What factors shape female phenotypes of a poleward‐moving damselfly at the edge of its range?

Individuals at the expansion front during a climate‐driven range expansion are expected to differ phenotypically from those individuals in core populations. Little information is known about the joint, potentially opposing, effects of stressful conditions at the range edge versus evolutionary changes that take place during range expansion in shaping the phenotypes at the range front. We investigated the effect of range expansion on immune function, body condition and flight‐related morphology (flight muscle ratio, wing loading, and wing aspect ratio) of field‐collected females of the poleward‐moving damselfly Coenagrion scitulum. Individuals at the expansion front had a lower body condition, which indicated more stressful conditions at the range edge. Despite the counteracting effect of the shorter growth season, the higher flight muscle ratios at the expansion front indicated a strong selection for dispersal ability during range expansion. The current study suggests that models need to incorporate the interplay of stressful conditions and evolutionary processes at the expansion front to arrive at robust predictions of future species distributions under global warming. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 556–568.     

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.     

Wing dimorphism in the migratory locust,Locusta migratoria: differentiation of wing morph and phase polyphenism

A short‐winged morph was recently discovered in the migratory locust, Locusta migratoria. It is different from the normal, long‐winged morph not only in forewing length but also in hind femur length, displaying a dimorphism. To understand the significance of this dimorphism, other morphological characters were compared between the two morphs, and the time of differentiation of wing‐pad length was investigated. Wing weights were heavier in the long‐winged morph than in the short‐winged morph. This result showed that the short‐winged morph is not formed by a failure of wing expansion. No obvious morph‐specific differences were observed in wing venation, but wing allometry studies indicated that the distal areas of the fore‐ and hindwings were disproportionally reduced in the short‐winged morph compared to the long‐winged morph. The morphological differentiation of the wing pad between the two morphs was observed at the penultimate nymphal stage. The flight muscle was well developed in the two morphs, and no sign of flight muscle histolysis was detected in either morph after adult emergence. An analysis of adult body dimensions suggested that the density‐dependent phase shifts known for the long‐winged morph of this locust were also exhibited by the short‐winged morph, demonstrating that these shifts are not specific to the migratory long‐winged morph.     

Soaring and manoeuvring flight of a steppe eagle Aquila nipalensis

We used an onboard inertial measurement unit, together with onboard and ground‐based video cameras, to record the movements of the body, wings and tail of a steppe eagle Aquila nipalensis during wide‐ranging flight. The eagle's flight consisted of a more or less continuous sequence of banked turns, interrupted by occasional wing tucks and roll‐over manoeuvres, and ultimately terminated by a wing‐over manoeuvre leading in to a diving landing approach. The flight configuration of the bird, and its pattern of movement during angular perturbations, together suggest that the eagle is inherently stable in pitch and yaw, and perhaps also in roll. The control inputs used to generate roll moments during banked turns were too subtle to be detected. Control of yaw and pitch during banked turns involved a consistent pattern of tail movement, wherein the tail was spread and depressed immediately before the turn, and then overbanked with respect to the bird during the latter part of the turn. Differential adjustment of wing posture is probably also involved in the control of banked turns, but it was only consistently apparent during more extreme roll manoeuvres. For example, roll‐over and wing‐over manoeuvres were both accomplished by differential changes in the angle of incidence and spread of the wings. In general, however, the bird appeared to maintain positive loading on its wings at all times, except during extreme flight manoeuvres.     

