Functional Ecology

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.

Nutrition plays a fundamental role in insect physiology, yet whether growth, immunity and stress responses share a common nutritional optimum remains poorly understood. Using the Geometric Framework for Nutrition, we investigated how dietary protein and carbohydrate balance regulate multiple physiological functions in the specialist herbivore Plutella xylostella. Growth was jointly determined by protein and carbohydrate intake, with maximal biomass accumulation constrained primarily by protein availability. Immune traits showed distinct nutritional optima: total hemocyte counts and phenoloxidase activity were maximised at intermediate protein-carbohydrate (P:C) ratios, whereas survival following Bacillus thuringiensis challenge depended predominantly on protein intake. Antioxidant and detoxification responses likewise varied non-linearly across the nutrient landscape, with individual enzyme systems each exhibiting distinct nutrient-dependent optima. Importantly, no single macronutrient balance simultaneously maximised growth, immune performance and oxidative homeostasis, demonstrating that these functions have divergent nutritional requirements. Together, these findings show that macronutrient balance generates competing physiological demands, offering an integrated view of nutrient-dependent regulation of growth, immunity and stress physiology in a specialist herbivore.

Diet can influence lifespan and age-dependent physiological function, but how nutrition during development and adulthood affects relationships among stress resistance traits across age remains unclear. Using an isocaloric full-factorial larval x adult diet design in outbred Drosophila melanogaster, we tested how protein- and carbohydrate-biased diets affected starvation, desiccation, cold, osmotic, and heat resistance across the adult life course and sex. We also measured whole-body triacylglyceride, protein, body weight, and water content, and compared responses with matched lifespan and reproductive-output data. Resistance declined across the adult life course, but the traits did not vary as a single phenotype: among the traits included in the covariance analysis, desiccation, osmotic, and heat resistance formed a tightly correlated core, whereas starvation resistance remained comparatively peripheral in both sexes. Developmental diet produced trait-specific effects, whereas protein-biased adult diet favoured heat and osmotic resistance, paralleling effects on lifespan and reproductive output, but had opposite effects on starvation and cold resistance in specific contexts. Thus, nutrition alters rather than uniformly improves stress resistance, and covariance among resistance traits does not necessarily predict how those traits respond to nutritional perturbation.

Ecosystem functions can be regulated by biodiversity ranging from broad to narrow scales, including variation within a species. For most multicellular life, intraspecific variation is determined not only by an organism's phenotype and genotype, but also variation imparted by their microbiome. Here, we investigate how leaf decomposition in rivers is affected by phenotypic variation in plant secondary metabolites (PSMs) and antibiotic-induced variation within the gut microbiomes of aquatic macroinvertebrate decomposers. We found that consumption by Dicosmoecus caddisflies was inhibited by the presence of dietary PSMs from riparian Alnus rubra trees, irrespective of the state of their gut microbiome. We further compared consumption of diets prepared from 20 different local and non-local A. rubra trees. We found that antibiotic-treated and control caddisflies consumed similar amounts of diets containing local A. rubra PSMs. However, compared to caddisflies with an intact gut microbiome, antibiotic-treated caddisflies consumed less of non-local A. rubra diets. Thus, decomposition by caddisflies could be regulated by both host adaptation to local resources and bacterial-mediated digestion of non-local resources. Overall, our study suggests that gut microbiomes can facilitate adjustment of macroinvertebrate decomposers to novel plant phenotypes, which may arise from shifting spatial distributions of plants in response to global change.

