Design and evaluation of a deformable wing configuration for economical hovering flight of an insect-like tailless flying robot.

Studies on wing kinematics indicate that flapping insect wings operate at higher angles of attack (AoAs) than conventional rotary wings. Thus, effectively flying an insect-like flapping-wing micro air vehicle (FW-MAV) requires appropriate wing design for achieving low power consumption and high force generation. Even though theoretical studies can be performed to identify appropriate geometric AoAs for a wing for achieving efficient hovering flight, designing an actual wing by implementing these angles into a real flying robot is challenging. In this work, we investigated the wing morphology of an insect-like tailless FW-MAV, which was named KUBeetle, for obtaining high vertical force/power ratio or power loading. Several deformable wing configurations with various vein structures were designed, and their characteristics of vertical force generation and power requirement were theoretically and experimentally investigated. The results of the theoretical study based on the unsteady blade element theory (UBET) were validated with reference data to prove the accuracy of power estimation. A good agreement between estimated and measured results indicated that the proposed UBET model can be used to effectively estimate the power requirement and force generation of an FW-MAV. Among the investigated wing configurations operating at flapping frequencies of 23 Hz to 29 Hz, estimated results showed that the wing with a suitable vein placed outboard exhibited an increase of approximately 23.7% ± 0.5% in vertical force and approximately 10.2% ± 1.0% in force/power ratio. The estimation was supported by experimental results, which showed that the suggested wing enhanced vertical force by approximately 21.8% ± 3.6% and force/power ratio by 6.8% ± 1.6%. In addition, wing kinematics during flapping motion was analyzed to determine the reason for the observed improvement.

Assessment of the Hindlimb Membrane Musculature of Bats: Implications for Active Control of the Calcar.

The striking postcranial anatomy of bats reflects their specialized ecology; they are the only mammals capable of powered flight. Bat postcranial adaptations include a series of membranes that connect highly-modified, or even novel, skeletal elements. While most studies of bat postcranial anatomy have focused on their wings, bat hindlimbs also contain many derived and functionally important, yet less studied, features. In this study, we investigate variation in the membrane and limb musculature associated with the calcar, a neomorphic skeletal structure found in the hindlimbs of most bats. We use diffusible iodine-based contrast-enhanced computed tomography and standard histological techniques to examine the calcars and hindlimb membranes of three bat species that vary ecologically (Myotis californicus, a slow-flying insectivore; Molossus molossus, a fast-flying insectivore; and Artibeus jamaicensis, a slow-flying frugivore). We also assess the level of mineralization of the calcar at muscle attachment sites to better understand how muscle contraction may enable calcar function. We found that the arrangement of the calcar musculature varies among the three bat species, as does the pattern of mineral content within the calcar. M. molossus and M. californicus exhibit more complex calcar and calcar musculature morphologies than A. jamaicensis, and the degree of calcar mineralization decreases toward the tip of the calcar in all species. These results are consistent with the idea that the calcar may have a functional role in flight maneuverability. Anat Rec, 301:441-448, 2018. © 2018 Wiley Periodicals, Inc.

Selective attention in peacocks during assessment of rival males.

Males in many species compete intensely for access to females. In order to minimize costly interactions, they can assess their rivals' competitive abilities by evaluating traits and behaviors. We know little about how males selectively direct their attention to make these assessments. Using Indian peafowl (Pavo cristatus) as a model system, we examined how males visually assess their competitors by continuously tracking the gaze of freely moving peacocks during the mating season. When assessing rivals, peacocks selectively gazed toward the lower display regions of their rivals, including the lower eyespot and fishtail feathers, dense feathers, body and wings. Their attention was modified based on the rivals' behavior such that they spent more time looking at rivals when rivals were shaking their wings and moving. The results indicate that peacocks selectively allocate their attention during rival assessment. The gaze patterns of males assessing rivals were largely similar to those of females evaluating mates, suggesting that some male traits serve a dual function in both intra- and intersexual selection. However, males spent more time than females looking at the upper eyespots and this could indicate that the upper eyespots function more in close-up rival assessment than mate choice.

Assessment of the correlation between wing size and body weight in captive Culex quinquefasciatus.

