Flight Metabolic Rate Research Articles

Experimental elevated temperature affects bumblebee foraging and flight speed.

Global warming threatens wild bees and their interaction with plants. While earlier studies have highlighted the negative effects of elevated temperatures on bee-plant interactions, we still lack knowledge about how they impact the foraging behaviours that are central to bee pollination activities. To address this knowledge gap, we investigated how ambient temperature affected the foraging behaviours of the bumblebee Bombus terrestris. We allowed the bumblebees to forage freely on artificial flowers in two climate-controlled rooms set at 24°C and 32°C. The colonies were alternated between the two temperatures every week. We recorded the flower visitation rate, flight speed, total foraging time and number of foraging trips. In addition, we measured flight metabolic rate across a range of temperatures to assess its potential as an underlying mechanism. In comparison to 24°C, at 32°C, flower visitation time decreased while flower visitation rate and flight speed increased. This is consistent with the reduction in flight metabolic rate recorded between these temperatures. At 32°C, the number of trips made by each worker decreased, suggesting that, despite the reduced energetic cost, flight in elevated temperatures may be stressful. Our results suggest that elevated temperatures affect bumblebee foraging behaviour and that this would likely disrupt plant-insect interactions.

Resting Metabolic Rate Does Not Predict Peak Metabolic Rate in the Glanville Fritillary Butterfly.

Peak metabolic rate reflects maximal performance and may have direct fitness consequences, whereas resting metabolic rate (RMR) represents the maintenance cost of the whole animal. These traits may be linked, which has significant implications for the evolution of both traits. In vertebrates, a positive correlation between RMR and aerobic capacity has been proposed to explain the origin of endothermy. However, as studies on the relationship between RMR and aerobic capacity have focused on vertebrates, we know much less about these traits in ectothermic insects. I measured RMR in the Glanville fritillary butterfly (Melitaea cinxia) using two configurations: one optimized for measuring flight metabolic rate and the other optimized for RMR. The relationship between RMR and body mass was similar for the two configurations. Body mass explained 82% of the variation in RMR when it was measured using the "flight" configuration at 32°C, and 91% when using the "rest" configuration at 23°C. The Q10 coefficient calculated based on the two RMR measurements was 2.8. Mass-independent RMR was positively correlated between measurements obtained using the two instrument configurations. However, neither measure of RMR was correlated with peak metabolic rate, which indicates that RMR cannot be used as a surrogate measure for aerobic capacity in the Glanville fritillary. Ectothermic insects may be able to combine high metabolic capacity with no apparent increase in maintenance cost. Even though RMR is among the most frequently measured physiological variables, it may have limited predictive power when it comes to questions related to activity or aerobic capacity, or in the case of butterflies, flight performance.

Effects of tag mass on the physiology and behaviour of common noctule bats

BackgroundExternal tags, such as transmitters and loggers, are often used to study bat movements. However, physiological and behavioural effects on bats carrying tags have rarely been investigated, and recommendations on the maximum acceptable tag mass are rather based on rules of thumb than on rigorous scientific assessment.MethodsWe conducted a comprehensive three-step assessment of the potential physiological and behavioural effects of tagging bats, using common noctules Nyctalus noctula as a model. First, we examined seasonal changes in body mass. Second, we predicted and then measured potential changes in flight metabolic rate in a wind tunnel. Third, we conducted a meta-analysis of published data to assess effects of different tag masses on the weight and behaviour of bats.ResultsIndividual body mass of common noctules varied seasonally by 7.0 ± 2.6 g (range: 0.5–11.5 g). Aerodynamic theory predicted a 26% increase in flight metabolic rate for a common noctule equipped with a 3.8 g tag, equating to 14% of body mass. In a wind tunnel experiment, we could not confirm the predicted increase for tagged bats. Our meta-analysis revealed a weak correlation between tag mass and emergence time and flight duration in wild bats. Interestingly, relative tag mass (3–19% of bat body mass) was not related to body mass loss, but bats lost more body mass the longer tags were attached. Notably, relatively heavy bats lost more mass than conspecifics with a more average body mass index.ConclusionBecause heavy tags (> 3 g) were generally used for shorter periods of time than lighter tags (~ 1 g), the long-term effects of heavy tags on bats cannot be assessed at this time. Furthermore, the effects of disturbance and resource distribution in the landscape cannot be separated from those of tagging. We recommend that tags weighing 5–10% of a bat’s mass should only be applied for a few days. For longer studies, tags weighing less than 5% of a bat's body mass should be used. To avoid adverse effects on bats, researchers should target individuals with average, rather than peak, body mass indices.

