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.

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. 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.

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.

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.

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.

The most studied symbiosis between land vertebrates and algae, is the one between Ambystoma maculatum and Oophila amblystomatis, where the latter colonizes the amphibian''s eggs. Although clear benefit given by algal growth for the salamander embryo has been observed, mutualism is not widely accepted. Mutualistic hypotheses posit benefits from the host to the algae which include protection, N- and C-sources, but no experimental evidence has been obtained. In fact, recent studies suggest that the algae are stressed during this relationship and propose fermentation rather than photosynthesis as main driver for algal growth. In this work, we analyzed embryonic host developmental rate, algal and embryo growth and photosynthetic function of eggs housed at different light intensities. We provide evidence that photosynthetic growth is the main driver for Oophila and that higher algal growth yields faster developmental rate and embryo size. Also, we provide strong evidence that rates of photosynthetic electron transport (and the concomitant O2 evolution) must be higher than those of consumption by the embryo for this symbiosis to be mutualistic. Since photosynthetic rate is dependent on light, this work illustrates the importance of light intensity in this symbiotic relationship.

Associations among species are hypothesized to become fewer and more specialized toward the tropics. However, evidence for the latitudinal specialization gradient (LSG) remains mixed and restricted to plant-animal and host-parasite models, with increasing recognition that our understanding of LSG is often complicated by differences in species richness and sampling effects. Here, we tested whether specialization between foliar fungal endophytes - fungi that live within plant leaves - and their plant hosts increased toward the tropics by surveying 1,539 plants that spanned a 55-degree latitudinal gradient in Central and North America. Contrary to expectations, endophytes and plants formed a greater number of associations that were more evenly distributed toward tropical latitudes and were thus less specialized, even after accounting for differences in endophyte and plant community structure. Endophyte-plant communities were most specialized where evaporative demand and plant richness were low, and where water availability and primary productivity were high. Our findings show a reversed LSG where the specialization between endophytes and plants decreased toward the tropics and provides evidence that many species of endophytes can coexist even when there is high overlap in their host niches.

Environmental variability influences the evolution of phenotypic plasticity and the ecological interactions among species in communities, but it is unclear how these processes interact to affect ecological systems across scales. We investigate models describing how plasticity evolves in competitive communities exposed to seasonal variation in abiotic conditions. We conducted eco-evolutionary analyses to identify communities of coexisting species with evolutionarily stable plasticity traits. These analyses revealed that the presence of interspecific competitors constrained the evolution of plasticity in some species, which instead evolved to be specialists. Plasticity often impaired species coexistence by allowing individual species to control more niche space. Communities containing plastic species were usually less species-rich but more productive and less variable. Similar results were obtained when plasticity rate was evolutionarily labile, but plasticity rate altered the patterning of temporal niche partitioning. This work shows how plasticity and competition combine to shape ecological and trait responses to environmental variability.

Heavy snowpack can be a severe constraint for alpine herbivores, yet adult survival in long-lived temperate ungulates is typically canalized against environmental variation. Using a 45-year individual-based dataset on bighorn sheep (Ovis canadensis) and historical snow reconstructions at Ram Mountain, Canada, we evaluated the direct and indirect effects of snow conditions on age- and sex-specific survival. For both sexes, survival of lambs and senescent individuals decreased with increasing average snow depth, whereas non-senescent adults (aged [&ge;] 1 year and < 8 years) showed high survival under mean conditions. However, this resilience broke down once specific thresholds of snow depth were crossed, with sharp survival declines beyond 90 cm of snow depth for females and 100 cm for males. Lamb survival, already reduced by increasing average snow depth, declined further once snow depth exceeded 90 cm. Crucially, the annual probability of snow depth exceeding 90 cm increased markedly over the study period, along with a density-independent temporal decline in survival across all demographic groups. Lamb survival declined by 0.37, and adult female survival, the most critical vital rate for population growth, dropped by 0.13 over the study period. Our findings challenge the established paradigm of demographic buffering in prime-aged adults, demonstrating that their resilience to environmental variability is fundamentally non-linear and breaks down once a changing climate regime breaches adult physiological boundaries, severely compromising population viability.

Understanding how species traits shape community properties is a central goal in ecology. But whether traits drive consistent patterns of dominance and assembly regardless of initial species abundances (evenness) remains unclear. We addressed this question using phytoplankton microcosm experiments to disentangle the effects of evenness and traits on community assembly and functioning, focusing on traits related to energy use (photosynthesis, size) alongside demographic parameters. Community assembly was highly deterministic, indicating that initial evenness had little influence on species dominance and final community structure. Instead, evenness drove community functioning. Demography was a stronger predictor of dominance than photosynthetic rates or individual size, suggesting that growth and competitive success depend on multiple metabolic processes that cannot be reduced to simpler proxies. These results show that community structure can emerge predictably across a range of initial conditions, and highlight the importance of demographic metrics over simpler metabolic descriptors for forecasting community outcomes.

Riverine and riparian food webs are connected via 'allochthonous' resource flows from neighboring ecosystems. These resources, such as terrestrial leaf litter, contribute to benthic macroinvertebrate diets, supporting higher trophic levels. While allochthony occurs in streams globally, its sensitivity to broad environmental gradients remains poorly understood. We explored the drivers of allochthony across six river basins within the contiguous United States, including climate, stream size, and flow permanence. We used Bayesian mixing models to estimate macroinvertebrate dietary proportions in four functional feeding groups. Air temperature best predicted allochthony, with consumers relying less on allochthonous resources with increasing temperature. Our models indicate that macroinvertebrate diets become primarily autochthonous at approximately 15{degrees}C mean annual air temperature, with leaf litter-consuming shredders showing a slightly higher threshold for this shift. We also found increased allochthony in smaller, non-perennial streams. These findings highlight the complexity of stream allochthony and responses to multi-scale environmental factors.

