Plant antagonistic facilitation across environmental gradients: a soil-resource ecosystem engineering model.

Theory questions the persistence of nonreciprocal interactions in which one plant has a positive net effect on a neighbor that, in return, has a negative net impact on its benefactor - a phenomenon known as antagonistic facilitation. We develop a spatially explicit consumer-resource model for belowground plant competition between ecosystem engineers, plants able to mine resources and make them available for any other plant in the community, and exploiters. We use the model to determine in what environmental conditions antagonistic facilitation via soil-resource engineering emerges as an optimal strategy. Antagonistic facilitation emerges in stressful environments where ecosystem engineers' self-benefits from mining resources outweigh the competition with opportunistic neighbors. Among all potential causes of stress considered in the model, the key environmental parameter driving changes in the interaction between plants is the proportion of the resource that becomes readily available for plant consumption in the absence of any mining activity. Our results align with theories of primary succession and the stress gradient hypothesis. However, we find that the total root biomass and its spatial allocation through the root system, often used to measure the sign of the interaction between plants, do not predict facilitation reliably.

Pneumococcus and the stress-gradient hypothesis: A trade-off links R0 and susceptibility to co-colonization across countries.

Modern molecular technologies have revolutionized our understanding of bacterial epidemiology, but reported data across studies and different geographic endemic settings remain under-integrated in common theoretical frameworks. Pneumococcus serotype co-colonization, caused by the polymorphic bacteria Streptococcus pneumoniae, has been increasingly investigated and reported in recent years. While the global genomic diversity and serotype distribution of S. pneumoniae have been well-characterized, there is limited information on how co-colonization patterns vary globally, critical for understanding the evolution and transmission dynamics of the bacteria. Gathering a rich dataset of cross-sectional pneumococcal colonization studies in the literature, we quantified patterns of transmission intensity and co-colonization prevalence variation in children populations across 17 geographic locations. Linking these data to an SIS model with cocolonization under the assumption of quasi-neutrality among multiple interacting strains, our analysis reveals strong patterns of negative co-variation between transmission intensity (R0) and susceptibility to co-colonization (k). In line with expectations from the stress-gradient-hypothesis in ecology (SGH), pneumococcus serotypes appear to compete more in co-colonization in high-transmission settings and compete less in low-transmission settings, a trade-off which ultimately leads to a conserved ratio of single to co-colonization µ=1/(R0-1)k. From the mathematical model's behavior, such conservation suggests preservation of 'stability-diversity-complexity' regimes in coexistence of similar co-colonizing strains. We find no major differences in serotype compositions across studies, pointing to adaptation of the same set of serotypes across variable environments as an explanation for their differential interaction in different transmission settings. Our work highlights that the understanding of transmission patterns of Streptococcus pneumoniae from global scale epidemiological data can benefit from simple analytical approaches that account for quasi-neutrality among strains, co-colonization, as well as variable environmental adaptation.

Positive interactions between corals and damselfish increase coral resistance to temperature stress.

By the century's end, many tropical seas will reach temperatures exceeding most coral species' thermal tolerance on an annual basis. The persistence of corals in these regions will, therefore, depend on their abilities to tolerate recurrent thermal stress. Although ecologists have long recognized that positive interspecific interactions can ameliorate environmental stress to expand the realized niche of plants and animals, coral bleaching studies have largely overlooked how interactions with community members outside of the coral holobiont shape the bleaching response. Here, we subjected a common coral, Pocillopora grandis, to 10 days of thermal stress in aquaria with and without the damselfish Dascyllus flavicaudus (yellowtail dascyllus), which commonly shelter within these corals, to examine how interactions with damselfish impacted coral thermal tolerance. Corals often benefit from nutrients excreted by animals they interact with and prior to thermal stress, corals grown with damselfish showed improved photophysiology (Fv /Fm ) and developed larger endosymbiont populations. When exposed to thermal stress, corals with fish performed as well as control corals maintained at ambient temperatures without fish. In contrast, corals exposed to thermal stress without fish experienced photophysiological impairment, a more than 50% decline in endosymbiont density, and a 36% decrease in tissue protein content. At the end of the experiment, thermal stress caused average calcification rates to decrease by over 80% when damselfish were absent but increase nearly 25% when damselfish were present. Our study indicates that damselfish-derived nutrients can increase coral thermal tolerance and are consistent with the Stress Gradient Hypothesis, which predicts that positive interactions become increasingly important for structuring communities as environmental stress increases. Because warming of just a few degrees can exceed corals' temperature tolerance to trigger bleaching and mortality, positive interactions could play a critical role in maintaining some coral species in warming regions until climate change is aggressively addressed.

