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Background: Controlling the spread of arboviral diseases remains a considerable challenge due to the rapid development of insecticide resistance in Aedes mosquitoes. This study evaluated the effects of boric acid-containing toxic sugar bait (TSB) on field populations of resistant Aedes aegypti mosquitoes. In addition, this study examined the flight activity and wing beat frequency and amplitude of males and the flight activity, fecundity, and insemination of females after pairing with males exposed to TSB. The population dynamics of Aedes mosquitoes under imbalanced sex ratios were examined to simulate realistic field conditions for male suppression under the effect of TSB., Results: The mortality of male mosquitoes was consistently high within 24 h after exposure. By contrast, the mortality of female mosquitoes was inconsistent, with over 70% mortality observed at 168 h. The flight activity and wing beat amplitude of treated males were significantly lower than those of controls, but no significant difference in wing beat frequency was detected. The fecundity and insemination of treated female mosquitoes were lower than those of controls. A simulation study indicated that considerably low male population densities led to mating failures, triggering a mate-finding Allee effect and resulting in persistently low population levels., Conclusion: Boric acid-containing TSB could effectively complement current chemical intervention approaches to control resistant mosquito populations. TSB is effective in reducing field male populations and impairing male flight activity and female-seeking behavior, resulting in decreased fecundity and insemination. Male suppression due to TSB potentially results in a small mosquito population. © 2024 Society of Chemical Industry., (© 2024 Society of Chemical Industry.)

Long noncoding RNAs (lncRNAs) are transcribed elements increasingly recognized for their roles in regulating gene expression. Thus far, however, we have little understanding of how lncRNAs contribute to evolution and adaptation. Here, we show that a conserved lncRNA, ivory , is an important color patterning gene in the buckeye butterfly Junonia coenia . ivory overlaps with cortex , a locus linked to multiple cases of crypsis and mimicry in Lepidoptera. Along with a companion paper by Livraghi et al., we argue that ivory , not cortex , is the color pattern gene of interest at this locus. In J. coenia , a cluster of cis -regulatory elements (CREs) in the first intron of ivory are genetically associated with natural variation in seasonal color pattern plasticity, and targeted deletions of these CREs phenocopy seasonal phenotypes. Deletions of different ivory CREs produce other distinct phenotypes as well, including loss of melanic eyespot rings, and positive and negative changes in overall wing pigmentation. We show that the color pattern transcription factors Spineless, Bric-a-brac, and Ftz-f1 bind to the ivory promoter during wing pattern development, suggesting that they directly regulate ivory . This case study demonstrates how cis -regulation of a single noncoding RNA can exert diverse and nuanced effects on the evolution and development of color patterns, including modulating seasonally plastic color patterns., Competing Interests: Competing interests statement:The authors declare no competing interest.

There is limited literature regarding wing tip edema (WTE) in raptors, and much of our current understanding of the condition is based on anecdotal reports. The aims of this retrospective study were to describe the clinical features of WTE in birds of prey, to identify prognostic factors for return to flight and patient survival following diagnosis, and to develop and assess the clinical significance of a novel WTE grading system. Between 2004 and 2022, 41 cases of WTE were identified in 39 captive birds. No cases were found in wild birds. Harris's hawks ( Parabuteo unicinctus ), lanner falcons ( Falco biarmicus ), and peregrine falcons ( Falco peregrinus ) had the highest frequencies of WTE, and all cases presented between October and May. Increasing days of air frost per month and colder median monthly temperatures were significant risk factors for the development of WTE. Of the cases where patient outcomes were known, 23/31 (74.2%) cases returned to normal flight and 29/34 (85.3%) cases survived. End-stage disease, represented by primary flight feather loss and metacarpal ischemic (dry) gangrene, and enalapril use were associated with poor patient outcomes. Presentation within 24 hours of disease onset, isoxsuprine use, and physiotherapy were associated with improved patient outcomes. This study showed that WTE is an infrequently encountered but clinically significant condition in captive raptors and is associated with an overall high morbidity and moderate mortality risk.

