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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).)

In dictating the relative distances between the elbow, shoulder and wrist, avian brachial index (BI = humerus/radius-ulna length) likely influences wing kinematics and, therefore, might predict extinct avian flight capability. This underpins the hypothesis that non-neornithine Mesozoic avialans with relatively low BIs (associated with improved flight capabilities) restricted neornithine diversification until after the Cretaceous-Paleogene boundary. Here, correlations between flight metrics (wingbeat frequency (f), stroke angle (θ), wing loading (Q) and aspect ratio) and BI were investigated and vice versa. Additionally, the evolutionary model best describing the phylogenetic distribution of BI, and the temporal patterns in BI, flight metrics, body mass (M b ), and size-corrected humerus (L h ) and radius-ulna (L ru ) length were determined. BI was best described by Ornstein-Uhlenbeck processes, with low α values indicating a gradual shift towards a future theoretical optimum. BI also decreased overall through evolutionary time with the flight metrics mirroring temporal patterns of change in BI. M b , L h and L ru overall decreased apart from increases in L h and L ru following the middle-late Miocene (also leading to BI increasing) due to diversifications of the Anatinae and Sphenisciformes. L h overall decreased further than L ru. Consequently, decreasing L h mainly contributed to decreasing BI through evolutionary time, implying flight performance increased through neornithine evolution. However, the timings of radiations in these variables implies an Eocene radiation of neornthine flight ecology rather than a rapid expansion into niches vacated by non-neornithine Mesozoic avialians following the Cretaceous-Paleogene boundary. Multiple regressions showed f, θ and Q explained 60% of variation in BI. However, unequivocally evaluating whether BI is related to wing movement (and flight capability) requires direct measures of wing movement for many species, which are currently unavailable. Finally, the findings here and previously observed clade-specificity in BI, suggest flight ecology may also be clade-specific. Hence, the utility of phylogeny in predicting flight ecology requires exploration., (© 2021 Wiley Periodicals LLC.)

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.)

Domesticated brown trout Salmo trutta parr were subjected to increased, variable flow under controlled experimental conditions. Using geometric morphometric analyses, K ¯ (a mass-length index) and caudal fin area-body length ratio, this study assessed morphological responses in lateral body depth, growth and robustness and propulsive potential, respectively, of parr over the course of 32 weeks. Geometric morphometric analyses did not reveal an effect of exercise on either lateral body depth or caudal fin area. However, improved overall robustness and growth trajectories in exercised parr showed a positive adaptive response to the enriched habitat. Exercise and habitat heterogeneity thus have the potential to improve survivability of domesticated salmonids in the wild., (© 2018 The Authors/World Fish. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of The Fisheries Society of the British Isles.)

Competing Interests: Conflict of interest The authors declare that they have no conflict of interest.

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.)

Climate change and urbanisation of watercourses affect water temperatures and current flow velocities in river systems on a global scale. This represents a particularly critical issue for migratory fish species with complex life histories that use rivers to reproduce. Salmonids are migratory keystone species that provide substantial economical value to ecosystems and human societies. Consequently, a comprehensive understanding of the effects of environmental stressors on their reproductive success is critical in order to ensure their continued abundance during future climatic change. Salmonids are capital breeders, relying entirely on endogenous energy stores to fuel return migration to their natal spawning sites and reproduction upon arrival. Metabolic rates and cost of transport en-route increase with temperature and at extreme temperatures, swimming is increasingly fuelled anaerobically, resulting in an oxygen debt and reduced capacity to recover from exhaustive exercise. Thermally challenged salmonids also produce less viable gametes, which themselves are affected by water temperature after release. Passage through hydrological barriers and temperature changes both affect energy expenditure. As a result, important energetic tradeoffs emerge between extra energy used during migration and that available for other facets of the reproductive cycle, such as reproductive competition and gamete production. However, studies identifying these tradeoffs are extremely sparse. This review focuses on the specific locomotor responses of salmonids to thermal and hydrological challenges, identifying gaps in our knowledge and highlighting the potential implications for key aspects of their reproduction., (Copyright © 2015 The Authors. Published by Elsevier Inc. All rights reserved.)

This study examined how acute warming of water temperature affects the mechanical efficiency of swimming and aerobic capabilities of the brown trout Salmo trutta. Swimming efficiency was assessed using the relationship between swimming kinematics and forward speed (U), which is thought to converge upon an optimum range of a dimensionless parameter, the Strouhal number (St ). Swim-tunnel intermittent stopped-flow respirometry was used to record kinematics and measure oxygen consumption (ṀO2) of S. trutta during warming and swimming challenges. Salmo trutta maintained St between 0·2 and 0·3 at any given U over a range of temperatures, irrespective of body size. The maintenance of St within the range for maximum efficiency for oscillatory propulsion was achieved through an increase in tail-beat frequency (ftail) and a decrease in tail-beat amplitude (A) as temperature increased. Maintenance of efficient steady-state swimming was fuelled by aerobic metabolism, which increased as temperature increased up to 18° C but declined above this temperature, decreasing the apparent metabolic scope. As St was maintained over the full range of temperatures whilst metabolic scope was not, the results may suggest energetic trade-offs at any given U at temperatures above thermal optima., (© 2015 The Fisheries Society of the British Isles.)

