Functional Ecology

Urbanization creates novel environments that can drive phenotypic and behavioural responses, yet how multiple traits respond across spatial scales remains poorly understood. In particular, elevated ambient temperatures via the urban heat island effect may drive morphological and behavioural responses. We investigated body size, abdominal colouration, microhabitat use, behavioural thermoregulation and thermal offset relative to ambient air in the orb-weaving spider Araneus diadematus across rural-urban gradients in northern Belgium. Contrary to predictions from the temperature-size rule, body size increased with urbanization at large spatial scales, whereas size-corrected abdomen area--reflecting body condition and reproductive investment--declined with urbanization, with strongest support at local spatial scales. Abdominal colouration showed no response to urbanization despite evidence for both carotenoid-like pigments and melanin-associated structures. Nevertheless, body size and colouration covaried, with sites containing larger spiders tending to harbour darker individuals, whereas within sites larger individuals were slightly brighter than smaller conspecifics. Thermal responses showed little variation along the urbanization gradient. Retreats were consistently warmer than web hubs, and spiders maintained body temperatures above both their immediate microhabitat and ambient air. Only retreat-associated behavioural thermoregulation showed a weak decline with urbanization at local spatial scales. Our results reveal contrasting trait responses to urbanization across spatial scales and demonstrate that size-colour covariation can persist despite divergent responses of individual traits. These findings highlight the importance of considering multiple traits, their covariation and spatial scale to accurately understand and predict ecological responses of ectotherms to urban environments.

Climate warming is altering the timing of seasonal events across ecosystems, impacting the temporal synchrony of interactions among species1,2. For trophic interactions, the match-mismatch hypothesis predicts that when consumers become phenologically asynchronous with key ephemeral resources their fitness will decline3-5. Most studies of mismatch focus on single resource-consumer species pairs, and implicitly assume trophic specialisation. However, many consumers exploit more than one resource species, giving rise to several mechanisms whereby the negative impacts of mismatch on individuals and populations could be buffered6. Here we experimentally manipulate phenological asynchrony across 48 plant-caterpillar interactions in a spring woodland food-web system and assay caterpillar performance. As asynchrony increases, we find strong evidence for a decline in survival that generalises across host-caterpillar interactions, whereas caterpillar growth and development are largely unaffected. We also show that focus in the literature on a single model interaction (Oak-Winter Moth)7,8 has likely overestimated the general impact asynchrony in this system. The strength of the effect of mismatch varies markedly among host-plants, caterpillars, and their interactions--with a small number of interactions showing little or no decline in consumer performance despite substantial asynchrony. Our results demonstrate that the fitness consequences of phenological mismatch are widespread but interaction-specific, revealing substantial heterogeneity in how trophic interactions are expected to respond to climate-driven shifts in seasonal timing. This variation in response could allow resource diversity and resource switching to buffer consumer guilds against the phenological impacts of ongoing climate change, stabilising the abundance of caterpillars for higher trophic levels.

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

The exchange of energy and organisms across habitat boundaries links aquatic and terrestrial ecosystems and sustains ecosystem functioning. Although disturbance may disrupt these linkages, the mechanisms at play remain poorly understood. Here, we investigated the extent to which flow intermittency may disrupt riparian-aquatic ecosystem linkages by altering consumer communities in the recipient ecosystem or by altering resource quality in the donor ecosystem. We ran an experiment in an intermittent river network in California, focusing on a critical forest-to-river subsidy (organic matter in the form of leaf litter), its transformation, and its reciprocal benefit (aquatic insect production). Using three riparian species (willow, cottonwood, and oak) at sites spanning a gradient of flow permanence, we quantified intraspecific plasticity in leaf traits (specific leaf area, nitrogen and phosphorus concentrations, and {delta}13C), measured decomposition rates, and estimated the secondary production of aquatic shredders (Plecoptera). Across all leaf species, decomposition rates were 16-36% lower at intermittent than perennial sites, an effect largely driven by intraspecific leaf trait plasticity rather than changes in consumer abundance. At high flow intermittency, willow experienced water stress (enriched {delta}13C) and reduced specific leaf area, while cottonwood showed primarily stoichiometric responses (reduced leaf nitrogen and phosphorus). Despite these divergent strategies, all species produced lower-quality litter at intermittent sites. Variance partitioning confirmed that initial litter quality uniquely explained 51.5% of variation in decomposition rates, more than double the contribution of invertebrate community metrics; and structural equation modeling revealed that both leaf traits and stonefly (Plecoptera) secondary production significantly predicted decomposition rates, with leaf traits exerting the stronger effect. Notably, stonefly secondary production was 37-98% lower at intermittent sites across leaf species. Because these insects later emerge as terrestrial adults, they provide a significant energy flux to riparian predators, and, thus, impoverished litter quality suppresses the reciprocal transfer of energy back to terrestrial food webs. As drought intensifies globally, the decoupling of terrestrial-aquatic linkages may begin in the riparian canopy.