Butterfly wings possess distinct micro/nanostructures that contribute to their vibrant coloration, light-trapping capabilities, and sensitivity to various stimuli. These complex features have inspired the creation of diverse devices and systems, such as sensors, photovoltaics, photocatalysis, and robotics. Specifically, the wing scales of the Emerald Swallowtail (Papilio palinurus) display iridescent, polarization-sensitive, and retroreflective colors due to their hierarchical structures. However, current technologies fail to mimic these natural designs fully, limiting their practical application in everyday life. In this study, we introduce a groundbreaking method for fabricating artificial wing scales that emulate the biological structure's functionality with a much simpler geometry. By integrating self-graded lossy media into metallic micro-concavity arrays, we achieve pronounced iridescent effects in both coaxial and non-coaxial arrangements, while preserving retroreflective properties. In particular, the simplified design allows for switchable color patterns based on the viewing angle. Demonstrating the concept, we successfully employ these conspicuous retroreflectors in hydrogen gas detection and the bi-directional/switchable recognition of patterned signals., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Bats stand out among mammalian species for their exceptional traits, including the capacity to navigate through flight and echolocation, conserve energy through torpor/hibernation, harbor a multitude of viruses, exhibit resistance to disease, survive harsh environmental conditions, and demonstrate exceptional longevity compared to other mammals of similar size. In vivo studies of bats are challenging for several reasons, such as difficulty in locating and capturing them in their natural environments, limited accessibility, low sample size, environmental variation, long lifespans, slow reproductive rates, zoonotic disease risks, species protection, and ethical concerns. Thus, establishing alternative laboratory models is crucial for investigating the diverse physiological adaptations observed in bats. Obtaining quality cells from tissues is a critical first step for successful primary cell derivation. However, it is often impractical to collect fresh tissue and process the samples immediately for cell culture due to the resources required for isolating and expanding cells. As a result, frozen tissue is typically the starting resource for bat primary cell derivation, but cells in frozen tissue are usually damaged and have low integrity and viability. Isolating primary cells from frozen tissues thus poses a significant challenge. Herein, we present a successfully developed protocol for isolating primary dermal fibroblasts from frozen bat wing biopsies. This protocol marks a significant milestone, as this is the first protocol specifically focused on fibroblast isolation from bat frozen tissue. We also describe methods for primary cell characterization, genetic manipulation of primary cells through lentivirus transduction, and the development of stable cell lines. © 2024 Wiley Periodicals LLC. Basic Protocol 1: Bat wing biopsy collection and preservation Support Protocol 1: Blood collection from bat venipuncture Basic Protocol 2: Isolation of primary fibroblasts from adult bat frozen wing biopsy Support Protocol 2: Primary fibroblast culture and subculture Support Protocol 3: Determination of growth curve and doubling time Support Protocol 4: Cell banking and thawing of primary fibroblasts Basic Protocol 3: Lentiviral transduction of bat primary fibroblasts Basic Protocol 4: Bat stable fibroblast cell line development Support Protocol 5: Bat fibroblast validation by immunofluorescence staining Basic Protocol 5: Chromosome counting., (© 2024 Wiley Periodicals LLC.)

The black soldier fly (BSF) Hermetia illucens (Diptera: Stratiomyidae) plays a significant role at the larval stage in the circular economy due to its ability to convert organic waste into valuable products for energy, food, feed, and agricultural applications. Many data are available on larval development and biomass generation, but basic research on this species is lacking and little is known about adult biology, in particular about the cues involved in sexual recognition. In the present study, using various instruments (stereomicroscope, scanning and transmission electron microscope, hyperspectral camera and spectrophotometer), wing ultrastructure of both sexes was analysed, reflectance and transmission spectra of the wings were measured and behavioural bioassays were carried out to measure male response to specific visual stimuli. The collected data showed the existence of sexual dimorphism in the wings of H. illucens due to iridescent structural colouration generated by a multilayer of melanin located in the dorsal lamina of the central part of the wing. Wing sexual dimorphism is particularly evident regarding the strong emission of blue light of female wings. Blue colour induces in males a strong motivation to mate. The obtained results can help to improve and optimize the breeding techniques of BSF., (© 2024. The Author(s).)