Animals reproduce across a spectrum from oviparity to viviparity. Internal gestation and live birth have inherent advantages despite their costs. However, comparative analyses of key drivers, such as resilience to dehydration, lack taxonomic breadth. In this study, we assessed whether live birth improves the ability to proliferate in environments with limited access to water. To do so, we used the viviparous cockroach, Diploptera punctata, as a model of viviparity to assess dehydration-induced damage during extended periods of water deprivation. During pregnancy, maternal survival did not decline during dry periods compared with non-pregnant individuals, suggesting that pregnancy has minimal impact on resistance to dehydration stress. When mothers are deprived of water for 2 weeks, they show an increase in osmolality after one week, while their embryos show no change in osmolality until after two weeks, at which point abortions occur. Periods of dehydration increased the duration of pregnancy, likely due to a slower production of in utero milk, but there was no change in the size of progeny. After birth, viviparous D. punctata newborns showed increased survival under dry conditions compared to two other ovoviviparous species, which are much smaller and dehydrate more quickly. The combined impact of increased dehydration resistance during pregnancy and during the first independent phase indicates a benefit of prolonged viviparity in Diploptera punctata. This study provides evidence that viviparity in animal systems could enable progeny survival during drought.

The biogeochemical niche hypothesis (BNH) proposes the multi-elemental composition of organisms - their elementome - as a new ecological dimension. However, which ecological factors shape elementome assembly remains little known, especially in animals. Here, we studied the mandibular elementome of two sympatric small mammals - Apodemus flavicollis and Clethrionomys glareolus - to assess how intraspecific variability (ontogenetic changes in body mass and sex under the vertebrate bone hypothesis; VBH) and environmental filtering (season and habitat) shape essential and non-essential elementome assembly. Species showed moderate elementome segregation and seasonal niche partitioning, with implications for coexistence. Ontogenetic body mass predicted elemental variation and calcium substitution, with several hypermetric scalings in autumn indicating strong departures from mass-invariant homeostasis. Finally, our results suggest a dichotomy: essential elementomes were mainly driven by intraspecific variation, whereas non-essential elementomes were rather shaped by environmental filtering. Our results position animal elementomes as an integrative ecological dimension linking organismal biology, species interactions, and environmental filtering across individuals, populations, and species.

Litter invertebrates mediate a substantial proportion of decomposition and nutrient cycling in terrestrial ecosystems, yet how moisture availability structures their communities remains poorly understood. We propose that moisture availability could shape litter faunal communities through two alternate pathways: (1) a productivity effect: greater productivity and detrital inputs in wetter forests should increase litter-faunal abundance, (2) a trait effect: phylogenetic differences in water balance should filter communities according to taxon-specific moisture tolerances. We tested these expectations across 100 forest sites spanning a moisture gradient in south-eastern Australia, from dry sclerophyll forest to cool temperate rainforest. Measuring 11 focal invertebrate taxa, total litter-faunal abundance increased threefold across the moisture gradient, from 400,000 individuals per hectare of dry forest to 1.2 million individuals per hectare of rainforest - consistent with greater resource availability in wetter and more productive habitats. However, this overall abundance trend masked pronounced variation in community composition across habitat types. While moisture-sensitive crustaceans and myriapods (particularly Isopoda and Chilopoda) were concentrated in rainforest litter, two groups of hexapods (Blattodea and Embioptera) were most abundant in dry forest litter and declined sharply toward rainforest habitat. These contrasting responses indicate that dry forests do not simply represent depauperate versions of wet-forest communities but harbour their own distinct and abundant litter invertebrate assemblages. Thus, moisture structures litter faunal communities via both proposed mechanisms: (1) a productivity effect and (2) a trait filtering effect. Given that moisture fundamentally structures litter faunal communities, shifts in forest moisture regimes under climate change may substantially alter detritivore communities, with consequences for decomposition processes and nutrient cycling globally.

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

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.

