Thermoregulatory strategies in two closely related sympatric Scarabaeus species (Coleoptera: Scarabaeinae)

Abstract. The thermoregulation strategies of Scarabaeus sacer L. and Scarabaeus cicatricosus Lucas were studied in the Doñana National Park, Spain. In this area, both species coexist, showing the same habitat and food preferences. However, S. cicatricosus is active during warmer parts of the day compared to S. sacer. Both species thermoregulate their thoracic temperature but, whereas the abdomen of S. sacer is a passive thermal window, S. cicatricosus actively thermoregulates abdominal temperature by increasing heat transfer from the thorax to the abdomen at high T_a values. In the case of S. sacer, their endothermy indicates an adaptive capacity to thorax heat retention, as occurs mainly in winter‐flying insects. This mechanism, possibly related to the aerodynamic flight posture in Scarabaeinae, could be an effective barrier to retard the rate of abdominal heat loss during flight. This endothermic strategy makes flight difficult at higher temperatures, although it allows flight during cooler periods of the day. On the other hand, S. cicatricosus showed a different adaptive behaviour to S. sacer. In this case, a significant decrease in abdominal heat loss at higher ambient temperatures would indicate a decrease in heat transfer from the thorax to the abdomen, as occurs in some desert and semiarid insects. This ‘heat exchanger’ mechanism observed in S. cicatricosus could be due to the irregular posture adopted during flight, with the posterior legs clearly extended and separate from the body. This behaviour increases turbulence and convective cooling, favouring exposure of the soft abdominal tergal cuticle and, subsequently, water loss. Thus, for S. cicatricosus, the well‐adapted ‘heat exchanger’ permits flight during periods of the day when temperatures would possibly be lethal for those species with high endothermy. From an adaptive viewpoint, these mechanisms of thermoregulation may explain how both closely‐related sympatric species respond in different ways to environmental temperature, favouring their coexistence.     

Body temperature and thermoregulation in two species of yellowjackets,Vespula germanica and V. maculifrons

In spite of the abundance and broad distribution of social wasps, little information exists concerning thermoregulation by individuals. We measured body temperatures of the yellowjackets Vespula germanica and V. maculifrons and examined their thermoregulatory mechanisms. V. germanica demonstrated thermoregulation via a decreasing gradient between thorax temperature and ambient temperature as ambient temperature increased. V. maculifrons exhibited a constant gradient at lower ambient temperatures but thorax temperature was constant at high ambient temperatures. Head temperature exhibited similar patterns in both species. In spite of low thermal conductances, a simple heat budget model predicts substantial heat loads in warm conditions in the absence of thermoregulation. Both species regurgitated when heated on the head. A smaller volume of regurgitant was produced at lower head temperatures and a larger volume at higher head temperatures. Small regurgitations resulted in stabilization of head temperature, while large ones resulted in 4°C decreases in head temperature. Regurgitation was rare when wasps were heated upon the thorax. Abdomen temperature was 3–4°C above ambient temperature, and approached ambient temperature under the hottest conditions. No evidence was found for shunting of hot hemolymph from thorax to abdomen as a cooling mechanism. The frequency of regurgitation in workers returning to the nest increased with ambient temperature. Regurgitation may be an important thermoregulatory strategy during heat stress, but is probably not the only mechanism used in yellowjackets.Abbreviations M _b body mass - M _th thorax mass - T _a ambient temperature - T _ab abdomen temperature - T _b body temperature - T _h head temperature - T _th thorax temperature - C _t thermal conductance     

Thermoregulatory physiology of the carpenter bee,Xylocopa varipuncta

Summary The carpenter beesXylocopa varipuncta maintain thoracic temperatures of 33.0°C to 46.5°C during continuous free flight from 12°C to 40°C. Since the thoracic temperature excess is not constant (decreasing from 24°C at low air temperatures to 6°C at high) the bees are thermoregulating. We document physiological transfer of relatively large amounts of heat to the abdomen and to the head during pre-flight warm-up and during artificial thoracic heating. Most of the temperature increase of the head is due to passive conduction, while that of the abdomen is due to active physiological heat transfer despite a series of convolutions of the aorta in the petiole that anatomically conform to a counter-current heat exchanger. Although the thermoregulatory mechanisms during flight are far from clarified, our data suggest that thermoregulation involves a strong reliance on active convective cooling through increased flight speed.     

