Ecology

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.

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.

Defended prey are often thought to rely on conspicuous warning signals to distinguish themselves from undefended prey, facilitating predator learning and avoidance. Yet many chemically defended organisms are inconspicuous and difficult to distinguish from palatable prey, raising a fundamental question: how do predators avoid defended prey when prey themselves provide little or no reliable signals of defense? We tested whether host plants can function as indirect indicators of prey defense using Carolina wrens (Thryothorus ludovicianus), chemically defended Eumaeus atala larvae, undefended Galleria mellonella larvae, and two distinct plant species. Defended and undefended prey were experimentally made to appear visually identical, forcing predators to rely on host-plant associations as the only available source of information. Wrens rapidly altered both search behavior and attack decisions according to plant identity. Mechanistic attack models incorporating predator discrimination between defended and undefended prey were strongly favored when plants were present, with defended prey experiencing substantially lower attack risk than undefended prey. When plants were removed, support for discrimination weakened and attacks on both prey types increased. Spatial autocorrelation of host plants further shaped predator behavior, with strongly clustered plants hosting defended prey receiving disproportionately fewer visits. Together, these results suggest host plants can function as ecologically meaningful cues of prey defense, influencing both predator search patterns and attack decisions. More broadly, our findings suggest warning systems may extend beyond prey phenotype alone, and that ecological context (including host-plant identity and spatial structure) may contribute to the evolution and maintenance of aposematism.

Diverse ecological communities can reduce pathogen transmission; a phenomenon known as the dilution effect of biodiversity. This effect, in turn, depends on variation in host species' ability to spread disease (i.e., host competence) and which host species persist under biodiversity loss. Mutualistic symbionts can influence host competence and are sensitive to variation in host biodiversity; yet whether mutualists can modify dilution effects is unknown, partially because quantifying host competence during epidemics remains challenging. We experimentally tested whether mutualists could modify dilution effects by establishing gradients of host species richness, community competence, and access to the below-ground arbuscular mycorrhizal fungus (AMF), Rhizoglomus irregulare. Plant hosts were then exposed to the fungal root pathogen, Rhizoctonia solani, and disease spread was monitored over the course of an epidemic. Overall, host richness did not affect epidemic outcomes. Instead, AMF inoculation increased disease risk via species-specific increases in host competence. Moreover, communities containing highly competent host species experienced more disease, but only in the presence of mutualistic AMF. These results underscore the importance of plant community composition in mycorrhiza-mediated disease dynamics and suggest that predicting disease in our changing world might require a deeper understanding of host-symbiont and pathogen-symbiont interactions under global change.

1. Detecting causal relationships from ecological time series remains challenging because community dynamics are often nonlinear, high-dimensional, noisy, and strongly synchronized. These characteristics can reduce the reliability of existing causal inference methods and complicate the identification of ecological interactions in complex natural systems. 2. I introduce Forest-weighted Causal Correlation (FORCC), a machine-learning-based framework that infers directional causality from adaptive neighborhood structures learned by random forests. Rather than evaluating causal direction through cross-map reconstruction, FORCC quantifies directional dependence from the preservation of neighborhood structure using rank-based associations among forest-weight matrices. 3. Simulations of coupled nonlinear systems showed that FORCC accurately recovered causal direction while substantially reducing synchronization-induced false-positive detections relative to conventional convergent cross mapping. Chatterjee's {xi} exhibited the strongest resistance to synchronization-induced errors among the evaluated rank-based statistics. 4. Application to a 12-year marine community time series from Maizuru Bay, Japan, identified asymmetric relationships between individual taxa and community dynamics, revealing candidate driver-like and passenger-like roles that were not readily explained by seasonal synchrony alone. 5. By combining adaptive neighborhood learning with rank-based causal inference, FORCC provides a scalable framework for causal discovery in ecological time series. The method offers a practical tool for screening causal relationships, generating ecological hypotheses, and investigating complex interactions in multivariate natural systems.

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.

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.

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.

