Ecological Monographs

Embedding thermal tolerance metrics into population dynamics offers a promising toolkit for understanding the impacts of stressful heat events, but the time-dependence of thermal stress rarely factors into such assessments. Within organisms, heat causes damage that alternatively accumulates under stressful temperatures and is repaired under permissive ones. While risk under exclusively stressful temperatures is well characterized by thermal death time models, resilience to fluctuating temperatures is less clear. Understanding how this damage accumulation within organisms scales up to affect population thermal tolerance is a necessary step for predicting population dynamics and extinction risk. We address this gap by embedding a model of organismal stress and recovery into population dynamics, yielding time-dependent thermal performance curves of population growth rates. By parameterizing this novel framework with existing data from Drosophila melanogaster, we explore how integrating thermal stress across biological scales shapes the consequences of organismal stress on population outcomes under realistic thermal fluctuations.

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

Ecologists have long investigated how community complexity affects ecological stability, yet how community complexity influences multistability, defined as the presence of alternative stable states, remains poorly understood. We developed a novel framework integrating stochastic community assembly with stability landscape analysis to quantify multistability from species interaction matrices. Using this framework, we systematically explored how species interaction properties shape the relationship between species diversity (species pool size) and the number of alternative stable states. Mean interaction strength was the primary determinant: competitive interactions amplified the positive relationship between species diversity and the number of alternative stable states. In competitive communities, a greater number of alternative stable states was associated with lower community uncertainty, a measure of the long-term unpredictability of community assembly dynamics. These results highlight the importance of characterizing the entire stability landscape. Our framework provides a general approach for understanding and quantifying multistability in complex ecological communities.

Backgrounds and Aims: Partitioning of biomass to various organs is fundamental to growth and survival strategies of plant species, with broad-scale ecological implications from competition to carbon flux of ecosystems. Although partitioning patterns have been studied extensively in embryophytes, little is known about the variation and evolutionary drivers of biomass partitioning strategies in marine plants, such as seaweeds, that form marine forests analogous to those on land. Methods: We measured developmental variation in standing organ biomass for 18 species of kelp and used a previously published dataset on land plants to draw comparisons between kelps and embryophytes, with respect to developmental allometry, interspecific allometry and the relationship between the two measures. Key results: We report that kelps exhibit large variation in partitioning patterns, with some species differing by more than an order of magnitude in holdfast and stipe investment when controlling for size. The range of biomass partitioning patterns exhibited by kelps includes values not found among embryophytes, highlighting differences in selective pressures on land and in the ocean. However, as previously demonstrated, empirical values of interspecific scaling curves closely match those of land plants. This suggests that despite fundamentally different physical constraints, evolutionary scaling has converged on nearly identical interspecific exponents. We further demonstrate that allometric covariation exists between support organs (stipe and holdfast), highlighting the importance of physical forces in driving functional diversification of marine plants and supporting the notion that covariation of traits is a hallmark of botanical structure and function. Conclusions: The results of our study illuminate the variation and evolutionary patterns of biomass partitioning in marine primary producers. Moreover, these results highlight that universal evolutionary scaling relationships can emerge from fundamentally divergent developmental and biomechanical pathways.

The performance of any natural population in variable environments depends on contemporaneous changes in its vital rates (e.g., survival, reproduction) as well as legacies carried by its population structure. Yet whether the relative contribution of these two pathways can be predicted from life history remains unknown. Here, we use stochastic simulations of 1,986 matrix population models from 137 species to quantify the contribution of transient dynamics to variation in population growth rate, and test its associations with key life history traits. Longer generation times were associated with reductions in transient contributions, contrary to theoretical expectations. Greater stage-specific survival heterogeneities were associated with increases in transient contributions, whereas greater iteroparity was associated with decreases in plants but increases in animals. These associations were robust to body size, phylogenetic relationships, and vital-rate variability. Life history traits therefore provide a strong predictor for when population structure shapes population responses to environmental variability.

