Ecology

Predation risk varies through space and time due to changing refuge quality, predator communities, and prey traits. Despite this, ecological research is often focused on measuring average predation risk at the community level. While this can give important information about overall trophic transfer and ecological efficiency, it ignores differences in predation risk among prey species within a community, which may be important determinants of species coexistence and local diversity. We used crustaceans associated with temperate seagrass in Northern California to explore the relationship between seasonal variation in among-species and community-level predation risk for a community of morphologically distinct prey. We measured predation risk of the four most abundant and widespread prey species at six field sites every two to six weeks for one year. At the community level, sites differed significantly in their annual variation in predation risk, and these differences were correlated with the amount of variation in the among-species predation risk. When there was more within-year variation in predation risk among the four prey species, predation risk at the community level was more stable across the year. On the other hand, when each prey species in the community had similar levels of predation risk throughout the year, predation as a community-level process was much more seasonal and variable. Variation in predation risk also changed across a gradient of seagrass cover, a proxy for refuge quality. Sites with greater seagrass cover had less annual variation in community-level predation risk and more variation in predation risk among the four species at any given time point. In contrast, at sites with less eelgrass, all species were consumed at the same rate throughout the year, suggesting previously demonstrated differences in antipredator strategies among species are less relevant in the absence of habitat-forming species. We suggest that larger species-specific differences in predation risk throughout a year result in a more stable level of predation risk for the whole community, and that this may be driven by increased refuge provided by seagrass habitat mediating different prey species relative levels of susceptibility to predation.

Intraspecific behavioral variation in prey could alter predator-prey interactions, yet its effects on temporal dynamics and food-web persistence remain underexplored. Pea aphids (Acyrthosiphon pisum) exhibit dropping behavior in response to predators like the seven-spot ladybird (Coccinella septempunctata). This response could be an effective anti-predator defense but could be costly in terms of energy expenditure and time not available for feeding. To investigate the impact of behavioral variation on food-chain persistence and dynamics, we used a tri-trophic experimental system with Vicia faba (plant), pea aphids (prey), and seven-spot ladybirds (predator), implementing three aphid behavioral treatments: droppers, non-droppers, and a mix of both droppers and non-droppers. To minimize genetic differences, we used clonal aphid populations across all treatments. We then tracked predator-prey population dynamics and species persistence over 25 days. Our results showed that aphid dropping behavior reduced food-chain persistence, with extinction risk significantly higher in dropper treatments than in the mix or non-dropper treatments. Ladybirds persisted across treatments, although they showed a steeper decline in abundance in the dropper treatment. In the mixed behavioral treatment, they had an intermediate persistence, suggesting a buffering effect of behavioral variation. Trophic food-chain state transitions also differed by treatment, with tri-trophic states most stable in the non-dropper, and least frequent in the dropper treatment. Furthermore, our results showed a trend of dropper treatments becoming more stable and robust towards the end of the experiment. These results demonstrate that prey behavior influences the persistence and dynamics of food-chains, with important implications for behavior-driven community dynamics.

Time-series data for ecological communities are increasingly available from long-term studies designed to track species responses to environmental change. However, classical multivariate methods for analyzing community composition have limited applicability for time series, as they do not account for temporal autocorrelation in community-member abundances. Furthermore, traditional approaches often obscure the connections between responses at the community level and those for individual taxa, limiting their capacity to infer mechanisms of community change. We show how linear mixed models that account for group-specific temporal autocorrelation and observation error can be used to infer both taxon- and community-level responses to environmental predictors from replicated time-series data. Variation in taxon-specific responses to predictors is modeled using random effects, which can be used to characterize variation in community composition. Moreover, the degree of autocorrelation is estimated separately for each taxon, since this is likely to vary due to differences in their underlying population dynamics. We illustrate the utility of the approach by analyzing the response of a predatory arthropod community to spatiotemporal variation in allochthonous resources in a subarctic landscape. Our results show how mixed models with temporal autocorrelation provide a unified approach to characterizing taxon- and community-level responses to environmental variation through time.

