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Migratory distances and stopover locations are changing for many passerines in response to climate change. Morphological changes have been linked to rising global temperatures in both migrants and residents, but the implications of these changes on fuel loads, and associated flight ranges are little studied. Wing length and body mass changes between 1964 and 2020 were calculated for 15 migrant and partially migrant passerines in Britain. Changes in fuel load and lean body mass were also estimated and used to predict flight ranges. Twelve of the species have undergone morphological changes and eight species, estimated fuel load changes. Nine species were estimated to have reduced flight ranges, indicating that the morphological changes have not compensated fully for the reduction in flight range experienced since 1964. Partial migrants showed greater decreases in flight ranges than did full migrants, which may indicate greater behavioural plasticity in the former. Those species which do not adapt morphologically or behaviourally may be unable to complete long migrations, resulting in restriction to sub-optimal breeding/wintering habitats, or a need for a sooner first stop and more stops en route. This highlights the importance of conserving migratory stopover sites, particularly in the Mediterranean and North Africa that immediately precede major geographical barriers, as-well-as breeding and wintering grounds., (© 2024. The Author(s).)

The European swallowtail butterfly ( Papilio machaon ) is so named, because of the long and narrow prominences extending from the trailing edge of their hindwings and, although not a true tail, they are referred to as such. Despite being a defining feature, an unequivocal function for the tails is yet to be determined, with predator avoidance (diverting an attack from the rest of the body), and enhancement of aerodynamic performance suggested. The swallowtail, however, is sexually size dimorphic with females larger than males, but whether the tail is also sexually dimorphic is unknown. Here, museum specimens were used to determine whether sexual selection has played a role in the evolution of the swallowtail butterfly tails in a similar way to that seen in the tail streamers of the barn swallow ( Hirundo rustica ), where the males have longer streamers than those of the females. Previously identified sexual dimorphism in swallowtail butterfly size was replicated, but no evidence for dimorphism in tail length was found. If evolved to mimic antennae and a head to divert a predatory attack, and if an absolute tail size was the most effective for this, then the tail would probably be invariant with butterfly hindwing size. The slope of the relationship between tail length and size, however, although close to zero, was nonetheless statistically significantly above (tail length ∝ hindwing area 0.107 ± 0.011 ). The slope also did not equate to that expected for geometric similarity (tail length ∝ hindwing area 1/2 ) suggesting that tail morphology is not solely driven by aerodynamics. It seems likely then, that tail morphology is primarily determined by, and perhaps a compromise of several, factors associated with predator avoidance (e.g. false head mimicry and a startling function). Of course, experimental data are required to confirm this., Competing Interests: The authors have no conflicts of interest to declare., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Climate change is predicted to impact freshwater aquatic environments through changes to water temperature ( T water ), river flow and eutrophication. Riverine habitats contain many economically and ecologically important fishes. One such group is the migratory salmonids, which are sensitive to warm T water and low O 2 (hypoxia). While several studies have investigated the independent effects of T water and hypoxia on fish physiology, the combined effects of these stressors is less well known. Furthermore, no study has investigated the effects of T water and O 2 saturation levels within the range currently experienced by a salmonid species. Thus, the aim of this study was to investigate the simultaneous effects of T water and O 2 saturation level on the energetics and kinematics of steady-state swimming in brown trout, Salmo trutta . No effect of O 2 saturation level (70 and 100% air saturation) on tail-beat kinematics was detected. Conversely, T water (10, 14, 18 and 22°C) did affect tail-beat kinematics, but a trade-off between frequency ( f tail ) and amplitude ( A , maximum tail excursion) maintained the Strouhal number (St =  f tail • A / U , where U is swimming speed) within the theoretically most mechanically efficient range. Swimming oxygen consumption rate ([Formula: see text]) and cost of transport increased with both U and T water . The only effect of O 2 saturation level was observed at the highest T water (22°C) and fastest swimming speed (two speeds were used-0.6 and 0.8 m s -1 ). As the extremes of this study are consistent with current summer conditions in parts of UK waterways, our findings may indicate that S. trutta will be negatively impacted by the increased T water and reduced O 2 levels likely presented by anthropogenic climate change., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Research into the terrestrial locomotion of birds is often based upon laboratory treadmill experiments. However, it is unclear how transposable these results are for birds moving in the wild. Here, using video recordings, we compared the kinematics of locomotion (stride frequency, stride length, stance phase, swing phase, duty factor) and speed range of Svalbard rock ptarmigan (Lagopus muta hyperborea) under field and laboratory treadmill conditions. Our findings indicate that the kinematics of walking and aerial running are conserved when moving on the treadmill and in the field. Differences, however, were found when grounded running under the two conditions, linked to substrate. Substrate effects were confirmed by analysing trials only moving over very hard snow. In line with laboratory treadmill energetic predictions, wild ptarmigan have a preferred speed during walking and to a lesser extent when aerial running but not when moving with a grounded running gait. The birds were also capable of a higher top speed in the field than that observed during treadmill studies. Our findings demonstrate that laboratory treadmill research provides meaningful information relevant to wild birds while highlighting the importance of understanding the substrate the animals are moving over.