How emerging adaptive variants interact is an important factor in the evolution of wild populations, but the opportunity to empirically study this interaction is rare. We recently documented the emergence of an adaptive phenotype "curly-wing" in Hawaiian populations of field crickets ( Teleogryllus oceanicus ). Curly-wing inhibits males' ability to sing, protecting them from eavesdropping parasitoid flies ( Ormia ochracea ). Surprisingly, curly-wing co-occurs with similarly protective silent "flatwing" phenotypes in multiple populations, in which neither phenotype has spread to fixation. These two phenotypes are frequently coexpressed, but since either sufficiently reduces song amplitude to evade the fly, their coexpression confers no additional fitness benefit. Numerous "off-target" phenotypic changes are known to accompany flatwing, and we find that curly-wing, too, negatively impacts male courtship ability and affects mass and survival of females under lab conditions. We show through crosses and genomic and mRNA sequencing that curly-wing expression is associated with variation on a single autosome. In parallel analyses of flatwing, our results reinforce previous findings of X-linked single-locus inheritance. By combining insights into the genetic architecture of these alternative phenotypes with simulations and field observations, we show that the co-occurrence of these two adaptations impedes either from fixing, despite extreme fitness benefits, due to fitness epistasis. This co-occurrence of similar adaptive forms in the same populations might be more common than is generally considered and could be an important force inhibiting adaptive evolution in wild populations of sexually reproducing organisms., Competing Interests: Competing interests statement:The authors declare no competing interest.

To accommodate growth, insects must periodically shed their exoskeletons. In Manduca sexta, Drosophila melanogaster and Tribolium castaneum, Bursicon (Burs)/ Partner of bursicon (Pburs)-LGR2 signal is an indispensable component for the proper execution of ecdysis behavior during adult eclosion. Nevertheless, the behavioral events and the roles of bursicon signaling in other insects deserve further exploration. In the current paper, we found that the pupal-adult ecdysis in Henosepilachna vigintioctomaculata could be divided into three distinct stages, preecdysis, ecdysis and postecdysis. Preecdysis behavioral sequences included abdomen twitches, dorsal-ventral contractions and air filling that function to loosen the old cuticle. Ecdysis events began with anterior-posterior contractions that gradually split the old integument along the dorsal body midline, followed by freeing of legs and mouthparts, and culminated in detachment from pupal cuticle. Postecdysis behavioral processes contained three actions: perch selection and stretching of elytra and hindwings. RNA interference for HvBurs, HvPburs or Hvrk (encoding LGR2) strongly impaired wing expansion actions, and slightly influenced preecdysis and ecdysis behaviors. The RNAi beetles failed to extend their elytra and hindwings. In addition, injected with dsrk also caused kinked femurs and tibia. Our findings establish that bursicon pathway is involved in regulation of adult eclosion behavior, especially wing expansion motor programs. Given that wings facilitate food foraging, courtship, predator avoidance, dispersal and migration, our results provide a potential target for controlling H. vigintioctomaculata., Competing Interests: Declaration of competing interest The authors declare no competing financial interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Indicator models of sexual selection posit that females choose males on the basis of traits that reveal male genetic quality and thereby enjoy increased offspring production. Here, we report that females of the butterfly Eurema hecabe receive indirect benefits from choosing males based on their ultraviolet (UV) wing coloration, a heritable and condition-dependent trait in this species. We first used a large laboratory-bred pedigree to demonstrate a per-family association between inbreeding and male UV trait value. Females exerted choice for UV-bright males within this protocol, and the average male UV trait value increased over six consecutive generations, presumably due to such selection and despite an increasing rate of pedigree-wide inbreeding. We then experimentally imposed a standard strength of inbreeding upon lines of divergent male UV trait values. Inbreeding depressed the siring performance of low UV treatment males more severely and resulted in a marginal reduction of their UV brightness, which rebounded sharply following subsequent outcrossing. These findings are consistent with the ornament-based signaling of genetic quality as a function of underlying individual-level mutational load., (© 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.)

