Oikos

1. Plant roots employ multiple strategies to acquire soil resources, including the outgrowth of root hairs and the association with arbuscular mycorrhizal fungi (AMF). These structures represent alternative pathways to access soil resources. Evidence is derived mostly from studies within single species or multi-species experiments under controlled conditions. It remains unclear whether this trade-off holds under natural conditions and whether it manifests consistently across different scales of biological organization. 2. We analysed 81 grassland species representing grasses, forbs and legumes, sampled under heterogeneous field conditions in three regions across Germany, allowing for analyses (i) across different species (interspecific), (ii) across communities (intercommunity), (iii) within plant species (intraspecific) and (iv) within root systems of individual plants (intra-individual). Fine root traits related to AMF colonization (including scoring of fungal hyphae and arbuscules) and root hair growth (including length and incidence) were quantified. 3. Comparing plant species, we observed a strong negative correlation between mycorrhizal colonization and root hair traits, confirming previous studies working under controlled conditions. The negative relationship between root hair length and mycorrhization was also evident when comparing mean trait values at the plant community scale. Among different individuals of the same species (i.e. intraspecific scale), there was no consistent pattern of trait-trait relationship. When comparing within roots of the same plant (i.e. intra-individual scale), co-occurrence of root hairs and mycorrhiza was less likely than what would be expected by chance, with forbs showing the most consistent negative pattern. 4. Our study demonstrates that the hypothesized trade-off between AMF and root hairs is a robust pattern under field conditions at the community and interspecific scale. At the intraspecific scale, no uniform trade-off pattern can be discerned, and within single root systems the negative trade-off pattern is widespread but weak. This indicates that ecological and evolutionary constraints shape trait-trait relationships at higher scale, whereas physiological limits do not restrict trait co-occurrence at lower scales.

In natural ecosystems, community dynamics are driven by diverse interactions operating through both ecological and reproductive processes. Although theory predicts that these processes jointly generate complex and contingent dynamics through the interplay of stabilizing (negative frequency dependence arising from niche differences) and destabilizing effects (positive frequency dependence caused by reproductive interference), empirical tests of this prediction remain scarce. Here, we combined experimental and theoretical approaches to investigate the dynamics of two congeneric annual plant species (Commelina) experiencing both ecological competition and reproductive interference. Our empirically parameterized model revealed a large niche difference and mutual reproductive interference. However, the reproductive interference was not strong enough to generate pronounced contingency of community dynamics on initial conditions. Instead, coexistence was the most likely outcome regardless of initial species composition. Parameter exploration indicated that the weak contingency arose because reproductive assurance strategies of Commelina species buffered the minority disadvantage caused by reproductive interference. These findings provide empirical evidence that reproductive interference does not necessarily destabilize coexistence and that it may be outweighed by ecological niche and fitness differences. Our study highlights the importance of integrating ecological and reproductive processes towards better prediction of community dynamics.

Environmental variability influences the evolution of phenotypic plasticity and the ecological interactions among species in communities, but it is unclear how these processes interact to affect ecological systems across scales. We investigate models describing how plasticity evolves in competitive communities exposed to seasonal variation in abiotic conditions. We conducted eco-evolutionary analyses to identify communities of coexisting species with evolutionarily stable plasticity traits. These analyses revealed that the presence of interspecific competitors constrained the evolution of plasticity in some species, which instead evolved to be specialists. Plasticity often impaired species coexistence by allowing individual species to control more niche space. Communities containing plastic species were usually less species-rich but more productive and less variable. Similar results were obtained when plasticity rate was evolutionarily labile, but plasticity rate altered the patterning of temporal niche partitioning. This work shows how plasticity and competition combine to shape ecological and trait responses to environmental variability.

