Oikos

Habitat fragmentation, driven by human activities, disrupts habitat connectivity and alters ecological processes through geometric and demographic fragmentation effects. Dispersal plays a fundamental role in shaping the distribution, abundance, and persistence of species in modified landscapes. While previous research looked at the evolution of dispersal strategies at the species level, community-level dynamics remain underexplored. Species exhibit diverse dispersal strategies to persist in modified landscapes, yet predicting how these strategies interact at the community level requires a more integrated approach. This study employed an individual-based simulation model to explore how fragmentation and other landscape characteristics influence community-level dispersal strategies. We tested the effects of varying fragmentation levels, environmental autocorrelation, habitat amount, and disturbance levels on the emerging distribution of dispersal distances within a community in modified and continuous landscapes. We hypothesised that fragmentation and other spatial patterns would significantly shape community composition, favouring particular dispersal strategies under specific environmental conditions. The findings reveal that higher disturbance levels and greater habitat amount increased the community-weighted mean of dispersal distance, while fragmentation showed only minor variation. Additionally, low autocorrelation was associated with the highest community-weighted mean of dispersal distance. These results highlight the importance of considering community-level dynamics when predicting ecosystem responses to landscape modification. By clarifying how landscape structure and disturbance shape community-level dispersal strategies, this study advances our understanding of the mechanisms underlying species persistence and community structure in modified landscapes.

The phenology of organisms worldwide is shifting in response to changes in environmental conditions. There is growing concern that resulting timing mismatches among interacting species will negatively impact system-level properties, yet there is no general framework for evaluating community responses to changes in phenology. To address this gap, we developed a mathematical framework based on local stability analysis and used it to assess the resilience implications of phenological perturbations with a multi-year, highly time-resolved empirical dataset on subalpine plant-pollinator communities. The forecasted effects of phenological perturbations were largely independent of perturbations to species densities, indicating the potential for even small changes in phenology to disrupt the functioning of ecosystems that are otherwise highly stable.

Nutrient availability, ecosystem productivity, and consumer assemblages are intricately linked through complex interactions and feedbacks. Nutrients influence the diversity and functional roles of consumers via shifts in resource quality and quantity, and consumers can alter ecosystem production and nutrient availability. However, our understanding of how characteristics of consumers respond to and influence concomitant shifts in nutrient availability and production is limited. We quantified the response of well-studied consumer assemblages (benthic invertebrates and zooplankton) to realistic nutrient loads that altered gross primary production (GPP) and ecosystem respiration (ER). We fertilized 14 outdoor experimental ponds for 2 months and monitored total water column carbon (TC), nitrogen (TN), and phosphorus (TP), GPP, ER, and net ecosystem production (NEP) weekly. Then, we evaluated how fertilization and the variation in nutrients and metabolism caused by fertilization were related to shifts in consumer assemblages. Fertilization increased water column TN and TP and reduced TC:TP ratios, TN:TP ratios, and rates of GPP and ER. However, consumer assemblages were more tightly linked to variation in nutrient availability and production across ponds than to fertilization. Greater declines in benthic diversity occurred in ponds with higher average TN:TP ratios during the experiment. Consistent with predicted effects of cladocerans on nutrient availability, shifts in cladoceran abundances were positively associated with average water column TN:TP ratios during the experiment. Finally, elevated GPP and ER were associated with greater increases in the abundance of benthic invertebrate predators, suggesting the possibility of top-down control. Our study highlights the critical role of consumer-mediated processes in the interaction between nutrient availability and production. Manuscript HighlightsO_LIFertilization reduced pond gross primary production and ecosystem respiration rates. C_LIO_LIInvertebrate predator abundance was inversely related to gross primary production. C_LIO_LIShifts in consumer assemblages were tightly linked to nutrients and production. C_LI

