Origin of feathered flight

The origin of flight in birds and theropod dinosaurs is a many-sided and debatable problem. We develop a new approach to the resolution of this problem, combining terrestrial and arboreal hypotheses of the origin of flight. The bipedalism was a key adaptation for the development of flight in both birds and theropods. The bipedalism dismissed the forelimbs from the supporting function and promoted transformation into wings. For the development of true flapping avian flight, a key role was played by the initial universal anisodactylous foot of birds. This foot pattern provided a firm support on both land and trees. Theropod dinosaurs, archaeopteryxes, and some other early feathered creatures had a pamprodactylous foot and, hence, they developed only gliding descent. Early birds descended by flattering parachuting with the use of incipient wings; this gave rise to true flight. Among terrestrial vertebrates, only bats, pterosaurians, and birds developed true flapping flight, although they followed different morphofunctional pathways when solving this task. However, it remains uncertain what initiated the adaptation of the three groups for the air locomotion. Nevertheless, the past decade has provided unexpectedly abundant paleontological data, which facilitate the resolution of this question with reference to birds.     

Hip extensor EMG and forelimb/hind limb weight support asymmetry in primate quadrupeds

Higher weight support on the hind limb than forelimb is among the distinctive characteristics of primate quadrupeds. Although often assumed to be due to a more posteriorly positioned whole body center of mass, there are little data to support such a difference. Reynolds (1985. Am J Phys Anthropol 67:335-349) notes that the distribution of forces on the limbs can also be influenced by average limb posture, but suggests that this effect is too small to account for the asymmetry in weight support observed in primates. Instead, he proposes that high hind limb forces are brought about by an active process of shifting weight off the forelimbs and onto the hind limbs through use of hind limb retractors. In this study, we use video records of walking animals to explore the degree to which average limb posture in primates and other quadrupedal mammals deviates from vertical, and use electromyography to test Reynolds' model of hind limb retractor activity and posterior weight shift. The limb posture results indicate that primate forelimbs oscillate about a vertical or slightly retracted axis, and though the hind limbs are slightly protracted, the magnitude of deviation from vertical is too small to have a major effect on weight support distribution. The electromyographic results reveal higher levels of hip extensor activity in antipronograde primates that bear a higher proportion of weight on their hind limbs. This lends support to Reynolds' suggestion that some primates use muscles to actively shift weight onto hind limbs to relieve stresses on forelimbs less well structured for weight support.     

Stresses on the limbs of quadrupedal primates

Data is presented from eight primates on the ground reaction forces on the limbs during locomotion. These subjects supported from 30 to 45% of their body weight on their forelimbs. Other quadrupedal mammals support 55-60% of their body weight on their forelimbs. The increase of peak vertical force with speed varies greatly between the subjects. The variation in weight supported by the forelimbs and the peak forces on the forelimbs is proposed to correlate with variation in locomotor adaptations. It is suggested that the occurrence of bipedalism in primates represents the extreme expression of the tendency in primates to reduce the compressive forces on their forelimbs.     

Integration and dissociation of limb elements in flying vertebrates: a comparison of pterosaurs, birds and bats

Flapping flight has evolved independently in three vertebrate clades: pterosaurs, birds and bats. Each clade has a unique flight mechanism involving different elements of the forelimb. Here, patterns of limb integration are examined using partial correlation analysis within species and matrix correlation analysis across species to test whether the evolution of flapping flight has involved developmental dissociation of the serial homologues in the fore- and hind limb in each clade. Our sample included seven species of birds, six species of bats, and three species of pterosaurs for which sufficient sample sizes were available. Our results showed that, in contrast to results previously reported for quadrupedal mammals, none of the three clades demonstrated significant integration between serial homologues in the fore- and hind limb. Unexpectedly, there were few consistent patterns of within-forelimb correlations across each clade, suggesting that wing integration is not strongly constrained by functional relationships. However, there was significant integration within the hind limbs of pterosaurs and birds, but not bats, possibly reflecting the differing functions of hind limbs (e.g. upright support vs. suspension) in these clades.     

