Oikos

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.

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.

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 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.

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.

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.

By concentrating rodents along verges, roads can reshape rodent-mediated seed dispersal, yet empirical tests remain scarce. We conducted a two-year field experiment in Mediterranean oak woodlands in southern Portugal to test how seed dispersal varies with distance from roads across road type (paved vs. unpaved) and road-forest context (edge vs. non-edge). We tracked labeled holm oak acorns, recording dispersal distances and the number of dispersal events. The two metrics responded differently to road distance. Dispersal distances changed little with distance from roads in non-edge contexts but increased in edge road-forest contexts (2x longer at 400 m than at 10 m) and showed a year x distance-to-road interaction, with longer dispersal distances farther from roads in the second year (a poor mast year). Dispersal distances were also longer when acorns were deposited under shrubs and in areas of higher tree density, and decreased with greater natural acorn availability. In contrast, the number of dispersal events declined with distance from roads (30% more events at 10 m than at 400 m) and was higher along unpaved than paved roads (39% more events). Dispersal frequency also increased in the poor mast year and with shrub cover. No acorns crossed the road. Thus, road verges can concentrate rodent seed handling but do not increase dispersal distances near roads nor provide cross-road seed connectivity; instead, dispersal outcomes depend on edge context, road type, and microhabitat structure. Management that retains structural cover at verges and the adjacent forest edge (e.g., shrub patches and non-uniform clearing) can harness verge-associated activity to increase acorn deposition in sheltered microsites and promote regeneration farther into forest interiors in roaded landscapes.

Theory suggests that different components of environmental fluctuations, from daily and seasonal cycles to multidecadal trends, can have distinct and even opposing effects on species abundances and community dynamics, depending on their specific adaptations. But empirical research that deconstructs the influence of these different cycles on communities is lacking. Here, we used long-term biological monitoring data together with flow records of rivers across New Zealand to (i) investigate the role of fast, slow, and seasonal river-flow fluctuations in structuring macroinvertebrate communities; and (ii) to assess whether life-history and mobility traits mediate the response. Using joint species distribution models, we found striking differences in taxon and community responses to the different components of river flow variation. Responses to slow fluctuations were generally stronger and better predicted by traits, while responses to seasonal fluctuations were highly heterogeneous. Fast increases in flow, typical of flooding events, had pervasive negative effects on species abundances, but the severity of impact partly depended on mobility traits. Our results suggest that different ecological mechanisms underpin the response to distinct environmental fluctuations, highlighting the value of jointly considering multiple temporal scales of variation and species functional traits to understand and predict how communities reorganise under fluctuating environmental regimes.

The Allee effect is a phenomenon where individual fitness is reduced in small populations, for example because of mate-finding difficulties or increased predation. Allee effects matter in conservation biology because they can drive small populations to extinction. The severity of Allee effects can depend on traits such as mate-search rate and defense against predators. Many natural populations exhibit considerable intraspecific trait variation (ITV) in such traits, but most studies so far assume these traits to be constant. Thus the impact of ITV on populations with Allee effect is largely unknown. Here we create two individual-based stochastic models that simulate a small population experiencing either a mate-finding Allee effect or a predator-driven Allee effect. We analyze how ITV, trait inheritance, and mutation affect the proportion of surviving populations. Under the mate-finding Allee effect, higher ITV hindered population survival and increased Allee thresholds. This can be explained by Jensens inequality and the negative curvature of the mate-finding function. Under the predator-driven Allee effect, ITV effects were weak, but higher mutation standard deviations were beneficial, likely because they provided more substrate for selection to act on. We thus recommend to take into account ITV when dealing with threatened populations with an Allee effect.