Insect migrations have ecological and economic impacts, particularly in agriculture. However, there is limited knowledge about the migratory movements of pests at the continental scale, which is an important factor influencing the spread of resistance genes. Understanding the migratory patterns of economic pests, like Helicoverpa zea (Boddie), is essential for improving Integrated Pest Management (IPM) and Insect Resistance Management (IRM) strategies. In this study, we used stable hydrogen isotopic ratios in wing tissue as a biogeochemical marker to examine migratory patterns and estimate the native origins of H. zea individuals collected across a wide latitudinal gradient in North America. Samples collected at higher latitudes (Ontario, Canada and Minnesota, USA) exhibited a greater proportion (60%-96%) of nonlocal individuals, with an increased probability of origin from the southeastern United States. Populations from mid-latitudes (Florida, North Carolina, and South Carolina) showed a blend of local and nonlocal (40%-60%) individuals. Finally, 15% of the southernmost population individuals (Puerto Rico) were classified as migratory, with some having a probability of origin at higher latitudes (>30°). Overall, our results provide evidence of a northward spring/summer migration of H. zea in North America and underscore the significance of the southeastern United States as a hub for genetic flow. In addition, based on stable hydrogen isotopic ratios, there is strong evidence of reverse (southward) migration of H. zea from the continental United States to Puerto Rico. Our study highlights the implications for IPM and IRM programs and the need for management strategies that account for both northward and southward migration patterns., (© The Author(s) 2024. Published by Oxford University Press on behalf of Entomological Society of America.)

The Spalt transcriptional regulators participate in a variety of cell fate decisions during multicellular development. Vertebrate Spalt proteins have been mostly associated to the organization of heterochromatic regions, but they also contribute regulatory functions through binding to A/T rich motives present in their target genes. The developmental processes in which the Drosophila spalt genes participate are well known through genetic analysis, but the mechanism by which the Spalt proteins regulate transcription are still unknown. Furthermore, despite the prominent changes in gene expression associated to mutations in the spalt genes, the specific DNA sequences they bind are unknow. Here, we analyze a DNA fragment present in the regulatory region of the knirps gene. Spalt proteins are candidate repressors of knirps expression during the formation of the venation pattern in the wing disc, and we identified a minimal conserved 30bp sequence that binds to Spalt major both in vivo and in vitro. This sequence mediates transcriptional repression in the central region of the wing blade, constituting the first confirmed case of a direct regulatory interaction between Spalt major and its target DNA in Drosophila. Interestingly, we also find similar sequences in a set of eight novel candidate Spalt target genes, pointing to a common mechanism of transcriptional repression mediated by Spalt proteins., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Morphogenetic programs coordinate cell signaling and mechanical interactions to shape organs. In systems and synthetic biology, a key challenge is determining optimal cellular interactions for predicting organ shape, size, and function. Physics-based models defining the subcellular force distribution facilitate this, but it is challenging to calibrate parameters in these models from data. To solve this inverse problem, we created a Bayesian optimization framework to determine the optimal cellular force distribution such that the predicted organ shapes match the experimentally observed organ shapes. This integrative framework employs Gaussian Process Regression, a non-parametric kernel-based probabilistic machine learning modeling paradigm, to learn the mapping functions relating to the morphogenetic programs that maintain the final organ shape. We calibrated and tested the method on Drosophila wing imaginal discs to study mechanisms that regulate epithelial processes ranging from development to cancer. The parameter estimation framework successfully infers the underlying changes in core parameters needed to match simulation data with imaging data of wing discs perturbed with collagenase. The computational pipeline identifies distinct parameter sets mimicking wild-type shapes. It enables a global sensitivity analysis to support the regulation of actomyosin contractility and basal ECM stiffness to generate and maintain the curved shape of the wing imaginal disc. The optimization framework, combined with experimental imaging, identified that Piezo, a mechanosensitive ion channel, impacts fold formation by regulating the apical-basal balance of actomyosin contractility and elasticity of ECM. This workflow is extensible toward reverse-engineering morphogenesis across organ systems and for real-time control of complex multicellular systems., (© 2024. The Author(s).)

