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Biology is a wellspring of inspiration in engineering design. This paper delves into the application of elastic instabilities-commonly used in biological systems to facilitate swift movement-as a power-amplification mechanism for soft robots. Specifically, inspired by the nonlinear mechanics of the hummingbird beak-and shedding further light on it-we design, build and test a novel, rapid-response, soft end effector. The hummingbird beak embodies the capacity for swift movement, achieving closure in less than [Formula: see text]. Previous work demonstrated that rapid movement is achieved through snap-through deformations, induced by muscular actuation of the beak's root. Using nonlinear finite element simulations coupled with continuation algorithms, we unveil a representative portion of the equilibrium manifold of the beak-inspired structure. The exploration involves the application of a sequence of rotations as exerted by the hummingbird muscles. Specific emphasis is placed on pinpointing and tailoring the position along the manifold of the saddle-node bifurcation at which the onset of elastic instability triggers dynamic snap-through. We show the critical importance of the intermediate rotation input in the sequence, as it results in the accumulation of elastic energy that is then explosively released as kinetic energy upon snap-through. Informed by our numerical studies, we conduct experimental testing on a prototype end effector fabricated using a compliant material (thermoplastic polyurethane). The experimental results support the trends observed in the numerical simulations and demonstrate the effectiveness of the bio-inspired design. Specifically, we measure the energy transferred by the soft end effector to a pendulum, varying the input levels in the sequence of prescribed rotations. Additionally, we demonstrate a potential robotic application in scenarios demanding explosive action. From a mechanics perspective, our work sheds light on how pre-stress fields can enable swift movement in soft robotic systems with the potential to facilitate high input-to-output energy efficiency.

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There is general consensus among evolutionary biologists that natural selection drives phenotypic modifications within populations over generational time. How to reconstruct this historical process, however, has been discussed mostly in theoretical terms, and recommendations and explanations on how to translate such theoretical insights into practice are needed. The present study aims at providing a theory-supported practical guide on how to reconstruct historical evolutionary processes by applying a morphology-centered approach through a series of interdependent steps of descriptive morphology, functional analysis, ecological observation, integration of paleoecological data, and evolutionary synthesis. Special attention is given to the development of tests regarding the accuracy, closeness to reality, and plausibility of the hypotheses at every level of the reconstructive process. This morphology-centered approach had its beginnings in the wake of the evolutionary synthesis and is part of the scientifically necessary process of reciprocal testing of hypotheses generated by different methods and data for the reconstruction of evolutionary history.

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The larvae of Neuroptera are predators that feed by injecting bioactive compounds into their prey and then suctioning the fluids through modified mouthparts. We explore the evolutionary history of this feeding structure through the examination of a new fossil larva preserved in Late Cretaceous Kachin amber, which we describe as new genus and species, Electroxipheus veneficus gen et sp. nov. X-ray phase-contrast microtomography enabled us to study the anatomy of the larva in 3D, including the structure of the mouthparts and that of the venom delivery system. The specimen exhibited a unique combination of morphological traits not found in any known fossil or extant lacewing, including an unusual structure of the antenna. Phylogenetic analyses, incorporating a selection of living and fossil larval Neuroptera and enforcing maximum parsimony and Bayesian inference, identified the larva as belonging to the stem group Mantispoidea. The larva shows that the anatomy of the feeding and venom-delivery apparatus has remained unchanged in Neuroptera from the Cretaceous to the present. The morphology of the specimen suggests that it was an active predator, in contrast with the scarcely mobile, specialized relatives, like mantispids and berothids.

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The flow path of vertebrate hearts has a looped configuration characterized by curved (sigmoid) and twisted (chiral) components. The looped heart design is phylogenetically conserved among vertebrates and is thought to represent a significant determinant of cardiac pumping function. It evolves during the embryonic period of development by a process called "cardiac looping". During the past decades, remarkable progress has been made in the uncovering of genetic, molecular, and biophysical factors contributing to cardiac looping. Our present knowledge of the functional consequences of cardiac looping lags behind this impressive progress. This article provides an overview and discussion of the currently available information on looped heart design and its implications for the pumping function. It is emphasized that: (1) looping seems to improve the pumping efficiency of the valveless embryonic heart. (2) bilaterally asymmetric (chiral) looping plays a central role in determining the alignment and separation of the pulmonary and systemic flow paths in the multi-chambered heart of tetrapods. (3) chiral looping is not needed for efficient pumping of the two-chambered hearts of fish. (4) it is the sigmoid curving of the flow path that may improve the pumping efficiency of lower as well as higher vertebrate hearts.

