Evolutionary Ecology

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

Conspicuous coloration in animals is generally thought to evolve and be maintained through inter- or intraspecific interactions such as mate choice, but this might not always be the case. The sight-line hypothesis proposes that conspicuous light-dark contrast in front of the eyes (hereafter, eyeline) evolves and is maintained due to viability selection, enhancing an individual visual acuity and thus evolutionarily associated with a particular foraging behavior that requires accurate aiming. However, empirical evidence that supports the sight-line hypothesis is virtually absent, with no studies demonstrating the key prediction that the direction of eyelines matters. Here, I tested the sight-line hypothesis using macroevolutionary analyses in terns and allies, which are a suitable study system, because they have variation in facial color patterns, including presence/absence and, if any, various angles of eyelines. They also have a large variation in foraging behavior, including picking, plunge diving, and skimming. As predicted by the sight-line hypothesis, tern lineages that require accurate aiming at foraging (e.g., plunge diving) are more likely to have eyelines. In addition, the evolutionary transition to the state with eyelines and these foraging behaviors was more likely to occur than the reverse transition. Furthermore, as expected by the fact that the direction of travel is upwardly deviated from the direction of the bills during skimming, the eyeline angle from bills was evolutionarily positively associated with the occurrence of skimming behavior. To my knowledge, the current study is the first to demonstrate that the direction of the eyeline matters, thereby strongly supporting the sight-line hypothesis.

Sarita Lake, British Columbia houses a distinctive population of threespine stickleback (Gastrosteus aculeatus L.) with a phenotype characterized by unusually large individuals relative to nearby conspecifics. We tested the hypothesis that members of this population are not isometrically larger but rather exhibit variation in allometric trajectories that reflect changes in developmental timing impacting the developmental-genetic architecture of the phenotype. We used 3D geometric morphometrics to characterize the size and shape of skulls, pectoral girdles and pelvic girdles from a sample of individuals from nearby freshwater and marine populations and compare them to a sample from Sarita Lake. We showed that individuals from the Sarita Lake population are larger in each body region compared to most other populations examined. Further, these individuals have dorsally expanded skulls and relatively robust pelvic armour. We also showed that the relationship between size and shape is differently structured among body regions and is heavily influenced by non-uniform sexually-mediated variation across populations sampled. Our results reflect complex underlying developmental trajectories, and we suggest that the large phenotype observed may be driven by fecundity selection on female size in combination with a limnetic trophic niche and relatively increased predation pressure in Sarita Lake.

O_LISeasonal patterns of species appearances constitute a major component of diversity variation. Theory attributes this phenological structuring of communities to the alignment of life cycles to suitable moments and to constraints of seasonality on development, yet the specific mechanisms operating across taxa remain largely unresolved. In insects, body size and colour are key functional traits that contribute to driving spatial community assembly through their link to thermoregulatory performance and development. C_LIO_LIHere we analyse variation in mean body size and colour lightness of 483 butterfly assemblages across Great Britain and throughout the season to test whether trait alignment with seasonal environment and developmental constraints may shape the phenological structuring of communities. C_LIO_LIBoth body size and body colour varied more along season than across space, emphasizing the importance of phenology on diversity variation. Body size was larger early and late in the season, i.e. under conditions of low temperature and solar radiation. This pattern contrasted with the spatial trends found and was driven by species overwintering as adults, which we interpret as being likely due to energetic constraints. Body colour, conversely, was darker early and late in the season, mirroring the spatial pattern found, and suggesting a thermoregulatory alignment with seasonal conditions. Furthermore, covariation between body size and colour suggests a thermoregulatory interaction between both traits. C_LIO_LIOur findings suggest that life-cycle constraints and seasonal thermoregulatory alignment contribute to shaping the phenological structure of insect communities. C_LI

Predator-prey interactions are key drivers of behavioural and life-history evolution, yet their mechanisms remain difficult to study in natural contexts. The nematode Pristionchus pacificus is a model predator, but most studies exploring its behaviours use Caenorhabditis elegans as prey, a species that it likely only rarely encountered in nature. Here, we examine predation within nematode communities associated with beetle carcasses, the native necromenic habitat of P. pacificus. We identify Oscheius myriophilus as a cohabiting species, likely representing natural prey. Using predatory assays, automated tracking, and machine-learning-based behavioural analysis, we show that P. pacificus actively kills and consumes O. myriophilus. Strikingly, predation rates are lower than those observed for C. elegans, suggesting partial resistance or reciprocal adaptation in O. myriophilus. Consistent with this, O. myriophilus exhibits a mixed reproductive strategy, with early oviposition followed by ovoviviparity and matricide. As later developmental stages are more resistant to predation, internal hatching may protect offspring while providing maternal resources for development. These findings establish these nematodes as a tractable model for investigating predator-prey interactions and their evolutionary consequences, highlighting how behavioural strategies and life-history traits can co-evolve in natural communities.

