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Oxygen limitation has been proposed as one of the key factors that limits body size at high temperatures (the oxygen-temperature hypothesis). Geographic patterns in body size are thought to be driven in part by the effects of temperature on oxygen supply and demand, particularly when the increased oxygen demand of tissues at higher temperatures outpaces the ability of large organisms to supply internal tissues with oxygen. We tested the effects of temperature on the rate of oxygen consumption of two temperate sea spider (Pycnogonida) species, Achelia chelata and Achelia gracilipes, across a range of body sizes. We measured oxygen consumption at 5 temperatures: 12, 16, 20, 24, and 28 [degrees]C. Oxygen consumption of both species increased significantly with temperature, but the effect did not depend on body size; thus, we found no evidence to support the oxygen-temperature hypothesis. While previous interspecific studies on Antarctic pycnogonids have found that larger-bodied animals have more porous cuticles, thus potentially offsetting their higher aerobic metabolic demand by increasing oxygen diffusivity, the pore area of the cuticle of the two temperate species did not change with body size. This suggests that the generally small size of warm-water sea spiders may be due to selective factors other than oxygen limitation., Introduction Almost all aspects of an organism's physiology are influenced by body size (Kingsolver and Huey, 2008). As a result, the physiological and environmental factors that underlie evolutionary or acclimatory [...]

A huge amount of research attention has focused on the evolution of life histories, but most research focuses on dominant individuals that acquire a disproportionate level of reproductive success, while the life histories and reproductive tactics of subordinate individuals have received less attention. Here, we review the links between early life adversity and performance during adulthood in birds, and highlight instances in which subordinate individuals outperform dominant conspecifics. Subordinate individuals are those from broods raised under high risk of predation, with low availability of food, and/or with many parasites. Meanwhile, the broods of many species hatch or are born asynchronously and mitigation of the asynchrony is generally lacking from variation in maternal effects such as egg size and hormone deposition or genetic effects such as offspring sex or parentage. Subordinate individuals employ patterns of differential growth to attempt to mitigate the adversity they experience during early life, yet they overwhelmingly fail to overcome their initial handicap. In terms of surviving through to adulthood, subordinate individuals employ other "suboptimal" tactics, such as adaptively timing foraging behaviors to avoid dominant individuals. During adulthood, meanwhile, subordinate individuals rely on "suboptimal" tactics, such as adaptive dispersal behaviors and competing for partners at optimal times, because they represent the best options available to them to acquire copulations whenever possible. We conclude that there is a gap in knowledge for direct links between early life adversity and subordination during adulthood, meaning that further research should test for links. There are instances, however, where subordinate individuals employ "suboptimal" tactics that allow them to outperform dominant conspecifics during adulthood. [ABSTRACT FROM AUTHOR]

A study published in Functional Ecology explores the constraints on the dispersal of forest-dwelling tropical birds. The researchers found that species with long, slender, and pointed wings, as well as proportionally small eyes, were better able to fly across open habitats. The study also revealed that environmental filtering played a role in shaping the phenotypes of bird species in fragmented forests. The findings highlight the need for further research on the biomechanical and physiological mechanisms that affect dispersal ability in tropical birds. [Extracted from the article]

The evolution of nest site use and nest architecture in the non-avian ancestors of birds remains poorly understood because nest structures do not preserve well as fossils. Nevertheless, the evidence suggests that the earliest dinosaurs probably buried eggs below ground and covered them with soil so that heat from the substrate fuelled embryo development, while some later dinosaurs laid partially exposed clutches where adults incubated them and protected them from predators and parasites. The nests of euornithine birds—the precursors to modern birds—were probably partially open and the neornithine birds—or modern birds—were probably the first to build fully exposed nests. The shift towards smaller, open cup nests has been accompanied by shifts in reproductive traits, with female birds having one functioning ovary in contrast to the two ovaries of crocodilians and many non-avian dinosaurs. The evolutionary trend among extant birds and their ancestors has been toward the evolution of greater cognitive abilities to construct in a wider diversity of sites and providing more care for significantly fewer, increasingly altricial, offspring. The highly derived passerines reflect this pattern with many species building small, architecturally complex nests in open sites and investing significant care into altricial young. This article is part of the theme issue 'The evolutionary ecology of nests: a cross-taxon approach'. [ABSTRACT FROM AUTHOR]