Interaction networks, in which nodes represent species and edges represent direct interactions between species, have a long and impactful history in community ecology. However, co-occurrence networks, where edges represent statistical relationships among species presences or abundances, are often easier to construct from lab and field data. It is clear that co-occurrence edges often do not represent direct interactions, but frameworks for the interpretation of co-occurrence networks have not kept pace with their generation. It is therefore unclear when and how these networks can be used to gain insight into community dynamics. Here, we use a Generalized Lotka-Volterra-based model to explore the contexts in which emergent properties of species interaction networks are identifiable in their resulting co-occurrence networks. We find that, in spite of many differences in direct edges, key features of the true interaction network, such as unipartite modularity, high-degree nodes (hubs), and bipartite modularity and nestedness, can be preserved in co-occurrence networks. In contrast, node degree distributions are not preserved even in the most idealized scenarios. We propose that networks derived from large co-occurrence datasets could therefore be used in future empirical work to test existing hypotheses of how emergent network structures drive ecological community dynamics.

Metacommunity theory explains how species distributions arise from local population dynamics and dispersal between habitat patches. Four conceptual paradigms--patch dynamics, species sorting, mass effects, and demographic stochasticity--have emerged as frameworks for understanding metacommunity structure and dynamics, but their integration remains an open problem. Here we introduce a probabilistic-stochastic-deterministic (PSD) modelling framework that unifies these paradigms within a single mathematical description. PSD approximates individual-based models (IBM) with computational efficiency comparable to ordinary differential equations (ODE) while capturing demographic stochasticity and permitting analytical treatment. Through validation against IBM simulations in single-patch communities and spatially explicit metacommunities with rock-paper-scissors dynamics, we demonstrate that PSD accurately reproduces IBM behaviour where ODE models fail, specifically when demographic stochasticity dominates during immigration. For metacommunities with long-distance dispersal, we analytically derive the period of a slow collective oscillations, revealing body-mass and dispersal-rate dependencies invisible to ODE theory. Our analysis shows that the four paradigms represent valid descriptions in different regions of parameter space, controlled by individual body-mass, immigration rate, and regional species richness. The PSD framework thus provides both a practical simulation tool and an analytical machinery for predicting metacommunity dynamics across ecological regimes.

Patterns in the vegetation across arid and semiarid regions may be explained as a form of self-organization driven by water scarcity, and are often modeled through reaction-diffusion dynamics. Recent work has shown that similar mathematical models generate patterns on networks. However, these studies have focused on idealized topologies with no reference to natural pattern-forming systems. Our study aims at bridging these two fields: we employ a physical reaction-diffusion vegetation model, and gradually modify the topology of the diffusion network by adding random shortcuts over a 2-dimensional grid, interpolating between a regular lattice and a random network. We found that network topology strongly shapes both the resulting vegetation patterns and the precipitation range that supports them. Three behavioral regimes emerge. On a regular lattice, high-regularity patterns develop reflecting local diffusion processes. On a random network, the system is dominated by global pressure towards homogenization yielding either a uniform state or a single patch. In the intermediate shortcut density range, as the network topology resembles a small world network, the interaction between the two scales of diffusion generates two kinds of disordered patterns: low-regularity patterns with a well-defined characteristic wavelength, and irregular patterns characterized by a broad patch size distribution. These disordered patterns resemble real-world observations and, in our model, they show different responses to changing precipitation. Although we focused on dryland vegetation, we suggest that network-mediated diffusion could lead to similar mechanisms in a wide variety of pattern-forming systems. HighlightsO_LIWe study vegetation pattern formation over different diffusion network topologies. C_LIO_LITwo kinds of stable disordered patterns states develop over small world topologies. C_LIO_LILow-regularity patterns with a well-defined characteristic wavelength. C_LIO_LIIrregular patterns characterized by a broad patch size distribution. C_LIO_LIThese different kinds of disordered states show different relations to precipitation. C_LI

