Stress Gradient Hypothesis Research Articles

A size-gradient hypothesis for alpine treeline ecotones

Research on the stress gradient hypothesis recognizes that positive (i.e. facilitative) and negative (i.e. competitive) plant interactions change in intensity and effect relative to abiotic stress experienced on a gradient. Motivated by observations of alpine treeline ecotones, we suggest that this switch in interaction could operate along a gradient of relative size of individual plants. We propose that as neighbors increase in size relative to a focal plant they improve the environment for that plant up to a critical point. After this critical point is surpassed, however, increasing relative size of neighbors will degrade the environment such that the net interaction intensity becomes negative. We developed a conceptual (not site or species specific) individual based model to simulate a single species with recruitment, growth, and mortality dependent on the environment mediated by the relative size of neighbors. Growth and size form a feedback. Simulation results show that the size gradient model produces metrics similar to that of a stress gradient model. Visualizations reveal that the size gradient model produces spatial patterns that are similar to the complex ones observed at alpine treelines. Size-mediated interaction could be a mechanism of the stress gradient hypothesis or it could operate independent of abiotic stress.

Phosphorus availability and microbial community in the rhizosphere of intercropped cereal and legume along a P-fertilizer gradient

Positive below-ground interactions (facilitation) should be more pronounced when resources limit crop growth, according to the stress-gradient hypothesis. Our aim was to test this hypothesis for intercropped durum wheat and faba bean along a P-fertilizer gradient. A field experiment was conducted in a long-term P-fertilizer trial with three rates of P-fertilization (No, Low and High P). Microbial biomass was assessed by chloroform fumigation-extraction. Quantitative PCR was applied to evaluate the abundance of relevant microbial groups. Phosphorus availability and microbial biomass systematically increased in the rhizosphere compared to bulk soil. P-fertilization resulted in higher abundance of targeted bacterial phyla, whole bacterial and fungal communities, and depressed mycorrhizal colonization of durum wheat, but not faba bean. Microbial biomass carbon significantly increased in the rhizosphere only in P-fertilized treatments, pointing to P limitation of microbial communities. Intercropping yielded a significant effect on rhizosphere microbial properties only at High P. Microbial biomass P increased in the rhizosphere of intercropped faba bean only at No P level, and was thus the sole finding supporting the stress-gradient hypothesis. P-fertilization was the main driver of microbial communities in this field trial, and P-fertilizer application modulated the species-specific effect in the intercrop. Plant performance did not validate the stress-gradient hypothesis as positive plant-plant interactions occurred regardless of the level of P-fertilization.

Enhanced facilitation at the extreme end of the aridity gradient in the Atacama Desert: a community-level approach.

Plant facilitation is now recognized as an important process in severe environments. However, there is still no agreement on how facilitation changes as conditions become increasingly severe. The classic stress gradient hypothesis (SGH) predicts a monotonic increase in facilitation, which rises in frequency as conditions approach the extreme end of the environmental gradient. However, few studies have evaluated the validity of the SGH at the community level, the level at which it was formulated. Moreover, few studies have tested the SGH at either extreme of the gradient, and very few have excluded the effect of livestock on community response to stress. In line with the SGH, we hypothesized that several spatial pattern summary statistics would change monotonically from the least to the most arid sites, indicating increasingly aggregated patterns. In this study, we performed an evaluation of the SGH both within communities of shrub species and across a large portion of the Atacama Desert, and we isolated the abiotic component of the SGH. Our environmental gradient covered an extreme aridity gradient (< 20-130 mm annual precipitation). To perform point pattern analysis, we established 13 sites with environmental conditions representing four distinct levels of this gradient. Further, we conducted species co-occurrence analyses at 19 sites along the gradient. Both sets of analyses showed stronger positive spatial associations among plants at the most extreme end of the gradient. This was true regardless of whether we included all individuals, only small individuals located around large ones, or individuals in species pairs. Moreover, species tended to show greater co-occurrence as environmental severity increased. This increase in aggregation in the plant community seems to correlate with an increase in the strength of positive interspecific interactions, rather than greater clustering within each species. These monotonic increases in species co-occurrence and spatial association in more severe environments are consistent with some of the predictions of SGH, and collectively these results suggest that as the climate becomes more arid, positive species pairs interactions tend to be prevalent in the community.