1. Temperature shapes life histories through its effects on energy allocation among growth, reproduction and maintenance. The resulting temperature dependence of performance is commonly characterised using thermal performance curves (TPCs), which are often treated as static descriptions that apply uniformly throughout life. Yet the response to thermal variability may change across ontogeny. 2. We recorded growth, reproduction and survival throughout the lifetimes of individual Daphnia magna under four thermal regimes: Constant Low (15C), Constant Rearing (20C), Constant High (25C) and Variable (15C to 25C). The mean temperature under Variable closely matched that under Constant Rearing. Using an allometric energy-allocation model, we derived age specific and lifetime performance measures and quantified their uncertainty with a stratified individual-level bootstrap. Our principal quantity was the Jensen contrast, defined as performance under Variable minus that under Constant Rearing. 3. Age specific point estimates of the Jensen contrast changed sign across ontogeny. A positive aggregate contrast in full trajectory growth concealed a negative mid-life contrast; reproductive point estimates were positive during early and mid life but negative later. Bootstrap support was strong for the positive early life and negative mid-life growth contrasts, and for the positive early life reproductive contrast. Under a second-order approximation, this pattern of estimated signs is consistent with ontogenetic changes in effective thermal curvature. 4. The positive early life reproductive response did not carry through to the lifetime outcome: expected lifetime reproduction was lower under Variable than under Constant Rearing, with strong bootstrap support for the negative lifetime contrast. Because lifetime reproduction is survival weighted, lower survival under Variable reduced later reproductive contributions; across the estimated trajectories, the accumulated negative late life contribution out-weighed the earlier positive contributions. 5. These results show why forecasts based on mean temperature, a single life stage or an aggregate TPC may misrepresent lifetime performance under variable thermal regimes. Accounting for both ontogeny and temperature variability may therefore improve forecasts of life-history performance in increasingly variable thermal environments.

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.

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.

Exotic species can reshape native communities, in the extreme, to local extirpation. Before that, natives may adjust where and when they use the landscape, in ways that allow coexistence and may be the earliest sign of trouble. Distinguishing coexistence built on such adjustment from one built on pre-existing niche differences requires comparing a native where the invader is absent with the same native where it is present, across sites at different invasion stages. Using eight years (2018-2026) of camera-trap data from 103 stations across Uruguay (154,590 camera-nights), we asked how the invasive axis deer (Axis axis) affects the space and time use of two native cervids: the forest-dwelling brown brocket (Subulo gouazoubira) and the grassland-restricted pampas deer (Ozotoceros bezoarticus). Activity-overlap, directional-avoidance, hierarchical activity, and two-species occupancy analyses showed sharply divergent dyads. A. axis and S. gouazoubira shared forest yet segregated in space and along the understory stratum the brocket requires and the invader avoids, and shifted their diel activity by context; understory cover fell with A. axis abundance, consistent with habitat modification. For A. axis and O. bezoarticus, segregation was intrinsic, opposite diel niches and habitats, with no support for a detectable shift in pampas deer behavior. The only behavioral signal was asymmetric: the invader, rather than the native, appeared to adjust, though few shared sites warrant caution. The same invader thus drives different mechanisms, but coexistence is provisional: A. axis is still expanding, far from equilibrium, so these shifts read best as early warning signals.

Heavy snowpack can be a severe constraint for alpine herbivores, yet adult survival in long-lived temperate ungulates is typically canalized against environmental variation. Using a 45-year individual-based dataset on bighorn sheep (Ovis canadensis) and historical snow reconstructions at Ram Mountain, Canada, we evaluated the direct and indirect effects of snow conditions on age- and sex-specific survival. For both sexes, survival of lambs and senescent individuals decreased with increasing average snow depth, whereas non-senescent adults (aged [&ge;] 1 year and < 8 years) showed high survival under mean conditions. However, this resilience broke down once specific thresholds of snow depth were crossed, with sharp survival declines beyond 90 cm of snow depth for females and 100 cm for males. Lamb survival, already reduced by increasing average snow depth, declined further once snow depth exceeded 90 cm. Crucially, the annual probability of snow depth exceeding 90 cm increased markedly over the study period, along with a density-independent temporal decline in survival across all demographic groups. Lamb survival declined by 0.37, and adult female survival, the most critical vital rate for population growth, dropped by 0.13 over the study period. Our findings challenge the established paradigm of demographic buffering in prime-aged adults, demonstrating that their resilience to environmental variability is fundamentally non-linear and breaks down once a changing climate regime breaches adult physiological boundaries, severely compromising population viability.