INTRODUCTION: Mass production of mosquitoes under laboratory conditions allows implementing methods to control vector mosquitoes. Colony development depends on mosquito size and weight. Body size can be estimated from its correlation with wing size, whereas weight is more difficult to determine. Our goal was to test whether wing size can predict the weight. METHODS: We compared dry weight and wing centroid size of Culex quinquefasciatus reared at different temperatures and four diets. RESULTS: Weight and wing size were strongly correlated. The diets did not influence wing size. CONCLUSIONS: Wing centroid size is a good predictor of Cx. quinquefasciatus body weight.

Assessment of Aedes albopictus (Skuse) (Diptera: Culicidae) clutch size in wild and laboratory populations.

Aedes albopictus (Skuse) is an invasive mosquito species found across the southern U.S. with range expansion into many northern states. Intra- and interspecific larval competition have been evaluated for Ae. albopictus with respect to subsequent adult size, immature and adult survivability, and its capacity to vector pathogens as an adult. However, limited data are available on egg production as related to larval rearing conditions. Because Ae. albopictus is a container-inhabiting mosquito that oviposits in resource-limited habitats, it is found under variable density-dependent conditions. Therefore, we examined the impact of specific rearing conditions on Ae. albopictus clutch size and adult body size; comparing the egg production values and wing lengths from known developmental densities to those from field-collected populations. Field populations varied significantly among collection sites in mean clutch size (23 to 46). These clutch sizes were comparable to the mean clutch sizes of females reared at the larval densities of nine (20 eggs) and three (53 eggs) larvae per 3 ml of water in the laboratory. Field populations experienced density-dependent effects impacting adult mosquito size. Mosquitoes from the four sample sites had mean wing lengths of 1.99, 2.47, 2.51, and 2.54 mm, which were less than the mean wing length of mosquitoes reared at larval densities of three larvae per 3 ml of water (2.57 mm).

Evidence of selection on phenotypic plasticity and cost of plasticity in response to host-feeding sources in the major Chagas disease vector Triatoma infestans.

Phenotypic plasticity is the ability of a genotype to display alternative phenotypes in different environments. Understanding how plasticity evolves and the factors that favor and constrain its evolution have attracted great interest. We investigated whether selection on phenotypic plasticity and costs of plasticity affect head and wing morphology in response to host-feeding sources in the major Chagas disease vector Triatoma infestans. Full-sib families were assigned to blood-feeding on either live pigeons or guinea pigs throughout their lives. We measured diet-induced phenotypic plasticity on wing and head size and shape; characterized selection on phenotypic plasticity for female and male fecundity rates, and evaluated costs of plasticity. Wing size and shape variables exhibited significant differences in phenotypic plasticity associated with host-feeding source in female and male bugs. Evidence of selection on phenotypic plasticity was detected in head size and shape for guinea pig-fed females. A lower female fecundity rate was detected in more plastic families for traits that showed selection on plasticity. These results provide insights into the morphological phenotypic plasticity of T. infestans, documenting fitness advantages of head size and shape for females fed on guinea pigs. This vector species showed measurable benefits of responding plastically to environmental variation rather than adopting a fixed development plan. The presence of cost of plasticity suggests constraints on the evolution of plasticity. Our study indicates that females fed on guinea pigs (and perhaps on other suitable mammalian hosts) have greater chances of evolving under selection on phenotypic plasticity subject to some constraints.

Estimating sampling error of evolutionary statistics based on genetic covariance matrices using maximum likelihood.

We explore the estimation of uncertainty in evolutionary parameters using a recently devised approach for resampling entire additive genetic variance-covariance matrices (G). Large-sample theory shows that maximum-likelihood estimates (including restricted maximum likelihood, REML) asymptotically have a multivariate normal distribution, with covariance matrix derived from the inverse of the information matrix, and mean equal to the estimated G. This suggests that sampling estimates of G from this distribution can be used to assess the variability of estimates of G, and of functions of G. We refer to this as the REML-MVN method. This has been implemented in the mixed-model program WOMBAT. Estimates of sampling variances from REML-MVN were compared to those from the parametric bootstrap and from a Bayesian Markov chain Monte Carlo (MCMC) approach (implemented in the R package MCMCglmm). We apply each approach to evolvability statistics previously estimated for a large, 20-dimensional data set for Drosophila wings. REML-MVN and MCMC sampling variances are close to those estimated with the parametric bootstrap. Both slightly underestimate the error in the best-estimated aspects of the G matrix. REML analysis supports the previous conclusion that the G matrix for this population is full rank. REML-MVN is computationally very efficient, making it an attractive alternative to both data resampling and MCMC approaches to assessing confidence in parameters of evolutionary interest.

