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Prey often rely on multiple defences against predators, such as flight speed, attack deflection from vital body parts, or unpleasant taste, but our understanding on how often and why they are co-exhibited remains limited. Eudaminae skipper butterflies use fast flight and mechanical defences (hindwing tails), but whether they use other defences like unpalatability (consumption deterrence) and how these defences interact have not been assessed. We tested the palatability of 12 abundant Eudaminae species in Peru, using training and feeding experiments with domestic chicks. Further, we approximated the difficulty of capture based on flight speed and quantified it by wing loading. We performed phylogenetic regressions to find any association between multiple defences, body size, and habitat preference. We found a broad range of palatability in Eudaminae, within and among species. Contrary to current understanding, palatability was negatively correlated with wing loading, suggesting that faster butterflies tend to have lower palatability. The relative length of hindwing tails did not explain the level of butterfly palatability, showing that attack deflection and consumption deterrence are not mutually exclusive. Habitat preference (open or forested environments) did not explain the level of palatability either, although butterflies with high wing loading tended to occupy semi-closed or closed habitats. Finally, the level of unpalatability in Eudaminae is size dependent. Larger butterflies are less palatable, perhaps because of higher detectability/preference by predators. Altogether, our findings shed light on the contexts favouring the prevalence of single versus multiple defensive strategies in prey., (© The Author(s) 2024. Published by Oxford University Press on behalf of the European Society of Evolutionary Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Coffee berry borer (Hypothenemus hampei Ferrari) (Coleoptera: Curculionidae) is the most damaging insect pest of coffee worldwide, causing significant losses in coffee yields and quality. Knowledge of vertical and temporal flight patterns in coffee berry borer could be used to optimize spray timing and precision targeting of areas within the coffee tree, which may be more susceptible. In the present study, we estimated the vertical distribution of coffee berry borer females using traps set at 1-m intervals up to 5 m in height. We also quantified coffee berry borer infestation in the low, mid, and high canopy and documented fruit availability. Temporal flight patterns were estimated using timer traps, and correlation analyses were conducted to determine the relationship between the timing of daily flight and weather variables. Across the 4 study sites, we observed that 77%-84% of the trap catch was at 1 m, 11%-20% was at 2 m, and 1%-4% was at 3-5 m in height. Fruit infestation was significantly higher in the low branches (35%) relative to the high branches (17%). Flight height remained the same year-round, regardless of fruit availability. Coffee berry borer flew in low numbers during the day and night but peaked from 12 to 4 PM. Daily flight was positively correlated with an increase in air temperature and wind speed and negatively correlated with relative humidity. Findings from this study suggest that pesticide sprays should target low- to mid-level branches at 1-2 m in height and aim to be conducted in the early afternoon when coffee berry borer are actively flying and most vulnerable to chemical controls., (© The Author(s) 2024. Published by Oxford University Press on behalf of Entomological Society of America.)

Flight behavior is an important component to understand in the context of pest management. However, because of their small size, little is known about the flight capacity of most stored-product insects, and when a flight has been assessed, it usually consists of a propensity for initiating flight. Despite a priori expectations of the importance of flight for moths, there are no data about the flight capacity and little on the flight behavior of the Mediterranean flour moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae). As a result, the objective of the current study was to (i) characterize the baseline flight capacity of E. kuehniella and (ii) determine how flight capacity is affected by the presence of kairomonal, pheromonal, or no stimuli. We found adult E. kuehniella flew a mean of 24-34 km in a 24-h period, and the distance flown per bout increased from 91 to 207 m in the presence of pheromones but decreased to 41 m when food was nearby compared to a negative control. The total number of flight bouts was 1.6-fold higher in the presence of pheromone compared to the negative control, but E. kuehniella flew significantly slower with pheromone and food cues present, suggesting they may be exhibiting an optimal foraging strategy. Our data on flight capacity results in qualitatively and quantitatively different conclusions about flight than those conclusions formed if only flight initiation is considered. Overall, this novel information is useful for understanding the spread within facilities and in the landscape (between facilities), as well as parameterizing ecological modeling., (Published by Oxford University Press on behalf of Entomological Society of America 2024.)

Birds and bats have long lifespans relative to their body size compared with non-flying animals. However, the genomic basis associated with longer lifespan of flying species despite their higher metabolism was unclear. In this study, we hypothesized that genes involved in the regulation of metabolism and lifespan changed with the acquisition of flight and searched for genes that show specific evolutionary patterns in flying species. As a result, we identified several genes that show different evolutionary rates in bird and bat lineages. Genes in pathways involved in lifespan regulation were conserved in birds, while they evolved at an accelerated rate in bats. We also searched for genes in which convergent amino acid substitutions occurred in birds and bats and found such substitutions in genes involved in cancer, reactive oxygen species control and immunity. Our study revealed genomic changes associated with the acquisition of flight in birds and bats and suggested that multiple genes involved in the regulation of lifespan and metabolism support both high metabolism and longevity in flying species.

During spring migration, nocturnal migrants attempt to minimize their travel time to reach their breeding grounds early. However, how they behave and respond to unfavourable conditions during their springtime travels is much less understood. In this study, we reveal the effects of atmospheric factors on nocturnal bird migration under adverse conditions during spring and autumn, based on one of the most detailed bird migration studies globally, using radar data from 13 deployments over a period of seven years (2014-2020) in the Levant region. Using ERA5 reanalysis data, we found that migratory birds maintain similar ground speeds in both autumn and spring migrations, but during spring, when encountering unfavourable winds, they put more effort into maintaining their travel speed by increasing self-powered airspeed by 18%. Moreover, we report for the first time that spring migrants showed less selectivity to wind conditions and migrated even under unfavourable headwind and crosswind conditions. Interestingly, we discovered that temperature was the most important weather parameter, such that warm weather substantially increased migration intensities in both seasons. Our results enhance our understanding of bird migration over the Levant region, one of the world's largest and most important migration flyways, and the factors controlling it. This information is essential for predicting bird migration, which-especially under the ongoing anthropogenic changes-is of high importance.

