Diel patterns of activity of Thymelicus lineola adults (Lepidoptera: Hesperiidae) in relation to weather *

ABSTRACT. 1. Daily patterns of activity, thoracic temperature (T_th) and thermoregulatory behaviour in relation to environmental conditions were studied in the European skipper Thymelicus lineola (Ochsenheimer) adults. 2. Daily activity was limited mainly by T_th, which in turn was dependent on air temperature (T_a) and sunlight. However, when light intensity fell to < 100 W m^?2, skippers also became inactive. 3. T_th was sometimes as much as 12°C above T_a, and this was most pronounced under cool conditions when basking activity predominated. 4. Black globe temperature (T_bg) which, in the absence of wind, is influenced by both T^a, and radiant heat load in a manner similar to a basking butterfly, was used to approximate the maximum T_th attained by T.lineola through basking. 5. Both males and females basked at T_bg >20°C, and if this temperature was not attained, skippers remained inactive. As T_bg increased, basking activity declined more rapidly in males than in females. Basking bouts were also shorter in males. 6. Males also flew, fed and courted females at T_bg >20°C, while females only fed and flew at T_bg >22°C. The percentage of both sexes feeding and flying, and courting in the case of males, were positively correlated with T_bg. 7. In warm weather, males divided their time equally between flying and feeding, while females spent the majority of their time feeding. Feeding bouts were shorter and flying bouts were longer in males than in females. Flight duration in males was positively correlated with T_bg. 8. Skippers avoided, and, consequently laid fewer eggs in, shady areas. 9. Sexual differences in activity patterns in relation to weather reflect differences in the reproductive requirements of the two sexes.     

Thermoregulation in the primitively eusocial paper wasp, Polistes dominulus

Regulation of wing muscle temperature is important for sustaining flight in many insects, and has been well studied in honeybees. It has been much less well studied in wasps and has never been demonstrated in Polistes paper wasps. We measured thorax, head, and abdomen temperatures of inactive Polistes dominulus workers as they warmed after transfer from 8 to ~25°C ambient temperature, after removal from hibernacula, and after periods of flight in a variable temperature room. Thorax temperature (T _th) of non-flying live wasps increased more rapidly than that of dead wasps, and T _th of some live wasps reached more than 2°C above ambient temperature (T _a), indicating endothermy. Wasps removed from hibernacula had body region temperatures significantly above ambient. The T _th of flying wasps was 2.5°C above ambient at T _a = 21°C, and at or even below ambient at T _a = 40°C. At 40°C head and abdomen temperatures were both more than 2°C below T _a, indicating evaporative cooling. We conclude that P. dominulus individuals demonstrate clear, albeit limited, thermoregulatory capacity.     

Thermoregulation of water foraging honeybees—Balancing of endothermic activity with radiative heat gain and functional requirements

Foraging honeybees are subjected to considerable variations of microclimatic conditions challenging their thermoregulatory ability. Solar heat is a gain in the cold but may be a burden in the heat. We investigated the balancing of endothermic activity with radiative heat gain and physiological functions of water foraging Apis mellifera carnica honeybees in the whole range of ambient temperatures (T_a) and solar radiation they are likely to be exposed in their natural environment in Middle Europe.The mean thorax temperature (T_th) during foraging stays was regulated at a constantly high level (37.0-38.5 °C) in a broad range of T_a (3-30 °C). At warmer conditions (T_a = 30-39 °C) T_th increased to a maximal level of 45.3 °C. The endothermic temperature excess (difference of T_body − T_a of living and dead bees) was used to assess the endogenously generated temperature elevation as a correlate of energy turnover. Up to a T_a of ∼30 °C bees used solar heat gain for a double purpose: to reduce energetic expenditure and to increase T_th by about 1-3 °C to improve force production of flight muscles. At higher T_a they exhibited cooling efforts to get rid of excess heat. A high T_th also allowed regulation of the head temperature high enough to guarantee proper function of the bees’ suction pump even at low T_a. This shortened the foraging stays and this way reduced energetic costs. With decreasing T_a bees also reduced arrival body weight and crop loading to do both minimize costs and optimize flight performance.     