Any mechanistic biodiversity-ecosystem functioning theory must explain how biodiversity in natural communities arises, which requires an evolutionary metacommunity perspective. Using a model of adaptive evolution of competitive metacommunities in landscapes with spatially heterogeneous supply of two resources, we studied the evolution of resource-uptake specialization as a function of resource type (substitutable/essential) and trade-off shape between resource-uptake affinities (generalist-/specialist-favouring). We identified "critical essentiality" as a key phenomenon separating parameter regions with fundamentally different biodiversity--ecosystem function relationships. Specifically, while greater spatial heterogeneity in resource supply increases biodiversity, this comes at the cost of reduced productivity when resources are essential. Moreover, when resources are essential, both generalist- and specialist-favoring trade-offs select for phenotypes that prevent efficient resource partitioning, leading to low resource use efficiency and consumer biomass - despite often high species richness. In contrast, when resources are substitutable, a specialist-favoring affinity trade-off selects for phenotypes that partition resources efficiently, promoting a metacommunity of only two species characterized by high functional diversity, resource use efficiency, and biomass. Overall, our analyses thus suggest that conditions promoting the evolution of diverse consumer communities often do not engender higher functional richness, consumer biomass, or resource use efficiency.

Understanding how species traits shape community properties is a central goal in ecology. But whether traits drive consistent patterns of dominance and assembly regardless of initial species abundances (evenness) remains unclear. We addressed this question using phytoplankton microcosm experiments to disentangle the effects of evenness and traits on community assembly and functioning, focusing on traits related to energy use (photosynthesis, size) alongside demographic parameters. Community assembly was highly deterministic, indicating that initial evenness had little influence on species dominance and final community structure. Instead, evenness drove community functioning. Demography was a stronger predictor of dominance than photosynthetic rates or individual size, suggesting that growth and competitive success depend on multiple metabolic processes that cannot be reduced to simpler proxies. These results show that community structure can emerge predictably across a range of initial conditions, and highlight the importance of demographic metrics over simpler metabolic descriptors for forecasting community outcomes.

Litter invertebrates mediate a substantial proportion of decomposition and nutrient cycling in terrestrial ecosystems, yet how moisture availability structures their communities remains poorly understood. We propose that moisture availability could shape litter faunal communities through two alternate pathways: (1) a productivity effect: greater productivity and detrital inputs in wetter forests should increase litter-faunal abundance, (2) a trait effect: phylogenetic differences in water balance should filter communities according to taxon-specific moisture tolerances. We tested these expectations across 100 forest sites spanning a moisture gradient in south-eastern Australia, from dry sclerophyll forest to cool temperate rainforest. Measuring 11 focal invertebrate taxa, total litter-faunal abundance increased threefold across the moisture gradient, from 400,000 individuals per hectare of dry forest to 1.2 million individuals per hectare of rainforest - consistent with greater resource availability in wetter and more productive habitats. However, this overall abundance trend masked pronounced variation in community composition across habitat types. While moisture-sensitive crustaceans and myriapods (particularly Isopoda and Chilopoda) were concentrated in rainforest litter, two groups of hexapods (Blattodea and Embioptera) were most abundant in dry forest litter and declined sharply toward rainforest habitat. These contrasting responses indicate that dry forests do not simply represent depauperate versions of wet-forest communities but harbour their own distinct and abundant litter invertebrate assemblages. Thus, moisture structures litter faunal communities via both proposed mechanisms: (1) a productivity effect and (2) a trait filtering effect. Given that moisture fundamentally structures litter faunal communities, shifts in forest moisture regimes under climate change may substantially alter detritivore communities, with consequences for decomposition processes and nutrient cycling globally.

Ecological stability and function emerge from the many and diverse types of interactions among species. Ecological networks represent the structure and strength of species interactions, but their study is often restricted to single interaction-types, likely due to an absence of models for plausibly (i.e., nonrandomly) structured 'multiplex networks' of multiple interaction types. Here, we present a general but mechanistic model that simulates multiplex networks using the different, but correlated, traits that determine different species interactions. We show that ecological stability varies systematically with these trait correlations by controlling the emergent complexity and positioning of strong interactions in the community. Our model provides a flexible but power tool to study multiplex networks under alternative trait-based hypotheses, offering new insights into the structure and stability of complex natural systems.