Host‐plant dependent wing phenotypic variation in the neotropical butterfly Heliconius erato

Most phytophagous insects feed on a single plant during development, and this may influence not only performance‐linked traits, but also more subtle morphological differences. Insect–plant interactions are thus valuable for studying environmental influences on phenotypes. By using geometric morphometrics, we investigated the variation in forewing size and shape in the butterfly Heliconius erato phyllis reared on six species of passion vines (Passiflora spp.). We detected wing shape sexual dimorphism, for which the adaptive significance deserves further investigation. There was size as well as wing shape variation among individuals fed on different hosts. These subtle differences in shape were interpreted as environmental effects on development, which should be under weak natural selection for these traits, and therefore not strongly canalized. This result reinforces the role of plasticity on host‐plant use, as well as the corresponding consequences on developmental variability among phytophagous insects. We propose that this variation can be an important factor in resource specialization and partner recognition, possibly triggering reproductive isolation and sympatric speciation in phytophagous insects. This interaction also shows itself as a good model for studying the role of environmental and interaction diversity in evolution. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 765–774.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Phylogenetic reassessment of the five forewing radial-veined tribes of Riodininae (Lepidoptera: Riodinidae)

The relationships among the genera and tribal groupings of Riodininae with five forewing radial veins, and between these and tribes with four forewing radial veins, were examined using a phylogenetic analysis. Using the type species from all sixteen genera in the tribal groupings Eurybiini, Mesosemiini and incertae sedis (a presumed paraphyletic group of loosely related genera), and representatives from the four forewing radial‐veined riodinine tribes, thirty‐five new and traditional characters were coded from adult ecology, wing venation and pattern, the adult head and body, male and female genitalia, and early stage ecology and morphology. The majority of characters are illustrated. Phylogenetic analysis of these data produced five equally most parsimonious cladograms using equal weights and after successive weighting. The strict consensus of these confirms the monophyly of Eurybiini and Mesosemiini as currently conceived, but also indicates several higher‐level relationships not previously hypothesized. Mesosemiini is here more broadly defined to also include the entire incertae sedis section, and the tribe is divided into Mesosemiina, for the previously delimited Mesosemiini plus Eunogyra and Teratophthalma, and Napaeina, subtr.n. for the incertae sedis section minus these two genera. The following hypothesis of relationships is tentatively proposed for the basal clades of Riodininae: Mesosemiini + (Eurybiini + remainder of Riodininae). These new hypotheses, and the characters supporting them, are discussed and compared with those previously proposed.     

Direct and correlated responses to artificial selection on flight activity in the oriental fruit moth (Lepidoptera: Tortricidae)

The ability of a sufficient number of individuals to disperse is crucial for long‐term survival of populations. However, dispersal is often energetically costly, and thus is expected to trade‐off against other life‐history traits. In insect pest species, the occurrence of individuals with high flight activity challenges management practices. We performed artificial selection on flight activity and measured correlated responses to selection in the oriental fruit moth, Grapholita (= Cydia) molesta, a widely distributed and expanding lepidopteran pest of fruit crops. Both sexes rapidly responded to the imposed regime of divergent selection, indicating an adaptive potential of flight activity in this species. Upward‐selected moths died sooner than downward‐selected ones, providing evidence for a cost of flight activity to adult survival, reputedly associated with enhanced metabolic rates. Oppositely‐selected females had similar total reproductive output, disproving a trade‐off between dispersal and reproduction, although females with higher flight activity laid their eggs sooner. The ratio of body weight to forewing surface (forewing loading) did not significantly differ between selected lines. The present study contributes to the understanding of dispersal evolution, and also provides new insights into life‐history theory as well as important baseline data for the improvement of pest management practices. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 879–889.     

Subtle variation in size and shape of the whole forewing and the red band among co‐mimics revealed by geometric morphometric analysis in Heliconius butterflies