Thermoregulation of water foraging wasps (Vespula vulgaris and Polistes dominulus)

A comparison of the thermoregulation of water foraging wasps (Vespula vulgaris, Polistes dominulus) under special consideration of ambient temperature and solar radiation was conducted. The body surface temperature of living and dead wasps was measured by infrared thermography under natural conditions in their environment without disturbing the insects’ behaviour. The body temperature of both of them was positively correlated with T_a and solar radiation. At moderate T_a (22–28 °C) the regression lines revealed mean thorax temperatures (T_th) of 35.5–37.5 °C in Vespula, and of 28.6–33.7 °C in Polistes. At high T_a (30–39 °C) T_th was 37.2–40.6 °C in Vespula and 37.0–40.8 °C in Polistes. The thorax temperature excess (T_th–T_a) increased at moderate T_a by 1.9 °C (Vespula) and 4.4 °C (Polistes) per kW⁻¹ m⁻². At high T_a it increased by 4.0 °C per kW⁻¹ m⁻² in both wasps. A comparison of the living water foraging Vespula and Polistes with dead wasps revealed a great difference in their thermoregulatory behaviour. At moderate T_a (22–28 °C) Vespula exhibited distinct endothermy in contrast to Polistes, which showed only a weak endothermic activity. At high T_a (30–39 °C) Vespula reduced their active heat production, and Polistes were always ectothermic. Both species exhibited an increasing cooling effort with increasing insolation and ambient temperature.     

Energy metabolism and body temperature in 13 sunbird species (Nectariniidae)

• 1.1. Diurnal cycles of body temperature, T_b, and energy metabolism, M, at different ambient temperatures (T_a: +5 −+ 32°C) were tested in 13 sunbird species from various habitats and of different body masses (5.2–14.2 g) including one of the smallest passerines, Aethopyga christinae.
• 2.2. Resting M-level (night) reaches T_a-dependent mean values of 54% (+5°C) and 49% (+25°C) of activity M-levels (day). Expected level is ca 75%.
• 3.3. Resting metabolic rate of sunbirds lies within the range of theoretically expected values for birds.
• 4.4. Mean linear metabolism-weight regression of the night values follows: M = 0.102 × W^0.712 (M = energy metabolism in kJ/hr and W = body mass in g).
• 5.5. Thermal conductances, T_c, are lower (−24%) than the predicted values. This is caused by a decrease of T_b at low T_a. Mean nocturnal T_c is 3.2 J/g × hr × °C, mean day-time value is 4.3 J/g × hr × °C.
• 6.6. The zone of thermoneutrality is, in most species, within a T_a-range of 24–28°C.
• 7.7. Normal day and night levels of T_b are in the same range as reported for other birds of the same weight class. T_b decreases slightly with falling T_a (partial heterothermia). Lowest recorded T_b was 34.2°C.
• 8.8. No species tested showed any sign of torpor at night, independent of T_a, body mass or habitat origin.

     

Determinants of bovine thermal response to heat and solar radiation exposures in a field environment

Continuous exposure of cattle to summer heat in the absence of shade results in significant hyperthermia and impairs growth and general health. Reliable predictors of heat strain are needed to identify this condition. A 12-day study was conducted during a moderate summer heat period using 12 Angus x Simmental (Bos taurus) steers (533 ± 12 kg average body weight) to identify animal and ambient determinations of core body temperature (T _core) and respiration rate (RR) responses to heat stress. Steers were provided standard diet and water ad libitum, and implanted intraperitoneally with telemetric transmitters to monitor T _core hourly. Visual count of flank movement at 0800 and 1500 hours was used for RR. Dataloggers recorded air temperature (T _a), and black globe temperatures (T _bg) hourly to assess radiant heat load. Analysis was across four periods and 2 consecutive days averaged within each period. Average T _a and T _bg increased progressively from 21.7 to 30.3°C and 25.3 to 34.0°C, respectively, from the first to fourth periods. A model utilizing a quadratic function of T _a explained the most variation in T _core (R ² = 0.56). A delay in response from 1 to 3 h did not significantly improve R ² for this relationship. Measurements at 0800 and 1500 hours alone are sufficient to predict heat strain. Daily minimum core body temperature and initial 2-h rise in T _a were predictors of maximum core temperature and RR. Further studies using continuous monitoring are needed to expand prediction of heat stress impact under different conditions.     

Is foraging in the desert ant,Messor pergandei (Hymenoptera: Formiceidae), limited by water?

Abstract.