Flight plays a crucial role in the fitness of insect pollinators, such as bumblebees. Despite their relatively large body size compared with their wings, bumblebees can fly under difficult ambient conditions, such as cooler temperatures. While their body size is often positively linked to their foraging range and flight ability, the influence of age remains less explored. Here, we studied the flight performance (distance, duration and speed) of ageing bumblebee workers using tethered flight mills. Additionally, we measured their intertegular distance and dry mass as proxies for their body size. We found that the flight distance and duration were predominantly influenced by age, challenging assumptions that age does not play a key role in foraging and task allocation. From the age of 7 to 14 days, flight distance and duration increased sixfold and fivefold, respectively. Conversely, the body size primarily impacted the maximum and average flight speed of workers. Our findings indicate that age substantially influences the flight distance and duration in bumblebee workers, affecting foraging performance and potentially altering task allocation strategies. This underscores the importance of considering individual age and physiological changes alongside body size/mass in experiments involving bumblebee workers.

In the current biodiversity crisis, populations of many species have alarmingly declined, and insects are no exception to this general trend. Biodiversity monitoring has become an essential asset to detect biodiversity change but remains patchy and challenging for organisms that are small, inconspicuous or make (nocturnal) long-distance movements. Radars are powerful remote-sensing tools that can provide detailed information on intensity, timing, altitude and spatial scale of aerial movements and might therefore be particularly suited for monitoring aerial insects and their movements. Importantly, they can contribute to several essential biodiversity variables (EBVs) within a harmonized observation system. We review existing research using small-scale biological and weather surveillance radars for insect monitoring and outline how the derived measures and quantities can contribute to the EBVs 'species population', 'species traits', 'community composition' and 'ecosystem function'. Furthermore, we synthesize how ongoing and future methodological, analytical and technological advancements will greatly expand the use of radar for insect biodiversity monitoring and beyond. Owing to their long-term and regional-to-large-scale deployment, radar-based approaches can be a powerful asset in the biodiversity monitoring toolbox whose potential has yet to be fully tapped. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Radar networks hold great promise for monitoring population trends of migrating insects. However, it is important to elucidate the nature of responses to environmental cues. We use data from a mini-network of vertical-looking entomological radars in the southern UK to investigate changes in nightly abundance, flight altitude and behaviour of insect migrants, in relation to meteorological and celestial conditions. Abundance of migrants showed positive relationships with air temperature, indicating that this is the single most important variable influencing the decision to initiate migration. In addition, there was a small but significant effect of moonlight illumination, with more insects migrating on full moon nights. While the effect of nocturnal illumination levels on abundance was relatively minor, there was a stronger effect on the insects' ability to orientate close to downwind: flight headings were more tightly clustered on nights when the moon was bright and when cloud cover was sparse. This indicates that nocturnal illumination is important for the navigational mechanisms used by nocturnal insect migrants. Further, our results clearly show that environmental conditions such as air temperature and light levels must be considered if long-term radar datasets are to be used to assess changing population trends of migrants. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Georgina Mills discusses new research looking at how hummingbirds use somatosensation to adjust their flight., (© 2024 British Veterinary Association.)

Potato leafhopper (PLH), Empoasca fabae Harris (Hemiptera: Cicadellidae), is an economic pest of a variety of crops that migrates between overwintering sites in the southern United States and northern breeding grounds. Since 2005, the Midwest Suction Trap Network (STN) has monitored the magnitude and timing of aerially dispersing aphids' activity, but the potential of the network to monitor other taxa is only beginning to be explored. Here, we use the Midwest STN to examine how the magnitude and timing of PLH activity vary with weather, cropland cover, and time of year. We found that weekly PLH activity increased early in the season (May-June) with increasing degree day accumulation and decreased mid-season (July-August) with increasing occurrence of rain. The first detections occurred earlier in southern latitudes, while the last detections occurred sooner, when there was more surrounding potato land cover, and later over time between 2018 and 2021 and in southern latitudes. PLH activity was thus longer in duration in southern latitudes and has continued to extend later into the year overall. Resolving uncertainty about how well the Midwest STN captures migratory activity and how closely suction trap detections reflect local population densities in crop fields remain important research priorities before the potential of the Midwest STN for PLH monitoring can be realized. Still, observed patterns suggest that PLH could increase in economic importance as insects disperse over larger portions of the growing season in the warming, agriculturally productive US Midwest and that the STN can become a useful tool to monitor these changes., (© The Author(s) 2024. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

Halteres are multifunctional mechanosensory organs unique to the true flies (Diptera). A set of reduced hindwings, the halteres beat at the same frequency as the lift-generating forewings and sense inertial forces via mechanosensory campaniform sensilla. Though haltere ablation makes stable flight impossible, the specific role of wing-synchronous input has not been established. Using small iron filings attached to the halteres of tethered flies and an alternating electromagnetic field, we experimentally decoupled the wings and halteres of flying Drosophila and observed the resulting changes in wingbeat amplitude and head orientation. We find that asynchronous haltere input results in fast amplitude changes in the wing (hitches), but does not appreciably move the head. In multi-modal experiments, we find that wing and gaze optomotor responses are disrupted differently by asynchronous input. These effects of wing-asynchronous haltere input suggest that specific sensory information is necessary for maintaining wing amplitude stability and adaptive gaze control.