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

Climate change is advancing spring phenology in temperate systems, with the potential to disrupt synchrony between trophic levels. Predicting these shifts requires understanding not only direct plastic responses to temperature, but also how plasticity at one stage carries over to shape timing at subsequent stages. We experimentally quantified direct thermal plasticity in phenology and its carry-over effects across the full life cycle of the winter moth (Operophtera brumata), a holometabolous insect whose fitness relies on synchrony with host plant budburst. Using a fully factorial split-clutch rearing experiment across four temperature treatments, we exposed individuals to contrasting conditions at each life stage and used structural equation modelling to partition direct and carry-over effects on phenology. Each life stage showed distinct plastic responses to temperature. Carry-over effects transmitted approximately 0.38 days/day of plastic advance to the next life stage on average, with the remainder absorbed by compensatory changes in the duration of the subsequent life stage. Together, these results show that carry-over effects propagate plastic responses across the life cycle, which are partially buffered by compensatory changes in developmental duration. Accurate predictions of phenological shifts under climate change therefore require models that account for carry-over effects and developmental compensation across life stages.

The consequences of maternal effects for offspring fitness can depend on the environmental conditions offspring encounter and the fitness traits considered. We used a transgenerational experiment in Daphnia magna to test how maternal exposure to warming and elevated salinity influenced offspring survival, reproduction and body size across matching, mismatching and multiple-stressor environments. Maternal exposure to salinity enhanced offspring survival under matching salinity, and benefits extended to mismatching environments. In contrast, maternal exposure to warming provided no survival benefit under matching warming and reduced survival in mismatching environments, while accelerating reproduction across contexts. Multiple-stressor conditions revealed pronounced maternal effects, while contrasting responses across survival, reproduction and body size showed that benefits to one fitness trait did not extend to others. Our findings demonstrate that environmental matching alone is insufficient to predict maternal effects, whose consequences depend on offspring environmental complexity and the fitness trait considered.

Urbanisation imposes novel environmental conditions that can shape phenotypic traits including plumage, yet, its effects on characteristics beyond colouration remain poorly understood. Feather microstructure plays an important role in thermoregulation, protection from solar radiation and waterproofing, and its expression is sensitive to environmental conditions. Here, we quantified feather barbule density, a key component of feather microstructure, in great tits (Parus major) from five urban-rural paired populations across Europe to assess its association with urbanisation, temperature and elevation. We found no significant effects of the urban environment or temperature. However, feather microstructure significantly varied between the studied areas and was positively associated with elevation and ultraviolet B (UV-B) radiation, suggesting a potential adaptive response to variation in solar radiation.

Dispersal differences between hosts and their symbionts can generate mismatched population structure, potentially destabilizing beneficial interactions across space. We tested this possibility in the carnivorous pitcher plant Darlingtonia californica and its obligate arthropod associates, the midge Metriocnemus edwardsi and the mite Sarraceniopus darlingtoniae, sampled across sites spanning the hosts patchy range in Oregon and northern California, USA. Comparing nuclear and chloroplast genomic data from D. californica with mitochondrial COI data from both arthropods, we tested how range position, landscape connectivity, and dispersal mode influence population genetic structure across this mutualistic metacommunity. Host plant populations supported the central-marginal hypothesis: nuclear diversity declined toward the range margins, and marginal populations showed greater nuclear genetic differentiation. Chloroplast variation was more weakly structured, most clearly separating the northern Oregon Coast populations and revealing cytonuclear discordance consistent with historical seed-mediated movement or chloroplast capture near the boundary between neighboring regions. Landscape connectivity estimated from an ecological niche model was also associated with genetic exchange. Circuit-theoretic current flow was positively related to effective migration inferred independently from plant genotypes. Further, landscape resistance explained variation in plant and mite differentiation beyond geographic distance alone. Both arthropods showed significant spatial congruence with the host plant but not with one another, a pattern inconsistent with co-dispersal and suggesting that each associate tracks the shared landscape according to its own dispersal biology. These results show that regional genetic concordance among obligate ecological partners can coexist with substantial differences in the processes governing their movement and local connectivity.