• 1 The seed-harvesting ant Messor (Veromessor) prrgandei (Mayr) is a common inhabitant of southwestern deserts of the U.S.A. Foragers vary in size from less than 1 mg to more than 10 mg in body mass and may travel over 80 m on a single foraging trip. Their small size, long foraging range, and hot, arid habitat suggest that water stress may limit foraging activity. We examined intercolony and interindividual variation in water loss of M.pergandei foragers under several different situations in the field.
• 2 Colonies differed significantly in minimum critical water content (W_c) of individual foragers (water content below which foragers are incapable of normal locomotion). In one colony small workers had disproportionately higher W_c than larger workers; in the other colony W_c was isometric with body size.
• 3 Groups of workers confined in the field approached W_c only after normal foraging stopped and substrate temperatures exceeded 45°C, while water content of individual foragers did not approach the W_c during normal foraging periods. Moreover, seed load and distance travelled did not negatively affect forager water content, as measured on return to the nest: indeed, our results suggest that forager hydration level may influence load selection and/or foraging distance. We conclude that, under normal circumstances, foraging in M.pergandei is not water-limited.

     

Seasonal control of energy requirements for thermoregulation in the djungarian hamster (Phodopus sungorus), living in natural photoperiod

Summary Djungarian dwarf hamsters,Phodopus s. sungorus, were kept in natural photoperiodic conditions throughout the year, either inside at a constantT _a of 23°C or outside subjected to seasonally varyingT _a. Comparisons were made between hamsters from both conditions to evaluate the significance of seasonal changes in photoperiod and/orT _a as environmental cues for seasonal acclimatization inPhodopus. Basal metabolic rate was lowest in July (1.68 ml/g·h) and highest in January (2.06 ml/g·h inPhodopus living outside), combined with a decrease inT _1c from 26°C in July to 20°C in January. This was parallelled by seasonal changes in body weight (summer 42 g, winter 25g), fur colouration, fur depth and the occurrence of short daily torpor.AtT _a below thermoneutrality total energy requirements for thermoregulation in winter acclimatizedPhodopus were found 36% lower than summer values (e.g. at O°CT _a in summer 1,160 mW, in winter 760 mW), which were effected by a combined strategy of reducing body weight (19%) together with improvements of thermal insulation of the body surface (17%). All seasonal changes were similar inPhodopus living inside or outside, suggesting that seasonal changes in photoperiod and not seasonal changes inT _a is the overriding controller for the environmental cueing of seasonality in energy requirements for thermoregulation.This research was supported by the Deutsche Forschungsgemeinschaft (He 990)     

Strategies of thermoregulation and foraging in two vespid wasps,Dolichovespula maculata andVespula vulgaris

Summary D. maculata, the white-faced hornet, stabilized (regulated) thoracic temperature (T _Th) over wide ranges of ambient temperature (T _a), whileV. vulgaris, the common yellowjacket, regulatedT _Th poorly. The hornets also maintained a higherT _Th than the wasps, sometimes heating 38°C aboveT _a. Attacking individuals of both species had higherT _Th than those either leaving or returning to the nest from foraging. The hornets, who are primarily hunters of live prey, showed peak activity near dawn, and they were as active atT _a=2°C as at 20°C. Being able to regulate theirT _Th and fly at the lowT _a should enhance their ability to capture small insects that are usually torpid at theseT _a. The yellowjacket wasps, on the other hand, who are scavengers as well as hunters, did not leave the nest at 2°C; their activity decreased greatly with decreasingT _a. Differences in the foraging technique of the two vespids may be related to their different abilities to thermoregulate.     

Brain temperature regulation of panting and non-panting pigeons exposed to extreme thermal conditions

• 1.1. Brain (hypothalamic), skin and body temperatures were measured in hand-reared acclimated (Acc, n = 5) and non-acclimated (NAcc, n =7) rock pigeons (Columba livia, mean body mass 237 g) exposed to increasing ambient temperatures (T_a) (30–60°C) and low humidities.
• 2.2. In non-panting Acc birds, brain temperature gradually increased from 40.1 ± 0.4°C at 30°C to 41.2 ± 0.4°C at 60°C T_a. A mean body temperature (T_b) of 41.2 ± 0.2°C was measured at T_a up to 50°C; an increase of 1.1°C was observed at 60°C (T_b 42.2 ±0.6°C).
• 3.3. In Acc panting birds exposed for 2 hr to 60°C, T_hy was 41.9 ± 0.8°C and T_s was somewhat (but insignificantly) higher, i.e., 42.2 ± 0.7°C. It looks as if both values were increased as a result of a slight hyperthermia that developed (T_b = 43.5 ± 0.9°C).
• 4.4. The significance of the present results for evaluating neuronal thermoresponsiveness of birds' hypothalamus is discussed.