In autumn 1950 David and Elizabeth Lack chanced upon a huge migration of insects and birds flying through the Pyrenean Pass of Bujaruelo, from France into Spain, later describing the spectacle as combining both grandeur and novelty. The intervening years have seen many changes to land use and climate, posing the question as to the current status of this migratory phenomenon. In addition, a lack of quantitative data has prevented insights into the ecological impact of this mass insect migration and the factors that may influence it. To address this, we revisited the site in autumn over a 4 year period and systematically monitored abundance and species composition of diurnal insect migrants. We estimate an annual mean of 17.1 million day-flying insect migrants from five orders (Diptera, Hymenoptera, Hemiptera, Lepidoptera and Odonata) moving south, with observations of southward 'mass migration' events associated with warmer temperatures, the presence of a headwind, sunlight, low windspeed and low rainfall. Diptera dominated the migratory assemblage, and annual numbers varied by more than fourfold. Numbers at this single site hint at the likely billions of insects crossing the entire Pyrenean mountain range each year, and we highlight the importance of this route for seasonal insect migrants.

The use of Meliponini for crop pollination in protected environments is practically non-existent. One of the reasons is the difficulty of acclimatizing Meliponini to the temperature and light conditions inside greenhouses. We investigated how covering materials used in greenhouses, which filter different intensities of ultraviolet (UV) light, affect the foraging behaviors, flight orientation, attraction to walls and ceilings, and mortality of Scaptotrigona cf. postica (Letreille), Frieseomelitta varia (Lepeletier), and Melipona quadrifasciata (Lepeletier). The experiments were conducted in 5.3 m 3 arenas covered with four types of plastic films that do not polarize sunlight, with UV transmittance levels ranging from 0.1 to 54%, compared to a transparent glass control. The temperature inside the arenas varied between treatments, from 27 ± 3°C to 31 ± 2°C. All three species collected resources and returned to the colony, regardless of the covering material. However, the proportion of this behavior, the number of bees attracted to the ceiling and wall, and mortality varied among treatments and/or throughout the confinement days for each species. Melipona quadrifasciata and F. varia acclimatized better to the confined environments than S. cf. postica and showed consistent resource collection behavior throughout the confinement days in all tested materials, except for the one that filtered around 90% of UV. In all three species, the mortality gradually decreased throughout the confinement days. The results indicate that the choice of covering material, considering its optical characteristics, can be crucial to ensure greater effectiveness of the pollination services provided by stingless bees in protected systems., (© 2024. Sociedade Entomológica do Brasil.)

Optimal migration theory prescribes adaptive strategies of energy, time or mortality minimization. To test alternative hypotheses of energy- and time-minimization migration we used multisensory data loggers that record time-resolved flight activity and light for positioning by geolocation in a long-distance migratory shorebird, the little ringed plover, Charadrius dubius . We could reject the hypothesis of energy minimization based on a relationship between stopover duration and subsequent flight time as predicted for a time minimizer. We found seasonally diverging slopes between stopover and flight durations in relation to the progress (time) of migration, which follows a time-minimizing policy if resource gradients along the migration route increase in autumn and decrease in spring. Total flight duration did not differ significantly between autumn and spring migration, although spring migration was 6% shorter. Overall duration of autumn migration was longer than that in spring, mainly owing to a mid-migration stop in most birds, when they likely initiated moult. Overall migration speed was significantly different between autumn and spring. Migratory flights often occurred as runs of two to seven nocturnal flights on adjacent days, which may be countering a time-minimization strategy. Other factors may influence a preference for nocturnal migration, such as avoiding flight in turbulent conditions, heat stress and diurnal predators.

An insect's wingbeat frequency is a critical determinant of its flight performance and varies by multiple orders of magnitude across Insecta. Despite potential energetic benefits for an insect that matches its wingbeat frequency to its resonant frequency, recent work has shown that moths may operate off their resonant peak. We hypothesized that across species, wingbeat frequency scales with resonance frequency to maintain favourable energetics, but with an offset in species that use frequency modulation as a means of flight control. The moth superfamily Bombycoidea is ideal for testing this hypothesis because their wingbeat frequencies vary across species by an order of magnitude, despite similar morphology and actuation. We used materials testing, high-speed videography and a model of resonant aerodynamics to determine how components of an insect's flight apparatus (stiffness, wing inertia, muscle strain and aerodynamics) vary with wingbeat frequency. We find that the resonant frequency of a moth correlates with wingbeat frequency, but resonance curve shape (described by the Weis-Fogh number) and peak location vary within the clade in a way that corresponds to frequency-dependent biomechanical demands. Our results demonstrate that a suite of adaptations in muscle, exoskeleton and wing drive variation in resonant mechanics, reflecting potential constraints on matching wingbeat and resonant frequencies.

Anthropogenic activities have decimated the Atlantic Forest domain (AF) and increased the pressure on freshwater biota, such as Trichoptera, which is the most affected order by the current insect decline. Adult mobility is crucial for the colonisation of new environments unconnected by water sources. In this article, we describe the assemblage of caddisflies in a preserved AF fragment related to their functional feeding group and provide empirical data on the patterns of horizontal and vertical flight. Adults were collected using white sheet and light attraction traps, placed at different distances and heights from a stream in Pernambuco, Brazil. A total of 2934 specimens of 15 species from five families were collected, mostly collector-filterers. Horizontal flight was limited, with 80% of the abundance concentrated up to 20 m. Vertical stratification was also concentrated at lower heights. A female-biased proportion was observed at higher strata. The richness and abundance of species decreased with increasing distances and heights from the stream. Overall, Chimarra sp. and Macrostemum scharfi were the dominant species. Trichoptera is a key taxon used as a biological indicator of water quality, and here, knowledge on the diversity and flight patterns of adults is expanded. At the risk of intensive pollution of rivers in Atlantic forests, data on the adult dispersal can be incorporated in the assessment of endangerment status and in conservation strategies., (© 2024. Sociedade Entomológica do Brasil.)

