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Although theoretical work on optimal migration has been largely restricted to birds, relevant free-flight data are now becoming available for migratory insects. Here we report, for the first time in passion-vine butterflies, that Heliconius sara migrates directionally. To test optimal migration models for insects, we quantified the aerodynamic power curve for free-flying H. sara as they migrated across the Panama Canal. Using synchronized stereo-images from high-speed video cameras, we reconstructed three-dimensional flight kinematics of H. sara migrating naturally across the Panama Canal. We also reconstructed flight kinematics from a single-camera view of butterflies flying through a flight tunnel. We calculated the power requirements for flight for H. sara over a range of flight velocities. The relationship between aerodynamic power and velocity was "J"-shaped across the measured velocities with a minimum power velocity of 0.9 m/s and a maximum range velocity of 2.25 m/s. Migrating H. sara did not compensate for crosswind drift. Changes in airspeed with tailwind drift were consistent with the null hypothesis that H. sara did not compensate for tailwind drift, but they were also not significantly different from those predicted to maximize the migratory range of the insects.

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Mayfly larvae generally are aquatic, but some madicolous taxa (i.e., living in thin water films) crawl over rocks within streams and waterfalls. When startled, these larvae can break the water film, jump, and enter an aerial phase of locomotion. Because mayfly larvae have been suggested as potential exemplars for the origin of insect wings as tracheal gills, and furthermore represent the most basal extant lineage of pterygotes, we analyzed jumping behavior and free-fall trajectories for one such species of mayfly (Myobaetis ellenae, Baetidae) in Costa Rica. Jumping was commonplace in this taxon, but was undirected and was characterized by body spinning at high angular velocities. No aerodynamic role for the tracheal gills was evident. By contrast, jumping by a sympatric species of bristletail (Meinertellus sp., Archaeognatha) consistently resulted in head-first and stable body postures during aerial translation. Although capable of intermittently jumping into the air, the mayfly larvae could neither control nor target their aerial behavior. By contrast, a stable body posture during jumps in adult bristletails, together with the demonstrated capacity for directed aerial descent in arboreal representatives of this order, support ancestrally terrestrial origins for insect flight within the behavioral context of either jumping or falling from heights.

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Gliding ants avoid predatory attacks and potentially mortal consequences of dislodgement from rainforest canopy substrates by directing their aerial descent towards nearby tree trunks. The ecologically relevant measure of performance for gliding ants is the ratio of net horizontal to vertical distance traveled over the course of a gliding trajectory, or glide index. To study variation in glide index, we measured three-dimensional trajectories of Cephalotes atratus ants gliding in natural rainforest habitats. We determined that righting phase duration, glide angle, and path directness all significantly influence variation in glide index. Unsuccessful landing attempts result in the ant bouncing off its target and being forced to make a second landing attempt. Our results indicate that ants are not passive gliders and that they exert active control over the aerodynamic forces they experience during their descent, despite their apparent lack of specialized control surfaces.

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The tempo of species diversification in large clades can reveal fundamental evolutionary mechanisms that operate on large temporal and spatial scales. Hummingbirds have radiated into a diverse assemblage of specialized nectarivores comprising 338 species, but their evolutionary history has not, until now, been comprehensively explored. We studied hummingbird diversification by estimating a time-calibrated phylogeny for 284 hummingbird species, demonstrating that hummingbirds invaded South America by ∼22 million years ago, and subsequently diversified into nine principal clades (see [5-7]). Using ancestral state reconstruction and diversification analyses, we (1) estimate the age of the crown-group hummingbird assemblage, (2) investigate the timing and patterns of lineage accumulation for hummingbirds overall and regionally, and (3) evaluate the role of Andean uplift in hummingbird speciation. Detailed analyses reveal disparate clade-specific processes that allowed for ongoing species diversification. One factor was significant variation among clades in diversification rates. For example, the nine principal clades of hummingbirds exhibit ∼15-fold variation in net diversification rates, with evidence for accelerated speciation of a clade that includes the Bee, Emerald, and Mountain Gem groups of hummingbirds. A second factor was colonization of key geographic regions, which opened up new ecological niches. For example, some clades diversified in the context of the uplift of the Andes Mountains, whereas others were affected by the formation of the Panamanian land bridge. Finally, although species accumulation is slowing in all groups of hummingbirds, several major clades maintain rapid rates of diversification on par with classical examples of rapid adaptive radiation.

