New enantiornithine diversity in the Hell Creek Formation and the functional morphology of the avisaurid tarsometatarsus.

Enantiornithines were the most diverse group of birds during the Cretaceous, comprising over half of all known species from this period. The fossil record and subsequently our knowledge of this clade is heavily skewed by the wealth of material from Lower Cretaceous deposits in China. In contrast, specimens from Upper Cretaceous deposits are rare and typically fragmentary, yet critical for understanding the extinction of this clade across the K-Pg boundary. The most complete North American Late Cretaceous enantiornithine is Mirarce eatoni, a member of the diverse clade Avisauridae. Except for Mirarce, avisaurids are known only from isolated hindlimb elements from North and South America. Here we describe three new enantiornithines from the Maastrichtian Hell Creek Formation, two of which represent new avisaurid taxa. These materials represent a substantial increase in the known diversity of Enantiornithes in the latest Cretaceous. Re-examination of material referred to Avisauridae through phylogenetic analysis provides strong support for a more exclusive Avisauridae consisting of six taxa. Exploration of the functional morphology of the avisaurid tarsometatarsus indicates potential strong constriction and raptorial attributes. The lower aspect ratio of the tarsometatarsus facilitates a more biomechanically efficient lever system which in extant birds of prey equates to lifting proportionally heavier prey items. In addition, the proportional size and distal position of the m. tibialis cranialis tubercle of the tarsometatarsus is similar to the morphology seen in extant birds of prey. Together with the deeply-grooved metatarsal trochlea facilitating robust and likely powerful pedal digits, morphologies of the hindlimb suggest avisaurids as Late Cretaceous birds of prey.

Long- and short-term responses to climate change in body and appendage size of diverse Australian birds.

Changes to body size and shape have been identified as potential adaptive responses to climate change, but the pervasiveness of these responses has been questioned. To address this, we measured body and appendage size from 5013 museum bird skins of 78 ecologically and evolutionary diverse Australian species. We found that morphological change is a shared response to climate change across birds. Birds increased relative bill surface area, tarsus length, and relative wing length through time, consistent with expectations of increasing appendage size as climates warm. Furthermore, birds decreased in absolute wing length, consistent with the expectation of decreasing body size in warmer climates. Interestingly, these trends were generally consistent across different diets and migratory and thermoregulatory behaviors. Shorter term responses to higher temperatures were contrary to long-term effects for appendages, wherein relative appendage size decreased after hotter years, indicating the complex selective pressures acting on birds as temperatures rise with climate change. Overall, our findings support the notion that morphological adaptation is a widespread response to climate change in birds that is independent of other ecological traits.

Agile perching maneuvers in birds and morphing-wing drones.

Avian perching maneuvers are one of the most frequent and agile flight scenarios, where highly optimized flight trajectories, produced by rapid wing and tail morphing that generate high angular rates and accelerations, reduce kinetic energy at impact. While the behavioral, anatomical, and aerodynamic factors involved in these maneuvers are well described, the underlying control strategies are poorly understood. Here, we use optimal control methods on an avian-inspired drone with morphing wing and tail to test a recent hypothesis derived from perching maneuver experiments of Harris' hawks that birds minimize the distance flown at high angles of attack to dissipate kinetic energy before impact. The resulting drone flight trajectories, morphing sequence, and kinetic energy distribution resemble those measured in birds. Furthermore, experimental manipulation of the wings that would be difficult or unethical with animals reveals the morphing factors that are critical for optimal perching maneuver performance of birds and morphing-wing drones.

Species morphology better predicts plant-hummingbird interactions across elevations than nectar traits.