Intraspecific interactions shift from competitive to facilitative across a low to high disturbance gradient in a salt marsh

Disturbances are ubiquitous among salt marshes, and disturbances such as litter removal may form stressful environmental conditions. The stress-gradient hypothesis (SGH) predicts that the relative importance of facilitation and competition between species will vary inversely to each other across abiotic stress gradients. However, how intraspecific interactions vary across disturbance levels, and whether they follow the SGH has not been investigated. To test the SGH with intraspecific interactions within Suaeda salsa individuals, we assessed individual survival in established fully mapped salt marsh plots with two treatments, disturbed, litter removal and control, undisturbed plots. Recently developed spatial statistics were applied to distinguish between random processes, intraspecific facilitation, scramble competition, and contest competition underlying the spatial patterns at different spatial scales, growth stages, and disturbance levels. We found evidence that intraspecific interactions among S. salsa individuals across disturbance levels tended to support the SGH, but this support depended on time and space. Intraspecific interactions were more likely to show positive density dependence in disturbed plots, particularly at the seedling period and when individuals were separated by < 9 cm. However, positive density dependence was not detected in undisturbed plots. The protective effects of litter on seedlings, which were lost in undisturbed sites, may have been at least partly simulated at high seedling densities. Mortality in disturbed sites was not random, and facilitation had a more important influence than competition on population dynamics in high disturbance sites. However, the influence of competition on population dynamics, specifically scramble competition was also clearly important.

Arbuscular mycorrhizal fungi in changing mountain grassland ecosystems: a challenge for research

Even though arbuscular mycorrhizal fungi (AMF) are present from foothills to all alpine habitats, research on their role in mountain ecosystems remains incomplete. Here we provide a literature review of the ecology and functioning of AMF in mountain ecosystems, as well as their response to global change. We investigated how AM fungal abundance, community composition, and fungal traits are studied under field conditions and are affected by altitude, habitat patchiness, succession, host identity, seasonality, and interaction with other living organisms. The effects of climate change, nutrient enrichment, land-use change, and their interactions are also addressed in this review, as well as the possible applications of AMF for grassland restoration. We show that altitudinal effects on AMF are blurred by other environmental gradients and by host-plant identity. The benefits to plants and possible facilitation effects by AMF in mountain ecosystems have not yet been identified. Based on the stress-gradient hypothesis, the symbiosis between plants and AMF should become more mutualistic with increasing environmental stress. We propose a working hypothesis for the functioning of the AMF–plant interaction along altitudinal gradients by grading it along the mutualism–parasitism continuum, considering current and global climate and environmental change conditions. We conclude by suggesting further research directions.

Tree species richness induces strong intraspecific variability of beech (Fagus sylvatica) leaf traits and alleviates edaphic stress

Manipulating stand composition is an important management tool that foresters can use to affect the nature of forests and ecosystem processes. In mixed stands, interspecific interactions among trees can cause changes in tree performances. Nevertheless, these interactions are context dependent (cf. stress-gradient hypothesis, SGH). We thus investigated how intraspecific functional changes in leaf trait (19 traits) of European beech (Fagus sylvatica) were influenced by stand composition. We compared pure beech stands with four mixed stands containing from one to three additional tree species along a gradient of edaphic stress (gradient of soil water-holding capacity and rooting depth). First, we demonstrated that stand composition induced strong intraspecific leaf trait variation in beech for LDMC, LMA, phenolic compounds, leaf pH and magnesium concentration, suggesting higher nutrient acquisition by more diverse stands. Nevertheless, these results were modulated by edaphic stress. Mixed stands only conferred an advantage in relatively stressed sites (luvisol and leptosol). Besides, the addition of oak to beech stands had unexpected negative effects in sites with less severe stress (cambisol) as indicated by the null or positive LogRR of LMA, LDMC and phenolics. This study found that stand composition is an important though often-overlooked driver of intraspecific variability in leaf quality, and potentially reflects changes in beech tree physiology and productivity. Our results also suggest that positive interactions prevail in sites with stressful conditions. Such validation of the SGH is rare in natural or managed mature forests. Lastly, we strongly recommend that forest managers consider stand composition and abiotic factors when implementing forest growth models to improve their yield predictions.

