Comparative functional morphology of the wings of Diptera

Wings of representative species of the order Diptera were compared with a simple model structure in which corrugated spars diverge from a V-shaped leading edge spar. Both develop torsion and camber when subjected to aerodynamic loads, forming a propeller shape. Both the leading edge and the cubitus of flies' wings twist basally, allowing camber to be set up as the media hinges up or down at the arculus. Three different wing types were identified: stiff wings possessing two or three main spars; and wings capable of ventral flexion. In wings possessing only two spars, found mainly in the Nematocera, control of camber is achieved largely by the use of cross veins. Wing control and flight are generally imprecise. The third spar, found in most Brachycera, in the Syrphidae and in the Conopidae controls camber and helps support a broader wing. Finer control of camber is exerted by marginal cross veins, and these insects generally have precise, darting flight. Ventral flexion mechanisms are found in the Simuliidae, the Stratiomyiidae, and widely in the Schizophora. Control of ventral flexion, which occurs at the end of the downstroke, allows fast, unpredictable manoeuvres. Functional similarities indicate either phylogenetic relationship or convergence.     

Comparing wing loading,flight muscle and lipid content in ant‐attended and non‐attended Tuberculatus aphid species

Although all Tuberculatus aphids possess wings, some species associated with ants exhibit extremely low levels of dispersal compared with those not associated with ants. Furthermore, phylogenetic interspecific comparisons find significantly higher wing loading (i.e. higher ratio of body volume to wing area) in ant‐attended species. This observation indicates that ant‐attended species may allocate more of their body resources to reproductive traits (i.e. embryos) rather than flight apparatus (i.e. wings, flight muscle and lipid). The present study focuses on two sympatric aphid species and aims to investigate the hypothesized trade‐off in resource investment between fecundity and the flight apparatus; specifically, the ant‐attended Tuberculatus quercicola (Matsumura) and non‐attended Tuberculatus paiki Hille Ris Lambers. Species differences are compared in: (i) morphology, (ii) embryo production, (iii) triacylglycerol levels and (iv) wing loading and flight muscle. The results show that T. quercicola has a larger body volume, higher fecundity and higher wing loading compared with T. paiki, which has a smaller, slender‐shaped body, lower fecundity and lower wing loading. No significant difference is found between the species with respect to the percentage of triacylglycerol content in dry body weight. The flight muscle development is significantly lower in T. quercicola than in T. paiki. These results indicate that the additive effect of higher wing loading and the lower amount of flight muscle development in T. quercicola may increase the physical difficulty of flight, and hence be responsible for its lower dispersal ability. The trade‐off between fecundity and dispersal documented in wing‐dimorphic insects may therefore be applicable to T. quercicola, which has fully developed wings.     

昆虫翅型分化的调控及翅多型性的进化

翅多型现象普遍存在于各昆虫类群,一些学者就昆虫翅多型进行了大量的研究工作。根据昆虫翅型的分化,可划分为长翅型和短翅型,长翅型具飞行能力,而短翅型不能飞行。一些昆虫种类,如蚜虫,出现无翅个体,被称为无翅型。除飞行能力外,长翅型和短翅型在行为、生理等方面也存在差异。文章主要就环境因素对翅型分化的影响、翅多型的内分泌控制机理、翅多型的遗传机制及其进化等作一概述。     

食物胁迫对翅二型丽斗蟋飞行肌和繁殖发育的影响

前期研究表明,在食物充足的条件下,翅二型丽斗蟋雌成虫长、短翅型间存在着资源投入和收益的权衡关系(trade-off);而雄成虫长短翅型间不存在此类权衡关系。在自然条件下,昆虫可能遭受食物缺乏的胁迫,因而进一步就食物胁迫对丽斗蟋飞行肌和繁殖发育的影响进行了研究。结果表明,在食物胁迫的条件下,长翅雌虫仍维持飞行肌的发育,但繁殖发育受到显著的抑制;而短翅雌虫飞行肌显著降解,繁殖发育亦维持在较高水平。说明即使是营养缺乏时,其雌成虫长、短翅型也依然存在资源配置的差异,具飞行肌和繁殖发育的权衡关系。长翅雄虫飞行肌的重量与食物充足组并无显著差异,但精巢的干重显著降低;而短翅雄虫在食物胁迫条件下飞行肌显著降解,但其精巢重量与食物充足组并无显著差异。可以认为,丽斗蟋雄虫的长、短翅型间也存在飞行肌和繁殖发育的权衡关系。     

