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1.
Butterflies display extreme variation in wing shape associated with tremendous ecological diversity. Disentangling the role of neutral versus adaptive processes in wing shape diversification remains a challenge for evolutionary biologists. Ascertaining how natural selection influences wing shape evolution requires both functional studies linking morphology to flight performance, and ecological investigations linking performance in the wild with fitness. However, direct links between morphological variation and fitness have rarely been established. The functional morphology of butterfly flight has been investigated but selective forces acting on flight behaviour and associated wing shape have received less attention. Here, we attempt to estimate the ecological relevance of morpho‐functional links established through biomechanical studies in order to understand the evolution of butterfly wing morphology. We survey the evidence for natural and sexual selection driving wing shape evolution in butterflies, and discuss how our functional knowledge may allow identification of the selective forces involved, at both the macro‐ and micro‐evolutionary scales. Our review shows that although correlations between wing shape variation and ecological factors have been established at the macro‐evolutionary level, the underlying selective pressures often remain unclear. We identify the need to investigate flight behaviour in relevant ecological contexts to detect variation in fitness‐related traits. Identifying the selective regime then should guide experimental studies towards the relevant estimates of flight performance. Habitat, predators and sex‐specific behaviours are likely to be major selective forces acting on wing shape evolution in butterflies. Some striking cases of morphological divergence driven by contrasting ecology involve both wing and body morphology, indicating that their interactions should be included in future studies investigating co‐evolution between morphology and flight behaviour.  相似文献   

2.
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.  相似文献   

3.
Wing morphology is known to strongly affect flight performance by affecting lift and drag during flight. Performance may consequently deteriorate during feather moult due to the creation of feather gaps in the wing. Since wing gap size may directly affect the extent of reduced flight capacity, rapid moult involving the creation of large feather gaps is expected to substantially impair flight compared with the small gaps induced by a slower moult. To examine the factors affecting wing-feather moult speed, we studied adults of nineteen resident or very short-distance migrant passerine species during their post-breeding moult using a model-selection framework following a phylogenetically controlled analysis. We examined the speed of wing-feather moult in relation to each species’ flight distance index that was estimated based on local foraging movements rather than on longer flights (e.g., migration), assessed by the Delphi technique of expert evaluation. Moult speed was also examined with respect to six morphometric variables: body mass, wing loading, the feather comprising the tip of the wing, aspect ratio, wing span, and wing area. Our results suggest that flight distance index is the most important factor determining the speed of wing-feather moult in songbirds. Species that regularly fly a shorter distance were found to moult quickly, and those that take relatively longer flights moult slowly. These results suggest that the aerodynamic cost of wing area reduction due to feather moult shapes the evolution of annual routine processes by dictating a slower moult speed (resulting in small wing gaps) for species that regularly fly long distances and consequently may be affected more substantially by large wing gaps compared with short distance flyers.  相似文献   

4.
The selective pressures associated with flight are significant factors in shaping the morphology of volant forms. Tropical seabirds are of particular interest because of their long foraging bouts, which can last hundreds of kilometers in search of unpredictable (spatially and temporally) resources. Here, we contrast wing loading (WL), aspect ratio (AR), and planform shape among five pelecaniform seabirds and correlate morphological diversity with known differences in flight strategies. Overall, WL and AR scaled with body mass. The Great Frigratebird had lower WL than that predicted, whereas the Red-tailed Tropicbird had higher WL than that predicted. The tropicbird also exhibited a lower AR than that predicted. Visualization of planform shape was accomplished by using Thin-plate spline relative warp analysis (TPS/RWA), and three major regions of variations were discovered: wing base, mid-wing, and distal wing/wing tip. As expected, the three boobies were more similar than either the tropicbird or the frigatebird. The tropicbird had a broader distal wing and more rounded wing tip, associated with its greater use of flapping flight. The frigatebird showed the greatest deviation in the distal wing and wing tip associated with the high maneuverability required for aerial pursuit and kleptoparasitism. By using TPS/RWA, important differences were detected in planform shape that would have otherwise gone unnoticed when using only WL and AR. These differences correlated strongly with parameters such as maneuverability, flapping, and soaring flight.  相似文献   

