Carnivorous bats?

Only large bats can take large prey but size alone does not identify 'carnivorous bats' (those including small terrestrial vertebrates in their diets). Morphological data, including body mass, aspect ratio and relative wing loading, along with information about orientation and foraging strategies can be used to characterize a suite of features which identifies carnivorous bats. We use the available data to make predictions about which large Microchiroptera will be found to be carnivorous. A combination of morphological features including body mass (^0.017 kg), low aspect ratio (<6.3), and low relative wing loading (<36) significantly identifies carnivorous species from among other animal-eating forms. Some carnivorous species use short, low intensity, high frequency, broadband echolocation cells but rely on prey generated cues to locate their targets. Other carnivorous species are facultative echolocators. The available data lead to the prediction that Phyllostomus hastatus and Hipposideros diadema are not regularly carnivorous, while Otonycteris hemprichi may be. Large species with echolocation calls adapted for flutter detection (rhinolophids and hipposiderids) or those with long narrowband calls and high aspect ratio wings with high relative wing loading (for example molossids, some emballonurids and some vespertilionids) chase airborne prey in the open; neither of these approaches involves prey other than arthropods.     

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.     

Effects of body-size variation on flight-related traits in latitudinal populations of Drosophila melanogaster

In the present study, we tested the hypothesis whether flight-related traits such as wing area, flight-muscle ratio, wing loading and dispersal yield evidence of geographical variation in nine wild-collected as well as laboratory-reared (at 21°C) latitudinal populations of Drosophila melanogaster from the Indian subcontinent. We observed positive clinal variation in the wing–thorax ratio, wing aspect ratio and wing area, along a latitudinal gradient for both the sexes. In contrast, geographical changes in three parameters of flight ability, i.e. flight-muscle ratio, wing loading and dispersal, showed negative correlation withlatitude. On the basis of isofemale line variability, we observed positive correlation of wing loading with flight-muscle ratio as well as dispersal behaviour in both the sexes. We also found positive correlation between duration of development and wing area. Interestingly, southern populations of D. melanogaster from warm and humid habitats exhibited higher flight-muscle ratio as well as the higher wing loading than northern populations which occur in cooler and drier climatic conditions. Laboratory tests for dispersal-related walking behaviour showed significantly higher values for southern populations compared with northern populations of D. melanogaster. Multiple regression analysis of geographical changes in flight-muscle ratio, wing loading as well as walking behaviour as a function of average temperature and relative humidity of the origin of populations in wild-collected flies have suggested adaptive changes in flight-related traits in response to steeper gradients of climatic factors in the Indian subcontinent. Finally, adaptive latitudinal variations in flight-related traits in D. melanogaster are consistent with results of other studies from different continents despite differences due to specific climatic conditions in the Indian subontinent.     

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.     

Ecomorphology and foraging behaviour of Pacific boobies

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.     

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.     

The wing morphology of limnephilid caddisflies in relation to their habitat preferences

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Reproductive variability in Pinus sylvestris in southern France: Implications for invasion

Abstract. Though fecundity and dispersal have been recognized as major factors in most invasion models, their ecological determinants are still poorly known. This paper aims to identify the main sources of variation in seed production and dispersal distance in a naturally expanding Pinus sylvestris population. We propose some tree measurements that may be related to their contribution to population expansion. We quantified cone and seed production and measured three seed characteristics related to their dispersal ability: mass, wing area and wing‐loading (mass:area ratio) in cones sampled at different relative heights and aspects in the canopy and in trees of different age and competitive status. Results showed that isolated trees had a much more abundant cone production, which was more evenly distributed in the canopy than trees within stands. Age was also positively related to cone production. Seed dimensions varied between and within trees but we found no effect of isolation or age. The strong positive correlation between wing area and seed mass leads to a limited variability of seed wing loading. Seed characteristics may thus play a minor role in individual dispersal ability and relative tree position in the stand that is strongly linked to tree fecundity should be more appropriate for estimating the individual contribution to the whole population expansion. Our results also highlight the importance of obtaining demographic data in low‐density populations to estimate the invasive potential of a species.     