Hummingbirds have evolved to hover and manoeuvre with exceptional flight control. This is enabled by their musculoskeletal system that successfully exploits the agile motion of flapping wings. Here, we synthesize existing empirical and modelling data to generate novel hypotheses for principles of hummingbird wing actuation. These may help guide future experimental work and provide insights into the evolution and robotic emulation of hummingbird flight. We develop a functional model of the hummingbird musculoskeletal system, which predicts instantaneous, three-dimensional torque produced by primary (pectoralis and supracoracoideus) and combined secondary muscles. The model also predicts primary muscle contractile behaviour, including stress, strain, elasticity and work. Results suggest that the primary muscles (i.e. the flight 'engine') function as diverse effectors, as they do not simply power the stroke, but also actively deviate and pitch the wing with comparable actuation torque. The results also suggest that the secondary muscles produce controlled-tightening effects by acting against primary muscles in deviation and pitching. The diverse effects of the pectoralis are associated with the evolution of a comparatively enormous bicipital crest on the humerus. [ABSTRACT FROM AUTHOR]

Much of the awe that humans have for the flight of birds derives from our earthbound habits and our bias toward emphasizing visual cues for interpreting processes in the world. Although we move through it and breathe it, air is vastly less dense than our bodies, so it is fanciful to imagine moving our limbs in a manner that would enable us to support our weight in the air. Moreover, air is invisible to us unless we use special tools to reveal its flow patterns. As evidence of our visual bias, contemplate a strong wind. You probably form a mental image of leaves moving on tree branches, or dust swirling about in a tornado, in both cases, solids moving in response to the force of the wind rather than the essence of the wind itself. The aerodynamics of bird flight are complex, particularly when the wings are dynamically changing shape and when flapping velocities exceed the forward velocity of the bird. In this primer, Bret W. Tobalske explores how wing function varies with flight speed, the formation of tip and root vortices that are useful to researchers but represent a constraint on span efficiency, and tail function. He encourages new research into flow sensing and extending existing knowledge into phylogenetic and ontogenetic contexts. [ABSTRACT FROM AUTHOR]

Diving birds are regarded as a classic example of morphological convergence. Divers tend to have small wings extending from rotund bodies, requiring many volant species to fly with rapid wingbeats, and rendering others flightless. The high wing-loading of diving birds is frequently associated with the challenge of using forelimbs adapted for flight for locomotion in a "draggier" fluid, but this does not explain why species that rely exclusively on their feet to dive should have relatively small wings, as well. Therefore, others have hypothesized that ecological factors shared by wing-propelled and foot-propelled diving birds drive the evolution of high wing-loading. Following a reexamination of the aquatic habits of birds, we tested between hypotheses seeking to explain high wing-loading in divers using new comparative data and phylogenetically informed analyses. We found little evidence that wing-propelled diving selects for small wings, as wing-propelled and foot-propelled species share similar wing-loadings. Instead, our results suggest that selection to reduce buoyancy has driven high wing-loading in divers, offering insights for the development of bird-like aquatic robots. [ABSTRACT FROM AUTHOR]

