Erosion of community diversity and stability by herbivore removal under warming

Climate change has the potential to influence the persistence of ecological communities by altering their stability properties. One of the major drivers of community stability is species diversity, which is itself expected to be altered by climate change in many systems. The extent to which climatic effects on community stability may be buffered by the influence of species interactions on diversity is, however, poorly understood because of a paucity of studies incorporating interactions between abiotic and biotic factors. Here, I report results of a 10-year field experiment, the past 7 years of which have focused on effects of ongoing warming and herbivore removal on diversity and stability within the plant community, where competitive species interactions are mediated by exploitation through herbivory. Across the entire plant community, stability increased with diversity, but both stability and diversity were reduced by herbivore removal, warming and their interaction. Within the most species-rich functional group in the community, forbs, warming reduced species diversity, and both warming and herbivore removal reduced the strength of the relationship between diversity and stability. Species interactions, such as exploitation, may thus buffer communities against destabilizing influences of climate change, and intact populations of large herbivores, in particular, may prove important in maintaining and promoting plant community diversity and stability in a changing climate.     

Habitat complexity facilitates coexistence in a tropical ant community

The role of habitat complexity in the coexistence of ant species is poorly understood. Here, we examine the influence of habitat complexity on coexistence patterns in ant communities of the remote Pacific atoll of Tokelau. The invasive yellow crazy ant, Anoplolepis gracilipes (Smith), exists in high densities on Tokelau, but still coexists with up to seven other epigeic ant species. The size-grain hypothesis (SGH) proposes that as the size of terrestrial walking organisms decreases, the perceived complexity of the environment increases and predicts that: (1) leg length increases allometrically with body size in ants, and (2) coexistence between ant species is facilitated by differential habitat use according to body size. Analysis of morphological variables revealed variation inconsistent with the morphological prediction of the SGH, as leg length increased allometrically with head length only. We also experimentally tested the ability of epigeic ants in the field to discover and dominate food resources in treatments of differing rugosity. A. gracilipes was consistently the first to discover food baits in low rugosity treatments, while smaller ant species were consistently the first to discover food baits in high rugosity treatments. In addition, A. gracilipes dominated food baits in planar treatments, while smaller ant species dominated baits in rugose treatments. We found that the normally predictable outcomes of exploitative competition between A. gracilipes and other ant species were reversed in the high rugosity treatments. Our results support the hypothesis that differential habitat use according to body size provides a mechanism for coexistence with the yellow crazy ant in Tokelau. The SGH may provide a mechanism for coexistence in other ant communities but also in communities of other terrestrial, walking insects that inhabit a complex landscape.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Revisiting projected shifts in the climate envelopes of North American trees using updated general circulation models

Global climate models are constantly being upgraded, but it is often not clear what these changes have on climate change impact projections. We used difference maps to directly compare downscaled projections of temperature and precipitation across North America for two versions (or generations) of three different Atmospheric‐Ocean General Circulation Models (AOGCM)s. We found that AOGCM versions differed in their projections for the end of the current century by up to 4 °C for annual mean temperature and 60% for annual precipitation. To place these changes in an ecological context, we reanalyzed our work on shifts in tree climate envelopes (CEs) using the newer‐generation AOGCM projections. Based on the updated AOGCMs, by the 2071–2100 period, tree CEs shifted up to 2.4 degrees further north or 2.6 degrees further south (depending on the AOGCM) and were about 10% larger in size. Despite considerable differences between versions of a given AOGCM, projections made by the newer version of each AOGCM were in general agreement, suggesting convergence across the three models studied here. Assessing the AOGCM outputs in this way provides insight into the magnitude and importance of change associated with AOGCM upgrades as they continue to evolve through time.     

The future of terrestrial mammals in the Mediterranean basin under climate change

The Mediterranean basin is considered a hotspot of biological diversity with a long history of modification of natural ecosystems by human activities, and is one of the regions that will face extensive changes in climate. For 181 terrestrial mammals (68% of all Mediterranean mammals), we used an ensemble forecasting approach to model the future (approx. 2100) potential distribution under climate change considering five climate change model outputs for two climate scenarios. Overall, a substantial number of Mediterranean mammals will be severely threatened by future climate change, particularly endemic species. Moreover, we found important changes in potential species richness owing to climate change, with some areas (e.g. montane region in central Italy) gaining species, while most of the region will be losing species (mainly Spain and North Africa). Existing protected areas (PAs) will probably be strongly influenced by climate change, with most PAs in Africa, the Middle East and Spain losing a substantial number of species, and those PAs gaining species (e.g. central Italy and southern France) will experience a substantial shift in species composition.     