Phylogenetic Relatedness Influences Plant Interspecific Interactions Across Stress Levels in Coastal Ecosystems: a Meta-Analysis

Positive and negative interactions can occur simultaneously between plant species. According to the stress gradient hypothesis (SGH), species interactions shift towards more facilitative interactions or reductions in competition with increasing stress, whereas debate continues over whether evolutionary history influences the strength of species interactions. However, few studies have investigated the effects of phylogenetic relatedness (i.e., the sum of branch lengths separating species on a phylogeny) on the outcomes of interspecific interactions across stress levels. Therefore, we conducted a Bayesian meta-analysis on data collected from publications on plant interactions within coastal ecosystems in order to investigate the effects of phylogenetic relatedness on interspecific interactions across different stress levels. These analyses showed the effect sizes of species interactions on survival and growth to increase with stress increment, supporting the SGH in coastal ecosystems. However, phylogenetic relatedness did not lead to these differences of interspecific interactions between low and high stress. We found that species interactions affecting plant survival were not significantly influenced by phylogenetic relatedness; however, when evolutionary relationships of target and neighbor species were more phylogenetically distant, their interactions were more likely to facilitate growth of target species. Furthermore, the effect of the interaction between phylogenetic distance and stress on species interactions was negative. This suggests the observed net effects of phylogenetically distant neighbor species on target species were not due to true facilitation but reductions in competition when moving from low stress to high stress environments. According to these results, phylogenetic relatedness should be considered in choosing species for restoration of coastal ecosystem plant communities. Specifically, increasing the phylogenetic breadth of the assemblage is more likely to include species that have evolved to reduce stress on surrounding species through modification of the environment.

Is biotic resistance to invaders dependent upon local environmental conditions or primary productivity? A meta-analysis

Biotic resistance is an important component to limiting the spread of invasive plants into native communities. However, under certain conditions native plants may also facilitate invasion, a process identified as biotic assistance. Identifying the conditions that associate with resistance and assistance is needed to better understand the factors driving species invasion.Substantial theory and empirical work suggests net effects of neighbors may be dependent upon habitat productivity and environmental conditions. How that applies to the interaction among the resident community and the invaders is largely untested. Here we compiled data from 23 articles, which had experimentally determined the strength of biotic resistance and/or assistance. We then combined these data with remote-sensing estimates of productivity, precipitation and temperature at each study site. Using standard meta-analytical techniques we determined the overall effect resident communities had on the emergence, growth, reproduction and survival of non-native invaders. Further, we tested whether the interaction between resident communities and invasive species was influenced by primary productivity, temperature and precipitation.Across all sites, we found broad support for biotic resistance, while evidence for biotic assistance was rare. However, we found the relative magnitude of biotic resistance on invaders increased with temperature or precipitation; a pattern consistent with the stress gradient hypothesis. In contrast we found no evidence that the strength of biotic resistance varied as a function of primary productivity. Further evaluation of the relationship between productivity and environmental conditions on the direction and strength of the effect of resident species on invaders may help predict invasion establishment and success. Understanding or predicting the susceptibility of communities to invasion may help prioritize management efforts.

Evolution of resource use along a gradient of stress leads to increased facilitation

The stress‐gradient hypothesis (SGH) posits that the relative importance of facilitative interactions versus negative interactions increases as levels of abiotic stress increase. Originally formulated in empirical studies of plant populations, in recent years the SGH has been found to describe how interactions change in response to stress in a wide range of species including algae, mussels and moths. However, there has been little theory attempting to predict patterns from first principles in relation to different types of interactions. Here, we use mathematical models of microbial populations to investigate whether patterns consistent with the SGH arise when species interact through resource use and allelopathy. Evolution alters the degree to which competition for resource use versus facilitation (cross‐feeding) occurs. Our results are consistent with the SGH; species interactions evolve to be more facilitative as average stress intensifies. This occurs because at greater stress the species evolve to become specialists on either of the two resources thereby decreasing overlap in resource use and increasing facilitation through cross‐feeding. In addition, the production of toxic allelopathic compounds decreases as stress intensifies due to density‐dependent effects. Our results suggest that the SGH could arise through fundamental interactions that are common to many organisms and therefore that the SGH could be a more widespread phenomenon than previously recognised.

