Wing-assisted Incline Running Research Articles

Mind the ramp: Association between early life ramp use and spatial cognition in laying hen pullets.

Ramps facilitate earlier access to complex environments and increase early life voluntary exercise, which may positively affect the cognitive development of chickens. This study focused on quantifying individual differences in ramp use and its impact on spatial cognition of laying hen pullets. Sixteen identical pens were housed with Lohmann Selected Leghorn (LSL) chicks of which eight chicks from each pen were colour marked from one day of age (DoA) to serve as focal birds. We quantified overall ramp use (walk/run, wing-assisted incline running, and jump/fly to and from ramps) by scan sampling recorded videos for 6, 10, 12, 20, 27, 41, and 55 DoA for all focal birds. From 56 to 95 DoA, long and short-term spatial memory of three focal birds per pen were assessed in a holeboard test in three consecutive phases: cued, uncued and reversal. Mixed model analysis showed that the spatial cognitive abilities of the birds were linked to differences in ramp use frequency averaged across all observation days. Birds with higher ramp use made fewer reference (Estimate ± Confidence Interval = 0.94 [0.88, 0.99], p = 0.08) and working memory errors (Est ± CI = 0.77 [0.59, 1.00], p = 0.06) in the cued phase than birds with lower ramp use. In contrast, birds with higher ramp use made more reference memory errors (Est ± CI = 1.10 [1.01, 1.20], p = 0.05) in the reversal phase. Birds with higher ramp use also made more reference memory errors compared to birds with lower ramp use as the phases changed from cued to uncued (p = 0.001). Our results indicate that there might be a relationship between early life ramp use and spatial cognition of laying hens.

A wing-assisted incline running exercise regime during rearing increases initial flight velocity during descent in adult white- and brown-feathered laying hens

Domestic laying hens rely primarily on their hindlimbs for terrestrial locomotion. Although they perform flapping flight, they appear to use maximal power during descent and thus may lack control for maneuvering and avoiding injuries on landing. This in turn may result in injury in open rearing systems. Wing-assisted incline running (WAIR) requires a bird to use its wings to assist the hindlimbs during climbing of an incline, and training in WAIR may therefore provide a useful method to increase a hen's power reserve and control for flight. We subjected hens to an exercise regimen involving inclines to induce WAIR for 16 wk during rearing. We then measured wing and body kinematics during aerial descent from a 155 cm platform. We hypothesized that birds reared with exercise would be better able to modulate their wing and body kinematics for making slower, more-controlled descent and landing. Brown-feathered birds exhibited greater wing beat frequencies than white-feathered birds, which is consistent with the higher wing loading of brown-feathered birds and WAIR-trained birds exhibited greater initial flight velocities compared to control birds. This may indicate that WAIR training provided an improved capacity to modulate flight velocity and strengthen the leg muscles. Providing incline exercises during rearing may therefore improve welfare for adult laying hens as greater initial flight velocity should reduce the power required for supporting body weight in the air and allow a hen to direct her excess power toward maneuvering.

Multi-Modal Mobility Morphobot (M4) with appendage repurposing for locomotion plasticity enhancement

Robot designs can take many inspirations from nature, where there are many examples of highly resilient and fault-tolerant locomotion strategies to navigate complex terrains by recruiting multi-functional appendages. For example, birds such as Chukars and Hoatzins can repurpose wings for quadrupedal walking and wing-assisted incline running. These animals showcase impressive dexterity in employing the same appendages in different ways and generating multiple modes of locomotion, resulting in highly plastic locomotion traits which enable them to interact and navigate various environments and expand their habitat range. The robotic biomimicry of animals’ appendage repurposing can yield mobile robots with unparalleled capabilities. Taking inspiration from animals, we have designed a robot capable of negotiating unstructured, multi-substrate environments, including land and air, by employing its components in different ways as wheels, thrusters, and legs. This robot is called the Multi-Modal Mobility Morphobot, or M4 in short. M4 can employ its multi-functional components composed of several actuator types to (1) fly, (2) roll, (3) crawl, (4) crouch, (5) balance, (6) tumble, (7) scout, and (8) loco-manipulate. M4 can traverse steep slopes of up to 45 deg. and rough terrains with large obstacles when in balancing mode. M4 possesses onboard computers and sensors and can autonomously employ its modes to negotiate an unstructured environment. We present the design of M4 and several experiments showcasing its multi-modal capabilities.