Fish introductions reveal the temperature dependence of species interactions

A major area of current research is to understand how climate change will impact species interactions and ultimately biodiversity. A variety of environmental conditions are rapidly changing owing to climate warming, and these conditions often affect both the strength and outcome of species interactions. We used fish distributions and replicated fish introductions to investigate environmental conditions influencing the coexistence of two fishes in Swedish lakes: brown trout (Salmo trutta) and pike (Esox lucius). A logistic regression model of brown trout and pike coexistence showed that these species coexist in large lakes (more than 4.5 km²), but not in small, warm lakes (annual air temperature more than 0.9–1.5°C). We then explored how climate change will alter coexistence by substituting climate scenarios for 2091–2100 into our model. The model predicts that brown trout will be extirpated from approximately half of the lakes where they presently coexist with pike and from nearly all 9100 lakes where pike are predicted to invade. Context dependency was critical for understanding pike–brown trout interactions, and, given the widespread occurrence of context-dependent species interactions, this aspect will probably be critical for accurately predicting climate impacts on biodiversity.     

Density-dependent diversification in North American wood warblers

Evidence from both molecular phylogenies and the fossil record suggests that rates of species diversification often decline through time during evolutionary radiations. One proposed explanation for this pattern is ecological opportunity, whereby an initial abundance of resources and lack of potential competitors facilitate rapid diversification. This model predicts density-dependent declines in diversification rates, but has not been formally tested in any species-level radiation. Here we develop a new conceptual framework that distinguishes density dependence from alternative processes that also produce temporally declining diversification, and we demonstrate this approach using a new phylogeny of North American Dendroica wood warblers. We show that explosive lineage accumulation early in the history of this avian radiation is best explained by a density-dependent diversification process. Our results suggest that the tempo of wood warbler diversification was mediated by ecological interactions among species and that lineage and ecological diversification in this group are coupled, as predicted under the ecological opportunity model.     

Predicting changes in Fennoscandian vascular-plant species richness as a result of future climatic change

It is anticipated that future climatic warming following the currently enhanced greenhouse effect will change the distribution limits of many vascular plant species. Using annual accumulated respiration equivalents, calculated from January and July mean temperatures and total annual precipitation, simple presence–absence response surface plots are constructed for 1521 native vascular-plant species in 229 75×75-km grid squares within Fennoscandia. The contemporary occurrences in relation to present-day climate and to predicted changes in climate (and hence annual accumulated respiration equivalents) are used to predict possible future immigrations and extinctions within each grid square. The percentage of potential change in species richness for each grid square is estimated from these predictions. Results from this study suggest a mean increase in species richness per grid square of 26%. Increases in species richness are greatest in the southern parts of the alpine/boreal regions in Fennoscandia. There are ten species that potentially may become extinct in Fennoscandia as a result of predicted climatic warming. Possible conservation strategies to protect such endangered species are outlined.     

Combining a regional climate model with a phytoplankton community model to predict future changes in phytoplankton in lakes

1. Linking a regional climate model (RCM) configured for contemporary atmospheric greenhouse gas concentrations, with a phytoplankton community model (PROTECH) produced realistic simulations of 20 years of recent phytoplankton data from Bassenthwaite Lake, in the North‐West of England. 2. Meteorological drivers were derived from the RCM to represent a future climate scenario involving a 1% per annum compound increase in atmospheric CO₂ concentrations until 2100. Using these drivers, PROTECH was run for another 20 year period representing the last two decades of the 21st century. 3. Comparison of these present and future simulations revealed likely impacts on the current seasonal phytoplankton development. Under future climate conditions, the simulated spring bloom showed an increase in cyanobacteria dominance caused by greater success of Planktothrix. Also, the summer cyanobacteria bloom declined earlier because of nutrient limitation caused by the increased spring growth. Overall productivity in the lake did not change. 4. Analysis showed that these predicted changes were driven by changes in water temperature, which were in turn triggered by the higher air temperatures predicted by the RCM.     

Predicting ecosystem shifts requires new approaches that integrate the effects of climate change across entire systems

Most studies that forecast the ecological consequences of climate change target a single species and a single life stage. Depending on climatic impacts on other life stages and on interacting species, however, the results from simple experiments may not translate into accurate predictions of future ecological change. Research needs to move beyond simple experimental studies and environmental envelope projections for single species towards identifying where ecosystem change is likely to occur and the drivers for this change. For this to happen, we advocate research directions that (i) identify the critical species within the target ecosystem, and the life stage(s) most susceptible to changing conditions and (ii) the key interactions between these species and components of their broader ecosystem. A combined approach using macroecology, experimentally derived data and modelling that incorporates energy budgets in life cycle models may identify critical abiotic conditions that disproportionately alter important ecological processes under forecasted climates.     