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

As one of few animals with the capability to execute agile yawing maneuvers, it is quite desirable to take inspiration from hummingbird flight aerodynamics. To understand the wing and body kinematics and associated aerodynamics of a hummingbird performing a free yawing maneuver, a crucial step in mimicking the biological motion in robotic systems, we paired accurate digital reconstruction techniques with high-fidelity computational fluid dynamics (CFD) simulations. Results of the body and wing kinematics reveal that to achieve the pure yaw maneuver, the hummingbird utilizes very little body pitching, rolling, vertical, or horizontal motion. Wing angle of incidence, stroke, and twist angles are found to be higher for the inner wing (IW) than the outer wing (OW). Unsteady aerodynamic calculations reveal that drag-based asymmetric force generation during the downstroke (DS) and upstroke (US) serves to control the speed of the turn, a characteristic that allows for great maneuvering precision. A dual-loop vortex formation during each half-stroke is found to contribute to asymmetric drag production. Wake analysis revealed that asymmetric wing kinematics led to leading-edge vortex strength differences of around 59% between the IW and OW. Finally, analysis of the role of wing flexibility revealed that flexibility is essential for generating the large torque necessary for completing the turn as well as producing sufficient lift for weight support. [ABSTRACT FROM AUTHOR]

Author(s): Douglas R. Warrick (corresponding author) [1]; Bret W. Tobalske [2]; Donald R. Powers [3] Despite profound musculoskeletal differences, hummingbirds (Trochilidae) are widely thought to employ aerodynamic mechanisms similar to [...]

The shape of remiges (primary and secondary feathers) is constrained and stereotyped by the demands of flight, but members of the subfamily of New World ground doves (Peristerinae) possess many atypical remex shapes with which they produce sonations of alarm. Within the genus Columbina specifically, the seventh primary feathers (P7) have elongated barbs that create a protrusion on the trailing vane which varies in size and shape between species. These feathers are hypothesized to have been coopted to produce communicative sounds (i.e. sonations) during flight, but the mechanism of this sound production is unknown. We tested the sound-producing capabilities of spread wing specimens from three species of ground doves (C. inca , C. passerina , and C. talpacoti) in a wind tunnel. High speed video and audio analyses indicated that all wings of adult birds produced buzzing sounds in the orientation and flow velocity of mid-upstroke. These buzzing sounds were produced as the protrusion of elongated barbs fluttered and collided with adjacent P6 feathers at a fundamental frequency of 200 and 400 Hz, respectively. Wings from juvenile C. inca produced significantly quieter buzzes and most (three of four individuals) lacked the elongated barbs that are present in adults. Buzzing sounds produced in the wind tunnel were similar to those produced by wild birds indicating that these P7 feathers have been coopted to produce acoustic signals (sonations) during flight. The shape and mechanism of sound production described here in Columbina appear to be unique among birds. [ABSTRACT FROM AUTHOR]

Developmental responses can help young animals reduce predation risk but can also yield costs to performance and survival in subsequent life stages with major implications for lifetime fitness. Compensatory mechanisms may evolve to offset such costs, but evidence from natural systems is largely lacking.In songbirds, increased nest predation risk should favour reduced provisioning, but also young that fledge (leave their nest) at an earlier age. Both responses can result in fledglings with shorter wings, reduced mobility and decreased survival. Young may compensate for shorter wings developmentally by reallocating resources towards feather development or behaviourally by adjusting flight kinematics or habitat use. However, underfed young may lack the capacity to express these phenotypes due to insufficient resources or an inability to adjust allocation of resources.Using predation risk experiments and 29 years of observational field data, we test whether increased nest predation risk reduces flight performance and survival during the fledgling stage and explore potential mechanisms that might underlie these effects. We show that young from high‐risk nests did not leave the nest earlier on average, but wing growth was slower likely due to observed reductions in parental feeding rates. Wings were shorter in high‐risk nests when fledglings left the nest early. Yet, fledglings from high‐risk nests showed improved flight performance for a given wing length such that flight performance at fledging did not differ between young from high‐risk and low‐risk nests. Young from high‐risk nests may have offset the costs of shorter wings on flight performance by accelerating the emergence of flight feathers from their sheaths to reduce wing porosity, though evidence for this mechanism was mixed. Fledglings from high‐risk nests also selected habitat with denser woody vegetation compared with young from low‐risk nests.Together, these developmental and behavioural responses seem to mitigate the expected effects of increased nest predation risk on fledgling survival. Ultimately, our results show that offspring predation risk can affect parental provisioning and offspring morphology without major implications for performance and survival in subsequent life stages. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