Osmia solitary bees are important pollinators of various crops worldwide. Refrigeration has been widely used to synchronize the emergence time of Osmia species from cocoons with the blooming time of different crops, but the fitness of Osmia after refrigeration remains unknown. Here, the effects of long-term refrigeration at 0 °C on the vitality, flight ability, and metabolism of Osmia excavata, which is known as the "king of pollination" in China, were studied. The survival rate (>90% before 120 d), weight loss rate (<15% after 170 d), and mean flight speed of O. excavata were not greatly affected after long-term refrigeration. The content of fats, which have antifreeze and energy storage properties, was not significantly altered in O. excavata before 130 d of refrigeration, which might explain why the survival rates and flight speed of O. excavata remained high after long-term refrigeration. However, the flight duration and distance decreased significantly (P < 0.05), and both were positively correlated with the reduced trehalose levels in O. excavata (r = [+0.69] - [+0.71]; P < 0.05). Overall, these findings indicate that the pollination potential of O. excavata for various crops with different flowering periods is high after long-term refrigeration; however, long-term refrigeration may decrease pollination efficiency. Our findings highlight new research directions that could improve the ecological service function of refrigerated O. excavata., (© 2023 Institute of Zoology, Chinese Academy of Sciences.)

Insect flight is a complex trait involved in different behaviors, from the search for sexual partners, food, or breeding sites. Many studies have postulated the adaptive advantages of certain morphological traits in relation to increased flight capacity, such as low values of wing loading or high values of wing:thorax ratio and wing-aspect ratio. However, few studies have evaluated the relationship between variables related to flight and morphological traits in Drosophila. This work aimed to study morphological traits in males and females of two pairs of sibling species: Drosophila buzzatii Patterson and Wheeler-Drosophila koeferae Fontdevila and Wasserman, and Drosophila melanogaster Meigen-Drosophila simulans Sturtevant, and to analyze its relationship with flight. We detected the highest proportion of flight time in D. koepferae and D. simulans compared to D. buzzatii and D. melanogaster, respectively. Our results also revealed sexual dimorphism, with males exhibiting a higher proportion of flight time than females. Surprisingly, we did not find a general pattern to explain the relationship between morphology and the proportion of flight time because associations varied depending upon the analyses (considering all groups together or each sex-species combination separately). Moreover, these associations explained a low percentage of variation, suggesting that other nonmorphological components related to flight, such as physiological variables, should be taken into account. This work allowed us to show the variability and complexity of an aspect of flight, suggesting that the adaptive role of the morphological traits studied might have been overestimated., (© 2023 Institute of Zoology, Chinese Academy of Sciences.)

Background: Flight can drastically enhance dispersal capacity and is a key trait defining the potential of exotic insect species to spread and invade new habitats. The phytophagous European spongy moths (ESM, Lymantria dispar dispar) and Asian spongy moths (ASM; a multi-species group represented here by L. d. asiatica and L. d. japonica), are globally invasive species that vary in adult female flight capability-female ASM are typically flight capable, whereas female ESM are typically flightless. Genetic markers of flight capability would supply a powerful tool for flight profiling of these species at any intercepted life stage. To assess the functional complexity of spongy moth flight and to identify potential markers of flight capability, we used multiple genetic approaches aimed at capturing complementary signals of putative flight-relevant genetic divergence between ESM and ASM: reduced representation genome-wide association studies, whole genome sequence comparisons, and developmental transcriptomics. We then judged the candidacy of flight-associated genes through functional analyses aimed at addressing the proximate demands of flight and salient features of the ecological context of spongy moth flight evolution., Results: Candidate gene sets were typically non-overlapping across different genetic approaches, with only nine gene annotations shared between any pair of approaches. We detected an array of flight-relevant functional themes across gene sets that collectively suggest divergence in flight capability between European and Asian spongy moth lineages has coincided with evolutionary differentiation in multiple aspects of flight development, execution, and surrounding life history. Overall, our results indicate that spongy moth flight evolution has shaped or been influenced by a large and functionally broad network of traits., Conclusions: Our study identified a suite of flight-associated genes in spongy moths suited to exploration of the genetic architecture and evolution of flight, or validation for flight profiling purposes. This work illustrates how complementary genetic approaches combined with phenotypically targeted functional analyses can help to characterize genetically complex traits., (© 2024. His Majesty the King in Right of Canada as represented by the Minister of Natural Resources and Julien Prunier, Richard Hamelin, Francois Oliver Hebert, Roger C. Levesque, Ilga Porth. Parts of this work were authored by US Federal Government authors and are not under copyright protection in the US; foreign copyright protection may apply.)