A thermal performance curve perspective explains decades of disagreements over how air temperature affects the flight metabolism of honey bees.

While multiple studies have shown that honey bees and some other flying insects lower their flight metabolic rates when flying at high air temperatures, critics have suggested such patterns result from poor experimental methods as, theoretically, air temperature should not appreciably affect aerodynamic force requirements. Here, we show that apparently contradictory studies can be reconciled by considering the thermal performance curve of flight muscle. We show that prior studies that found no effects of air temperature on flight metabolism of honey bees achieved flight muscle temperatures that were near or on equal, opposite sides of the thermal performance curve. Honey bees vary their wing kinematics and metabolic heat production to thermoregulate, and how air temperature affects the flight metabolic rate of honey bees is predictable using a non-linear thermal performance perspective of honey bee flight muscle.

Enhanced dispersal capacity in edge population individuals of a rapidly expanding butterfly.

Natural range shifts offer the opportunity to study the phenotypic and genetic changes contributing to colonization success. The recent range shift of the Southern small white butterfly (Pieris mannii) from the South to the North of Europe offers a prime example to examine a potential dispersal syndrome in range-expanding individuals. We compared butterflies from the core and edge populations using a multimodal approach addressing behavioral, physiological, and morphological traits related to dispersal capacity. Relative to individuals from the core range (France), individuals from the edge (Germany) showed a higher capacity and motivation to fly, and a higher flight metabolic rate. They were also smaller, which may enhance their flight maneuverability and help them cope with limited resource availability, thereby increasing their settlement success in novel environments. Altogether, the behavioral, physiological, and morphological differences observed between core and edge populations in P. mannii suggest the existence of a dispersal syndrome in range-expanding individuals. Whether these differences result from genetic and/or phenotypic responses remains, however, to be determined.

Flying, nectar-loaded honey bees conserve water and improve heat tolerance by reducing wingbeat frequency and metabolic heat production

Heat waves are becoming increasingly common due to climate change, making it crucial to identify and understand the capacities for insect pollinators, such as honey bees, to avoid overheating. We examined the effects of hot, dry air temperatures on the physiological and behavioral mechanisms that honey bees use to fly when carrying nectar loads, to assess how foraging is limited by overheating or desiccation. We found that flight muscle temperatures increased linearly with load mass at air temperatures of 20 or 30 °C, but, remarkably, there was no change with increasing nectar loads at an air temperature of 40 °C. Flying, nectar-loaded bees were able to avoid overheating at 40 °C by reducing their flight metabolic rates and increasing evaporative cooling. At high body temperatures, bees apparently increase flight efficiency by lowering their wingbeat frequency and increasing stroke amplitude to compensate, reducing the need for evaporative cooling. However, even with reductions in metabolic heat production, desiccation likely limits foraging at temperatures well below bees' critical thermal maxima in hot, dry conditions.

Divergent impacts of the neonicotinoid insecticide, clothianidin, on flight performance metrics in two species of migratory butterflies.

Long-distance flight is crucial for the survival of migratory insects, and disruptions to their flight capacity can have significant consequences for conservation. In this study, we examined how a widely used insecticide, clothianidin (class: neonicotinoid), impacted the flight performance of two species of migratory butterflies, monarchs (Danaus plexippus) and painted ladies (Vanessa cardui). To do this, we quantified the free-flight energetics and tethered-flight velocity and distance of the two species using flow-through respirometry and flight mill assays. Our findings show differential effects of the pesticide on the two species. For painted ladies, we found that clothianidin exposure reduced average free-flight metabolic rates, but did not affect either average velocity or total distance during tethered flight. Other studies have linked low flight metabolic rates with reduced dispersal capacity, indicating that clothianidin exposure may hinder painted lady flight performance in the wild. Conversely, for monarchs, we saw no significant effect of clothianidin exposure on average free-flight metabolic rates but did observe increases in the average velocity, and for large individuals, total distance achieved by clothianidin-exposed monarchs in tethered flight. This suggests a potential stimulatory response of monarchs to low-dose exposures to clothianidin. These findings indicate that clothianidin exposure has the potential to influence the flight performance of butterflies, but that not all species are impacted in the same way. This highlights the need to be thoughtful when selecting performance assays, as different assays can evaluate fundamentally distinct aspects of physiology, and as such may yield divergent results.