     

Diurnal body temperature patterns in free‐ranging populations of two southern African arid‐zone nightjars

Endotherms allocate large amounts of energy and water to the regulation of a precise body temperature (T_b), but can potentially reduce thermoregulatory costs by allowing T_b to deviate from normothermic levels. Many data on heterothermy at low air temperatures (T_a) exist for caprimulgids, whereas data on thermoregulation at high T_a are largely absent, despite members of this taxon frequently roosting and nesting in sites exposed to high operative temperatures. We investigated thermoregulation in free‐ranging rufous‐cheeked nightjars Caprimulgus rufigena and freckled nightjars Caprimulgus tristigma in the southern African arid zone. Individuals of both species showed labile T_b fluctuating around a single modal T_b (T_b‐mod). Average T_b‐mod was 39.7°C for rufous‐cheeked nightjars and 39.0°C for freckled nightjars. In both species, diurnal T_b increased with increasing T_a. At T_a ≥ 38°C, rufous‐cheeked nightjar mean T_b increased to 42°C, equivalent to 2.3°C above T_b‐mod. Under similar conditions, freckled nightjar T_b was on average only 1.1°C above T_b‐mod, with a mean T_b of 40.0°C. Freckled nightjars are one of the most heterothermic caprimulgids investigated to date, but our data suggest that during hot conditions this species maintains T_b within a narrow range above T_b‐mod, possibly reflecting an evolutionary tradeoff between decreased thermal sensitivity to lower T_b but increased sensitivity to high T_b. These findings reveal how general thermoregulatory patterns at similar T_a can vary even among closely related species.     

Implications of climate change on the habitat shifts of tropical lizards

The effect of temperature on the distributions of ectothermic vertebrates is well documented. Despite the increase of 6°C expected in the next 60 years in South America, numerous vertebrates are still considered as ‘Least Concern’ species by the IUCN due to their large distribution, insufficient widespread threats and insignificant population decline. One example is the lizard Tropidurus torquatus (Squamata: Tropiduridae), commonly found thermoregulating in anthropic environments throughout the Brazilian Cerrado, but restricted to gallery forests in the equator‐ward localities. The urban areas in this warmer region have been colonised by other closely related congeners (e.g. Tropidurus oreadicus). This study aimed to understand this divergence of habitat selection by these tropirudids that may explain some of the species responses to past and future climate warming. We collected body temperatures (T_b), micro‐environmental temperatures (T_a) and operative (T_e) temperatures in four sites along a latitudinal gradient: a pole‐ward and two central sites where T. torquatus inhabit urban areas and one equator‐ward site where T. torquatus and T. oreadicus occur in the gallery forest and in urban microhabitats, respectively. All three populations of T. torquatus present similar T_b (35.5–36°C) and shared microhabitats with a similar T_a (34–37.3°C). The T_e in the equator‐ward urban site was considerably higher than in the gallery forest. Tropidurus oreadicus T_b was 38.2 °C (30.1–41.3°C) and was active at a T_a of 30.5–42.3°C. The overlap between the genus T_b, T_a and T_e highlights a decrease in the hours of activity that lizards would experience under climate warming. The reduction of hours of activity together with the devastation of natural habitats represents threats and an alarming scenario especially for the equator‐ward populations.     

Dynamics and precision of thermoregulatory responses of eastern skunk cabbage Symplocarpus foetidus

Inflorescences of the arum lily Symplocarpus foetidus are thermogenic and thermoregulatory. The spadix increases respiratory heat production rate as ambient temperature decreases. This study examined the relationships between spadix temperature (T_s), respiration rate () and ambient temperature (T_a) at equilibrium and during transient responses to step changes in T_a. Intact inflorescences inside a miniature constant temperature cabinet in the field showed the most precise temperature regulation yet recorded; over a 37.4 °C range in T_a (?10.3 to 27.1 °C), T_s changed only 3.5 °C (22.7 to 26.2 °C). Regulated temperatures were not related to spadix size (1.9–7.3 g) or circadian cycle. Dynamic responses to step changes in T_a involved a phasic change in T_s, first in the same direction as T_a, then reversing at 38.3 min, and finally approaching equilibrium at 87.6 min, on average. Meanwhile changed in a monotonic curve toward equilibrium. Models revealed that the dynamics of temperature change were inconsistent with simply a physical lag in the system, but involved some form of biochemical regulation, possibly by changes in activity of a rate‐limiting functional protein.     