Temporal stability of plant populations is a key determinant of species coexistence and ecosystem functioning, yet its trait-based explanation has remained largely aboveground. Whether root functional traits contribute to long-term population stability remains poorly understood, despite roots mediating water and nutrient acquisition. We combined 17 years of vegetation monitoring from 150 temperate grassland plots across three regions in Germany with root functional trait data for 72 common grassland species. Temporal stability was computed as coefficient of variation (CV), with lower CV indicating higher stability. We tested whether root traits predict stability, and specifically which belowground dimensions from resource acquisition/conservation or soil exploration dominate, and whether differences in land use intensity modifies these relationships. Root traits were stronger and more consistent predictors of temporal stability than the classical aboveground traits specific leaf area (SLA) and leaf dry matter content (LDMC). Higher average root diameter (AD), root hair incidence (RHI), specific root length (SRL), and root tissue density (RTD) were generally associated with higher temporal stability, although the strength of these relationships varied among regions. Land use modified selected trait and stability relationships, with the strongest shifts occurring along the mowing gradient. Increasing mowing strengthened the associations of higher RHI and RTD with temporal stability, whereas the associations of AD and root N shifted towards lower stability. Our results identify belowground functional traits as an important missing component of trait-based temporal stability research and advance our understanding of the functional strategies underlying population temporal stability in managed semi-natural grasslands.

Global biodiversity recovery demands restoration at landscape scales, where ecological outcomes are shaped not only by habitat amount but also spatial arrangement. Alternative landscape restoration strategies can increase habitat amount, improve quality, and alter configuration in fundamentally different ways and may therefore vary in their efficiency. Here, we conceptualise five landscape restoration strategies and use a spatially explicit metapopulation framework to compare their ability to increase predicted species persistence across heterogeneous landscapes and trait space defined by dispersal ability and specialisation. We find that restoration outcomes are strongly trait-dependent, with restoration of existing degraded habitats yielding the highest benefits - particularly for specialists. Improvement of the landscape matrix to create low-quality habitat primarily benefits generalists, while strategies creating new habitat patches or enlarging patches provide more consistent but modest gains. These results reveal fundamental trade-offs among restoration strategies and highlight the utility of trait-explicit approaches to inform their implementation and evaluation.

Shifts in predation and human harvest often coincide with changes in life history trait distributions in prey populations. These changes may arise via evolution or plasticity, and extrinsic environmental effects may affect such traits. This complexity prompts questions about how trait distributions are shaped by multiple factors' interactions. We hypothesized that food availability and climate affect both the magnitude, and mechanism (evolution vs. plasticity), by which traits change in response to harvest's intensity and size selectivity. We tested these hypotheses using individual-based bioenergetic population models allowing both plasticity and evolution. We exposed genetically diverse fish populations to gradients of productivity and stochasticity in food availability, crossed with gradients of selective or nonselective fishing mortality. Food availability shaped traits both plastically and evolutionarily, and modulated how harvesting influenced those responses. Food effects can suppress harvest impacts altogether, or obscure evolution's effects on phenotypes. However, when climatic factors shape food-driven changes by affecting both fish and their prey, either synergistic or antagonistic interactions can emerge between food variation and harvest in determining phenotype distributions. Our research suggests that understanding consumer-resource dynamics, and climatic processes shaping them, is essential to interpreting how harvest can shape traits relevant to population viability and fisheries sustainability.