Heliconius are unpalatable butterflies that exhibit remarkable intra‐ and interspecific variation in wing color pattern, specifically warning coloration. Species that have converged on the same pattern are often clustered in Müllerian mimicry rings. Overall, wing color patterns are nearly identical among co‐mimics. However, fine‐scale differences exist, indicating that factors in addition to natural selection may underlie wing phenotype. Here, we investigate differences in shape and size of the forewing and the red band in the Heliconius postman mimicry ring (H. erato phyllis and the co‐mimics H. besckei, H. melpomene burchelli, and H. melpomene nanna) using a landmark‐based approach. If phenotypic evolution is driven entirely by predation pressure, we expect nonsignificant differences among co‐mimics in terms of wing shape. Also, a reinforcement of wing pattern (i.e., greater similarity) could occur when co‐mimics are in sympatry. We also examined variation in the red forewing band because this trait is critical for both mimicry and sexual communication. Morphometric results revealed significant but small differences among species, particularly in the shape of the forewing of co‐mimics. Although we did not observe greater similarity when co‐mimics were in sympatry, nearly identical patterns provided evidence of convergence for mimicry. In contrast, mimetic pairs could be distinguished based on the shape (but not the size) of the red band, suggesting an “advergence” process. In addition, sexual dimorphism in the red band shape (but not size) was found for all lineages. Thus, we infer that natural selection due to predation by birds might not be the only mechanism responsible for variation in color patterns, and sexual selection could be an important driver of wing phenotypic evolution in this mimicry ring.     

New fossil mousebird (Aves: Coliiformes) with feather preservation provides insight into the ecological diversity of an Eocene North American avifauna

Coliiformes (mousebirds) are represented by just six extant species. These species, restricted to sub‐Saharan Africa, are all primarily frugivorous and are among the most sedentary of living birds. Previously described fossil Coliiformes preserving feather traces share the short, rounded wing shape of extant mousebirds. Along with osteological evidence, these observations have been proposed to support poor sustained flight capabilities across the stem mousebird lineage. We report a new species of Coliiformes from the early Eocene (51.66 ± 0.09 Ma) Fossil Butte Member of the Green River Formation, represented by one of the comparatively few fossils from these deposits preserving carbonized traces of the wing and tail feathering. Feather traces indicate an elongate, tapering wing shape similar to that of some extant aerial insectivores, and suggestive of a capacity for sustained and agile open‐air flight. Traces of the rectrices reveal the tail accounted for approximately two‐thirds of the total length of the bird, a proportion similar to that in living mousebirds. Phylogenetic analysis places the new species as a stem representative of Coliiformes, demonstrating for the first time that the two major clades of Coliiformes – Sandcoleidae and Colii – co‐occurred at Fossil Lake. Based on the recovered phylogeny, as well as the osteology and feathering of extant and fossil Coliiformes, the wing shape of the new species is interpreted as apomorphic. In addition to documenting unexpected morphological specialization within stem‐lineage Coliiformes, the new species adds yet another taxon to the emerging reconstruction of the diverse Paleogene avifauna from the tightly dated and nearly synchronous fossiliferous deposits of the Fossil Butte Member. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 685–706.     

The effects of latitude,body size,and sexual selection on wing shape in a damselfly

Under natural selection, wing shape is expected to evolve to optimize flight performance. However, other selective factors besides flight performance may influence wing shape. One such factor could be sexual selection in wing sexual ornaments, which may lead to alternative variations in wing shape that are not necessarily related to flight performance. In the present study, we investigated wing shape variations in a calopterygid damselfly along a latitudinal gradient using geometric morphometrics. Both sexes show wing pigmentation, which is a known signal trait at intra‐ and interspecific levels. Wing shape differed between sexes and, within the same sex, the shape of the hind wing differed from the front wing. Latitude and body size explained a high percentage of the variation in wing shape for female front and hind wings, and male front wings. In male hind wings, wing pigmentation explained a high amount of the variation in wing shape. On the other hand, the variation in shape explained by pigmentation was very low in females. We suggest that the conservative morphology of front wings is maintained by natural selection operating on flight performance, whereas the sex‐specific differences in hind wings most likely could be explained by sexual selection. The observed sexual dimorphism in wing shape is likely a result of different sex‐specific behaviours. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 263–274.     