Forecasting infectious disease dynamics is important for ecological management and emerging disease preparedness, yet many systems lack the long-term datasets required to develop reliable predictions. Spatial replication may provide an alternative by allowing information to be shared among populations, but the extent to which this improves forecasting remains unclear. Here, I used epidemic and temperature data from 20 replicated semi-natural Daphnia-parasite pond populations monitored across four seasons (80 epidemics) to test whether information from multiple populations improves forecasts and whether environmental similarity identifies informative populations. I forecasted disease prevalence, infected host density, and healthy host density using a benchmark model, autoregressive integrated moving average (ARIMA), and time-series regression models. Models were trained using focal-population data alone, mean dynamics across other populations, or temperature-weighted mean dynamics from environmentally similar populations. Epidemics showed strong seasonal structure, with prevalence and infected host density generally peaking during the warmest period of the season. However, no forecasting approach consistently outperformed others across all variables. Temperature improved forecasts of prevalence and infected host density but not healthy host density. Multi-population training improved forecasts in some cases, particularly for ARIMA models, whereas temperature-weighted averaging provided no additional benefit over simple averaging. Regression models consistently produced the most accurate forecasts of healthy host density. These results show that borrowing information among populations can improve forecasts under specific conditions but is not universally beneficial, highlighting the importance of matching forecasting approaches and data sources to the ecological processes being predicted.

The density-yield relationship of wild plant populations, which in agricultural systems is typically consistent with the constant final yield hypothesis, has remained largely unexplored in a restoration context despite its potential importance. In degraded California shrublands, non-native annual grasses are associated with reduced native shrub recruitment. By studying their density-yield relationship, we can potentially optimize current restoration practices by identifying the level of grass control needed to reduce the competitive pressure they create for woody plants. This in turn could enhance woody plant establishment. Here, we conducted field experiments at a degraded chaparral site in Santa Barbara County, California. We manipulated the density of Bromus diandrus, a dominant invasive annual grass, through 0-100% removal treatments to test its density-yield relationship. Simultaneously, we outplanted Ceanothus megacarpus seedlings as phytometers while also monitoring soil moisture and light availability throughout the growing season. Our results demonstrate that the Bromus diandrus density-yield relationship follows the constant final yield hypothesis, with strong intraspecific competition observed from moderate to high grass density contributing to a constant population biomass and a density dependence allometric exponent comparable to previous agricultural studies. A reduction in grass biomass (yield) was observed at 75% grass removal, yet an increase in resource availability (i.e., light and soil moisture) was only observed at 100% grass removal, as indicated by both direct measurements and enhanced phytometer performance. Applied to restoration in California Chaparral and other Mediterranean ecosystems, where the reduction of "weed" biomass is a common management goal, our findings suggest full removal of non-native annual grasses is likely needed to aid in the restoration of native woody seedlings.

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.

Ecological uniqueness, commonly quantified by local contribution to beta diversity (LCBD), is important for understanding and conserving biodiversity. Its widely documented negative relationship with species richness has led to the view that species-poor communities may be disproportionately important for regional diversity. Here, we derive neutral expectations for richness-uniqueness relationships and show that negative slopes can arise from neutral assembly rather than necessarily reflecting an ecological signal of uniqueness. Neutral simulations validate the analytical expectations across abundance- and incidence-based LCBD formulations. Applying the framework to the Barro Colorado Island tropical forest and North American Breeding Bird Survey, we show that accounting for neutral sampling expectations substantially changes the ranking and spatial distribution of ecologically unique sites. By separating neutral sampling expectations from excess uniqueness, our framework provides a principled basis for interpreting ecological uniqueness and informing biodiversity conservation.

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.

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.

Ecological responses to environmental stressors are increasingly shaped by interactions among anthropogenic and climatic drivers, yet the influence of prior exposure history on subsequent stress responses remains poorly integrated into multiple stressor ecology. Using the blue mussel (Mytilus edulis) as a model system, we experimentally tested whether prior exposure to environmental nanoplastics modifies responses to subsequent stressors, including copper contamination, nutrient enrichment, and salinity fluctuations, individually and in combination. Nanoplastic pre-exposure reduced survival across subsequent treatments and altered stressor interaction outcomes at the community level, shifting responses toward synergistic effects that amplified metabolic costs. These results demonstrate that exposure history can modify both the magnitude and direction of multi-stressor effects, with consequences extending beyond individual performance to ecosystem functioning. By incorporating temporal exposure dynamics into multiple stressor research, our study highlights how ecological responses to global change depend not only on the identity and intensity of stressors, but also on prior environmental conditions shaping subsequent organismal and ecosystem health status and responses.