1. Grassland ecosystems are pivotal for global biodiversity, carbon (C) storage and food security. This biome, however, together with the ecosystem services it provides, is threatened by global change perturbations. These perturbations rarely occur in isolation and most often affect environments simultaneously, which is a pressing issue in ecology. Nevertheless, most of the research conducted so far has focussed on single simultaneous perturbations or pairs thereof. The few examples considering the effect of more than two simultaneous perturbations focussed only on plants or soil separately, which ignores the potential buffering or feedback effects on ecosystem functioning induced by plant-soils interactions. 2. To tackle this knowledge gap, we used an experimental setup involving mesocosms (soil and plants intact) collected in an extensively managed permanent grassland. Mesocosms were arranged in a greenhouse experiment where we applied individual and combined global change perturbations (up to ten in total) including application of fertilisers, pesticides and antibiotics, simulated trampling and grazing, heat stress and drought. Measured ecosystem functions and properties included a large set of biological, physical and chemical soil and plant associated variables. 3. Results showed that multiple perturbations in most cases had negative effects irrespective of the direction found for individual perturbations. However, in contrast to previous bare soil experiments, we only found scarce evidence for substantial synergistic effects of simultaneous perturbations in our experiment conducted with mesocosms that contained vegetation. This suggests a mitigating impact of plants on soil responses to global change. Moreover, we found a strong dominative effect of drought on ecosystem functioning, driving most of the effects observed. 4. Synthesis: Our results highlight the importance of evaluating global change impacts under more realistic settings, i.e., applying multiple perturbations simultaneously and including plant-soil interactions. Mitigation of global change in terrestrial ecosystems will only be possible once we holistically understand how ecosystems respond to altered conditions.

Environmental change alters species growth rates and, through species interactions, can produce complex effects on community functioning and composition. Because functional traits mediate both environmental responses and competition, rapid trait adaptation may modify these effects, but its consequences in multispecies communities remain unclear. We used a trait-based model of competitive communities to test how adaptation affects the mean and diversity of species growth responses to environmental change, and how this influences functional and compositional resistance (how unaffected the community is by the environmental change) and resilience (how well it recovers after it has acclimated to the altered environment). Adaptation altered response variance more strongly than the mean by changing the balance between environmental optimisation and competition-driven trait divergence. As a result, adaptation strongly increased compositional stability, but had less clear effects on abundance. Adaptation also generated a trade-off between resistance and resilience: faster adaptation improved resistance but reduced resilience as communities were displaced less by the environmental change and thus recovered less. Overall, adaptation can be stabilising, but higher adaptation rates do not necessarily produce greater stability across all stability components.

Beta diversity is a fundamental component of regional biodiversity that accumulates from compositional differences between local species assemblages. Although beta diversity is increasingly applied across spatial scales, its scale dependence remains incompletely synthesized, particularly concerning variation in spatial scale used to define alpha diversity (grain size). Using tree community data from Barro Colorado Island, we find that both additive and multiplicative beta diversity are consistently higher for unequal than equal grain divisions across different spatial extents and numbers of grains. These results can be explained by Jensen's inequality based on the concave-down relationship of alpha diversity with area that in turn increases beta diversity through the scaling relationship between alpha and gamma diversity. Combined with our additional analysis showing that this prediction is robust to more general models of species richness, including species--area and species--sampling-effort curves, these results reveal a previously unrecognized, predictable range of beta diversity variation.

Ecological coarse-graining of individuals or species into aggregate variables, such as total biomass or regional abundance, is common practice, yet differences among species can still influence aggregate dynamics. We propose a metric for quantifying this irreducible dynamical heterogeneity. For small fluctuations near a stable equilibrium, we compare the aggregate fluctuation spectrum of the full dynamical system with that of the best Ornstein-Uhlenbeck model having the same dimension as the aggregate. The remaining spectral mismatch quantifies the dynamical effects of hidden differences that the fitted aggregate model cannot capture. For weakly interacting random community models at fixed collective interaction strength, we show that increasing species richness can either reduce or increase this heterogeneity, depending on how species interact. Our framework helps understand when ecological and evolutionary differences average out versus when their dynamical effects persist.