Identifying how species interact in ecological networks is key to understanding species' contributions to ecosystem functioning, but can be difficult to establish in species-rich communities. Functional niches, which capture broader ecological strategies, are often easier to assess and could help approximate species' interaction roles. We tested this by integrating species interaction and leaf functional trait data of tree species and leaf-associated interaction types from a large-scale forest biodiversity experiment. We found that identity, variability, and similarities of functional niches shaped corresponding aspects of interaction roles. Interestingly, however, this association inversed in the case of functional niche variability, which decreased instead of increased the variability of species interactions. This suggests a potential trade-off between processes that determine variability in functional and interaction space. Furthermore, effects were not constrained within aspects, with similarities of interactions being particularly responsive to most tested niche aspects, indicating that interaction-based competition among plants is particularly sensitive to functional niche differences. Moreover, many effects of functional niches depended on tree species biomass and tree species richness, highlighting the importance of the larger ecological context for determining species interaction roles. Together, our findings show that species' interaction roles, while clearly altered by, are not simply a consequence of functional niches. Instead, interaction roles integrate multiple aspects of functional niches depending on species performances and community composition and can therefore be crucial for anticipating ecosystem change.

Long-lived plants facing anthropogenic disturbance often exhibit recruitment failure despite persistent reproductive adults, generating extinction debt masked by longevity. However, whether adult presence reliably reflects environmental suitability for recruitment remains unclear. Here, we examine ontogenetic niche differentiation and its consequences for recruitment in Jubaea chilensis, an endangered long-lived Mediterranean palm with an aging population. We assigned individuals within the largest known population to four ontogenetic stages and characterized their environmental niches using climatic, edaphic, topographic, and vegetation variables. We then applied spatial and multivariate analyses, including Random Forest models to evaluate environmental segregation and identify predictors of seedling establishment. Age classes occupied significantly different environmental niches, with the greatest differentiation between seedlings and reproductive adults. Saplings and adult differentiation reflected mainly topographic variables at landscape scale, whereas seedling establishment was primarily predicted by microhabitat conditions (vegetation cover heterogeneity, east-facing slope orientation, and soil texture). This pattern is consistent with niche reconfiguring throughout the life cycle, suggesting that adult occurrence and recruitment suitability respond to distinct environmental conditions. Over one-fifth of sampled individuals occupied high-suitability sites without recruitment, suggesting that ontogenetic niche shifts are associated with a spatial decoupling between adult persistence and recruitment, consistent with demographic collapse independent of habitat degradation. This failure is likely mediated by insufficient effective seed dispersal, as the sole disperser (Octodon degus) preys on most seeds before dispersal. Conservation strategies based solely on adult distribution may therefore overestimate effective habitat and underestimate extinction risk in long-lived species.

Understanding when species diversification and phenotypic evolution are coupled remains a central challenge in macroevolution. Such coupling may depend on the traits considered, phenotypic integration, or the geographic context in which diversification occurs. Here, we investigate these relationships in Ithomiini, a diverse Neotropical butterfly tribe whose diversification is strongly associated with the Andes-Amazonia system. We quantified forewing and hindwing size and shape for 1,618 individuals representing 291 species using geometric morphometrics, and combined these data with phylogenetic comparative methods, diversification-rate estimates, and spatial analyses of species distributions. Evolutionary rate shifts were largely trait- and wing-specific, indicating limited coupling between forewing and hindwing evolution. Across the phylogeny, wing size evolution and extant wing size were not associated with species diversification, nor was hindwing shape evolution. By contrast, forewing shape evolutionary rates showed a positive association with diversification. Spatial analyses revealed that this coupling was geographically concentrated in the Andes, where high diversification rates coincided with elevated forewing size and shape diversity and increased forewing evolutionary rates. Hindwing diversity, by contrast, more closely tracked phylogenetic diversity and showed weaker or opposing relationships with diversification. Our results show that diversification and phenotypic evolution in Ithomiini are neither generally coupled nor fully independent, but linked in trait-specific and geographically structured ways. More broadly, they highlight the need to decompose complex phenotypes and integrate spatial context when testing macroevolutionary coupling.