Temperature is a ubiquitous environmental factor affecting physiological processes of ectotherms. Due to the effects of climate change on global air and water temperatures, predicting the impacts of changes in environmental thermal conditions on ecosystems is becoming increasingly important. This is especially crucial for migratory fish, such as the ecologically and economically vital salmonids, because their complex life histories make them particularly vulnerable. Here, we addressed the question whether temperature affects the morphology of brown trout, Salmo trutta L. spermatozoa. The fertilising ability of spermatozoa is commonly attributed to their morphological dimensions, thus implying direct impacts on the reproductive success of the male producing the cells. We show that absolute lengths of spermatozoa are not affected by temperature, but spermatozoa from warm acclimated S. trutta males have longer flagella relative to their head size compared to their cold acclimated counterparts. This did not directly affect sperm swimming speed, although spermatozoa from warm acclimated males may have experienced a hydrodynamic advantage at warmer temperatures, as suggested by our calculations of drag based on head size and sperm swimming speed. The results presented here highlight the importance of increasing our knowledge of the effects of temperature on all aspects of salmonid reproduction in order to secure their continued abundance., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Climate change alters the thermal habitat of aquatic species on a global scale, generating novel environmental challenges during all life stages, including reproduction. Changes in water temperature profoundly influence the performance of ectothermic aquatic organisms. This is an especially crucial issue for migratory fish, because they traverse multiple environments in order to reproduce. In externally fertilizing migratory fish, gametes are affected by water temperature indirectly, within the reproductive organ in which they are produced during migration, as well as directly, upon release into the surrounding medium at the spawning grounds. Both direct (after release) and indirect (during production) thermal impacts on gamete quality have been investigated, but never in conjunction. Here, we assessed the cumulative influence of temperature on brown trout, Salmo trutta , sperm quality during sperm production (male acclimation temperature) as well as upon release (sperm activation water temperature) on two consecutive dates during the brown trout spawning season. Early in the season, warm acclimation of males reduced their fertilization probability (lower sperm velocity) when compared with cold-acclimated males, especially when the activation water temperature was also increased beyond the thermal optimum (resulting in a lower proportion of motile sperm with lower velocity). Later in the season, sperm quality was unaffected by acclimation temperature and thermal sensitivity of sperm was reduced. These results give novel insights into the complex impacts of climate change on fish sperm, with implications for the reproduction and management of hatchery and wild trout populations in future climate scenarios., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

The diversity of wing morphologies in birds reflects their variety of flight styles and the associated aerodynamic and inertial requirements. Although the aerodynamics underlying wing morphology can be informed by aeronautical research, important differences exist between planes and birds. In particular, birds operate at lower, transitional Reynolds numbers than do most aircraft. To date, few quantitative studies have investigated the aerodynamic performance of avian wings as fixed lifting surfaces and none have focused upon the differences between wings from different flight style groups. Dried wings from 10 bird species representing three distinct flight style groups were mounted on a force/torque sensor within a wind tunnel in order to test the hypothesis that wing morphologies associated with different flight styles exhibit different aerodynamic properties. Morphological differences manifested primarily as differences in drag rather than lift. Maximum lift coefficients did not differ between groups, whereas minimum drag coefficients were lowest in undulating flyers (Corvids). The lift to drag ratios were lower than in conventional aerofoils and data from free-flying soaring species; particularly in high frequency, flapping flyers (Anseriformes), which do not rely heavily on glide performance. The results illustrate important aerodynamic differences between the wings of different flight style groups that cannot be explained solely by simple wing-shape measures. Taken at face value, the results also suggest that wing-shape is linked principally to changes in aerodynamic drag, but, of course, it is aerodynamics during flapping and not gliding that is likely to be the primary driver., Competing Interests: The authors declare there are no competing interests.