5.
Most studies of lean mass dynamics in free-living passerine birds have focused on Old World species at geographical barriers where they are challenged to make the longest non-stop flight of their migration. We examined lean mass variation in New World passerines in an area where the distribution of stopover habitat does not require flights to exceed more than a few hours and most migrants stop flying well before fat stores near exhaustion. We used either quantitative magnetic resonance (QMR) analysis or a morphometric model to measure or estimate, respectively, the fat and lean body mass of migrants during stopovers in New York, USA. With these data, we examined (1) variance in total body mass explained by lean body mass, (2) hourly rates of fat and lean body mass change in single-capture birds, and (3) net changes in fat and lean mass in recaptured birds. Lean mass contributed to 50% of the variation in total body mass among white-throated sparrows Zonotrichia albicollis and hermit thrushes Catharus guttatus. Lean mass of refueling gray catbirds Dumetella carolinensis and white-throated sparrows, respectively, increased 1.123 and 0.320 g h−1. Lean mass of ovenbirds Seiurus aurocapillus accounted for an estimated 33–40% of hourly gains in total body mass. On average 35% of the total mass gained among recaptured birds was lean mass. Substantial changes in passerine lean mass are not limited to times when birds are forced to make long, non-stop flights across barriers. Protein usage during migration is common across broad taxonomic groups, migration systems, and migration strategies.  相似文献   

6.
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.  相似文献   

7.
The optimum body mass of passerine birds typically represents a trade‐off between starvation risk, which promotes fat gain, and predation pressure, which promotes fat loss to maintain maneuvrability. Changes in ecological factors that affect either of these variables will therefore change the optimum body masses of populations of passerine birds. This study sought to identify and quantify the effects of changing temperatures and predation pressures on the body masses and wing lengths of populations of passerine birds throughout Britain and Ireland over the last 50 years. We analyzed over 900,000 individual measurements of body mass and wing length of blue tits Cyanistes caeruleus, coal tits Periparus ater, and great tits Parus major collected by licenced bird ringers throughout Britain and Ireland from 1965 to 2017 and correlated these with publicly available temperature data and published, UK‐wide data on the abundance of a key predator, the sparrowhawk Accipiter nisus. We found highly significant, long‐term, UK‐wide decreases in winter body masses of adults and juveniles of all three species. We also found highly significant negative correlations between winter body mass and winter temperature, and between winter body mass and sparrowhawk abundance. Independent of these effects, body mass further correlated negatively with calendar year, suggesting that less well understood dynamic factors, such as supplementary feeding levels, may play a major role in determining population optimum body masses. Wing lengths of these birds also decreased, suggesting a hitherto unobserved large‐scale evolutionary adjustment of wing loading to the lower body mass. These findings provide crucial evidence of the ways in which species are adapting to climate change and other anthropogenic factors throughout Britain and Ireland. Such processes are likely to have widespread implications as the equilibria controlling evolutionary optima in species worldwide are upset by rapid, anthropogenic ecological changes.  相似文献   

8.
The wing morphology of bats is very diverse, and may correlate with energetic, behavioural, and ecological demands. If these demands conflict, wing shape may reflect compromise solutions. In this study, we compared the wing morphology of two bats,Tadarida brasiliensis (Geoffroy, 1824) andMyotis chiloensis (Waterhouse, 1828), that differ in body size, habitat, and foraging behaviour. We analyzed features of biomechanical and energetic relevance, and sought evidence of compromise solutions to energetic, ecological, and behavioural demands. We found that wing span of both species conformed to expectations based on allometric relationships, but that although the wing area ofM. chiloensis did not differ from predictions, the wing area ofT. brasiliensis was lower.M. chiloensis possessed an unusually low second moment of area of the humerus. Wing form ofM. chiloensis is consistent with highly maneuverable flight needed to live between shrubs and wooded habitats, and its low aspect ratio and low wing loading indicate a high energetic cost and a low flight speed, respectively. The low humeral second moment of area may be related to a reduction of wing mass and may result in decreased inertial power. In contrast,T. brasiliensis showed high aspect ratio and wing loading, characteristic of high speed, energetically economic flight.  相似文献   

9.
The morphology of the avian skeleton is often studied in the context of adaptations for powered flight. The effects of other evolutionary forces, such as sexual selection, on avian skeletal design are unclear, even though birds produce diverse behaviors that undoubtedly require a variety of osteological modifications. Here, we investigate this issue in a family of passerine birds called manakins (Pipridae), which have evolved physically unusual and elaborate courtship displays. We report that, in species within the genus Manacus, the shaft of the radius is heavily flattened and shows substantial solidification. Past work anecdotally notes this morphology and attributes it to the species' ability to hit their wings together above their heads to produce loud mechanical sonations. Our results show that this feature is unique to Manacus compared to the other species in our study, including a variety of taxa that produce other sonations through alternate wing mechanisms. At the same time, our data reveal striking similarities across species in total radius volume and solidification. Together, this suggests that supposedly adaptive alterations in radial morphology occur within a conserved framework of a set radius volume and solidness, which in turn is likely determined by natural selection. Further allometric analyses imply that the radius is less constrained by body size and the structural demands that underlie powered flight, compared to other forelimb bones that are mostly unmodified across taxa. These results are consistent with the idea that the radius is more susceptible to selective modification by sexual selection. Overall, this study provides some of the first insight into the osteological evolution of passerine birds, as well as the way in which opposing selective forces can shape skeletal design in these species. J. Morphol. 277:766–775, 2016. © 2016 Wiley Periodicals, Inc.  相似文献   