O_LISpecies diversity potentially has a dual effect on communities: a generally positive effect on overall community biomass, reflecting the expression of species response and interaction traits, and a poorly characterised effect on mass-specific species contribution to ecosystem functions, reflecting the expression of their effect traits. Disentangling the effects of biodiversity on total biomass from those on effect trait expression would help settle a long-standing debate by clarifying how biodiversity relates to both facets of species effects on ecosystem functioning. C_LIO_LIFollowing the classical BEF approach, we calculate expected ecosystem function based on observed functioning in monoculture. We then derive a net biodiversity effect (NBE) and decompose it into four components: the classical complementarity and selection effects on total community biomass, and complementarity and selection effects on effect trait expression. The latter two reflect, respectively, a complementarity or facilitation in how effect traits influence the function, and how species with the highest potential for increasing the function become dominant in the community. C_LIO_LIWe illustrate this NBE decomposition with three ecosystem functions (nitrogen retention capacity, soil hydraulic conductivity improvement, and forage digestibility) measured in assembled communities under controlled experimental conditions of perennial grassland plants. Regarding nitrogen retention, we find a positive complementary effect via total biomass, but a negative biodiversity effect via effect trait expression. For hydraulic conductivity improvement, biodiversity effects are mostly mediated by total biomass. As for forage digestibility, we found a positive complementarity effect on trait expression, outweighed however by a negative selection effect. This analysis reveals how biodiversity may have contrasting effects on ecosystem functions via its impact on biomass and effect trait expression. C_LI SynthesisSeparating between the effect of biodiversity on plant community biomass and on effect trait expression at the community level is one important step towards understanding the pathways by which diverse plant communities drive ecosystem functioning.

The ongoing decline in biodiversity highlights the need for understanding the causes of population changes. This study uses 25-year, large-scale monitoring dataset to investigate the influence of climate and landscape structure on the annual population growth rates of 84 bird species across Poland. Our methodological framework involves the spatiotemporal decomposition of these environmental drivers to decouple demographic effects of long-term carrying capacities from the short-term effects of environmental perturbations. Using species-specific demographic models followed by a community-wide meta-analysis, we evaluated how individual species responses scale up to shape community-level dynamics. The results reveal significant variation in species-specific responses to individual drivers. At the community level, our findings suggest that bird populations are mainly regulated by the long-term spatial constraints rather than short-term disturbances. Persistent environmental heterogeneity had the strongest positive demographic effect on birds, followed by temperature, forest dominance over croplands, and precipitation. In contrast, rapid temporal shifts in environmental heterogeneity and precipitation anomalies negatively affected population growth, whereas urbanisation consistently exerted a negative effect across both spatiotemporal dimensions. Our results highlight the significance of protecting existing heterogeneous and ecotonal habitats, as well as the need to incorporate features that enhance habitat heterogeneity into urban development. Article impact statementPreserving heterogeneous habitats is essential for the conservation of bird populations.

Human activities are driving unprecedented environmental change, yet assessments of ecosystem resilience often overlook the rapid pace of change in the Anthropocene. Predator-prey systems are sensitive to the rate of environmental change and the whole system can collapse if predator population fail to promptly adjust to environmentally-driven shifts in resource population. Here, we investigate how different combinations of predator responsiveness and rates of environmental change influence the system vulnerability to critical transitions, explicitly addressing its interplay with magnitude of change. We found that, as predator responsiveness decreases, relatively slower rates and smaller magnitudes of environmental change leads to system collapse. Hence, even low and seemingly inoffensive total magnitudes of environmental change can be catastrophic if the rate of change is beyond a critical threshold. We propose considering predator responsiveness and current rates of environmental change as crucial factors in predicting the Anthropocenes impact on ecosystems.

The Biodiversity-Ecosystem Functioning (BEF) literature has shown species diversity to be essential for ecosystem functioning and services. Yet although acquiring information through interspecific networks can impact ecosystem functioning, it is unclear how it is modulated by species diversity. Eliciting vocal responses using predator models across a latitudinal gradient, we first show that the species diversity of birds increases public information about predation both in the low-cost system of mobbing and in the higher-cost system of alarm calls. A similar result was also found across a fragment area gradient for mobbing; this system was then used to test how species diversity affects interspecific information flow in mobbing communities. We set up two BEF playback experiments, manipulating the species richness level of the playback sound files by varying the number of species producing mobbing calls (one, two, four, eight species). In an experiment in which the call rate across treatments was held constant, and only heterospecific responses were counted, increasing species richness of the sound files increased the number of species and individuals responding, the number of calls produced and their frequency range, and decreased latency to call. An experiment in which call rate increased with the addition of species in each treatment showed a similar, but stronger pattern. There was little evidence that the signals of one particular species changed responses. This supports the hypothesis that the species diversity of a community is a key component influencing the quantity and quality of information flow inside it.