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 material properties and morphologies of the modified integumentary organs of birds (the keratinous bills, claws and feathers) have evolved to withstand the variety of mechanical stresses imposed by their interaction with the environment. These stresses are likely to vary temporally in seasonal environments and may also differ between the sexes as a result of behavioural dimorphism. Here we investigate the morphology and material properties of the claws of male and female Svalbard ptarmigan (Lagopus muta hyperborea) during the summer and winter using nanoindentation. Despite differences in locomotor demands between the sexes and pronounced seasonal differences in environmental conditions, like ground substrate, ambient temperature and day length, there was no significant difference in Young׳s modulus or hardness between the seasons for each sex. However, when comparing males and females, female claws were significantly harder than those of males and both sexes had significantly wider claws during winter. We propose that wider claws may follow winter claw moulting as the claws are regrown and form an important part of the ptarmigan׳s snowshoe-like foot that is an adaptation to locomotion on snow. Future work focusing on growth rates and more broad measures of material properties in both captive and wild birds is required to determine the extent of seasonal and sex differences in the material properties of their keratinous structures., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

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.

The relative length proportions of the three bony elements of the pelvic (femur, tibiotarsus and tarsometatarsus) and pectoral (humerus, ulna and manus) limbs of the early Cretaceous bird Gansus yumenensis, a well-represented basal ornithuromorph from China, are investigated and compared to those of extant taxa. Ternary plots show that the pectoral limb length proportions of Gansus are most similar to Apodiformes (swifts and hummingbirds), which plot away from all other extant birds. In contrast, the pelvic limb length proportions of Gansus fall within the extant bird cluster and show similarities with the neornithine families Podicipedidae (grebes), Diomedeidae (albatross) and Phalacrocoracidae (cormorants). Although it does have some of the pelvic limb features of grebes and cormorants, the femur of Gansus is more gracile and is thus more consistent with an albatross-like shallow-diving mode of life than a strong foot-propelled diving movement pattern. The position of Gansus in pectoral limb ternary morphospace is largely due to its elongated manus. In contrast to apodiformes, where the humerus and ulna are short and robust, an adaptation, which provides a stiff wing for their demanding fast agile and hovering flight (respectively), the wing-bones of Gansus are slender, indicating a less vigorous flapping flight style. The suite of characters exhibited by Gansus mean it is difficult to completely interpret its likely ecology. Nevertheless, our analyses suggest that it is probable that this bird was both volant and capable of diving to some degree using either foot-propelled or, perhaps, both its wings and its feet for underwater locomotion., (© 2012 The Authors. Journal of Evolutionary Biology © 2012 European Society For Evolutionary Biology.)

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.

The primary feathers of birds are subject to cyclical forces in flight causing their shafts (rachises) to bend. The amount the feathers deflect during flight is dependent upon the flexural stiffness of the rachises. By quantifying scaling relationships between body mass and feather linear dimensions in a large data set of living birds, we show that both feather length and feather diameter scale much closer to predictions for geometric similarity than they do to elastic similarity. Scaling allometry also indicates that the primary feathers of larger birds are relatively shorter and their rachises relatively narrower, compared to those of smaller birds. Two-point bending tests indicated that larger birds have more flexible feathers than smaller species. Discriminant functional analyses (DFA) showed that body mass, primary feather length and rachis diameter can be used to differentiate between different magnitudes of feather bending stiffness, with primary feather length explaining 63% of variance in rachis stiffness. Adding fossil measurement data to our DFA showed that Archaeopteryx and Confuciusornis do not overlap with extant birds. This strongly suggests that the bending stiffness of their primary feathers was different to extant birds and provides further evidence for distinctive flight styles and likely limited flight ability in Archaeopteryx and Confuciusornis., (© 2012 The Authors. Journal of Evolutionary Biology © 2012 European Society For Evolutionary Biology.)

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.

We examine the relationships between primary feather length (f(prim)) and total arm length (ta) (sum of humerus, ulna and manus lengths) in Mesozoic fossil birds to address one aspect of avian wing shape evolution. Analyses show that there are significant differences in the composition of the wing between the known lineages of basal birds and that mean f(prim) (relative to ta length) is significantly shorter in Archaeopteryx and enantiornithines than it is in Confuciusornithidae and in living birds. Based on outgroup comparisons with nonavian theropods that preserve forelimb primary feathers, we show that the possession of a relatively shorter f(prim) (relative to ta length) must be the primitive condition for Aves. There is also a clear phylogenetic trend in relative primary feather length throughout bird evolution: our analyses demonstrate that the f(prim)/ta ratio increases among successive lineages of Mesozoic birds towards the crown of the tree ('modern birds'; Neornithes). Variance in this ratio also coincides with the enormous evolutionary radiation at the base of Neornithes. Because the f(prim)/ta ratio is linked to flight mode and performance in living birds, further comparisons of wing proportions among Mesozoic avians will prove informative and certainly imply that the aerial locomotion of the Early Cretaceous Confuciusornis was very different to other extinct and living birds., (© 2011 The Authors. Journal of Evolutionary Biology © 2011 European Society For Evolutionary Biology.)