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

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

Many sea anemones reproduce asexually by pedal laceration, shedding fragments of the pedal disk that regenerate into polyps. How feeding affects the amount of this reproduction differs among species, but whether feeding also sets its timing has rarely been examined. We fed the sea anemone Cylista elegans (formerly Sagartia elegans) daily, every second day, every fourth day, or not at all, measured growth, laceration, and survival over 35 days, then reassigned anemones to new schedules to test whether laceration follows the current or previous schedule. Growth rose with feeding but saturated, animals fed daily and every second day growing at similar rates. Total laceration depended on whether an anemone was fed rather than how often. Every fed schedule produced more lacerates than starvation. The timing, by contrast, tracked the schedule closely. Laceration was suppressed for about a day after each meal and recovered before the next, so anemones fed every second or every fourth day lacerated on matching two- and four-day rhythms, while daily-fed animals were arrhythmic. When moved to a new schedule, the period shifted to match it, not the old one. Only starvation caused death. Asexual reproduction in C. elegans is therefore bound closely to feeding, which fuels laceration yet suppresses it during digestion, confining it between meals. This coupling distinguishes C. elegans from anemones with symbiotic microalgae, where starvation rather than feeding drives reproduction, and links it to the feeding-driven clonal proliferation of the invasive sea anemone Diadumene lineata.

SummaryO_LIIn woody perennials, reproductive allocation considered as a fraction of NPP (RA) has been rarely quantified, especially tracked across plant lifetimes, measured in biologically meaningful currencies, and separated from standing biomass. C_LIO_LIWe addressed this gap by measuring dry mass, nitrogen, and phosphorus for every aboveground tissue type for 14 iteroparous perennial shrubs tracked across their full lifetimes, calculating RA using four accounting schemes and three currencies. C_LIO_LIRA was substantially higher by mid-life than typical estimates from ecosystem-scale studies. Distinguishing standing biomass from yearly production further revised RA upwards. Using a nutrient currency (especially P) increased the perceived costs of reproduction. Propagules were highly nutrient-enriched and reproductive accessory costs consumed more nutrients than did the propagules themselves. Considering N and P resorption from senescing leaves and wood shrank the effective vegetative nutrient budget, further concentrating net annual nutrient demand in reproductive tissues. C_LIO_LIOur results highlight high investment in RA for woody perennials, especially using nutrient currencies. Broadly similar allocation patterns were observed across species with different functional traits and lifespans, suggesting generality that may apply across biomes. Widespread underestimation of RA in forest growth models likely overestimates the proportion of NPP available for vegetative growth, leading to substantial errors in predictions. C_LI

