Wing length allometry in Odonata: differences between families in relation to migratory behaviour

In insects, wing shape and body size are correlated with several aspects of behaviour, and the optimal morphology of wings is a trade-off between a number of functional demands in relation to behaviour (e.g. foraging, migration and sexual display). Dragonflies are spectacularly skilful flyers and present a range of different wing shapes, but to date, no detailed studies have been conducted in this group on wing length allometry in relation to body size. In this paper, we use published data on body length and wing length in all European and North American dragonflies to investigate differences in wing length allometries among Odonata taxa (suborders and families) and to relate these to behavioural patterns. We found different wing allometries between Zygoptera and Anisoptera, which are probably related to the flight mode and wing form of the two suborders. Among the Anisoptera, the Libellulidae showed a distinct wing length allometry from all other anisopteran families and migrants differed from non-migrant species. The first dichotomy is likely to reflect the adaptation of wing morphology of Libellulidae to sit-and-wait behaviour and to brief foraging flights (most species of this family are perchers) with respect to all other families, members of which are typically flyers. The second dichotomy reflects the trend of migrating species to have relatively longer wings than non-migrating members of the same family. Finally, wing length allometry differed among all the zygopteran families analysed, and this pattern suggested that each family evolved a particular wing morphology in response to peculiarities in behaviour, habitat and flight mode.     

Flight behaviour of seabirds in relation to wind direction and wing morphology

We studied flight direction relative to wind direction (hereafter referred to as "flight direction"), the relation between wing morphology and flight behaviour and interspecies relationships in flight behaviour among all major seabird taxa. We calculated wing loading and aspect ratios for 98 species from 1029 specimens. Species were sorted into 13 groups on the basis of similarity in patterns of flight direction. The primary flight direction of Pelecaniformes and Charadriiformes was into and across headwinds. The most common flight direction of Procellariiformes was across wind. Seabirds avoided flying with tailwinds. Wing loading and aspect ratios were positively correlated in Procellariiformes, Pelecaniformes and alcids but negatively correlated in larids. In Procellariiformes, incidence of headwind flight and that of tailwind flight were significantly correlated with wing loading and aspect ratio; species with higher wing loading and aspect ratios flew more often into headwinds and less often with tailwinds. In contrast, the proportion of Pelecaniformes and Charadriiformes flying with tailwinds increased significantly with increased wing loading. Our results demonstrate a close link in seabirds between flight behaviour, wing morphology and natural history patterns in terms of distribution, colony location, dispersal and foraging behaviour.     

Trophic ecology of adult male Odonata. I. Dietary niche metrics by foraging guild,species, body size,and location

1. Information on the dietary niches of adult odonates is sparse, as they are highly mobile and evasive animals, which makes them difficult to observe in their natural habitat. Moreover, there is a lack of knowledge on how varying behavioural traits of odonates relate to phenomena like niche partitioning. 2. This study investigated niche partitioning amongst odonate species, foraging guilds and size classes in a riverine system in the Eastern Cape province of South Africa. A combination of stable isotope and fatty acid‐based niches was used to infer odonate feeding. 3. Both fatty acid and stable isotope‐based niches showed that there was niche separation amongst odonates that forage in flight (fliers) and those that forage from a perch (perchers), amongst odonates of different size classes (damselflies, medium‐ and large‐sized dragonflies), and amongst species, although varying levels of niche overlap were observed in each case. 4. Niche sizes of odonates varied between an upstream and a downstream site. Generally greater niche overlap was recorded at the narrow upstream site (associated with low insect emergence rates) than the wider downstream site (associated with high insect emergence rates), indicating that a greater degree of resource sharing occurred at the upstream site where aquatic food was less abundant. 5. The findings of this study suggest that dietary niches of odonates can be influenced by foraging guild, body size, and/or environmental conditions, and additional study in a variety of regions is recommended to determine the greater applicability of these findings.     

Peculiarities of structural-functional organization of the motor neuropil in the dragonfly thoracic ganglia

The study considers structural-functional relations in motor neuropil of the thoracic ganglia in dragonflies-insects capable of performing very complex and fast maneuvering in flight. The motor neuropil in dragonflies was shown to be more differentiated than in less mobile insects, while its motor nuclei are more outlined and approached to each other. There were revealed dendrites of the leg muscle motoneurons (intermediate nucleus), running to the anterior and posterior nuclei that contain dendrites of the wing muscle motoneurons. A possible role of such a dendrite approaching is discussed for close functional cooperation of wing and leg muscles essential for dragonflies to catch a large prey in flight by using their legs. Peculiarities of structural organization of the wing muscle motoneurons in dragonflies and locusts are considered to suggest the greater functional capabilities of motoneurons in the dragonfly motor apparatus.     

