Evolutionary Ecology

The connection between female host plant preference and offspring performance is important for understanding how relationships between plants and phytophagous insects have evolved. According to the preference-performance hypothesis, female insects should evolve to oviposit on host plants on which offspring performance is the highest. Here, we examined the preference-performance hypothesis in the northern brown argus (Aricia artaxerxes) butterfly in the province of Uppland, Sweden, by comparing female host plant preference and larval growth between the host plant species wood cranesbill (Geranium sylvaticum) and bloody cranesbill (G. sanguineum). We also investigated if host plant preference in A. artaxerxes was related to the geographic distribution of A. artaxerxes and its host plants in the province Uppland. We found that the A. artaxerxes females, contrary to the preference-performance hypothesis, preferred ovipositing on G. sylvaticum, even though larvae feeding on G. sylvaticum were slightly smaller than those feeding on G. sanguineum. Since G. sylvaticum is more abundant and probably more utilized than G. sanguineum in Uppland, an explanation for this negative preference-performance connection may be that there are advantages associated with utilizing a more common host plant species, even though larvae feeding on this plant show reduced growth rates. Overall, the results show that factors other than offspring performance, such as geographic distribution, may influence female host plant preference in A. artaxerxes.

In many tropical areas, seasonal rainfall leads to distinct dry and wet seasons. Many butterflies developing under wet season conditions develop into adults with large ventral eyespots on the wing margins, whereas those developing under dry season conditions have smaller or no eyespots. In greener, wet season habitats, larger eyespots can divert predator attacks toward the wing margins, while reduced eyespot size improves camouflage in the dry leaf litter-dominated habitat during the dry season. However, the dry season is also characterised by higher desiccation stress than the wet season. We hypothesised that larvae developing under dry season conditions develop into adults with higher desiccation tolerance than those reared under wet season conditions. We tested this by rearing larvae of the butterfly Mycalesis mineus under simulated dry and wet season conditions and assaying the desiccation tolerance of the resulting adults. Butterflies reared in dry conditions survived longer under desiccation stress, lost lesser water during pupal-adult metamorphosis, and were heavier than those reared in wet conditions. We also tested the correlation between eyespot size and desiccation tolerance. A negative correlation between the traits would be expected if similar developmental pathways regulate them. Consistent with this expectation, individuals with smaller eyespots had higher desiccation tolerance. Our results demonstrate plasticity in desiccation tolerance, and suggest that predator avoidance and desiccation tolerance traits may share similar developmental pathways.

Behaviour is a key yet often overlooked component of animal camouflage and how it evolves alongside colour and morphology remains poorly understood. The repeated evolution of stick-like postures in spiders offers a useful framework for investigating the importance of behaviour for concealment, since matching the environment should rely on specific body forms and postures, not just colouration. We hypothesised that when spiders behaviourally align their body with the background orientation it should influence the shape, posture and colouration that best enhances camouflage. To test this, we used a genetic algorithm and human observers to evolve digital spiders to be harder to find. We evaluated how selection under three behavioural orientation treatments (aligned, random, and evolvable orientation) influenced spider capture time, background match (lightness and colour), posture, and body (cephalothorax and abdomen) dimensions. We found that spiders that behaviourally aligned with the background took substantially longer to find through evolving a better background match, and a more elongated posture and body shape than randomly orientated spiders. Our spiders mirrored the shape and posture adopted by numerous clades, illustrating how behavioural camouflage represents a key concealment strategy in structurally complex habitats, part of an interacting suite of traits that encompass successful concealment.

Biotic interactions can drive evolutionary diversification, but the underlying mechanisms differ depending on the type of interaction. For instance, Ehrlich and Ravens escape-and-radiate coevolution provides a pathway of diversification in antagonistic interactions, whereas in mutualistic networks, coevolution is hypothesized to result in trait convergence rather than diversification. The combined effect of mutualism and antagonism on diversification remains unclear, even though organisms naturally engage in multiple types of interactions simultaneously. Using an eco-evolutionary simulation model, we investigate diversification in tripartite ecological networks such as plant-pollinator-herbivore networks. We find that diversification patterns vary according to the way mutualism and antagonism are connected on the trait level. If the two interactions are governed by uncorrelated plant traits, we observe little diversification in the mutualistic and substantial diversification in the antagonistic subnetwork. By contrast, if the same plant trait mediates both mutualism and antagonism (an example of ecological pleiotropy), diversification rates in all guilds become interdependent. In this case, even the mutualistic guild diversifies considerably when antagonism is strong, while strong mutualism restricts diversification also in the antagonistic guild. Our study underlines that the inclusion of multiple interaction types is necessary to advance our understanding of evolutionary dynamics in ecological networks.