Reversal of intraspecific interactions by an ecosystem engineer leads to variable seedling success along a stress gradient

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 543:163-171 (2016) - DOI: https://doi.org/10.3354/meps11561 Reversal of intraspecific interactions by an ecosystem engineer leads to variable seedling success along a stress gradient S. Yang1,2,*, J. HilleRisLambers1, J. L. Ruesink1 1Department of Biology, University of Washington, Box 351800, Seattle, WA 98195-1800, USA 2Present address: Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, WA 98221, USA *Corresponding author: sylvia.yang@wwu.edu ABSTRACT: By definition, ecosystem engineers can have large effects on resource flows and habitat structure, with impacts on other organisms ranging from facilitative to inhibitory. The stress gradient hypothesis posits that facilitative interactions predominate in conditions of environmental stress and switch to competitive interactions in benign environments. We conducted an empirical test of the stress gradient hypothesis for a marine ecosystem engineer, eelgrass Zostera marina L., across a hydrodynamic gradient. Eelgrass meadows perform important functions in coastal ecosystems and have experienced contemporary declines worldwide with variable recruitment success. In this study, we first determine whether eelgrass modified the local hydrodynamic conditions and then how the presence of an adult eelgrass canopy affected the plants’ own relative seedling success over the hydrodynamic gradient. Eelgrass reduced water motion significantly, which provided a positive feedback on survival and growth for conspecific seedlings, but only at the stressful end of the hydrodynamic gradient. By contrast, adult conspecifics negatively impacted seedling recruitment and growth in calmer environments, presumably due to intraspecific competition. Thus, eelgrass ecosystem engineering does not always facilitate the plant’s own performance, and our results support the stress gradient hypothesis as an explanation for this context dependence. The balance of these complex interactions may provide a mechanism for delayed or failed recovery of eelgrass in habitats exposed to high hydrodynamic exposure. KEY WORDS: Context dependence · Density dependence · Habitat modification · Seagrass · Stress gradient hypothesis Full text in pdf format PreviousNextCite this article as: Yang S, HilleRisLambers J, Ruesink JL (2016) Reversal of intraspecific interactions by an ecosystem engineer leads to variable seedling success along a stress gradient. Mar Ecol Prog Ser 543:163-171. https://doi.org/10.3354/meps11561 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 543. Online publication date: February 03, 2016 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2016 Inter-Research.

A Review of Processes Behind Diversity—Productivity Relationships in Forests

Through complementarity interactions, mixed-species forests can be more productive than monocultures, and this effect can increase with tree-species richness. However, this is not always the case. This review examines the processes and stand structural attributes that can influence diversity-productivity relationships (DPRs); how they influence resource availability, resource uptake, and resource-use efficiency; and also describes some important differences between tree-diversity versus grassland-diversity experiments. The size of the complementarity effects caused by these processes and stand structures varies along spatial and temporal gradients in resource availability and climate. These spatial and temporal dynamics have now been examined in many studies, and the general patterns are summarized using a simple framework; complementarity is predicted to increase as the availability of resource “X” declines (or climatic condition X becomes harsher) if the species interactions improve the availability, uptake, or use efficiency of resource X (or interactions improve climatic condition X). Importantly, this framework differs from the stress-gradient hypothesis to account for a wider range of inter-specific plant interactions (not only facilitation) by considering contrasting methods used to quantify species interactions while accounting for stand structure. In addition, complementarity (as opposed to facilitation) for a given species combination can increase as growing conditions improve in forests, contrary to predictions of the stress-gradient hypothesis with regards to facilitation. This review indicates that while the effects of tree-species diversity on growth and other forest functions are now receiving a lot of attention, far less is known about the effects of structural diversity on growth or forest functioning. Direct measurements of the processes, as opposed to focusing mainly on growth responses, could greatly contribute to our understanding of structural diversity effects.