Elevated air temperature alters an old-field insect community in a multifactor climate change experiment

To address how multiple, interacting climate drivers may affect plant–insect community associations, we sampled insects that naturally colonized a constructed old‐field plant community grown for over 2 years under simultaneous CO₂, temperature, and water manipulation. Insects were sampled using a combination of sticky traps and vacuum sampling, identified to morphospecies and the insect community with respect to abundance, richness, and evenness quantified. Individuals were assigned to four broad feeding guilds in order to examine potential trophic level effects. Although there were occasional effects of CO₂ and water treatment, the effects of warming on the insect community were large and consistent. Warming significantly increased Order Thysanoptera abundance and reduced overall morphospecies richness and evenness. Nonmetric multidimensional scaling found that only temperature affected insect community composition, while a Sørensen similarity index showed less correspondence in the insect community between temperature treatments compared with CO₂ or soil water treatments. Within the herbivore guild, elevated temperature significantly reduced richness and evenness. Corresponding reductions of diversity measures at higher trophic levels (i.e. parasitoids), along with the finding that herbivore richness was a significant predictor of parasitoid richness, suggest trophic‐level effects within the insect community. When the most abundant species were considered in temperature treatments, a small number of species increased in abundance at elevated temperature, while others declined compared with ambient temperature. Effects of temperature in the dominant insects demonstrated that treatment effects were limited to a relatively small number of morphospecies. Observed effects of elevated CO₂ concentration on whole‐community foliar N concentration did not result in any effect on herbivores, which are probably the most susceptible guild to changes in plant nutritional quality. These results demonstrate that climatic warming may alter certain insect communities via effects on insect species most responsive to a higher temperature, contributing to a change in community structure.     

Olive phenology as a sensitive indicator of future climatic warming in the Mediterranean

Experimental and modelling work suggests a strong dependence of olive flowering date on spring temperatures. Since airborne pollen concentrations reflect the flowering phenology of olive populations within a radius of 50 km, they may be a sensitive regional indicator of climatic warming. We assessed this potential sensitivity with phenology models fitted to flowering dates inferred from maximum airborne pollen data. Of four models tested, a thermal time model gave the best fit for Montpellier, France, and was the most effective at the regional scale, providing reasonable predictions for 10 sites in the western Mediterranean. This model was forced with replicated future temperature simulations for the western Mediterranean from a coupled ocean‐atmosphere general circulation model (GCM). The GCM temperatures rose by 4·5 °C between 1990 and 2099 with a 1% per year increase in greenhouse gases, and modelled flowering date advanced at a rate of 6·2 d per °C. The results indicated that this long‐term regional trend in phenology might be statistically significant as early as 2030, but with marked spatial variation in magnitude, with the calculated flowering date between the 1990s and 2030s advancing by 3–23 d. Future monitoring of airborne olive pollen may therefore provide an early biological indicator of climatic warming in the Mediterranean.     

Functional groups and patterns of organization in North American ant communities: a comparison with Australia

Ant distribution and behavioural dominance is examined at nine sites along an elevational gradient (1400–2600 m) in south eastern Arizona, in order to classify North American species according to a functional group scheme used extensively in Australia. The functional groups are then used as a basis for determining patterns of community structure along the environmental gradient, and for comparing community structure between Australia and North America. Quantitative information on species com- position was obtained from pitfall traps, and patterns of ant abundance at tuna baits were used to determine relative behavioural dominance among taxa. A total of eighty-three species from twenty-eight genera was recorded along the elevational gradient, with site species richness ranging from four (high elevation Douglas fir forest) to thirty-three (mid elevation oak–juniper woodland). There was a strong correlation between ant abundance and richness, which was not an artefact of sampling intensity. The most common ants were species of Forelius, Monomorium, Crematogaster and Pheidole at the three desert sites, species of Formica, Pheidole and Crematogaster at the three woodland sites, and species of Prenolepis and Formica at one forest site. No species were abundant at two other forest sites. The most common species in traps also tended to be the most common species at baits. In terms of behavioural dominance, highly competitive ants included species of Solenopsis, Forelius, Monomorium and Liometopum. Species of Pheidole and Crematogaster tended to be moderately competitive, whereas species of Dory- myrmex, Myrmica, Camponotus and Formica (fusca gp) had low competitive ability. On the basis of these results and on published records of other taxa, North American ants were assigned to functional groups as follows (major taxa only given here): Dominant Dolichoderinae—Forelius, Liome- topum; Subordinate Camponotini—Camponotus; Hot Climate Specialists—Pogonomyrmex, Myrmecocystus; Cold Climate Specialists—Formica (rufa, exsecta and microgyna groups), Leptothorax, Stenamma, Lasius, Prenolepis; Cryptic Species—Smithistruma, Solenopsis (subgenus Diplorhop- trum), Acanthomyops; Opportunists—Formica (fusca group), Myrmica, Paratrechina, Dorymyrmex; Generalized Myr- micinae—Pheidole, Crematogaster, Monomorium; Specialist Predators—no major taxa. Functional group composition varied systematically along the elevation gradient: Dominant Dolichoderinae, Generalized Myrà micinae and Hot Climate Specialists were predominant at desert sites; Generalized Myrmicinae and Opportunists were predominant at woodland sites; and Opportunists and Cold Climate Specialists were predominant at forest sites. These patterns are consistent with published studies from elsewhere in North America. Almost all North American taxa can be matched with what appear to be ecologically equivalent taxa in Australia, and biogeographic patterns of functional group composition are broadly similar across the two continents. The major differences are that Australian ant communities are far richer in species, and are almost always dominated by dolichoderines, particularly species of Iridomyrmex. Generalized myrmicines are subdominant to dolichoderines in Australia, but are the behaviourally dominant ants throughout the warmer parts of North America. In cool-temperate North America, species of Formica (especially rufa and exsecta groups) are behaviourally dominant, as they are throughout the Palearctic. Some major features of the North American fauna can be linked to its poor representation of Dominant Dolichoderinae, including (1) the relatively low degree of physiological, morphological and behavioural specialization of Hot Climate Specialists; (2) behavioural dominance by formicines in cool-temperate habitats; and (3) the susceptibility to invasion by behaviourally dominant species such as the imported fire ant Solenopsis invicta and the Argentine ant Linepithema humile.     