O_LITerrestrial insects are vulnerable to desiccation due to their small body size. Because insects lose most water through cuticular evaporation, cuticular traits strongly influence desiccation tolerance. Individuals with greater cuticular melanisation, i.e., darker ones, are hypothesised to tolerate desiccation better than less melanised ones. C_LIO_LIIn many butterflies, pupal colour is plastic - individuals pupating on leaves tend to be greener, while those that pupate away from leaves (off-leaf), such as on tree bark or defoliated twigs, tend to be browner. Brown pupae are hypothesised to have more cuticular melanin and are expected to experience higher desiccation stress than leaf-borne green pupae. Thus, plasticity in pupal melanisation may be an adaptation against desiccation. We tested this in the butterfly Eurema blanda. C_LIO_LIWe demonstrate that individuals pupating on on-leaf substrates are greener than those pupating on off-leaf substrates, and that desiccation stress is higher in the off-leaf substrates, a microenvironment typical of brown pupae, than in typical green pupae. Using Raman spectroscopy, we show that brown, but not green, pupal cuticles contain melanin. C_LIO_LIFollowing this, we obtained greener and browner pupae by manipulating substrate colour. When subjected to desiccation stress, browner pupae survived better than greener ones. There was no correlation between pupal colour and survival in the absence of desiccation stress. Thus, melanisation appears to confer a survival advantage to pupae by increasing desiccation tolerance. C_LIO_LISurvival under desiccating conditions was inversely related to water loss. Interestingly, melanisation did not correlate with water loss, suggesting that melanisation helps tolerate desiccation through physiological mechanisms not directly related to water loss reduction. C_LIO_LIOur findings reveal an additional, crucial, adaptive value of pupal colour plasticity, a trait that has been studied primarily from an anti-predatory perspective. C_LI

Reproductive Character Displacement (RCD) often occurs when species with mating-related polymorphism come into secondary contact, leading to divergence in reproductive traits. Ischnura elegans and Ischnura graellsii have formed two independent hybrid zones in Spain where reinforcement has strengthened a mechanical barrier, and RCD has shaped mating-related structures, although reinforcement is asymmetric only in gynochrome females. This study examines the link between asymmetric reinforcement and asymmetric RCD. Using geometric morphometrics, we analyze prothorax shape and size in both female morphs and males, and male caudal appendages, to assess morphological divergence, determine whether gynochrome females show stronger divergence, and test for morphological covariation between male traits involved in the tandem position. Our results reveal consistent patterns of size and shape variation across species and zones: in I. elegans, androchromes are larger and resemble males in size, with clear shape differentiation between female morphs that diminishes in hybrid zones. In contrast, I. graellsii shows less consistent size differences between males and morphs, and weaker shape differentiation. Our results confirm RCD in prothorax shape in I. elegans females from both hybrid zones, but reveal that RCD in prothorax size is asymmetric, occurring only in gynochrome females from the NC hybrid zone. We also detected RCD in the prothorax shape of I. elegans males from the NC hybrid zone, extending previous evidence of RCD in male caudal appendages, while morphological covariation between male cerci and the prothorax was limited to size in I. elegans. Together, these findings illustrate how hybridization may generate morph-specific patterns of reproductive divergence.