1. Plant-soil feedback (PSF) is increasingly recognized as an important mechanism influencing plant coexistence, but most studies have examined PSF in isolation under controlled conditions. Under natural conditions, PSF operates together with other processes such as resource competition, yet how these processes jointly shape plant coexistence across environmental gradients remains poorly understood. 2. We combined field and greenhouse experiments to assess the contribution of PSF to plant coexistence at low- and high-elevation sites in the Swiss Alps. In the field, we quantified total niche and fitness differences and predicted interaction outcomes for nine plant species pairs, integrating PSF, resource competition, and other processes. In a complementary greenhouse experiment, we used soil microbial communities conditioned during the field experiment to isolate PSF-driven niche and fitness differences for the same species pairs. 3. PSF alone generated strong fitness differences but weak niche differences in the greenhouse experiment, predominantly predicting competitive exclusion. Interaction outcomes predicted from PSF alone corresponded poorly with those in the field, suggesting that PSF in isolation poorly explain plant coexistence under natural conditions. Nevertheless, PSF-driven niche and fitness differences were related to those quantified in the field, which were likely driven largely by resource competition, suggesting that PSF may contribute to coexistence by reinforcing or counteracting other ecological processes. These relationships varied between elevations, highlighting the context dependence of PSF contributions to coexistence. 4. Synthesis. Our study shows that the impacts of plant-soil feedback on plant coexistence cannot be understood from their effects in isolation. Instead, their contribution depends on how they interact with other ecological processes and on the environmental context in which these processes operate. Together, our findings emphasize the need to move beyond studying PSF in isolation towards quantifying how it combines with other mechanisms that drives coexistence under natural conditions.

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

Anthropogenic climate change is threatening global biodiversity, with sea turtles particularly vulnerable as offspring sex and developmental success are strongly influenced by incubation temperature. Behavioral plasticity, including the seasonal distribution of reproductive output, may provide short-term mechanisms for mitigating these impacts. Here, we investigated season-wide hatchling sex ratios and emergence success in a small population of green turtles (Chelonia mydas), tracking individual females across their nesting seasons. Sex ratios varied markedly through the nesting season, with later nests producing a greater proportion of male hatchlings. Moreover, sex ratios were relatively consistent among nests laid by individual females. Overall, females producing more nests over a season also produced more male offspring, suggesting that both nesting phenology and reproductive output influence individual contributions to future population demographics. Mechanistic models indicate that hatchling sex ratios have trended towards female-biased ratios (>80% female) over the past 50 years, and are projected to approach complete feminization by 2100 under continued warming. Although emergence success currently remains high (>85%), it is predicted to decline sharply after mid-century, with viable hatchling production falling to [~]30% by the end of the century. Models further show that maintaining contemporary sex ratios and emergence success will require unrealistically large delays in nesting phenology, and that even extreme shifts in phenology become ineffective by 2100. Together, these findings demonstrate that individual females can increase male hatchling production by nesting later and producing more nests, but behavioral plasticity alone is unlikely to offset the accelerating impacts of climate change on this population.

Quantifying how individuals adjust the seasonal timings of life-history events (i.e. their phenology) in response to seasonal environmental conditions is fundamental to predicting impacts of changing seasonality. However, the current standard approach of inferring phenological responses by directly analysing observed event dates relies on foundational assumptions that could be invalidated by changing seasonality, and excludes individuals that do not express focal events, potentially generating misleading inferences. To overcome these limitations, we devised a Phenological Threshold Model (PTM) that conceptualizes phenological events as switches between discrete life-history states and primarily estimates variation in underlying latent dynamic responses to seasonal progression, from which event dates secondarily arise. We demonstrate that the PTM can accurately retrieve simulated parameter values, including given incomplete seasonal field observations. Application to multi-year data on annual breeding in black-legged kittiwakes (Rissa tridactyla) demonstrated substantial among-individual variation in latent responses to seasonally varying environmental drivers, and associated daily probabilities that individuals will initiate breeding, revealing how annual distributions of breeding dates and proportions of non-breeders jointly arise. We thereby formalise a new conceptual and statistical framework for quantifying the potential for, and constraints on, rapid plastic and micro-evolutionary changes in phenology in changing seasonal climatic and environmental regimes.