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.

Barnacle geese were walked on a treadmill at speeds ranging from 0.25 to 1.25 ms(-1), which was their highest sustainable speed. No evidence for a gait change was found. The gait of a barnacle goose appears to conform to the classical pendulum mechanics based model of walking, with the kinetic energy of forward motion (horizontal kinetic energy, E(kh)) out-of-phase with the sum of the gravitational potential (E(p)), and vertical kinetic (E(kv)) energies of the centre of mass at all speeds. Why barnacle geese are unable to aerial run when other 'waddling' species do show an aerial phase (e.g., mallard ducks) is unclear. Presumably, however, it is likely to relate to the amount of lateral kinetic energy generated, which is a feature of 'waddling'. We predict that lateral kinetic energy generated by barnacle geese and other waddling species that cannot aerial run, is higher than in those that can. Due to competing selection pressures for swimming and flight, barnacle geese are mechanically and energetically inefficient walkers relative to more specialist cursorial birds. Their upper walking speed, however, appears to be limited by morphology (via kinematics) and not metabolic capacity (energetics)., (Copyright 2010 Elsevier Inc. All rights reserved.)

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.

Although pterosaurs are a well-known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon-specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased - this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.

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 225 million-year-old reptile Sharovipteryx mirabilis was the world's first delta-winged glider; this remarkable animal had a flight surface composed entirely of a hind-limb membrane. We use standard delta-wing aerodynamics to reconstruct the flight of S. mirabilis demonstrating that wing shape could have been controlled simply by protraction of the femora at the knees, and by variation in incidence of a small forelimb canard. Our method has allowed us to address the question of how identifying realistic glide performance can be used to set limits on aerodynamic design in this small animal. Our novel interpretation of the bizarre flight mode of S. mirabilis is the first based directly on interpretation of the fossil itself and the first grounded in aerodynamics.

The limb proportions of the extinct flying pterosaurs were clearly distinct from their living counterparts, birds and bats. Within pterosaurs, however, we show that further differences in limb proportions exist between the two main groups: the clade of short-tailed Pterodactyloidea and the paraphyletic clades of long-tailed rhamphorhynchoids. The hindlimb to forelimb ratios of rhamphorhynchoid pterosaurs are similar to that seen in bats, whereas those of pterodactyloids are much higher. Such a clear difference in limb ratios indicates that the extent of the wing membrane in rhamphorhynchoids and pterodactyloids may also have differed; this is borne out by simple ternary analyses. Further, analyses also indicate that the limbs of Sordes pilosus, a well-preserved small taxon used as key evidence for inferring the extent and shape of the wing membrane in all pterosaurs, are not typical even of its closest relatives, other rhamphorhynchoids. Thus, a bat-like extensive hindlimb flight membrane, integrated with the feet and tail may be applicable only to a small subset of pterosaur diversity. The range of flight morphologies seen in these extinct reptiles may prove much broader than previously thought.

By analyzing a homogenous dataset we show, in contradiction to a previous study, that the scaling of body frontal area (S(b)) with body mass (m(b)) does not differ between passerine and nonpasserine birds. It is likely that comparison of data collected from live passerines with data collected from frozen nonpasserines had led to the incorrect conclusion that the scaling of S(b) varied between the taxa. We suggest that body dimensions collected from frozen specimens, or specimens stored in alcohol, are not applicable to live birds, and that both the current equations presented in the literature for predicting S(b) from m(b) may lead to inaccurate estimates. Using data from preserved specimens, we found that S(b) scales isometrically with m(b) (S(b) proportional, variant m(b) (0.66)), and therefore we found no evidence for larger birds being more streamlined than smaller birds. S(b) scales with negative allometry against wingspan (b), however, and b scales with positive allometry against m(b), so larger birds have smaller S(b) relative to b. In addition, it appears that dorsoventral flattening of the body is a general characteristic of bird's bodies but that it is more pronounced in larger birds, suggesting perhaps a function in terms of increased lift during forward flight. It appears that bird's bodies obey the surface-to-area geometric scaling law, but bird body shape may vary in relation to aerodynamic function. We suggest that a large-scale study, simultaneously measuring S(b) and m(b) in live passerines and nonpasserines, is required to improve the predictive power of S(b) upon m(b) scaling equations, which play a key role in the estimation of mechanical power consumption in flight in birds. Furthermore, the relations between bird body shape and axial skeleton dimensions, with reference to aerodynamic adaptation, warrant further investigation.

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.