The Effects of Established Trees on Woody Regeneration during Secondary Succession in Tropical Dry Forests

AbstractUnderstanding the mechanisms controlling secondary succession in tropical dry forests is important for the conservation and restoration of this highly threatened biome. Canopy‐forming trees in tropical forests strongly influence later stages of succession through their effect on woody plant regeneration. In dry forests, this may be complex given the seasonal interplay of water and light limitations. We reviewed observational and experimental studies to assess (1) the relative importance of positive and negative effects of established trees on regeneration; (2) the mechanisms underlying these effects; and (3) to test the ‘stress gradient hypothesis’ in successional tropical dry forests. The effects of established trees on seed dispersal, seed survival, and seed germination—either through direct changes to moisture and temperature regimes or mediated by seed dispersers and predators—are mainly positive. The balance between positive and negative effects on seedling establishment is more complex and depends on the season and leaf phenology of both trees and seedlings. Seedling survival is generally enhanced by established trees mitigating dry conditions. Established trees have counteracting effects on water and light availability that influence seedling growth. The probability of a positive effect of established trees on seedling survival decreases with increased rainfall, which supports the stress gradient hypothesis. Priorities for future research are experiments to test for facilitation and competition and their underlying mechanisms, long‐term studies evaluating how these effects change with ontogeny, and studies focusing on the species‐specificity of interactions.

Does the stress‐gradient hypothesis hold water? Disentangling spatial and temporal variation in plant effects on soil moisture in dryland systems

Summary The nature of the relationship between water limitation and facilitation has been one of the most contentious debates surrounding the stress‐gradient hypothesis (SGH), which states that plant‐plant interactions shift from competition to facilitation with increasing environmental stress. We take a closer look at the potential role of soil moisture in mediating plant‐plant interaction outcomes by assessing effects of climate and soil texture on plant modulation of soil moisture. Using an empirically‐parameterized soil moisture model, we simulated soil moisture dynamics beneath shrubs and in un‐vegetated coarse and fine soils for 1000 sites in the Western United States with &lt;700 mm mean annual precipitation. This threshold reflects the transition from dryland (&lt;600 mm precipitation) to mesic ecosystems. Positive effects of shrubs on shallow soil moisture (i.e. the difference between shrub and interspace soil moisture) decreased along the aridity gradient when long‐term average conditions were considered, contrary to expectations based on the SGH. Negative effects of shrubs on deeper soil moisture also increased with aridity. However, when extreme years were considered, positive effects of shrub on soil moisture were greatest at intermediate points along the spatial aridity gradient, consistent with a hump‐backed model of plant‐plant interactions. When viewed through time within a site, shrub effects on shallow soil moisture were positively related to precipitation, with more complex relationships exhibited in deeper soils Taken together, the results of this simulation study suggest that plant effects on soil moisture are predictable based on relatively general relationships between precipitation inputs and differential evaporation and transpiration rates between plant and interspace microsites that are largely driven by temperature. In particular, this study highlights the importance of differentiating between temporal and spatial variation in weather and climate, respectively, in determining plant effects on available soil moisture. Rather than focusing on the somewhat coarse‐scale predictions of the SGH, it may be more beneficial to explicitly incorporate plant effects on soil moisture into predictive models of plant‐plant interaction outcomes in drylands.

Species identity and neighbor size surpass the impact of tree species diversity on productivity in experimental broad-leaved tree sapling assemblages under dry and moist conditions.

Species diversity may increase the productivity of tree communities through complementarity (CE) and/or selection effects (SE), but it is not well known how this relationship changes under water limitation. We tested the stress-gradient hypothesis, which predicts that resource use complementarity and facilitation are more important under water-limited conditions. We conducted a growth experiment with saplings of five temperate broad-leaved tree species that were grown in assemblages of variable diversity (1, 3, or 5 species) and species composition under ample and limited water supply to examine effects of species richness and species identity on stand- and tree-level productivity. Special attention was paid to effects of neighbor identity on the growth of target trees in mixture as compared to growth in monoculture. Stand productivity was strongly influenced by species identity while a net biodiversity effect (NE) was significant in the moist treatment (mostly assignable to CE) but of minor importance. The growth performance of some of the species in the mixtures was affected by tree neighborhood characteristics with neighbor size likely being more important than neighbor species identity. Diversity and neighbor identity effects visible in the moist treatment mostly disappeared in the dry treatment, disproving the stress-gradient hypothesis. The mixtures were similarly sensitive to drought-induced growth reduction as the monocultures, which may relate to the decreased CE on growth upon drought in the mixtures.