Coracoid strength as an indicator of wing-beat propulsion in birds.

Birds generate a propulsive force by flapping their wings. They use this propulsive force for various locomotion styles, such as aerodynamic flight, wing-paddle swimming and wing-assisted incline running. It is therefore important to reveal the origin of flapping ability in the evolution from theropod dinosaurs to birds. However, there are no quantitative indices to reconstruct the flapping abilities of extinct forms based on their skeletal morphology. This study compares the section modulus of the coracoid relative to body mass among various extant birds to test whether the index is correlated with flapping ability. According to a survey of 220 historical bird specimens representing 209 species, 180 genera, 83 families and 30 orders, the section modulus of the coracoid relative to body mass in non-flapping birds was significantly smaller than that of flapping birds. This indicates that coracoid strength in non-flapping birds is deemphasised, whereas in flapping birds the strength is emphasisedto withstand the contractile force produced by powerful flapping muscles, such as the m. pectoralis and m. supracoracoideus. Therefore, the section modulus of the coracoid is expected to be a powerful tool to reveal the origin of powered flight in birds.

Apparent wing-assisted incline running in a Common Grackle (<i>Quiscalus quiscula</i>)

Wing-assisted incline running (WAIR) has been observed in bird taxa from multiple clades. Its wide phylogenetic distribution in modern birds suggests that it is an ancestral trait for class Aves. WAIR as a behaviour is speculated to predate the evolution of full-powered flight, and to have formed a behavioural and physiological stepping stone between terrestrial and aerial life. Here I report an observation of apparent WAIR in a Common Grackle (Quiscalus quiscula) photographed incidentally on a trail camera deployed in an urban backyard in Guelph, Ontario, Canada. To my knowledge this is the first documented observation of apparent WAIR for the family Icteridae. Furthermore, it highlights the value of non-systematic use of trail cameras for making unique natural history observations.

Where is WAIR (and other wing-assisted behaviours)? Essentially everywhere: a response to Kuznetsov and Panyutina (2022)

Abstract Kuznetsov and Panyutina (2022) offer a reanalysis of the kinematic and force plate data previously published by Bundle and Dial (2003). Their intention is to describe instantaneous wing forces during wing-assisted incline running (WAIR), focusing particularly on the upstroke phase. Based on their interpretation of wing forces and muscle function, the authors conclude that ‘WAIR is a very specialized mode of locomotion that is employed by a few specialized birds as an adaptation to a very specific environment and involving highly developed flying features of the locomotor apparatus’, and thus not relevant to the evolution of avian flight. Herein, we respond to the authors’ interpretations, offering an alternative perspective on WAIR and, more generally, on studies exploring the evolution of avian flight.

Where was WAIR in avian flight evolution?