Taxonomic assessment of Culicoides brunnicans, C. santonicus and C. vexans (Diptera: Ceratopogonidae) in France: Implications in systematics.

Culicoides brunnicans Edwards, 1939, Culicoides santonicus, Callot et al., 1966, and Culicoides vexans (Staeger, 1839) belong to the Vexans group of the subgenus Oecacta. These species had never been studied by molecular methods and their distribution in Western Europe overlapped. C. brunnicans and C. santonicus are two closely related species and their diagnoses are based on the wing pattern only. An integrative taxonomic approach was conducted on females of the Vexans group, Culicoides furens (the type species of the subgenus Oecacta), and Culicoides nubeculosus (as outgroup) using sequences of D1D2 rDNA domains, sequences of COI mtDNA, and wing morphometrics. Species of the Vexans group were discriminated by all traits, and were closely related in the phylogeny. Their taxonomic status and their relation to C. furens were re-evaluated.

Variable assessment of wing colouration in aerial contests of the red-winged damselfly Mnesarete pudica (Zygoptera, Calopterygidae).

Wing pigmentation is a trait that predicts the outcome of male contests in some damselflies. Thus, it is reasonable to suppose that males would have the ability to assess wing pigmentation and adjust investment in a fight according to the costs that the rival may potentially impose. Males of the damselfly Mnesarete pudica exhibit red-coloured wings and complex courtship behaviour and engage in striking male-male fights. In this study, we investigated male assessment behaviour during aerial contests. Theory suggests that the relationship between male resource-holding potential (RHP) and contest duration describes the kind of assessment adopted by males: self-assessment, opponent-only assessment or mutual assessment. A recent theory also suggests that weak and strong males exhibit variations in the assessment strategies adopted. We estimated male RHP through male body size and wing colouration (i.e. pigmentation, wing reflectance spectra and transmission spectra) and studied the relationship between male RHP and contest duration from video-documented behavioural observations of naturally occurring individual contests in the field. The results showed that males with more opaque wings and larger red spots were more likely to win contests. The relationships between RHP and contest durations partly supported the self-assessment and the mutual assessment models. We then experimentally augmented the pigmented area of the wings, in order to evaluate whether strong and weak males assess rivals' RHP through wing pigmentation. Our experimental manipulation, however, clearly demonstrated that strong males assess rivals' wing pigmentation. We finally suggest that there is a variation in the assessment strategy adopted by males.

Wing kinematics in a hovering dronefly minimize power expenditure.

When an insect hovers or performs constant-speed flight, its wings flap at certain amplitude, frequency, angle of attack, etc., and the flight is balanced (vertical force equals to the weight, and horizontal force and pitch moment are zero). It is possible that when some other sets of values of wing kinematical parameters are used, the force and moment balance conditions can still be satisfied. Does the wing kinematics used by a constant-speed flying insect minimize the power expenditure? In this study, whether the wing kinematics used by a freely hovering dronefly minimizes its energy expenditure was investigated. First, the power consumption using the set of values of wing kinematical parameters that was actually employed by the insect was computed. Then, the kinematical parameters were changed while keeping the equilibrium flight conditions satisfied, and the power consumption was recalculated. It was found that wing kinematical parameters used by the freely hovering dronefly are very close to that minimize its energy consumption, and they can ensure the margin of controllability from hovering to maneuvers. That is, slight change of wing kinematical parameters did not cause significant change of the specific power (maintained a relatively small value).

Transgenic Anopheles gambiae expressing an antimalarial peptide suffer no significant fitness cost.