O_LIChanges in habitat structure across species distributions may contribute to the generation and maintenance of range limits, but few studies have evaluated this by directly measuring habitat availability across relevant spatial scales. C_LIO_LIHere, we test the predictions that coarse-scale and patch-level habitat availability decline towards and beyond the northern range limit of Pacific coastal dune endemic Camissoniopsis cheiranthifolia. We used aerial imagery and geographic information system (GIS) tools to measure the coarse-scale availability of coastal dune habitat in California and Oregon. The availability of finer-scale habitat patches specifically suitable for C. cheiranthifolia was measured in a 2-generation field survey of > 4,200 5m x 5m plots randomly distributed across 1100 km of coastal dune habitat transcending the species northern range limit. At each plot, we estimated the proportion of area that contained suitable habitat as well as recorded occupancy by C. cheiranthifolia. As an alternative approach to visually estimating habitat suitability, we recorded plant community composition at each plot to predict beyond-range habitat suitability using a random forest model. C_LIO_LIContrary to our predictions, we found that coastal dune habitat, measured coarsely from aerial imagery, was more abundant and continuous towards and beyond the northern range limit. At the fine scale, however, the proportion of plots with suitable habitat (patch suitability) and the proportion of habitat within plots that was suitable (patch size) declined across the range limit. Moreover, patches were more isolated from one another and, in one survey year, less temporally stable towards and beyond the range limit. Finally, occupancy by C. cheiranthifolia was less likely in smaller, more isolated, and temporally unstable patches, providing mechanistic insight to the previously observed decline in occupancy towards the range limit. C_LIO_LISynthesis: Taken together, our results suggest that fine-scale habitat patch configuration changes in ways that likely impede patch colonization, thereby reducing occupancy and limiting the species northern distribution. Thus, consideration of geographic variation in patch and landscape structure, rather than only coarse-scale habitat availability, may be essential for understanding the processes that limit species ranges. C_LI

Artificial pond construction is widely used in wetland restoration, yet biodiversity outcomes depend strongly on design and subsequent management. We tested how different regimes (grazing, mowing, and no management) influence habitat structure, water conditions, and aquatic macroinvertebrate diversity in newly excavated experimental ponds within an eutrophic wetland in South Moravia (Czechia). Across four focal groups (Mollusca, Odonata, Coleoptera, Heteroptera), we observed rapid colonisation of the newly built ponds. Species richness and densities rose during early development, dropped after drying events, and then partially recovered, indicating repeated "resetting" of communities under fluctuating hydrology. Periodic drying also prevented fish stock establishment. Management significantly affected species composition and both grazed and mowed ponds displayed higher densities (abundances) than controls, but differed only slightly in terms of species richness. The grazed ponds were characterised by high sunlight exposure, reduced reed dominance, and trampling-generated high littoral heterogeneity. These ponds showed highest numbers of taxa adapted to shallow and warm waterbodies, muddy substrate, semiaquatic microhabitats, or newly emerged and disturbed habitats. The mowed ponds promoted dense submergent vegetation, supporting Odonata representation and other taxa requiring aquatic vegetation. The control ponds remained highly shaded by high-grown reed, organic-matter rich, hosting a set of taxa tolerant of low-light, low-oxygen conditions. At the wetland scale, multiple small ponds increased overall diversity through high between-pond heterogeneity. Our results highlight that pond construction alone is insufficient for wetland restoration: follow-up long-term management regimes, especially extensive grazing, can rapidly generate structural heterogeneity and sustain diverse aquatic invertebrate assemblages in eutrophic wetlands.

AimUnderstanding whether invasive species retain or shift their ecological niches has traditionally relied on scalar overlap metrics that quantify the magnitude of niche change, but not its structure. Here, we test whether biological invasions involve a reorganisation of the environmental axes along which native and invasive ranges are differentiated, and whether the dominant axes of this reorganisation are consistently associated with invasion pathway type (intercontinental vs. within-continent). LocationGlobal (North America, Europe, Africa, Asia, Australasia). Time periodContemporary (environmental variables representing long-term averages, 1980-2021). Major taxa studiedFreshwater crayfish (Decapoda: Astacidea): Procambarus clarkii, Faxonius limosus, Pacifastacus leniusculus, Faxonius virilis, Faxonius rusticus. MethodsWe analysed native and invasive occurrences for five globally important crayfish invaders using [~]400 hydrologically resolved environmental variables from the Global Crayfish Database of Geospatial Traits. Classification models were used to quantify environmental differentiation between native and invasive ranges, and feature contributions were aggregated by environmental domain (climate, topography, soil, land cover). Patterns were evaluated across intercontinental and within-continent invasion pathways and assessed for robustness using cross-validation, permutation tests, sample-size sensitivity, and comparisons with classical niche overlap metrics. ResultsNative and invasive occurrences were consistently distinguishable across all species (accuracy 96.5-99.9%). A pathway-dependent pattern emerged: intercontinental invaders were primarily differentiated along climatic dimensions (58-76% of model importance), whereas within-continent invaders showed a more balanced contribution of climatic and topographic variables ([~]42% each), including strong signals from river network position. This contrast was stable across cross-validation folds (SD < 1.6%), and supported by permutation tests (P = 0.001). Classical niche overlap metrics (Schoeners D = 0.30-0.62) did not capture this qualitative distinction. Main conclusionsBiological invasions involve not only changes in niche position but a reorganisation of the environmental axes that distinguish species distributions. Our results suggest that the dominant axes of this reorganisation differ systematically with invasion pathway, reflecting whether species encounter novel climatic regimes or primarily shift within existing climatic space along topographic and network-position gradients. By resolving which environmental dimensions underpin native-invasive differentiation, this approach provides a complementary perspective to scalar overlap metrics and a basis for more mechanistic interpretations of invasion processes.