Abstract The popular hypothesis that the flapping flight of birds evolved through the stage of wing-assisted incline running (WAIR) is tested through calculations of external forces acting on the wings in the downstroke and upstroke during WAIR activity of an adult chukar partridge (Alectoris chukar). Contrary to previous studies of WAIR, it is found that the total vertical impulse provided by the wings is negative (i.e. downward directed). Thus, the wings cancel out about half of the positive (upward-directed) impulse provided by the legs. The function of the wings in WAIR is not to lift the body, but to push it towards the steep slope. It is like a wing, or aerofoil, of a race car, which is tilted at a negative angle of attack to press the car to the ground for greater friction and adhesion of the wheels and to prevent the race car from becoming airborne. The calculated force pattern suggests that the dominant external forces acting on the wings in the sagittal plane during WAIR are not the virtual forces of inertia but the aerodynamic forces. More specifically, it is only the aerodynamic lift that can be responsible for the wing force component acting at right angles to the wingbeat plane in both the downstroke and the upstroke during WAIR. The lift towards the slope during the WAIR upstroke is ensured by separation of the primary flight feathers and their setting at a negative angle of attack. Analysis of the force pattern in the anatomical coordinate space of a bird’s body shows that the main wing muscles in the WAIR downstroke are the humeral retractor muscles and in the WAIR upstroke the humeral elevator muscles. The humeral protractor muscles are not required at all during WAIR. Therefore, the morphologically necessary protractor state of the supracoracoideus muscle in a ‘semi-flying’ ancestor, which is logically explained by the classical hypothesis of a gliding ancestor, does not fit the WAIR hypothesis of flapping flight origin. Wing-assisted incline running should be regarded as a crown locomotor specialization of birds and is not an appropriate model for locomotion in avian ancestors.

Aerodynamics Show Membranous-Winged Theropods Were a Poor Gliding Dead-End

The bizarre Scansoriopterygidae theropods Yi and Ambopteryx are the only dinosaurs thought to have had skin stretched between elongate fingers that form a potential membranous wing. While these structures have been suggested to have been used in aerial locomotion, this has never been shown to be the case. Demonstrating this function is of great importance because if these wings permitted flight it would support the growing view that theropod dinosaurs evolved flight multiple times in distinctive ways before the origin of modern birds. Here using Laser-Stimulated Fluorescence imaging we re-evaluate the anatomy of Yi and integrate this new information into several different equation-based aerodynamic calculations covering flight potential, methods of take-off (terrestrial-based running, arboreal launch), and powered versus gliding flight capabilities. For the first time we test whether Yi and Ambopteryx had flight capabilities and if so how efficient and effective it was. We also evaluate wing-based behaviours including wing-assisted incline running (WAIR). We find that Yi and Ambopteryx were highly unlikely to have any form of powered flight and had limited gliding abilities compared to similar-sized living and fossil gliders. We find that both taxa could not take-off from a terrestrial setting, supporting the idea that they were likely arboreal. Compared to similar-sized early-diverging fossil paravians, like Archaeopteryx and Microraptor, Scansoriopterygidae show significant deficiencies and differences in flapping-based locomotion models that indicate a much less competent flight stroke with limited aerial abilities. Combining these findings, our results show that the Scansoriopterygidae are not models for the early evolution of bird flight and that their structurally distinct and inefficient wings differed greatly from contemporaneous paravians including the earliest birds. Instead, we propose that Scansoriopterygidae represent a unique but failed flight architecture of non-avialan theropods. These findings conform with emerging data supporting multiple independent origins of flight in modern lineages. They show that the battle to capture the vertebrate aerial morphospace in the Middle to Late Jurassic was a dynamic and complex one not restricted to pterosaurs and only one theropod lineage.

Building a Bird: Musculoskeletal Modeling and Simulation of Wing-Assisted Incline Running During Avian Ontogeny

Flapping flight is the most power-demanding mode of locomotion, associated with a suite of anatomical specializations in extant adult birds. In contrast, many developing birds use their forelimbs to negotiate environments long before acquiring “flight adaptations,” recruiting their developing wings to continuously enhance leg performance and, in some cases, fly. How does anatomical development influence these locomotor behaviors? Isolating morphological contributions to wing performance is extremely challenging using purely empirical approaches. However, musculoskeletal modeling and simulation techniques can incorporate empirical data to explicitly examine the functional consequences of changing morphology by manipulating anatomical parameters individually and estimating their effects on locomotion. To assess how ontogenetic changes in anatomy affect locomotor capacity, we combined existing empirical data on muscle morphology, skeletal kinematics, and aerodynamic force production with advanced biomechanical modeling and simulation techniques to analyze the ontogeny of pectoral limb function in a precocial ground bird (Alectoris chukar). Simulations of wing-assisted incline running (WAIR) using these newly developed musculoskeletal models collectively suggest that immature birds have excess muscle capacity and are limited more by feather morphology, possibly because feathers grow more quickly and have a different style of growth than bones and muscles. These results provide critical information about the ontogeny and evolution of avian locomotion by (i) establishing how muscular and aerodynamic forces interface with the skeletal system to generate movement in morphing juvenile birds, and (ii) providing a benchmark to inform biomechanical modeling and simulation of other locomotor behaviors, both across extant species and among extinct theropod dinosaurs.