Positive interactions expand habitat use and the realized niches of sympatric species.

Niche theory, the oldest, most established community assembly model, predicts that in sympatry, the realized niche will contract due to negative interspecific interactions, but fails to recognize the effects of positive interactions on community assembly. The stress gradient hypothesis predicts that positive interactions expand realized niches in stressful habitats. We tested the predictions of the stress gradient hypothesis in a cobble beach model system across both physical and biological stress gradients. We transplanted seven common littoral species within, adjacent to, and below Spartina alterniflora cordgrass stands in control, cage control, predator exclusion cage, shade, and shaded predator exclusion cage treatments to test the hypothesis that cordgrass expands intertidal organism habitats. On cobble beaches, cordgrass ameliorates physical and predation stresses, expanding the distribution and realized niches of species to habitats in which they cannot live without facilitation, suggesting that niche theory and species distribution models should be amended to accommodate the role of positive interactions in community assembly.

Using positive interaction ecology to explain grassland degradation induced by overgrazing

Overgrazing of grassland ecosystems in the temperate steppe region of China has resulted in serious degradation and a distinct vegetation pattern. Grassland degradation is a major concern to ecologists and has been the focus of much recent research. A lot of work has been done on grassland degradation in grassland communities but few studies have explored the mechanisms of degradation from an ecological theory perspective. Positive interaction ecology theory has become one of the major themes in ecology, particularly the Stress Gradient Hypothesis. In the following study we attempt to explain the mechanisms of grassland degradation using positive interaction ecology and provide a new perspective for recognizing grassland degradation.

Competition with trees does not influence root characteristics of perennial grasses in semi-arid and arid savannas in South Africa

Savannas support mixed tree-grass communities and interactions between these are typically viewed as being competitive based on studies that focused on grass aboveground production. However, an important plant response to competition and resource limitation is an increase in root reserves. We investigated root characteristics of perennial grasses in the presence and absence of trees as a proxy of competition in South African savannas in three sites that differed in rainfall. We based our study on the hypothesis that competition from trees and water limitation will result in increased storage in roots of grasses under trees. Results indicate no significant effect of variation in rainfall of the different study locations on root characteristics of grasses. Furthermore, trees did not significantly influence most grass root characteristics that we measured. The only exception was nitrogen-content that showed an increase with rainfall and tree presence through potentially higher mineralization rates and nitrogen availability in the under-tree canopy environment. As the study sites are in the drier rainfall range in South Africa, it is likely that trees and grasses in these dry savannas may have a positive relationship conforming to the stress-gradient hypothesis. Alternatively, grasses and trees may be using complementary water and nutritional resources.

Climate modifies tree interactions in terms of basal area growth and mortality in monospecific and mixed Fagus sylvatica and Pinus sylvestris forests

Net interactions between trees vary depending on environmental conditions and other factors such as stand density, age, or between-species complementarity and/or facilitation. According to the stress gradient hypothesis, positive or facilitative interactions are more frequent in high-stress environments whereas negative or competitive interactions occur in benign environments, although recent studies highlight the influence of species composition, type of stress, ontogeny, etc. on the interaction–stress gradient pattern. The aim of this paper is to analyze whether site climatic variables are a key factor in tree interactions in mixed stands of beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.). To test how site climatic conditions modify the effect of inter-specific competition on tree basal area growth and tree mortality, growth and mortality models were fitted using monospecific and mixed sample plots located in matching site conditions selected from the Spanish National Forest Inventory in the Navarra Province. Tree competition status was broken down into four terms according to size-symmetry (size-symmetric vs. size-asymmetric) and species identity (intra-specific vs. inter-specific). The results showed that the size-symmetric inter-specific component was non-significant on beech basal area growth and had a negative effect on pine growth. The effect of size-asymmetric inter-specific competition was always non-significant, resulting in a higher basal area growth when the admixed species are of a larger size. The interaction between annual precipitation and this inter-specific competition effect was much more pronounced on beech than on pine. Inter-specific competition had a negative effect on pine growth under better climatic conditions and a positive effect at dryer sites, while beech always benefited from the presence of pine, although the benefit was greater where climatic site conditions were better. In tree mortality models, pine mortality increased with the proportion of beech, while beech mortality was lower as the proportion of pine increased. Precipitation modified the inter-specific competition effect on tree mortality although the site influence was less relevant than on tree growth. For pine mortality, the negative effect of beech admixture was stronger at lower mean annual precipitation, while in the case of beech the positive effect of pine increased at higher levels of precipitation. The influence of climate on the effect of competition, the variation in their strength depending on the mode of competition (size-symmetric or size-asymmetric), along with the inter-specific competition component, highlight the importance of considering the effect of site conditions on between-species interactions when modeling tree growth and mortality. The species-specific patterns of variation in tree interactions along climatic gradients and the differences in tree growth and mortality corroborate the need to consider the nature of stress-limiting factors and species composition and the importance of analyzing both dynamic processes simultaneously.