The evolution of body shape reflects the interplay between functional constraints and habitat structure. In fishes, cave environments are well known for promoting regressive traits such as eye and pigment loss, yet their influence on overall body form remains poorly understood. Here, we examine patterns of body shape variation in cave- and surface-dwelling trichomycterid catfishes from northeastern Colombia to assess whether consistent associations exist between habitat type and morphology. Using geometric morphometric analyses, we quantified differences in body shape among species inhabiting subterranean and surface environments. Our results reveal significant habitat-associated differentiation in body shape along the main axes of morphological variation. Cave-dwelling species exhibit more elongated and fusiform body shapes, whereas surface-dwelling species tend to show deeper and more robust morphologies. In a functional context, these contrasting body patterns suggest associations with differing locomotor demands imposed by subterranean versus surface habitats. Although we do not explicitly test convergence or performance, the recurrence of similar body shapes among species from different clades occupying comparable habitats is consistent with repeated morphological responses to shared ecological constraints. Research HighligthsO_LIMultivariate shape analyses reveal significant habitat-associated variation in trichomycterid fishes. Recurrent morphological patterns suggest repeated responses potentially mediated by habitat constraints. C_LIO_LIBody shape differs consistently between cave- and surface-dwelling trichomycterids. Cave species exhibit more elongated and fusiform forms, whereas surface species display deeper body configurations. C_LI

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.

Matching habitat choice provides a mechanism for individuals to maximise their expected fitness by selecting an environment that better fits their phenotype. Many animals choose their local environment by evaluating levels of perceived predation risk against possible resource gain. To test if predation risk is a major driver of habitat choice, we quantify scototaxis, or preference for dark versus light backgrounds, in juvenile guppies. As light backgrounds increase visibility to predators, this aspect of habitat choice captures variation in boldness in small fishes. By rearing and testing 586 fish descended from ten natural populations from Trinidad under common garden conditions, we first quantify (broad sense) heritable variation, i.e. evolutionary potential, within populations. Next, we test for evolutionary divergence among populations in mean preference, and if present, whether ancestral predation regime is a mediator of divergence. Finally, we ask whether families and/or populations differ in the amount of behavioural variation they contain. Habitat choice varied among families (12% of total variance), consistent with heritable variation (0.2). We also found mean preference varies among populations (11% of total variance explained). Evolutionary divergence among-populations is partly explained by ancestral predation regime, with populations from low-predation sites showing a stronger average preference for dark backgrounds than high-predation populations from the same river. Additionally, we find that within-population behavioural variation is greater in high-predation populations. We conclude that guppy populations contain heritable variation that could facilitate adaptive evolution if scototaxis is subject to natural selection. Furthermore, while genetic drift may also contribute to evolutionary divergence among-populations, observed patterns are qualitatively consistent with local adaption to predation regime. Our results suggests that high predation sites favour bolder habitat choice on average, but also that local predation regime shape the evolutionary dynamics of variation, perhaps by maintaining shy-bold variation among-individuals or by favouring individuals with less-predicable behaviour.

The ability of animals to innovate - solve novel problems - can shape their ecology and evolution. Here we investigate how individual traits and environmental complexity relate to successful solving of a novel problem. We presented foraging bumble bees (Bombus impatiens) with artificial flowers of not-previously-encountered shapes and recorded the bees latency to access nectar. We measured individual foraging traits across multiple trips with simple flowers that did not require innovation, and bees were foraging either in a simple or complex environment (cluttered flight arena). Bees in complex environments took longer to find and were less likely to land on novel flowers, indicating that environmental complexity may take up cognitive resources and make search more difficult. However, we did not find an effect of environmental treatment on the ability or time to access reward in novel flowers once bees had landed on them. In contrast, behavioral traits significantly predicted how quickly bees solved novel flowers. In particular, overall foraging tempo as well as routine formation, i.e. how much bees followed a fixed route on known flowers, predicted innovation - faster bees innovated faster, and bees with more repetitive foraging sequences were slower to solve the novel tasks. Overall, while the degree of evolutionary novelty in tasks or solutions is always hard to evaluate, our findings demonstrate that environment and individual traits may affect innovation in different ways. Individuals in simple environments may be more likely to detect, and individuals that are generally faster and have a lower tendency to develop fixed routines may be more likely to solve, novel tasks.