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Animal fliers frequently move through a variety of perturbed flows during their daily aerial routines. However, the extent to which these perturbations influence flight control and energetic expenditure is essentially unknown. Here, we evaluate the kinematic and metabolic consequences of flight within variably sized vortex shedding flows using five Anna's hummingbirds feeding from an artificial flower in steady control flow and within vortex wakes produced behind vertical cylinders. Tests were conducted at three horizontal airspeeds (3, 6 and 9 m s(-1)) and using three different wake-generating cylinders (with diameters equal to 38, 77 and 173% of birds' wing length). Only minimal effects on wing and body kinematics were demonstrated for flight behind the smallest cylinder, whereas flight behind the medium-sized cylinder resulted in significant increases in the variances of wingbeat frequency, and variances of body orientation, especially at higher airspeeds. Metabolic rate was, however, unchanged relative to that of unperturbed flight. Hummingbirds flying within the vortex street behind the largest cylinder exhibited highest increases in variances of wingbeat frequency, and of body roll, pitch and yaw amplitudes at all measured airspeeds. Impressively, metabolic rate under this last condition increased by up to 25% compared with control flights. Cylinder wakes sufficiently large to interact with both wings can thus strongly affect stability in flight, eliciting compensatory kinematic changes with a consequent increase in flight metabolic costs. Our findings suggest that vortical flows frequently encountered by aerial taxa in diverse environments may impose substantial energetic costs.

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Animals that sustain high levels of aerobic activity under hypoxic conditions (e.g., birds that fly at high altitude) face the physiological challenge of jointly optimizing blood-O2 affinity for O2 loading in the pulmonary circulation and O2 unloading in the systemic circulation. At high altitude, this challenge is especially acute for small endotherms like hummingbirds that have exceedingly high mass-specific metabolic rates. Here we report an experimental analysis of hemoglobin (Hb) function in South American hummingbirds that revealed a positive correlation between Hb-O2 affinity and native elevation. Protein engineering experiments and ancestral-state reconstructions revealed that this correlation is attributable to derived increases in Hb-O2 affinity in highland lineages, as well as derived reductions in Hb-O2 affinity in lowland lineages. Site-directed mutagenesis experiments demonstrated that repeated evolutionary transitions in biochemical phenotype are mainly attributable to repeated amino acid replacements at two epistatically interacting sites that alter the allosteric regulation of Hb-O2 affinity. These results demonstrate that repeated changes in biochemical phenotype involve parallelism at the molecular level, and that mutations with indirect, second-order effects on Hb allostery play key roles in biochemical adaptation.

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Hummingbirds (family Trochilidae) represent an extreme outcome in vertebrate physiological design and are the only birds capable of sustained hovering. The giant hummingbird (Patagona gigas) is the largest trochilid, with a mass of ~20 g, and is found over an altitudinal range from 0 to 4,500 m above sea level. We report here measurements of daily, basal, and hovering rates of oxygen consumption in the giant hummingbird; compare these values with data from smaller hummingbirds; and assess overall metabolic and allometric limits to trochilid body size. The sustained metabolic scope (i.e., the ratio of daily energy expenditure to basal metabolic rate) in the giant hummingbird is higher than that in smaller hummingbirds but lies below a proposed theoretical maximum value for endotherms. Scaling exponents in the allometric relationships for different modes of energetic expenditure were comparable, suggesting that the giant hummingbird, although a clear outlier in terms of body size, does not obviously deviate from metabolic relationships derived from other trochilid taxa.

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In contrast to the patagial membranes of gliding vertebrates, the aerodynamic surfaces used by falling wingless ants to direct their aerial descent are unknown. We conducted ablation experiments to assess the relative contributions of the hindlegs, midlegs and gaster to gliding success in workers of the Neotropical arboreal ant Cephalotes atratus (Hymenoptera: Formicidae). Removal of hindlegs significantly reduced the success rate of directed aerial descent as well as the glide index for successful flights. Removal of the gaster alone did not significantly alter performance relative to controls. Equilibrium glide angles during successful targeting to vertical columns were statistically equivalent between control ants and ants with either the gaster or the hindlegs removed. High-speed video recordings suggested possible use of bilaterally asymmetric motions of the hindlegs to effect body rotations about the vertical axis during targeting manoeuvre. Overall, the control of gliding flight was remarkably robust to dramatic anatomical perturbations, suggesting effective control mechanisms in the face of adverse initial conditions (e.g. falling upside down), variable targeting decisions and turbulent wind gusts during flight.

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Migrating insects use their sensory systems to acquire local and global cues about their surroundings. Previous research on tethered insects suggests that, in addition to vision and cephalic bristles, insects use antennal mechanosensory feedback to maintain their airspeeds. Owing to the large displacements of migratory insects and difficulties inherent in tracking single individuals, the roles of these sensory inputs have never been tested in freely migrating insects. We tracked individual uraniid moths (Urania fulgens) as they migrated diurnally over the Panama Canal, and measured airspeeds and orientation for individuals with either intact or amputated flagella. Consistent with prior observations that antennal input is necessary for flight control, 59 per cent of the experimental moths could not fly after flagella amputation. The remaining fraction (41%) was flight-capable and maintained its prior airspeeds despite severe reduction in antennal input. Thus, maintenance of airspeeds may not involve antennal input alone, and is probably mediated by other modalities. Moths with amputated flagella could not recover their proper migratory orientations, suggesting that antennal integrity is necessary for long-distance navigation.

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