Understanding habitat association of animals and how they change through ontogeny is critical to predict the likely effects of habitat change on populations. We investigated how fine scale habitat associations of three common coral reef damselfish species changed among life-stages on reefs surrounding Lizard Island, northern Great Barrier Reef. All three species showed distinct habitat selection at settlement, however the degree to which these initial associations changed through ontogeny were species specific. Pomacentrus amboinensis associated with sandy areas throughout all life-stages; Pomacentrus chrysurus settled to areas with high cover of sand and rubble, but displayed no clear habitat preferences as juveniles or adults. Pomacentrus moluccensis settled to areas with high cover of fine branching corals before shifting to areas with relatively high cover of soft corals as adults. We also compared two different approaches to estimate habitat selection; one that quantified the benthic composition within the approximate home range of individuals versus a more widely used approach of recording a single point underneath the focal individual when they were first observed. Although results were broadly similar, the benthic composition approach revealed details that was overlooked using the single point method. Decreases in the availability of any of these preferred benthic habitats may adversely affect future populations, therefore understanding habitat associations and their transitions among life stages will be crucial in predicting future reef fish communities under ongoing coral loss and habitat change. This will require to systematically study a broader range of species, integrating relevant spatial and temporal scales.

Interactions among co-circulating parasite species influence infection risk and disease progression. Such interactions can occur within hosts, for example altering susceptibility, or indirectly through host demography or movement, potentially affecting landscape-scale transmission. Despite their ubiquity, the spatial implications of these interactions have received limited attention. We combine spatially-explicit modelling with laboratory experiments to investigate how different parasite-parasite interactions influence disease spread. We model within-host, demographic, and dispersal-related interactions across a linear landscape, showing that within-host interactions modifying host susceptibility have the strongest effects on parasite prevalence, spatial heterogeneity, and rate of spread. Furthermore, these effects are amplified when parasites invade sequentially, generating pronounced patch-level spatial priority effects. We tested these predictions experimentally using a protist host (Paramecium caudatum) and two bacterial parasites (Holospora undulata and H. obtusa). Consistent with model predictions, we found that H. obtusa reduces prevalence and spatial spread of H. undulata through reductions in host susceptibility, and found evidence for spatial priority effects, observing reduced H. undulata prevalence when introduced after H. obtusa. Our theoretical and experimental results highlight that parasite-parasite interactions can have important implications for parasite spatial epidemiology, but the magnitude of those effects depend on the interaction type and the timing of invasion.