Species traits greatly influence interactions between plants and pollinators where floral nectar is the primary energy source fostering this mutualism. However, very little is known about how nectar traits mediate interactions in pollination networks compared with morphological traits. Here, we evaluated the role of morphological and nectar traits in shaping plant-hummingbird interaction networks along an elevation gradient. For this, we assessed patterns in floral phenotypic traits and network properties of plant species across elevations in Costa Rica. We also analysed whether plant species with generalized flower traits are ecological generalists and how morphological trait matching versus nectar traits affect interactions. We found marked variation in floral phenotypic traits and flower abundance of hummingbird-visited plant species across 10 sites along the elevation gradient. We did not find evidence for a relationship between flower morphology and nectar traits or between morphological and ecological generalization of plant species. Plant-hummingbird interaction frequency increased when the lengths of hummingbird bill and flower corolla were similar, indicating morphological matching, whereas nectar traits were unrelated to interactions. While nectar may play a difficult-to-detect secondary role within plant-hummingbird networks, our results reinforce the idea that morphological matching is an important factor in structuring ecological communities.

Muscle-controlled physics simulations of bird locomotion resolve the grounded running paradox.

Humans and birds use very different running styles. Unlike humans, birds adopt "grounded running" at intermediate speeds-a running gait where at least one foot always maintains ground contact. Avian grounded running is a paradox: Animals usually minimize locomotor energy expenditure, but birds prefer grounded running despite incurring higher energy costs. Using predictive gait simulations of the emu (Dromaius novaehollandiae), we resolve this paradox by demonstrating that grounded running represents an optimal gait for birds, from both energetics and muscle excitations perspectives. Our virtual experiments decoupled effects of posture and tendon elasticity, biomechanically relevant anatomical features that cannot be isolated in real birds. The avian body plan prevents (near) vertical leg postures, making the running style used by humans impossible. Under this anatomical constraint, grounded running is optimal if the muscles produce the highest forces in crouched postures, as is true in most birds. Shared anatomical features suggest that, as a behavior, avian grounded running first evolved within non-avian dinosaurs.

Tactile bill-tip organs in seabirds suggest conservation of a deep avian symplesiomorphy.

Birds' bills are their main tactile interface with the outside world. Tactile bill-tip organs associated with specialized foraging techniques are present in several bird groups, yet remain understudied in most clades. One example is Austrodyptornithes, the major seabird clade uniting Procellariiformes (albatrosses and petrels) and Sphenisciformes (penguins). Here, we describe the mechanoreceptor arrangement and neurovascular anatomy in the premaxillae of Austrodyptornithes. Using a wide phylogenetic sample of extant birds (361 species), we show that albatrosses and penguins exhibit complex tactile bill-tip anatomies, comparable to birds with known bill-tip organs, despite not being known to use tactile foraging. Petrels (Procellariidae, Hydrobatidae and Oceanitidae) lack these morphologies, indicating an evolutionary transition in bill-tip mechanosensitivity within Procellariiformes. The bill-tip organ in Austrodyptornithes may be functionally related to nocturnal foraging and prey detection under water, or courtship displays involving tactile stimulation of the bill. Alternatively, these organs may be vestigial as is likely the case in most palaeognaths (e.g. ostriches and emu). Ancestral state reconstructions fail to reject the hypothesis that the last common ancestor of Austrodyptornithes had a bill-tip organ; thus, tactile foraging may be ancestral for this major extant clade, perhaps retained from a deeper point in crown bird evolutionary history.

Investigation of Glandula Uropygialis in Different Avian Species Using Morphometric and Histological Methods.