Electric organ discharge (EOD) waveform diversity in African elephantfish is often attributed to sexual selection, yet EODs also mediate active electrolocation during prey detection, raising the possibility that natural selection on foraging ecology contributes to waveform divergence. Paramormyrops kingsleyae exhibits an intraspecific polymorphism where certain populations emit biphasic EODs whereas other populations emit triphasic waveforms. The genes underlying this polymorphism show signatures of selection; the polymorphism persists despite gene flow and is behaviorally discriminable by the fish themselves. If waveform differences influence prey detection during active electrolocation, biphasic and triphasic fish should consume systematically different prey. We tested this prediction using DNA metabarcoding of gut contents from 186 mormyrids representing 16 species across eight sites in Gabon, employing two independent COI primer sets for cross-validation and pairing dietary data with environmental invertebrate sampling to distinguish active prey preference from passive availability. At the community level in the diverse Bale Creek mormyrid assemblage, species identity was the dominant predictor of diet composition (R{superscript 2} {approx} 24%), consistent with phylogenetic signal in foraging ecology. Within P. kingsleyae, waveform type was the strongest independent predictor of dietary composition (R{superscript 2} = 5-6%), explaining variance independently of geographic region, sex, body size, and parasitism status -- a result concordant across both primer sets. Dietary differences were driven by prey species turnover rather than differential abundance of shared prey, and prey selectivity analyses confirmed that waveform types differ in which prey they actively prefer, not merely in what is locally available. These findings are consistent with natural selection on foraging ecology contributing to the maintenance of EOD waveform polymorphism, though the sensory mechanisms linking subtle waveform differences to prey detection remain an open question.

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.

Batesian mimicry is a defensive adaptation where predators learn to avoid aposematic prey and generalize their warning signals to phenotypically similar mimics. The phenotypic accuracy needed for mimics to benefit from this adaptation depends on the relative densities of models and mimics and the models unpalatability. As aposematic models become more unpalatable or more common relative to their mimics, warning signals become stronger, allowing even poor mimics to benefit. However, few studies have disentangled the importance of relative frequencies of models and mimics from absolute density of the prey community (both models and mimics) in driving relaxed selection on imperfect mimics. Here, we test the hypothesis that increasing model unpalatability and absolute prey community density accelerates predator avoidance learning and enhances protection for imperfect mimics. Using replicas of the model Adelpha iphiclus (Linnaeus), its imperfect mimic Adelpha serpa (Boisduval), and the palatable control Junonia evarete (Cramer), we conducted field experiments that enhanced model unpalatability and doubled absolute prey density while maintaining a constant ratio of model, mimic, and control phenotypes. We found that enhanced model unpalatability and increased absolute density significantly reduced predation on all species, highlighting absolute community density as an underappreciated mechanism shaping selection on imperfect Batesian mimics.

O_LIMany organisms alter phenotypically plastic traits in response to environmental cues to match their phenotypes with variable environments. In larval amphibians, development and growth rates respond to spatiotemporally variable mortality risk from predation, wetland drying, or resource limitation. However, these rates are also temperature-dependent for ectotherms. Although wild animals experience these factors simultaneously (e.g., thermal regimes, predation risk, resource limitation), most studies investigate their impacts in isolation, limiting our understanding of how they interact across ecological contexts. C_LIO_LIHere we simultaneously exposed larval Plains Leopard Frogs (Lithobates blairi) to varying resource levels and predation risk treatments across a thermal regime to investigate the joint effects of these ecological drivers on growth and development rates and their consequences for size and vagility after metamorphosis. We crossed two predation treatments (waterborne cues from Procambarus gracilis fed L. blairi larvae, control water) with three food resource levels (5%, 25%, 50% of body mass) and six thermal regimes (diel {+/-} 3{degrees}C cycles of 15, 20, 22, 24, 26, 28{degrees}C), replicating each combination five times for a total of 180 individuals. We recorded growth and development rates and completion of metamorphosis, then measured juvenile body size and jumping performance. C_LIO_LIThe number of larvae completing metamorphosis was primarily determined by temperature and temperature-dependent effects of resource limitation. Percent metamorphosis peaked at intermediate temperatures when resources were high and were higher in predation-risk treatments at the warmest temperatures. Under high resources, development and growth rates showed unimodal thermal responses that were absent when resources were constrained. Higher resources increased development rates, but proportional increases in growth maintained constant body size across temperatures. Post-metamorphic body size differed only by predation treatment, with predator-exposed individuals being smaller. Juvenile jumping performance increased with body size and individuals raised with high resources without predator cues exhibited the highest performance. C_LIO_LIThe absence of temperature effects on size at metamorphosis reflected unexpected coupling of growth and development rates across treatments, producing uniform body sizes. This pattern contrasts with the temperature-size rule and suggests that plastic responses may exhibit selection for a minimum viable size at metamorphosis. C_LI