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

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

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

According to sexual selection theory, males should benefit more from mating with multiple partners than females do, as male investment into offspring production is typically lower. For females, empirical evidence indeed often shows diminishing returns or even costs of mating multiply. For males, the assumption often seems to be "the more, the better" - i.e., a steady increase of male reproductive success with mate number - but experimental tests of it are rare. Here we used a laboratory experiment with Trinidadian guppies (Poecilia reticulata), known for being promiscuous, to assess how pairing males weekly with 4 vs. 7 females affects both sexes reproductive performance (n = 32 polygynous males and 170 monogamous females). Increased polygyny delayed females reproductive onset by 9% and tripled their risk of reproductive failure. High-polygyny males fathered offspring with 49% more females and had 73% higher daily reproductive output. Yet, they needed 19% longer to initiate pregnancy, and only accumulated more offspring than low-polygyny males after two months. This study suggests that male mating performance is not unlimited. Especially when high extrinsic mortality selects for fast reproduction, less polygyny might be advantageous, and the strength of sexual selection perhaps more similar between the sexes than often assumed.

In many systems, mutations can have background-dependent fitness effects due to genetic interactions between loci within a genome (intragenomic epistasis). In some cases, such as when species are coevolving, genetic interactions between loci can span across species; this is described as intergenomic epistasis. It is known that intragenomic epistasis can make adaptation more repeatable by constraining accessible mutational paths. Here, we investigate whether intergenomic epistasis leads to the same pattern of increased repeatability and how repeatability is influenced by the interplay of intra- and intergenomic epistasis. For this, we model a two-species system in which the fitness of a species depends on the combination of genotypes that are present in both species. We implement this system using an NKC model, which allows us to construct coevolutionary fitness landscapes on which we simulate adaptation by means of mutations in both species. To quantify the repeatability of adaptation, we track the realised endpoints of adaptive walks and record the distribution of fitnesses of the focal and partner species at these evolutionary endpoints. We find that intergenomic epistasis creates highly repeatable patterns of adaptation that depend on the underlying shape of the coevolutionary landscapes. The patterns of repeatability deviate from expectations based on intragenomic epistasis due to fitness trade-offs between species, which can lead to cycling and large co-evolutionary fitness loads.

Movement is costly, and animals are under strong selective pressure to move efficiently, yet, in patchy, dynamic landscapes, decision-making is inherently uncertain. We quantify the energetic savings achieved by using up-to-date information presented within social cues for reducing movement costs. We use an agent-based model, founded on realistic aeronautical rules and parametrised on the Andean condor (Vultur gryphus), to study movement in patchy landscapes. By explicitly considering altitude, flight results in a sequence of soaring and gliding in the 3D space. We investigate how the cost of movement to an overall goal varies when birds use social information from others that are either fixed in space or moving collectively to the common goal, and under different risk-taking speed strategies, from slow and cautious to fast and risky. The value of social information is operationalised as energetic savings in units of basal metabolic rate. Under low predictability, agents with intermediate risk and high social-information use exhibit lowest movement costs, with up to 41% energy savings over asocial movement. By extending classical aeronautical theory to social and variable environments we demonstrate the adaptive value of social information for efficient movement in patchy, unpredictable landscapes.

Resource competition and parasite exposure both present common density-dependent fitness costs for wild animals. Because launching effective immune responses is costly in terms of resources, parasites fitness costs should be further exacerbated in high-density, resource-depleted areas. To disentangle these relationships, we related density, parasitism, and resource availability to survival and fecundity across lifespan in a long-term study of wild red deer. All fitness measures declined with a combination of parasite count, greater density, and reduced resource availability. Beyond these relationships, as expected, local density and resource scarcity exacerbated survival costs of parasitism in calves, effectively undermining tolerance of infection. However, these synergistic relationships faded in yearlings and then reversed in adults, likely through age-structured selection biases. These findings emphasize that the costs of parasites and resource scarcity can be synergistic and intertwined with density in wild populations, accentuating the value of incorporating resource competition when examining parasite-dependent population regulation.

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

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

Alleles with opposing effects on fitness characters are said to exhibit selectional antagonistic pleiotropy (broadly construed so that effects are not necessarily confined to the same individual). A number of theoretical investigations considered the case where a pair of alleles at a locus influences two fitness components and derived the conditions giving rise to stable polymorphism under various assumptions about the mode of trait-interaction. Strikingly, many of these analyses concluded that the potential for maintaining polymorphism is strongly constrained by the joint influence of two factors: (1) the prevalence of weak selection coefficients over coefficients of large magnitude, and (2) the absence of beneficial dominance reversals (where the deleterious effects of each allele are partially or completely masked in the heterozygous genotype). Consequently, the conclusion that selective polymorphism is unlikely to be maintained by intralocus mechanisms of antagonistic pleiotropy has achieved widespread acceptance. Here we argue that such conclusions do not apply to any of the following models of antagonism: (i) additive trait-interaction, (ii) multiplicative trait-interaction, (iii) bivoltine selection, (iv) soft selection, (v) hard selection, and (vi) sexual antagonism. We demonstrate that the parameter space giving rise to stable allelic variation is quite large throughout, and moreover, the plenitude of suitable parameters neither depends on the strength of selection nor requires dominance reversal. Dominance coefficients associated with stringent conditions for stable polymorphism are shown to be atypical as compared to all feasible parameters, and best regarded as an outcome of adherence to a special relation: dominance with a constant magnitude and direction, which includes the case of additive allelic effects at a locus. Properties of single-locus equilibria (heterozygosity, allele frequency differentiation) are investigated, as well as the contribution of dominance schemes to the genetic variance in fitness characters in populations at multilocus linkage equilibrium. Author summaryAllelic variants at a locus with opposing effects on multiple fitness components (antagonistic fitness pleiotropy) have long been appreciated as a possible source of balancing selection. The prevalence of polymorphism owing to this form of natural selection, however, has been doubted on theoretical grounds due to the fact that standard assumptions of genetic models (namely, constant magnitudes for the dominance coefficients) are hardly conducive to the maintenance of polymorphism. The major exception to this conclusion lies with schemes that exhibit dominance reversal (where the direction of dominance for antagonistic alleles flips across fitness components). Here we conduct a geometric analysis of the space of polymorphism-promoting dominance parameters and conclude that the conditions for maintaining balanced alleles is unrestrictive, with non-reversals playing an underappreciated role.