Development of locomotion over inclined surfaces in laying hens

The purpose of the present study was to evaluate locomotor strategies during development in domestic chickens (Gallus gallus domesticus); we were motivated, in part, by current efforts to improve the design of housing systems for laying hens which aim to reduce injury and over-exertion. Using four strains of laying hens (Lohmann Brown, Lohmann LSL lite, Dekalb White and Hyline Brown) throughout this longitudinal study, we investigated their locomotor style and climbing capacity in relation to the degree (0 to 70°) of incline, age (2 to 36 weeks) and the surface substrate (sandpaper or wire grid). Chicks and adult fowl performed only walking behavior to climb inclines ⩽40° and performed a combination of wing-assisted incline running (WAIR) or aerial ascent on steeper inclines. Fewer birds used their wings to aid their hind limbs when climbing 50° inclines on wire grid surface compared with sandpaper. The steepness of angle achieved during WAIR and the tendency to fly instead of using WAIR increased with increasing age and experience. White-feathered strains performed more wing-associated locomotor behavior compared with brown-feathered strains. A subset of birds was never able to climb incline angles >40° even when using WAIR. Therefore, we suggest that inclines of up to 40° should be provided for hens in three-dimensional housing systems, which are easily negotiated (without wing use) by chicks and adult fowl.

Ontogeny of Flight Capacity and Pectoralis Function in a Precocial Ground Bird (Alectoris chukar)

Flight is the defining characteristic of birds, yet the mechanisms through which flight ability develops are only beginning to be understood. Wing-assisted incline running (WAIR) and controlled flapping descent (CFD) are behaviors that may offer significant adaptive benefits to developing birds. Recent research into these forms of locomotion has focused on species with precocial development, with a particularly rich data set from chukar partridge (Alectoris chukar). Here we briefly review the kinematics and aerodynamics of flight development in this species. We then present novel measurements of the development of pectoralis contractile behavior during the ontogenetic transition toward powered flight. To obtain these new empirical data, we used indwelling electromyography (EMG) and sonomicrometry and tested WAIR and CFD in seven age classes of chukar (n = 2-4 birds per age) from 5 days post hatching (dph) to adult (300+ dph). For each age class, we measured muscle activity during maximal performance, which was WAIR at 65° in birds 5 dph, CFD in birds 9 dph, WAIR at 80° in birds 14 dph, level flight in birds 25-61 dph, and ascending flight in adults. We also measured muscle activity during sub-maximal performance in all age classes. Flapping chukar chicks use near-continuous activation of their pectoralis at relatively low electromyography amplitudes for the first 8 days and progress to stereotypic higher-amplitude activation bursts by Day 12. The pectoralis undergoes increasing strain at higher strain rates with age, and length trajectory becomes more asymmetrical with greater variation in contractile velocity within the shortening phase of individual contractions. At 20-25 days (12-15% adult chukar mass), pectoralis activity and locomotor performance approaches that of adults, although strain rate exhibits a temporary decrease at 61 dph concurrent with using newly-replaced primary feathers. To better understand how these patterns relate to the evolution of life-history strategy and locomotion, we encourage future efforts to explore these behaviors in altricial and semi-altricial bird species.

The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents.