First record of a pterosaur landing trackway

The terrestrial progression of pterosaurs, the flying reptiles of the Mesozoic Era, has been debated for over two centuries. The recent discovery of quadrupedal pterodactyloid pterosaur tracks from Late Jurassic sediments near Crayssac, France, shows that the hindlimbs moved parasagittally, as in mammals, birds and other dinosaurs, and the hypertrophied forelimbs could make tracks both close to the body wall and far outside it. Their manus tracks are unique in form, position and kinematics, which would be expected because the forelimbs were used for flight. Here, we report the first record of a pterosaur landing track, which differs substantially from typical walking trackways. The individual landed on both hind feet in parallel fashion, dragged its toes slightly as it left the track, landed again almost immediately and placed the hindfeet parallel again, then placed its forelimbs on the ground, took another short step with both hindlimbs and adjusted its forelimbs, and then began to walk off normally. The trackway shows that pterosaurs stalled to land, a reflection of their highly developed capacity for flight control and manoeuverability.     

Flapping before Flight: High Resolution,Three-Dimensional Skeletal Kinematics of Wings and Legs during Avian Development

Some of the greatest transformations in vertebrate history involve developmental and evolutionary origins of avian flight. Flight is the most power-demanding mode of locomotion, and volant adult birds have many anatomical features that presumably help meet these demands. However, juvenile birds, like the first winged dinosaurs, lack many hallmarks of advanced flight capacity. Instead of large wings they have small “protowings”, and instead of robust, interlocking forelimb skeletons their limbs are more gracile and their joints less constrained. Such traits are often thought to preclude extinct theropods from powered flight, yet young birds with similarly rudimentary anatomies flap-run up slopes and even briefly fly, thereby challenging longstanding ideas on skeletal and feather function in the theropod-avian lineage. Though skeletons and feathers are the common link between extinct and extant theropods and figure prominently in discussions on flight performance (extant birds) and flight origins (extinct theropods), skeletal inter-workings are hidden from view and their functional relationship with aerodynamically active wings is not known. For the first time, we use X-ray Reconstruction of Moving Morphology to visualize skeletal movement in developing birds, and explore how development of the avian flight apparatus corresponds with ontogenetic trajectories in skeletal kinematics, aerodynamic performance, and the locomotor transition from pre-flight flapping behaviors to full flight capacity. Our findings reveal that developing chukars (Alectoris chukar) with rudimentary flight apparatuses acquire an “avian” flight stroke early in ontogeny, initially by using their wings and legs cooperatively and, as they acquire flight capacity, counteracting ontogenetic increases in aerodynamic output with greater skeletal channelization. In conjunction with previous work, juvenile birds thereby demonstrate that the initial function of developing wings is to enhance leg performance, and that aerodynamically active, flapping wings might better be viewed as adaptations or exaptations for enhancing leg performance.     

Flapping tail membrane in bats produces potentially important thrust during horizontal takeoffs and very slow flight

Historically, studies concerning bat flight have focused primarily on the wings. By analyzing high-speed video taken on 48 individuals of five species of vespertilionid bats, we show that the capacity to flap the tail-membrane (uropatagium) in order to generate thrust and lift during takeoffs and minimal-speed flight (<1 m ^s−1) was largely underestimated. Indeed, bats flapped the tail-membrane by extensive dorso-ventral fanning motions covering as much as 135 degrees of arc consistent with thrust generation by air displacement. The degree of dorsal extension of the tail-membrane, and thus the potential amount of thrust generated during platform launches, was significantly correlated with body mass (P = 0.02). Adduction of the hind limbs during upstrokes collapsed the tail-membrane thereby reducing its surface area and minimizing negative lift forces. Abduction of the hind limbs during the downstroke fully expanded the tail-membrane as it was swept ventrally. The flapping kinematics of the tail-membrane is thus consistent with expectations for an airfoil. Timing offsets between the wings and tail-membrane during downstrokes was as much as 50%, suggesting that the tail-membrane was providing thrust and perhaps lift when the wings were retracting through the upstoke phase of the wing-beat cycle. The extent to which the tail-membrane was used during takeoffs differed significantly among four vespertilionid species (P = 0.01) and aligned with predictions derived from bat ecomorphology. The extensive fanning motion of the tail membrane by vespertilionid bats has not been reported for other flying vertebrates.     