O_LIPhenological shifts are a major ecological consequence of climate change, yet studies often focus on single life stages meaning that the potential for carryover effects between life stages remains poorly understood. Failing to account for these effects may lead to inaccurate estimates of phenological shifts, with consequences for predicted synchrony among interacting species. This is especially relevant for temperate systems where climate warming is occurring unevenly across the year. C_LIO_LIHere, we investigated how temperature experienced the previous autumn and winter (during the pupal and egg stage) influences spring phenology in the winter moth (Operophtera brumata), a herbivorous insect with distinct life stages. Using 50 years of local climate data to create five experimental temperature regimes, we first quantified phenotypic plasticity in the duration and temporal variability of pupal and egg development. We then examined how timing of adult moth emergence affects timing of offspring hatching. C_LIO_LIWe found divergent effects of temperature on different life stages; pupal development time was shortest at intermediate temperatures while egg development time decreased linearly with increasing temperature. Furthermore, phenological shifts due to the conditions experienced by the mother were carried over to influence the phenology of her offspring. While this carryover effect was partially compensated during subsequent stages, compensation decreased under warming conditions. C_LIO_LIThese results refine our understanding of the sensitivity of the annual cycle of winter moth phenology to variation in temperature with potential implications for population dynamics and interspecific interactions. Overall, our findings highlight the need to consider the impacts of warming across multiple life stages so that carryover effects can be properly accounted for. Doing so will improve predictions of phenological shifts under future climates. C_LI

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

Judgment bias tasks are increasingly used to assess affective states in animals, yet the extent to which they might reflect transient states or stable traits remains unclear. Here, we tested bumblebees (Bombus terrestris) in an active choice task across three repeated sessions to assess individual consistency in the absence of any manipulation. Bees were trained to associate each of two colours with either a high or a low reward, presented in separate chambers. During testing, they were presented with ambiguous colours. Bees were more likely to choose the high reward chamber and to choose more quickly in response to colours closer to the positive colour. The latency to choose the cues showed significant and moderate repeatability across sessions, suggesting a stable, trait-like underlying component. In contrast, the repeatability of the chamber choices was negligible, indicating that such responses might be largely state-dependent and influenced by situational factors. These findings suggest that judgment biases, particularly as assessed through an active choice task reflect states affected by external factors. Active choice tasks may help disentangle stable behavioural traits from transient affective states in invertebrates.

Protogynous sex change, where individuals first function as females and later as males, is a key life-history strategy among polygynous reef fishes. In haremic systems, sex change is typically socially regulated, with dominants suppressing subordinates sex change through aggression. Females within a harem form a size-based hierarchy that can remain stable in most species through the threat of eviction. We studied a different situation in the cleaner wrasse Labroides dimidiatus, where larger females have incomplete control, as they spend most of their time alone at their own cleaning territory. We tracked over 400 individuals for 12 months, recording growth, behavior, social organization, and sex change. We confirmed earlier reports that both sexes direct aggression primarily at those ranked immediately below them. However, we observed 30 cases where smaller females outgrew larger ones, revealing hierarchy instability. Of 42 sex change events, 43% occurred in presence of the male, and half of these early sex changers were not the largest female, but individuals overlooked by the male. Fast growth relative to harem-mates and harem switching increased the likelihood of sex change. Local population densities also influenced growth and sex change, with individuals in high-density demes growing faster and changing sex at larger sizes. Our findings reveal flexible sex change dynamics in a system with incomplete social dominance. Such incomplete control and observations that becoming male confers both higher reproductive success and survival highlight the need to expand game-theoretical and life-history frameworks to encompass such strategic flexibility. Lay summaryDominant cleaner wrasse cannot fully control subordinates as individuals occupy distinct core areas. Tracking 400 fish for a year, we found that smaller females could outgrow initially larger ones, and early sex change despite a larger male. Fast growth and harem switching increased the chances of becoming male. Population density also shaped these strategies. Our findings reveal flexible sex change dynamics in a system where becoming male confers both higher reproductive success and survival.

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