Viability indicator traits are expected to be integrated extensively across the genome yet sex-limited to ensure that any benefits are sexually concordant. Understanding how such expectations are accommodated requires elucidating the quantitative genetic architecture of candidate traits in and across the sexes. Here we applied an animal modelling approach to partition the autosomal, allosomal, and direct maternal bases of variation in sexual versus non-sexual dorsal wing colouration in the butterfly Eurema hecabe. The sexual colour trait-coherently scattered ultraviolet that is under strong directional selection due to female choice-is brighter and more expansive in males, and overlays non-sexual pigmentary yellow markings that otherwise dominate both wing surfaces in each sex. Our modelling estimated high and sexually equivalent autosomal variances for ultraviolet reflectance (furnishing h2 ~ 0.58 overall and ~0.75 in males), accompanied by smaller but generally significant Z-linked and maternal components. By contrast, variation in non-sexual yellow was largely attributed to Z-linked sources. Intersexual genetic correlations based upon the major source of variation in each trait were high and not different from 1.0, implying regulation by a pool of genes common to each sex. An expansive autosomal basis for ultraviolet is consistent with its hypothesized role as a genome-wide viability indicator and ensures that both sons and daughters will inherit their father's attractiveness., (© 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.)

The Ornate Moth, Utetheisa ornatrix , has served as a model species in chemical ecology studies for decades. Like in the widely publicized stories of the Monarch and other milkweed butterflies, the Ornate Moth and its relatives are tropical insects colonizing whole continents assisted by their chemical defenses. With the recent advances in genomic techniques and evo-devo research, it is becoming a model for studies in other areas, from wing pattern development to phylogeography, from toxicology to epigenetics. We used a genomic approach to learn about Utetheisa's evolution, detoxification, dispersal abilities, and wing pattern diversity. We present an evolutionary genomic analysis of the worldwide genus Utetheisa , then focusing on U. ornatrix . Our reference genome of U. ornatrix reveals gene duplications in the regions possibly associated with detoxification abilities, which allows them to feed on toxic food plants. Finally, comparative genomic analysis of over 100 U. ornatrix specimens from the museum with apparent differences in wing patterns suggest the potential roles of cortex and lim3 genes in wing pattern formation of Lepidoptera and the utility of museum-preserved collection specimens for wing pattern research., Competing Interests: Competing interests statement:Drs. Mutanen, Hallwachs, and Janzen are co-authors of an unpublished manuscript with 73 authors (DOI: 10.21203/rs.3.rs-2180975/v1).

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.)

The wing is one of the most important parts of a bird's locomotor system and is the inspiration origination for bionic wing design. During wing motions, the wing shape is closely related to the rotation angles of wing bones. Therefore, the research on the law of bone movement in the process of wing movement can be good guidance for the design of the bionic morphing wing. In this paper, the skeletal posture of the peregrine falcon wing during the extension/flexion is studied to obtain critical data on skeletal posture. Since an elbow joint and a wrist joint rotate correlatively to drive a wing to flex/extend, the wing skeleton is simplified as a four-bar mechanism in this paper. The degree of reproduction of wing skeleton postures was quantitatively analyzed using the four-bar mechanism model, and the bionic wing skeleton was designed. It is found that the wing motions have been reproduced with high precision., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Tang et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

The early period of ontogeny is key to understanding the patterns of body plan formation in birds. Most studies of avian development have focused on the development of individual avian characters, leaving their developmental integration understudied. We explored the dynamics and integration of relative percentage increments in body mass, lengths of head, skeletal elements of wing and leg, and primary flight feathers in the embryonic and postnatal development of the Rook (Corvus frugilegus). The relative percentage increments were calculated according to Brody's equation. Groups of similar growing traits (modules) were determined using hierarchical cluster analysis, and the degree of correlation between modules was estimated by PLS analysis. The embryonic and postnatal periods demonstrate significant consistency both in the dynamics of changes in relative percentage increments of studied traits as well as in the clustering of individual modules. The modules mainly include the body mass and head length, as well as the elements that form the fore- and hind limbs. Differences were revealed in the combination of modules into clusters in embryonic and postnatal periods. Hind limb elements clustered together with wing elements in the embryonic period but with body mass and the head in the postnatal period. The strongest modularity was noted for the leg in embryogenesis, and for the wing in postnatal development. The forelimb and especially the primary feathers had more distinctive growth patterns. We suggest the changes in the degree of integration between locomotor modules in ontogenesis are connected with the earlier functioning of the legs in the postnatal period and with the preparation of the wings for functioning after a chick leaves the nest., (© 2024 Wiley Periodicals LLC.)