The differing limb dynamics and postures of small and large terrestrial animals may be mechanisms for minimising metabolic costs under scale-dependent muscle force, work and power demands; however, empirical evidence for this is lacking. Leghorn chickens (Gallus gallus domesticus) are highly dimorphic: males have greater body mass and relative muscle mass than females, which are permanently gravid and have greater relative intestinal mass. Furthermore, leghorns are selected for standard (large) and bantam (small) varieties and the former are sexually dimorphic in posture, with females having a more upright limb. Here, high-speed videography and morphological measurements were used to examine the walking gaits of leghorn chickens of the two varieties and sexes. Hindlimb skeletal elements were geometrically similar among the bird groups, yet the bird groups did not move with dynamic similarity. In agreement with the interspecific scaling of relative duty factor (DF, the proportion of a stride period with ground contact for any given foot) with body mass, bantams walked with greater DF than standards, and females walked with greater DF than males. Greater DF in females than in males was achieved via variety-specific kinematic mechanisms, associated with the presence/absence of postural dimorphism. Females may require greater DF in order to reduce peak muscle forces and minimise power demands associated with lower muscle to reproductive tissue mass ratios and smaller body size. Furthermore, a more upright posture observed in the standard, but not bantam, females, may relate to minimising the work demands of being larger and having proportionally larger reproductive tissue volume. Lower DF in males relative to females may also be a work-minimising strategy and/or due to greater limb inertia (as a result of greater pelvic limb muscle mass) prolonging the swing phase., (© 2016. Published by The Company of Biologists Ltd.)

Leghorn (layer) chickens (Gallus gallus domesticus) differ in locomotor morphology and performance due to artificial selection for standard (large) and bantam (small) varieties, sexual dimorphisms and ontogenetic stage. Here, the hind limb skeletal muscle architectural properties of mature and juvenile standard breeds and mature bantams are compared and linked to measures of locomotor performance. Mature males possessed greater relative muscle physiological cross-sectional areas (PCSAs) than their conspecific females, indicative of greater force-generating capacity, and in line with their greater maximum sustainable speeds compared with females. Furthermore, some of the relative fascicle lengths of the pennate muscles were greater in mature males than in mature females, which may permit greater muscle contractibility. Immature standard leghorns, however, did not share the same dimorphisms as their mature forms. The differences in architectural properties between immature and mature standard males indicate that with the onset of male sexual maturity, concomitant with increasing muscle mass in males, the relative fascicle lengths of pennate muscles and the relative PCSAs of the parallel-fibred muscles also increase. The age-related differences in standard breed male muscle architecture are linked to the presence and absence of sex differences in maximum aerobic speeds. Males of bantam and standard varieties shared similar muscle proportions (% body mass), but exhibited intrinsic muscle differences with a tendency for greater force-generating capabilities in bantams and greater contractile capabilities in standards. The metabolic costs associated with the longer fascicle lengths, together with more crouched limbs in standard than in bantam males may explain the lack of allometry in the minimum metabolic cost of transport between these birds of different size., (© 2016 Anatomical Society.)

In leghorn chickens (Gallus gallus domesticus) of standard breed (large) and bantam (small) varieties, artificial selection has led to females being permanently gravid and sexual selection has led to male-biased size dimorphism. Using respirometry, videography and morphological measurements, sex and variety differences in metabolic cost of locomotion, gait utilisation and maximum sustainable speed (Umax) were investigated during treadmill locomotion. Males were capable of greater Umax than females and used a grounded running gait at high speeds, which was only observed in a few bantam females and no standard breed females. Body mass accounted for variation in the incremental increase in metabolic power with speed between the varieties, but not the sexes. For the first time in an avian species, a greater mass-specific incremental cost of locomotion, and minimum measured cost of transport (CoTmin) were found in males than in females. Furthermore, in both varieties, the female CoTmin was lower than predicted from interspecific allometry. Even when compared at equivalent speeds (using Froude number), CoT decreased more rapidly in females than in males. These trends were common to both varieties despite a more upright limb in females than in males in the standard breed, and a lack of dimorphism in posture in the bantam variety. Females may possess compensatory adaptations for metabolic efficiency during gravidity (e.g. in muscle specialization/posture/kinematics). Furthermore, the elevated power at faster speeds in males may be linked to their muscle properties being suited to inter-male aggressive combat., (© 2015. Published by The Company of Biologists Ltd.)