The natural world is filled with substrates of varying properties that challenge locomotor abilities. Birds appear to transition smoothly from aerial to terrestrial environments during take-offs and landings using substrates that are incredibly variable. It may be challenging to control movement on and off compliant (flexible) substrates such as twigs, yet birds routinely accomplish such tasks. Previous research suggests that birds do not use their legs to harness elastic recoil from perches. Given avian mastery of take-off and landing, we hypothesized that birds instead modulate wing, body and tail movements to effectively use compliant perches. We measured take-off and landing performance of diamond doves (Geopelia cuneata (N = 5) in the laboratory and perch selection in this species in the field (N = 25). Contrary to our hypothesis, doves do not control take-off and landing on compliant perches as effectively as they do on stiff perches. They do not recover elastic energy from the perch, and take-off velocities are thus negatively impacted. Landing velocities remain unchanged, which suggests they may not anticipate the need to compensate for compliance. Legs and wings function as independent units: legs produce lower initial velocities when taking off from a compliant substrate, which negatively impacts later flight velocities. During landing, significant stability problems arise with compliance that are ameliorated by the wings and tail. Collectively, we suggest that the diamond dove maintains a generalized take-off and landing behavior regardless of perch compliance, leading us to conclude that perch compliance represents a challenge for flying birds. Free-living diamond doves avoid the negative impacts of compliance by preferentially selecting perches of larger diameter, which tend to be stiffer. [ABSTRACT FROM AUTHOR]

The environmental conditions animals experience during development can have sustained effects on morphology, physiology, and behavior. Exposure to elevated levels of stress hormones (glucocorticoids, GCs) during development is one such condition that can have long-term effects on animal phenotype. Many of the phenotypic effects of GC exposure during development (developmental stress) appear negative. However, there is increasing evidence that developmental stress can induce adaptive phenotypic changes. This hypothesis can be tested by examining the effect of developmental stress on fitness-related traits. In birds, flight performance is an ideal metric to assess the fitness consequences of developmental stress. As fledglings, mastering takeoff is crucial to avoid bodily damage and escape predation. As adults, takeoff can contribute to mating and foraging success as well as escape and, thus, can affect both reproductive success and survival. We examined the effects of developmental stress on flight performance across life-history stages in zebra finches ( Taeniopygia guttata). Specifically, we examined the effects of oral administration of corticosterone ( CORT, the dominant avian glucocorticoid) during development on ground-reaction forces and velocity during takeoff. Additionally, we tested for associations between flight performance and reproductive success in adult male zebra finches. Developmental stress had no effect on flight performance at all ages. In contrast, brood size (an unmanipulated variable) had sustained, negative effects on takeoff performance across life-history stages with birds from small broods performing better than birds from large broods. Flight performance at 100 days posthatching predicted future reproductive success in males; the best fliers had significantly higher reproductive success. Our results demonstrate that some environmental factors experienced during development (e.g. clutch size) have stronger, more sustained effects than others (e.g. GC exposure). Additionally, our data provide the first link between flight performance and a direct measure of reproductive success. [ABSTRACT FROM AUTHOR]

Hummingbirds are nature's masters of aerobatic manoeuvres. Previous research shows that hummingbirds and insects converged evolutionarily upon similar aerodynamic mechanisms and kinematics in hovering. Herein, we use three-dimensional kinematic data to begin to test for similar convergence of kinematics used for escape flight and to explore the effects of body size upon manoeuvring. We studied four hummingbird species in North America including two large species (magnificent hummingbird, Eugenes fulgens, 7.8 g, and blue-throated hummingbird, Lampornis clemenciae, 8.0 g) and two smaller species (broad-billed hummingbird, Cynanthus latirostris, 3.4 g, and black-chinned hummingbirds Archilochus alexandri, 3.1 g). Starting from a steady hover, hummingbirds consistently manoeuvred away from perceived threats using a drastic escape response that featured body pitch and roll rotations coupled with a large linear acceleration. Hummingbirds changed their flapping frequency and wing trajectory in all three degrees of freedom on a stroke-by-stroke basis, likely causing rapid and significant alteration of the magnitude and direction of aerodynamic forces. Thus it appears that the flight control of hummingbirds does not obey the 'helicopter model' that is valid for similar escape manoeuvres in fruit flies. Except for broad-billed hummingbirds, the hummingbirds had faster reaction times than those reported for visual feedback control in insects. The two larger hummingbird species performed pitch rotations and global-yaw turns with considerably larger magnitude than the smaller species, but roll rates and cumulative roll angles were similar among the four species. [ABSTRACT FROM AUTHOR]