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.)

How a developing organ robustly coordinates the cellular mechanics and growth to reach a final size and shape remains poorly understood. Through iterations between experiments and model simulations that include a mechanistic description of interkinetic nuclear migration, we show that the local curvature, height, and nuclear positioning of cells in the Drosophila wing imaginal disc are defined by the concurrent patterning of actomyosin contractility, cell-ECM adhesion, ECM stiffness, and interfacial membrane tension. We show that increasing cell proliferation via different growth-promoting pathways results in two distinct phenotypes. Triggering proliferation through insulin signaling increases basal curvature, but an increase in growth through Dpp signaling and Myc causes tissue flattening. These distinct phenotypic outcomes arise from differences in how each growth pathway regulates the cellular cytoskeleton, including contractility and cell-ECM adhesion. The coupled regulation of proliferation and cytoskeletal regulators is a general strategy to meet the multiple context-dependent criteria defining tissue morphogenesis., (© 2024. The Author(s).)

Flight was a key innovation in the adaptive radiation of insects. However, it is a complex trait influenced by a large number of interacting biotic and abiotic factors, making it difficult to unravel the evolutionary drivers. We investigate flight patterns in neotropical heliconiine butterflies, well known for mimicry of their aposematic wing color patterns. We quantify the flight patterns (wing beat frequency and wing angles) of 351 individuals representing 29 heliconiine and 9 ithomiine species belonging to ten color pattern mimicry groupings. For wing beat frequency and up wing angles, we show that heliconiine species group by color pattern mimicry affiliation. Convergence of down wing angles to mimicry groupings is less pronounced, indicating that distinct components of flight are under different selection pressures and constraints. The flight characteristics of the Tiger mimicry group are particularly divergent due to convergence with distantly related ithomiine species. Predator-driven selection for mimicry also explained variation in flight among subspecies, indicating that this convergence can occur over relatively short evolutionary timescales. Our results suggest that the flight convergence is driven by aposematic signaling rather than shared habitat between comimics. We demonstrate that behavioral mimicry can occur between lineages that have separated over evolutionary timescales ranging from <0.5 to 70 My., Competing Interests: Competing interests statement:The authors declare no competing interest.

Dispersal and establishment strategies are highly variable. Each strategy is associated with specific costs and benefits, and understanding which factors favour or disfavour a strategy is a key issue in ecology and evolution. Ants exhibit several strategies of establishment, i.e. of colony foundation. Some species rely on winged queens that found new colonies alone when others found with accompanying workers (colony fission). The benefits conferred by these workers have been little studied and quantified, because comparing the costs and benefits of solitary foundation vs. colony fission is difficult when comparing different species. We investigated this using the ant Myrmecina graminicola, one of the few species that use both strategies. Young mated queens were allowed to found new colonies in the laboratory, with either zero (solitarily), two or four workers (colony fission). The presence of workers increased both survival and growth of the foundations over the first year, with more workers yielding higher growth. Few workers (as little as two workers) were sufficient to provide benefits, suggesting that in M. graminicola the strategy of colony fission may not dramatically decrease the number of new colonies produced compared to solitary foundation. Because queens performing solitary foundation or colony fission differ in dispersal (by flight vs. on foot), our results support the hypothesis that these two strategies of foundation coexist along a competition-colonization trade-off, where solitary foundation offers a colonization advantage, while colony fission has a competitive advantage., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

We describe the new genus and species Stolleagrion foghnielseni n. gen. et sp. from the Fur Formation in northwestern Denmark based on a single fossil wing. This is the first odonatan described from the earliest part of the PETM recovery phase of the early Eocene. A combination of nine wing character states are considered to be diagnostic of the Dysagrionidae Cockrell only together with the cephalozygopteran head; however, the combination of these nine plus the presence of Ax0 is also diagnostic without the head. By this, we assign Stolleagrion foghnielseni to the Dysagrionidae and reassess the position of other odonates previously treated as cf. Dysagrionidae.

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.)

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.)