The minimum metabolic cost of transport (CoTmin; J kg(-1) m(-1)) scales negatively with increasing body mass (∝Mb (-1/3)) across species from a wide range of taxa associated with marked differences in body plan. At the intraspecific level, or between closely related species, however, CoTmin does not always scale with Mb. Similarity in physiology, dynamics of movement, skeletal geometry and posture between closely related individuals is thought to be responsible for this phenomenon, despite the fact that energetic, kinematic and morphometric data are rarely collected together. We examined the relationship between these integrated components of locomotion in leghorn chickens (Gallus gallus domesticus) selectively bred for large and bantam (miniature) varieties. Interspecific allometry predicts a CoTmin ∼16% greater in bantams compared with the larger variety. However, despite 38% and 23% differences in Mb and leg length, respectively, the two varieties shared an identical walking CoTmin, independent of speed and equal to the allometric prediction derived from interspecific data for the larger variety. Furthermore, the two varieties moved with dynamic similarity and shared geometrically similar appendicular and axial skeletons. Hip height, however, did not scale geometrically and the smaller variety had more erect limbs, contrary to interspecific scaling trends. The lower than predicted CoTmin in bantams for their Mb was associated with both the more erect posture and a lower cost per stride (J kg(-1) stride(-1)). Therefore, our findings are consistent with the notion that a more erect limb is associated with a lower CoTmin and with the previous assumption that similarity in skeletal shape, inherently linked to walking dynamics, is associated with similarity in CoTmin., (© 2015. Published by The Company of Biologists Ltd.)

Take-off in bats is separated into two distinct phases: an initial jump and a subsequent wing powered acceleration. Here, using footage from a high-speed camera, the first comparative study of the performance during the wing induced phase of take-off in six insectivorous bat species is described. Despite distinct differences in foraging strategy, the mass specific power generated by the bats during wing induced take-off did not differ between species, with the exception of Myotis capaccinii. This suggests that differences in take-off performance may only be evident in bats that exhibit particularly unusual foraging strategies, such as the trawling behaviour of M. capaccinii - with differences in the remaining species only manifesting in subtler aspects of flight performance such as agility or manoeuvrability. The poorer take-off performance of M. capaccinii could be related to either a reduction in wing-stroke amplitude to stop the wings hitting the water's surface during foraging or perhaps an effect of having very large feet. No scaling relationship between body mass and mass-specific take-off power was found, which supports earlier research on birds and insects, suggesting that the mass-specific muscle power available for flight is broadly similar across a large range of body sizes and species., (© 2014. Published by The Company of Biologists Ltd.)

The relationship between tail (or wing) beat frequency (f(tail)), amplitude (A) and forward velocity (U) in animals using oscillatory propulsion, when moving at a constant cruising speed, converges upon an optimum range of the Strouhal number (St = f(tail) · A/U). Previous work, based on observational data and supported by theory, shows St falling within the broad optimum range (0.2

Fat storage is essential to the survival of many bird species, providing energy reserves, but can have an effect on locomotor performance with an associated potential increase in predation risk. In particular, the ability to initiate flight through jumping is critical to predator avoidance and may be influenced by changes in body mass (Mb). Here we investigate seasonal differences in the jump take-off performance of high Arctic Svalbard rock ptarmigan (Lagopus muta hyperborea) resulting from around a 50% increase in Mb during winter as a result of fat deposition. Using force-plate data and videography, we reveal that, in the absence of alterations to take-off angle, winter Svalbard rock ptarmigan are unable to increase hind-limb power output during jumping to compensate for their increased Mb. As a result, peak take-off velocity is reduced by 42% and jump duration is also extended during winter. The consequences of reduced jumping performance upon Svalbard ptarmigan during winter may be relatively small given their low risk of predation during this season. It may be, however, that the observed reduction in jumping performance when fat may contribute to the sub-maximal pattern of fat acquisition observed in other bird species.

Recently it was proposed that the primary feathers of Archaeopteryx lithographica (HMN1880) were overlaid by long covert feathers, and that a multilayered feathered wing was a feature of early fossils with feathered forelimbs. The proposed long covert feathers of Archaeopteryx were previously interpreted as dorsally displaced remiges or a second set of impressions made by the wing. The following study shows that the qualitative arguments forwarded in support of the elongated covert hypothesis are neither robust nor supported quantitatively. The idea that the extant bird wing with its single layer of overlapping primaries evolved from an earlier multilayered heavily coveted feathered forelimb as seen in Anchiornis huxleyi is reasonable. At this juncture, however, it is premature to conclude unequivocally that the wing of Archaeopteryx consisted of primary feathers overlaid with elongated coverts.