Effects of flight behaviour on body temperature and kinematics during inter-male mate competition in the solitary desert bee Centris pallida

Abstract. Body temperatures and kinematics are measured for male Centris pallida bees engaged in a variety of flight behaviours (hovering, patrolling, pursuit) at a nest aggregation site in the Sonoran Desert. The aim of the study is to test for evidence of thermoregulatory variation in convective heat loss and metabolic heat production and to assess the mechanisms of acceleration and forward flight in field conditions. Patrolling males have slightly (1–3 °C) cooler body temperatures than hoverers, despite similar wingbeat frequencies and larger body masses, suggesting that convective heat loss is likely to be greater during patrolling flight than during hovering. Comparisons of thorax and head temperature as a function of air temperature (T_a) indicate that C. pallida males are thermoregulating the head by increasing heat transfer from the thorax to the head at cool T_a. During patrolling flight and hovering, wingbeat frequency significantly decreases as T_a increases, indicating that variation in metabolic heat production contributes to thermal stability during these behaviours, as has been previously demonstrated for this species during flight in a metabolic chamber. However, wingbeat frequency during brief (1–2 s) pursuits is significantly higher than during other flight behaviours and independent of T_a. Unlike most other hovering insects, C. pallida males hover with extremely inclined stroke plane angles and nearly horizontal body angles, suggesting that its ability to vary flight speed depends on changes in wingbeat frequency and other kinematic mechanisms that are not yet described.     

EVOLUTION AND COADAPTATION OF THERMOREGULATORY BEHAVIOR AND WING PIGMENTATION PATTERN IN PIERID BUTTERFLIES

This paper addresses the question of how the relationship between morphological structure and functional performance differs in related groups of organisms. I describe the relationship between a suite of phenotypic characters (behavioral posture and the pattern of wing pigmentation) and one function of these characters (thermoregulatory performance) for two groups of butterflies in the family Pieridae, focusing on how behavior and wing pattern interact to affect specific aspects of thermoregulation. Using both natural and experimentally created variation in wing-melanization patterns, I develop and test a series of predictions about the relations among thermoregulatory posture, melanization pattern, body temperature, and flight activity. Results show that increased melanization in different wing regions has positive, negative, or neutral effects in increasing body temperature of Pieris butterflies. The angle of the wings used during basking alters the relative importance of different modes of heat transfer and thereby determines the contribution of different dorsal wing regions to thermoregulation. Experimentally increased dorsal melanization can either increase or decrease the onset of flight activity and can directly alter thermoregulatory posture. For Pieris, dorsal melanization affects basking and flight, while ventral melanization primarily affects overheating. These results are used to generate a functional map relating melanization pattern to thermoregulatory performance in Pieris. Reflectance-basking posture, white background color, and melanization pattern represent coadapted characters in Pieris that interact to determine thermoregulatory performance. The differences in thermoregulatory posture and background color between pierid butterflies in the subfamilies Pierinae and Coliadinae have led to a reorganization and partial reversal of the thermoregulatory effects of melanization pattern. I suggest that this change in the physical mechanism of thermoregulatory adaption in pierids has qualitatively altered the nature of selection on wing-melanization pattern.     

A role for thermoregulation in the <Emphasis Type="Italic">Polistes dominulus</Emphasis> invasion: a comparison of the thermoregulatory abilities of the invasive wasp <Emphasis Type="Italic">P. dominulus</Emphasis> and the native wasp <Emphasis Type="Italic">P. fuscatus</Emphasis>

Social insects are excellent invaders that have had negative impacts on native species and humans. Many invasive species move from warmer to cooler climates. For these species, thermal adaptations may both be important for their ability to invade and to limit their invasion range. The invasion of Polistes dominulus into North America provides an example of a primitively eusocial invader from a warmer climate. We studied the differences in thermoregulation between P. dominulus and the native P. fuscatus. We found that, during flight, thorax temperature in P. fuscatus was less affected by ambient temperature than thorax temperature of P. dominulus. We also found that P. dominulus and P. fuscatus showed different patterns of warming after removal from a cold environment. Unlike P. dominulus, live P. fuscatus never fully cooled down in a cold environment. P. fuscatus also reached their relative minimum flight temperatures earlier than P. dominulus, but P. dominulus maintained higher elevated temperatures for longer. These differences in thermoregulatory ability suggest that the lower winter survival of P. dominulus could be offset by a greater thermal tolerance during flight, while the lower thermal tolerance of P. fuscatus in flight is offset somewhat by better thermoregulatory ability.     

Small bees overheat in sunlit flowers: do they make cooling flights?