Growth temperature and phenotypic plasticity in two Drosophila sibling species: probable adaptive changes in flight capacities

In the sibling species Drosophila melanogaster and D. simulans, growth and development at constant temperatures, from 12 to 30 °C, resulted in extensive variations of adult size and flight parameters with significant differences between species. Changes in body weight, thorax length and wing length were nonlinear, with maximum values of each trait at lower temperatures for D. simulans than for its sibling species. By contrast, the wing/thorax ratio and the wing loading varied monotonically with growth temperature. These traits were negatively correlated, the wing/thorax ratio decreasing with growth temperature while the wing loading increased. Wing/thorax ratio, which is easier to measure, thus appears as a convenient predictor of wing loading. During tethered flight at the same ambient temperature, the wingbeat frequency changed linearly as a function of the wing moment of inertia. More interestingly, the beat rate was strongly correlated with the increase of wing loading at growth temperature above 13 °C. The likely adaptive significance of these morphometrical changes for flight efficiency is discussed.     

Landscape structure and phenotypic plasticity in flight morphology in the butterfly Pararge aegeria

In evolutionary time, varying environments may lead to different morphs as a result of genetic adaptation and divergence or phenotypic plasticity. Landscapes that differ in the extent of habitat fragmentation may provide different selection regimes for dispersal, but also for other ecological functions. Several studies on flying insects have shown differences in flight morphology between landscapes, but whether such differences result from plastic responses have rarely been tested. We did a reciprocal transplant experiment with offspring of speckled wood butterfly females (Parargeaegeria) from three types of landscape differing in fragmentation: woodland landscape, landscape with woodland fragments and agricultural landscape with only hedgerows. Young caterpillars were allowed to grow individually on potted host grasses in small enclosures under the three landscape conditions (split‐brood design). Mortality in caterpillars was much higher in agricultural landscape compared to the other landscapes. Additive to the effect of landscape of development, landscape of origin also affected mortality rate in a similar way. Flight morphology of the adults resulting from the experiment differed significantly with landscape. Independent of the landscape of origin, males and females that developed in agricultural landscape were the heaviest and had the greatest wing loadings. Females that developed in agricultural landscape had higher relative thorax mass (i.e. greater flight muscle allocation) in line with adaptive predictions on altered dispersal behaviour with type of landscape. In males, relative thorax mass did not respond significantly relative to landscape of development, but males originating from landscape with woodland fragments allocated more into their thorax compared to males from the other types. We found significant G×E interactions for total dry mass and wing loading. Our results suggest the existence of phenotypic plasticity in butterfly flight morphology associated with landscape structure.     

Evolution of flight morphology in a butterfly that has recently expanded its geographic range

Individuals colonizing unoccupied habitats typically possess characters associated with increased dispersal and, in insects, colonization success has been related to flight morphology. The speckled wood butterfly, Pararge aegeria, has undergone recent major expansions in its distribution: in the north of its range, P. aegeria has colonized many areas in north and east England, and in the south, it was first recorded on Madeira in 1976. We examined morphological traits associated with flight and reproduction in the northern subspecies tircis, and in the southern subspecies aegeria, from sites colonized about 20 years ago in northern England and on Madeira, respectively. Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were from individuals reared in a common environment and there were significant family effects in most of the variables measured. Compared with individuals from sites continuously occupied in recent history, colonizing individuals were larger (adult live mass). In the subspecies tircis, colonizing individuals also had relatively larger thoraxes and lower wing aspect ratios indicating that evolutionary changes in flight morphology may be related to colonization. However, sex by site interactions in analyses of thorax mass and abdomen mass suggest different selection pressures on flight morphology between the sexes in relation to colonization. Overall, the subspecies aegeria was smaller (adult live mass) and had a relatively larger thorax and wings, and smaller abdomen than subspecies tircis. Evolutionary changes in flight morphology and dispersal rate may be important determinants of range expansion, and may affect responses to future climate change. Received: 1 March 1999 / Accepted: 30 June 1999     

The effect of habitat amount on flight-related traits in the butterfly Hamadryas februa is sex-dependent