O_LIThe enemy release hypothesis is a popular hypothesis to explain the success of invasive plants. Enemy release studies typically focus on single species or types of communities, feature indirect experimental manipulations that apply pesticides to whole communities not individual species, and only examine responses of established plants or plant populations, limiting their generality. Using a novel species-specific approach, we examine whether enemy release can enhance seedling survival and recruitment of 16 grassland species by experimentally linking enemy release with enhanced plant performance. C_LIO_LIWe planted seedlings of 16 native grassland species from two functional groups (C4 grasses and non-legume forbs) into two grassland sites (early and mid succession). We hand-painted 1,548 individual seedlings with pesticides (insecticide and fungicide) over the course of a growing season to enforce aboveground species-specific enemy release, and tested whether it enhanced survival relative to untreated controls. Using native species enabled us to directly test effects of enemy release, while avoiding confounding factors like unknown invasion histories. Of the 16 native study species, 13 are naturalised/invasive outside of their native ranges. C_LIO_LIRelease from insects increased seedling survival by 80% on average, with no additional benefit of release from fungal pathogens. This effect was consistent across functional groups and community successional stages, and was strongest in resource-acquisitive species. The size of species performance benefits from enemy release were positively correlated with the number of regions globally where each species has been introduced and naturalised. C_LIO_LISynthesis. Previous studies of enemy release have centred on adults and findings have varied among species. We found a positive effect of release from insect herbivores early in colonisation - a trend that held across functional groups and types of resident community. We posit that the consistent vulnerability of seedlings vis-a-vis later life stages leads to this more ubiquitous benefit of enemy release. Enemy release may therefore aid initial recruitment of most, if not all, plants during the invasion process, even if enemies rapidly accumulate. The positive correlations between the benefits of enemy release for seedlings, species life history strategies and global naturalisation patterns provide compelling hypotheses for future research. C_LI

There is considerable interest in understanding patterns of {beta}-diversity that measure the amount of change in species composition through space or time. Most hypotheses for {beta}-diversity evoke nonrandom processes that generate spatial and temporal within species aggregation; however, {beta}-diversity can also be driven by random sampling processes. Here, we describe a framework based on rarefaction curves that quantifies the non-random contribution of species compositional differences across samples to {beta}-diversity. We isolate the effect of within-species spatial or temporal aggregation on beta-diversity using a coverage standardized metric of {beta}-diversity ({beta}C). We demonstrate the utility of our framework using simulations and an empirical case study examining variation in avian species composition through space and time in engineered versus natural riparian areas. The primary strengths of our approach are that it provides an intuitive visual null model for expected patterns of biodiversity under random sampling that allows integrating analyses across -, {gamma}-, and {beta}-scales. Importantly, the method can accommodate comparisons between communities with different species pool sizes, and can be used to examine species turnover both within and between meta-communities. Open Research statement: all code and data used in this manuscript are available at the following link: https://github.com/MoBiodiv/beta_concept

O_LIPlant community succession is structured by priority effects, plant consumer pressure, and soil resource supply. Importantly, these drivers may interact, their effects may vary temporally, and they may influence different facets of plant community diversity by promoting different plant tradeoff strategies.\nC_LIO_LIIn an herbaceous successional system, we manipulated priority effects by altering initial plant richness, consumer pressure via pesticide spraying, and soil resource supply via fertilization. We examined how these processes jointly influenced succession, including taxonomic diversity and functional traits, over four years.\nC_LIO_LIDiversity decreased in different years in response to more diverse priority effects, lower consumer pressure, and increased soil resource supply. Functionally, higher soil resource supply increased community height, SLA, and seed mass; higher consumer pressure decreased intraspecific community height, and increased interspecific SLA; priority effects led to decreased seed mass only when plots were unplanted.\nC_LIO_LIOur results suggest species resource strategies underlie plant diversity responses. Resource addition promoted resource-acquisitive species, consumer pressure disadvantaged resource-conservative species, and diversity of priority effects altered subsequent community composition through persistence of early residents, not via traits. We show that community responses to drivers of succession depend on underlying trait tradeoffs of resident species, and these tradeoffs influence community diversity across succession.\nC_LI