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Phyllostomidae, the most diverse family of Neotropical bats, encompass 230 species with varied dietary habits and food acquisition methods. Their feeding niche diversification has shaped skull and wing morphologies through natural selection, reflecting food processing and flight strategies. Yet, evolution of bat hindlimbs, especially in phyllostomids, remains little understood. Previous studies highlighted the femur's morphology as a key to understanding the evolution of quadrupedalism in yangochiropteran bats, including the adept walking observed in vampire bats (Desmodontinae). Here, we aimed to describe the femoral morphological variation in Phyllostomidae, correlating this with body size and assessing the effects of phylogenetic history, dietary habits, and hindlimb usage. Analyzing 15 femoral traits from 45 species across 9 subfamilies through phylogenetically informed methods, we discovered a significant phylogenetic structure in femoral morphology. Allometric analysis indicated that body mass accounts for about 85% of the variance in phyllostomid femoral size and about 11% in femoral shape. Relatively smaller femurs showed to be typical in Stenodermatinae, Lonchophyllinae, and Glossophaginae, in contrast to the larger femurs of Phyllostominae, Desmodontinae, Micronycterinae, and Lonchorrhininae. Furthermore, extensive femur shape variation was detected, with the most distinct morphologies in vampire bats, followed by frugivorous species. Adaptive evolutionary models related to diet more effectively explained variations in femoral relative size and shape than stochastic models. Contrary to the conventional belief of limited functional demand on bat femurs, our findings suggest that femoral morphology is significantly influenced by functional demands associated with diet and food capture, in addition to being partially structured by body size and shared evolutionary history.

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Essential for sustaining a high metabolic rate is the efficient fragmentation of food, which is determined by molar morphology and the movement of the jaw. The latter is related to the jaw morphology and the arrangement of the masticatory muscles. Soricid jaw apparatuses are unique among mammals, as the articulation facet on the condylar process is separated into a dorsal and a ventral part, which has often been linked to more differentiated jaw motions. Soricidae also possess a remarkably elongated angular process. However, the precise function of the unique morphology of soricid jaw apparatuses has not been fully understood yet. By digitally reconstructing the masticatory musculature via the diffusible iodine-based contrast-enhanced computed tomography technique, we show how the unique jaw morphology is reflected in the spatial organization as well as the inner architecture and respective fascicle orientations of the muscles. From the lines of action of the m. masseter and the m. pterygoideus internus, both muscles inserting on the lateral and medial side of the angular process, respectively, we infer that the angular process is substantial for roll and yaw rotations of the mandible. The m. masseter is subdivided into four and the m. pterygoideus internus into five subunits, each exhibiting a slightly different line of action and torque. This enables Soricidae to adjust and adapt these rotational movements according to the properties of the ingested food, allowing for more efficient fragmentation. Additionally, those guided rotational motions allow for precise occlusion despite tooth wear. The temporalis is the largest of the adductor muscles and is mainly responsible for exerting the bite force. Overall, the unique jaw bone morphology in conjunction with the complex muscle arrangement may contribute towards a more efficient energy gain and the maintenance of a high metabolic rate, which is crucial for small-bodied mammals such as shrews.

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The physical interactions between organisms and their environment ultimately shape diversification rates, but the contributions of biomechanics to evolutionary divergence are frequently overlooked. Here, we estimated a performance landscape for biting in an adaptive radiation of Cyprinodon pupfishes, including scale-biting and molluscivore specialists, and compared performance peaks with previous estimates of the fitness landscape in this system. We used high-speed video to film feeding strikes on gelatin cubes by scale eater, molluscivore, generalist and hybrid pupfishes and measured bite dimensions. We then measured five kinematic variables from 227 strikes using the SLEAP machine-learning model. We found a complex performance landscape with two distinct peaks best predicted gel-biting performance, corresponding to a significant non-linear interaction between peak gape and peak jaw protrusion. Only scale eaters and their hybrids were able to perform strikes within the highest performance peak, characterized by larger peak gapes and greater jaw protrusion. A performance valley separated this peak from a lower performance peak accessible to all species, characterized by smaller peak gapes and less jaw protrusion. However, most individuals exhibited substantial variation in strike kinematics and species could not be reliably distinguished by their strikes, indicating many-to-many mapping of morphology to performance. The two performance peaks observed in the lab were partially consistent with estimates of a two-peak fitness landscape measured in the wild, with the exception of the new performance peak for scale eaters. We thus reveal a new bimodal non-linear biomechanical model that connects morphology to performance to fitness in a sympatric radiation of trophic niche specialists.