Predator foolhardiness and morphological evolution in 17-year cicadas (Magicicada spp.)

Periodical cicadas in the genus Magicicada have an unusual life history that includes an exceptionally long life cycle and a massive, synchronized emergence. Considerable effort has been put into research aimed at understanding the evolutionary history of periodical cicadas, but surprisingly little attention has been given to their morphological evolution. Their slow flight and approachability have been described as 'predator-foolhardy' behaviour. We quantified flight speeds for M. cassini, M. septendecim, and Tibicen chloromera (a nonperiodical cicada species) , and interpreted them in terms of thorax musculature, body proportions and wing size and shape in relation to body size. On average, T. chloromera flew three to four times faster than did the two Magicicada species. Using empirical relationships between flight speed and body length, body mass or wing loading, we determined M. cassini and M. septendecim to be unusually slow fliers for their body size, whereas T. chloromera was not. The relatively slow flight speeds of Magicicada species could be largely accounted for by relatively small thoracic muscle masses, as indicated by thorax length × width measurements, and low wing loadings. Aspect ratio differences were contributing factors. Male Magicicada and female Tibicen were more active in mate searching than was the opposite sex, and correspondingly had relatively large aspect ratios. We interpret the morphological traits responsible for the slow flight of Magicicada species as being adaptations to searching for mates in dense aggregations around the canopy of trees, relatively unconstrained by the per-capita risk of predation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 1–13.     

Effects of different methods of non‐lethal tissue sampling on butterflies

1. We investigated the effects of two methods of non‐lethal tissue sampling on post‐release flight behaviour (short‐term response) and survival (long‐term response) of two butterflies, Pieris rapae and Coenonympha tullia, within the same natural habitat. We applied three treatments: control (no tissue removal), wing clipping, and leg removal. Our study is the first to directly compare the effects of these common sampling methods. 2. We monitored the flight behaviour of the butterflies by following individuals immediately after their release. In 99 behaviour trials of P. rapae and 101 of C. tullia we found no significant differences in proportion of time spent flying or displacement per unit time among treatment groups in either species. 3. We used standard mark–recapture techniques continuously throughout the flight season to compare the survival of individuals. We marked a total of 687 P. rapae and 490 C. tullia butterflies. We found no significant differences in survival among treatments in either species. 4. We detected differences between the sexes in survival in P. rapae and flight behaviour in C. tullia. In addition to indicating differences in ecology between the sexes, these results also suggest that our analyses were sufficiently powerful to detect a significant effect of tissue removal had such an effect existed. 5. Our work is an important addition to the accumulating evidence that these methods of non‐lethal tissue sampling are generally not detrimental. These sampling techniques closely mimic conditions in the wild, as wing wear and leg losses occur naturally.     

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.     

Structural-functional peculiarities of the wing apparatus of insects that do not have and do have the maneuvering flight

The work considers character of behavior in flight and discusses peculiarities of structural-functional organization of the wing apparatus of two representatives of insects—the migratory Asian locust Locusta migratoria (a low-maneuvering insect) and the dragonfly-darner Aeshna sp. (an insect able to perform complex maneuvers in air). The main principles underlying the insect wing apparatus activity are considered and the mechanisms allowing the dragonflies to perform complex maneuvers in the flight are analyzed in detail.     

Initiation of flight and pre-flight behaviour of anisopterous dragonflies Aeshna spp.

The start of flight of mature adult dragonflies (Aeshna spp.) was investigated using electrophysiological and photographic techniques already described. Flight begins with either an upwards or a downwards movement of the wings. Neuromuscular activity and wing vibrations resembling ‘warm-up’ in moths is reported in two species of Aeshna.     

Mechanism of the wing colouration in the dragonfly Zenithoptera lanei (Odonata: Libellulidae) and its role in intraspecific communication

Zenithoptera dragonflies are known for their remarkable bluish colouration on their wings and unique male behaviour of folding and unfolding their wings while perching. However, nothing is known about the optical properties of such colouration and its structural and functional background. In this paper, we aimed to study the relationship between the wing membrane ultrastructure, surface microstructure and colour spectra of male wings in Zenithoptera lanei and test the hypothesis that colouration functions as a signal in territorial fights between males. The results show that the specific wing colouration derives from interference in alternating layers of melanized and unmelanized cuticle in the wing membrane, combined with diffuse scattering in two different layers of wax crystals on the dorsal wing surface, one lower layer of long filaments, and one upper layer of leaf-shaped crystals. The results also show that the thicker wax coverage of the dorsal surface of the wings results in increased brightness and reduced chroma. In the field experiments, we have demonstrated that there is a reduction of aggressive reactions of rivals towards individuals with experimentally reduced amount of blue wing colouration.     