2The genotype-to-phenotype architecture (GPA), defined by complex interactions such as pleiotropy, epistasis, and regulatory control, is a fundamental yet often overlooked driver of biodiversity dynamics. While empirical evidence suggests that traits mediating species interactions (biotic) and environmental responses (abiotic) are frequently correlated, most eco-evolutionary theories treat these traits as independent, leaving a gap in our understanding of how genomic architecture influences community-level outcomes. In this study, we contrast two distinct GPAs, modular (independent trait evolution) and correlated (integrated trait evolution), within a spatially explicit multilayer network framework. We evaluate their impact on biodiversity across varying regimes of selection, migration, and biotic and environmental filtering. Our results reveal a hierarchy of drivers: selection strength dictates the absolute magnitude of the architectural effect, while migration and context-dependent biotic and abiotic effects determine which architecture yields a diversity advantage. Correlated GPAs enhance species coexistence and diversity in low-migration landscapes characterized by strong selection and moderate, balanced biotic and abiotic pressures. In these contexts, trait integration serves as a buffer against selective noise. Conversely, modular GPAs support higher diversity under high migration and strong biotic interactions, where the decoupling of trait modules provides the adaptive flexibility necessary to navigate spatially conflicting selective pressures. Our findings demonstrate that genomic architecture acts as a critical filter for environmental perturbations. Integrating complex GPAs into multispecies models is essential for quantifying the co-evolutionary feedbacks among traits, population adaptation, and species persistence. Our framework provides a path for predicting how biodiversity emerges and persists across biological scales, from genomics to communities and food webs, under the accelerating pressures of global change. 1 ConclusionsO_LIWe integrate trait architecture to spatial biodiversity to show biodiversity patterns are not merely products of ecological interactions, but are fundamentally constrained by Genotype-to-Phenotype Architecture (GPA). By linking GPA to biodiversity we show the interplay between the complexity of an organism and community structure in determining diversity patterns. C_LIO_LIThe hierarchy of Eco-Evolutionary Drivers: We establish a new conceptual hierarchy where selection strength acts as the fundamental governor of architectural impact, while the specific architecture predicting higher diversity (Correlational vs. Modular) is dictated by the interplay of migration scales and context-dependent biotic and abiotic dynamics. C_LIO_LISelection-Migration contingency for coexistence: We provide a new hypothesis for species coexistence: Correlational selection serves as a stabilizing force under dispersal limitation, whereas Modular trait architecture provides the adaptive flexibility to maintain diversity in high-migration, spatially heterogeneous landscapes. C_LIO_LIAdaptive decoupling as a diversity engine: We propose that trait modularity functions as a "buffer" against extinction by decoupling phenotypic responses. This allows populations to navigate conflicting selective pressures, effectively facilitating evolutionary rescue in complex biotic environments. C_LIO_LIMethodological framework for empirical inference: To bridge the gap between theory and data, we provide a novel likelihood-based framework. This enables researchers to infer latent trait architectures from population genomic samplings, turning GPA from a theoretical construct into a measurable sampling variable in natural populations. C_LIO_LIWe define a new roadmap for the next generation of eco-evolutionary modeling. By identifying the gaps between existing simulation engines, we provide a conceptual "blueprint" for a digital ecosystem that fully integrates complex genetic architecture with global bio-diversity dynamics. C_LI

Honeypot ants represent an example of convergent evolution, where a group of workers specialized in storing liquid food in their crops (i.e., stomach) has independently evolved multiple times across different ant genera. While seasonal resource scarcity and arid conditions are thought to drive the evolution of repletism, the role of environmental variables in this process has not been tested. With this is mind, species ensemble models were computed to assess suitability and richness areas, and the importance of predictors. Predictor importance was compared between genera and groups occupying a similar geographical area. Niche overlap and similarity between honeypot ant species were also evaluated to determine whether they occupy similar environmental spaces. Similarity was mainly found within genera, and Leptomyrmex and Myrmecocystus showed striking niche differences. Overall, Leptomyrmex distribution was mainly influenced by atmospheric bioclimatic variables like precipitation and temperature, while Myrmecocystus had soil bioclimatic variables as the most important predictors for their current distribution. Our results indicate that honeypot ants species currently do not occupy the same environmental space, and are not experiencing the same contemporary environmental stressors. While our results suggest that contemporary environmental factors cannot explain the convergence of honeypot ants, future research will examine past climatic conditions along with investigations into the ant genomes to understand more about the causes and consequences of the convergence.