Determining the drivers of ecological stability amid accelerating global environmental change is a critical goal of contemporary ecology. Various candidate drivers have been suggested, with recent attention turning to response diversity--the variation among organism-environment responses. However, despite conceptual interest in response diversity as a driver of stability, there remain few field tests of this relationship. Using multi-species competitive communities of floating aquatic macrophytes as an experimental model for measuring temporal stability and response diversity to nutrient loading, we show that response diversity does not promote temporal stability of total macrophyte cover, but that communities with an uneven distribution of species responses were more resistant to an exogenous shock. To quantify macrophyte composition and growth dynamics from photographic time series of our experimental communities, we developed an open-source, scalable, machine learning workflow (LeafMosaic) capable of classifying four species from noisy field data including variable lighting, resolution, and plant morphology. We measured response diversity as the balance of positive and negative biomass growth responses to dissolved nitrate concentration, weighted by species relative contributions to biomass, and tested its effect on temporal stability and resistance to an unexpected pulse disturbance (a large typhoon that disrupted our outdoor mesocosms). Response imbalance predicted typhoon resistance, but species asynchrony and mean population stability best predicted community stability, with no direct or indirect effect of species responses. Overall, our results provide new experimental evidence for how the structure of species responses promotes stability, and we aim our LeafMosaic workflow to empower future field experiments using floating macrophytes to study response diversity and ecological stability.

O_LIBiodiversity loss can alter interactions not only through changes in tree species richness, but also through the loss of particular functional strategies from ecological communities. Working in a subtropical forest diversity experiment we asked whether tree species richness effects on arthropod herbivory and leaf pathogen infestation depend on community functional diversity, and whether trait dissimilarity-based, non-random species loss alters these relationships compared to random loss. To address this, we combined already established planted scenarios with newly constructed extinction pathways. C_LIO_LIWe tested the responses of herbivory and leaf pathogen infestation (i) to tree species richness, functional diversity (Raos Q), community structure and resource strategies (i.e. ever-greenness) and community-weighted trait means as well as predation, and (ii) trait dissimilarity-based extinction pathway analyses that contrasted directed loss of functionally similar versus functionally distinct tree species. C_LIO_LIHerbivory increased with tree species richness and this increase was significantly stronger in communities with higher tree functional diversity. Under directed species loss scenarios, herbivory differed most strongly from random-loss expectations when similar tree species were lost first. By contrast, losing functionally distinct species first produced richness effects that were much closer to the random-loss scenarios. Trait-based species loss will therefore modify trophic interactions more strongly than random loss. For pathogen infestation tree richness effects depended on evergreenness and among planted extinction scenarios (three-way interaction), with only minor deviations of trait-based extinction path-ways from random-loss expectations. Pathogen infestation also tended to increase with community-weighted mean leaf nitrogen. Predation showed no clear relationship with tree species richness or functional diversity but was positively associated with herbivory. The strength of this association differed among extinction scenarios, providing no evidence for consistent top-down regulation. C_LI SynthesisThe ecological consequences of biodiversity loss for leaf damage depend on which functional strategies are lost, not only on how many tree species remain. By integrating ob-served tree diversity gradients with trait-based extinction pathways, this study shows that functional diversity and host redundancy help explain why herbivores and pathogens are shaped by the same changes in tree diversity through different functional constraints and im-prove predictions of interaction strength under non-random species loss.

Host and parasite richness are generally positively correlated, but the stability of this relationship during community assembly remains untested. The composition of host communities can alter parasite transmission, and the relationship between host and parasite richness is sensitive to parasite transmission. Thus, changes in composition during host community assembly could strengthen or weaken the relationship between host and parasite richness. Host community assembly, in turn, can be driven by many processes, including resource enrichment. To test the hypothesis that host community assembly can alter the relationship between host and parasite richness, we experimentally crossed host diversity and resource supply to hosts, then allowed communities to assemble. As previously shown, initial host diversity and resource supply determined the trajectory of host community assembly, altering post-assembly host species richness, richness-independent host phylogenetic diversity, and colonization by exotic host species. Throughout community assembly, host richness predicted parasite richness. As predicted, this effect was moderated by exotic abundance: communities dominated by exotic species exhibited a stronger positive relationship between post-assembly host and parasite richness. Ultimately, these results suggest that, by modulating parasite transmission, community assembly can modify the relationship between host and parasite richness, providing a novel mechanism to explain contingencies in this relationship.