Evidence of facultative daytime hypothermia in a small passerine wintering at northern latitudes

The use of hypothermia as a means to save energy is well documented in birds. This energy‐saving strategy is widely considered to occur exclusively at night in diurnally active species. However, recent studies suggest that facultative hypothermia may also occur during the day. Here, we document the use of daytime hypothermia in foraging Black‐capped Chickadees Poecile atricapillus wintering in eastern Canada. We measured the body temperature (T_b) of 126 individuals (plus 48 repeated measures) during a single winter and related values to ambient temperature (T_a) at the time of capture. We also tested whether daytime hypothermia was correlated with the size of body reserves (residuals of mass on structural size and fat score) and levels of metabolic performance (basal metabolic rate and maximum thermogenic capacity). We found that T_b of individual birds was lower when captured at low T_a, reaching values as low as 35.5 °C in actively foraging individuals. T_b was unrelated to metabolic performance or measures of body reserves. Therefore, daytime hypothermia does not result from individuals being unable to maintain T_b during cold spells or to a lack of body reserves. Our data also demonstrated a high level of individual variation in the depth of hypothermia, the causes of which remain to be explored.     

Daily torpor and energetics in a tropical mammal, the northern blossom-bat Macroglossus minimus (Megachiroptera)

Little is known about torpor in the tropics or torpor in megachiropteran species. We investigated thermoregulation, energetics and patterns of torpor in the northern blossom-bat Macroglossus minimus (16 g) to test whether physiological variables may explain why its range is limited to tropical regions. Normothermic bats showed a large variation in body temperature (T _b) (33 to 37 °C) over a wide range of ambient temperatures (T _as) and a relatively low basal metabolic rate (1.29 ml O₂ g⁻¹h⁻¹). Bats entered torpor frequently in the laboratory at T _as between 14 and 25 °C. Entry into torpor always occurred when lights were switched on in the morning, independent of T _a. MRs during torpor were reduced to about 20–40% of normothermic bats and T _bs were regulated at a minimum of 23.1 ± 1.4 °C. The duration of torpor bouts increased with decreasing T _a in non-thermoregulating bats, but generally terminated after 8 h in thermoregulating torpid bats. Both the mean minimum T _b and MR of torpid M. minimus were higher than that predicted for a 16-g daily heterotherm and the T _b was also about 5 °C higher than that of the common blossom-bat Syconycteris australis, which has a more subtropical distribution. These observations suggest that variables associated with torpor are affected by T _a and that the restriction to tropical areas in M. minimus to some extent may be due to their ability to enter only very shallow daily torpor. Accepted: 22 September 1997     

Thermoregulation in the cicada Platypedia putnami variety lutea (Homoptera: Tibicinidae) with a test of a crepitation hypothesis

• 1.Measurements of body temperature (T_b) in the field demonstrate that Platypedia putnami var. lutea Davis regulates T_b through behavioral mechanisms.
• 2.Thermal responses (minimum flight temperature 17.3°C, maximum voluntary tolerance-temperature 32.5°C, and heat torpor temperature 44.4°C) of P. putnami var. lutea are related to the altitude of their habitat.
• 3.Water loss rates increase with ambient temperature (T_a). Water loss rates are not significantly different at the extremes of the active T_b range but increase significantly when exposed to elevated T_a.
• 4.Acoustic activity was restricted at 6.7°C T_b range. This is similar to the lower end of the T_b range for singing measured in cicada species that produce sound with a timbal mechanism.
• 5.The use of the wing musculature to produce acoustic signals in P. putnami var. lutea does not increase the T_b range over which the species can call compared to timbal calls produced by other cicada species.

     

Taking the heat: thermoregulation in Asian elephants under different climatic conditions

Some mammals indigenous to desert environments, such as camels, cope with high heat load by tolerating an increase in body temperature (T _b) during the hot day, and by dissipating excess heat during the cooler night hours, i.e., heterothermy. Because diurnal heat storage mechanisms should be favoured by large body size, we investigated whether this response also exists in Asian elephants when exposed to warm environmental conditions of their natural habitat. We compared daily cycles of intestinal T _b of 11 adult Asian elephants living under natural ambient temperatures (T _a) in Thailand (mean T _a ~ 30°C) and in 6 Asian elephants exposed to cooler conditions (mean T _a ~ 21°C) in Germany. Elephants in Thailand had mean daily ranges of T _b oscillations (1.15°C) that were significantly larger than in animals kept in Germany (0.51°C). This was due to both increased maximum T _b during the day and decreased minimum T _b at late night. Elephant’s minimum T _b lowered daily as T _a increased and hence entered the day with a thermal reserve for additional heat storage, very similar to arid-zone ungulates. We conclude that these responses show all characteristics of heterothermy, and that this thermoregulatory strategy is not restricted to desert mammals, but is also employed by Asian elephants.     