Accurate species identification is crucial to assess the medical relevance of a mosquito specimen, but requires intensive experience of the observers and well-equipped laboratories. In this proof-of-concept study, we developed a convolutional neural network (CNN) to identify seven Aedes species by wing images, only. While previous studies used images of the whole mosquito body, the nearly two-dimensional wings may facilitate standardized image capture and reduce the complexity of the CNN implementation. Mosquitoes were sampled from different sites in Germany. Their wings were mounted and photographed with a professional stereomicroscope. The data set consisted of 1155 wing images from seven Aedes species as well as 554 wings from different non-Aedes mosquitoes. A CNN was trained to differentiate between Aedes and non-Aedes mosquitoes and to classify the seven Aedes species based on grayscale and RGB images. Image processing, data augmentation, training, validation and testing were conducted in python using deep-learning framework PyTorch. Our best-performing CNN configuration achieved a macro F1 score of 99% to discriminate Aedes from non-Aedes mosquito species. The mean macro F1 score to predict the Aedes species was 90% for grayscale images and 91% for RGB images. In conclusion, wing images are sufficient to identify mosquito species by CNNs., (© 2024. The Author(s).)

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.)

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).)

Recent studies have revealed taxonomic signals within the wing cells of certain mosquito species. In our study, wing cell differentiation among mosquito vectors from the Tanaosri mountain range in Thailand was evaluated using the outline-based geometric morphometric (GM) approach. Our focus was on four specific wing cells for GM analysis: the wing contour (external cell), the second submarginal cell (internal cell 1), the first posterior cell (internal cell 2), and the third posterior cell (internal cell 3). Before proceeding with the GM approach, the identity of seven mosquito genera and 21 species was confirmed using molecular techniques. Our validated classification tests demonstrated that the performance of mosquito species classification varies according to genus. Notably, three Aedes species exhibited the highest accuracy for both internal cell 2 and internal cell 3, each registering a score of 93.20 %. In the case of two Mansonia species, the wing contour displayed a remarkable accuracy of 98.57 %. Consequently, we suggest the use of the outline-based GM approach, particularly focusing on the wing contour, for differentiating Mansonia annulifera and Mansonia uniformis. In contrast, the highest accuracy for classifying Culex species was found in internal cell 1, at 75.51 %, highlighting the challenges due to similarities in wing cells within this genus. These findings provide a guideline for future applications of the outline-based GM approach, focusing on wing cells, as an alternative method to classify mosquito vector species., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Butterfly wings display rich phenotypic diversity and are associated with complex biological functions, thus serving as an important evolutionary system to address the genetic basis and evolution of phenotypic diversification. We review recent butterfly studies that revealed complex functions underlying diversified wing patterns and describe the genetic and environmental factors involved in wing pattern determinations. These factors lead to inter-specific divergence, genetic polymorphism, and phenotypic plasticity, which in many cases are decided by several key genes. We also summarize the research advances on gene co-option as an important origin of functional complexity and evolutionary novelty. These findings reveal a pattern of evolutionary innovation within a constrained developmental framework during butterfly wing morphogenesis, but further research is required to gain a systematic and comprehensive understanding., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Inc. All rights reserved.)

How can highly dispersive species give rise to genetically distinct populations? This seemingly paradoxical pattern is common among insular birds, but not in those with continental distributions. Broyles and Myers sequence the genomes of almost 150 individuals from the island-dwelling Pacific swallow (Hirundo tahitica) and its continental counterpart, the welcome swallow (H. neoxena). They find strong population structure only among island populations and attribute this to a behavioral reduction in dispersal propensity following island colonization. However, wing shape remains consistent across populations, suggesting it might not accurately reflect dispersal propensity in this group. This study illustrates the interplay between dispersal, isolation, and divergence, offering insights into how geographic factors affect speciation and population differentiation on islands., (© The Author(s) 2023. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE). All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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.)