10.
We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.  相似文献   

11.
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.  相似文献   

12.
We studied the response in wing size to rearing at different temperatures of nine strains of Drosophila representing six species. The species varied in their natural habitats from tropical to temperate and one cosmopolitan. The evolutionary divergence of the species spans 50 million years. While some quantitative differences were found, all species responded to temperature very similarly: females increased an average of ∼11% and males ∼14% when reared at 19 °C compared to 25 °C. The phenotypic plasticity in wing size in response to temperature appears to be a fixed trait in Drosophila across long evolutionary time and diverse ecological settings. This likely reflects the close relationship between wing area (and thus wing loading) and insect body mass that is a crucial factor for flight regardless of ecology and is, thus, maintained across long evolutionary time.  相似文献   

13.
The power that the birds can use for flight (available power) and the power required for flight according to physical laws (requisite power) grow with an increase in body mass, the exponents of the corresponding functions being different. Small birds can follow different strategies, either improving the aerodynamic quality of the body (thereby saving the excess available power) or sacrifice aerodynamic quality in favor of morphological adaptation to factors other than the demands of flight proper, which provides the possibility of utilizing a wider range of ecological niches. A hypothesis is proposed that the high metabolic rate of passerine birds, compared to representatives of other bird orders, is an adaptation to maneuverable (i.e., relatively low-speed) flight necessary for successful colonization of forest habitats. The speed that birds of such size can develop according to the scaling theory is too high for nesting and foraging in tree crowns, and its reduction is possible in two ways: by increasing air drag or by changing the style of flight (by analogy with airplane vs. helicopter). The first way is feasible, but a high air drag due to morphological modifications (e.g., in the size of the tail or characteristics of the wing) interferes with the possibility of long-distance migration flight, as energy expenditures for it will exceed the energy potential of the bird. This is why migratory nonpasserine birds, which have used this strategy, are practically absent in forests of the temperate zone. Therefore, more promising is the second way involving transition to a new flight style and, in a certain sense, to a new morphophysiological organization. Passerines have achieved this by changing their flight style so that the wing actively generates forces (lift and thrust) only in downstroke. Such a flight requires more energy, and, to provide it in sufficient amounts, passerine birds have increased their basal metabolic rate (BMR). Thus, both their flight energy expenditures and BMR are higher than in nonpasserines. Remarkably, among approximately 8660 extant bird species known today, more than half (about 5100 species) belong to the order Passeriformes. Such a ratio, unknown in any other vertebrate class, is evidence that passerines have gained a considerable biological advantage over all other birds due to their increased BMR.  相似文献   

14.
Bai M  McCullough E  Song KQ  Liu WG  Yang XK 《PloS one》2011,6(6):e21600
This study examines the evolution hindwing shape in Chinese dung beetle species using morphometric and phylogenetic analyses. Previous studies have analyzed the evolution of wing shape within a single or very few species, or by comparing only a few wing traits. No study has analyzed wing shape evolution of a large number of species, or quantitatively compared morphological variation of wings with proposed phylogenetic relationships. This study examines the morphological variation of hindwings based on 19 landmarks, 119 morphological characters, and 81 beetle species. Only one most parsimonious tree (MPT) was found based on 119 wing and body characters. To better understand the possible role of the hindwing in the evolution of Scarabaeinae, additional phylogenetic analyses were proposed based on the only body features (106 characters, wing characters excluded). Two MPT were found based on 106 body characters, and five nodes were collapsed in a strict consensus. There was a strong correlation between the morphometric tree and all phylogenetic trees (r>0.5). Reconstructions of the ancestral wing forms suggest that Scarabaeinae hindwing morphology has not changed substantially over time, but the morphological changes that do occur are focused at the base of the wing. These results suggest that flight has been important since the origin of Scarabaeinae, and that variation in hindwing morphology has been limited by functional constraints. Comparison of metric disparity values and relative evolutionary sequences among Scarabaeinae tribes suggest that the primitive dung beetles had relatively diverse hindwing morphologies, while advanced dung beetles have relatively similar wing morphologies. The strong correlation between the morphometric tree and phylogenetic trees suggest that hindwing features reflect the evolution of whole body morphology and that wing characters are suitable for the phylogenetic analyses. By integrating morphometric and cladistic approaches, this paper sheds new light on the evolution of dung beetle hind wings.  相似文献   