The aim of this study was to identify the glandula uropygialis's macroanatomical and histological structure in male and female birds (stork, goose, eagle, pigeon, crow and sparrowhawk) that belong to different populations. A total of 58 specimens were used in the study. The study materials were delivered to our laboratory by nature conservation and national parks, and no animals were euthanised for this study. The longitudinal and transversal lengths, dorsoventral heights and feather length of the glandula uropygialis and papilla uropygialis were measured with a digital calliper. Haematoxylin-eosin and Masson trichrome staining were performed for histological examinations. The shape of the glandula uropygialis was found to be pear-shaped in pigeons and sparrowhawks, heart-shaped in geese, kidney-shaped in eagles and oval-fascule-shaped in storks. In the crow, the shape of the glandula uropygialis was different from the other species in that it was located dorso-caudally. Histomorphological examination of the glands of these species revealed that the gland generally consisted of two lobes, right and left, surrounded by connective tissue from the outside. The parenchyma of these lobes consisted of tubulo-alveolar secretory glands with a radial arrangement from the periphery to the centre. The glands within the lobes differed from the periphery to the centre and although the general histomorphological appearance was similar between bird species, some differences were observed between species. Similar stromal structure was observed in all bird species evaluated in the study. However, the reticular connective tissue forming the roof of the gland was found to be more developed in pigeons, especially in Masson's trichroma staining. The degenerative layer in the glands was more prominent in the eagle, followed by the goose and crow. Although a common general histological structure was observed among bird species, simple histomorphological differences were found between these formations, but no differences were found between the sexes. The results obtained will be compared with the findings of the aves class and will provide a data source for this special gland specific to birds. It is thought that the results obtained may help to determine the functional properties of the gland and contribute to the science of ornithology.

Direct evidence of frugivory in the Mesozoic bird Longipteryx contradicts morphological proxies for diet.

Diet is one of the most important aspects of an animal's ecology, as it reflects direct interactions with other organisms and shapes morphology, behavior, and other life history traits. Modern birds (Neornithes) have a highly efficient and phenotypically plastic digestive system, allowing them to utilize diverse trophic resources, and digestive function has been put forth as a factor in the selectivity of the end-Cretaceous mass extinction, in which only neornithine dinosaurs survived.1 Although diet is directly documented in several early-diverging avian lineages,2 only a single specimen preserves evidence of diet in Enantiornithes, the dominant group of terrestrial Cretaceous birds.3 Morphology-based predictions suggest enantiornithines were faunivores,4,5,6 although the absence of evidence contrasts with the high preservation potential and relatively longer gut-retention times of these diets. Longipteryx is an unusual Early Cretaceous enantiornithine with an elongate rostrum; distally restricted dentition7; large, recurved, and crenulated teeth8; and tooth enamel much thicker than other paravians.9 Statistical analysis of rostral length, body size, and tooth morphology predicts Longipteryx was primarily insectivorous.4,5 Contrasting with these results, two new specimens of Longipteryx preserve gymnosperm seeds within the abdominal cavity interpreted as ingesta. Like Jeholornis, their unmacerated preservation and the absence of gastroliths indicate frugivory.10 As in Neornithes,11 complex diets driven by the elevated energetic demands imposed by flight, secondary rostral functions, and phylogenetic influence impede the use of morphological proxies to predict diet in early-diverging avian lineages.

Endocranial development in non-avian dinosaurs reveals an ontogenetic brain trajectory distinct from extant archosaurs.

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form.

A Large Marine Bird (Aves: Procellariiformes) from the Eocene of Western Siberia.

The article describes the first find of a bird from the Paleogene of Siberia. A fragment of tibiotarsus from the Eocene Tavda Formation of the Tyumen Region (Western Siberia) is assigned to Procellariiformes. The bird is morphologically closer to Procellariidae, but comparable in size to albatrosses (Diomedeidae) and is assumed to represent the stem members of the family. The find indicated for the first time that either stem albatrosses or similar large Procellariiformes could have had a worldwide distribution as early as the Eocene.

Comparative Computed Tomographic Measurements of the Synsacrum in Some Birds / Mediciones Tomográficas Computarizadas Comparativas del Sinsacro en Algunas Aves

SUMMARY: The Synsacrum is constructed of pelvic bones, sacral, lumbar vertebrae and some thoracic vertebrae, and it takes the form of a thin sheet of bone which is curled downwards at the sides. This study is designed to assess the morphometric specialization of the lumbosacral portion of the vertebral section in flying and non-flying winged creatures. Several parameters (ratio of cranial/caudal parts and diameter/length) of the lumbosacral vertebral column were calculated in computed tomography (CT). The ratio of diameter/length of the synsacrum was the narrowest in ostrich while the widest was in pigeon. While the cranial/caudal ratio was the smallest in ostrich and the largest in pigeon. The parameters of penguin were closed to that of pigeon. It is supposed that this ratio is related to the bird locomotion, flying, swimming or/and strolling.