Mosquito-borne diseases present some of the greatest health challenges faced by the world today. In many cases, existing control measures are compromised by insecticide resistance, pathogen tolerance to drugs and the lack of effective vaccines. In light of these difficulties, new genetic tools for disease control programmes, based on the deployment of genetically modified mosquitoes, are seen as having great promise. Transgenic strains may be used to control disease transmission either by suppressing vector populations or by replacing susceptible with refractory genotypes. In practice, the fitness of the transgenic strain relative to natural mosquitoes will be a critical determinant of success. We previously described a transgenic strain of Anopheles gambiae expressing the Vida3 peptide into the female midgut following a blood-meal, which exhibited significant protection against malaria parasites. Here, we investigated the fitness of this strain relative to non-transgenic controls through comparisons of various life history traits. Experiments were designed, as far as possible, to equalize genetic backgrounds and heterogeneity such that fitness comparisons focussed on the presence and expression of the transgene cassette. We also employed reciprocal crosses to identify any fitness disturbance associated with inheritance of the transgene from either the male or female parent. We found no evidence that the presence or expression of the effector transgene or associated fluorescence markers caused any significant fitness cost in relation to larval mortality, pupal sex ratio, fecundity, hatch rate or longevity of blood-fed females. In fact, fecundity was increased in transgenic strains. We did, however, observe some fitness disturbances associated with the route of inheritance of the transgene. Maternal inheritance delayed male pupation whilst paternal inheritance increased adult longevity for both males and unfed females. Overall, in comparison to controls, there was no evidence of significant fitness costs associated with the presence or expression of transgenes in this strain.

Wing geometric morphometrics and molecular assessment of members in the Albitarsis Complex from Colombia.

Malaria parasites are transmitted to humans by female mosquitoes of the genus Anopheles. The Albitarsis Complex harbours at least eight species not readily differentiable by morphology. This complicates the determination of those species involved in malaria transmission and the implementation of targeted and effective vector control strategies. In Colombia, there is little information about the identity and distribution of the Albitarsis Complex members. In this work, COI DNA barcoding was used to assign specimens Anopheles albitarsis s.l. to any of the previously designated species of the Albitarsis Complex. Two molecular operational taxonomic units (MOTUs), differentially distributed in Colombia, were detected, A. albitarsis I in the NW and NE, and A. albitarsis F, E and NE Colombia. In contrast, nuclear white gene and ITS2 sequence analyses did not allow differentiating between the MOTUs. Wing landmark-based geometric morphometrics applied to explore intertaxa phenotypic heterogeneity showed a subtle but significant difference in size, while shape did not allow the separation of the MOTUs. In general, the multiple marker analysis was not supportive of the existence in Colombia of more than one species of the Albitarsis Complex.

Backward flight in hummingbirds employs unique kinematic adjustments and entails low metabolic cost.

Backward flight is a frequently used transient flight behavior among members of the species-rich hummingbird family (Trochilidae) when retreating from flowers, and is known from a variety of other avian and hexapod taxa, but the biomechanics of this intriguing locomotor mode have not been described. We measured rates of oxygen uptake (V(O2)) and flight kinematics of Anna's hummingbirds, Calypte anna (Lesson), within a wind tunnel using mask respirometry and high-speed videography, respectively, during backward, forward and hovering flight. We unexpectedly found that in sustained backward flight is similar to that in forward flight at equivalent airspeed, and is about 20% lower than hovering V(O2). For a bird that was measured throughout a range of backward airspeeds up to a speed of 4.5 m s(-1), the power curve resembled that of forward flight at equivalent airspeeds. Backward flight was facilitated by steep body angles coupled with substantial head flexion, and was also characterized by a higher wingbeat frequency, a flat stroke plane angle relative to horizontal, a high stroke plane angle relative to the longitudinal body axis, a high ratio of maximum:minimum wing positional angle, and a high upstroke:downstroke duration ratio. Because of the convergent evolution of hummingbird and some hexapod flight styles, flying insects may employ similar kinematics while engaged in backward flight, for example during station keeping or load lifting. We propose that backward flight behavior in retreat from flowers, together with other anatomical, physiological, morphological and behavioral adaptations, enables hummingbirds to maintain strictly aerial nectarivory.