Although diurnality is widespread across Lepidoptera and has evolved many times independently, its causes and ecological implications are yet poorly understood. The "Salient Aroma Hypothesis" (SAH) postulates that diurnal insect herbivores are overall more specialized in dietary breadth than species active at night. It is furthermore assumed that diurnality evolved more frequently in species that live in cooler environments. Using European geometrid moths as a model group, we tested whether diurnal activity in adults is associated with an increased larval dietary breadth as predicted by the SAH. We further investigated whether species that exclusively occur in colder regions or whose flight period is restricted to cool seasons are more likely to exhibit a diurnal flight activity. Contrary to expectation, we found no consistent differences in larval dietary breadth between diurnal and nocturnal species, and thus no support for the SAH. Diurnal activity occurred more frequently in species restricted to cold regions but not in species restricted to cool seasons. We conclude that diurnality could serve as an advantageous adaptation in cold environments, depending on further factors such as resource availability or predation pressure, but has no immediate consequences for larval dietary breadth., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE). All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

We present a draft genome of the little bush moa ( Anomalopteryx didiformis )-one of approximately nine species of extinct flightless birds from Aotearoa, New Zealand-using ancient DNA recovered from a fossil bone from the South Island. We recover a complete mitochondrial genome at 249.9× depth of coverage and almost 900 megabases of a male moa nuclear genome at ~4 to 5× coverage, with sequence contiguity sufficient to identify more than 85% of avian universal single-copy orthologs. We describe a diverse landscape of transposable elements and satellite repeats, estimate a long-term effective population size of ~240,000, identify a diverse suite of olfactory receptor genes and an opsin repertoire with sensitivity in the ultraviolet range, show that the wingless moa phenotype is likely not attributable to gene loss or pseudogenization, and identify potential function-altering coding sequence variants in moa that could be synthesized for future functional assays. This genomic resource should support further studies of avian evolution and morphological divergence.

Challenges imposed by geographical barriers during migration are selective agents for animals. Juvenile soaring landbirds often cross large water bodies along their migratory path, where they lack updraft support and are vulnerable to harsh weather. However, the consequences of inexperience in accomplishing these water crossings remain largely unquantified. To address this knowledge gap, we tracked the movements of juvenile and adult black kites Milvus migrans over the Strait of Gibraltar using high-frequency tracking devices in variable crosswind conditions. We found that juveniles crossed under higher crosswind speeds and at wider sections of the strait compared with adults during easterly winds, which represent a high risk owing to their high speed and steady direction towards the Atlantic Ocean. Juveniles also drifted extensively with easterly winds, contrasting with adults who strongly compensated for lateral displacement through flapping. Age differences were inconspicuous during winds with a west crosswind speed component, as well as for airspeed modulation in all wind conditions. We suggest that the suboptimal sea-crossing behaviour of juvenile black kites may impact their survival rates, either by increasing chances of drowning owing to exhaustion or by depleting critical energy reserves needed to accomplish their first migration.

Flying animals often encounter winds during visually guided landings. However, how winds affect their flight control strategy during landing is unknown. Here, we investigated how sidewind affects the landing performance and sensorimotor control of foraging bumblebees (Bombus terrestris). We trained bumblebees to forage in a wind tunnel, and used high-speed stereoscopic videography to record 19,421 landing maneuvers in six sidewind speeds (0 to 3.4 m s-1), which correspond to winds encountered in nature. Bumblebees landed less often in higher windspeeds, but the landing durations from free flight were not increased by wind. By testing how bumblebees adjusted their landing control to compensate for adverse effects of sidewind on landing, we showed that the landing strategy in sidewind resembled that in still air, but with important adaptations. Bumblebees landing in a sidewind tended to drift downwind, which they controlled for by performing more hover maneuvers. Surprisingly, the increased hover prevalence did not increase the duration of free-flight landing maneuvers, as these bumblebees flew faster towards the landing platform outside the hover phases. Hence, by alternating these two flight modes along their flight path, free-flying bumblebees negated the adverse effects of high windspeeds on landing duration. Using control theory, we hypothesize that bumblebees achieve this by integrating a combination of direct aerodynamic feedback and a wind-mediated mechanosensory feedback control, with their vision-based sensorimotor control loop. The revealed landing strategy may be commonly used by insects landing in windy conditions, and may inspire the development of landing control strategies onboard autonomously flying robots., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

The silent flight of barn owls is associated with wing and feather specialisations. Three special features are known: a serrated leading edge that is formed by free-standing barb tips which appears as a comb-like structure, a soft dorsal surface, and a fringed trailing edge. We used a model of the leading edge comb with 3D-curved serrations that was designed based on 3D micro-scans of rows of barbs from selected barn-owl feathers. The interaction of the flow with the serrations was measured with Particle-Image-Velocimetry in a flow channel at uniform steady inflow and was compared to the situation of inflow with freestream turbulence, generated from the turbulent wake of a cylinder placed upstream. In steady uniform flow, the serrations caused regular velocity streaks and a flow turning effect. When vortices of different size impacted the serrations, the serrations reduced the flow fluctuations downstream in each case, exemplified by a decreased root-mean-square value of the fluctuations in the wake of the serrations. This attenuation effect was stronger for the spanwise velocity component, leading to an overall flow homogenization. Our findings suggest that the serrations of the barn owl provide a passive flow control leading to reduced leading-edge noise when flying in turbulent environments., (© 2024 IOP Publishing Ltd.)

Some insects have a frustrating knack for avoiding a swatter. A new study shows that mosquitos not only evade the visual image of the looming threat, they also surf the wave of air the swatter creates., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

The scaling of the energetic cost of locomotion with body mass is well documented at the interspecific level. However, methodological restrictions limit our understanding of the scaling of flight metabolic rate (MR) in free-flying insects. This is particularly true at the intraspecific level, where variation in body mass and flight energetics may have direct consequences for the fitness of an individual. We applied a 13 C stable isotope method to investigate the scaling of MR with body mass during free-flight in the beetle Batocera rufomaculata. This species exhibits large intraspecific variation in adult body mass as a consequence of the environmental conditions during larval growth. We show that the flight-MR scales with body mass to the power of 0.57, with smaller conspecifics possessing up to 2.3 fold higher mass-specific flight MR than larger ones. Whereas the scaling exponent of free-flight MR was found to be like that determined for tethered-flight, the energy expenditure during free-flight was more than 2.7 fold higher than for tethered-flight. The metabolic cost of flight should therefore be studied under free-flight conditions, a requirement now enabled by the 13 C technique described herein for insect flight., (© 2023 The Authors. Insect Science published by John Wiley & Sons Australia, Ltd on behalf of Institute of Zoology, Chinese Academy of Sciences.)