Spatial heterogeneity of abiotic resources is essential for species coexistence. Ecological theory often assumes predefined heterogeneity of resources that constrains community dynamics, but the recent developments of meta-ecosystem ecology and zoogeochemistry highlight nutrient patterns could result from the interactions between the activities and movements of organisms and their abiotic environment. Here we investigate the mechanisms by which biotic-abiotic feedbacks could generate nutrient spatial heterogeneity in a simple plant-herbivore occupancy model where populations forage, recycle, and disperse in a homogenous landscape. By systematically varying organisms ranges of foraging and dispersal, and recycling levels, we found that limited dispersal of plants plays a key role on the emergence of nutrient patchiness by favoring small clusters of vegetation that shape their environment through consumption and recycling. However, herbivores could also create nutrient spatial heterogeneity when large foraging and dispersal ranges, and high recycling, allow them to efficiently track plant hot spots and to increase population persistence. Unexpectedly, strong aggregation of herbivore populations did not necessarily result in nutrient clustering. Rather than via recycling, herbivores mainly affected nutrient distribution indirectly, through their top-down impact on plant distribution. When evenly spread in the landscape, herbivore populations with large foraging ranges created areas of strong herbivory pressure unfavorable to plant colonization where nutrient can accumulate. These results can help understand the dynamical feedback between biota and abiotic resources. In a context where human activities alter both nutrient distribution and species abundances, a better understanding of this biotic-abiotic feedback will be key to anticipate the response of ecosystems to current perturbations.

The ecological and evolutionary processes determining species range limits remain poorly understood. Ultimately, range limits depend on the species abilities to persist under heterogeneous conditions, by adaptive differentiation and phenotypic plasticity, including transgenerational effects. To investigate ecological differentiation and transgenerational effects in the clonal invasive knotweed, Reynoutria japonica, in Europe, we conducted a two-phase transplant experiment: plants sampled along the entire latitudinal gradient were planted in three sites located at the northern range margin, mid-range and near the southern range margin, and then re-transplanted among all three sites after two years. Biomass production and allocation were generally not associated with latitude of origin and previous growth at the same site did not promote performance. We therefore find no evidence that adaptive differentiation or transgenerational effects contribute to the wide distribution of R. japonica in Europe. However, at the northern site, with a 25% shorter season, knotweed plants invested much less biomass below-ground, and the pattern was further strengthened in plants that had grown in the northern site in the previous generation. Overwintering below-ground rhizomes are essential for survival and spread. We further explored limiting climate conditions in a species distribution model for the European range and found that mean annual temperature and temperature annual range are the main predictors of the European distribution of R. japonica. Taken together, our study suggests that low temperatures and associated short seasons may pose a limit to the broad environmental tolerance of R. japonica and restrict its northward spread by reducing below-ground biomass accumulation.