Heritabilities of Directional Asymmetry in the Fore- and Hindlimbs of Rabbit Fetuses

Directional asymmetry (DA), where at the population level symmetry differs from zero, has been reported in a wide range of traits and taxa, even for traits in which symmetry is expected to be the target of selection such as limbs or wings. In invertebrates, DA has been suggested to be non-adaptive. In vertebrates, there has been a wealth of research linking morphological asymmetry to behavioural lateralisation. On the other hand, the prenatal expression of DA and evidences for quantitative genetic variation for asymmetry may suggest it is not solely induced by differences in mechanic loading between sides. We estimate quantitative genetic variation of fetal limb asymmetry in a large dataset of rabbits. Our results showed a low but highly significant level of DA that is partially under genetic control for all traits, with forelimbs displaying higher levels of asymmetry. Genetic correlations were positive within limbs, but negative across bones of fore and hind limbs. Environmental correlations were positive for all, but smaller across fore and hind limbs. We discuss our results in light of the existence and maintenance of DA in locomotory traits.     

The role of wing length in the evolution of avian flightlessness

Flightlessness has evolved independently in at least 11 extant avian families. A number of hypotheses have been proposed to explain these transitions in individual families, including release from predation on oceanic islands, energetic costs of flight and use of forelimbs for activities other than flying. Few studies have sought to explore factors common to all families containing flightless species, which may explain the taxonomic distribution of flightlessness. In this study, we found that for all eight avian families which contain both flightless and flighted species, the flighted species have shorter wing lengths relative to body mass than their sister families. This result is not biased by taxon size. Models of avian aerodynamics predict that birds with relatively short wings pay a high energetic cost of flight. We suggest that these increased energetic costs of flying predispose these avian families to evolve flightless species. The various causes for the shortening of wings among flighted species of birds and the possibility of future transitions to flightlessness are discussed.     

Mechanics of the limbs of the walrus (Odobenus rosmarus) and the california sea lion (Zalophus californianus)

Moment arms and cross-sectional areas of muscles from the forelimbs and hind limbs of Odobenus and Zalophus were used to estimate relative torques associated with several locomotor movements. Odobenus uses predominantly the hind limbs for aquatic propulsion, while Zalophus only uses the forelimbs. Torques contributing to locomotor movements used during the power and recovery strokes were substantially greater for the hind limbs of Odobenus and for the forelimbs of Zalophus. Apparently, the lineages of these two genera shared an aquatic ancestor during the Miocene; therefore, they also shared a common method of aquatic propulsion. This common method of aquatic propulsion could have been either forelimb, hind limb, or both. Since the Miocene then, the method of aquatic propulsion and the mechanical characteristics of the limbs have diverged for at least one of these two lineages. A model is advanced to explain key mechanical differences between Odobenus and Zalophus that enable Odobenus to maneuver effectively in a head-down posture on the ocean floor.     

Flow Visualization of Rhinoceros Beetle (Trypoxylus dichotomus) in Free Flight

Aerodynamic characteristics of the beetle,Trypoxylus dichotomus,which has a pair of elytra (forewings) and flexible hind wings,are investigated.Visualization experiments were conducted for various flight conditions of a beetle,Trypoxylus dichotomus:free,tethered,hovering,forward and climbing flights.Leading edge,trailing edge and tip vortices on both wings were observed clearly.The leading edge vortex was stable and remained on the top surface of the elytron for a wide interval during the downstroke of free forward flight.Hence,the elytron may have a considerable role in lift force generation of the beetle.In addition,we reveal a suction phenomenon between the gaps of the hind wing and the elytron in upstroke that may improve the positive lift force on the hind wing.We also found the reverse clap-fling mechanism of the T.dichotomus beetle in hovering flight.The hind wings touch together at the beginning of the upstroke.The vortex generation,shedding and interaction give a better understanding of the detailed aerodynamic mechanism of beetle flight.     

The effects of latitude,body size,and sexual selection on wing shape in a damselfly

Under natural selection, wing shape is expected to evolve to optimize flight performance. However, other selective factors besides flight performance may influence wing shape. One such factor could be sexual selection in wing sexual ornaments, which may lead to alternative variations in wing shape that are not necessarily related to flight performance. In the present study, we investigated wing shape variations in a calopterygid damselfly along a latitudinal gradient using geometric morphometrics. Both sexes show wing pigmentation, which is a known signal trait at intra‐ and interspecific levels. Wing shape differed between sexes and, within the same sex, the shape of the hind wing differed from the front wing. Latitude and body size explained a high percentage of the variation in wing shape for female front and hind wings, and male front wings. In male hind wings, wing pigmentation explained a high amount of the variation in wing shape. On the other hand, the variation in shape explained by pigmentation was very low in females. We suggest that the conservative morphology of front wings is maintained by natural selection operating on flight performance, whereas the sex‐specific differences in hind wings most likely could be explained by sexual selection. The observed sexual dimorphism in wing shape is likely a result of different sex‐specific behaviours. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 263–274.     