Site-occupancy in relation to flight-morphology in caddisflies

1. The relationship between morphology and site‐occupancy provides opportunities to infer differences in dispersal and flight ability, but empirical data for aquatic insects is limited. 2. In this study, 17 species of caddisflies from 10 families were collected from springs, streams and lakes, and total body mass, relative thorax mass, relative wing area (wing loading), and the aspect ratio of the fore and hind wings (combined) were measured. 3. Partial least‐squares regression analysis of two independent distributional data sets produced significant models within which total body mass, relative thorax mass and wing loading were positively associated with site‐occupancy, whereas aspect ratio was negatively associated with site‐occupancy. 4. These results suggest that the faunal composition of streams is influenced by species dispersal abilities.     

Wetting Characteristics of Insect Wing Surfaces

Biological tiny structures have been observed on many kinds of surfaces such as lotus leaves,which have an effect on thecoloration of Morpho butterflies and enhance the hydrophobicity of natural surfaces.We investigated the micro-scale andnano-scale structures on the wing surfaces of insects and found that the hierarchical multiple roughness structures help in enhancingthe hydrophobicity.After examining 10 orders and 24 species of flying Pterygotan insects,we found that micro-scaleand nano-scale structures typically exist on both the upper and lower wing surfaces of flying insects.The tiny structures such asdenticle or setae on the insect wings enhance the hydrophobicity,thereby enabling the wings to be cleaned more easily.And thehydrophobic insect wings undergo a transition from Cassie to Wenzel states at pitch/size ratio of about 20.In order to examinethe wetting characteristics on a rough surface,a biomimetic surface with micro-scale pillars is fabricated on a silicon wafer,which exhibits the same behavior as the insect wing,with the Cassie-Wenzel transition occurring consistently around apitch/width value of 20.     

Differences in wing morphology between juvenile and adult European Turtle Doves Streptopelia turtur: implications for migration and predator escape

Behaviour has direct links to wing morphology in bird species. Many studies have postulated migration to be one of the most important forces of selection acting on wing morphology, particularly in relation to wing pointedness. Studies in passerines have found that adults have longer and more pointed wings than juveniles, especially in migratory species. We analysed differences in wing morphology between age groups of the European Turtle Dove, a non‐passerine migratory species that benefits from rounded wings during their daily activity, due to its ground‐feeding behaviour and acrobatic flight style. Our results show that adults of this species have longer but more rounded wings than juveniles. This suggests that in this species wing morphology in juveniles is selected to facilitate the first migration, whereas other selection forces (e.g. flight manoeuvrability) are more important after the first moult. These data also explain why juveniles are not as adept at escaping from predators or hunters as adults.     

Relationship between body size and flying-related structures in Neotropical social wasps (Polistinae,Vespidae, Hymenoptera)

Body size influences wing shape and associated muscles in flying animals which is a conspicuous phenomenon in insects, given their wide range in body size. Despite the significance of this, to date, no detailed study has been conducted across a group of species with similar biology allowing a look at specific relationship between body size and flying structures. Neotropical social vespids are a model group to study this problem as they are strong predators that rely heavily on flight while exhibiting a wide range in body size. In this paper we describe the variation in both wing shape, as wing planform, and mesosoma muscle size along the body size gradient of the Neotropical social wasps and discuss the potential factors affecting these changes. Analyses of 56 species were conducted using geometric morphometrics for the wings and lineal morphometrics for the body; independent contrast method regressions were used to correct for the phylogenetic effect. Smaller vespid species exhibit rounded wings, veins that are more concentrated in the proximal region, larger stigmata and the mesosoma is proportionally larger than in larger species. Meanwhile, larger species have more elongated wings, more distally extended venation, smaller stigmata and a proportionally smaller mesosoma. The differences in wing shape and other traits could be related to differences in flight demands caused by smaller and larger body sizes. Species around the extremes of body size distribution may invest more in flight muscle mass than species of intermediate sizes.     