Background: Powered flight is implicated as a major driver for the success of birds. Here we examine the effectiveness of three hypothesized pathways for the evolution of the flight stroke, the forelimb motion that powers aerial locomotion, in a terrestrial setting across a range of stem and basal avians: flap running, Wing Assisted Incline Running (WAIR), and wing-assisted leaping.Methods: Using biomechanical mathematical models based on known aerodynamic principals and in vivo experiments and ground truthed using extant avians we seek to test if an incipient flight stroke may have contributed sufficient force to permit flap running, WAIR, or leaping takeoff along the phylogenetic lineage from Coelurosauria to birds.Results: None of these behaviours were found to meet the biomechanical threshold requirements before Paraves. Neither was there a continuous trend of refinement for any of these biomechanical performances across phylogeny nor a signal of universal applicability near the origin of birds. None of these flap-based locomotory models appear to have been a major influence on pre-flight character acquisition such as pennaceous feathers, suggesting non-locomotory behaviours, and less stringent locomotory behaviours such as balancing and braking, played a role in the evolution of the maniraptoran wing and nascent flight stroke. We find no support for widespread prevalence of WAIR in non-avian theropods, but can’t reject its presence in large winged, small-bodied taxa like Microraptor and Archaeopteryx.Discussion: Using our first principles approach we find that “near flight” locomotor behaviors are most sensitive to wing area, and that non-locomotory related selection regimes likely expanded wing area well before WAIR and other such behaviors were possible in derived avians. These results suggest that investigations of the drivers for wing expansion and feather elongation in theropods need not be intrinsically linked to locomotory adaptations, and this separation is critical for our understanding of the origin of powered flight and avian evolution.

New Perspectives on the Ontogeny and Evolution of Avian Locomotion.

Close correspondence between form and function is a central tenet of natural selection. One of the most striking, textbook cases for form-function congruence is the evolution of flight and the body plan of birds: compared with other tetrapods, extant adult birds have highly modified integuments and skeletons, and it has traditionally been assumed that many of these modifications are adaptations or exaptations for flight. However, developing birds that lack many of the morphological signatures of flight capacity nevertheless use their developing wings for a variety of flapping behaviors, such as wing-assisted incline running and even brief flight. Immature birds thereby demonstrate that rudimentary "flight" apparatuses are more functional than traditional assumptions about form-function relationships would predict. Here, I review the ontogeny of avian locomotion, highlighting how the developmental acquisition of flight in extant birds can improve our understanding of form-function relationships in the avian body plan, and provide insight into the evolutionary origin of flight among extinct non-avian theropod dinosaurs.

Ontogeny of aerial righting and wing flapping in juvenile birds

Mechanisms of aerial righting in juvenile chukar partridge (Alectoris chukar) were studied from hatching to 14 days-post-hatching (dph). Asymmetric movements of the wings were used from 1 to 8 dph to effect progressively more successful righting behaviour via body roll. Following 8 dph, wing motions transitioned to bilaterally symmetric flapping that yielded aerial righting via nose-down pitch, along with substantial increases in vertical force production during descent. Ontogenetically, the use of such wing motions to effect aerial righting precedes both symmetric flapping and a previously documented behaviour in chukar (i.e. wing-assisted incline running) hypothesized to be relevant to incipient flight evolution in birds. These findings highlight the importance of asymmetric wing activation and controlled aerial manoeuvres during bird development and are potentially relevant to understanding the origins of avian flight.

Creating testing regimes for terrestrial origins of the avian flight stroke (919.8)

While the ancestry of birds has been firmly established within maniraptoran theropod dinosaurs, the exact timing of the origins of the flight stroke remains elusive. Debate rages as to the drivers of this critical step in the evolution of powered flight, but little biomechanical modeling has been done to test the various hypothesis proposed. Here we seek to test three proposed pathways for a terrestrial origin of flight: flap running, Wing Assisted incline Running (W.A.I.R.) and burst leaping, to examine where, if at all, on the non‐avian to avian theropod tree the aerodynamic forces required by each are possible. Based on figures generated for in‐vivo experimental data from the literature we find that W.A.I.R. is restricted, conservatively, to the sister clade to birds while flap running is biomechanically unlikely. These results will help to shape future discussion on the ecology and drivers of the origin of flight by placing theses behavioral possibilities in their evolutionary context.