Leaf carbon isotope tracks the facilitation pattern of legume shrubs shaped by water availability and species replacement along a large elevation gradient in Trans-Himalayas.

BACKGROUND AND AIMS: Understanding patterns and mechanisms of nurse plant facilitation is important to predict the resilience of arid/semi-arid ecosystems to climate change. We investigate whether water availability and nurse species turnover interact to shape the facilitation pattern of widespread legume shrubs along a large elevation gradient. We also investigate whether leaf δ13C of nurse plants can track the facilitation pattern. METHODS: We measured the relative interaction index (RII) of the number of species within and outside the canopy of two widespread legume shrub species (Caragana gerardiana and Caragana versicolor) alternatively distributed along a large elevation gradient in the Trans-Himalayas. We also assessed the proportional increase of species richness (ISR) at the community level using the paired plot data. To determine site-specific water availability, we measured the leaf δ13C of nurse shrubs and calculated the Thornthwaite moisture index (MI) for each elevation site. KEY RESULTS: Elevational variations in RII, ISR and δ13C were mainly explained by the MI when the effects of soil nitrogen and plant traits (leaf nitrogen and shrub size) were controlled. Variations in RII and ISR across the two nurse species were explained better by δ13C than by smoothly changing climatic factors along elevation. At the transition zone between the upper limit of C. gerardiana (4100 m) and the lower limit of C. versicolor (4200 m), RII and ISR were much higher in C. versicolor than in C. gerardiana under a similar MI. Such an abrupt increase in facilitation induced by nurse species replacement was well tracked by the variation of δ13C. CONCLUSIONS: Water availability and nurse species replacement are crucial to shaping facilitation patterns by legume shrubs along a large elevation gradient in dry mountainous regions, such as the Trans-Himalayas. Turnover in nurse species under global change might significantly alter the pattern of nurse plant facilitation associated with water availability, which can be well tracked by leaf δ13C.

Ecosystem engineering and leaf quality together affect arthropod community structure and diversity on white oak (Quercus alba L.).

Shelter building caterpillars act as ecosystem engineers by creating and maintaining leaf shelters, which are then colonized by other arthropods. Foliage quality has been shown to influence initial colonization by shelter-building caterpillars. However, the effects of plant quality on the interactions between ecosystem engineers and their communities have yet to be studied at the whole plant level. We examined how leaf tying caterpillars, as ecosystem engineers, impact arthropod communities on Quercus alba (white oak), and the modifying effect of foliage quality on these interactions. We removed all leaf tying caterpillars and leaf ties on 35 Q. alba saplings during the season when leaf tying caterpillars were active (June-September), and compared these leaf tie removal trees to 35 control trees whose leaf ties were left intact. Removal of these ecosystem engineers had no impact on overall arthropod species richness, but reduced species diversity, and overall arthropod abundance and that of most guilds, and changed the structure of the arthropod community as the season progressed. There was an increase in plant-level species richness with increasing number of leaf ties, consistent with Habitat Diversity Hypothesis. In turn, total arthropod density, and that of both leaf tying caterpillars and free-feeding caterpillars were affected by foliar tannin and nitrogen concentrations, and leaf water content. The engineering effect was greatest on low quality plants, consistent with the Stress-Gradient Hypothesis. Our results demonstrate that interactions between ecosystem engineering and plant quality together determine community structure of arthropods on Q. alba in Missouri.

Biodiversity promotes ecosystem functioning despite environmental change.