Lotic cyprinid communities can be structured as nest webs and predicted by the stress-gradient hypothesis.

Little is known about how positive biotic interactions structure animal communities. Nest association is a common reproductive facilitation in which associate species spawn in nests constructed by host species. Nest-associative behaviour is nearly obligate for some species, but facultative for others; this can complicate interaction network topology. Nest web diagrams can be used to depict interactions in nesting-structured communities and generate predictions about those interactions, but have thus far only been applied to cavity-nesting vertebrate communities. Likewise, the stress-gradient hypothesis (SGH) predicts that prevalent biotic interactions shift from competition to facilitation as abiotic and biotic stress increase; this model has been hardly applied to animal communities. Here, both of these models were applied to nest-associative fish communities and extended in novel ways to broaden their applicability. A nest web was constructed using spawning observations over 3 years in several streams in south-western Virginia, USA. Structural equation modelling (SEM) was then implemented through an information-theoretic framework to identify the most plausible nest web topology in stream fish communities at 45 sites in the New River basin of the central Appalachian Mountains, USA. To test the SGH, the per-nest reproductive success of 'strong' (nearly obligate) nest associates was used to represent interaction importance. Eigenvectors were extracted from a principal coordinate analysis (PCoA) of proportional species abundances to represent community structure. Both of these metrics were regressed on physical stress, a combination of catchment-scale agricultural land use and stream size (representing spatiotemporal habitat variability). Seventy-one per cent of SEM model evidence supported a parsimonious interaction topology in which strong associates rely on a single host (Nocomis), but not other species. PCoA identified a gradient of community structure dominated by Nocomis and associates, to communities dominated by other reproductive groups. Both metrics of interaction importance responded positively to physical stress. This study demonstrates that nest webs can be useful in a variety of systems and that SEM can be a quantitative extension of this framework. Likewise, the SGH can be used to understand positive interactions in animal communities and can be extended to predict proportional representation of facilitating and beneficiary species in communities.

Diversity differs among three variations of the stress gradients hypothesis in two representations of niche space

How does the stress-gradient hypothesis affect coexistence in relation to established theory? For two orthogonal stress gradients, a spatially explicit agent based simulation is used to project diversity for simple competitive and facilitative interactions and for three variations of the stress-gradient hypothesis: intraspecific and interspecific competitive and facilitative interactions are a function of the abiotic environment; interactions are relative to species-specific fitness along gradients; or interaction is fixed by species regardless of the abiotic environment. Simulations are run with two orthogonal environmental gradients for two representations of niche. Facilitation can increase diversity by maintaining larger source populations and thus higher establishment rates and sink populations. With species hierarchically related in niche space, the simulations show that positive interactions and changing interactions along a stress gradient maintain greater diversity through intraspecific competition that is effective where dominance would occur and through facilitation where stress is high. A changing environment that favors some species and harms others decreases diversity in the hierarchical cases, where poor competitors most likely subject to interspecific interaction respond most strongly. Diversity outcomes differ among the three stress gradient variations because the intensity of interactions differs across the environmental gradients, not because of change in the environment.