Extinct animal behavior has often been inferred from qualitative assessments of relative brain region size in fossil endocranial casts. For instance, flight capability in pterosaurs and early birds has been inferred from the relative size of the cerebellar flocculus, which in life protrudes from the lateral surface of the cerebellum. A primary role of the flocculus is to integrate sensory information about head rotation and translation to stabilize visual gaze via the vestibulo-occular reflex (VOR). Because gaze stabilization is a critical aspect of flight, some authors have suggested that the flocculus is enlarged in flying species. Whether this can be further extended to a floccular expansion in highly maneuverable flying species or floccular reduction in flightless species is unknown. Here, we used micro computed-tomography to reconstruct "virtual" endocranial casts of 60 extant bird species, to extract the same level of anatomical information offered by fossils. Volumes of the floccular fossa and entire brain cavity were measured and these values correlated with four indices of flying behavior. Although a weak positive relationship was found between floccular fossa size and brachial index, no significant relationship was found between floccular fossa size and any other flight mode classification. These findings could be the result of the bony endocranium inaccurately reflecting the size of the neural flocculus, but might also reflect the importance of the flocculus for all modes of locomotion in birds. We therefore conclude that the relative size of the flocculus of endocranial casts is an unreliable predictor of locomotor behavior in extinct birds, and probably also pterosaurs and non-avian dinosaurs.

Using open-flow respirometry and video footage (25 frames s(-1)), the energy expenditure and hindlimb kinematics of barnacle geese, Branta leucopsis, were measured whilst they were exercising on a treadmill at gradients of +7 and -7 deg, and on a level surface. In agreement with previous studies, ascending a gradient incurred metabolic costs higher than those experienced on level ground at comparable speeds. The geese, however, are the first species to show an increased duty factor when ascending a gradient. This increased duty factor was accompanied by a longer stance time, which was probably to enable the additional force required for ascending to be generated. Contrary to previous findings, the geese did not experience decreased metabolic costs when descending a gradient. For a given speed, the geese took relatively shorter and quicker strides when walking downhill. This 'choppy' stride and perhaps a lack of postural plasticity (an inability to adopt a more crouched posture) may negate any energy savings gained from gravity's assistance in moving the centre of mass downhill. Also contrary to previous studies, the incremental increase in metabolic cost with increasing speed was similar for each gradient, indicating that the efficiency of locomotion (mechanical work done/chemical energy consumed) is not constant across all walking speeds. The data here suggest that there are species-specific metabolic responses to locomotion on slopes, as well as the established kinematics differences. It is likely that a suite of factors, such as ecology, posture, gait, leggedness and foot morphology, will subtly affect an organism's ability to negotiate gradients.

Little is known regarding the physiological consequences of the behavioural and morphological differences that result from sexual selection in birds. Male and female Svalbard rock ptarmigans (Lagopus muta hyperborea) exhibit distinctive behavioural differences during the breeding season. In particular, males continuously compete for and defend territories in order to breed successfully, placing large demands on their locomotor system. Here, we demonstrate that male birds have improved locomotor performance compared with females, showing both a lower cost of locomotion (CoL) and a higher top speed. We propose that the observed sex differences in locomotor capability may be due to sexual selection for improved male performance. While the mechanisms underlying these energetic differences are unclear, future studies should be wary when pooling male and female data.

Here we report the resting metabolic rate in barnacle geese (Branta leucopsis) and provide evidence for the significant energetic effect of posture. Under laboratory conditions flow-through respirometry together with synchronous recording of behaviour enabled a calculation of how metabolic rate varies with posture. Our principal finding is that standing bipedally incurs a 25% increase in metabolic rate compared to birds sitting on the ground. In addition to the expected decrease in energy consumption of hindlimb postural muscles when sitting, we hypothesise that a change in breathing mechanics represents one potential mechanism for at least part of the observed difference in energetic cost. Due to the significant effect of posture, future studies of resting metabolic rates need to take into account and/or report differences in posture.

Svalbard rock ptarmigans were walked and run upon a treadmill and their energy expenditure measured using respirometry. The ptarmigan used three different gaits: a walking gait at slow speeds (less than or equal to 0.75 m s(-1)), grounded running at intermediate speeds (0.75 m s(-1) < U < 1.67 m s(-1)) and aerial running at high speeds (greater than or equal to 1.67 m s(-1)). Changes of gait were associated with reductions in the gross cost of transport (COT; J kg(-1) m(-1)), providing the first evidence for energy savings with gait change in a small crouched-postured vertebrate. In addition, for the first time (excluding humans) a decrease in absolute metabolic energy expenditure (rate of O(2) consumption) in aerial running when compared with grounded running was identified. The COT versus U curve varies between species and the COT was cheaper during aerial running than grounded running, posing the question of why grounded running should be used at all. Existing explanations (e.g. stability during running over rocky terrain) amount to just so stories with no current evidence to support them. It may be that grounded running is just an artefact of treadmill studies. Research investigating the speeds used by animals in the field is sorely needed.