Identifying and classifying species of stingless Neotropical bees is not a trivial task and requires the help of taxonomists and substantial study and training time. Also, there is a lack of taxonomically useful characters to differentiate among the megadiverse Neotropical group of stingless bees, and to recognize variation. Based on that, we have been testing alternatives to a character-based, efficient taxonomic determination of stingless bees, and herein we performed exploratory analyses of wing shape variation using geometric morphometrics. Thus, we built a data set of photographs of the right anterior wing of 1628 individuals belonging to 11 species in the genus Melipona (Illiger 1806) taken from collection material deposited in entomological collections. We then conducted a Procrustes analysis, followed by a Principal Component Analysis (PCA) and by a Canonical Variable Analyses (CVA). The two first principal components of the PCA accounted for 68% of the variation of the wing shape, and the ordination displayed by the first component separated species of the subgenus Melikerria from the others. In the CVA, the first two canonical axes explained 88% of the wing shape variation found among species, and Melikerria appears as a separate group, apart from the other subgenera in the first canonical axis. Along the second axis species belonging to Eomelipona and Michmelia, and among the Michmelia species, and its species group fuliginosa formed well-separated clusters. The wing shape variation of Melipona supports the recognition of subgenera as currently proposed for Melipona.

What regulates organ size and shape remains one fundamental mystery of modern biology. Research in this area has primarily focused on deciphering the regulation in time and space of growth and cell division, while the contribution of cell death has been overall neglected. This includes studies of the Drosophila wing, one of the best-characterized systems for the study of growth and patterning, undergoing massive growth during larval stage and important morphogenetic remodeling during pupal stage. So far, it has been assumed that cell death was relatively neglectable in this tissue both during larval stage and pupal stage, and as a result, the pattern of growth was usually attributed to the distribution of cell division. Here, using systematic mapping and registration combined with quantitative assessment of clone size and disappearance as well as live imaging, we outline a persistent pattern of cell death and clone elimination emerging in the larval wing disc and persisting during pupal wing morphogenesis. Local variation of cell death is associated with local variation of clone size, pointing to an impact of cell death on local growth that is not fully compensated by proliferation. Using morphometric analyses of adult wing shape and genetic perturbations, we provide evidence that patterned death locally and globally affects adult wing shape and size. This study describes a roadmap for precise assessment of the contribution of cell death to tissue shape and outlines an important instructive role of cell death in modulating quantitatively local growth and morphogenesis of a fast-growing tissue., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 The Author(s). Published by Elsevier Inc. All rights reserved.)

The initial process by which novel sexual signals evolve remains unclear, because rare new variants are susceptible to loss by drift or counterselection imposed by prevailing female preferences. 1 , 2 , 3 , 4 We describe the diversification of an acoustic male courtship signal in Hawaiian populations of the field cricket Teleogryllus oceanicus, which was brought about by the evolution of a brachypterous wing morph ("small-wing") only 6 years ago. 5 Small-wing has a genetic basis and causes silence or reduced-amplitude signaling by miniaturizing male forewings, conferring protection against an eavesdropping parasitoid, Ormia ochracea. 5 We found that wing reduction notably increases the fundamental frequency of courtship song from an average of 5.1 kHz to 6.4 kHz. It also de-canalizes male song, broadening the range of peak signal frequencies well outside normal song character space. As courtship song prompts female mounting and is sexually selected, 6 , 7 , 8 , 9 we evaluated two scenarios to test the fate of these new signal values. Females might show reduced acceptance of small-wing males, imposing counterselection via prevailing preferences. Alternatively, females might accept small-wing males as readily as long-wing males if their window of preference is sufficiently wide. Our results support the latter. Females preferred males who produced some signal over none, but they mounted sound-producing small-wing males as often as sound-producing long-wing males. Indiscriminate mating can facilitate the persistence of rare, novel signal values. If female permissiveness is a general characteristic of the earliest stages of sexual signal evolution, then taxa with low female mate acceptance thresholds should be more prone to diversification via sexual selection., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 The Authors. Published by Elsevier Inc. All rights reserved.)

First Upper Triassic dipterans (Nematocera) are described based on isolated wings from Fuchsberg near Seinstedt (Sevatian, uppermost Norian of Lower Saxony, Germany): Crivoptychoptera nebrias gen. et sp. nov. (Psychodomorpha, Ptychopteridae), Ptychopteridae incertae sedis, Dohloboyia triassica gen. et sp. nov. (Bibionomorpha, Boholdoyidae) and Chironomidae incertae sedis (Culicomorpha). All taxa under discussion present the oldest occurrences of their families. Geological background and composition of the Upper Triassic insect fauna of Seinstedt are briefly discussed.

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.)