Understanding spatial and temporal variation in community composition is central to protecting and monitoring biodiversity. Similarity in the composition of two communities decreases along spatial and temporal gradients, a relationship known as distance decay. Comparing distance-decay relationships across taxonomic, functional, and bioclimatic facets of diversity provides a means of interrogating niche-based versus neutral influences on community assembly. Furthermore, the context in which these facets yield overlapping or complementary information about global change impacts remains poorly understood. In this study, we examined spatiotemporal distance-decay relationships for taxonomic, functional, and bioclimatic facets of alpine plant diversity leveraging elevation gradients on five peaks in the White Mountains, California, USA, sampled at 3-4 time points between 2007 and 2024. Using multivariate Bayesian beta regression models, we estimated the slopes of elevation distance decay over multiple surveys and across taxonomic, functional and bioclimatic diversity to understand how spatial distance-decay relationships are changing through time and across different facets of diversity. We found strong spatial distance decay of taxonomic similarity across all peaks, with the highest elevation peak showing the strongest decay. Functional and bioclimatic similarities decayed more weakly than taxonomic similarity, suggesting environmental filtering shapes trait and niche distributions. Critically, spatial decay patterns showed no temporal signal across facets of diversity, indicating community stasis rather than biotic homogenization or differentiation. This study adds further evidence that xeric alpine plant communities are resisting climate change impacts through time.

Understanding how species interactions, ecological drift, and immigration jointly shape community composition is a central goal of community ecology. These processes determine not only average community composition, but also how often different community states occur. Here, we ask how competition reshapes the neutral distribution of community composition generated by drift and immigration. We analyze a stochastic two-species competition model with immigration and derive the stationary distribution of relative species composition near neutrality. The stationary distribution shows when communities are usually balanced and when they are instead dominated by one species. Species dominance can become common even when the corresponding deterministic model predicts stable coexistence, particularly in smaller communities. Balanced and dominance states can also occur together, producing a trimodal distribution. Competitive asymmetry further biases the distribution toward the competitively favored species. Thus, the interplay between competition, ecological drift, and immigration determines not only how much community composition varies, but also which community states are most likely to be observed. Our results provide an analytical bridge between neutral and competition theory and a framework for interpreting variation in community composition.

As ecosystems face rapid environmental change, understanding the interplay between landscape structure, genetic diversity, and adaptive potential is critical for predicting resilience. We investigated how biome-specific connectivity and environmental heterogeneity mediate adaptive strategies and resilience in the tropical rainbowfish (Melanotaenia splendida splendida) across contrasting rainforest and savannah ecosystems in tropical Australia. Using an integrative framework combining genomic, morphological, and environmental analyses, we inferred that hydroclimatic variation has been a key driver of genetic and phenotypic divergence. Rainforest populations exhibited strong local adaptation to fragmented, stable environments, while savannah populations showed less genomic differentiation and morphological divergence, patterns consistent with weaker local adaptation. We hypothesise that greater reliance on phenotypic plasticity may facilitate adaptive responses in savannah populations, reflecting their exposure to more connected and variable environments. Variance partitioning highlighted the contrasting roles of neutral and adaptive processes across biomes, with implications for managing resilience in freshwater systems under climate change. Our findings underscore the trade-offs between connectivity, local adaptation, and plasticity, providing a model for understanding adaptation and resilience across diverse ecosystems.