Thermoregulation and activity patterns of two syntopic cicadas,Tibicen chiricahua and T. duryi (Homoptera: Cicadidae), in central New Mexico

Summary Tibicen chiricahua and T. duryi are cicada species that are active as adults early each summer in central New Mexico, and are often syntopic in pinyon-juniper woodlands. Both species regulate thoracic temperature (T_th) within fairly narrow limits by utilizing behavioral mechanisms and evaporative cooling. However, syntopic populations of these two species were found to regulate at different T_th despite having synchronous annual and daily activity periods; overall mean T_th of T. chiricahua was 3.1°C higher than it was for T. duryi. Interspecific differences in evaporative cooling abilities and rates of passive heat exchange could not account for this difference in T_th. Part of the difference in T_th resulted from the fact that individuals of the two species were active in thermally distinct microhabitats. Within each species, mean T_th varied among behavior categories, and differences in how the two species allocated their time between activities also contributed to the interspecific difference in T_th. Though T. duryi is restricted to pinyon-juniper habitats such as the one in this study, T. chiricahua is also found in warmer habitats. The difference in T_th in the syntopic populations probably reflects interspecific differences in thermal preferences and thermal optima that are adaptive over their respective habitat ranges. The degree of dependence of T_th on ambient temperature (T_am) varied between activities within both species, with the least dependence exhibited during singing. Singing involves intense activity of tymbal muscles, which apparently can only function effectively over a relatively narrow range of temperature.     

Thermal niche partitioning in the grasshoppers Arphia conspersa and Trimerotropis suffusa from a montane habitat in central Colorado

ABSTRACT.

• 1 Two species of grasshoppers, Arphia conspersa and Trimerotropis suffusa, coexist in a montane habitat in central Colorado.
• 2 Field-recordings of body temperature revealed that A.conspersa has a significantly lower mean body temperature (T_b), sexual display temperature (T_d) and minimum flying temperature (MFT) than T.suffusa.
• 3 A test of the maximum voluntarily tolerated temperature (MVT) showed that T.suffusa has a higher MVT than A.conspersa.
• 4 Thermal niche breadth, as indexed by the difference between MVT and MFT and the range of environmental temperatures over which each species is active, is broader in the eurythermic A.conspersa than in the stenothermic T. suffusa.
• 5 Thermoregulatory ability, as evaluated by regression analysis of T_b on T_a, was shown to be better in T.suffusa than in A.conspersa and in displaying grasshoppers of both species than in non-displaying ones. The significance of these findings with respect to a cost-benefit model of behavioural thermoregulation in ectotherms is discussed.
• 6 Based on these data and observations it was concluded that A.conspersa and T.suffusa occupy different thermal niches and that thermal considerations may be importantly related to habitat preference, daily activity patterns, and consequent ecological separation.

     

Evaporative and excretory water loss during free flight in pigeons

Body water conservation is important in flying birds because the very high metabolic demands and heat dissipation requirements during flight depend on plasma-volume integrity. Wind tunnel experiments and theoretical model predictions show that evaporative water loss (EWL) depends on air temperature (T _a) and water vapor density (ρ_a), but these relationships have not been examined in free-flying birds. The contribution of excretory water loss to the total water loss of a flying bird is thought to be negligible but this assumption is untested. To study the dependence of water losses on environmental conditions in free-flying birds and to quantify the contribution of excretory water loss to total water loss, we estimated evaporative and excretory water losses in 16 trained, free-flying tippler pigeons (Columba livia, 250–340 g). We collected excreta by attaching a light latex, water-impermeable receptacle around each bird's vent. By gravimetry, we measured evaporative and excretory water losses of birds for eight flights at different T _as and compared flying to resting (control) birds for two of these flights. EWL was constant with respect to T _a when less than 15 °C, and increased with increasing T _a above 19 °C, indicating that evaporative cooling was invoked when the heat load increased. EWL increased with increasing ρ_a, possibly due to the strong correlation between ρ_a and T _a. Excretory water loss was independent of ρ_a or T _a and averaged almost 10% of the total water loss. Measurements of EWL made on pigeons during wind tunnel experiments and previous free-flight studies are consistent with our free-flight measurements made at similar T _a s. Accepted: 13 April 1999