Relationship between morphology,dispersal and habitat distribution in three species of Libellula (Odonata: Anisoptera)

Morphology is an important determinant of flight performance and can shape species’ dispersal behaviour. This study contrasted the morphology of flight-related structures in dragonfly species with different dispersal behaviours to gain insights into the relationship between morphology and dispersal behaviour. Specifically, wing size, wing shape and thorax size were compared in three co-occurring species from different clades within the genus Libellula (Odonata: Anisoptera: Libellulidae) to assess how these morphological traits are related to differences in dispersal behaviour and to how broadly their larvae occur across a habitat gradient. Two species had broad larval habitat distributions as well as high rates and distances of dispersal. These two species had relatively larger wings and thoraces than the third species, which was found only in permanent lakes and had limited dispersal. The hind-wings of more dispersive species also had lower aspect ratios and a relatively wider basal portion of the wing than the less dispersive species. Broad hind-wings may facilitate the use of gliding flight and reduce the energetic costs of dispersal. Determining the morphological traits associated with alternative dispersal behaviours may be a useful tool to assess the differential dispersal capacities of species or populations.     

Wing‐beat characteristics of birds recorded with tracking radar and cine camera

This study presents wing‐beat frequency data measured mainly by radar, complemented by video and cinematic recordings, for 153 western Palaearctic and two African species. Data on a further 45 Palaearctic species from other sources are provided in an electronic appendix. For 41 species with passerine‐type flight, the duration of flapping and pausing phases is given. The graphical presentations of frequency ranges and wing‐beat patterns show within‐species variation and allow easy comparison between species, taxonomic groups and types of flight. Wing‐beat frequency is described by Pennycuick (J. Exp. Biol. 2001; 204: 3283–3294) as a function of body‐mass, wing‐span, wing‐area, gravity and air density; for birds with passerine‐type flight the power‐fraction has also to be considered. We tested Pennycuick’s general allometric model and estimated the coefficients based on our data. The general model explained a high proportion of variation in wing‐beat frequency and the coefficients differed only slightly from Pennycuick’s original values. Modelling continuous‐flapping flyers alone resulted in coefficients not different from those predicted (within 95% intervals). Doing so for passerine‐type birds resulted in a model with non‐significant contributions of body‐mass and wing‐span to the model. This was mainly due to the very high correlation between body‐mass, wing‐span and wing‐area, revealing similar relative scaling properties within this flight type. However, wing‐beat frequency increased less than expected with respect to power‐fraction, indicating that the drop in flight level during the non‐flapping phases, compensated by the factor (g/q)^0.5 in Pennycuick’s model, is smaller than presumed. This may be due to lift produced by the body during the bounding phase or by only partial folding of the wings.     

Measurement on Camber Deformation of Wings of Free-flying Dragonflies and Beating-flying Dragonflies

1　IntroductionNumerouskinematicparameters,includingwing beatfrequency ,wingorientation ,andbothspan andchord wisedeformation ,arerelevanttotheaerodynam icanalysisofinsectflight[1,2 ] .Althoughnearlyalltherecentstudiesofinsectflightaerodynamics[3,4 ] haveidentifiedthatthemechanismsrequireflowseparationattheleadingedge ,andcamberisnotexpectedtohaveanysignificantinfluenceonthemagnitudeoftheforcecoefficient,someinsects ,suchasdragonfliesandbut terflies,frequently glideusinglowanglesofattack ,lead…     