Diversity of wing patterns and abdomen‐generated substrate sounds in 3 European scorpionfly species

In the genus Panorpa (Insecta: Mecoptera), also known as scorpionflies, premating behavior includes repeated sequences of slow wing movements (waving, fanning, flagging) which are accompanied by rapid abdomen vibrations that generate substantial substrate‐borne sound. It is still unknown whether wing patterns or vibratory signals contain information about species identity, sex and/or the quality of potential mating partners. Besides species‐specific pheromones, these multimodal signals may be of particular importance for the maintenance of reproductive isolation in sympatrically occurring scorpionfly species. Here, we analyzed phyologenetic relationships among, and the pattern of forewings as well as substrate‐borne sound in 3 different sympatric Central‐European scorpionfly species (P. communis, P. germanica, and P. alpina). Divergence time estimates, based on 879 bp of the mitochondrial COI gene, indicate longstanding separate evolutionary histories for the studied Panorpa species. Morphological analysis revealed that wing length as an indicator of body size increased in the following order: P. alpina < P. germanica < P. communis. Individuals can be assigned to the correct species and sex with high accuracy just by evaluation of the number of dark spots and the proportion of wing pigmentation. Despite high variability of interpulse period at an individual level, across species analysis revealed a positive correlation of average interpulse period as well as mean signal amplitude with forewing length. These results suggest wing patterns, but less likely vibratory signals, to contain information about species identity. Furthermore, receivers may be able to estimate the body size of a signaler solely on the basis of substrate‐borne sound.     

High lift function of the pteroid bone and forewing of pterosaurs

The pteroid bone is a rod-like element found only in pterosaurs, the flying reptiles of the Mesozoic. It articulated at the wrist, and supported a membranous forewing in front of the inner part of the wing spar. The function of this bone, particularly its orientation, has been much debated. It is widely believed that it pointed towards the body, and that the forewing was relatively narrow. An alternative hypothesis states that it was directed forwards during flight, resulting in a much broader forewing that acted as a leading edge flap. We tested scale models in a wind tunnel to determine the aerodynamic consequences of these conflicting hypotheses, and found that performance is greatly improved if the pteroid is directed forwards: the lift: drag ratios are superior and the maximum lift is exceptionally high in comparison with conventional aerofoils. This high lift capability may have enabled even the largest pterosaurs to take off and land without difficulty.     

Lack of morphological coevolution between male forelegs and female wings in Themira (Sepsidae: Diptera: Insecta)

The complex, species-specific foreleg armature in males of the genus Themira (Diptera: Sepsidae) provides an ideal system for testing competing hypotheses for the evolution of sexually dimorphic character divergence. In sepsid flies, the male holds onto the female by clasping her wing base with his modified forelegs. In the present study, we document the male leg and the female wing morphology using scanning electron microscopy and confocal microscopy. We use a phylogenetic tree for Themira to reconstruct male foreleg and female wing evolution and demonstrate that the male legs have evolved elaborate structures with little or no corresponding changes in wing morphology. This lack of interspecific variation in female wings is not in agreement with the hypothesis of a morphological 'evolutionary arms race' between males and females. However, there is also no evidence for sex-specific wing differences in sensory organs on the wing base that may explain how females could assess males according to Eberhard's 'cryptic female choice' hypothesis. Finally, our study reveals the function of several novel morphological clasping structures and documents that the male foreleg characters in Themira are highly homoplastic. Male forelegs in two clades evolve considerably faster than in other species or clades. These two clades include Themira superba and Themira leachi , species that have some of the most dramatically modified forelegs known in Diptera.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 227–238.     

Sexual dimorphism and intraspecific variation in wing size and shape of Tongeia fischeri (Lepidoptera: Lycaenidae)

Wing morphological variations are described here for the lycaenid butterfly Tongeia fischeri. A landmark‐based geometric morphometric approach based on wing venation of 197 male and 187 female butterflies collected in Japan was used to quantify wing size and shape variations between sexes and among populations. Sexual dimorphism in wing size and shape was detected. Females had significantly larger wings than males, while males showed a relatively elongated forewing with a longer apex and narrower wing tornus in comparison to females. Intraspecific variations in wing morphology among populations were revealed for the wing shape, but not wing size. Distinct wing shape differences were found in the vein intersections area around the distal part of the discal cell where median veins originated in the forewing and around the origin of the CU1 vein in the hindwing. In addition, phenotypic relationships inferred from wing shape variations grouped T. fischeri populations into three groups, reflecting the subspecies classification of the species. The spatial variability and phenotypic relationships between conspecific populations of T. fischeri detected here are generally in agreement with the previous molecular study based on mitochondrial and nuclear sequences, suggesting the presence of a phylogenetic signal in the wing shape of T. fischeri, and thus having taxonomic implications.     