The superiormanoeuvrability of hummingbirds emerges fromcomplex interactions of specialized neural and physiological processes with the unique flight dynamics of flapping wings. Escape manoeuvring is an ecologically relevant, natural behaviour of hummingbirds, from which we can gain understanding into the functional limits of vertebrate locomotor capacity. Here, we extend our kinematic analysis of escape manoeuvres from a companion paper to assess two potential limiting factors of the manoeuvring performance of hummingbirds: (1) muscle mechanical power output and (2) delays in the neural sensing and control system. We focused on the magnificent hummingbird (Eugenes fulgens, 7.8 g) and the black-chinned hummingbird (Archilochus alexandri, 3.1 g), which represent large and small species, respectively. We first estimated the aerodynamic forces, moments and the mechanical power of escape manoeuvres using measured wing kinematics. Comparing active-manoeuvring and passive-damping aerodynamic moments, we found that pitch dynamics were lightly damped and dominated by the effect of inertia, while roll dynamics were highly damped. To achieve observed closed-loop performance, pitch manoeuvres required faster sensorimotor transduction, as hummingbirds can only tolerate half the delay allowed in roll manoeuvres. Accordingly, our results suggested that pitch control may require a more sophisticated control strategy, such as those based on prediction. For the magnificent hummingbird, we estimated that escape manoeuvres required muscle mass-specific power 4.5 times that during hovering. Therefore, in addition to the limitation imposed by sensorimotor delays, muscle power could also limit the performance of escape manoeuvres. [ABSTRACT FROM AUTHOR]

Birds morph their wings during a single wingbeat, across flight speeds and among flight modes. Such morphing may allow them to maximize aerodynamic performance, but this assumption remains largely untested. We tested the aerodynamic performance of swept and extended wing postures of 13 raptor species in three families (Accipitridae, Falconidae and Strigidae) using a propeller model to emulate mid-downstroke of flapping during take-off and a wind tunnel to emulate gliding. Based on previous research, we hypothesized that (1) during flapping, wing posture would not affect maximum ratios of vertical and horizontal force coefficients (CV:CH), and that (2) extended wings would have higher maximum CV:CH when gliding. Contrary to each hypothesis, during flapping, extended wings had, on average, 31% higher maximum CV:CH ratios and 23% higher CV than swept wings across all biologically relevant attack angles (α), and, during gliding, maximum CV:CH ratios were similar for the two postures. Swept wings had 11% higher CV than extended wings in gliding flight, suggesting flow conditions around these flexed raptor wings may be different from those in previous studies of swifts (Apodidae). Phylogenetic affiliation was a poor predictor of wing performance, due in part to high intrafamilial variation. Mass was only significantly correlated with extended wing performance during gliding. We conclude that wing shape has a greater effect on force per unit wing area during flapping at low advance ratio, such as takeoff, than during gliding. [ABSTRACT FROM AUTHOR]