Ectomycorrhizal (EcM) fungi are well-recognized symbionts impacting tree health and ecosystem functioning globally, yet understanding of their timing of proliferation in soils across seasons and years remains limited. We analyzed monthly patterns of EcM fungal abundance and community structure over two years in five temperate monodominant forest plots via quantitative PCR and Illumina sequencing. We found that the phenological dynamics of EcM fungi differed significantly by host tree leaf habit, fungal exploration type, fungal genus, and soil moisture. Overall, total EcM fungal abundances based on qPCR consistently peaked in autumn, and were more dynamic in evergreen than deciduous plots, supporting ideas of surplus carbon and asymmetric above-belowground dynamics. Longer-distance exploration types peaked earlier and were more stable than shorter-distance types, suggesting an independent and supportive role in releasing spring nutrients. About half of 20 focal taxa consistently peaked in either autumn, summer, or spring, while others were either host- and/or year-dependent. Our findings highlight that phenology is a key EcM fungal trait best explained by both host and fungal contributions, and future studies across biomes should consider seasonal shifts and sampling to elucidate phenological traits. Summary- The timing of belowground production and seasonal community dynamics remain poorly understood for ectomycorrhizal (EcM) fungi. - We collected soils monthly for two years from five temperate monodominant forest plots. - Fungal production peaked in autumn, shorter-distance and evergreen-associated spanned wider ranges, and half of focal fungal genera showed seasonal preference, emphasizing autumn surplus carbon and spring nutrients from long-distance types. - Future studies should consider seasonal shifts when sampling EcM fungal communities, and forest carbon models should include asymmetric above-belowground phenology. Translated Summary (Spanish)- La fenologia de la produccion y composicion de comunidades de hongos ectomicorrizicos (EcM) es poco estudiada. - Recolectamos suelos mensualmente por dos anos de cinco parcelas mono-dominantes templados. - Produccion maxima de hongos ocurrio en otono, hongos asociados con arboles siempreverdes y de exploracion de corta-distancia observaron rangos mas amplios, y la mitad de generos de hongos focales observaron preferencia estacional, enfatizando extra carbono en otono y nutrientes en primavera de tipos larga-distancia. - Estudios deben considerar cambios estacionales para el muestreo de hongos EcM, y modelos de carbono deben incluir fenologia asimetrica entre hojas y hongos. Plain language summaryEctomycorrhizal fungi are critical for the global carbon cycle, but their seasonal and inter-annual growth patterns remain unclear. We sample soil DNA monthly over two years across five different monodominant temperate forest stands. We find an overall belowground peak in autumn, with significantly later growth under wetter conditions, more dynamism with evergreen trees, and distinct spring growth by longer-distance fungi.

Understanding the drivers of biodiversity in the worlds rivers, which are known to be hyperdiverse relative to their coverage area, has been an enduring goal in ecology. While regression-based empirical studies have identified a suite of environmental factors that are correlated with riverine fish biodiversity, these insights are often system-specific and inconsistent across regions. In contrast, a more limited body of studies have suggested that network connectivity of rivers affects fish biodiversity by limiting dispersal, and basin shape may modulate these relationships. The few theoretical papers that have explored basin morphology effects have tended to use extreme network shapes and inconsistent methods, thus limiting general insights. Here, we build on these results to demonstrate that river basin morphology, as measured by log aspect ratio, can alter both network connectivity and biodiversity in simulated, all-else-equal scenarios. First, we quantify variation in log aspect ratio across the worlds 100 largest rivers and use this empirical range of shape variation to guide synthetic experiments. In particular, we use Optimal Channel Networks (OCNs) constrained to basins with log aspect ratios within realistic range to study how shape alters connectivity profiles when node number and basin area are held constant. By coupling OCNs with dendritic neutral models, we demonstrate that variation in aspect ratio and concomitant changes in connectivity lead to substantial changes in simulated biodiversity. Finally, we use Earth Movers Distance to establish that basin-shape-induced changes in node-level connectivity distributions are predictive of transformations in node-level distributions of and {beta} diversity. Overall, elongated basins such as the Mekong River feature lower species richness (-diversity), higher turnover ({beta}-diversity), and less variable distributions of both quantities relative to a square reference basin. Furthermore, approximately one third of the worlds largest rivers are elongated enough to potentially feature statistically-detectable, shape-mediated variation in connectivity and biodiversity.