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

Biological rates, like growth, tend to have unimodal (hump-shaped) responses to temperature, and these relationships can vary among species and biological processes. In most systems, full thermal performance relationships are rarely characterized for interacting species (e.g., consumer-resource or host-pathogen pairs), making it challenging to predict how their interactions, and subsequently, how communities, will shift with climate change. We investigated how the thermal responses of eelgrass (Zostera marina, an important marine foundation species in the N. hemisphere) and an isopod grazer (Pentidotea resecata), which putatively acts as an indirect vector of eelgrass wasting disease, interact to affect eelgrass productivity and wasting disease dynamics. In a laboratory experiment crossing five temperatures, two grazing, and two disease exposure treatments, across various metrics, eelgrass growth responded unimodally to temperature in the absence of grazers. Grazers depressed plant growth and flattened its thermal performance curves. Thermal performance curves for isopods indicated that increases in grazing and survival at intermediate temperatures negated concurrent gains in plant growth at these temperatures, while decreased isopod survival at high temperatures reduced their top-down effect on eelgrass. Isopods had negligible effects on plant disease responses, but warming reduced the time to disease onset and increased final disease severity. Overall, whole-plant disease severity remained low and did not substantially affect eelgrass leaf elongation, net growth, or rhizome dry mass. However, disease-treatment plants grew more new leaves at intermediate temperatures, possibly to combat losses in photosynthetic capacity in diseased leaf tissue. These results indicate that climate change-associated warming will likely increase eelgrass vulnerability to wasting disease. However, in sublethal outbreaks, disease could have less of an impact on eelgrass productivity than warming-induced increases in grazing. Thus, ignoring grazer responses to temperature could result in unreliable predictions of eelgrass productivity under climate change.

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

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

Environmental warming is generally expected to increase metabolic demand in ectotherms. However, facultatively endothermic insects such as bumblebees regulate body temperature and may reduce thermogenic investment under warm conditions, potentially altering physiological performance and responses to climate change. We combined flow-through respirometry and infrared thermography to test how elevated ambient temperature (25 vs 35{degrees}C) affects feeding energetics and postprandial metabolism in the bumblebee Bombus terrestris. Bees maintained substantially lower thoracic temperature excess at 35{degrees}C than at 25{degrees}C, both before and during feeding. Feeding metabolic rate was also lower at 35{degrees}C and was strongly positively associated with thoracic temperature excess, indicating that feeding energetics were primarily explained by thermoregulatory state rather than ambient temperature alone. Elevated temperature reduced both the probability and energetic magnitude of specific dynamic action (SDA), including total SDA expenditure, early postprandial metabolism, and peak metabolic amplitude. In contrast, SDA duration and time to peak response showed little temperature dependence. Our results demonstrate that warming can suppress energetic expenditure in facultatively endothermic pollinators by limiting thermogenic investment and postprandial metabolic responses, potentially constraining the energetic flexibility underpinning foraging performance under climate warming.

For oviparous animals, the decision of where to lay eggs is critical, as offspring remain sessile from oviposition through hatching and are thus unable to escape unfavorable conditions. Consequently, females are expected to select oviposition sites that benefit embryo development and survival. This may be particularly relevant for arboreal frogs, which typically lay eggs on leaves overhanging water, where embryos are exposed to predation, desiccation, and other risks until hatching. Yet studies directly linking maternal substrate choice to embryo survival remain scarce. Here, we examine how oviposition substrate influences embryo survival in the Emerald glass frog (Espadarana prosoblepon), a species in which females deposit eggs on multiple substrates, providing a rare opportunity to test how oviposition decisions affect reproductive success. Monitoring clutches in situ, we compared microclimatic conditions, hatching success, and sources of embryo mortality between the most used substrates: the spike moss Selaginella diffusa and leaves. Additionally, we conducted a two-choice experiment in semi-captivity to test whether females preferentially select one substrate over the other. Although microclimatic conditions did not differ between substrates, hatching success was significantly higher on S. diffusa, which also experienced less predation. In the two-choice experiment, all females laid their eggs on S. diffusa, and those clutches had higher hatching success and faster embryonic development rates than those on leaves. Together, these results support the hypothesis that non-random oviposition site selection in E. prosoblepon is driven by the maximization of embryo survival, demonstrating that substrate choice has measurable fitness consequences for the offspring.