For centuries, naturalists have suggested that the tonal elements of pigeon wing sounds may be sonations (non-vocal acoustic signals) of alarm. However, spurious tonal sounds may be produced passively as a result of aeroelastic flutter in the flight feathers of almost all birds. Using mechanistic criteria emerging from recent work on sonations, we sought to: (1) identify characteristics of rock pigeon flight feathers that might be adapted for sound production rather than flight, and (2) provide evidence that this morphology is necessary for in vivo sound production and is sufficient to replicate in vivo sounds. Pigeons produce tonal sounds (700±50 Hz) during the latter two-thirds of each downstroke during take-off. These tones are produced when a small region of long, curved barbs on the inner vane of the outermost primary feather (P10) aeroelastically flutters. Tones were silenced in live birds when we experimentally increased the stiffness of this region to prevent flutter. Isolated P10 feathers were sufficient to reproduce in vivo sounds when spun at the peak angular velocity of downstroke (53.9-60.3 rad s-1), but did not produce tones at average downstroke velocity (31.8 rad s-1), whereas P9 and P1 feathers never produced tones. P10 feathers had significantly lower coefficients of resultant aerodynamic force (CR) when spun at peak angular velocity than at average angular velocity, revealing that production of tonal sounds incurs an aerodynamic cost. P9 and P1 feathers did not show this difference in CR. These mechanistic results suggest that the tonal sounds produced by P10 feathers are not incidental and may function in communication. [ABSTRACT FROM AUTHOR]

Slow flight is extremely energetically costly per unit time, yet highly important for takeoff and survival. However, at slow speeds it is presently thought that most birds do not produce beneficial aerodynamic forces during the entire wingbeat: instead they fold or flex their wings during upstroke, prompting the long-standing prediction that the upstroke produces trivial forces. There is increasing evidence that the upstroke contributes toforce production, but the aerodynamic and kinematic mechanisms remain unknown. Here, we examined the wingbeat cycle of two species: the diamond dove (Geopelia cuneata) and zebra finch (Taeniopygia guttata), which exhibit different upstroke styles - a wingtip-reversal and flexed-wing upstroke, respectively. We used a combination of particle image velocimetry and near-wake streamline measures alongside detailed 3D kinematics. We show that during the middle of the wingtip-reversal upstroke, the hand-wing has a high angular velocity (15.3±0.8 deg ms-1) and translational speed (8.4±0.6 m s-1). The flexed-wing upstroke, in contrast, has low wingtip speed during mid-upstroke. Instead, later in the stroke cycle, during the transition from upstroke to downstroke, it exhibits higher angular velocities (45.5±13.8 deg ms-1) and translational speeds (11.0±1.9 m s-1). Aerodynamically, the wingtip-reversal upstroke imparts momentum to the wake, with entrained air shed backward (visible as circulation of 14.4±0.09 m2 s-1). In contrast, the flexed-wing upstroke imparts minimal momentum. Clap and peel in the dove enhances the time course for circulation production on the wings, and provides new evidence of convergent evolution on time-varying aerodynamic mechanisms during flapping in insects and birds. [ABSTRACT FROM AUTHOR]

Transitions to and from the air are critical for aerial locomotion and likely shaped the evolution of flying animals. Research on take-off demonstrates that legs generate greater body accelerations compared with wings, and thereby contribute more to initial flight velocity. Here, we explored coordination between wings and legs in two species with different wingbeat styles, and quantified force production of these modules during the final phase of landing. We used the same birds that we had previously studied during take-off: zebra finch (Taeniopygia guttata, N=4) and diamond dove (Geopelia cuneata, N=3). We measured kinematics using high-speed video, aerodynamics using particle image velocimetry, and ground-reaction forces using a perch mounted on a force plate. In contrast with the first three wingbeats of take-off, the final four wingbeats during landing featured ~2 times greater force production. Thus, wings contribute proportionally more to changes in velocity during the last phase of landing compared with the initial phase of take-off. The two species touched down at the same velocity (~1 m s-1), but they exhibited significant differences in the timing of their final wingbeat relative to touchdown. The ratio of average wing force to peak leg force was greater in diamond doves than in zebra finches. Peak ground reaction forces during landing were ~50% of those during take-off, consistent with the birds being motivated to control landing. Likewise, estimations of mechanical energy flux for both species indicate that wings produce 3-10 times more mechanical work within the final wingbeats of flight compared with the kinetic energy of the body absorbed by legs during ground contact. [ABSTRACT FROM AUTHOR]