El sinsacro está formado por huesos pélvicos, sacros, vértebras lumbares y algunas vértebras torácicas, y toma la forma de una fina lámina de hueso curvada a los lados hacia abajo. Este estudio está diseñado para evaluar la especialización morfométrica de la porción lumbosacra de la sección vertebral en criaturas aladas voladoras y no voladoras. Se calcularon varios parámetros (relación de partes craneal/caudal y diámetro/longitud) de la columna vertebral lumbosacra en tomografía computarizada (TC). La relación diámetro/longitud del sinsacro fue la más estrecha en avestruz mientras que la más ancha fue en paloma. Mientras que la relación craneal/caudal fue la más pequeña en avestruz y la más grande en la paloma. Los parámetros del pingüino estaban cerrados en relación a los de la paloma. Se supone que esta relación está relacionada con la locomoción de las aves, volando, nadando y/o caminando.

Intersexual patterns of the digestive tract and body size are opposed in a large bird.

The appropriate structure of the digestive tract is crucial for individual adaptation to ecological conditions. In birds, the length of the small intestine, responsible for food absorption, is generally believed to be positively correlated with body size. In this study, we investigated the variation in small intestine length in the White Stork (Ciconia ciconia), a monomorphic species without visible sexual dimorphism, but characterized by differing parental efforts, which can be reflected by the small intestine lengths between the sexes. We examined the relationship between small intestine length and body size within the sexes. Our findings show that male White Storks have significantly shorter small intestines than females, despite having larger body sizes than the latter. Furthermore, we found a significant relationship between body size and small intestine length, but it was of a different nature in the two sexes. Males exhibited a previously unreported phenomenon, whereby increasing body size was associated with shortening small intestines, whereas females exhibited the opposite pattern. These novel findings shed light on the anatomical adaptations of the digestive tract in birds.

A new Paleogene fossil and a new dataset for waterfowl (Aves: Anseriformes) clarify phylogeny, ecological evolution, and avian evolution at the K-Pg Boundary.

Despite making up one of the most ecologically diverse groups of living birds, comprising soaring, diving and giant flightless taxa, the evolutionary relationships and ecological evolution of Anseriformes (waterfowl) remain unresolved. Although Anseriformes have a comparatively rich, global Cretaceous and Paleogene fossil record, morphological datasets for this group that include extinct taxa report conflicting relationships for all known extinct taxa. Correct placement of extinct taxa is necessary to understand whether ancestral anseriform feeding ecology was more terrestrial or one of a set of diverse aquatic ecologies and to better understand avian evolution around the K-T boundary. Here, we present a new morphological dataset for Anseriformes that includes more extant and extinct taxa than any previous anseriform-focused dataset and describe a new anseriform species from the early Eocene Green River Formation of North America. The new taxon has a mediolaterally narrow bill which is rarely found in previously described anseriform fossils. The matrix created to assess the placement of this taxon comprises 41 taxa and 719 discrete morphological characters describing skeletal morphology, musculature, syringeal morphology, ecology, and behavior. We additionally combine the morphological dataset with published sequences using Bayesian methods and perform ancestral state reconstruction for select morphological, ecological and behavioral characters. We recover the new Eocene taxon as the sister taxon to (Anseranatidae+Anatidae) across all analyses, and find that the new taxon represents a novel ecology within known Anseriformes and the Green River taxa. Results provide insight into avian evolution during and following the K-Pg mass extinction and indicate that Anseriformes were likely ancestrally aquatic herbivores with rhamphothecal lamellae..

Earliest evidence of avian primary feather moult.

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

Specialized diving traits in the generalist morphology of Fulica (Aves, Rallidae).