1. Although thermoregulation by large bees in cool climates has been well studied, less is known about the very different thermoregulatory strategies of small bees, especially those subjected to heat stress. 2. Studies were carried out on small (< 20 mg fresh weight), dark‐coloured, solitary bees (mostly halictids and hylaeine colletids) experiencing an extreme radiative heat load, enhanced by the high‐altitude location and by reflection of incident radiation by the high‐albedo petals of the flowers of Potentilla lancinata. 3. When foraging in the flowers, such bees experienced peak operative temperatures exceeding 44 °C. In these conditions, males largely stopped foraging but females continued, usually limiting their flower visits to a few seconds and making frequent short flights. These flights would cool the bees down, because bees suspended in air were cooler than bees in sunlit flowers, and convective cooling during flight would further enhance the cooling effect of departure from the flower. 4. As far as is known, cooling flights in small bees have not been proposed before, providing a new avenue for exploration of bee thermoregulatory strategies.     

Alterations in alpha-adrenergic and muscarinic cholinergic receptor binding in rat brain following nonionizing radiation

Microwave radiation produces hyperthermia. The mammalian thermoregulatory system defends against changes in temperature by mobilizing diverse control mechanisms. Neurotransmitters play a major role in eliciting thermoregulatory responses. The involvement of adrenergic and muscarinic cholinergic receptors was investigated in radiation-induced hyperthermia. Rats were subjected to radiation at 700 MHz frequency and 15 mW/cm2 power density and the body temperature was raised by 2.5 degrees C. Of six brain regions investigated only the hypothalamus showed significant changes in receptor states, confirming its pivotal role in thermoregulation. Adrenergic receptors, studied by [3H]clonidine binding, showed a 36% decrease in binding following radiation after a 2.5 degrees C increase in body temperature, suggesting a mechanism to facilitate norepinephrine release. Norepinephrine may be speculated to maintain thermal homeostasis by activating heat dissipation. Muscarinic cholinergic receptors, studied by [3H]quinuclidinyl benzilate binding, showed a 65% increase in binding at the onset of radiation. This may be attributed to the release of acetylcholine in the hypothalamus in response to heat cumulation. The continued elevated binding during the period of cooling after radiation was shut off may suggest the existence of an extra-hypothalamic heat-loss pathway.     

Can Newts Cope with the Heat? Disparate Thermoregulatory Strategies of Two Sympatric Species in Water

Many ectotherms effectively reduce their exposure to low or high environmental temperatures using behavioral thermoregulation. In terrestrial ectotherms, thermoregulatory strategies range from accurate thermoregulation to thermoconformity according to the costs and limits of thermoregulation, while in aquatic taxa the quantification of behavioral thermoregulation have received limited attention. We examined thermoregulation in two sympatric newt species, Ichthyosaura alpestris and Lissotriton vulgaris, exposed to elevated water temperatures under semi-natural conditions. According to a recent theory, we predicted that species for which elevated water temperatures pose a lower thermal quality habitat, would thermoregulate more effectively than species in thermally benign conditions. In the laboratory thermal gradient, L. vulgaris maintained higher body temperatures than I. alpestris. Semi-natural thermal conditions provided better thermal quality of habitat for L. vulgaris than for I. alpestris. Thermoregulatory indices indicated that I. alpestris actively thermoregulated its body temperature, whereas L. vulgaris remained passive to the thermal heterogeneity of aquatic environment. In the face of elevated water temperatures, sympatric newt species employed disparate thermoregulatory strategies according to the species-specific quality of the thermal habitat. Both strategies reduced newt exposure to suboptimal water temperatures with the same accuracy but with or without the costs of thermoregulation. The quantification of behavioral thermoregulation proves to be an important conceptual and methodological tool for thermal ecology studies not only in terrestrial but also in aquatic ectotherms.     

Sound production and endothermy in the horse botfly, Gasterophilus intestinalis

ABSTRACT. Adult Gasterophilus intestinalis (De Geer) frequently produce a 'buzzing' sound while stationary. This buzzing was always associated with heat production in the fly's thorax, although sometimes heat production occurred without audible buzzing. Thoracic temperature (Tth) could be elevated by as much as 12°C. As buzzing continued, the Tth rose, the pitch of the buzzing sound increased, the frequency being directly proportion to Tth. Periods of buzzing were usually, but not always, terminated by attempted flight. Often, flies showed long episodes of cycling, when periods of continuous buzzing were interspersed with periods of rest. Such cycling maintained Tth above ambient temperature for long periods. During sustained tethered flight, flies were able to maintain Tth at high, steady values for long periods. Heat loss from the thorax is restricted by a dense covering of hair, and also by active control over heat transfer between thorax and abdomen.     