Intraspecific scaling and early life history determine the cost of free-flight in a large beetle (Batocera rufomaculata).

The scaling of the energetic cost of locomotion with body mass is well documented at the interspecific level. However, methodological restrictions limit our understanding of the scaling of flight metabolic rate (MR) in free-flying insects. This is particularly true at the intraspecific level, where variation in body mass and flight energetics may have direct consequences for the fitness of an individual. We applied a 13C stable isotope method to investigate the scaling of MR with body mass during free-flight in the beetle Batocera rufomaculata. This species exhibits large intraspecific variation in adult body mass as a consequence of the environmental conditions during larval growth. We show that the flight-MR scales with body mass to the power of 0.57, with smaller conspecifics possessing up to 2.3 fold higher mass-specific flight MR than larger ones. Whereas the scaling exponent of free-flight MR was found to be like that determined for tethered-flight, the energy expenditure during free-flight was more than 2.7 fold higher than for tethered-flight. The metabolic cost of flight should therefore be studied under free-flight conditions, a requirement now enabled by the 13C technique described herein for insect flight.

Seasonal plasticity in morphology and metabolism differs between migratory North American and resident Costa Rican monarch butterflies.

Environmental heterogeneity in temperate latitudes is expected to maintain seasonally plastic life-history strategies that include the tuning of morphologies and metabolism that support overwintering. For species that have expanded their ranges into tropical latitudes, it is unclear the extent to which the capacity for plasticity will be maintained or will erode with disuse. The migratory generations of the North American (NA) monarch butterfly Danaus plexippus lead distinctly different lives from their summer generation NA parents and their tropical descendants living in Costa Rica (CR). NA migratory monarchs postpone reproduction, travel thousands of kilometers south to overwinter in Mexico, and subsist on little food for months. Whether recently dispersed populations of monarchs such as those in Costa Rica, which are no longer subject to selection imposed by migration, retain ancestral seasonal plasticity is unclear. To investigate the differences in seasonal plasticity, we reared the NA and CR monarchs in summer and autumn in Illinois, USA, and measured the seasonal reaction norms for aspects of morphology and metabolism related to flight. NA monarchs were seasonally plastic in forewing and thorax size, increasing wing area and thorax to body mass ratio in autumn. While CR monarchs increased thorax mass in autumn, they did not increase the area of the forewing. NA monarchs maintained similar resting and maximal flight metabolic rates across seasons. However, CR monarchs had elevated metabolic rates in autumn. Our findings suggest that the recent expansion of monarchs into habitats that support year-round breeding may be accompanied by (1) the loss of some aspects of morphological plasticity as well as (2) the underlying physiological mechanisms that maintain metabolic homeostasis in the face of temperature heterogeneity.

Size-dependent Scaling of Stingless Bee Flight Metabolism Reveals an Energetic Benefit to Small Body Size.

Understanding the effect of body size on flight costs is critical for development of models of aerodynamics and animal energetics. Prior scaling studies that have shown that flight costs scale hypometrically have focused primarily on larger (> 100 mg) insects and birds, but most flying species are smaller. We studied the flight physiology of thirteen stingless bee species over a large range of body sizes (1-115 mg). Metabolic rate during hovering scaled hypermetrically (scaling slope=2.11). Larger bees had warm thoraxes while small bees were nearly ecothermic; however, even controlling for body temperature variation, flight metabolic rate scaled hypermetrically across this clade. Despite having a lower mass-specific metabolic rate during flight, smaller bees could carry the same proportional load. Wingbeat frequency did not vary with body size, in contrast to most studies that find wingbeat frequency increases as body size decreases. Smaller stingless bees have greater relative forewing surface area which may help them reduce the energy requirements needed to fly. Further, we hypothesize that the relatively larger heads of smaller species may change their body pitch in flight. Synthesizing across all flying insects, we demonstrate that the scaling of flight metabolic rate changes from hypermetric to hypometric at approximately 58 mg body mass with hypermetic scaling below (slope=1.2) and hypometric scaling (slope=0.67) above 58 mg in body mass. The reduced cost of flight likely provides selective advantages for the evolution of small body size in insects. The biphasic scaling of flight metabolic rates and wingbeat frequencies in insects supports the hypothesis that the scaling of metabolic rate is closely related to the power requirements of locomotion and cycle frequencies.