Streambank erosion degrades both terrestrial and aquatic ecosystems, and riparian buffers are among the most widely used tools to control it. Yet most of what is known about how riparian vegetation controls erosion comes from one or two-species systems, which overlook the diversity of riparian communities and species interactions (competition and facilitation) that may shape the root systems that bind soil. We used a streambank mesocosm experiment to test whether plant species richness alters root traits, and whether root traits and species interactions reduce erosion. We established 85 mesocosms ranging from unvegetated controls to six-species polycultures, measured root biomass, length, volume, and diameter, quantified species interactions with relative yield totals (RYT), and measured erosion rate with a jet erosion test. Vegetated mesocosms eroded significantly less than unvegetated controls (t = 4.05, p = 0.023). Richness did not affect most root traits, although mean root diameter decreased as richness increased ({beta} = -0.0225, p = 0.047). RYT ranged from competition (RYT < 1) to facilitation (RYT >1), yet neither interactions nor any root trait was directly associated with erosion rate. However, RYT was negatively associated with soil moisture ({beta} = -0.196, p = 0.010). Piecewise structural equation models showed that erosion was best explained by a pathway linking species presence, root length, and soil moisture (AIC = 39.27; Fisher's C = 11.27, df = 12, p = 0.506). Panicum virgatum increased root length, whereas Lolium multiflorum and Echinochloa crus-galli reduced it; longer roots were associated with drier soil (standardized {beta} = -0.31, p = 0.005), and wetter soils eroded more slowly (standardized {beta} = -0.39, p < 0.001). Which species were present mattered more than how many, and plants were linked to erosion through soil moisture rather than root traits alone. Species selection for riparian buffers may need to consider effects on soil moisture as well as rooting extent, particularly as droughts followed by intense storms become more common under a changing climate.

Species at the highest trophic levels are generally the least resilient to perturbations, yet the factors that determine their persistence remain poorly understood. We ask how the intrinsic self-regulation of a basal host governs the persistence of the trophic levels above it. To address this question, we use a minimal discrete-time model of a host-parasitoid-hyperparasitoid system motivated by the biology of trigonalid wasps, obligate hyperparasitoids that complete development only when their caterpillar host is also attacked by a primary parasitoid. We compare two forms of host self-regulation in the absence of natural enemies: scramble (Ricker) fecundity, appropriate when the parasitoid is the host's principal enemy and which recovers the classical model of Beddington and Hammond as a special case; and contest (Beverton-Holt) fecundity, appropriate when the host is held in check by many additional enemies. Using numerical continuation, Lyapunov exponents, and direct computation of the model's trajectories, we find that the form of host regulation is decisive for persistence at the top trophic level. Under scramble host dynamics, coexistence is extremely fragile with respect to both parameter values and initial conditions. Under contest host dynamics, coexistence is robust across broad parameter ranges and loses stability only gradually, through a supercritical Neimark--Sacker bifurcation and a subsequent torus-doubling route to chaos. Therefore, in at least some trophic networks, the resilience of the highest levels can be determined at the base of the network.

1. Group foraging reduces the risks associated with foraging: larger groups dilute individual predation risk, allow group members to share the costs of vigilance, and provide a social structure for exchanging information about predators. The benefits of shared social information over simple safety in numbers should depend on the relationships among group members. Consequently, group composition should affect the selection of risky and safe habitats in social species. 2. We asked whether relationships among elk (Cervus canadensis) group members moving together influence their selection of risky habitats more than grouping alone. We tested three hypotheses. The familiarity hypothesis predicts that elk with close social ties are most likely to select risky habitats together. The inclusive fitness hypothesis predicts that the closest genetic relatives are most likely to select risky habitats together. The dilution hypothesis predicts that familiarity and genetic relationships do not influence habitat selection, with all group foraging benefits derived from proximity to others. 3. We used integrated Step Selection Analysis to test whether the identity of the closest group members influenced habitat selection. We tested whether the relationship of the nearest elk--either genetic or social--incentivized selection of risky habitat more than proximity to the nearest elk. 4. Group foraging clearly reduced risk, with elk more likely to select risky habitat when near others. Elk were also more likely to select risky habitats when near closer relatives, albeit with considerable variation among individuals. However, familiarity had a large and consistent effect: elk were far more likely to select risky habitats when near socially familiar group members. 5. Our results suggest that information shared within foraging groups, especially between socially familiar individuals, may lower the risks associated with habitat selection. Similarly, the benefits of reliable social information for reducing risk likely incentivizes the formation of stronger social ties within foraging groups. This positive feedback between social familiarity and habitat selection could thereby increase access to profitable habitats that are otherwise too risky, cementing what we term the social information landscape as a key driver of movement and habitat selection.