Zooplankton communities are influenced by multiple environmental factors, including temperature, nutrient and resource availability, which fluctuate seasonally and across years. While long-term average effects can identify overall drivers, they may overlook dynamic, context-dependent effects that govern short-term changes in diversity and abundance. Understanding and disentangling both perspectives is crucial for identifying and estimating the drivers that shape community structure under varying environmental states. Here, we applied Empirical Dynamic Modeling (CCM, SMap) to a 12-year weekly zooplankton time series to identify causal environmental drivers of taxonomic and morphological diversity and quantify how the influence of each driver shifts over time. We contrast these results with static long-term average effects inferred from Generalized Linear Models which included predictor sets identified using covariate adjustment and accounting for temporal autocorrelation. Drivers linked to long-term average associations differed from those regulating short-term zooplankton dynamics, revealing a decoupling between mean environmental effects and the drivers of temporal variability. Temperature emerged as a persistent regulator of zooplankton dynamics across multiple diversity dimensions, while variables commonly associated with background trophic conditions (e.g. particulate organic matter) were primarily associated with long-term patterns and showed limited dynamical relevance. Importantly, we find evidence for morphological homogenisation in response to short-term fluctuations in chlorophyll a, which was not detectable in long-term average relationships. This contrast highlights that mean environmental associations do not necessarily reflect the mechanisms governing community dynamics. Impacts might be underestimated if average effects appear weak, or misinterpreted if arising mainly from shared trends or seasonality rather than direct mechanisms Integrating both perspectives clarifies the identity and role of environmental drivers, improving inference and prediction of zooplankton community change through time.

Herbivory shapes plant invasion outcomes, yet its role in aquatic plant invasions under changing environmental conditions, such as artificial light at night (ALAN), remains poorly understood. We conducted three experiments using invasive macrophyte Myriophyllum aquaticum, a native macrophyte community (Vallisneria natans, Hydrilla verticillata, M. spicatum), invasive snail Pomacea canaliculata, and native snail Cipangopaludina chinensis to test the combined effects of ALAN and herbivory on non-native macrophyte invasions. ALAN increased M. aquaticum height and total biomass, but had no effect on native species growth. In feeding assays, P. canaliculata consumed all three native species but consistently avoided M. aquaticum under both light treatments. C. chinensis showed no feeding in no-choice assays, but in choice assays, consumed H. verticillata and M. spicatum under No-ALAN and only M. spicatum under ALAN. In community mesocosms, P. canaliculata reduced native macrophyte biomass by 48.0% under No-ALAN and 87.2% under ALAN without affecting M. aquaticum. This selective feeding increased M. aquaticums proportional biomass, with a stronger effect under ALAN than No-ALAN. These results suggest that ALAN can indirectly facilitate non-native macrophyte invasion by amplifying their relative biomass within native communities, particularly in the presence of invasive herbivores, and may promote invasional meltdown through altered feeding preferences.

Virulent pathogens commonly circulate in wildlife populations without causing mass mortality; the triggers of die-offs remain poorly understood. Prevailing frameworks emphasize individual host susceptibility, yet experimental manipulations of susceptibility factors often fail to predict population-level outcomes. We tracked ranavirus epizootics across 40 wood frog breeding ponds over three years, comparing lagged viral state variables against abiotic and host predictors at each epizootic stage. Lagged viral state--environmental DNA concentration and infection prevalence--outperformed abiotic and host predictors of transmission, intensification, and viral accumulation. Infected hosts shed virus into the water column throughout epizootics, but the reciprocal pathway, environmental virus driving new and more severe infections, activated only at the transition to die-off, consistent with a self-reinforcing feedback. The rate of viral accumulation discriminated die-offs, while no static pond or host feature was predictive, reframing mass mortality as an emergent property of pathogen accumulation in shared environments rather than of individual host susceptibility.