Stingless bees are key pollinators in tropical ecosystems, yet their ecological dynamics remain poorly understood in highly seasonal environments such as the seasonally dry tropical forests of Ecuador. These ecosystems experience pronounced climatic seasonality, with sharp transitions between dry and wet periods that strongly affect floral resource availability. Understanding interspecific competition and niche partitioning in such systems is critical, particularly given the global decline of pollinators. We investigated resource use and niche dynamics in two native stingless bees, Melipona mimetica and Scaptotrigona sp., by quantifying pollen, nectar, and resin collection across seasons. Log-linear models were used to test the effects of species, season, and their interaction on resource use, while non-metric multidimensional scaling (NMDS) assessed niche overlap. Contrary to the expectation that niche overlap increases under resource scarcity, we found greater overlap during the wet season, when resources are more abundant. This suggests that both species converge on high-quality floral resources during peak availability, reflecting an adaptive response to strong environmental seasonality. Pollen use remained stable across seasons, consistent with generalist foraging behavior. In contrast, nectar collection increased significantly during the wet season, while resin exhibited a shared seasonal peak, likely associated with synchronized nest construction or maintenance. These findings reveal context-dependent competition dynamics and highlight the role of environmental seasonality in shaping pollinator interactions. Our study provides new insights into the ecology of threatened stingless bees and contributes to their conservation in tropical dry forest ecosystems.

O_LIAgricultural intensification can support the expansion of introduced species which are highly adapted to human-modified landscapes, but the mechanisms by which this occurs are often unclear. C_LIO_LIHere we investigate the spatial ecology of a rapidly expanding introduced bumble bee (Bombus impatiens) and a native congener (B. mixtus) in agricultural landscapes of southwestern British Columbia, Canada. We used microsatellite genotyping and spatially explicit capture-recapture models to compare the foraging distance of the two species, and fitted hierarchical models to compare their abundance, behaviour (nest searching vs foraging), and lineage survival as a function of landscape composition and configuration. C_LIO_LIWe found that B. impatiens had a broader foraging range than B. mixtus, and that its colony/worker abundance were positively associated with the surrounding area of residential gardens, but decreased relative to B. mixtus abundance in response to increasing seminatural area. In contrast, B. mixtus colony abundance decreased in landscapes with a greater area of intensively managed berry crops. C_LIO_LIWe observed fewer B. impatiens queens per survey in landscapes with more low-disturbance landcover, and hypothesize space use of this species could be shaped by concentration on potential nesting habitat. Consistent with this observation, nest searching behaviour was more common for B. impatiens queens, while B. mixtus queens were primarily observed foraging, suggesting these two species derive different value from agricultural landscapes during colony establishment. C_LIO_LIFinally, we found that the rate of lineage re-capture between 2022 colonies and 2023 spring queens was nearly 10-fold higher for B. impatiens than for B. mixtus, indicating a greater capacity of the introduced species to complete its life cycle in agro-natural landscape mosaics. C_LIO_LIOur results suggest that differences in spatial ecology may contribute to the differential success of these two species in human-modified landscapes, and provide insight into the mechanisms by which land-use change shapes community composition. C_LI O_FIG O_LINKSMALLFIG WIDTH=184 HEIGHT=200 SRC="FIGDIR/small/723627v1_ufig1.gif" ALT="Figure 1"> View larger version (62K): org.highwire.dtl.DTLVardef@1e72eacorg.highwire.dtl.DTLVardef@a958a0org.highwire.dtl.DTLVardef@1f970b6org.highwire.dtl.DTLVardef@156f522_HPS_FORMAT_FIGEXP M_FIG C_FIG Graphical abstract. Coloured diagrams of B. mixtus and B. impatiens are credited to Elaine Evans and the Xerces Society, with permission.

Plants, as structural elements of habitats, contribute greatly to the maintenance of local biodiversity through their biological interactions. In this study we explore whether their rarity, according to Rabinowitzs (1981) three criteria, is related to the richness and diversity of arthropods and other plants they are associated to, in a gypsum-rich steppe. We first analysed whether the geographic abundance and ecological specialisation of 32 characteristic and dominant plant species are related to the diversity (richness and phylogenetic diversity (MPD)) and degree of local specialisation of arthropods associated with them (1,694 taxa). Then, we focused on a non endemic and non specialized plant in the study area (Krascheninnikovia ceratoides) to explore the effect of population size on two types of interactions: aerial arthropods and plant facilitation. Results indicate that: 1) plant species abundance (geographical range) is not related to the richness or MPD of communities of associated arthropods, 2) plant species ecological specialization (edaphic endemisms or gypsophiles) do not contribute differentially to the maintenance of singular arthropod communities, and 3) the community of aerial arthropods and plants interacting with K. ceratoides in a small population are not necessarily less diverse than those in patches of similar size in a large population. Results also revealed that the two plant species with fewer interactions (one rare, one widespread) do show the highest singularity in their interactions with arthropods. Our study illustrates the important contribution of rare plants to the conservation of local biodiversity.