From baby birds to feathered dinosaurs: incipient wings and the evolution of flight

Reconstructing the tree of life requires deciphering major evolutionary transformations and the functional capacities of fossils with “transitional” morphologies. Some of the most iconic, well-studied fossils with transitional features are theropod dinosaurs, whose skeletons and feathered forelimbs record the origin and evolution of bird flight. However, in spite of over a century of discussion, the functions of forelimb feathers during the evolution of flight remain enigmatic. Both aerodynamic and non-aerodynamic roles have been proposed, but few of the form-function relationships assumed by these scenarios have been tested. Here, we use the developing wings of a typical extant ground bird (Chukar Partridge) as possible analogues/homologues of historical wing forms to provide the first empirical evaluation of aerodynamic potential in flapping theropod “protowings.” Immature ground birds with underdeveloped, rudimentary wings generate useful aerodynamic forces for a variety of locomotor tasks. Feather development in these birds resembles feather evolution in theropod dinosaurs, and reveals a predictable relationship between wing morphology and aerodynamic performance that can be used to infer performance in extinct theropods. By spinning an ontogenetic series of spread-wing preparations on a rotating propeller apparatus across a range of flow conditions and measuring aerodynamic force, we explored how changes in wing size, feather structure, and angular velocity might have affected aerodynamic performance in dinosaurs choosing to flap their incipient wings. At slow angular velocities, wings produced aerodynamic forces similar in magnitude to those produced by immature birds during behaviors like wing-assisted incline running. At fast angular velocities, wings produced forces sufficient to support body weight during flight. These findings provide a quantitative, biologically relevant bracket for theropod performance and suggest that protowings could have provided useful aerodynamic function early in maniraptoran history, with improvements in aerodynamic performance attending the evolution of larger wings, more effective feather morphologies, and faster angular velocities.

Three-dimensional, high-resolution skeletal kinematics of the avian wing and shoulder during ascending flapping flight and uphill flap-running.

Past studies have shown that birds use their wings not only for flight, but also when ascending steep inclines. Uphill flap-running or wing-assisted incline running (WAIR) is used by both flight-incapable fledglings and flight-capable adults to retreat to an elevated refuge. Despite the broadly varying direction of travel during WAIR, level, and descending flight, recent studies have found that the basic wing path remains relatively invariant with reference to gravity. If so, joints undergo disparate motions to maintain a consistent wing path during those specific flapping modes. The underlying skeletal motions, however, are masked by feathers and skin. To improve our understanding of the form-functional relationship of the skeletal apparatus and joint morphology with a corresponding locomotor behavior, we used XROMM (X-ray Reconstruction of Moving Morphology) to quantify 3-D skeletal kinematics in chukars (Alectoris chukar) during WAIR (ascending with legs and wings) and ascending flight (AF, ascending with wings only) along comparable trajectories. Evidence here from the wing joints demonstrates that the glenohumeral joint controls the vast majority of wing movements. More distal joints are primarily involved in modifying wing shape. All bones are in relatively similar orientations at the top of upstroke during both behaviors, but then diverge through downstroke. Total excursion of the wing is much smaller during WAIR and the tip of the manus follows a more vertical path. The WAIR stroke appears “truncated” relative to ascending flight, primarily stemming from ca. 50% reduction in humeral depression. Additionally, the elbow and wrist exhibit reduced ranges of angular excursions during WAIR. The glenohumeral joint moves in a pattern congruent with being constrained by the acrocoracohumeral ligament. Finally, we found pronounced lateral bending of the furcula during the wingbeat cycle during ascending flight only, though the phasic pattern in chukars is opposite of that observed in starlings (Sturnus vulgaris).