1. Mobility in flying animals can be assessed by variations in morpho–ecological traits such as body, thorax and wing sizes, wing shape and the proportion between body mass and wing area. Habitat loss and fragmentation can promote phenotypic plasticity and microevolutionary divergencies in natural populations. In this context, sexual differences in physiology and behaviour can impose different selection pressure on morphological aspects related to flight.
2. We evaluated the relative impact of forest patch area and habitat amount in shaping flight-related morpho–ecological traits of the tropical butterfly Hamadryas februa. We find a marked sexual dimorphism in the species, with females being larger, having larger thorax, higher wing loadings and larger wing total area than males. These trait values indicate females as the more dispersive sex. We show that habitat amount modulates body mass allocations in both sexes, leading to an increase in thorax mass with decreasing habitat amount. The effect of habitat amount was more pronounced in females, which increased total mass and wing loading while decreasing thorax allocation with decreasing habitat amount. This outcome suggests that females increase abdominal mass in response to a reduction in habitat amount. The focal forest patch increasing area was linked to increases in hindwing lengths in both females and males.
3. We advocate that both landscape metrics (i.e., habitat amount and patch area) should be considered in studies evaluating landscapes' impacts on insect mobility. We discuss results in terms of the species' sexual differences in flight behaviour and the relative importance of both landscape metrics.

     

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.     

Flight mode affects allometry of migration range in birds

Billions of birds migrate to exploit seasonally available resources. The ranges of migration vary greatly among species, but the underlying mechanisms are poorly understood. I hypothesise that flight mode (flapping or soaring) and body mass affect migration range through their influence on flight energetics. Here, I compiled the tracks of migratory birds (196 species, weighing 12–10 350 g) recorded by electronic tags in the last few decades. In flapping birds, migration ranges decreased with body mass, as predicted from rapidly increasing flight cost with increasing body mass. The species with higher aspect ratio and lower wing loading had larger migration ranges. In soaring birds, migration ranges were mass‐independent and larger than those of flapping birds, reflecting their low flight costs irrespective of body mass. This study demonstrates that many animal‐tracking studies are now available to explore the general patterns and the underlying mechanisms of animal migration.     

Morphometrical evolution in a Drosophila clade: the Drosophila obscura group

Five morphometrical traits (wing and thorax length, ovariole number, and thoracic and female abdomen pigmentation) were investigated in laboratory stocks of 20 species belonging to the Drosophila obscura group (subgenus Sophophora). These species originated from four biogeographical regions and represent all five of the presently recognized, taxonomic subgroups. Size‐related traits (wing and thorax length) were highly variable across species, and interspecific variation explained more than 90% of total variability. In both traditional and phylogenetic analyses, wing size was positively correlated with latitude of origin. These interspecific correlations were however notably weaker than those for intraspecific correlations. Wing/thorax ratio, which may be related to flight capacity, showed little variation. Ovariole number was highly variable (range 27–53) both within and between species, and was positively correlated with the wing/thorax ratio, suggesting that species with relatively large ovaries have relatively low wing loading. Although many species are completely dark, 11 had some regions of light coloration. A light thorax with a median darkening was observed in six species. A variable pigmentation of abdominal tergites, in females only, was found in nine species, belonging to three subgroups only. With respect to both molecular phylogeny and morphometrical evolution, the D. obscura subgroup is probably now the best investigated clade in Drosophila.     

Drivers affecting habitat use in Afrotropical hipposiderid and pteropodid bats

Assessing how bats respond to habitat attributes requires an integrative approach to reliably predict direct community-level effects. We focused on hipposiderid and pteropodid bats because of their diverse resource use patterns, body size ranges, and dispersal abilities. We combined an array of bat species-level characteristics with key forest stand characteristics that may covary with habitat use. Twelve stations were sampled in the Lomami and Yangambi landscapes, Democratic Republic of the Congo. We investigated whether species-level flight ability of bats and forest stand characteristics can affect bat commuting flights and community-level estimates of both species detection and habitat occupancy. We captured bats for 108 trap-nights. Three sampling events (early evening, middle of the night, and early morning) were replicated for each survey night. Hipposiderids showed an early evening flight peak, while flight activity of pteropodids was constant throughout the night, but increased around the middle of the night. Species capture probability decreased with higher wing loading in hipposiderids and was negatively correlated with higher wing aspect ratio in pteropodids. Forest occupancy of hipposiderids increased along the gradient towards waterways, while pteropodid occurrence was not directly linked to measured forest stand variables. This suggests a consequence of habitat patterns at larger spatial scales, which would need clarifying through additional data collection. We discuss these findings in terms of resource-use strategies of clutter-tolerant and clutter-intolerant species. We argue that the occurrence of specific bat species and their habitat use patterns can serve as surrogate measures of ecosystem health.     

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.     