Pollination and seed dispersal by animals are key drivers of terrestrial biodiversity and ecosystem functioning. Effective mutualistic interactions rely heavily on temporal and functional- trait matching between plants and animals. While global warming is known to induce shifts in plant traits, the extent and direction in which higher temperatures may systematically alter flower and fruit traits across species in natural habitats remains poorly understood. Using elevation as a proxy for temperature across natural populations, we conducted a meta-analysis evaluating 21 quantitative functional traits (15 floral, 6 fruit) across 82 studies and 161 species. Standardized mixed-effects linear regression models revealed widespread, systemic responses to elevational temperature gradients in both reproductive structures. In flowers, higher temperatures were systematically associated with changes in petal and sepal width and length (and hence morphology), longevity, nectar volume, number of flowers, and inflorescence length. In fruits, elevation was associated with changes in vitamin C content, crop size, weight and width. Taken together, these results demonstrate that warming temperatures exert widespread, multi-axis effects on the morphology, availability, timing, and nutritional quality of both flowers and fleshy fruits. Given that flower and fruit traits are developmentally linked and co-determine animal visitor dynamics, these temperature-driven phenotypic shifts are likely to propagate cascading disruptions throughout plant-pollinator and plant-frugivore interaction networks under continued climate change.

Climate influences community assembly by constraining the conditions under which species can persist, but the consistency of macroecological processes across different ecosystems remains unclear. Interspecific variation in climatic niche properties that govern community assembly across climate gradients can be gleaned from species distributions. Species-climate associations, niche properties, and resulting assembly remains poorly understood for plants in tropical dry ecosystems, and among different life-forms. We examined how species-climate associations governed community assembly of four plant life-forms across 60,000 km2 of tropical dry ecosystem in peninsular India. In the 1750 km long Eastern Ghats mountain range, we surveyed vegetation in 2500 20 x 50 m plots. Across the north-south climatic gradient from cool, wet and seasonal sites to warm, dry and less-seasonal sites, from presence-absence data of 1601 species in four life forms (trees, shrubs, herbs, climbers), we performed Bayesian Hierarchical Modelling of Species Communities to predict conditions of occurrence for each species. From this, we derived species climatic niche optima and niche width, and examined the local-level prevalence of niche properties across the climate gradient to glean assembly. Most tree, shrub and climber species associated with warm-dry conditions had wider niches than species associated with cool-wet conditions, which also resulted in assemblages having wider niches at warmer sites. Herbaceous species, by contrast, had narrower niches when associated with warmer conditions, where narrow-niche species dominated the assemblage. In all life-forms, however, assemblages in both warm-dry and cool-humid conditions consisted of species primarily associated with and preferring (having their optima in) those conditions, suggestive of stress dominance. Further evidence for stress dominance in shrubs and climbers came from assemblages in the warmest and coolest sites having low richness and comprising mainly of species with wider niches. Tree richness increased at either end of the gradient, while herb richness increased in warm-dry conditions. Overall, our results suggest that plant life-forms differ in the processes driving assembly across broad climate gradients in a tropical dry ecosystem, but many herbs were specialized to warm-dry climates. To understand life-form-specific responses to environmental gradients in tropical dry ecosystems, future work should incorporate traits and evolutionary history.

Biological invasions are a major driver of global change, reshaping ecosystems and threatening biodiversity worldwide. Anticipating where invaders will establish and where they will exert the strongest ecological impacts are key challenges for early detection and targeted management. Although Species Distribution Models (SDMs) are widely used to forecast biological invasions, they often provide uncertain estimates of establishment ranges and limited insight into invader performance, making it difficult to anticipate ecological impacts. Here, we address these limitations by integrating physiological information on invader performance with SDMs to identify regions of high invasion risk. Using the brown alga Rugulopteryx okamurae, one of the most prominent marine invaders in Europe, we first test alternative hypotheses of northern establishment limits: (i) a cold-survival constraint driven by winter temperatures and (ii) a growth constraint derived from the species' thermal performance. To identify the more likely scenario, we combine cold-tolerance experiments with seasonal growth comparisons between the invader and a native macroalga Dictyota dichotoma, whose established distribution allows physiological performance to be directly related to realised presence. Finally, we project seasonal growth of the invader across the predicted establishment range as a proxy for biomass accumulation and potential ecological impacts. Our results indicate that northern limit in Europe will be more likely constrained by winter survival rather than growth, extending the potential establishment range of Rugulopteryx to mid-Norway. In contrast, the highest impacts are likely to remain concentrated in southern Europe, where thermal conditions sustain high year-round growth. Overall, our approach illustrates how understanding the physiological response of invaders to their environment can improve the interpretation of SDM outputs and help identify areas at greatest risk of impact within their potential establishment range.