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Unravelling the functional steps that underlie major transitions in the fossil record is a significant challenge for biologists owing to the difficulties of interpreting functional capabilities of extinct organisms. New computational modelling approaches provide exciting avenues for testing function in the fossil record. Here, we conduct digital bending experiments to reconstruct vertebral function in non-mammalian synapsids, the extinct forerunners of mammals, to provide insights into the functional underpinnings of the synapsid-mammal transition. We estimate range of motion and stiffness of intervertebral joints in eight non-mammalian synapsid species alongside a comparative sample of extant tetrapods, including salamanders, reptiles and mammals. We show that several key aspects of mammalian vertebral function evolved outside crown Mammalia. Compared to early diverging non-mammalian synapsids, cynodonts stabilized the posterior trunk against lateroflexion, while evolving axial rotation in the anterior trunk. This was later accompanied by posterior sagittal bending in crown mammals, and perhaps even therians specifically. Our data also support the prior hypothesis that functional diversification of the mammalian trunk occurred via co-option of existing morphological regions in response to changing selective demands. Thus, multiple functional and evolutionary steps underlie the origin of remarkable complexity in the mammalian backbone.

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Populations may adapt to similar environments via parallel or non-parallel genetic changes, but the frequency of these alternative mechanisms and underlying contributing factors are still poorly understood outside model systems. We used QTL mapping to investigate the genetic basis of highly divergent craniofacial traits between the scale-eater (Cyprinodon desquamator) and molluscivore (C. brontotheroides) pupfish adapting to two different hypersaline lake environments on San Salvador Island, Bahamas. We lab-reared F2 scale-eater x molluscivore intercrosses from two different lake populations, estimated linkage maps, scanned for significant QTL for 29 skeletal and craniofacial traits, female mate preference, and sex. We compared the location of QTL between lakes to quantify parallel and non-parallel genetic changes. We detected significant QTL for six craniofacial traits in at least one lake. However, nearly all shared QTL loci were associated with a different craniofacial trait within each lake. Therefore, our estimate of parallel evolution of craniofacial genetic architecture could range from one out of six identical trait QTL (low parallelism) to five out of six integrated trait QTL (high parallelism). We suggest that pleiotropy and trait integration can affect estimates of parallel evolution, particularly within rapid radiations. We also observed increased adaptive introgression in shared QTL regions, suggesting that gene flow contributed to parallel evolution. Overall, our results suggest that the same genomic regions may contribute to parallel adaptation across integrated suites of craniofacial traits, rather than specific traits, and highlight the need for a more expansive definition of parallel evolution.

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The ability to communicate through vocalization plays a key role in the survival of animals across all vertebrate groups. Although avian reptiles have received much attention relating to their stunning sound repertoire, non-avian reptiles have been wrongfully assumed to have less elaborate vocalization types, and little is known about the biomechanics of sound production and their underlying neural pathways in this group. We investigated alarm calls of Gekko gecko using audio and cineradiographic recordings. Acoustic analysis revealed three distinct call types: a sinusoidal call type (type 1); a train-like call type, characterized by distinct pulse trains (type 3); and an intermediate type, which showed both sinusoidal and pulse train components (type 2). Kinematic analysis of cineradiographic recordings showed that laryngeal movements differ significantly between respiratory and vocal behavior. During respiration, animals repeatedly moved their jaws to partially open their mouths, which was accompanied by small glottal movements. During vocalization, the glottis was pulled back, contrasting with what has previously been reported. In vitro retrograde tracing of the nerve innervating the laryngeal constrictor and dilator muscles revealed round to fusiform motoneurons in the hindbrain-spinal cord transition ipsilateral to the labeled nerve. Taken together, our observations provide insight into the alarm calls generated by G. gecko, the biomechanics of this sound generation and the underlying organization of motoneurons involved in the generation of vocalizations. Our observations suggest that G. gecko may be an excellent non-avian reptile model organism for enhancing our understanding of the evolution of vertebrate vocalization.