Three decades of research have demonstrated that biodiversity can promote the functioning of ecosystems. Yet, it is unclear whether the positive effects of biodiversity on ecosystem functioning will persist under various types of global environmental change drivers. We conducted a meta-analysis of 46 factorial experiments manipulating both species richness and the environment to test how global change drivers (i.e. warming, drought, nutrient addition or CO2 enrichment) modulated the effect of biodiversity on multiple ecosystem functions across three taxonomic groups (microbes, phytoplankton and plants). We found that biodiversity increased ecosystem functioning in both ambient and manipulated environments, but often not to the same degree. In particular, biodiversity effects on ecosystem functioning were larger in stressful environments induced by global change drivers, indicating that high-diversity communities were more resistant to environmental change. Using a subset of studies, we also found that the positive effects of biodiversity were mainly driven by interspecific complementarity and that these effects increased over time in both ambient and manipulated environments. Our findings support biodiversity conservation as a key strategy for sustainable ecosystem management in the face of global environmental change.

A case for associational resistance: Apparent support for the stress gradient hypothesis varies with study system.

According to the stress gradient hypothesis (SGH), ecological interactions between organisms shift positively as environmental stress increases. In the case of associational resistance, habitat is modified to ameliorate stress, benefitting other organisms. The SGH is contentious due to conflicting evidence and theoretical perspectives, so we adopted a meta-analytic approach to determine if it is widely supported across a variety of contexts, including different kingdoms, ecosystems, habitats, interactions, stressors, and life history stages. We developed an extensive list of Boolean search criteria to search the published ecological literature and successfully detect studies that both directly tested the hypothesis, and those that were relevant but never mentioned it. We found that the SGH is well supported by studies that feature bacteria, plants, terrestrial ecosystems, interspecific negative interactions, adults, survival instead of growth or reproduction, and drought, fire, and nutrient stress. We conclude that the SGH is indeed a broadly relevant ecological hypothesis that is currently held back by cross-disciplinary communication barriers. More SGH research is needed beyond the scope of interspecific plant competition, and more SGH research should feature multifactor stress. There remains a need to account for positive interactions in scientific pursuits, such as associational resistance in tests of the SGH.

Toxicity drives facilitation between 4 bacterial species.

Competition between microbes is extremely common, with many investing in mechanisms to harm other strains and species. Yet positive interactions between species have also been documented. What makes species help or harm each other is currently unclear. Here, we studied the interactions between 4 bacterial species capable of degrading metal working fluids (MWF), an industrial coolant and lubricant, which contains growth substrates as well as toxic biocides. We were surprised to find only positive or neutral interactions between the 4 species. Using mathematical modeling and further experiments, we show that positive interactions in this community were likely due to the toxicity of MWF, whereby each species' detoxification benefited the others by facilitating their survival, such that they could grow and degrade MWF better when together. The addition of nutrients, the reduction of toxicity, or the addition of more species instead resulted in competitive behavior. Our work provides support to the stress gradient hypothesis by showing how harsh, toxic environments can strongly favor facilitation between microbial species and mask underlying competitive interactions.

Conditional love? Co-occurrence patterns of drought-sensitive species in European grasslands are consistent with the stress-gradient hypothesis.

AIM: The stress-gradient hypothesis (SGH) postulates that species interactions shift from negative to positive with increasing abiotic stress. Interactions between species are increasingly being recognized as important drivers of species distributions, but it is still unclear whether stress-induced changes in interactions affect continental-to-global scale species distributions. Here, we tested whether associations of vascular plant species in dry grasslands in Europe follow the SGH along a climatic water deficit (CWD) gradient across the continent. LOCATION: Dry grasslands in Europe. TIME PERIOD: Present. MAJOR TAXA STUDIED: Vascular plants. METHODS: We built a context-dependent joint species distribution model (JSDM) to estimate the residual associations (i.e., associations that are not explained by the abiotic environment) of 161 plant species as a function of the CWD based on community data from 8,660 vegetation plots. We evaluated changes in residual associations between species for pairs and on the community level, and we compared responses for groups of species with different drought tolerances. RESULTS: We found contrasting shifts in associations for drought-sensitive and drought-tolerant species. For drought-sensitive species, 21% of the pairwise associations became more positive with increasing CWD, whereas 17% became more negative. In contrast, only 17% of the pairwise associations involving drought-tolerant species became more positive, whereas 27% became more negative in areas with a high CWD. Additionally, the incidence of positive associations increased with drought for drought-sensitive species and decreased for drought-tolerant species. MAIN CONCLUSIONS: We found that associations of drought-sensitive plant species became more positive with drought, in line with the SGH. In contrast, associations of drought-tolerant species became more negative. Additionally, changes in associations of single species pairs were highly variable. Our results indicate that stress-modulated species associations might influence the distribution of species over large geographical extents, thus leading to unexpected responses under climate change through shifts in species associations.