Understanding hind limb weight support in chimpanzees with implications for the evolution of primate locomotion

Most quadrupedal mammals support a larger amount of body weight on their forelimbs compared with their hind limbs during locomotion, whereas most primates support more of their body weight on their hind limbs. Increased hind limb weight support is generally interpreted as an adaptation that reduces stress on primates' highly mobile forelimb joints. Thus, increased hind limb weight support was likely vital for the evolution of primate arboreality. Despite its evolutionary importance, the mechanism used by primates to achieve this important kinetic pattern remains unclear. Here, we examine weight support patterns in a sample of chimpanzees (Pan troglodytes) to test the hypothesis that limb position, combined with whole body center of mass position (COM), explains increased hind limb weight support in this taxon. Chimpanzees have a COM midway between their shoulders and hips and walk with a relatively protracted hind limb and a relatively vertical forelimb, averaged over a step. Thus, the limb kinematics of chimpanzees brings their feet closer to the COM than their hands, generating greater hind limb weight support. Comparative data suggest that these same factors likely explain weight support patterns for a broader sample of primates. It remains unclear whether primates use these limb kinematics to increase hind limb weight support, or whether they are byproducts of other gait characteristics. The latter hypothesis raises the intriguing possibility that primate weight support patterns actually evolved as byproducts of other traits, or spandrels, rather than as adaptations to increase forelimb mobility. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Ontogeny of limb force distribution in squirrel monkeys (Saimiri boliviensis): insights into the mechanical bases of primate hind limb dominance

The distribution of peak vertical forces between the forelimbs and the hind limbs is one of the key traits distinguishing primate quadrupedal locomotion from that of other mammals. Whereas most mammals generate greater peak vertical forelimb forces, primates generate greater peak vertical hind limb forces. At the ultimate level, hind limb dominance in limb force distribution is typically interpreted as an adaptation to facilitate fine-branch arboreality. However, the proximate biomechanical bases for primate limb force distribution remain controversial. Three models have been previously proposed. The Center of Mass (COM) Position model attributes primates’ unique mode of limb loading to differences in the position of the whole-body COM relative to the hands and feet. The Active Weight Shift model asserts that primates actively redistribute body weight to their hind limbs by pitching the trunk up via the activation of hind limb retractor muscles. Finally, the Limb Compliance model argues that primates selectively mitigate forelimb forces by maintaining a compliant forelimb and a flat shoulder trajectory. Here, a detailed dataset of ontogenetic changes in morphology and locomotor mechanics in Bolivian squirrel monkeys (Saimiri boliviensis) was employed as a model system to evaluate each of these proposed models in turn. Over the first 10 months of life, squirrel monkeys transitioned from forelimb dominant infants to hind limb dominant juveniles, a change that was precipitated by decreases in peak vertical forelimb forces and increases in peak vertical hind limb forces. Results provided some support for all three of the models, although the COM Position and Active Weight Shift models were most strongly supported by the data. Overall, this study suggests that primates may use a variety of biomechanical strategies to achieve hind limb dominance in limb force distribution.     

Convergent morphological responses to loss of flight in rails (Aves: Rallidae)

The physiological demands of flight exert strong selection pressure on avian morphology and so it is to be expected that the evolutionary loss of flight capacity would involve profound changes in traits. Here, we investigate morphological consequences of flightlessness in a bird family where the condition has evolved repeatedly. The Rallidae include more than 130 recognized species of which over 30 are flightless. Morphological and molecular phylogenetic data were used here to compare species with and without the ability to fly in order to determine major phenotypic effects of the transition from flighted to flightless. We find statistical support for similar morphological response among unrelated flightless lineages, characterized by a shift in energy allocation from the forelimbs to the hindlimbs. Indeed, flightless birds exhibit smaller sterna and wings than flighted taxa in the same family along with wider pelves and more robust femora. Phylogenetic signal tests demonstrate that those differences are independent of phylogeny and instead demonstrate convergent morphological adaptation associated with a walking ecology. We found too that morphological variation was greater among flightless rails than flighted ones, suggesting that relaxation of physiological demands during the transition to flightlessness frees morphological traits to evolve in response to more varied ecological opportunities.     