The proximal costs of case construction in caddisflies: antioxidant and life history responses.

Animal construction allows organisms to cope with environmental variations but the physiological costs of such behaviour are still poorly understood. The aim of the present study was to measure the physiological cost of construction behaviour through the oxidative balance that is known to affect the ability of organs to function, stimulates senescence processes and ultimately impacts the fitness of the organism. We used larvae of caddisfly, Limnephilus rhombicus, by experimentally modifying the effort associated with case building. Larvae that were forced to build a new case showed a significant increase in both total antioxidant capacity and the specific activity of superoxide dismutase 48 and 72 h, respectively, after the initiation of the reconstruction. These results strongly suggest that the larval construction behaviour triggered the production of reactive oxygen species, but their effects were reversed 7 days after the reconstruction. In the animals that were forced to build a new case, oxidative stress appeared to be mitigated by a network of antioxidant defences because no oxidative damage was observed in proteins compared with the control larvae. At the adult stage, while longevity was not sex dependent and was not affected by the treatment, body mass and body size of adult males from the reconstruction treatment were significantly lower than the control values. This unexpected sex effect together with data on oxidative stress highlights the difficulty of determining the physiological cost associated with energy-demanding behaviours, implying a consideration of both their energetic and non-energetic components is required.

Size and asymmetry: are there costs to winning the royalty race?

Body size and morphology are key fitness-determining traits that can vary genotypically. They are likely to be important in social insect queens, which mate in swarms and found colonies independently, but genetic influences on queen morphology have been little investigated. Here, we show that the body size and morphology of queens are influenced by their genotype in the leaf-cutting ant Acromyrmex echinatior, a species in which certain lineages (patrilines) bias their development towards reproductive queens rather than sterile workers. We found no relationship between the queen-worker skew of patrilines and the size or morphology of queens, but there was a significant relationship with fluctuating asymmetry, which was greater in more queen-biased patrilines. Our results suggest that queen-biased patrilines do not incur a fitness cost in terms of body size, but may face more subtle costs in developmental stability. Such costs may constrain the evolution of royal cheating in social insects.

High manoeuvring costs force narrow-winged molossid bats to forage in open space.

Molossid bats are specialised aerial-hawkers that, like their diurnal ecological counterparts, swallows and swifts, hunt for insects in open spaces. The long and narrow wings of molossids are considered energetically adapted to fast flight between resource patches, but less suited for manoeuvring in more confined spaces, such as between tree-tops or in forest gaps. To understand whether a potential increase in metabolic costs of manoeuvring excludes molossids from foraging in more confined spaces, we measured energy costs and speed of manoeuvring flight in two tropical molossids, 18 g Molossus currentium and 23 g Molossus sinaloae, when flying in a ~500 m(3) hexagonal enclosure (~120 m(2) area), which is of similar dimensions as typical forest gaps. Flight metabolism averaged 10.21 ± 3.00 and 11.32 ± 3.54 ml CO(2) min(-1), and flight speeds 5.65 ± 0.47 and 6.27 ± 0.68 m s(-1) for M. currentium and M. sinaloae respectively. Metabolic rate during flight was higher for the M. currentium than for the similar-sized, but broader-winged frugivore Carollia sowelli, corroborating that broad-winged bats are better adapted to flying in confined spaces. These higher metabolic costs of manoeuvring flight may be caused by having to fly slower than the optimal foraging speed, and by the additional metabolic costs for centripetal acceleration in curves. This may preclude molossids from foraging efficiently between canopy trees or in forest gaps. The surprisingly brief burst of foraging activity at dusk of many molossids might be related to the cooling of the air column after sunset, which drives airborne insects to lower strata. Accordingly, foraging activity of molossids may quickly turn unprofitable when the abundance of insects decreases above the canopy.

Perch-hunting in insectivorous Rhinolophus bats is related to the high energy costs of manoeuvring in flight.