Morphology and evolution of the wing bullae in South American Leptophlebiidae (Ephemeroptera)

Insects were the first animals to take to the skies, and have been flying for over 320 million years. The order Ephemeroptera is, or at least is part of, the most early-diverging lineage of extant winged insects. The extant species present a very short adult life span, mainly dedicated to reproduction and dispersal of eggs. Mating and egg-laying behavior depend on flight. Wings are structures to fly and as such face a number of physical and physiological challenges. The convex curvature along the anterior–posterior axis of the wing generates a camber that must be carefully regulated. One of the most interesting ways of wing bending is provided by the bullae, which have been defined as short sections of flexible chitin, where the flexion lines cross veins. Although the bullae have been frequently used as taxonomic characters, there is no study focused on their morphology, although their prevalence on the wings of mayflies strongly suggests a role in flight. In order to identify evolutionary trends of these structures within Ephemeroptera, we constructed a matrix with comparative anatomy data of the bullae from whole mounts of the wings of 300 specimens belonging to 70 species of several mayfly families, as well as scanning microscopy samples of selected specimens. We also surveyed the number of bullae and their distribution in the wings of the different species within the South American Leptophlebiidae clade. We optimized the characters onto the latest published phylogeny for Leptophlebiidae.     

Upstroke wing flexion and the inertial cost of bat flight

Flying vertebrates change the shapes of their wings during the upstroke, thereby decreasing wing surface area and bringing the wings closer to the body than during downstroke. These, and other wing deformations, might reduce the inertial cost of the upstroke compared with what it would be if the wings remained fully extended. However, wing deformations themselves entail energetic costs that could exceed any inertial energy savings. Using a model that incorporates detailed three-dimensional wing kinematics, we estimated the inertial cost of flapping flight for six bat species spanning a 40-fold range of body masses. We estimate that folding and unfolding comprises roughly 44 per cent of the inertial cost, but that the total inertial cost is only approximately 65 per cent of what it would be if the wing remained extended and rigid throughout the wingbeat cycle. Folding and unfolding occurred mostly during the upstroke; hence, our model suggests inertial cost of the upstroke is not less than that of downstroke. The cost of accelerating the metacarpals and phalanges accounted for around 44 per cent of inertial costs, although those elements constitute only 12 per cent of wing weight. This highlights the energetic benefit afforded to bats by the decreased mineralization of the distal wing bones.     

The Role of Soft Vein Joints in Dragonfly Flight

Dragonflies are excellent flyers among insects and their flight ability is closely related to the architecture and material properties of their wings.The veins are main structure components of a dragonfly wing,which are found to be connected by resilin with high elasticity at some joints.A three-dimensional (3D) finite element model of dragonfly wing considering the soft vein joints is developed,with some simplifications.Passive deformation under aerodynamic loads and active flapping motion of the wing are both studied.The functions of soft vein joints in dragonfly flight are concluded.In passive deformation,the chordwise flexibility is improved by soft vein joints and the wing is cambered under loads,increasing the action area with air.In active flapping,the wing rigidity in spanwise direction is maintained to achieve the required amplitude.As a result,both the passive deformation and the active control of flapping work well in dragonfly flight.The present study may also inspire the design of biomimetic Flapping Micro Air Vehicles (FMAVs).     

Skipping flights in Ypthima butterflies (Lepidoptera: Nymphalidae)

The skipping flight patterns of three species of Ypthima (Lepidoptera: Nymphalidae) were analyzed using high‐speed video recordings to clarify how wings move and how driving forces are produced. All three species showed a flight pattern that includes a pause that accounts for about 50% of a flap cycle when their wings completely close after each upstroke. The observed pause causes the “skipping” flight trajectory based on the clap–fling mechanism. Pause duration was correlated with upstroke wing motion, suggesting the contribution of the latter to a long pause duration. This is also supported by the temporal relationship between the wing and body motions. The aerodynamic power necessary for the pause flight was calculated for the three species.     