Analysis of Maneuvering Flight of an Insect

Wing motion of a dragonfly in the maneuvering flight, which was measured by Wang et al. was investigated. Equations of motion for a maneuvering flight of an insect were derived. These equations were applied for analyzing the maneuvering flight. Inertial forces and moments acting on a body and wings were estimated by using these equations and the measured motions of the body and the wings. The results indicated the following characteristics of this flight: ( 1 ) The phase difference in flapping motion between the two fore wings and two hind wings, and the phase difference between the flapping motion and the feathering motion of the four wings are equal to those in a steady forward flight with the maximum efficiency. (2)The camber change and the feathering motion were mainly controlled by muscles at the wing bases.     

NOTES ON THE HABITS AND STRUCTURE OF THE GUACHARO STEATORNIS CARIPENSIS

1. Steatornis is peculiar, perhaps unique, among birds in possessing two specialized means, one auditory and the other tactile, of locating objects in total darkness.
2. Owing to the peculiar position of the hallux and the fact that the much-abbreviated metatarsus is more or less on the same plane as the toes, the hind limbs are ill-adapted for walking or for perching on a branch, but they afford the bird a secure foothold when resting longitudinally on a ledge.
3. In the embryo the abnormally short metatarsus is incorporated in the body of the foot. Although anatomically distinct in the adult, functionally it remains a part of the foot serving to support much of the bird's weight when resting in its normal crouching posture.     

A wing-assisted running robot and implications for avian flight evolution

DASH+Wings is a small hexapedal winged robot that uses flapping wings to increase its locomotion capabilities. To examine the effects of flapping wings, multiple experimental controls for the same locomotor platform are provided by wing removal, by the use of inertially similar lateral spars, and by passive rather than actively flapping wings. We used accelerometers and high-speed cameras to measure the performance of this hybrid robot in both horizontal running and while ascending inclines. To examine consequences of wing flapping for aerial performance, we measured lift and drag forces on the robot at constant airspeeds and body orientations in a wind tunnel; we also determined equilibrium glide performance in free flight. The addition of flapping wings increased the maximum horizontal running speed from 0.68 to 1.29 m s?1, and also increased the maximum incline angle of ascent from 5.6° to 16.9°. Free flight measurements show a decrease of 10.3° in equilibrium glide slope between the flapping and gliding robot. In air, flapping improved the mean lift:drag ratio of the robot compared to gliding at all measured body orientations and airspeeds. Low-amplitude wing flapping thus provides advantages in both cursorial and aerial locomotion. We note that current support for the diverse theories of avian flight origins derive from limited fossil evidence, the adult behavior of extant flying birds, and developmental stages of already volant taxa. By contrast, addition of wings to a cursorial robot allows direct evaluation of the consequences of wing flapping for locomotor performance in both running and flying.     

Insufficient visual information leads to spontaneous bipedal walking in Japanese monkeys

Japanese monkeys walked spontaneously on their hind limbs, when their vision was impaired either by narrowing the visual field or by reducing the incoming light. These variables were manipulated via goggles with translucent pipes and neutral density filters. The bipedal locomotion was observed more frequently as the impairment of the incoming visual information increased. It is very likely that facultative bipeds walk on their hind limbs when they feel the need to “free” their forelimbs to grope their way.     

How Could Beetle＇s Elytra Support Their Own Weight during Forward Flight？

The aerodynamic role of the elytra during a beetle＇s flapping motion is not well-elucidated, although it is well-recognized that the evolution of elytra has been a key in the success of coleopteran insects due to their protective function. An experimental study on wing kinematics reveals that for almost concurrent flapping with the hind wings, the flapping angle of the elytra is 5 times smaller than that of the hind wings. Then, we explore the aerodynamic forces on elytra in free forward flight with and without an effect of elytron-hind wing interaction by three-dimensional numerical simulation. The numerical results show that vertical force generated by the elytra without interaction is not sufficient to support even its own weight. However, the elytron-hind wing interaction improves the vertical force on the elytra up to 80%; thus, the total vertical force could fully support its own weight. The interaction slightly increases the vertical force on the hind wind by 6% as well.