Heritable bacterial symbionts are pervasive in terrestrial arthropods, often imposing reproductive manipulations to promote their own spread within host populations. Co-infections are common, potentially allowing symbiont co-infectors to hitchhike through a host population. However, adverse thermal conditions can disrupt these communities, particularly when co-infectors vary in their thermal sensitivity. We used a multi-generation experiment to test whether warm (29 {degrees}C) conditions disrupted spread of heritable symbionts through uninfected populations of the spider, Mermessus fradeorum. We tested two common infection combinations: a single infection with a cytoplasmic incompatibility (CI) inducing Rickettsiella or a feminizing co-infection that included a feminizing Wolbachia, the same Rickettsiella, and up to three apparent hitchhikers (two additional Wolbachia strains and Tisiphia). We initiated replicate populations with 1/3 of one infection type and 2/3 uninfected spiders, evaluating population infection rate over 5 spider generations under different temperature regimes. Under cool (21{degrees}C) conditions, Wolbachia feminization drove co-infection to 88% and Rickettsiella CI drove single infection to 83% of host populations. Vertical transmission for all symbionts was high (97-99%) and hitchhiking symbionts also spread effectively. Under warm conditions, feminization and CI efficacy were reduced, and symbionts suffered variably reduced vertical transmission. Warm conditions ultimately destroyed the co-infecting symbiont consortium and impeded symbiont spread. On its own, though, Rickettsiella was still able to increase, despite reduced strength of CI. We hypothesize that contrasting tensions between feminizing spread of the symbiont consortium versus environmentally driven loss of function and transmission may explain observed patterns of mixed infections in field populations of this spider.

The impacts of a changing abiotic environment on fitness and performance arise not only from low tolerance to new environmental conditions, but also from changes in the abundance and interaction intensity with other species. The strength of the interaction may itself depend on how well each species performs across environments, but there is a dearth of studies investigating how intrinsic fitness and interaction intensity covary across an abiotic environmental gradient. We addressed this question in a hypersaline consumer-resource system: the microalga Dunaliella spp. grazed by the brine shrimp Artemia franciscana. We exposed four Dunaliella strains to a range of salinities above seawater, with or without brine shrimps, and tracked their population sizes over time and the survival of their predators, to estimate basic parameters of a Lotka-Volterra model. We found that the intrinsic growth rate of algae, the survival rate of predators, and the per-capita predation rate, all varied with salinity and algal strain. Significant interactions between strain and salinity further revealed that these ecological responses to salinity are evolvable. Together with correlations between demographic parameters across salinity, this suggests that predation may influence the evolution of salinity tolerance curves, blurring the line between the fundamental and realized niches.

The timing of reproductive processes can influence how mating interactions shape reproductive strategies. We examined how female age correlates with receptivity and preference in the jumping spider Habronattus formosus, a system characterized by elaborate male courtship and strong female choice. To test how receptivity changes across the post-maturation period, we paired females of different ages with males and quantified male courtship displays and mating outcomes. Females were not receptive immediately after maturation and instead exhibited receptivity primarily 2-3 weeks after maturation. We further found that the full male courtship predicted mating success in older females, whereas only the second stage of the courtship predicted success in younger females. The delay in female receptivity is consistent with expectations for strong female choice systems, where females may have a greater opportunity to evaluate male courtship before mating. These results highlight the importance of age-dependent changes in female reproductive state and suggest that the timing of receptivity may shape how different components of male courtship influence mating success.