Foraging behaviour of bats is supposedly largely influenced by the high costs of flapping flight. Yet our understanding of flight energetics focuses mostly on continuous horizontal forward flight at intermediate speeds. Many bats, however, perform manoeuvring flights at suboptimal speeds when foraging. For example, members of the genus Rhinolophus hunt insects during short sallying flights from a perch. Such flights include many descents and ascents below minimum power speed and are therefore considered energetically more expensive than flying at intermediate speed. To test this idea, we quantified the energy costs of short manoeuvring flights (<2 min) using the Na-bicarbonate technique in two Rhinolophus species that differ in body mass but have similar wing shapes. First, we hypothesized that, similar to birds, energy costs of short flights should be higher than predicted by an equation derived for bats at intermediate speeds. Second, we predicted that R. mehelyi encounters higher flight costs than R. euryale, because of its higher wing loading. Although wing loading of R. mehelyi was only 20% larger than that of R. euryale, its flight costs (2.61 ± 0.75 W; mean ± 1 SD) exceeded that of R. euryale (1.71 ± 0.37 W) by 50%. Measured flight costs were higher than predicted for R. mehelyi, but not for R. euryale. We conclude that R. mehelyi face elevated energy costs during short manoeuvring flights due to high wing loading and thus may optimize foraging efficiency by energy-conserving perch-hunting.

Barriers to movement: Modelling energetic costs of avoiding marine wind farms amongst breeding seabirds.

Proposals for wind farms in areas of known importance for breeding seabirds highlight the need to understand the impacts of these structures. Using an energetic modelling approach, we examine the effects of wind farms as barriers to movement on seabirds of differing morphology. Additional costs, expressed in relation to typical daily energetic expenditures, were highest per unit flight for seabirds with high wing loadings, such as cormorants. Taking species-specific differences into account, costs were relatively higher in terns, due to the high daily frequency of foraging flights. For all species, costs of extra flight to avoid a wind farm appear much less than those imposed by low food abundance or adverse weather, although such costs will be additive to these. We conclude that adopting a species-specific approach is essential when assessing the impacts of wind farms on breeding seabird populations, to fully anticipate the effects of avoidance flights.

Cost of co-infection controlled by infectious dose combinations and food availability.

To what extent the combined effect of several parasite species co-infecting the same host (i.e. polyparasitism) affects the host's fitness is a crucial question of ecological parasitology. We investigated whether the ecological setting can influence the co-infection's outcome with the mosquito Aedes aegypti and two parasites: the microsporidium Vavraia culicis and the gregarine Ascogregarina culicis. The cost of being infected by the two parasites depended on the interaction between the two infectious doses and host food availability. The age at pupation of the mosquito was delayed most when the doses of the two parasites were highest and little food was available. As infectious dose increases with the parasites' prevalence and intensity of transmission, the cost of being co-infected depends on the epidemiological status of the two parasite species.

Reproductive skew and selection on female ornamentation in social species.

Male animals are typically more elaborately ornamented than females. Classic sexual selection theory notes that because sperm are cheaper to produce than eggs, and because males generally compete more intensely for reproductive opportunities and invest less in parental care than females, males can obtain greater fitness benefits from mating multiply. Therefore, sexual selection typically results in male-biased sex differences in secondary sexual characters. This generality has recently been questioned, because in cooperatively breeding vertebrates, the strength of selection on traits used in intrasexual competition for access to mates (sexual selection) or other resources linked to reproduction (social selection) is similar in males and females. Because selection is acting with comparable intensity in both sexes in cooperatively breeding species, the degree of sexual dimorphism in traits used in intrasexual competition should be reduced in cooperative breeders. Here we use the socially diverse African starlings (Sturnidae) to demonstrate that the degree of sexual dimorphism in plumage and body size is reduced in cooperatively breeding species as a result of increased selection on females for traits that increase access to reproductive opportunities, other resources, or higher social status. In cooperative breeders such as these, where there is unequal sharing of reproduction (reproductive skew) among females, and where female dominance rank influences access to mates and other resources, intrasexual competition among females may be intense and ultimately select for female trait elaboration. Selection is thereby acting with different intensities on males and females in cooperatively versus non-cooperatively breeding species, and female-female interactions in group-living vertebrates will have important consequences for the evolution of female morphological, physiological and behavioural traits.