Plant-plant co-occurrences under a complex land-use gradient in a temperate forest.

Land-use generates multiple stress factors, and we need to understand their effects on plant-plant interactions to predict the consequences of land-use intensification. The stress-gradient hypothesis predicts that the relative strength of positive and negative interactions changes inversely under increasing environmental stress. However, the outcome of interactions also depends on stress factor's complexity, the scale of analysis, and the role of functional traits in structuring the community. We evaluated plant-plant co-occurrences in a temperate forest, aiming to identify changes in pairwise and network metrics under increasing silvopastoral use intensity. Proportionally, positive co-occurrences were more frequent under high than low use, while negative co-occurrences were more frequent under low than high. Networks of negative co-occurrences showed higher centralization under low use, while networks of positive co-occurrences showed lower modularity and higher centralization under high use. We found a partial relationship between co-occurrences and key functional traits expected to mediate facilitation and competition processes. Our results shows that the stress-gradient hypothesis predicts changes in spatial co-occurrences even when two stress factors interact in a complex way. Networks of negative co-occurrences showed a hierarchical effect of dominant species under low use intensity. But positive co-occurrence network structure partially presented the characteristics expected if the facilitation was an important mechanism characterizing the community under high disturbance intensity. The partial relationship between functional traits and co-occurrences may indicate that other factors besides biotic interactions may be structuring the observed negative spatial associations in temperate Patagonian forests.

Earlier onset of flowering and increased reproductive allocation of an annual invasive plant in the north of its novel range.

BACKGROUND AND AIMS: It remains unclear whether invasive species can maintain both high biomass and reproductive output across their invaded range. Along latitudinal gradients, allocation theory predicts that faster flowering onset at high latitudes results in maturation at smaller size and thus reduced reproductive output. For annual invasive plants, more favourable environmental conditions at low latitudes probably result in stronger competition of co-occurring species, potentially driving selection for higher investment in vegetative biomass, while harsher climatic conditions and associated reproductive uncertainty at higher latitudes could reduce selection for vegetative biomass and increased selection for high reproductive investment (stress-gradient hypothesis). Combined, these drivers could result in increased or constant reproductive allocation with increasing latitude. METHODS: We quantified life-history traits in the invasive annual plant Impatiens glandulifera along a latitudinal gradient in Europe. By growing two successive glasshouse generations, we assessed genetic differentiation in vegetative growth and reproductive output across six populations, and tested whether onset of flowering drives this divergence. KEY RESULTS: Trait variation was mainly caused by genetic differentiation. As expected, flowering onset was progressively earlier in populations from higher latitudes. Plant height and vegetative biomass also decreased in populations from higher latitudes, as predicted by allocation theory, but their variation was independent of the variation in flowering onset. Reproductive output remained constant across latitudes, resulting in increased reproductive allocation towards higher latitudes, supporting the stress-gradient hypothesis. We also observed trait genetic differentiation among populations that was independent of latitude. CONCLUSIONS: We show that an annual invasive plant evolved several life-history traits across its invaded range in ~150 years. The evolution of vegetative and reproductive traits seems unconstrained by evolution of flowering onset. This genetic decoupling between vegetative and reproductive traits possibly contributes to the invasion success of this species.

Interactive range-limit theory (iRLT): An extension for predicting range shifts.