Mate preferences are often influenced by the magnitude of sexual signals, which are presumed to indicate underlying aspects of signaller quality. Although the perception of these signals depends on sensory processes, the role of perceptual adaptations and constraints in mate assessment is frequently overlooked. Many sensory systems follow Webers law of proportional processing, where discrimination between signals is based upon their proportional, or relative, difference rather than their absolute difference. Because preference strength varies with relative trait magnitude, Webers law could strongly influence sexual selection, changing the coevolution of traits and preferences. Here, we explore the consequences of Webers law for sexual selection using individual-based models, applying Scalar Utility Theory to mate choice. We investigate the coevolution of male ornaments and female preferences under both Fisherian and good genes scenarios, as well as scrutinizing the sexual selection of multiple ornaments and preferences. Including Webers law in these models either reduced ornament exaggeration, or promoted exaggeration and diversification of ornaments and preferences, depending on the costs of choice and how rapidly female survival decreases when preferences evolve away from the naturally selected optimum. These results highlight the importance of perception and cognitive processing in shaping sexual selection and its evolutionary impacts.

Understanding the fitness advantages conferred by eusociality remains a central challenge in behavioral ecology. One promising approach is to identify collective strategies that shift efficiency within social groups. Here, we test the hypothesis that reserve workforces in eusocial insect colonies represent an adaptive mechanism that enhances flexibility and foraging efficiency under fluctuating environmental conditions. We examine how such reserve workers modulate the departure and return rates of foragers and how these time-dependent dynamics shape the colonys overall energetic balance. By integrating an energetic-balance framework with stochastic search simulations inspired by empirical results from Aphaenogaster senilis, we quantify the energetic requirements for colony viability, incorporating energy intake, search costs, and basal metabolic demands. Our results show that as colonies grow, maintaining a positive energy balance requires a disproportionately larger relative workforce. By modulating departure and return rates over time, colonies control the synchrony of their collective search and efficiently activate or suppress their reserve workforce to scale foraging effort as needed. These findings suggest that the "lazy" or weakly engaged workers commonly observed in large colonies function as an essential reserve that stabilizes colony energetics and enhances responsiveness. Together, our results provide a functional explanation for sublinear metabolic scaling in eusocial groups and highlight workforce modulation as a key factor underlying their energetic stability and evolutionary success.

Interactions between cleaner fish Labroides dimidiatus and client fish, from which cleaners remove ectoparasites and mucus, represent a textbook example of mutualism involving sophisticated strategic decision-making. However, cleaners must also face intraspecific social challenges within a size-based hierarchy, where the largest females may eventually change sex and become males with higher reproductive rates. Following 540 individuals over 11 months, we found that, contrary to expectations, slow-growing females spent more time cleaning and cheated more frequently, without causing more negative client responses than fast-growing females did. Instead, variation in growth was best explained by social factors: fast-growing individuals experienced reduced social control, while slow growers spent more time in proximity to dominant individuals. As there was no evidence that spawning activity affected growth patterns, it appears that fast growth as a viable strategy for becoming a male largely depends on the lack of control by dominants.

Finding resources for the colony is one of the most difficult and risky tasks for a social insect worker. A worker on a foraging trip can face a number of challenges, including interference from other individuals, her own errors, and environmental disturbances. Collectively, colonies may use a variety of strategies to minimize the impact of such perturbations on the foraging process. Here, we investigated how individual Solenopsis xyloni ant workers react to perturbation of an established pheromone trail. We trained foragers from colonies in the field to either a low or high concentration sucrose solution in a feeder on a T-maze setup, then replaced a section of floor covering, removing a section of the pheromone trail previously laid. We found that while ants made correct choices on the T-maze when the trail was intact, their choices did not differ from chance when the trail was absent, indicating strong reliance on a pheromone trail (and not, for example, memory) to return to the resource. Moreover, when the trail was absent, we found that a majority of ants abandoned the resource, and that even the ants that were able to reach the resource did not repair the perturbed trail. However, with a high-quality resource, more ants persisted in attempting to reach it (instead of abandoning). We interpret these responses in the framework of robustness mechanisms discussed in systems biology. Our study thus links individual and collective responses to perturbations, and provides an empirical example of how information use interacts with system robustness. Statements and declarationsThe authors have no competing interests to declare that are relevant to the content of this article.

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