Metabolic rate governs the pace of life through the allocation and expenditure of energy, yet metabolic rate data are unavailable for most species. External morphological traits, such as gill slits, have been used to infer respiratory physiology and metabolism in sharks, yet, whether that pattern is generalizable across rays is unknown. In rays (Batoidea), spiracle length seems adaptive to oxygen uptake capacity, environmental oxygen availability, and activity level. Thus, we evaluate the allometry of external respiratory morphology (spiracles and gill slits) in rays using 550 museum specimens representing 88 species. We test whether spiracle length and gill slit height grow isometrically with body size (total length or disc width) within and across species. Overall, we find spiracles and gill slits are morphologically diverse, varying widely in shape within and between families. Spiracle length and gill slit height scale isometrically and are a consistent proportion of body size, despite such high morphological variability. While that proportion is consistent through ontogeny, each species proportion differs (e.g. gill slit height ranges from 0.93% of total length in Spotted Legskate to 2.93% in Pacific Electric Ray). Of the three swimming styles, pectoral oscillating species (Mobulid and Eagle Rays) may have steeper scaling of spiracle length with gill slit height - consistent with greater respiratory capacity in species with more active lifestyles. Our findings are consistent with literature hypothesizing external respiratory morphology (spiracles and gill slits) is tightly related to metabolic rate and activity level, and demonstrate the value of natural history collections in ecological research. Importantly, the isometry of both respiratory traits means that a single measure of any individual, irrespective of size, will be representative of the species thus providing a basis for expanding our understanding of metabolic morphology, enabling future work on life history, extinction risk, and ecological function.

Any mechanistic biodiversity-ecosystem functioning theory must explain how biodiversity in natural communities arises, which requires an evolutionary metacommunity perspective. Using a model of adaptive evolution of competitive metacommunities in landscapes with spatially heterogeneous supply of two resources, we studied the evolution of resource-uptake specialization as a function of resource type (substitutable/essential) and trade-off shape between resource-uptake affinities (generalist-/specialist-favouring). We identified "critical essentiality" as a key phenomenon separating parameter regions with fundamentally different biodiversity--ecosystem function relationships. Specifically, while greater spatial heterogeneity in resource supply increases biodiversity, this comes at the cost of reduced productivity when resources are essential. Moreover, when resources are essential, both generalist- and specialist-favoring trade-offs select for phenotypes that prevent efficient resource partitioning, leading to low resource use efficiency and consumer biomass - despite often high species richness. In contrast, when resources are substitutable, a specialist-favoring affinity trade-off selects for phenotypes that partition resources efficiently, promoting a metacommunity of only two species characterized by high functional diversity, resource use efficiency, and biomass. Overall, our analyses thus suggest that conditions promoting the evolution of diverse consumer communities often do not engender higher functional richness, consumer biomass, or resource use efficiency.

Foraging decisions often depend on net energy intake, but the need for specific micronutrients means that foods are not always comparable based on energy content. Although sodium is essential for many physiological functions, many organisms face challenges obtaining it, especially those relying on a plant-based diet. Flowering plants often lure pollinators by manipulating nectar carbohydrate content, but nectar sodium might also influence pollinator choice. We experimentally tested how sodium addition and nectar carbohydrate concentration interact to affect bumble bee foraging decisions by allowing bees to forage on artificial flowers containing either low- or high-sucrose nectar, with or without sodium. We also explored whether sugar and sodium content in floral nectar trade-off across plant species. Bees preferred sodium-enriched nectar and preference nearly doubled when sucrose concentration was low. Furthermore, bees switched more often from flowers without sodium to those with sodium when sucrose concentration was low. Given increased fidelity for artificial flowers with nectar sodium, sodium may enhance plant fitness by increasing conspecific pollen receipt. Yet despite behavioral evidence that sodium enhances the perceived quality of low-sucrose nectar for bees, we found no evidence for a nectar sugar-sodium trade-off across plant species. We discuss implications for plant-pollinator interactions and flower evolution.

1. Group foraging reduces the risks associated with foraging: larger groups dilute individual predation risk, allow group members to share the costs of vigilance, and provide a social structure for exchanging information about predators. The benefits of shared social information over simple safety in numbers should depend on the relationships among group members. Consequently, group composition should affect the selection of risky and safe habitats in social species. 2. We asked whether relationships among elk (Cervus canadensis) group members moving together influence their selection of risky habitats more than grouping alone. We tested three hypotheses. The familiarity hypothesis predicts that elk with close social ties are most likely to select risky habitats together. The inclusive fitness hypothesis predicts that the closest genetic relatives are most likely to select risky habitats together. The dilution hypothesis predicts that familiarity and genetic relationships do not influence habitat selection, with all group foraging benefits derived from proximity to others. 3. We used integrated Step Selection Analysis to test whether the identity of the closest group members influenced habitat selection. We tested whether the relationship of the nearest elk--either genetic or social--incentivized selection of risky habitat more than proximity to the nearest elk. 4. Group foraging clearly reduced risk, with elk more likely to select risky habitat when near others. Elk were also more likely to select risky habitats when near closer relatives, albeit with considerable variation among individuals. However, familiarity had a large and consistent effect: elk were far more likely to select risky habitats when near socially familiar group members. 5. Our results suggest that information shared within foraging groups, especially between socially familiar individuals, may lower the risks associated with habitat selection. Similarly, the benefits of reliable social information for reducing risk likely incentivizes the formation of stronger social ties within foraging groups. This positive feedback between social familiarity and habitat selection could thereby increase access to profitable habitats that are otherwise too risky, cementing what we term the social information landscape as a key driver of movement and habitat selection.