Bioinspired flapping-wing micro aerial vehicles (FWMAVs) have emerged over the last two decades as a promising new type of robot. Their high thrust-to-weight ratio, versatility, safety, and maneuverability, especially at small scales, could make them more suitable than fixed-wing and multi-rotor vehicles for various applications, especially in cluttered, confined environments and in close proximity to humans, flora, and fauna. Unlike natural flyers, however, most FWMAVs currently have limited take-off and landing capabilities. Natural flyers are able to take off and land effortlessly from a wide variety of surfaces and in complex environments. Mimicking such capabilities on flapping-wing robots would considerably enhance their practical usage. This review presents an overview of take-off and landing techniques for FWMAVs, covering different approaches and mechanism designs, as well as dynamics and control aspects. The special case of perching is also included. As well as discussing solutions investigated for FWMAVs specifically, we also present solutions that have been developed for different types of robots but may be applicable to flapping-wing ones. Different approaches are compared and their suitability for different applications and types of robots is assessed. Moreover, research and technology gaps are identified, and promising future work directions are identified., (Creative Commons Attribution license.)

Insects are excellent at flying in dense vegetation and navigating through other complex spatial environments. This study investigates the strategies used by honeybees ( Apis mellifera ) to avoid collisions with an obstacle encountered frontally during flight. Bees were trained to fly through a tunnel that contained a solitary vertically oriented cylindrical obstacle placed along the midline. Flight trajectories of bees were recorded for six conditions in which the diameter of the obstructing cylinder was systematically varied from 25 mm to 160 mm. Analysis of salient events during the bees' flight, such as the deceleration before the obstacle, and the initiation of the deviation in flight path to avoid collisions, revealed a strategy for obstacle avoidance that is based on the relative retinal expansion velocity generated by the obstacle when the bee is on a collision course. We find that a quantitative model, featuring a controller that extracts specific visual cues from the frontal visual field, provides an accurate characterization of the geometry and the dynamics of the manoeuvres adopted by honeybees to avoid collisions. This study paves the way for the design of unmanned aerial systems, by identifying the visual cues that are used by honeybees for performing robust obstacle avoidance flight.

This paper designs a bionic aircraft model equipped with multiple degrees of freedom to study the inertial force equation and the aerodynamic interaction between its forewings and hindwings. Each wing's phase difference angle (PDA) and stroke plane angle (SPA) are independently adjustable. Employing the kinematic equation of a single wing, we establish a model for the inertial force of the four-wing aircraft, validating its accuracy through experimental comparisons. Furthermore, we analyze various combinations of PDA and SPA parameters for the fore- and hindwings to ascertain the most efficient aerodynamic motion modes. Our findings reveal that aerodynamic interference between the fore- and hindwings tends to be unfavorable, predominantly due to the hindwings being exposed to the wake generated by the forewings, hindering their lift-capturing ability. Nevertheless, a specific PDA = 270° (forewing ahead of hindwing 270°) helps mitigate this interference across a wider range of SPA. Interestingly, when the stroke plane aligns parallel to the horizontal direction, asynchronous flapping of the fore- and hindwings, forming a lift mechanism akin to clap-and-fling wings, positively impacts lift. Consequently, staggered flapping of the fore- and hindwings reduces fuselage jitter and alleviates aerodynamic interference through specialized PDA, resulting in a temporary lift enhancement. The purpose of this study is to provide theoretical support for the longitudinal attitude control of four-wing aircraft., (© 2024 IOP Publishing Ltd.)

As a fundamental ecological aspect of most organisms, locomotor function significantly constrains morphology. At the same time, the evolution of novel locomotor abilities has produced dramatic morphological transformations, initiating some of the most significant diversifications in life history. Despite significant new fossil evidence, it remains unclear whether volant locomotion had a single or multiple origins in pennaraptoran dinosaurs and the volant abilities of individual taxa are controversial. The evolution of powered flight in modern birds involved exaptation of feathered surfaces extending off the limbs and tail yet most studies concerning flight potential in pennaraptorans do not account for the structure and morphology of the wing feathers themselves. Analysis of the number and shape of remex and rectrix feathers across a large dataset of extant birds indicates that the number of remiges and rectrices and the degree of primary vane asymmetry strongly correlate with locomotor ability revealing important functional constraints. Among these traits, phenotypic flexibility varies reflected by the different rates at which morphological changes evolve, such that some traits reflect the ancestral condition, whereas others reflect current locomotor function. While Mesozoic birds and Microraptor have remex morphologies consistent with extant volant birds, that of anchiornithines deviate significantly providing strong evidence this clade was not volant. The results of these analyses support a single origin of dinosaurian flight and indicate the early stages of feathered wing evolution are not sampled by the currently available fossil record., Competing Interests: Competing interests statement:The authors declare no competing interest.