The jump performance of five insectivorous bat species (Miniopterus schreibersii, Myotis blythii, Myotis capaccinii, Myotis myotis and Rhinolophus blasii) was filmed using a high-speed camera. All study bats jumped using a similar technique, with the wing musculature providing the force. The bats jumped off the wrist joint of their wings, typically with their feet already off the ground. Contrary to expectations, jump performance did not correlate with ecology and was instead strongly determined by body size. In general, the larger bats produced more jump force, left the ground at higher speeds and jumped higher than the smaller bats. The differences in force production disappeared when the data were corrected for body size, with the exception of Myotis capaccinii, which produced significantly less force. Scaling of jump performance with body size measured here was compared against two existing muscle performance scaling models. The model suggesting that muscle contraction velocity is proportional to muscle length was better supported than that based on muscle cross-sectional area. Both models, however, failed to accurately predict the scaling of jump forces, with the slope of the relationship being significantly steeper than predicted, highlighting the need for further investigations of vertebrate muscle performance scaling. The results of this study indicate that a bat's jumping ability is a secondary locomotor ability that uses the strongly selected-for flight apparatus with no apparent ecological trend present, i.e. flight so dominates bat locomotor morphology that other locomotor abilities tend to be derivative.

Wind tunnel tests conducted on a model based on the long-eared bat Plecotus auritus indicated that the positioning of the tail membrane (uropatagium) can significantly influence flight control. Adjusting tail position by increasing the angle of the legs ventrally relative to the body has a two-fold effect; increasing leg-induced wing camber (i.e., locally increased camber of the inner wing surface) and increasing the angle of attack of the tail membrane. We also used our model to examine the effects of flying with and without a tail membrane. For the bat model with a tail membrane increasing leg angle increased the lift, drag and pitching moment (nose-down) produced. However, removing the tail membrane significantly reduced the change in pitching moment with increasing leg angle, but it had no significant effect on the level of lift produced. The drag on the model also significantly increased with the removal of the tail membrane. The tail membrane, therefore, is potentially important for controlling the level of pitching moment produced by bats and an aid to flight control, specifically improving agility and manoeuvrability. Although the tail of bats is different from that of birds, in that it is only divided from the wings by the legs, it nonetheless, may, in addition to its prey capturing function, fulfil a similar role in aiding flight control.

The evolution of the avian wing has long fascinated biologists, yet almost no work includes the length of primary feathers in consideration of overall wing length variation. Here we show that the length of the longest primary feather (f(prim)) contributing to overall wing length scales with negative allometry against total arm (ta = humerus+ulna+manus). The scaling exponent varied slightly, although not significantly so, depending on whether a species level analysis was used or phylogeny was controlled for using independent contrasts: f(prim) is proportional to ta(0.78-0.82). The scaling exponent was not significantly different from that predicted (0.86) by earlier work. It appears that there is a general trend for the primary feathers of birds to contribute proportionally less, and ta proportionally more, to overall wingspan as this dimension increases. Wingspan in birds is constrained close to mass (M(1/3)) because of optimisation for lift production, which limits opportunities for exterior morphological change. Within the wing, variations in underlying bone and feather lengths nevertheless may, in altering the joint positions, permit a range of different flight styles by facilitating variation in upstroke kinematics.

The fossil birds Archaeopteryx and Confuciusornis had feathered wings resembling those of living birds, but their flight capabilities remain uncertain. Analysis of the rachises of their primary feathers shows that the rachises were much thinner and weaker than those of modern birds, and thus the birds were not capable of flight. Only if the primary feather rachises were solid in cross-section (the strongest structural configuration), and not hollow as in living birds, would flight have been possible. Hence, if Archaeopteryx and Confuciusornis were flapping flyers, they must have had a feather structure that was fundamentally different from that of living birds. Alternatively, if they were only gliders, then the flapping wing stroke must have appeared after the divergence of Confuciusornis, likely within the enantiornithine or ornithurine radiations.

A reanalysis of existing data suggests that the established tenet of increasing efficiency of transport with body size in terrestrial locomotion requires re-evaluation. Here, the statistical model that described the data best indicated a dichotomy between the data for small (<1 kg) and large animals (>1 kg). Within and between these two size groups there was no detectable difference in the scaling exponents (slopes) relating metabolic (E(met)) and mechanical costs (E(mech, CM)) of locomotion to body mass (M(b)). Therefore, no scaling of efficiency (E(mech, CM)/E(met)) with M(b) was evident within each size group. Small animals, however, appeared to be generally less efficient than larger animals (7% and 26% respectively). Consequently, it is possible that the relationship between efficiency and M(b) is not continuous, but, rather, involves a step-change. This step-change in the efficiency of locomotion mirrors previous findings suggesting a postural cause for an apparent size dichotomy in the relationship between E(met) and M(b). Currently data for E(mech, CM) is lacking, but the relationship between efficiency in terrestrial locomotion and M(b) is likely to be determined by posture and kinematics rather than body size alone. Hence, scaling of efficiency is likely to be more complex than a simple linear relationship across body sizes. A homogenous study of the mechanical cost of terrestrial locomotion across a broad range of species, body sizes, and importantly locomotor postures is a priority for future research.