Wing Dimorphisms and the Evolution of Migratory Polymorphisms among the Insecta

Many species of insects exhibit wing dimorphism, one morph havingfully developed wings and the other morph having reduced wingsand being incapable of flight. These wing dimorphisms providevisible manifestations of migratory polymorphisms. Since wingedindividuals do not, in principle, have to fly, the existenceof forms with reduced wings suggests that there is a tradeoffbetween flight capability and other fitness components. Comparisonsof the life histories of the fully winged and wing reduced morphsdemonstrate that this tradeoff is most commonly expressed asa decrease in the age of first reproduction and increased fecundityin the morph with reduced wings. Given these tradeoffs, theevolution of wing dimorphism will depend upon its genetic basis,including correlations with other life history components. Areview of the recent literature suggests that the heritabilityof wing morphology is high, and we suggest that this high heritabilitycould be maintained, in part, by antagonistic pleiotropy. In dimorphic species, the winged morph is generally consideredto be the migrant form. However, there are significant correlations,both within and among species, between the proportion of wingedindividuals, the proportion of winged individuals with functionalflight muscles, and the flight propensity of those individuals.This suggests that the proportion of winged individuals andthe propensity of the winged morph to migrate are intimatelyconnected at both the physiological and population level. Therefore,the study of the evolution of wing dimorphism is important notonly in its own right but also as a model of how migratory propensityevolves in monomorphically winged species.     

Habitat variation and wing coloration affect wing shape evolution in dragonflies

Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore‐ and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females.     

Dynamics of animal movement in an ecological context: dragonfly wing damage reduces flight performance and predation success

Much of our understanding of the control and dynamics of animal movement derives from controlled laboratory experiments. While many aspects of animal movement can be probed only in these settings, a more complete understanding of animal locomotion may be gained by linking experiments on relatively simple motions in the laboratory to studies of more complex behaviours in natural settings. To demonstrate the utility of this approach, we examined the effects of wing damage on dragonfly flight performance in both a laboratory drop–escape response and the more natural context of aerial predation. The laboratory experiment shows that hindwing area loss reduces vertical acceleration and average flight velocity, and the predation experiment demonstrates that this type of wing damage results in a significant decline in capture success. Taken together, these results suggest that wing damage may take a serious toll on wild dragonflies, potentially reducing both reproductive success and survival.     

Flight motor pattern in flying and non-flying Phasmida

Summary The insect order Phasmida comprises species with a broad spectrum of wing morphism and flight ability. By monitoring the electrical activity of several pterothoracic muscles the motor output during tethered flight was recorded for several Phasmida, ranging from excellent fliers to non-winged species. Both winged and non-winged species can generate a motor pattern as judged by criteria used to identify the locust flight pattern. However, in non-fliers the probability of expressing this pattern, its duration and precision are reduced. The antagonistic activity of the chosen muscle pairs is clearly different from the motor output during leg movements, which argues for specific motoneuronal coordination released for different behavioural performances. The demonstration of flight motor output in all tested Phasmida indicates that neural structures including their functional connectivity can be maintained independently of the appropriate peripheral structures. With respect to evolution this supports the idea that central neuronal interactions can be more conservative compared to changes in the periphery. Abbreviations of species names and indication of sexes are given in the first paragraph of Results     

alpha-glycerophosphate dehydrogenase activity in flight muscles of triatomine bugs Panstrongylus megistus and Triatoma sordida

The alpha-glycerophosphate dehydrogenase (alpha-GPDH) activity in flight muscles of Panstrongylus megistus and Triatoma sordida, vectors of Chagas disease in Brazil, was studied. Both species showed higher enzymatic activities in fliers than in non-fliers insects. T. sordida exhibited a higher proportion of flier insects than P. megistus. A possible role of alpha-GPDH on triatomines flight is discussed.     

Wing size but not wing shape is related to migratory behavior in a soaring bird

Both wing size and wing shape affect the flight abilities of birds. Intra and inter‐specific studies have revealed a pattern where high aspect ratio and low wing loading favour migratory behaviour. This, however, have not been studied in soaring migrants. We assessed the relationship between the wing size and shape and the characteristics of the migratory habits of the turkey vulture Cathartes aura, an obligate soaring migrant. We compared wing size and shape with migration strategy among three fully migratory, one partially migratory and one non‐migratory (resident) population distributed across the American continent. We calculated the aspect ratio and wing loading using wing tracings to characterize the wing morphology. We used satellite‐tracking data from the migratory populations to calculate distance, duration, speed and altitude during migration. Wing loading, but not aspect ratio, differed among the populations, segregating the resident population from the completely migratory ones. Unlike what has been reported in species using flapping flight during migration, the migratory flight parameters of turkey vultures were not related to the aspect ratio. By contrast, wing loading was related to most flight parameters. Birds with lower wing loading flew farther, faster, and higher during their longer journeys. Our results suggest that wing morphology in this soaring species enables lower‐cost flight, through low wing‐loading, and that differences in the relative sizes of wings may increase extra savings during migration. The possibility that wing shape is influenced by foraging as well as migratory flight is discussed. We conclude that flight efficiency may be improved through different morphological adaptations in birds with different flight mechanisms.