Migration is widely recognized as a strategy for animals to track seasonally shifting resources. Yet, seasonal and spatial dynamics of migration are challenging to study, particularly for difficult-to-track insects. Among insects, monarch butterflies (Danaus plexippus) have a well-documented fall migration, but spring breeding recolonization remains poorly understood, particularly for the western population. We conducted multi-year surveys across six regions in the western United States to characterize monarch breeding phenology and evaluate three related hypotheses: (i) the successive broods model, with discrete generations shifting activity across the breeding range, (ii) a diffusion-like expansion model with overlapping breeding periods, and (iii) a mid-summer lull model with temporary summer declines in breeding for areas near the overwintering habitat. Monarch immature presence served as an indicator of local breeding activity. Our results do not support the successive broods or mid-summer lull hypotheses. Breeding onset occurred earlier near overwintering areas and gradually expanded north-and eastward, with sustained activity in many regions throughout the season. Termination of breeding also occurred earlier at more distant sites, resulting in longer breeding activity nearer to overwintering habitat. Immature monarch density declined with distance from overwintering areas at onset and termination, suggesting delayed colonization of peripheral regions. Together, these results support a diffusion-like expansion of breeding rather than sequential generational replacement. Western monarchs also do not initiate or terminate breeding in close synchrony with host plant availability, contrary to predictions from resource-tracking theory. These findings highlight fundamental differences between western monarch breeding dynamics and paradigms for eastern monarchs, demonstrating that a single species can employ fundamentally different spatial strategies for recolonizing its breeding range in different regions. More generally, these results distinguish insect migration from systems with direct movements between wintering and breeding habitats, and underscore the value of long-term, landscape-scale monitoring for resolving habitat use across heterogeneous environments.

Climate change and anthropogenic pressures are reshaping marine food webs, altering prey availability and affecting top predators. The European Shag (Gulosus aristotelis), a coastal demersal seabird, provides a valuable model for examining environmentally mediated dietary variation, given its trophic plasticity and capacity to adjust prey use according to local availability, while also allowing assessment of potential demographic consequences. This study investigated spatial and temporal variation in diet at two Portuguese colonies (Berlengas and Arrabida) between 2016 and 2024 and assessed long-term reproductive productivity at Berlengas. A total of 468 regurgitated pellets were analysed, and diet composition was quantified using the Index of Relative Importance (IRI). Generalised additive models were applied to assess environmental, spatial, and period-specific effects on diet composition, while reproductive productivity was modelled in relation to prey biomass. Diet variation was primarily explained by environmental predictors, including sea surface temperature, chlorophyll-a concentration, and zooplankton, whereas year per se had no significant effect, indicating environmentally mediated bottom-up effects. Spatial differences between colonies reflected contrasting prey field structures, and period-specific patterns suggested increased specialisation during breeding. Higher biomass of sandeels (Ammodytidae) was positively associated with reproductive output, whereas shifts toward lower-energy prey were associated with reduced productivity. These findings demonstrate that environmentally driven dietary change has measurable demographic consequences, underscoring the importance of bottom-up processes in shaping seabird population dynamics and informing conservation strategies under ongoing climate change.

Habitat complexity (HC) in part determines the diversity, stability, and behavior of food webs and can influence predation according to a wide variety of functional relationships. Many aquatic species provide habitat complexity and are also consumed by other species (e.g., macrophytes, corals, mussels). However, food web theory does not readily account for these species that act as edible habitat complexity (EHC). Here, we combine existing theory on predator-prey interactions, HC, and prey switching to describe the role of EHC in benthic food web models. We dissect feedback loops in each model to demonstrate how self-regulation of the prey species, mediated by species densities and HC, drives that food webs behavior. HC can stabilize predator-prey interactions by coupling prey self-regulation with HC self-regulation. EHC can further stabilize predator-prey interactions across a wide variety of "HC functions" that relate HC to predation rates. Significance StatementHabitat complexity (HC) plays a critical role in trophic interactions, population dynamics, and food web stability. However, little theory exists to describe edible habitat complexity (EHC), where a species is both consumed and confers habitat complexity for other species. We provide a series of models demonstrating how HC and EHC alter the population dynamics and stability of simple aquatic food webs. HC is strongly stabilizing in food webs by providing safety in rarity for prey. EHC provides safety in rarity for both prey and the EHC species because their predators are omnivorous. Given the prevalence of EHC species in aquatic systems (e.g., macrophytes, corals, mussels), our models demonstrate the importance of maintaining EHC species in aquatic systems for stable food webs.