A central theme of range-limit theory (RLT) posits that abiotic factors form high-latitude/altitude limits, whereas biotic interactions create lower limits. This hypothesis, often credited to Charles Darwin, is a pattern widely assumed to occur in nature. However, abiotic factors can impose constraints on both limits and there is scant evidence to support the latter prediction. Deviations from these predictions may arise from correlations between abiotic factors and biotic interactions, as a lack of data to evaluate the hypothesis, or be an artifact of scale. Combining two tenets of ecology-niche theory and predator-prey theory-provides an opportunity to understand how biotic interactions influence range limits and how this varies by trophic level. We propose an expansion of RLT, interactive RLT (iRLT), to understand the causes of range limits and predict range shifts. Incorporating the main predictions of Darwin's hypothesis, iRLT hypothesizes that abiotic and biotic factors can interact to impact both limits of a species' range. We summarize current thinking on range limits and perform an integrative review to evaluate support for iRLT and trophic differences along range margins, surveying the mammal community along the boreal-temperate and forest-tundra ecotones of North America. Our review suggests that range-limit dynamics are more nuanced and interactive than classically predicted by RLT. Many (57 of 70) studies indicate that biotic factors can ameliorate harsh climatic conditions along high-latitude/altitude limits. Conversely, abiotic factors can also mediate biotic interactions along low-latitude/altitude limits (44 of 68 studies). Both scenarios facilitate range expansion, contraction or stability depending on the strength and the direction of the abiotic or biotic factors. As predicted, biotic interactions most often occurred along lower limits, yet there were trophic differences. Carnivores were only limited by competitive interactions (n = 25), whereas herbivores were more influenced by predation and parasitism (77%; 55 of 71 studies). We highlight how these differences may create divergent range patterns along lower limits. We conclude by (a) summarizing iRLT; (b) contrasting how our model system and others fit this hypothesis and (c) suggesting future directions for evaluating iRLT.

Plant interactions balance under biotic and abiotic stressors: the importance of herbivory in semi-arid ecosystems.

Biotic and abiotic stressors commonly co-occur in plant communities and influence interactions between plants. However, their combined effects on plant interactions have not been widely studied and are still unclear. Here, we assessed the balance of interactions between neighboring plants along a grazing gradient and under two water regimes. We conducted a three-year-field experiment in semi-arid central Argentina with transplants of the dominant palatable grass Piptochaetium napostaense growing in Baccharis ulicina and open microsites across a gradient of grazing pressure. Additionally, we established a water addition treatment along that gradient. We recorded herbivory, size, and fecundity of P. napostaense. During the first two years, P. napostaense was consumed less and was larger below Baccharis than in the open. These differences were greatest under high grazing pressure. Differences in fecundity between microsites were only detected under high and medium grazing pressure in the first two years. In the third year, Baccharis lost their leaves for unclear reasons and provided poor herbivory protection; hence, P. napostaense plants in Baccharis were larger than those in the open only under medium and low grazing pressure, and there were no differences in fecundity between microsites under any grazing condition. Water additions exerted no effect on plant interactions. The balance of interactions changed from positive under heavy grazing to neutral at low and no grazing and water availability did not alter that balance. We conclude that herbivore pressure is an important driver of the balance of plant interactions in semi-arid environments.

Community Response to Extreme Drought (CRED): a framework for drought-induced shifts in plant-plant interactions.

Contents Summary 52 I. Introduction 52 II. The Community Response to Extreme Drought (CRED) framework 55 III. Post-drought rewetting rates: system and community recovery 61 IV. Site-specific characteristics influencing community resistance and resilience 63 V. Conclusions 64 Acknowledgements 65 References 66 SUMMARY: As climate changes, many regions of the world are projected to experience more intense droughts, which can drive changes in plant community composition through a variety of mechanisms. During drought, community composition can respond directly to resource limitation, but biotic interactions modify the availability of these resources. Here, we develop the Community Response to Extreme Drought framework (CRED), which organizes the temporal progression of mechanisms and plant-plant interactions that may lead to community changes during and after a drought. The CRED framework applies some principles of the stress gradient hypothesis (SGH), which proposes that the balance between competition and facilitation changes with increasing stress. The CRED framework suggests that net biotic interactions (NBI), the relative frequency and intensity of facilitative (+) and competitive (-) interactions between plants, will change temporally, becoming more positive under increasing drought stress and more negative as drought stress decreases. Furthermore, we suggest that rewetting rates affect the rate of resource amelioration, specifically water and nitrogen, altering productivity responses and the intensity and importance of NBI, all of which will influence drought-induced compositional changes. System-specific variables and the intensity of drought influence the strength of these interactions, and ultimately the system's resistance and resilience to drought.