The thermal performance curve for aerobic metabolism of a flying endotherm.

Performance benefits of stable, warm muscles are believed to be important for the evolution of endothermy in mammals, birds and flying insects. However, thermal performance curves have never been measured for a free-flying endotherm, as it is challenging to vary body temperatures of these animals, and maximal flight performance is difficult to elicit. We varied air temperatures and gas densities to manipulate thoracic temperatures of flying honeybees from 29°C to 44°C, with low air densities used to increase flight metabolic rates to maximal values. Honeybees showed a clear thermal performance curve with an optimal temperature of 39°C. Maximal flight metabolic rates increased by approximately 2% per 1°C increase in thoracic temperature at suboptimal thoracic temperatures, but decreased approximately 5% per 1°C increase as the bees continued to heat up. This study provides the first quantification of the maximal metabolic performance benefit of thermoregulation in an endotherm. These data directly support aerobic capacity models for benefits of thermoregulation in honeybees, and suggest that improved aerobic capacity probably contributes to the multiple origins of endothermic heterothermy in bees and other insects.

Flying on empty: reduced mitochondrial function and flight capacity in food-deprived monarch butterflies.

Mitochondrial function is fundamental to organismal performance, health and fitness - especially during energetically challenging events, such as migration. With this investigation, we evaluated mitochondrial sensitivity to ecologically relevant stressors. We focused on an iconic migrant, the North American monarch butterfly (Danaus plexippus), and examined the effects of two stressors: 7days of food deprivation and infection by the protozoan parasite Ophryocystis elektroscirrha (known to reduce survival and flight performance). We measured whole-animal resting metabolic rate (RMR) and peak flight metabolic rate, and mitochondrial respiration of isolated mitochondria from the flight muscles. Food deprivation reduced mass-independent RMR and peak flight metabolic rate, whereas infection did not. Fed monarchs used mainly lipids in flight (respiratory quotient 0.73), but the respiratory quotient dropped in food-deprived individuals, possibly indicating switching to alternative energy sources, such as ketone bodies. Food deprivation decreased mitochondrial maximum oxygen consumption but not basal respiration, resulting in lower respiratory control ratio (RCR). Furthermore, food deprivation decreased mitochondrial complex III activity, but increased complex IV activity. Infection did not result in any changes in these mitochondrial variables. Mitochondrial maximum respiration rate correlated positively with mass-independent RMR and flight metabolic rate, suggesting a link between mitochondria and whole-animal performance. In conclusion, low food availability negatively affects mitochondrial function and flight performance, with potential implications for migration, fitness and population dynamics. Although previous studies have reported poor flight performance in infected monarchs, we found no differences in physiological performance, suggesting that reduced flight capacity may be due to structural differences or low energy stores.

Carry‐over effects of larval food stress on adult energetics and life history in a nectar‐feeding butterfly

Abstract Stressful juvenile developmental conditions can affect performance and fitness later in life. In holometabolous insects such as butterflies, development under stressful conditions may lead to smaller adult size, lower reproductive output, and shorter lifespan. However, how larval developmental stress affects energy intake and expenditure in adult individuals is poorly understood. We subjected last‐instar larvae of Speyeria mormonia Edwards (Lepidoptera: Nymphalidae) to periodic dietary restriction (DR) to examine the allocation of energy and nutrients among different life history processes. We measured adult food intake, resting metabolic rate (RMR), metabolic flight capacity, lifespan, and reproductive output. Consistent with pressure to disperse from a poor environment while maintaining offspring number, we predicted that stressed individuals would have increased adult food intake and higher flight capacity. Adult body size was strongly reduced. Contrary to predictions, we found no compensatory adult feeding. Mass‐adjusted flight metabolic rate was reduced, suggesting poor dispersal capacity. Larval DR did not affect adult lifespan, nor did the rate of metabolic senescence change. Larval DR did affect RMR, as stressed females had a steeper slope between RMR and body mass, which may reflect differences in physiological activity due to condition. Fecundity decreased less than predicted based on body mass. Instead of investing in flight capacity, females increased relative allocation to reproduction, which may partly buffer against poor environmental conditions. Understanding the interplay of energy acquisition and allocation to life history traits across the life cycle is vital for predicting responses to environmental change.

Insect flight metabolic rate revealed by bolus injection of the stable isotope 13C.