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OBJECTIVES: The elbow of Nacholapithecus has been extensively described qualitatively, however its ulnar morphology has never been the focus of an in-depth quantitative analysis before. Hence, our main aim is quantifying the proximal ulnar morphology in Nacholapithecus and exploring whether it is similar to those of Equatorius and Griphopithecus as previously reported. MATERIALS AND METHODS: We compared Nacholapithecus proximal ulnar morphology with a sample of extant and extinct anthropoids through principal component analysis and agglomerative hierarchical cluster analysis. Moreover, we calculated the Cophenetic Correlation Coefficient and checked for taxonomical group mean differences through MANOVA and pairwise post-hoc comparisons, as well as the phylogenetic signal in the variables used. RESULTS: The Nacholapithecus ulna displays a moderately long and relatively narrow olecranon, a relatively wide trochlear surface-radial notch width, and a relatively thin sigmoid notch depth. These features resemble those of large papionins and chimpanzees, and some extinct taxa, mainly Equatorius. DISCUSSION: Results presented here reinforce previous inferences on the functional morphology of the Nacholapithecus elbow, showing adaptations for general quadrupedal behaviors. However, other derived features (e.g., a relatively wide trochlear surface) might be associated with the ape-like traits described for its distal humerus (e.g., wide trochlear groove), thus displaying a combination of primitive and derived features in the proximal ulna. Finally, affinities with large papionins could suggest the presence of some terrestrial habits in Nacholapithecus. However, the lack of evidence in the rest of the skeleton prevents us from suggesting terrestrial affinities in this taxon in a conclusive manner.

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Talpid moles (Talpidae, Eulipotyphla) are mammals highly specialised in burrowing using their forelimbs. Fossoriality has allowed moles to expand their ecological niche by enabling access to subterranean resources and spaces. This specialisation in burrowing has led to adaptations in the forelimb bones of moles for humeral rotation digging, a distinctive strategy unparalleled among other diggers. While bone robustness has been examined in moles through external morphology, the adaptation of bone microstructure to digging strategy remains unclear. Based on two assumptions, (1) the humerus of moles is subjected to a torsional load due to humeral rotation digging, and (2) the magnitude of torsional load correlates with the compactness of the substrate in which the individuals can dig, we hypothesised that humeral rotation digging influences bone microstructure. Comparative analyses of transverse sections from the humeri and femora of three mole species (Mogera imaizumii, Mogera wogura and Urotrichus talpoides; Talpidae) and an outgroup eulipotyphlan (Suncus murinus; Soricidae) revealed that (1) vascular canals distributed in the humeri of moles align more predominantly circumferential along the bone walls, indicating an adaptation to the torsion generated by humeral rotation digging, and (2) the laminarity of vascular canals, particularly in Mogera species compared with Urotrichus, potentially reflects differences in the magnitude of load due to substrate compactness during digging. The aligned vascular canals are distinctive traits not observed in mammals employing other digging strategies. This suggests that vascular canal laminarity can be an indicator of not only humeral rotation digging in fossorial animals, but also the variation of eco-spaces in talpid species.

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The subgenual organ complex of stick insects has a unique neuroanatomical organisation with two elaborate chordotonal organs, the subgenual organ and the distal organ. These organs are present in all leg pairs and are already developed in newly hatched stick insects. The present study analyses for the first time the morphology of sensory organs in the subgenual organ complex for a membrane connecting the two sensory organs in newly hatched insects (Sipyloidea chlorotica (Audinet-Serville 1838)). The stick insect legs were analysed following hatching by axonal tracing and light microscopy. The subgenual organ complex in first juvenile instars shows the sensory organs and a thin membrane connecting the sensory organs resembling the morphology of adult animals. Rarely was this membrane not detected, where it is assumed as not developed during embryogenesis. The connection appears to influence the shape of the subgenual organ, with one end extending towards the distal organ as under tension. These findings are discussed for the following functional implications: (1) the physiological responses of the subgenual organ complex to mechanical stimuli after hatching, (2) the influence of the membrane on the displacement of the sensory organs, and (3) the connection between the subgenual organ and distal organ as a possible functional coupling.