To better understand the secret of natural flying vertebrates such as how humming-birds twist their wings to achieve superb flight ability, we presented a numerical investigation of dynamic twisting based on a hummingbird-like flapping wing model. Computational fluid dynamic simulations were performed to examine the effects of dynamic twisting on the unsteady flow field, the generation of instantaneous aerodynamic forces, and the time-averaged aerodynamic performance. This research reveals the details of leading-edge vortices (LEVs) and the underlying mechanisms behind the positive effects of wing torsion. The results demonstrated that wing torsion can effectively maintain the favorable distribution of effective angle of attack along the wing spanwise, resulting in a higher time-averaged thrust and vertical force. Further, the proper parameters of dynamic twisting can also improve the propulsive efficiency in forward flight. Dynamic twisting also showed a superior ability in controlling the airflow separation over the wing surface and maintaining the stability of the LEV. The amplitudes of effective angle of attack associated with the highest peak thrust and the maximum thrust-to-power at different advanced ratios were also explored, and it was found that the amplitudes decrease with increasing advanced ratio. To improve the efficiency during larger advanced ratio, specific modifications to the pitching of the wing were proposed in this work. The research in this paper has promising implications for the bio-inspired flapping wing., (© 2024 IOP Publishing Ltd.)

Enhancing the aerodynamic performance of bristled wings is an important topic for small flying robotics. This paper numerically investigates this situation at very low Reynolds numbers by using elliptic cylinders as the bristles instead of circular cylinders. Optimal configuration of the bristled wing with five elliptic cylinders is obtained, which corresponds to the maximum lift. The results show that, compared with the case of circular cylindrical bristles, the aerodynamic performance of the elliptical bristles can be enhanced effectively. The enhancement can be more significant as the aspect ratio of the ellipses increases and the gap width decreases. The bristled wing generates more lift compared to a flat-plate wing with a length five times that of the major axis of an ellipse. For the cases that the attack angle α for the whole wing is equal to those for the elliptical bristles θ , the optimal attack angle for ellipses maximizing the total lift force of the five-bristle model is between 40° and 45°. For α ≠ θ with the Reynold number Re ≪ 0.1, the optimal ellipse attack angle is between 40° and 45°. For α ≠ θ with Re ∼ 1, the optimal ellipse attack angle deviates heavier from the range between 40° and 45° at some α values and reaches approximately 32° at α = 20°. This paper can lay a foundation for optimal design of small flying robotics and enhancement of flow through porous structures in future., (© 2024 IOP Publishing Ltd.)

The femoral lobes of the orchid mantis give this fierce predator a flower-like appearance, but they also assist in gliding, showing that form can match function in more ways than one., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

In this work, we explored a bioinspired method for underwater object sensing based on active proprioception. We investigated whether the fluid flows generated by a robotic flapper, while interacting with an underwater wall, can encode the distance information between the wall and the flapper, and how to decode this information using the proprioception within the flapper. Such touchless wall-distance sensing is enabled by the active motion of a flapping plate, which injects self-generated flow to the fluid environment, thus representing a form of active sensing. Specifically, we trained a long short-term memory (LSTM) neural network to predict the wall distance based on the force and torque measured at the base of the flapping plate. In addition, we varied the Rossby number ( Ro , or the aspect ratio of the plate) and the dimensionless flapping amplitude (A∗) to investigate how the rotational effects and unsteadiness of self-generated flow respectively affect the accuracy of the wall-distance prediction. Our results show that the median prediction error is within 5% of the plate length for all the wall-distances investigated (up to 40 cm or approximately 2-3 plate lengths depending on the Ro ); therefore, confirming that the self-generated flow can enable underwater perception. In addition, we show that stronger rotational effects at lower Ro lead to higher prediction accuracy, while flow unsteadiness (A∗) only has moderate effects. Lastly, analysis based on SHapley Additive exPlanations (SHAP) indicate that temporal features that are most prominent at stroke reversals likely promotes the wall-distance prediction., (Creative Commons Attribution license.)

This paper discusses the wing inertial effects and the important role of the mean stroke angle on the pitch dynamics of hovering insects. The paper also presents a dynamic model appropriate for averaging and discusses the pitch stability results derived from the model. The model is used to predict the body angle of five insect species during hover, which are in good agreement with the available experimental results from different literature. The results suggest that the wing inertial forces have a considerable effect on pitch dynamics of insect flight and should not be ignored in dynamic analysis of hovering insects. The results also suggest that the body of hovering insects can not be vibrationally stabilized in a non-vertical orientation. Instead, the pitch angle of a hovering insect's body is mainly due to a balance of the moment of the insect's weight and the aerodynamic moment due to flapping kinematics with a nonzero mean stroke angle. Experiments with a flapping wing device confirm this results. To clearly explain the used model and clarify the difference between vibrational and non-vibrational stabilization, first this paper discusses the vibrational control of a three-degree-of-freedom force-input pendulum with its pivot moving in a vertical plane., (© 2024. The Author(s).)

Thermal soaring conditions above the sea have long been assumed absent or too weak for terrestrial migrating birds, forcing obligate soarers to take long detours and avoid sea-crossing, and facultative soarers to cross exclusively by costly flapping flight. Thus, while atmospheric convection does develop at sea and is used by some seabirds, it has been largely ignored in avian migration research. Here, we provide direct evidence for routine thermal soaring over open sea in the common crane, the heaviest facultative soarer known among terrestrial migrating birds. Using high-resolution biologging from 44 cranes tracked across their transcontinental migration over 4 years, we show that soaring performance was no different over sea than over land in mid-latitudes. Sea-soaring occurred predominantly in autumn when large water-air temperature difference followed mid-latitude cyclones. Our findings challenge a fundamental migration research paradigm and suggest that obligate soarers avoid sea-crossing not due to the absence or weakness of thermals but due to their low frequency, for which they cannot compensate with prolonged flapping. Conversely, facultative soarers other than cranes should also be able to use thermals over the sea. Marine cold air outbreaks, imperative to global energy budget and climate, may also be important for bird migration.