Mass coral bleaching events driven by marine heatwaves are increasing in frequency and severity, yet the long-term recovery trajectories of surviving corals remain poorly understood. Here, leveraging a cohort of individual coral colonies with a decade of tracked environmental and bleaching history, we captured structural shifts in the coral thermal performance landscape. Specifically, we quantified thermal performance curves (TPCs) for photosynthesis and calcification in bleaching-resistant and bleaching-susceptible colonies of two ecologically dominant reef-building corals (Montipora capitata and Porites compressa) at four and six years following the 2019 marine heatwave in K[a]neohe Bay, Hawaii (2023 and 2025, respectively). Coral thermal performance shifted substantially between 2023 and 2025, and these shifts differed between species, bleaching phenotypes, and traits. In P. compressa, photosynthetic thermal optimum (Topt) shifted downward over time by 1.6{degrees}C across both phenotypes, suggesting recalibration toward prevailing conditions at the potential cost of future heat tolerance. In M. capitata, photosynthetic performance was lower in bleaching-susceptible corals in 2023 but converged by 2025, suggesting susceptible colonies recovered. In contrast, Topt of photosynthesis remained persistently higher in bleaching-resistant colonies, likely reflecting established symbiont communities. Critically, photosynthesis and calcification did not recover in parallel. Calcification TPCs for both phenotypes of M. capitata were stable across both timepoints, whereas calcification TPCs in P. compressa continued to change through 2025. These findings demonstrate that coral thermal performance is not static following heatwaves but continues to be reshaped over multi-year recovery periods, and that species-specific traits and strategies can fundamentally constrain the pace and coupling of physiological recovery. Furthermore, elevated thermal tolerance acquired through a heatwave can erode during prolonged periods of ambient temperatures. As recurrent bleaching events shorten recovery windows, understanding these dynamic physiological trajectories are essential for understanding and forecasting reef futures.

While mutualistic symbioses with nitrogen-fixing bacteria enable plants to access fixed nitrogen, they also require substantial carbon investment. Under carbon limitation, such as shading, shifts in biomass allocation can decouple symbiotic investment from leaf and root growth, potentially compromising plant nitrogen status. Because shading shifts biomass allocation toward light acquisition, it could influence nitrogen fixing symbiosis in two opposing ways. If nodulation remains coupled to leaves rather than roots, nitrogen status should be maintained despite reduced root growth. Alternatively, if root growth constrains nodulation, nitrogen status should decline. We tested these hypotheses by manipulating light availability (full sunlight vs. 50% shade) and quantifying biomass allocation and symbiotic nodulation. Under shading, plants allocated proportionally more biomass to shoots than to roots and invested less biomass in root nodules. Relationships between nodulation and leaf or root biomass differed between treatments but converged with increasing plant size, although shaded plants never attained the root biomass observed in full sunlight. Leaf nitrogen concentration was maintained under shading because nodulation remained coupled to leaf investment despite reduced root allocation. These findings highlight that, under carbon limitation, maintaining leaf and nodule coupling enables plants to reduce nodule investment without compromising the nitrogen benefits of symbiosis.

Flight feather vanes are the primary aerodynamic surface of the avian wing. Because loading varies across the wing, vane macrostructure should co-vary with local mechanical demands, yet comparative data on how barb and barbule traits change among remiges and between vane surfaces remain scarce. We quantified barb density, barbule density, barb angle, barb length, and vane width on both vanes at three measurement positions along the rachis of all remiges in four species with contrasting flight modes (white stork, common buzzard, house sparrow, pygmy cormorant), generating over 40,000 measurements across 15 response variables from 992 feathers of 41 individuals. Two complementary generalised additive models characterised variation along the spanwise, inter-vane, and longitudinal axes, and compared outer primaries, inner primaries, and secondaries as functional wing regions. Feather macrostructure varied along all three axes and outer primaries represent the most distinctive region, with lower leading-vane barb density, reduced barb angles, and vane width asymmetry two to three times higher than in inner primaries or secondaries. House sparrow exhibited the densest vane architecture and the highest vane width asymmetry, whereas the low wing-beat frequency species showed complex nonlinear spanwise patterns undetectable by single-feather sampling. Pygmy cormorant barbule density was 39-53% lower than in all other species, matching its wettable plumage strategy. Longitudinal gradients in barb density and barb angle (22-31% decline) were conserved across species. The avian wing is thus functionally regionalised at the macrostructural level, with vane architecture reflecting both aerodynamic and ecological pressures. Summary statementFine-scale vane measurements across all remiges in four species show macrostructural regionalisation of the avian wing, with outer primaries showing the most distinctive vane architecture.