Digestive mutualism in a pitcher plant supports the monotonic rather than hump-shaped stress-gradient hypothesis model.

The stress-gradient hypothesis (SGH) predicts that the strength and frequency of facilitative interactions increase monotonically with increasing environmental stress, but some empirical studies have found this decrease at extreme stress levels, suggesting a hump-shaped SGH instead. However, empirical studies of the SGH are often hindered by confounding resource and non-resource stress gradients. Nepenthes pitcher plants trap animal prey using modified-leaf pitfall traps which are also inhabited by organisms known as inquilines. Inquilines may assist pitchers in the digestion of trapped prey. This interaction is known as a digestive mutualism and is both mutualistic and facilitative by definition. Inquiline species may also facilitate each other via processing chain commensalisms. We used in vitro experiments to examine the isolated effect of resource stress on the outcomes of two facilitative interactions: (i) digestive mutualism-facilitation of pitcher nutrient sequestration by two inquiline dipteran larvae, culicids and phorids; (ii) processing chain commensalism-facilitation between these two inquiline taxa. The net nutritional benefit of phorids on N. gracilis was found to conform more to a monotonic rather than hump-shaped SGH model. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were weak. These findings provide compelling evidence that changes in facilitation along an isolated resource stress gradient conform to the predictions of the monotonic SGH model rather than that of the revised hump-shaped model, and highlight the importance of isolating stress gradients in empirical tests of the SGH.

Geographic location, local environment, and individual size mediate the effects of climate warming and neighbors on a benefactor plant.

Predictions of plant responses to global warming frequently ignore biotic interactions and intraspecific variation across geographical ranges. Benefactor species play an important role in plant communities by protecting other taxa from harsh environments, but the combined effects of warming and beneficiary species on their performance have been largely unexamined. We analyzed the joint effects of elevated temperature and neighbor removal on the benefactor plant Silene acaulis, in factorial experiments near its low- and high-latitude range limits in Europe. We recorded growth, probability of reproduction and fruit set during 3 years. The effects of enhanced temperature were positive near the northern limit and negative in the south for some performance measures. This pattern was stronger in the presence of neighbors, possibly due to differential thermal tolerances between S. acaulis and beneficiary species in each location. Neighbors generally had a negative or null impact on S. acaulis, in agreement with previous reviews of overall effects of plant-plant interactions on benefactors. However, small S. acaulis individuals in the north showed higher growth when surrounded by neighbors. Finally, the local habitat within each location influenced some effects of experimental treatments. Overall, we show that plant responses to rising temperatures may strongly depend on their position within the geographic range, and on species interactions. Our results also highlight the need to consider features of the interacting taxa, such as whether they are benefactor species, as well as local-scale environmental variation, to predict the joint effects of global warming and biotic interactions on species and communities.

Evolutionary Responses to Conditionality in Species Interactions across Environmental Gradients.

The outcomes of many species interactions are conditional on the environments in which they occur. Often, interactions grade from being more positive under stressful or low-resource conditions to more antagonistic or neutral under benign conditions. Here, we take predictions about two well-supported ecological theories on conditionality-limiting resource models and the stress-gradient hypothesis-and combine them with those from the geographic mosaic theory of coevolution (GMTC) to generate predictions for systematic patterns of adaptation and coadaptation between partners along abiotic gradients. When interactions become more positive in stressful environments, mutations that increase fitness in one partner may also increase fitness in the other; because fitnesses are aligned, selection should favor greater mutualistic adaptation and coadaptation between interacting species in stressful ends of environmental gradients. As a corollary, in benign environments antagonistic coadaptation could result in Red Queen or arms-race dynamics or the reduction of antagonism through character displacement and niche partitioning. Here, we distinguish between generally mutualistic or antagonistic adaptation (i.e., mutations in one partner that have similar effects across multiple populations of the other) and specific adaptations to sympatric partners (local adaptation), which can occur either alone or simultaneously. We then outline the kinds of data required to test these predictions, develop experimental designs and statistical methods, and demonstrate these using simulations based on GMTC models. Our methods can be applied to a range of conditional outcomes and may also be useful in assisted translocation approaches in the face of climate change.