In many ecosystems, anthropogenic warming is reshaping thermal regimes, leading to resource quality declines and imposing a dual constraint for ectotherms: elevated metabolic demand coupled with reduced assimilable energy. We tested whether plasticity in gut morphology and gut microbiome can buffer amphibian larvae against these concurrent stressors. Common frog (Rana temporaria) tadpoles were reared at two temperatures (18 vs. 24.5{degrees}C) crossed with three food-quality treatments (low, medium, high). We quantified growth and developmental rates, critical thermal limits (CTmax, CTmin), gut morphology (mass, relative length), and gut bacterial diversity and composition, together with predicted functional pathways. Warming accelerated growth and development and increased CTmax. Food quality increased growth and development, with temperature-dependent effects on developmental rate and CTmax. Gut mass declined at higher temperature and low-quality diets, but relative gut length showed only modest diet effects and no temperature dependence. Bacterial community composition and structure shifted with temperature and food quality. Predicted pathways suggest functional reconfiguration under warming and low food quality, consistent with sustaining energy acquisition and mitigating metabolic and oxidative stress. Together, these results implicate microbiome plasticity, rather than gut morphological plasticity, as a candidate mechanism supporting larval performance and heat-tolerance acclimation under warming and low food quality.

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.

Global biodiversity loss is restructuring ecological communities and compromising ecosystem functioning. Yet, we still lack an empirical understanding of how specific trait combinations regulate ecosystem processes or how their functional expression is organised within the immediate environment. Here, we investigated trait-expression across 17 functionally contrasting sediment-dwelling marine invertebrate species. We show that while sediment reworking and burrow ventilation are depth dependent, their influence on nutrient concentrations are highly nutrient-specific. Our analyses reveal that nutrient concentrations are better explained by continuous, spatially explicit trait combinations than by categorical bioturbation modes. Surface modification was associated with higher ammonium concentrations, moderated by ventilation, while deeper sediment reworking and biomass were associated with higher nitrite concentrations. Phosphate relationships were weak, and nitrate was not explained by the measured traits. These results establish a mechanistic baseline for species-specific trait-nutrient relationships under controlled conditions. This shift from categorical trait assignments to a spatially explicit, empirically resolved trait-interaction framework provides a mechanistic foundation for predicting how changes in species composition may alter benthic nutrient cycling.

Nectar is a primary floral reward, which can also deter non-specialist pollinators; yet, the role of complex nectar composition in plant-pollinator interactions and its relationship to other floral traits remain understudied, particularly in natural communities. We profiled the nectar metabolome of 43 co-occurring species in an alpine grassland to evaluate relationships among nectar chemistry, phylogeny, floral display traits and pollinator visitation. Field observations revealed a dominant bee vs fly visitation gradient. Several aggregated nectar traits tracked this gradient independently of plant phylogeny, indicating functional convergence: specifically, bee visitation was associated with higher nectar sucrose proportion, greater compound richness, and more divergent profiles of uncommon sugars. Nectar traits exhibited variable relationships to floral display traits, suggesting partial inter-trait phenotypic integration. Together, these findings indicate that detailed features of nectar composition, including phytochemical diversity, play a functional role in floral phenotypes, with previously unappreciated implications for plant-pollinator community ecology.

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.

Resting metabolic rate, the basal energy required for an organism's maintenance, is central to predicting the eco-evolutionary consequences of environmental and anthropogenic pressures on marine ectotherms. Yet, direct measurements remain extremely scarce, covering less than 1% of marine fish species. Building on the Pace Of Life Syndrome (POLS) hypothesis, which posits a positive correlation between metabolism and life-history speed, we inferred temperature- and mass-specific resting metabolic rate for 18,214 fish species (16,998 Teleostei and 1,216 Elasmobranchii). Estimates were derived from relationships with 15 functional traits and phylogeny using phylogenetic structural equation modelling, a novel approach combining phylogenetic comparative methods with structural equation models. Cross-validations indicated high predictive performance, although some clades require cautious interpretation due to limited empirical data. In line with expectations from life-history theory, ecological niche and morphology, species with the highest metabolic rates were small, narrow, shallow-bodied, and long-jawed pelagic fish with high mortality, short lifespan and rapid growth - traits typical of small epipelagics. The metabolic rate variation aligned with the slow-fast life-history continuum, supporting the POLS hypothesis, but was equally explained by reproductive strategy: highly fecund, fast species exhibited the highest metabolic rate independent of mass and temperature, whereas low-fecundity slow species had the lowest. By filling a critical physiological data gap, our dataset provides a foundation to test long-standing ecological and evolutionary hypotheses such as POLS, and offers a powerful resource to improve models of species and community responses to climate change and exploitation, ultimately supporting physiology-informed ecosystem management.