Ontogenetic and behavioral studies using birds currently do not document the early evolution of flight because birds (including juveniles) used in such studies employ forelimb oscillation frequencies over 10 Hz, forelimb stroke-angles in excess of 130 degrees , and possess uniquely avian flight musculatures. Living birds are an advanced morphological stage in the development of flapping flight. To gain insight into the early stages of flight evolution (i.e., prebird), in the absence of a living analogue, a new approach using Strouhal number was used. Strouhal number is a nondimensional number that describes the relationship between wing-stroke amplitude (A), wing-beat frequency (f), and flight speed (U). Calculations indicated that even moderate wing movements are enough to generate rudimentary thrust and that a propulsive flapping flight-stroke could have evolved via gradual incremental changes in wing movement and wing morphology. More fundamental to the origin of the avian flapping flight-stroke is the question of how a symmetrical forelimb posture-required for gliding and flapping flight-evolved from an alternating forelimb motion, evident in all extant bipeds when running except birds.

Ecogeographical rules provide potential to describe how organisms are morphologically constrained to climatic conditions. Allen's rule (relatively shorter appendages in colder environments) remains largely unsupported and there remains much controversy whether reduced surface area of appendages provides energetic savings sufficient to make this morphological trend truly adaptive. By showing for the first time that Allen's rule holds for closely related endothermic species, we provide persuasive support of the adaptive significance of this trend for multiple species. Our results indicate that reduction of thermoregulatory cost during the coldest part of the breeding season is the most likely mechanism driving Allen's rule for these species. Because for 54% of seabird species examined, rise in seasonal maximum temperature over 100 years will exceed that for minimum temperatures, an evolutionary mismatch will arise between selection for limb length reduction and ability to accommodate heat stress.

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.

Here we show, by qualitative free- and tethered-flight flow visualization, that dragonflies fly by using unsteady aerodynamic mechanisms to generate high-lift, leading-edge vortices. In normal free flight, dragonflies use counterstroking kinematics, with a leading-edge vortex (LEV) on the forewing downstroke, attached flow on the forewing upstroke, and attached flow on the hindwing throughout. Accelerating dragonflies switch to in-phase wing-beats with highly separated downstroke flows, with a single LEV attached across both the fore- and hindwings. We use smoke visualizations to distinguish between the three simplest local analytical solutions of the Navier-Stokes equations yielding flow separation resulting in a LEV. The LEV is an open U-shaped separation, continuous across the thorax, running parallel to the wing leading edge and inflecting at the tips to form wingtip vortices. Air spirals in to a free-slip critical point over the centreline as the LEV grows. Spanwise flow is not a dominant feature of the flow field--spanwise flows sometimes run from wingtip to centreline, or vice versa--depending on the degree of sideslip. LEV formation always coincides with rapid increases in angle of attack, and the smoke visualizations clearly show the formation of LEVs whenever a rapid increase in angle of attack occurs. There is no discrete starting vortex. Instead, a shear layer forms behind the trailing edge whenever the wing is at a non-zero angle of attack, and rolls up, under Kelvin-Helmholtz instability, into a series of transverse vortices with circulation of opposite sign to the circulation around the wing and LEV. The flow fields produced by dragonflies differ qualitatively from those published for mechanical models of dragonflies, fruitflies and hawkmoths, which preclude natural wing interactions. However, controlled parametric experiments show that, provided the Strouhal number is appropriate and the natural interaction between left and right wings can occur, even a simple plunging plate can reproduce the detailed features of the flow seen in dragonflies. In our models, and in dragonflies, it appears that stability of the LEV is achieved by a general mechanism whereby flapping kinematics are configured so that a LEV would be expected to form naturally over the wing and remain attached for the duration of the stroke. However, the actual formation and shedding of the LEV is controlled by wing angle of attack, which dragonflies can vary through both extremes, from zero up to a range that leads to immediate flow separation at any time during a wing stroke.