1. Biotic insular systems differ from conventional islands because patch attributes change dynamically as patch-forming organisms develop. It therefore remains unclear whether the assembly mechanisms predicted by island biogeography theory (IBT) operate in such systems. Here, using epiphytic birds nest ferns (BNFs, Asplenium nidus) as a model biotic island system, we tested whether fungal and bacterial community diversity conform to species-area relationships predicted by IBT. With a stratified sampling scheme, we further evaluated the underlying mechanisms (passive sampling, disproportionate effects, and environmental heterogeneity) of species-area relationships, and assessed isolation effects using distance-decay patterns in community similarity. 2. We treated each BNF individual as a microbial island and categorized 24 BNFs into three size classes. Microbial and humus samples from multiple litter layers within each BNF individual were collected; microbial communities were characterized using next-generation sequencing, and humus chemical properties (pH and C:N ratio) were measured to characterize microhabitat conditions. To investigate mechanisms underlying species-area relationships, we applied a multi-scale rarefaction framework to partition diversity components. Spatial distances among BNFs were quantified to evaluate isolation effects. 3. Consistent with IBT predictions, both fungal and bacterial communities exhibited positive species-area relationships, indicating that larger BNFs harbored greater microbial richness. Diversity partitioning suggested that fungal richness increased through both disproportionate effects and environmental heterogeneity, whereas bacterial richness was primarily driven by environmental heterogeneity. Within larger ferns, greater heterogeneity in litter pH was associated with increased species turnover across litter layers, suggesting that decomposition-driven pH gradients create diverse microhabitats that promote microbial diversity. In addition, both microbial communities exhibited distance-decay patterns, indicating that isolation contributes to community assembly through dispersal limitation. 4. Synthesis. Our results demonstrate that BNFs function as a biotic insular system, in which both patch size and spatial isolation structure microbial diversity, consistent with predictions from IBT. Furthermore, we show that environmental heterogeneity generated by the growth of the habitatforming BNF mechanistically links island area to microbial diversity. Our study integrates both local habitat heterogeneity and regional spatial structure, highlighting the potential to extend IBT and metacommunity theory to organism-formed habitats.

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.

Biodiversity patterns in tropical freshwater ecosystems remain unevenly understood, particularly in high-integrity regions such as Indigenous territories. In this study, we assessed taxonomic and functional beta diversity of Ephemeroptera, Plecoptera, and Trichoptera (EPT) in Amazonian streams located within the Panara Indigenous Territory, Brazil. We evaluated the relative contributions of local environmental variables, spatial processes, and landscape context to beta-diversity patterns. We disentangled the roles of replacement and richness differences across taxonomic and functional dimensions. EPT larvae were sampled in 31 streams during the dry season. Beta diversity was quantified using Sorensen-based dissimilarity indices, and functional dissimilarity was calculated from seven ecological traits using Gower distances. Taxonomic beta diversity was dominated by genus replacement and was jointly structured by local habitat variables and spatial components, indicating the combined influence of environmental filtering and dispersal limitation. In contrast, functional beta diversity was higher than taxonomic beta diversity and was predominantly structured by richness differences, with significant effects of local environmental variables but no detectable influence of spatial processes. This pattern indicates a decoupling between taxonomic and functional dimensions, suggesting high levels of functional redundancy among EPT genera across streams. Our findings demonstrate that Amazonian streams within Indigenous territories provide key systems for understanding community assembly processes under low levels of direct anthropogenic disturbance. By revealing contrasting mechanisms underlying taxonomic and functional beta diversity, this study underscores the importance of integrating multiple facets of biodiversity and reinforces the role of Indigenous territories as strategic landscapes for safeguarding Amazonian freshwater biodiversity.