Foot-propelled diving comprises the primary locomotion-based feeding strategy for many birds, including families such as Phalacrocoracidae, Anhingidae, Podicipedidae, Gaviidae, and the diving ducks within Anatidae. While the morphology of specialized divers is well known, the corresponding morphology is less known for birds not as specialized but capable of diving, such as the coots (Rallidae, Fulica spp.). To compare the osteology of Fulica with other (non-diving) Rallidae, and with foot-propelled diving birds that are distantly related, we considered osteological characters, as well as the proportion of the hind limb bones and the femoral splay angle to construct a phylomorphospace, and to perform a comparative disparity analysis considering ecomorphologically relevant characters related to swimming and diving. Coots resulted to be significantly disparate from other Rallidae showing many traits of specialized foot-propelled divers, but only noticeable when compared with other rallids, as the degree of development of these traits is markedly less than in loons, grebes, or cormorants. This may correspond to a stabilizing selection of characteristics associated with a generalist morphology in Fulica. Studying adaptation in generalist taxa broadens our understanding of ecomorphologically significant features, thereby enabling us to generalize their evolutionary patterns.

Insights into microstructure and expression of markers of proliferation, apoptosis and T cells in the spleen of cattle egret (Bubulcus ibis).

The spleen is the largest secondary lymphoid organ with significant roles in pathogen clearance. It is involved in several avian diseases. The cattle egret is a wild insectivorous bird of agricultural and socioeconomic importance. Data related to microstructural features of cattle egret spleen are lacking. The present study investigated the gross anatomical, histological and immunohistochemical characteristics of the cattle egret spleen. Proliferation (PCNA and PHH3), apoptosis (cleaved caspase 3, C.CASP3) and T-cell (CD3 and CD8) markers were assessed. Grossly, the spleen appeared brownish red, oval-shaped and located at the oesophago-proventricular junction. Histologically, the spleen was surrounded by a thin capsule sending a number of trabeculae which contained branches of the splenic vessels. The white pulp consisted of the periarteriolar lymphoid sheath and periellipsoidal lymphatic sheath (PELS). The red pulp was formed of sinusoids and cords. The penicillar capillaries, which represent the terminal segments of the splenic arterial tree were highly branched, wrapped by prominent ellipsoids and directly connected to the splenic sinusoids, suggesting a closed type of circulation. Immunohistochemically, proliferating cell nuclear antigen (PCNA)-expressing cells were distributed with high counts throughout the splenic parenchyma, being highest within the splenic cords and PELS. Both PHH3- and C.CASP3-expressing cells revealed a similar pattern to that of PCNA, although with fewer counts. Large numbers of T cells were observed throughout the splenic parenchyma, mainly within the cords, as revealed by CD3 and CD8 immunoreaction. The present study provides a clear insight into the precise structure of the spleen in cattle egrets and thus improves our understanding about birds' immunity.

Early Jurassic origin of avian endothermy and thermophysiological diversity in dinosaurs.

A fundamental question in dinosaur evolution is how they adapted to long-term climatic shifts during the Mesozoic and when they developed environmentally independent, avian-style acclimatization, becoming endothermic.1,2 The ability of warm-blooded dinosaurs to flourish in harsher environments, including cold, high-latitude regions,3,4 raises intriguing questions about the origins of key innovations shared with modern birds,5,6 indicating that the development of homeothermy (keeping constant body temperature) and endothermy (generating body heat) played a crucial role in their ecological diversification.7 Despite substantial evidence across scientific disciplines (anatomy,8 reproduction,9 energetics,10 biomechanics,10 osteohistology,11 palaeobiogeography,12 geochemistry,13,14 and soft tissues15,16,17), a consensus on dinosaur thermophysiology remains elusive.1,12,15,17,18,19 Differential thermophysiological strategies among terrestrial tetrapods allow endotherms (birds and mammals) to expand their latitudinal range (from the tropics to polar regions), owing to their reduced reliance on environmental temperature.20 By contrast, most reptilian lineages (squamates, turtles, and crocodilians) and amphibians are predominantly constrained by temperature in regions closer to the tropics.21 Determining when this macroecological pattern emerged in the avian lineage relies heavily on identifying the origin of these key physiological traits. Combining fossils with macroevolutionary and palaeoclimatic models, we unveil distinct evolutionary pathways in the main dinosaur lineages: ornithischians and theropods diversified across broader climatic landscapes, trending toward cooler niches. An Early Jurassic shift to colder climates in Theropoda suggests an early adoption of endothermy. Conversely, sauropodomorphs exhibited prolonged climatic conservatism associated with higher thermal conditions, emphasizing temperature, rather than plant productivity, as the primary driver of this pattern, suggesting poikilothermy with a stronger dependence on higher temperatures in sauropods.