1Ecosystem functioning can be maintained in species-rich communities even under fairly severe perturbations. This is because communities with high richness include variation both in species functional roles (functional diversity) and in their responses to environmental changes (response diversity). Response diversity has been proposed as a key mechanism underpinning the stabilising role of biodiversity in variable environments. However, less understood is the role of response diversity in stabilising communities against perturbations and thus preserving ecosystem function. Here, we employ community data from 76 reservoirs across the broad latitudinal gradient of the Japanese archipelago to show that zooplankton assemblages with higher variability in responses to environmental variables can retain functional diversity as species are removed, but that the result depends on the variable examined. We combine empirically derived occurrence data for 47 zooplankton species with biotic and abiotic variables in a joint species distribution model to derive species-specific environmental responses, then measure response diversity to environmental axes including fish community structure and water temperature. We also measure functional trait diversity of zooplankton assemblages and simulate sequential species extinctions, capturing the extinction thresholds beyond which half the functional diversity is lost. Finally, we combine these data streams to show that response diversity can predict higher functional robustness in zooplankton assemblages, but not consistently. The role of response diversity in predicting functional robustness was contingent on the specific metric and environmental variable considered. We found that a balance of positive and negative species responses to water temperature was a significant predictor of robustness, though other metrics and environmental variables mainly yielded non-significant relationships. Overall, we show that response diversity can confer stability to perturbations such as species extinctions, and we demonstrate the utility of species distribution models for measuring response diversity, overcoming mechanistic data limitations and expanding the toolkit available for studying response diversity in natural systems.

AimThree fundamental and inter-related concepts have accrued debates: ecological communities, beta diversity ({beta}), and spatial scale. Spatial scaling of {beta} informs the community concept because the scale of maximal {beta} corresponds to the most apparent size of an ecological community (without invoking external features of habitat, etc.). Here we test five alternative hypotheses about spatial scaling of {beta} for ants, birds, diatoms, and trees across the contiguous USA, using spatial grains from 1 to 106 km2. We compare {beta} scaling among clades and test hypotheses about repeatability where data permit for: (a) summer and winter bird {beta} in six consecutive years; (b) trees through time (4 years, spaced 5 years apart). Finally, we compare different forms of {beta} (i.e., observed and deviations from null models based on spatial heterogeneity and spatial homogeneity). LocationThe contiguous United States of America Time PeriodRecent but varying with clade Taxa Studiedants, birds, diatoms, and trees MethodsWe obtained data from publicly-available sources and assigned point locations to hexagonal grids ranging from 1 to 106 km2. At each spatial grain, we calculated mean pairwise {beta} between each hexagon and its neighboring grids. We also compared alternative {beta} measures and evaluated potential confounding effects of neighborhood size and species richness on results. ResultsSpatial scaling of {beta} repeatedly supported the regional community concept among clades, though with different spatial scales per clade. Based on peak mean {beta}, community size for trees ([~]300 km2) < winter birds ([~]500 km2) < summer birds ([~]2000 km2) {approx} ants ([~]2000 km2) < diatoms ([~]11,000 km2). We note that community scales represent peaks on gradients rather than definitive one-size-fits-all scales. Spatial scaling of {beta} was sensitive to seasonality (birds) and consistent among years for both birds and trees. Also, {beta} deviation from a null model based on spatial heterogeneity adjusted observed {beta} but was less sensitive to neighborhood size and species richness than {beta} deviation based on spatial homogeneity. Main conclusionsResults here indicate that: (a) similar patterns should occur across the tree of life; (b) local ecological and evolutionary forces scale up to form repeatable regional community patterns in ways not yet fully understood; (c) local biodiversity conservation efforts need to be coordinated at biogeographical scales to best achieve goals; and (d) a recent method to calculate {beta} deviation from a null model based on spatial heterogeneity improves {beta} research.