Thermoregulation of foraging honeybees on flowering plants: seasonal variability and influence of radiative heat gain

1. During nectar and pollen foraging in a temperate climate, honeybees are exposed to a broad range of ambient temperatures, challenging their thermoregulatory ability. The body temperature that the bees exhibit results from endothermic heat production, exogenous heat gain from solar radiation, and heat loss. In addition to profitability of foraging, season was suggested to have a considerable influence on thermoregulation. To assess the relative importance of these factors, the thermoregulatory behaviour of foragers on 33 flowering plants in dependence on season and environmental factors was investigated.2. The bees (Apis mellifera carnica Pollman) were always endothermic. On average, the thorax surface temperature (T(th)) was regulated at a high and rather constant level over a broad range of ambient temperatures (T(th) = 33.7-35.7°C, T(a) = 10-27°C). However, at a certain T(a), T(th) showed a strong variation, depending on the plants from which the bees were foraging. At warmer conditions (T(a) = 27-32°C) the T(th) increased nearly linearly with T(a) to a maximal average level of 42.6 °C. The thorax temperature excess decreased strongly with increasing T(a) (T(th)-T(a) = 21.6 - 3.6°C).3. The bees used the heat gain from solar radiation to elevate the temperature excess of thorax, head, and abdomen. Seasonal dependance was reflected in a 2.7 °C higher mean T(th) in the spring than in the summer. An anova revealed that season had the greatest effect on T(th), followed by T(a) and radiation.4. It was presumed the foragers' motivational status to be the main factor responsible for the variation of T(th) between seasons and different plants.     

Vasopressin in thermoregulation--competitive demands: experimental evidence and theoretical considerations.

Previous studies have substantiated the antipyretic role played by extrahypothalamic limbic system (EXHY-LS) AVP during fever. Repeated attempts to elucidate other thermoregulatory functions of this hormone have failed. Circumstantial evidence, however, suggest central role for this hormone in thermoregulation under hypohydration. Hypohydration, hyperosmolarity and hypovolaemia induced upward shifts in temperature thresholds for activation of heat dissipating mechanisms. When hypovolaemia is superimposed on hyperosmolarity these shifts are additive. Analogously, these two stressors when combined, decrease the osmotic threshold for AVP release. In rats, the elevated temperature thresholds for evaporative cooling and peripheral vasodilation occurring with hypohydration are positively correlated with lower Hypothalamic/EXHY-LS AVP ratio. Reciprocal relations between limbic system and blood AVP contents suggest competitive interaction between central and peripheral demands. Hypothesis for the possible mode of action of central AVP in thermoregulation under hypohydration is discussed.     

Ontogeny of thermoregulation in precocial birds

The aim of this paper is to summarise the results of earlier experiments on thermoregulation and heat balance in birds, to present new results concerning thermoregulation during the perinatal period in precocial embryos and to develop a model of the ontogeny of thermoregulation over the whole lifespan of birds. The ontogeny of thermoregulation in precocial birds is characterised by three phases with different efficiency of the system. In the prenatal phase, all control elements of the thermoregulatory system can function, but the efficiency of the system is low. It is postulated that endothermic reactions during the prenatal period do not have a proximate (immediate), but rather an ultimate influence on the efficiency of thermoregulation. They may support adaptivity to expected environmental conditions and may be involved in epigenetic adaptation processes. During the early postnatal phase, the thermoregulatory system develops and matures. Summit metabolism and resting metabolic rate and their thermoregulatory set points increase. Preferred temperature is significantly different during different behavioural activities. The phase of full-blown homeothermy starts at approximately the 10th day of life. It is characterised by an activation order of thermoregulatory control elements and by secondary chemical thermoregulation. The influence of thermal and non-thermal climatic factors on heat production and heat loss may be described by mathematical models.     

THERMOREGULATORY ADAPTATIONS IN MARINE MAMMALS: INTERACTING EFFECTS OF EXERCISE AND BODY MASS. A REVIEW1

A review of thermoregulation in marine mammals led to the following conclusions: very little is known about thermoregulation in large cetaceans. The only measured value for the metabolic rate of a whale, albeit a young one, was substantially higher than the predicted value for a terrestrial mammal of similar size. Very small and newborn marine mammals rely on a high metabolic heat production to sustain their body temperature during exposure to cold or in the water. The considerable insulation of some adult marine mammals may absolve them from the need for a high level of heat production. One marine mammal in tropical or subtropical waters is hypometabolic. There is evidence for a powerful control of thermoregulatory mechanisms by the anterior hypothalamic/preoptic region of the brain in two species. Thermoregulation in marine mammals during exercise remains paradoxical.     

Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies

In nectar-feeding butterflies, reproductive potential is usually thought to depend on the size of the reproductive reserves in the abdomen, the adult food quality and, for females, the amount of resources received in the spermatophores at mating. Recent findings show that thorax mass and nitrogen content decrease with age in some butterfly species, and that thorax resources may be used for reproduction in the butterfly Pieris napi , just as in some other insects. In order to determine whether this is a general pattern and ascertain how it relates to the investment of resources in reproduction we studied the dynamics of thorax and abdomen mass changes in 11 Swedish butterfly species. By regressing thorax and abdomen mass on age of field-collected specimens, we show that loss of mass from both the thorax and the abdomen is a common phenomenon among nectar-feeding temperate zone butterflies under natural conditions. We argue that our results indicate that resources from flight muscles can be reallocated to reproduction by these butterflies, thus increasing their reproductive potential. Within species, females use proportionately more resources from the thorax than do males, as expected from the difference in investment of resources in reproduction. Among males we expect species with a higher reproductive investment to have a larger decrease in thorax and abdomen mass, and our data indicate that this is the case. Looking at the change in relative thorax mass, our results suggest that the use of resources from the thorax does not affect flight performance negatively, something that could constrain the use of muscle resources.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 363–380.     

Thermoregulation and microhabitat use in mountain butterflies of the genus Erebia: Importance of fine-scale habitat heterogeneity

Mountain butterflies have evolved efficient thermoregulation strategies enabling their survival in marginal conditions with short flight season and unstable weather. Understanding the importance of their behavioural thermoregulation by habitat use can provide novel information for predicting the fate of alpine Lepidoptera and other insects under ongoing climate change. We studied the link between microhabitat use and thermoregulation in adults of seven species of a butterfly genus Erebia co-occurring in the Austrian Alps. We captured individuals in the field and measured their body temperature in relation to microhabitat and air temperature. We asked whether closely related species regulate their body temperature differently, and if so, what is the effect of behaviour, species traits and individual traits on body to air and body to microhabitat temperature differences. Co-occurring species differed in mean body temperature. These differences were driven by active microhabitat selection by individuals and also by species–specific habitat preferences. Species inhabiting grasslands and rocks utilised warmer microclimates to maintain higher body temperature than woodland species. Under low air temperatures, species of rocky habitats heated up more effectively than species of grasslands and woodlands which allowed them to stay active in colder weather. Species morphology and individual traits play rather minor roles in the thermoregulatory differences; although large species and young individuals maintained higher body temperature. We conclude that diverse microhabitat conditions at small spatial scales probably contribute to sympatric occurrence of closely related species with different thermal demands and that preserving heterogeneous conditions in alpine landscapes might mitigate detrimental consequences of predicted climate change.     

Thermoregulation in rapid growing broiler chickens is compromised by constraints on radiative and convective cooling performance

Broiler chickens are selected to undergo a rapid six-week hatch-to-slaughter growth phase to attain large body and muscle mass. Broilers have relatively high resting and locomotor metabolic costs suggesting that adaptive thermoregulatory mechanisms are required to dissipate excess heat. Using thermal imaging in the growing broiler we characterised the trajectory of radiative and convective cooling in still air across broiler development. Scaling of head, tarsus and toe surface area did not deviate from body mass^2/3 while torso area increased with positive allometry, body mass^0.82, reflecting increased feather coverage and/or disproportionate abdominal/thoracic growth. Despite relatively increased area, the body became less effective for heat transfer presumably due to increasing feather coverage. Conversely, the magnitude of heat exchange from the distal hindlimbs was improved in larger birds. Overall capacity to transfer heat by convection and radiation in still air was attenuated over development, since the proportion of resting metabolic rate accounted for decreased in standing and sitting postures. This physiological constraint could be ameliorated by increased latent heat transfer or provision of environmental ventilation, which we modelled according to industrial guidelines. Based on models, higher airspeeds coincided with improved convective cooling that assisted in maintaining the proportion of RMR accounted for by convective and radiative heat transfer. These data highlight the potentially adverse thermoregulatory effects of rapid growth rate and body mass increases, which may contribute to the increased sedentary resting and decreased locomotor behaviour observed in large broilers.     