Facilitation costs and benefits function simultaneously on stress gradients for animals.

Understanding the variation in species interactions along environmental stress gradients is crucial for making robust ecological predictions about community responses to changing environmental conditions. The facilitation-competition framework has provided a strong basis for predictions (e.g. the stress-gradient hypothesis, SGH), yet the mechanisms behind patterns in animal interactions on stress gradients are poorly explored in particular for mobile animals. Here, we proposed a conceptual framework modelling changes in facilitation costs and benefits along stress gradients and experimentally tested this framework by measuring fitness outcomes of benefactor-beneficiary interactions across resource quality levels. Three arthropod consumer models from a broad array of environmental conditions were used including aquatic detritivores, potato moths and rainforest carrion beetles. We detected a shift to more positive interactions at increasing levels of stress thereby supporting the application of the SGH to mobile animals. While most benefactors paid no significant cost of facilitation, an increase in potato moth beneficiary's growth at high resource stress triggered costs for benefactors. This study is the first to experimentally show that both costs and benefits function simultaneously on stress gradients for animals. The proposed conceptual framework could guide future studies examining species interaction outcomes for both animals and plants in an increasingly stressed world.

Reduced competition enhances community temporal stability under conditions of increasing environmental stress.

The stress-gradient hypothesis predicts a switch from competition to facilitation, under increasing environmental stress. However, it is unclear how important is the change in competition-facilitation balance (i.e., the net outcome of plant-plant interactions) along the stress gradient in the regulation of community temporal stability (i.e., the inverse of temporal variability in total biomass). Increasing environmental stress may enhance community temporal stability by reduced competition or eventually by leading to facilitative interactions between the dominant and subordinate species. Here, we present the results of a 5-yr mesocosm experiment that demonstrates the effects of interspecific interactions on the temporal stability of a riparian community across different drought-stress scenarios. We constructed artificial communities of dominant species (Carex elata) and three subordinate species to simulate the independent effects of environmental stress and interspecific interactions. Using removal of the dominant species, we evaluated the interplay of various mechanisms regulating the temporal stability of the subordinate species (competition-facilitation balance, species asynchrony, and dominant species stability). By simultaneous testing of these stabilizing mechanisms, we show their importance differs depending on environmental variability and harshness. The predominant role is taken by species asynchrony in a seasonally dry environment, whereas in a permanently dry environment, the importance of reduced competition increases. Reduced competition was stabilizing, in particular through increased total community biomass, whereas species asynchrony increased total community biomass and decreased biomass variation. These results suggest experiments and simulations that exclude interspecific interactions may not offer realistic predictions of the effects of changing hydrological regimes on ecosystem functioning.

Effects of biotic interactions on tropical tree performance depend on abiotic conditions.

Predicting biotic responses to environmental change requires understanding the joint effects of abiotic conditions and biotic interactions on community dynamics. One major challenge is to separate the potentially confounding effects of abiotic environmental variation and local biotic interactions on individual performance. The stress gradient hypothesis (SGH) addresses this issue directly by predicting that the effects of biotic interactions on performance become more positive as the abiotic environment becomes more stressful. It is unclear, however, how the predictions of the SGH apply to plants of differing functional strategies in diverse communities. We asked (1) how the effect of crowding on performance (growth and survival) of trees varies across a precipitation gradient, and (2) how functional strategies (as measured by two key traits: wood density and leaf mass per area, LMA) mediate average demographic rates and responses to crowding across the gradient. We built trait-based neighborhood models of growth and survival across a regional precipitation gradient where increasing precipitation is associated with reduced abiotic stress. In total, our dataset comprised ~170,000 individual trees belonging to 252 species. The effect of crowding on tree performance varied across the gradient; crowding negatively affected growth across plots and positively affected survival in the wettest plot. Functional traits mediated average demographic rates across the gradient, but we did not find clear evidence that the strength of these responses depends on species' traits. Our study lends support to the SGH and demonstrates how a trait-based perspective can advance these concepts by linking the diversity of species interactions with functional variation across abiotic gradients.