Non-Jumping Take off Performance in Beetle Flight （Rhinoceros Beetle Trypoxylus dichotomus）

In recent decades, the take-off mechanisms of flying animals have received much attention in insect flight initiation. Most of previous works have focused on the jumping mechanism, which is the most common take-off mechanism found in flying animals. Here, we presented that the rhinoceros beetle, Trypoxylus dichotomus, takes offwithout jumping. In this study, we used 3-Dimensional （3D） high-speed video techniques to quantitatively analyze the wings and body kinematics during the initiation periods of flight. The details of the flapping angle, angle of attack of the wings and the roll, pitch and yaw angles of the body were investigated to understand the mechanism of take-off in T. dichotomus. The beetle took off gradually with a small velocity and small acceleration. The body kinematic analyses showed that the beetle exhibited stable take-off. To generate high lift force, the beetle modulated its hind wing to control the angle of attack; the angle of attack was large during the upstroke and small during the downstroke. The legs of beetle did not contract and strongly release like other insects. The hind wing could be con- sidered as a main source of lift for heavy beetle.     

The genus Huperzia (Lycopodiaceae) in the Azores and Madeira

The taxonomy and nomenclature of the genus Huperzia Bernh. in the Azores and Madeira have been reviewed. Plants collected in the Azores and Madeira were characterized morphologically. The independence between two endemic species common to Madeira and the Azores Islands –Huperzia suberecta (Lowe) Tardieu and Huperzia dentata (Herter) Holub – is clearly shown. A clear‐cut morphological separation between these taxa and Huperzia selago (L.) Bernh. ex Schrank & Mart. of continental Europe is established. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 522–533.     

Mate-locating strategies are related to relative body length and wing colour in the speckled wood butterfly Pararge aegeria

1. The mate-locating strategies of Pararge aegeria (L.) males were studied in relation to adult morphology (dorsal wing colour, forewing length, body length and forewing length : body length ratio) and generation.
2. Males locate females either by perching and defending territories, or by patrolling. Individuals were more consistent in their mating strategies than expected by chance.
3. Forewing length : body length ratio was positively correlated with thorax mass : body mass; relatively short-bodied males had relatively heavy thoraxes. Therefore, forewing length : body length ratio was an index of mass allocation.
4. Perching males had higher forewing length : body length ratios and were paler than patrolling males.
5. The higher forewing length : body length ratio was due to the differences in body length and not wing length. Perchers had shorter bodies than patrollers.     

Wing morphology is related to host plants in cactophilic Drosophila gouveai and Drosophila antonietae (Diptera,Drosophilidae)

A central issue in evolutionary biology is to understand the mechanisms promoting morphological evolution during speciation. In a previous study, we showed that the Neotropical cactophilic sibling species Drosophila gouveai and Drosophila antonietae can be reared in media prepared with their presumptive natural host plants (Pilosocereus machrisis and Cereus hildmaniannus) and that egg to adult viability is not independent of the cactus host. In the present study, we investigate the effects of ecological and genetic factors on interspecific divergence in wing morphology, in relation to the pattern of wing venation and phenotypic plasticity in D. gouveai and D. antonietae, by means of the comparative analysis of isofemale lines reared in the two cactus hosts. The species differed significantly in wing size and shape, although specific differences were mainly localized in a particular portion of the wing. We detected significant variation in form among lines, which was not independent of the breeding cactus, suggesting the presence of genetic variation for phenotypic plasticity and wing shape variation in both species. We discuss the results considering the plausible role of host plant use in the evolutionary history of cactophilic Drosophila inhabiting the arid zones of South America. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 655–665.