Measuring metabolic rate (MR) poses a formidable challenge in free-flying insects who cannot breathe into masks or be trained to fly in controlled settings. Consequently, flight MR has been predominantly measured on hovering or tethered insects flying in closed systems. Stable isotopes such as labelled water allow measurement of MR in free-flying animals but integrates the measurement over long periods exceeding the average flight duration of insects. Here, we applied the ‘bolus injection of isotopic 13C Na-bicarbonate’ method to insects to measure their flight MR and report a 90% accuracy compared to respirometry. We applied the method on two beetle species, measuring MR during free flight and tethered flight in a wind tunnel. We also demonstrate the ability to repeatedly use the technique on the same individual. Therefore, the method provides a simple, reliable and accurate tool that solves a long-lasting limitation on insect flight research by enabling the measurement of MR during free flight.

Varied Air Temperature and Helium‐Nitrogen Ratios Reveal the Thermal Performance Curve of a Free‐Flying Endotherm

While thermal performance curves are central to understanding the thermal biology of ectotherms and are key parameters for predicting effects of climate change, a thermal performance curve has never been described for free flight of any endotherm. This is likely the case for two reasons: 1) it is challenging to vary the body temperature of most flying endotherms, and 2) thermal performance curves require assessment of maximal performance at each body temperature, and maximal performance is challenging to elicit during fight. We overcame both of these challenges by flying honey bees at 23°C and 35°C air temperatures and at air densities ranging from 1.288-0.441 kg·m-3 (21% O2/79% N2 to 21% O2/79% He). Decreasing air density increases lift requirements, providing a way to test maximal aerobic performance during hovering flight. Helium has a six-fold higher thermal conductance than nitrogen, so interactively changing air temperature and He-N2 content provides a system for varying body temperature. Decreasing air density produced an increase in flight metabolic rate of up to 40% (Fig. 1), and the range of air temperatures and He-N2 content produced thorax temperatures that ranged from 29-44°C (Fig. 2). Plotting the maximal performance at each thorax temperature revealed a clear thermal performance curve, with an optimal temperature of 39°C, and maximal performance decreasing by about 5% with increasing or decreasing 1°C thorax temperature (Fig. 2). These findings help resolve past disagreements about how honey bees thermoregulate during hovering flight; some prior studies have found that flight metabolic rates were constant across air temperatures, while others have reported that decreased metabolic heat production at higher air temperatures contributes to thermoregulation. We found that the range of thorax temperatures differed in these prior studies, and our results suggest that honey bees use different mechanisms to thermoregulate during hovering depending on thorax temperature. Because maximal flight performance will determine capacities to carry loads and fly in challenging environments, the thermal performance curve during flight likely affects the fitness of honey bee colonies, and thus may be useful for developing predictions for effects of climate change on these important pollinators. Also, documentation of the strong effects of thorax temperature on flight performance provides a quantitative measure of the benefits of thermoregulation for this partially homeothermic periodic endotherm.

Consumption of field-realistic doses of a widely used mito-toxic fungicide reduces thorax mass but does not negatively impact flight capacities of the honey bee (Apis mellifera)

Commercial beekeepers in many locations are experiencing increased annual colony losses of honey bees (Apis mellifera), but the causes, including the role of agrochemicals in colony losses, remain unclear. In this study, we investigated the effects of chronic consumption of pollen containing a widely-used fungicide (Pristine®), known to inhibit bee mitochondria invitro, which has recently been shown to reduce honey bee worker lifespan when field-colonies are provided with pollen containing field-realistic levels of Pristine®. We fed field colonies pollen with a field-realistic concentration of Pristine® (2.3ppm) and a concentration two orders of magnitude higher (230ppm). To challenge flight behavior and elicit near-maximal metabolic rate, we measured flight quality and metabolic rates of bees in two lower-than-normal air densities. Chronic consumption of 230 but not 2.3ppm Pristine® reduced maximal flight performance and metabolic rates, suggesting that the observed decrease in lifespans of workers reared on field-realistic doses of Pristine®-laced pollen is not due to inhibition of flight muscle mitochondria. However, consumption of either the 230 or 2.3ppm dose reduced thorax mass (but not body mass), providing the first evidence of morphological effects of Pristine®, and supporting the hypothesis that Pristine® reduces forager longevity by negatively impacting digestive or nutritional processes.

Sex-related interannual plasticity in wing morphological design in Heliconius charithonia enhances flight metabolic performance.