Nectar and floral morphology differ in evolutionary potential in novel pollination environments.

Plants can evolve rapidly after pollinator changes, but the response of different floral traits to novel selection can vary. Floral morphology is often expected to show high integration to maintain pollination accuracy, while nectar traits can be more environmentally sensitive. The relative role of genetic correlations and phenotypic plasticity (PP) in floral evolution remains unclear, particularly for nectar traits, and can be studied in the context of recent pollinator changes. Digitalis purpurea shows rapid recent evolution of corolla morphology but not nectar traits following a range expansion with hummingbirds added as pollinators. We use this species to compare PP, heritability, evolvability and integration of floral morphology and nectar in a common garden. Morphological traits showed higher heritability than nectar traits, and the proximal section of the corolla, which regulates access to nectar and underwent rapid change in introduced populations, presented lower integration than the rest of the floral phenotype. Nectar was more plastic than morphology, driven by highly plastic sugar concentration. Nectar production rate showed high potential to respond to selection. These results explain the differential rapid evolution of floral traits previously observed in this species and show how intrafloral modularity determines variable evolutionary potential in morphological and nectar traits.

Comparative study of the ciliary body and iris morphology in the anterior eye chamber of five different vertebrate classes.

One crucial component of the optical system is the ciliary body (CB). This body secretes the aqueous humour, which is essential to maintain the internal eye pressure as well as the clearness of the lens and cornea. The histological study was designed to provide the morphological differences of CB and iris in the anterior eye chambers of the following vertebrate classes: fish (grass carp), amphibians (Arabian toad), reptiles (semiaquatic turtle, fan-footed gecko, ocellated skink, Egyptian spiny-tailed lizard, Arabian horned viper), birds (common pigeon, common quail, common kestrel), and mammals (BALB/c mouse, rabbit, golden hamster, desert hedgehog, lesser Egyptian jerboa, Egyptian fruit bat). The results showed distinct morphological appearances of the CB and iris in each species, ranging from fish to mammals. The present comparative study concluded that the morphological structure of the CB and iris is the adaptation of species to either their lifestyle or survival in specific habitats.

Small body size is associated with increased evolutionary lability of wing skeleton proportions in birds.

Birds are represented by 11,000 species and a great variety of body masses. Modular organisation of trait evolution across birds has facilitated simultaneous adaptation of different body regions to divergent ecological requirements. However, the role modularity has played in avian body size evolution, especially small-bodied, rapidly evolving and diverse avian subclades, such as hummingbirds and songbirds, is unknown. Modularity is influenced by the intersection of biomechanical restrictions, adaptation, and developmental controls, making it difficult to uncover the contributions of single factors such as body mass to skeletal organisation. We develop a novel framework to decompose this complexity, assessing factors underlying the modularity of skeletal proportions in fore-limb propelled birds distributed across a range of body masses. We demonstrate that differences in body size across birds triggers a modular reorganisation of flight apparatus proportions consistent with biomechanical expectations. We suggest weakened integration within the wing facilitates radiation in small birds. Our framework is generalisable to other groups and has the capacity to untangle the multi-layered complexity intrinsic to modular evolution.