Factors shaping the ontogeny of vocal signals in a wild parrot

Parrots rely heavily on vocal signals to maintain their social and mobile lifestyles. We studied vocal ontogeny in nests of wild green-rumped parrotlets (Forpus passerinus) in Venezuela. We identified three successive phases of vocal signaling that corresponded closely to three independently derived phases of physiological development. For each ontogenetic phase, we characterized the relative importance of anatomical constraints, motor skills necessary for responding to specific contexts of the immediate environment, and the learning of signals that are necessary for adult forms of communication. We observed shifts in the relative importance of these three factors as individuals progressed from one stage to the next; there was no single fixed ratio of factors that applied across the entire ontogenetic sequence. The earliest vocalizations were short in duration, as predicted from physical constraints and under-developed motor control. Calls became longer and frequency modulated during intermediate nestling ages in line with motor skills required for competitive begging. In the week before fledging, calls drastically shortened in accordance with the flight-constrained short durations of adult contact calls. The latter constraints were made evident by the demonstrated links between wing-assisted incline running, a widespread prelude to avian flight, just before the shift from long-duration begging calls to short-duration contact calls. At least in this species, the shifting emphases of factors at different ontogenetic stages precluded the morphing of intermediate-stage begging calls into adult contact calls; as shown previously, the latter are influenced by sample templates provided by parents.

When hatchlings outperform adults: locomotor development in Australian brush turkeys (Alectura lathami, Galliformes)

Within Galliformes, megapods (brush turkey, malleefowl, scrubfowl) exhibit unique forms of parental care and growth. Hatchlings receive no post-hatching parental care and exhibit the most exaggerated precocial development of all extant birds, hatching with fully developed, flight-capable forelimbs. Rather than flying up to safety, young birds preferentially employ wing-assisted incline running. Newly hatched Australian brush turkeys (Alectura lathami) are extraordinarily proficient at negotiating all textured inclined surfaces and can flap-walk up inclines exceeding the vertical. Yet, as brush turkeys grow, their forelimb-dependent locomotor performance declines. In an attempt to elucidate how hatchlings perform so well, we analysed hindlimb forces and forelimb kinematics. We measured ground reaction forces (GRFs) for animals spanning the entire growth range (110-2000 g) as they ascended a variably positioned inclined ramp that housed a forceplate. These data are compared with a similar dataset for a chukar partridge (Alectoris chukar) that exhibit a growth strategy typical of most other Galliformes and that demonstrate improved incline performance with increasing age. The brush turkeys' ontogenetic decline in incline running performance is accompanied by loss of traction at steep angles, reduced GRFs and increased wing-loading. We hypothesize that Australian brush turkeys, in contrast to other Galliformes, develop from forelimb-dominated young that exploit a variable terrain (e.g. mound nests, boulders, embankments, cliffs, bushes and trees) into hindlimb-dominated adults dependent on size and running speed to avoid predation.

Precocial development of locomotor performance in a ground-dwelling bird (Alectoris chukar): negotiating a three-dimensional terrestrial environment

Developing animals are particularly vulnerable to predation. Hence, precocial young of many taxa develop predator escape performance that rivals that of adults. Ontogenetically unique among vertebrates, birds transition from hind limb to forelimb dependence for escape behaviours, so developmental investment for immediate gains in running performance may impair flight performance later. Here, in a three-dimensional kinematic study of developing birds performing pre-flight flapping locomotor behaviours, wing-assisted incline running (WAIR) and a newly described behaviour, controlled flapping descent (CFD), we define three stages of locomotor ontogeny in a model gallinaceous bird (Alectoris chukar). In stage I (1-7 days post-hatching (dph)) birds crawl quadrupedally during ascents, and their flapping fails to reduce their acceleration during aerial descents. Stage II (8-19 dph) birds use symmetric wing beats during WAIR, and in CFD significantly reduce acceleration while controlling body pitch to land on their feet. In stage III (20 dph to adults), birds are capable of vertical WAIR and level-powered flight. In contrast to altricial species, which first fly when nearly at adult mass, we show that in a precocial bird the major requirements for flight (i.e. high power output, wing control and wing size) convene by around 8 dph (at ca 5% of adult mass) and yield significant gains in escape performance: immature chukars can fly by 20 dph, at only about 12 per cent of adult mass.