Morphometrical and front wing abrasion analysis of a Hungarian cotton bollworm <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> (Lepidoptera: Noctuidae) population

To determine the cotton bollworm migrating population rate in Hungary, we examined the weights and the front wing morphological feautures of trapped moths. We used sex pheromone traps to monitor field populations during the maize vegetation cycle period in 2008. We examined moths trapped at various times, and measured their body mass (m) and morphological features, namely the front wing quotient (fWQ = quotient of length of front wing/width of front wing), modified wing loading (WL = weight of moth/surface of front wing), and the relative thorax size (RTS = width of thorax/width of head). The data were analysed by Student t-test, anterior wing abrasion and darkness were analysed by a Adobe Photoshop 7.0 software. The Hungarian appearance of three cottom bollworm generations in 2008 was also observed. Based on the examined morphological features we found regularity in body mass, front wing quotient and modified wing loading changes during the flight period. The specimens trapped in the first and third part of the flight period had lower body mass, larger wing surface, longer wings and more favourable modified wing loading than the specimens trapped in the middle of the flight period. The abrasion and colour of the anterior wings of cotton bollworms were concordant to morphometric investigations. The abrasion in darker spots E1 and E3 clearly showed a more intensive usage of the wings in case of specimens trapped at the beginning and at the end of the flight period.     

Flight machinery dimensions of honeybees, Apis mellifera

The flight muscles of different honeybee subspecies are known to have different allozymes of malate dehydrogenase (Mdh) which in turn are correlated with differences in mass-specific metabolic flight rate. Flight capacity is also affected by dimensional, morphological relationships of mass and area which allow an estimation of an “excess power index”. The dimensions of the flight machinery of honeybees (based on our own data) were coupled with the frequency distributions of Mdh (taken from the literature) to compare nine subspecies of African and nine European honeybees, Apis mellifera as miniature aircraft. The two groups differed significantly for five dimensions of flight machinery and in the distribution frequencies of Mdh phenotypes. In the African group, northern and southern subgroups occurred which significantly differed in body mass and excess power index, while flight engine and body mass varied proportionately. In the European group, wing surface was nearly constant but body mass and the thorax/body mass ratio varied significantly resulting in significantly differing wing loading values. The final excess power index (modified for allozyme phenotype) of the European bees reflected both flight machinery and allozymic differences. Mdh allozymic phenotype frequencies were correlated with the dimensional morphological components of the excess power index. As a group, the European subspecies of honeybees were 33% heavier and had 15% more wing surface area than the African group. However, the former have a thorax/body mass ratio of 0.45 and wing loading value of 0.48 against the latter's 0.53 and 0.35 respectively. This confers an advantage on the African group solely on the grounds of dimensions because there was proportionately less mass per unit area of wing surface and so lower lift requirement. The better engine to aircraft mass ratio provides greater power per unit mass in the African group taken as miniature aircraft. Differences in metabolic capacity associated with Mdh allozymes (taken from the literature) finally result in an excess power index that is 38% greater in the African than European subspecies of honeybees. Accepted: 22 December 1998     

Sexual differences rather than flight performance underlie movement and exploration in a tropical butterfly

1. Individual movement behaviour governs several routine processes, and may scale up to important ecological processes, including dispersal. However, movement is affected by a wealth of factors, including abiotic conditions, flight performance, and behavioural traits. Although it has been historically assumed that insect flight is in the first place ruled by physiology and morphology, researchers have only recently begun to understand the potentially important role of behavioural traits. 2. This study aims to disentangle the relative importance of thermal conditions during development, and especially flight performance (capacity), versus behaviour (intrinsic motivation) in relation to movement attributes (i.e. time until take‐off, number of positions visited) under controlled laboratory conditions in the tropical butterfly Bicyclus anynana. 3. As predicted, links were found between flight performance (forced flight) and morphological traits (body size). However, this link was less pronounced for movement and exploratory behaviour, suggesting a more pronounced role of intrinsic motivation on the actual decision to move, or not. Thus, flight performance and movement may not be intimately associated. 4. Flight behaviour was mainly determined by sexual differences, with males showing better flight performance, higher mobility, and enhanced exploration than females. 5. Lower developmental temperatures increased thorax–abdomen ratio, thorax mass, and exploratory behaviour, and decreased wing loading. This may potentially aid flight capacity under thermally challenging conditions. 6. This study adds to the growing evidence that behavioural traits should not be neglected when investigating movement and dispersal, as they may well play a crucially important role.