1. The physical structure of an environment potentially influences feeding interactions among organisms, for instance, by providing refuge for prey. We examined how habitat complexity affects the functional feeding response of an ambush predator (damselfly larvae Ischnura elegans) and a pursuit predator (backswimmer Notonecta glauca) feeding on the isopod Asellus aquaticus. 2. We ran experiments in aquatic microcosms with an increasing number of structural elements (0, 2, or 3 rings of plastic plants in different spatial configurations), resulting in five habitat complexity levels. Across these levels, predators were presented with different prey densities to determine the functional response pattern. The experimental design and analysis allowed us to test for effects of structure presence, amount, and complexity level on functional response in one pass, without confounding predictors. 3. The feeding for both predators across all complexity levels was best described by a type II functional response model, and habitat drove feeding strength. Regarding the latter, the predators showed different responses to the complexity treatments. The overall feeding rate of I. elegans was mainly explained by the absence vs. presence of structure. Yet, in the case of N. glauca, feeding rate was strongly dependent on habitat complexity with the predator showing a unique maximum feeding rate (i.e. the inverse of the handling time) for each complexity level and a decreasing attack rate with increasing amount of habitat. 4. On average, prey consumption by both predators was reduced when complex structures were present, compared to the no habitat structure environment (e.g. consumption more than halved for some treatments). Our findings demonstrate that habitat complexity dampens feeding rates and therefore plays a key role in the stability of freshwater ecosystems.

Biological rates, like growth, tend to have unimodal (hump-shaped) responses to temperature, and these relationships can vary among species and biological processes. In most systems, full thermal performance relationships are rarely characterized for interacting species (e.g., consumer-resource or host-pathogen pairs), making it challenging to predict how their interactions, and subsequently, how communities, will shift with climate change. We investigated how the thermal responses of eelgrass (Zostera marina, an important marine foundation species in the N. hemisphere) and an isopod grazer (Pentidotea resecata), which putatively acts as an indirect vector of eelgrass wasting disease, interact to affect eelgrass productivity and wasting disease dynamics. In a laboratory experiment crossing five temperatures, two grazing, and two disease exposure treatments, across various metrics, eelgrass growth responded unimodally to temperature in the absence of grazers. Grazers depressed plant growth and flattened its thermal performance curves. Thermal performance curves for isopods indicated that increases in grazing and survival at intermediate temperatures negated concurrent gains in plant growth at these temperatures, while decreased isopod survival at high temperatures reduced their top-down effect on eelgrass. Isopods had negligible effects on plant disease responses, but warming reduced the time to disease onset and increased final disease severity. Overall, whole-plant disease severity remained low and did not substantially affect eelgrass leaf elongation, net growth, or rhizome dry mass. However, disease-treatment plants grew more new leaves at intermediate temperatures, possibly to combat losses in photosynthetic capacity in diseased leaf tissue. These results indicate that climate change-associated warming will likely increase eelgrass vulnerability to wasting disease. However, in sublethal outbreaks, disease could have less of an impact on eelgrass productivity than warming-induced increases in grazing. Thus, ignoring grazer responses to temperature could result in unreliable predictions of eelgrass productivity under climate change.

The maintenance of parasite diversity has historically taken a host-centric approach. Yet, many parasites are host generalists, and most parasites spend at least some time outside of their hosts. So, what mechanisms besides host-associated niche partitioning allow parasites to coexist? Using a year-long field survey and lab mesocosms, we examined whether environmental niche partitioning or aggregation could enable coexistence among soil-dwelling entomopathogenic nematodes. Field patterns along an elevational gradient reveal that species abundances differentiate with soil structure and moisture levels. Yet, most species strongly overlap within-sites throughout the year. Thus, niche partitioning alone is not sufficient to explain the coexistence of these species and other mechanisms are necessary to explain their coexistence. Aggregation at the within-site scale provides evidence for such a mechanism. Each species showed significant intraspecific clumping and largely random associations with other species. A mesocosm test of the consequences of intraspecific aggregation found that parasites at low or high densities limit their own population growth. Aggregation can promote negative feedback facilitating species coexistence. Our findings offer field-based evidence that spatiotemporal niche partitioning and aggregation both play a critical role in maintaining parasite species diversity, illustrating the importance of extending out view of parasites beyond their hosts.