Mechanisms of Thermoregulation in Flying Bees

SYNOPSIS. Thermoregulation of elevated thorax temperatures isnecessary for bees to achieve the high rates of power productionrequired for flight, and is a key factor allowing them to occupywidely varying thermal environments. However, the mechanismsby which bees thermoregulate during flight are poorly understood.Thermoregulation is accomplished by balancing heat gain andheat loss via the following routes: convection, evaporation,and metabolic heat production. There appears to be a diversityof thermoregulatory mechanisms employed during flight amongbee species. Some species, particularly Bombus spp., activelyincrease the distribution of thoracic heat to the abdomen duringflight as air temperature (T_a) rises, and apparently thermoregulateby varying convective heat loss. However, thermal variationin convection has not been directly measured for any free-flyingbee. Above 33°C, flying Apis mellifera greatly increaseevaporative heat loss with T_a, and many other species "tongue-lash"during flight at high T_as or when artificially heated. Thus,evaporation seems to be important for preventing overheatingduring flight at very high T_as. Flying A. mellifera and Centrispallida strongly decrease metabolic rate as T_a increases, suggestingthat they are varying metabolic heat production for thermoregulationand not aerodynamic requirements. Variation in metabolic heatproduction appears to be mediated by changes in wingbeat kinematics,since wingbeat frequency decreases with T_a for A. melliferaand Centris spp. It is unknown if the decrease in flight metabolicrate at higher T_as occurs secondarily as a consequence of greaterefficiency or if it is truly an active response.     

Thermoregulation and the effect of body temperature on call temporal parameters in the cicada Diceroprocta olympusa (Homoptera: Cicadidae)

We investigated the thermoregulatory behavior, thermal responses (minimum flight, maximum voluntary tolerance and heat torpor temperatures) and the effect of body temperature (T(b)) on call parameters in the cicada Diceroprocta olympusa (Walker). Regression of T(b) as a function of ambient (T(a)) or perch temperatures (T(p)) suggests thermoregulation is occurring. Thermoregulation occurs through behavioral changes that alter the uptake of solar radiation. T(p) is a better predictor of T(b) than is T(a). Thermal responses (minimum flight temperature 20.4 degrees C, maximum voluntary tolerance temperature 37 degrees C, and heat torpor temperature 46.7 degrees C) may be related to the humid, grassland habitat of the species. In contrast to other acoustic insects, no significant relationship was found between the temporal parameters of the calling song and T(b) within the population of D. olympusa.     

Thermal physiological ecology of Colias butterflies in flight

Summary As a comparison to the many studies of larger flying insects, we carried out an initial study of heat balance and thermal dependence of flight of a small butterfly (Colias) in a wind tunnel and in the wild.Unlike many larger, or facultatively endothermic insects, Colias do not regulate heat loss by altering hemolymph circulation between thorax and abdomen as a function of body temperature. During flight, thermal excess of the abdomen above ambient temperature is weakly but consistently coupled to that of the thorax. Total heat loss is best expressed as the sum of heat loss from the head and thorex combined plus heat loss from the abdomen because the whole body is not isothermal. Convective cooling is a simple linear function of the square root of air speed from 0.2 to 2.0 m/s in the wind tunnel. Solar heat flux is the main source of heat gain in flight, just as it is the exclusive source for warmup at rest. The balance of heat gain from sunlight versus heat loss from convection and radiation does not appear to change by more than a few percent between the wings-closed basking posture and the variable opening of wings in flight, although several aspects require further study. Heat generation by action of the flight muscles is small (on the order of 100 m W/g tissue) compared to values reported for other strongly flying insects. Colias appears to have only very limited capacity to modulate flight performance. Wing beat frequency varies from 12–19 Hz depending on body mass, air speed, and thoracic temperature. At suboptimal flight temperatures, wing beat frequency increases significantly with thoracic temperature and body mass but is independent of air speed. Within the reported thermal optimum of 35–39°C, wing beat frequency is negatively dependent on air speed at values above 1.5 m/s, but independent of mass and body temperature. Flight preference of butterflies in the wind tunnel is for air speeds of 0.5–1.5 m/s, and no flight occurs at or above 2.5 m/s. Voluntary flight initiation in the wild occurs only at air speeds 1.4 m/s.In the field, Colias fly just above the vegetation at body temperatures of 1–2°C greater than when basking at the top of the vegetation. These measurements are consistent with our findings on low heat gain from muscular activity during flight. Basking temperatures of butterflies sheltered from the wind within the vegetation were 1–2°C greater than flight temperatures at vegetation height.