The evolution of flight is a rare event in vertebrate history, and one that demands functional integration across multiple anatomical/physiological systems. The neuroanatomical basis for such integration and the role that brain evolution assumes in behavioural transformations remain poorly understood. We make progress by (i) generating a positron emission tomography (PET)-based map of brain activity for pigeons during rest and flight, (ii) using these maps in a functional analysis of the brain during flight, and (iii) interpreting these data within a macroevolutionary context shaped by non-avian dinosaurs. Although neural activity is generally conserved from rest to flight, we found significant increases in the cerebellum as a whole and optic flow pathways. Conserved activity suggests processing of self-movement and image stabilization are critical when a bird takes to the air, while increased visual and cerebellar activity reflects the importance of integrating multimodal sensory information for flight-related movements. A derived cerebellar capability likely arose at the base of maniraptoran dinosaurs, where volumetric expansion and possible folding directly preceded paravian flight. These data represent an important step toward establishing how the brain of modern birds supports their unique behavioural repertoire and provide novel insights into the neurobiology of the bird-like dinosaurs that first achieved powered flight.

The agamid lizards of the genus Draco are undoubtedly the most renown reptilian gliders, using their rib-supported patagial wings as lifting surfaces while airborne. Recent investigations into these reptiles highlighted the role of body posture during gliding, however, the aerodynamics of postural changes in Draco remain unclear. Here, we examine the aerodynamics and gliding performances of Draco volans using a numerical approach focusing on three postural changes: wing expansion, body camber, and limb positioning. To this aim, we conducted 70 three-dimensional steady-state computational fluid dynamics simulations of gliding flight and 240 two-dimensional glide trajectory calculations. Our results demonstrate that while airborne, D. volans generates a separated turbulent boundary layer over its wings characterized by a large recirculation cell that is kept attached to the wing surface by interaction with wing-tip vortices, increasing lift generation. This lift generating mechanism may be controlled by changing wing expansion and shape to modulate the generation of aerodynamic force. Furthermore, our trajectory simulations highlight the influence of body camber and orientation on glide range. This sheds light on how D. volans controls its gliding performance, and conforms to the observation that these animals plan their glide paths prior to take off. Lastly, D. volans is mostly neutral in pitch and highly maneuverable, similar to other vertebrate gliders. The numerical study presented here thus provides a better understanding of the lift generating mechanism and the influence of postural changes in flight in this emblematic animal and will facilitate the study of gliding flight in analogous gliding reptiles for which direct observations are unavailable., (Creative Commons Attribution license.)

The four-winged form of dragonfly and damselfly allows them to fly with great agility and endurance, which are accomplished by independently controlling the kinematics of each wing. In this study, we performed numerical simulations of two tandem airfoils oscillating along an inclined stroke plane atRe=157. We investigated the effects of the stroke plane angle (β)of forefoil and hindfoil on the aerodynamic performance of dragonflies (or damselflies) hovering flight. Simulations were conducted for parallel and non-parallel stroke planes of forefoil and hindfoil oscillating with three phase differences:ϕ=0o,90oand180o. For parallel stroke planes, the results show that the total lift increases withβ, whereas the total thrust decreases in same condition. In addition, the total lift and thrust decrease with an increase inϕ. The forefoil performance is affected by the flow induced by hindfoil's leading-edge vortices, and the hindfoil interactions with forefoil's wake vortices significantly affect the hindfoil's performance. The findings demonstrate that non-parallel stroke planes exert detrimental effects on the total lift forϕ=0oand90o. However, forϕ=180o, the lift augmentation of 46% is obtained in the case where the forefoil stroke plane angle is smaller than the hindfoil. The results obtained during this investigation can help in optimizing the wing kinematics during the micro air vehicles development., (© 2024 IOP Publishing Ltd.)

We present models of bat motion derived from radio-tracking data collected over 14 nights. The data presents an initial dispersal period and a return to roost period. Although a simple diffusion model fits the initial dispersal motion we show that simple convection cannot provide a description of the bats returning to their roost. By extending our model to include non-autonomous parameters, or a leap frogging form of motion, where bats on the exterior move back first, we find we are able to accurately capture the bat's motion. We discuss ways of distinguishing between the two movement descriptions and, finally, consider how the different motion descriptions would impact a bat's hunting strategy., (© 2024. The Author(s).)

To glide in forest canopies, arboreal vertebrates evolved various skin-derived aerodynamic structures, such as patagial membranes or webbing, but no comparable structure has been reported from wingless arboreal arthropods. 1 , 2 , 3 Orchid mantises (Hymenopus coronatus) have been traditionally considered a textbook example of flower mimicry for ∼200 years due to their highly expanded, petal-shaped femoral lobes. However, the empirical evidence substantiating the petal-mimicry function of the femoral lobes has not been entirely conclusive. 4 , 5 , 6 Observational and experimental evidence suggests that these lobes do not contribute to flower mimicry for luring pollinators 6 , 7 and likely serve other functions. 7 , 8 After observing their aerial escape initiated with active jumping, we hypothesized that orchid mantises can glide and that their femoral lobes are used for gliding. Through behavioral investigations and morphological analyses, we show that orchid mantis nymphs are excellent gliders, exhibiting the shallowest gliding trajectories observed in terrestrial invertebrates. 9 , 10 , 11 , 12 , 13 The lobe extensions on their femoral segments are cambered airfoils, which increase the mantis projected area by ∼36% and play a vital role in the aerodynamic underpinning of the observed gliding. Despite a 165-fold increase in body mass throughout ontogeny, older female mantis nymphs maintained a persistent gliding capability. We further showed a notable 40%-56% reduction in wing loading attributed to the positive size allometry of these lobes, indicating a clear promotion of gliding throughout ontogeny. This is the first documentation of gliding-adapted "leg wings" in a wingless arthropod. The evolution of such structures is potentially common among arboreal arthropods and demands a systematic re-examination., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)