Precocious reproduction, prior to the acquisition of the final adult morphology, occurs in a variety of animals and represents a distinct life-history strategy with important consequences for growth and survival. Larval reproduction is far rarer and usually restricted to groups that go through metamorphosis. In ctenophores, Carl Chun described dissogeny more than 130 years ago, a distinctive life-history pattern involving larval and adult reproductive periods separated by gonadal regression. Yet, the potential plasticity of larval reproduction in ctenophores remains unresolved. Here, we developed a method to follow and quantify reproductive output in Mnemiopsis leidyi larvae and tested environmental factors that may regulate precocious reproduction. We found that conspecific density strongly influences its onset: low-density, well-fed animals can bypass reproduction at the cydippid stage and develop rapidly into the lobed form, whereas higher density promotes precocious reproduction. Food availability and associated growth rate also contribute but cannot alone explain the observed differences. Once reproduction is established, however, high conspecific density suppresses reproductive output, an effect reproduced by exposure to seawater incubated with high M. leidyi densities, consistent with water-borne chemical communication. Salinity and light conditions further modulate spawning. Together, our results reveal substantial environmental plasticity in precocious reproduction in M. leidyi, reconcile long-standing contradictory observations of ctenophore development, and highlight how environmental cues can reshape reproductive life-history transitions.

Heat-induced sterility is widespread across ectotherm species, yet its underlying mechanisms remain poorly understood. In mated females, thermal stress can disrupt female reproductive physiology while also affecting sperm stored within the reproductive tract, making the mechanisms of fertility loss difficult to distinguish. Here, we investigated the extent and recovery dynamics of female heat-induced fertility loss across seven Drosophila species. We tracked fertility after acute heat exposure, tested whether remating restored fertility, quantified stored sperm in two closely related species, and examined whether reproductive thermal sensitivity covaried with adult heat tolerance. We found that heat exposure reduced egg-hatching success in all seven species, but recovery differed among species. Fertility recovered within several days in most species, whereas D. simulans showed persistent declines and D. melanogaster showed limited recovery. Remating restored or maintained fertility in both species. Stored sperm declined through time in both D. simulans and D. mauritiana, but D. mauritiana retained more sperm at later sampling points, consistent with their contrasting patterns of fertility decline and maintenance. Across species, greater adult heat tolerance was associated with smaller immediate reductions in fertility, but not with cumulative reproductive loss across subsequent days. Together, these findings show that similar initial fertility losses can lead to markedly different recovery trajectories and highlight the importance of comparative, multi-species approaches that integrate female physiology, sperm-storage dynamics, mating behavior, and whole-organism thermal tolerance when assessing reproductive resilience to climate warming.

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.

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.

In natural ecosystems, community dynamics are driven by diverse interactions operating through both ecological and reproductive processes. Although theory predicts that these processes jointly generate complex and contingent dynamics through the interplay of stabilizing (negative frequency dependence arising from niche differences) and destabilizing effects (positive frequency dependence caused by reproductive interference), empirical tests of this prediction remain scarce. Here, we combined experimental and theoretical approaches to investigate the dynamics of two congeneric annual plant species (Commelina) experiencing both ecological competition and reproductive interference. Our empirically parameterized model revealed a large niche difference and mutual reproductive interference. However, the reproductive interference was not strong enough to generate pronounced contingency of community dynamics on initial conditions. Instead, coexistence was the most likely outcome regardless of initial species composition. Parameter exploration indicated that the weak contingency arose because reproductive assurance strategies of Commelina species buffered the minority disadvantage caused by reproductive interference. These findings provide empirical evidence that reproductive interference does not necessarily destabilize coexistence and that it may be outweighed by ecological niche and fitness differences. Our study highlights the importance of integrating ecological and reproductive processes towards better prediction of community dynamics.