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Epiphytes develop anatomical features to improve efficiency of the uptake of water and nutrients, such as absorptive foliar scales or a velamen radicum. Despite substantial studies on the occurrence, morphology, development and phylogeny of the velamen, most of the available literature is focused on Orchidaceae, making current knowledge on velamen clearly biased. A recent publication firmly established that velamina are common in Anthurium species. Thus, this study provides further insights by describing velamen morphological characteristics of Anthurium species and classifying them into different velamen types. Furthermore, we investigate if the different velamen morphological traits are clade-specific and phylogenetically conserved within the genus. Using SEM, we performed a morphological study on 89 Anthurium species, describing six micromorphological traits of velamen and exodermis, following traits used to classify Orchidaceae velamen by Porembski & Barthlott (1988). We distinguished nine velamen types, including two that are unique to Anthurium and not similar to any type found in Orchidaceae. Comparing velamen morphology within the phylogenetic tree of Anthurium revealed clear phylogenetic signals. This study provides detailed morphological descriptions among 89 species of Anthurium from the Araceae, and substantially broadens our knowledge of this tissue. However, velamen function has been even less studied, with hardly anything known about functional significance of having secondary cell wall thickening and perforations on velamen cell walls. Therefore, a logical next step would be to connect these anatomical features to their functions.

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The mechanism by which insects achieve attachment and locomotion across diverse substrates has long intrigued scientists, prompting extensive research on the functional morphology of attachment pads. In stick insects, attachment and locomotion are facilitated by two distinct types of smooth cuticular attachment pads: the primary adhesion force-generating arolium and the friction force-generating euplantulae. They are both supported by an adhesive secretion delivered into the interspace between the attachment pads and the substrate. In this study, we analysed and compared internal morphology, material composition and ultrastructure, as well as the transportation pathways in both adhesive organs in the stick insect Medauroidea extradentata using scanning electron microscopy, micro-computed tomography, light microscopy, and confocal laser scanning microscopy. Our observations revealed structural differences between both attachment pads, reflecting their distinct functionality. Furthermore, our results delineate a potential pathway for adhesive secretions, originating from exocrine epidermal cells and traversing various layers before reaching the surface. Within the attachment pad, the fluid may influence the viscoelastic properties of the pad and control the attachment/detachment process. Understanding the material composition of attachment pads and the distribution process of the adhesive secretion can potentially aid in the development of more effective artificial attachment systems.

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The neurocranial elevation generated by axial muscles is widespread among aquatic gnathostomes. The mechanism has two functions: first, it contributes to the orientation of the mouth gape, and second, it is involved in suction feeding. To provide such mobility, anatomical specialization of the anterior part of the vertebral column has evolved in many fish species. In modern chimaeras, the anterior part of the vertebral column develops into the synarcual. Possible biological roles of the occipital-synarcual joint have not been discussed before. Dissections of the head of two species of ratfishes (Chimaera monstrosa and Chimaera phantasma) confirmed the heterocoely of the articulation surface between the synarcual and the neurocranium, indicating the possibility of movements in the sagittal and frontal planes. Muscles capable of controlling the movements of the neurocranium were described. The m. epaxialis is capable of elevating the head, the m. coracomandibularis is capable of lowering it if the mandible is anchored by the adductor. Lateral flexion is performed by the m. lateroventralis, for which this function was proposed for the first time. The first description of the m. epaxialis profundus is given, its function is to be elucidated in the future. Manipulations with joint preparations revealed a pronounced amplitude of movement in the sagittal and frontal planes. Since chimaeras generate weak decrease in pressure in the oropharyngeal cavity when sucking in prey, we hypothesised the primary effect of neurocranial elevation, in addition to the evident lateral head mobility, is accurate prey targeting.

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The long, gracile morphology of the limb bones of the Late Miocene hyaenid Ictitherium ebu has led to the hypothesis that this animal was cursorial. The forelimb and femur of the holotype were compared with specimens of extant Hyaenidae and Canidae. Two morphometric methods were used. The first used measurements to calculate indices of different morphological characters. The second method involved capturing photographs of the anterior distal humerus of each specimen, mapping six landmarks on them, and calculating truss distances. These distances represent a schematic reproduction of the elbow. Multivariate statistical analysis primarily separated the data based on taxonomy, yet locomotor and habitat categories were also considered. Ictitherium ebu has an overall morphology similar to that of the maned wolf and a distal humerus reminiscent of that of the aardwolf. The long, gracile limb bones of I. ebu are suggested to be adaptations for pouncing on prey, for locomotor efficiency, and for looking over the tall grass of the open environments the animal lived in, much like the present-day maned wolf.