Flight morphological variations and its consequences on animal performance are common in winged insects. In the butterfly Heliconius charithonia, sex-related differences in the wing morphological design have been described resulting in differences in foraging behavior, daily flight distances and flight aerodynamics. It has been suggested that these differences should be reflected in the metabolic capacities and energetic budgets associated with flight in both sexes. In this study, we analyzed the relationship between wing morphological variation and metabolic performance, flight aerodynamics and energetic reserves in females and males of Heliconius charithonia over two years. The results confirm the presence of wing shape sexual dimorphism, but also show an unexpected sex-related annual variation in wing shape, mirrored in the metabolic condition (resting metabolic rate) of individuals. However, contrary to expectation, intersexual variations in wing shape are not related to differences between the sexes in terms of flight aerodynamics, flight metabolic rates, or energetic reserves (carbohydrates, lipids and proteins). Our results indicate a considerable plasticity in H. charithonia wing shape, which we suggest is determined by a trade-off between environmental pressures and reproductive restriction of each sex, maintaining an optimum flight design. Finally, similarities in metabolic rates between young and older males and females in both years may be a consequence of the ability of Heliconius species to feed on pollen.

Host plant defense produces species-specific alterations to flight muscle protein structure and flight-related fitness traits of two armyworms.

Insects manifest phenotypic plasticity in their development and behavior in response to plant defenses, via molecular mechanisms that produce tissue-specific changes. Phenotypic changes might vary between species that differ in their preferred hosts and these effects could extend beyond larval stages. To test this, we manipulated the diet of southern armyworm (SAW; Spodoptera eridania) and fall armyworm (FAW; Spodoptera frugiperda) using a tomato mutant for jasmonic acid plant defense pathway (def1), and wild-type plants, and then quantified gene expression of Troponin t (Tnt) and flight muscle metabolism of the adult insects. Differences in Tnt spliceform ratios in insect flight muscles correlate with changes to flight muscle metabolism and flight muscle output. We found that SAW adults reared on induced def1 plants had a higher relative abundance (RA) of the A isoform of Troponin t (Tnt A) in their flight muscles; in contrast, FAW adults reared on induced def1 plants had a lower RA of Tnt A in their flight muscles compared with adults reared on def1 and controls. Although mass-adjusted flight metabolic rate showed no independent host plant effects in either species, higher flight metabolic rates in SAW correlated with increased RA of Tnt A Flight muscle metabolism also showed an interaction of host plants with Tnt A in both species, suggesting that host plants might be influencing flight muscle metabolic output by altering Tnt This study illustrates how insects respond to variation in host plant chemical defense by phenotypic modifications to their flight muscle proteins, with possible implications for dispersal.

The highs and lows of bird flight.

Bar-headed geese lower their flight metabolic rates to fly in low-oxygen conditions.

Effects of flight and food stress on energetics, reproduction, and lifespan in the butterfly Melitaea cinxia

Environmental change can have drastic effects on natural populations. To successfully predict such effects, we need to understand how species that follow different life-history strategies respond to stressful conditions. Here I focus on two stressors, increased flight and dietary restriction, and their effects on bioenergetics and life-history. Using the Glanville fritillary butterfly (Melitaea cinxia), I subjected mated females to three treatments: (1) control conditions, (2) repeated forced flight with unlimited food, and (3) repeated forced flight coupled with food restriction. Interestingly, flight increased fecundity: females in both flight treatments initiated oviposition earlier, laid more egg clutches, and had higher total fecundity than control females. However, food-restriction by 50% reduced clutch size and resulted in an approximately 25% decrease in total fecundity compared to flown females with unlimited food. There were no differences in egg wet mass, water content or hatching success. Flown females with unlimited food appeared to exhibit a trade-off between reproduction and lifespan: they had higher mass-independent resting metabolic rate and shorter lifespan than females in the other treatments. Mass-independent flight metabolic rate, reflecting flight capacity, did not differ among the treatments. There were no differences in the rate of metabolic senescence across the treatments. The current findings suggest a mechanistic link between flight and reproduction, potentially mediated by juvenile hormone signalling. It appears that this wing-monomorphic butterfly does not show an oogenesis-flight trade-off often found in wing-dimorphic insects. Nevertheless, nectar-feeding is needed for achieving maximum reproductive output, suggesting that diminishing nectar resources may negatively impact natural populations.