Wings versus legs in the avian bauplan: Development and evolution of alternative locomotor strategies

Wings have long been regarded as a hallmark of evolutionary innovation, allowing insects, birds, and bats to radiate into aerial environments. For many groups, our intuitive and colloquial perspective is that wings function for aerial activities, and legs for terrestrial, in a relatively independent manner. However, insects and birds often engage their wings and legs cooperatively. In addition, the degree of autonomy between wings and legs may be constrained by tradeoffs, between allocating resources to wings versus legs during development, or between wing versus leg investment and performance (because legs must be carried as baggage by wings during flight and vice versa). Such tradeoffs would profoundly affect the development and evolution of locomotor strategies, and many related aspects of animal ecology. Here, we provide the first evaluation of wing versus leg investment, performance and relative use, in birds—both across species, and during ontogeny in three precocial species with different ecologies. Our results suggest that tradeoffs between wing and leg modules help shape ontogenetic and evolutionary trajectories, but can be offset by recruiting modules cooperatively. These findings offer a new paradigm for exploring locomotor strategies of flying organisms and their extinct precursors, and thereby elucidating some of the most spectacular diversity in animal history.     

Antagonistic natural and sexual selection on wing shape in a scrambling damselfly

Wings are a key trait underlying the evolutionary success of birds, bats, and insects. For over a century, researchers have studied the form and function of wings to understand the determinants of flight performance. However, to understand the evolution of flight, we must comprehend not only how morphology affects performance, but also how morphology and performance affect fitness. Natural and sexual selection can either reinforce or oppose each other, but their role in flight evolution remains poorly understood. Here, we show that wing shape is under antagonistic selection with regard to sexual and natural selection in a scrambling damselfly. In a field setting, natural selection (survival) favored individuals with long and slender forewings and short and broad hindwings. In contrast, sexual selection (mating success) favored individuals with short and broad forewings and narrow‐based hindwings. Both types of selection favored individuals of intermediate size. These results suggest that individuals face a trade‐off between flight energetics and maneuverability and demonstrate how natural and sexual selection can operate in similar directions for some wing traits, that is, wing size, but antagonistically for others, that is, wing shape. Furthermore, they highlight the need to study flight evolution within the context of species’ mating systems and mating behaviors.     

On the sex-specific mechanisms of butterfly flight: flight performance relative to flight morphology, wing kinematics, and sex in Pararge aegeria

Many evolutionary ecological studies have documented sexual dimorphism in morphology or behaviour. However, to what extent a sex-specific morphology is used differently to realize a certain level of behavioural performance is only rarely tested. We experimentally quantified flight performance and wing kinematics (wing beat frequency and wing stroke amplitude) and flight morphology (thorax mass, body mass, forewing aspect ratio, and distance to centre of forewing area) in the butterfly Pararge aegeria (L.) using a tethered tarsal reflex induced flight set-up under laboratory conditions. On average, females showed higher flight performance than males, but frequency and amplitude did not differ. In both sexes, higher flight performance was partly determined by wing beat frequency but not by wing stroke amplitude. Dry body mass, thorax mass, and distance to centre of forewing area were negatively related to wing beat frequency. The relationship between aspect ratio and wing stroke amplitude was sex-specific: females with narrower wings produced higher amplitude whereas males show the opposite pattern. The results are discussed in relation to sexual differences in flight behaviour.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 675–687.     

Kinematic compensation for wing loss in flying damselflies

Flying insects can tolerate substantial wing wear before their ability to fly is entirely compromised. In order to keep flying with damaged wings, the entire flight apparatus needs to adjust its action to compensate for the reduced aerodynamic force and to balance the asymmetries in area and shape of the damaged wings. While several studies have shown that damaged wings change their flapping kinematics in response to partial loss of wing area, it is unclear how, in insects with four separate wings, the remaining three wings compensate for the loss of a fourth wing. We used high-speed video of flying blue-tailed damselflies (Ischnura elegans) to identify the wingbeat kinematics of the two wing pairs and compared it to the flapping kinematics after one of the hindwings was artificially removed. The insects remained capable of flying and precise maneuvering using only three wings. To compensate for the reduction in lift, they increased flapping frequency by 18 ± 15.4% on average. To achieve steady straight flight, the remaining intact hindwing reduced its flapping amplitude while the forewings changed their stroke plane angle so that the forewing of the manipulated side flapped at a shallower stroke plane angle. In addition, the angular position of the stroke reversal points became asymmetrical. When the wingbeat amplitude and frequency of the three wings were used as input in a simple aerodynamic model, the estimation of total aerodynamic force was not significantly different (paired t-test, p = 0.73) from the force produced by the four wings during normal flight. Thus, the removal of one wing resulted in adjustments of the motions of the remaining three wings, exemplifying the precision and plasticity of coordination between the operational wings. Such coordination is vital for precise maneuvering during normal flight but it also provides the means to maintain flight when some of the wings are severely damaged.     