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The family Bovidae [Mammalia: Artiodactyla] is speciose and has extant representatives on every continent, forming key components of mammal communities. For these reasons, bovids are ideal candidates for studies of ecomorphology. In particular, the morphology of the bovid humerus has been identified as highly related to functional variables such as body mass and habitat. This study investigates the functional morphology of the bovid distal humerus in isolation due to its increased likelihood of preservation in the fossil record, and the resulting opportunity for a better understanding of the ecomorphology of extinct bovids. A landmark scheme of 30 landmarks was used to capture the 3D distal humerus morphology in 111 extant bovid specimens. We find that the distal humerus has identifiable morphologies associated with body mass, habitat preference and tribe affiliation and that some characteristics are shared between high body mass bovids and those living on hard, flat terrain which is likely due to the high stress on the bone in both cases. We directly apply our findings regarding extant bovids to the extinct alcelaphine bovid, Rusingoryx atopocranion from the mid to late Pleistocene (>33-45 ka) Lake Victoria region of Kenya. This species is known for some peculiar morphologies including a domed cranium with hollow nasal crests, and having small hooves for a bovid of its size. Another interesting aspect of Rusingoryx's skeletal morphology which has not been addressed is an unusual protrusion on the lateral epicondyle of the distal humerus. Despite considerable individual variation in the Rusingoryx specimens, we find evidence to support its historical assignment to the tribe Alcelaphini, and that it likely preferred open grassland habitats, which is consistent with independent reconstructions of the palaeoenvironment. We also provide the most accurate body mass estimate for Rusingoryx to date, based on distal humerus centroid size. Overall, we are able to conclude that the distal humerus in extant bovids is highly informative regarding body mass, habitat preference and tribe, and that this can be applied directly to a fossil taxon with promising results.

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New World porcupines (Erethizontinae) originated in South America and dispersed into North America as part of the Great American Biotic Interchange (GABI) 3-4 million years ago.1 Extant prehensile-tailed porcupines (Coendou) today live in tropical forests of Central and South America.2,3 In contrast, North American porcupines (Erethizon dorsatum) are thought to be ecologically adapted to higher-latitude temperate forests, with a larger body, shorter tail, and diet that includes bark.4,5,6,7 Limited fossils8,9,10,11,12,13 have hindered our understanding of the timing of this ecological differentiation relative to intercontinental dispersal during the GABI and expansion into temperate habitats.14,15,16,17,18 Here, we describe functionally important features of the skeleton of the extinct Erethizon poyeri, the oldest nearly complete porcupine skeleton documented from North America, found in the early Pleistocene of Florida. It differs from extant E. dorsatum in having a long, prehensile tail, grasping foot, and lacking dental specializations for bark gnawing, similar to tropical Coendou. Results from phylogenetic analysis suggest that the more arboreal characteristics found in E. poyeri are ancestral for erethizontines. Only after it expanded into temperate, Nearctic habitats did Erethizon acquire the characteristic features that it is known for today. When combined with molecular estimates of divergence times, results suggest that Erethizon was ecologically similar to a larger species of Coendou when it crossed the Isthmus of Panama by the early Pleistocene. It is likely that the range of this more tropically adapted form was limited to a continuous forested biome that extended from South America through the Gulf Coast.

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Aquatic birds represent diverse ecologies and locomotion types. Some became flightless or lost the ability for effective terrestrial locomotion, yet, certain species excel in water, on land, and in air, despite differing physical characteristics associated with each medium. In this exploratory study, we intend to quantitatively analyze the morphological variety of multiple limb bones of aquatic birds using 3D geometric morphometrics. Morphological variation is mainly driven by phylogeny, which also affects size and locomotion. However, the shape of the ulna, including the proportion and orientation of the epiphyses is influenced by size and aquatic propulsive techniques even when phylogeny is taken into consideration. Certain trends, possibly linked to functions, can be observed too in other bones, notably in cases where phylogenetic and functional signals are probably mixed when some taxa only englobe species with similar functional requirements: penguins exhibit the most distinctive wing bone morphologies, highly adapted to wing-propulsion; advanced foot-propellers exhibit femur morphology that reduces proximal mobility but supports stability; knee structures, like cnemial crests of varied sizes and orientations, are crucial for muscle attachments and efficient movement in water and on land; taxa relying on their feet in water but retaining terrestrial abilities share features enabling swimming and walking postures. Size-linked changes distinguish the wing bones of non-wing-propelled taxa. For hindlimbs, larger size relates to robust bones probably linked to terrestrial abilities, but robustness in femora can be connected to foot-propulsion. These results help us better understand birds' skeletal adaptation and can be useful inferring extinct species' ecology.