15.
Monarch butterflies (Danaus plexippus) are parasitized by the protozoan Ophryocystis elektroscirrha throughout their geographical range. Monarchs inhabiting seasonally fluctuating environments migrate annually, and parasite prevalence is lower among migratory relative to non‐migratory populations. One explanation for this pattern is that long‐distance migration weeds out infected animals, thus reducing parasite prevalence and transmission between generations. In this study we experimentally infected monarchs from a migratory population and recorded their long‐distance flight performance using a tethered flight mill. Results showed that parasitized butterflies exhibited shorter flight distances, slower flight speeds, and lost proportionately more body mass per km flown. Differences between parasitized and unparasitized monarchs were generally not explained by individual variation in wing size, shape, or wing loading, suggesting that poorer flight performance among parasitized hosts was not directly caused by morphological constraints. Effects of parasite infection on powered flight support a role for long‐distance migration in dramatically reducing parasite prevalence in this and other host–pathogen systems.  相似文献   

16.
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.  相似文献   

17.
Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)1/6 and (wing loading)1/2 among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, Ue) of 138 species, ranging 0.01–10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of Ue in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in Ue. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in Ue. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading.  相似文献   

18.
Wing size and shape, expressed as wing loading and aspect ratio respectively, together with bill morphology are parameters that can reveal differences related to the foraging ecology of seabirds. Six species of booby (Sulidae) that inhabit the Pacific are the focus of this study: four mainly pelagic species, Masked Booby Sula dactylatra, Nazca Booby Sula granti, Red‐footed Booby Sula sula and Brown Booby Sula leucogaster, and two coastal species, Blue‐footed Booby Sula nebouxii and Peruvian Booby Sula variegata. Pelagic boobies showed segregation among species in body mass and relative bill size, and they differed in wing morphology (wing loading and aspect ratio) from the coastal boobies. The coastal Peruvian and Blue‐footed Boobies are largely allopatric but overlap in northern Peru. In their area of sympatry, they showed evidence of character displacement in body size and in wing and bill morphology, which suggests that competition plays an important role in sympatry. This study improves our understanding of ecological interactions among Pacific boobies and of how selective pressures have shaped their ecomorphology and foraging behaviours.  相似文献   

19.
Wing morphology and flight kinematics profoundly influence foraging costs and the overall behavioral ecology of hummingbirds. By analogy with helicopters, previous energetic studies have applied the momentum theory of aircraft propellers to estimate hovering costs from wing disc loading (WDL), a parameter incorporating wingspan (or length) and body mass. Variation in WDL has been used to elucidate differences either among hummingbird species in nectar-foraging strategies (e.g., territoriality, traplining) and dominance relations or among gender-age categories within species. We first demonstrate that WDL, as typically calculated, is an unreliable predictor of hovering (induced power) costs; predictive power is increased when calculations use wing length instead of wingspan and when actual wing stroke amplitudes are incorporated. We next evaluate the hypotheses that foraging strategy and competitive ability are functions of WDL, using our data in combination with those of published sources. Variation in hummingbird behavior cannot be easily classified using WDL and instead is correlated with a diversity of morphological and physiological traits. Evaluating selection pressures on hummingbird wings will require moving beyond wing and body mass measurements to include the assessment of the aerodynamic forces, power requirements, and power reserves of hovering, forward flight, and maneuvering. However, the WDL-helicopter dynamics model has been instrumental in calling attention to the importance of comparative wing morphology and related aerodynamics for understanding the behavioral ecology of hummingbirds.  相似文献   

20.
Wing venation has long been used for insect identification. Lately, the characterization of venation shape using geometric morphometrics has further improved the potential of using the wing for insect identification. However, external factors inducing variation in wing shape could obscure specific differences, preventing accurate discrimination of species in heterogeneous samples. Here, we show that interspecific difference is the main source of wing shape variation within social wasps. We found that a naive clustering of wing shape data from taxonomically and geographically heterogeneous samples of workers returned groups congruent with species. We also confirmed that individuals can be reliably attributed to their genus, species and populations on the basis of their wing shape. Our results suggested that the shape variation reflects the evolutionary history with a potential influence of other factors such as body shape, climate and mimicry selective pressures. However, the high dimensionality of wing shape variation may have prevented absolute convergences between the different species. Wing venation shape is thus a taxonomically relevant marker combining the accuracy of quantitative characters with the specificity required for identification criteria. This marker may also highlight adaptive processes that could help understand the wing's influence on insect flight.  相似文献   

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