimates have been lost in regions now experiencing cool climates. Molecular phylogenies were used to classify species as "basal" and "derived" based on their family, and their richness patterns were contrasted. Path models also evaluated environmental predictors of richness patterns. As predicted, basal clades are more diverse in the lowland tropics, whereas derived clades are more diverse in the extratropics and high-altitude tropics. Seventy-four percent of the variation in bird richness was explained by environmental variables, but models differed for basal and derived groups. The overall gradient is described by the spatial pattern of basal clades, although there are differences in the Old and New Worlds. We conclude that in ecological time, the global richness gradient reflects birds' responses to climatic gradients, partially operating via plants. Over evolutionary time, the gradient primarily reflects the extirpation of species in older clades from parts of the world that have become cooler in the present. A strong secondary effect arises from dispersal of clades from centers of origin and subsequent radiations. Overall, the diversity gradient is well explained by niche conservatism and the "time-for-speciation" hypothesis.

A global evaluation of metabolic theory as an explanation for terrestrial species richness gradients.

We compiled 46 broadscale data sets of species richness for a wide range of terrestrial plant, invertebrate, and ectothermic vertebrate groups in all parts of the world to test the ability of metabolic theory to account for observed diversity gradients. The theory makes two related predictions: (1) In-transformed richness is linearly associated with a linear, inverse transformation of annual temperature, and (2) the slope of the relationship is near -0.65. Of the 46 data sets, 14 had no significant relationship; of the remaining 32, nine were linear, meeting prediction 1. Model I (ordinary least squares, OLS) and model II (reduced major axis, RMA) regressions then tested the linear slopes against prediction 2. In the 23 data sets having nonlinear relationships between richness and temperature, split-line regression divided the data into linear components, and regressions were done on each component to test prediction 2 for subsets of the data. Of the 46 data sets analyzed in their entirety using OLS regression, one was consistent with metabolic theory (meeting both predictions), and one was possibly consistent. Using RMA regression, no data sets were consistent. Of 67 analyses of prediction 2 using OLS regression on all linear data sets and subsets, two were consistent with the prediction, and four were possibly consistent. Using RMA regression, one was consistent (albeit weakly), and four were possibly consistent. We also found that the relationship between richness and temperature is both taxonomically and geographically conditional, and there is no evidence for a universal response of diversity to temperature. Meta-analyses confirmed significant heterogeneity in slopes among data sets, and the combined slopes across studies were significantly lower than the range of slopes predicted by metabolic theory based on both OLS and RMA regressions. We conclude that metabolic theory, as currently formulated, is a poor predictor of observed diversity gradients in most terrestrial systems.

Macroevolutionary dynamics in environmental space and the latitudinal diversity gradient in New World birds.

Correlations between species richness and climate suggest non-random occupation of environmental space and niche evolution through time. However, the evolutionary mechanisms involved remain unresolved. Here, we partition the occupation of environmental space into intra- and inter-clade components to differentiate a model based on pure conservation of ancestral niches with higher diversification rates in the tropics, and an adaptive radiation model based on shifts in adaptive peaks at the family level allowing occupation of temperate regions. We examined these mechanisms using within- and among-family skewness components based on centroids of 3560 New World bird species across four environmental variables. We found that the accumulation of species in the tropics is a result of both processes. The components of adaptive radiation have family level skewness of species' distributions strongly structured in space, but not phylogenetically, according to the integrated analyses of spatial filters and phylogenetic eigenvectors. Moreover, stronger radiation components were found for energy variables, which are often used to argue for direct climatic effects on diversity. Thus, the correspondence between diversity and climate may be due to the conservation of ancestral tropical niches coupled with repeated broad shifts in adaptive peaks during birds' evolutionary history more than by higher diversification rates driven by more energy in the tropics.

The mid-domain effect and diversity gradients: is there anything to learn?

The mid-domain effect (MDE) has been proposed as a null model for diversity gradients and an explanation for observed patterns. Here we respond to a recent defense of the concept, explaining that it cannot represent a viable model in either real or null worlds. First, the MDE misrepresents the nature of species ranges. There is also an internal logical inconsistency underlying the MDE because the range size frequency distribution, necessary to generate a hump-shaped pattern under randomization, cannot exist in the absence of environmental gradients and is generated by the ecological and historical processes that the MDE claims to exclude.

A test of multiple hypotheses for the species richness gradient of South American owls.

Many mechanisms have been proposed to explain broad scale spatial patterns in species richness. In this paper, we evaluate five explanations for geographic gradients in species richness, using South American owls as a model. We compared the explanatory power of contemporary climate, landcover diversity, spatial climatic heterogeneity, evolutionary history, and area. An important aspect of our analyses is that very different hypotheses, such as history and area, can be quantified at the same observation scale and, consequently can be incorporated into a single analytical framework. Both area effects and owl phylogenetic history were poorly associated with richness, whereas contemporary climate, climatic heterogeneity at the mesoscale and landcover diversity explained ca. 53% of the variation in species richness. We conclude that both climate and environmental heterogeneity should be retained as plausible explanations for the diversity gradient. Turnover rates and scaling effects, on the other hand, although perhaps useful for detecting faunal changes and beta diversity at local and regional scales, are not strong explanations for the owl diversity gradient.

Herbivore responses to plant secondary compounds: a test of phytochemical coevolution theory.

Literature data were collected on the floristic distribution and toxicity of phytochemicals to herbivores and on herbivore specialization in order to test phytochemical coevolution theory. The theory makes four predictions that can be tested with this information. Herbivores can adapt to novel, more toxic chemicals by becoming specialists, or they can become generalists but at the cost of lower feeding success on any particular host. Thus, the first two predictions are as follows: herbivores should do better on chemicals that are present in their normal host, and this pattern should be stronger for specialists than for generalists. The "escape and radiation" aspect of the theory holds that if a plant taxon with a novel defense chemical diversifies, the chemical will become widespread. Eventually, herbivores will adapt to and disarm it. So the third prediction is that more widespread chemicals are less toxic than more narrowly distributed ones. Because generalists should not do as well as specialists on chemicals disarmed by the latter, the fourth prediction is that the third prediction should be more true for generalists than specialists and should depend on presence/absence of the chemical in the normal host. Multiple regressions of toxicity (herbivore mortality and final weight) on three predictor variables (chemical presence/absence in the normal host, specialism, and chemical floristic distribution) and relevant interactions were used to test these predictions. Chemical presence/absence in the normal host, the interaction between this variable and specialism, and chemical floristic distribution had significant effects on both measures of toxicity, supporting the first three predictions of the model. Support for the fourth prediction (a three-way interaction among all predictor variables) was evident for final weight but not mortality, perhaps because growth is more responsive to toxicity differences than survival. In short, the phytochemistry literature provides broad support for the phytochemical coevolution model.