The redder the better: wing color predicts flight performance in monarch butterflies

The distinctive orange and black wings of monarchs (Danaus plexippus) have long been known to advertise their bitter taste and toxicity to potential predators. Recent work also showed that both the orange and black coloration of this species can vary in response to individual-level and environmental factors. Here we examine the relationship between wing color and flight performance in captive-reared monarchs using a tethered flight mill apparatus to quantify butterfly flight speed, duration and distance. In three different experiments (totaling 121 individuals) we used image analysis to measure body size and four wing traits among newly-emerged butterflies prior to flight trials: wing area, aspect ratio (length/width), melanism, and orange hue. Results showed that monarchs with darker orange (approaching red) wings flew longer distances than those with lighter orange wings in analyses that controlled for sex and other morphometric traits. This finding is consistent with past work showing that among wild monarchs, those sampled during the fall migration are darker in hue (redder) than non-migratory monarchs. Together, these results suggest that pigment deposition onto wing scales during metamorphosis could be linked with traits that influence flight, such as thorax muscle size, energy storage or metabolism. Our results reinforce an association between wing color and flight performance in insects that is suggested by past studies of wing melansim and seasonal polyphenism, and provide an important starting point for work focused on mechanistic links between insect movement and color.     

若干蛾类翅面正投影形状聚类分析鳞翅目：缰翅亚目)

本文以翅面正投影形状的特征参数为指标,对20科71种鳞翅目蛾类昆虫进行系统聚类。结果,粘虫Mythimna separata、小地老虎Agrotis ypsilon、稻纵卷叶螟Cnaphalocrocismedinalis等迁飞昆虫集中地归于一类,表明具有远距离迁飞行为的蛾类翅面几何形状相似,存在区别于其它种类的共同特征,即前翅较窄长,翅前缘较平直,外侧宽阔。这可能是适应远距离迁飞的特征。     

A mechanical model of wing and theoretical estimate of taper factor for three gliding birds

We tested a mechanical model of wing, which was constructed using the measurements of wingspan and wing area taken from three species of gliding birds. In this model, we estimated the taper factors of the wings for jackdaw (Corrus monedula), Harris’ hawk (Parabuteo unicinctas) and Lagger falcon (Falco jugger) as 1.8, 1.5 and 1.8, respectively. Likewise, by using the data linear regression and curve estimation method, as well as estimating the taper factors and the angle between the humerus and the body, we calculated the relationship between wingspan, wing area and the speed necessary to meet the aerodynamic requirements of sustained flight. In addition, we calculated the relationship between the speed, wing area and wingspan for a specific angle between the humerus and the body over the range of stall speed to maximum speed of gliding flight. We then compared the results for these three species of gliding birds. These comparisons suggest that the aerodynamic characteristics of Harris’ hawk wings are similar to those of the falcon but different from those of the jackdaw. This paper also presents two single equations to estimate the minimum angle between the humerus and the body as well as the minimum span ratio of a bird in gliding flight.     

Wing flexibility enhances load-lifting capacity in bumblebees

The effect of wing flexibility on aerodynamic force production has emerged as a central question in insect flight research. However, physical and computational models have yielded conflicting results regarding whether wing deformations enhance or diminish flight forces. By experimentally stiffening the wings of live bumblebees, we demonstrate that wing flexibility affects aerodynamic force production in a natural behavioural context. Bumblebee wings were artificially stiffened in vivo by applying a micro-splint to a single flexible vein joint, and the bees were subjected to load-lifting tests. Bees with stiffened wings showed an 8.6 per cent reduction in maximum vertical aerodynamic force production, which cannot be accounted for by changes in gross wing kinematics, as stroke amplitude and flapping frequency were unchanged. Our results reveal that flexible wing design and the resulting passive deformations enhance vertical force production and load-lifting capacity in bumblebees, locomotory traits with important ecological implications.