Climate and wildfires in the North American boreal forest

The area burned in the North American boreal forest is controlled by the frequency of mid-tropospheric blocking highs that cause rapid fuel drying. Climate controls the area burned through changing the dynamics of large-scale teleconnection patterns (Pacific Decadal Oscillation/El Niño Southern Oscillation and Arctic Oscillation, PDO/ENSO and AO) that control the frequency of blocking highs over the continent at different time scales. Changes in these teleconnections may be caused by the current global warming. Thus, an increase in temperature alone need not be associated with an increase in area burned in the North American boreal forest. Since the end of the Little Ice Age, the climate has been unusually moist and variable: large fire years have occurred in unusual years, fire frequency has decreased and fire–climate relationships have occurred at interannual to decadal time scales. Prolonged and severe droughts were common in the past and were partly associated with changes in the PDO/ENSO system. Under these conditions, large fire years become common, fire frequency increases and fire–climate relationships occur at decadal to centennial time scales. A suggested return to the drier climate regimes of the past would imply major changes in the temporal dynamics of fire–climate relationships and in area burned, a reduction in the mean age of the forest, and changes in species composition of the North American boreal forest.     

Vulnerability of carbon storage in North American boreal forests to wildfires during the 21st century

The boreal forest contains large reserves of carbon. Across this region, wildfires influence the temporal and spatial dynamics of carbon storage. In this study, we estimate fire emissions and changes in carbon storage for boreal North America over the 21st century. We use a gridded data set developed with a multivariate adaptive regression spline approach to determine how area burned varies each year with changing climatic and fuel moisture conditions. We apply the process‐based Terrestrial Ecosystem Model to evaluate the role of future fire on the carbon dynamics of boreal North America in the context of changing atmospheric carbon dioxide (CO₂) concentration and climate in the A2 and B2 emissions scenarios of the CGCM2 global climate model. Relative to the last decade of the 20th century, decadal total carbon emissions from fire increase by 2.5–4.4 times by 2091–2100, depending on the climate scenario and assumptions about CO₂ fertilization. Larger fire emissions occur with warmer climates or if CO₂ fertilization is assumed to occur. Despite the increases in fire emissions, our simulations indicate that boreal North America will be a carbon sink over the 21st century if CO₂ fertilization is assumed to occur in the future. In contrast, simulations excluding CO₂ fertilization over the same period indicate that the region will change to a carbon source to the atmosphere, with the source being 2.1 times greater under the warmer A2 scenario than the B2 scenario. To improve estimates of wildfire on terrestrial carbon dynamics in boreal North America, future studies should incorporate the role of dynamic vegetation to represent more accurately post‐fire successional processes, incorporate fire severity parameters that change in time and space, account for human influences through increased fire suppression, and integrate the role of other disturbances and their interactions with future fire regime.     

Experimental evidence of reduced diversity of seedlings due to climate modification in a Mediterranean-type community

We are still lacking in experimental evidence of the effects of climate change on the richness of plant species under field conditions. We report a decrease in the species richness of recruited seedlings in a Mediterranean shrubland in experimentally induced drought and warming over 4 consecutive years. Drought decreased the number of emerging seedlings and their respective species richness. Warming also decreased seedling species richness, but it did not affect the number of emerging seedlings. Species that produce fewer recruits are more likely to disappear in drier or warmer scenarios. However, when the effect of induced climate treatment was greatest, the more abundant species in control stands were not necessarily the ones least affected by treatment; in other words, species‐idiosyncratic responses may occur. These results show that demographic processes are sensitive to minor climate changes, with probable consequences on the diversity and structure of the future plant communities.