Global biodiversity is declining at rates faster than at any other point in human history. Experimental manipulations of biodiversity at small spatial scales have demonstrated that communities with fewer species consistently produce less biomass than higher diversity communities. However, understanding how the global extinction crisis is likely to impact global ecosystem functioning will require applying these local and largely experimental findings to natural systems at substantially larger spatial and temporal scales. Here we propose that we can use two simple macroecological patterns - the species area curve and the biomass-area curve - to upscale the species richness-biomass relationship. We demonstrate that at local spatial scales, each additional species will contribute more to biomass production with increasing area sampled because the species-area curve saturates and the biomass-area curve increases monotonically. We use species-area and biomass-area curves from a Minnesota grassland and a Panamanian tropical dry forest to examine the species richness - biomass relationship at three and ten sampling extents, respectively. In both datasets, the observed relationship between biodiversity and biomass production at every sampling extent was predicted from simple species-area and biomass-area relationships. These findings suggest that macroecological patterns like the species-area curve underpin the scaling of biodiversity-ecosystem functioning research and can be used to predict these relationships at the global scales where they are relevant for species loss.

Species tolerating the same environmental conditions can potentially colonize and thrive in the same habitats and eco-regions. Are any pair of those species equally probable to co-occur in the same community? Can we quantify the propensity of two species to co-occur together? Here, we focus on a simple but largely overlooked community-level pattern: the co-occurrence-occupancy curve, which relates the tendency of species to co-occur with others to their total occupancy across sites. We first define this empirical curve and then derive its expected shape under a random null model that assumes site equivalence and species independence. Building on these results, we introduce the Species Association Index (SAI), an occupancy-standardized measure that quantifies the tendency of a species to associate with others independently of its overall frequency of occurrence. The SAI enables meaningful comparisons among species with contrasting occupancies and provides a transparent benchmark against which departures from neutrality can be assessed. We illustrate the approach using two contrasting systems--tropical rain forest trees on Barro Colorado Island and organisms from Mediterranean rocky shores--highlighting both the generality of the co-occurrence-occupancy framework and its limitations.

The exchange of energy and organisms across habitat boundaries links aquatic and terrestrial ecosystems and sustains ecosystem functioning. Although disturbance may disrupt these linkages, the mechanisms at play remain poorly understood. Here, we investigated the extent to which flow intermittency may disrupt riparian-aquatic ecosystem linkages by altering consumer communities in the recipient ecosystem or by altering resource quality in the donor ecosystem. We ran an experiment in an intermittent river network in California, focusing on a critical forest-to-river subsidy (organic matter in the form of leaf litter), its transformation, and its reciprocal benefit (aquatic insect production). Using three riparian species (willow, cottonwood, and oak) at sites spanning a gradient of flow permanence, we quantified intraspecific plasticity in leaf traits (specific leaf area, nitrogen and phosphorus concentrations, and {delta}13C), measured decomposition rates, and estimated the secondary production of aquatic shredders (Plecoptera). Across all leaf species, decomposition rates were 16-36% lower at intermittent than perennial sites, an effect largely driven by intraspecific leaf trait plasticity rather than changes in consumer abundance. At high flow intermittency, willow experienced water stress (enriched {delta}13C) and reduced specific leaf area, while cottonwood showed primarily stoichiometric responses (reduced leaf nitrogen and phosphorus). Despite these divergent strategies, all species produced lower-quality litter at intermittent sites. Variance partitioning confirmed that initial litter quality uniquely explained 51.5% of variation in decomposition rates, more than double the contribution of invertebrate community metrics; and structural equation modeling revealed that both leaf traits and stonefly (Plecoptera) secondary production significantly predicted decomposition rates, with leaf traits exerting the stronger effect. Notably, stonefly secondary production was 37-98% lower at intermittent sites across leaf species. Because these insects later emerge as terrestrial adults, they provide a significant energy flux to riparian predators, and, thus, impoverished litter quality suppresses the reciprocal transfer of energy back to terrestrial food webs. As drought intensifies globally, the decoupling of terrestrial-aquatic linkages may begin in the riparian canopy.