The wing kinematics of birds vary systematically with body size, but we still, after several decades of research, lack a clear mechanistic understanding of the aerodynamic selection pressures that shape them. Swimming and flying animals have recently been shown to cruise at Strouhal numbers (St) corresponding to a regime of vortex growth and shedding in which the propulsive efficiency of flapping foils peaks (St approximately fA/U, where f is wingbeat frequency, U is cruising speed and A approximately bsin(theta/2) is stroke amplitude, in which b is wingspan and theta is stroke angle). We show that St is a simple and accurate predictor of wingbeat frequency in birds. The Strouhal numbers of cruising birds have converged on the lower end of the range 0.2 < St < 0.4 associated with high propulsive efficiency. Stroke angle scales as theta approximately 67b-0.24, so wingbeat frequency can be predicted as f approximately St.U/bsin(33.5b-0.24), with St0.21 and St0.25 for direct and intermittent fliers, respectively. This simple aerodynamic model predicts wingbeat frequency better than any other relationship proposed to date, explaining 90% of the observed variance in a sample of 60 bird species. Avian wing kinematics therefore appear to have been tuned by natural selection for high aerodynamic efficiency: physical and physiological constraints upon wing kinematics must be reconsidered in this light.

Analysis of a comprehensive dataset demonstrates that the brachial index (BI = humerus length/ulna length) of modern birds (Neornithes) varies significantly between clades at all taxonomic levels, yet is strongly correlated with recent phylogenetic hypotheses. Variance in BI at the infraclass level is low, but increases rapidly during the proposed major radiation of neornithines in the Palaeocene and Eocene. Although a BI of greater than 1 is primitive for Neornithes, more basal groups of Mesozoic birds (Confuciusornithidae and some members of the diverse Enantiornithidae) had BIs comparable with those of 'higher' modern clades. It is possible that occupation of ecological niches by these Mesozoic clades precluded the divergence of some groups of neornithines until after the Cretaceous-Tertiary boundary. We suggest that with further analysis and data collection the relationships between flight behaviour, ecology and BI can be determined. Hence, BI may provide a useful tool for characterizing the ecology of fossil birds.

Daily energy expenditure (DEE) of male barn swallows (Hirundo rustica), measured using the doubly labelled water technique, correlated with streamer length. Contrary to predictions derived from previous findings, neither a positive linear nor a u-shaped relationship was found between DEE and streamer length. Instead, an n-shaped curve showed that the highest DEE corresponded to an intermediate streamer length of 119 mm. A model incorporating estimates of resting metabolism and flight energy expenditure from aerodynamics models suggested that variation in individual tail dynamics was the most likely explanation for the observed DEE. We suggest that streamer length is coupled to foraging strategy, because tail dynamics control flight performance, which in turn influences flight behaviour.

Dimensionless numbers are important in biomechanics because their constancy can imply dynamic similarity between systems, despite possible differences in medium or scale. A dimensionless parameter that describes the tail or wing kinematics of swimming and flying animals is the Strouhal number, St = fA/U, which divides stroke frequency (f) and amplitude (A) by forward speed (U). St is known to govern a well-defined series of vortex growth and shedding regimes for airfoils undergoing pitching and heaving motions. Propulsive efficiency is high over a narrow range of St and usually peaks within the interval 0.2 < St < 0.4 (refs 3-8). Because natural selection is likely to tune animals for high propulsive efficiency, we expect it to constrain the range of St that animals use. This seems to be true for dolphins, sharks and bony fish, which swim at 0.2 < St < 0.4. Here we show that birds, bats and insects also converge on the same narrow range of St, but only when cruising. Tuning cruise kinematics to optimize St therefore seems to be a general principle of oscillatory lift-based propulsion.

The doubly-labeled water technique and video were used to measure the effect of mass loading on energy expenditure and takeoff performance in zebra finches, Taeniopygia guttata, that were making routine (nonalarm) short flights. Finches that carried 27% additional mass did not expend more energy during flight than unloaded controls. Carrying additional mass, however, led to a reduced body mass and a decreased velocity during takeoffs (by 12%). Calculations of instantaneous mechanical power indicated that energy expended by unloaded and loaded finches at takeoff was similar, due to the observed decrease in velocity by mass-loaded finches and a lowering of their body mass. During routine short flights, zebra finches appear to maintain their metabolic power input and mechanical power output regardless of mass loading. Here, the costs of carrying additional mass during routine short flights were revealed to be behavioral and not energetic.

Many small birds perform short flights, for which take-offs, ascents and descents form a large component of the total flight time and which are characterised by low airspeeds. Using the doubly-labelled water technique, zebra finches Taeniopygia guttata engaging in repeated short flights were found to expend 13.65 kJ more than 'non-flying' controls, which equated to a flight expenditure of 27.8 times their basal metabolic rate. This is over three times the predicted flight expenditure derived from existing aerodynamic models. These data were used to determine a coefficient (0.11) for converting the mechanical power derived from aerodynamic models into metabolic power. An equation is presented, based on body mass, which can be used to predict the costs of short flights in ecological and behavioural studies of birds.