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Overwintering insects are facing energetic challenges because of food shortage, low temperature, and desiccation stress. Paper wasps of the genus Polistes overwinter as mated adults (gynes) in hibernacula protecting them from predation, snow, and rain but barely from low environmental temperature. In different climates, they face differing overwintering temperature regimes, and therefore they may differ in their energy use. We investigated how much of energy resources built up until autumn is used during diapause dormancy in natural hibernacula by measuring lipid, glycogen, and free carbohydrate content in autumn and early spring in Polistes dominula from temperate European (Austrian) and warm Mediterranean (Italian) climate and Polistes gallicus from Mediterranean climate. Winter energy consumption amounted to ~ 339 and ~ 310 J per wasp in the Austrian and Italian Polistes dominula populations. The smaller Italian Polistes gallicus consumed ~ 247 J. This amounts to 2.62, 2.35, and 1.79 J per day. Of this, the energy demand was mainly fuelled by lipids (84%, 93%, and 90%, respectively), but glycogen stores contributed also considerably (16%, 6%, and 9%). Free carbohydrates decreased only by 0.7%, 1%, and 0.8%. While fat stores seem still sufficient in spring, the wasps depleted most of their carbohydrates. The energy reserves of 396, 400, and 147 J per wasp remaining in spring in the three populations seem sufficient to fuel rest or simple brood care activities for a whole summer but restrict foraging flights to a few hours (~ 3.5–6 h). Results suggest that energy supply might become challenging in expected future climate scenarios. [ABSTRACT FROM AUTHOR]

During foraging honeybees are always endothermic to stay ready for immediate flight and to promote fast exploitation of resources. This means high energetic costs. Since energy turnover of foragers may vary in a broad range, energetic estimations under field conditions have remained uncertain. We developed an advanced model, combining the benefits of mechanistic and correlative models, which enables estimation of the energy turnover of stationary foragers from measurements of body surface temperature, ambient air temperature and global radiation. A comprehensive dataset of simultaneously measured energy turnover (ranging from 4 to 85 mW) and body surface temperature (thorax surface temperature ranging from 33.3 to 45 °C) allowed the direct verification of model accuracy. The model variants enable estimation of the energy turnover of stationary honeybee foragers with high accuracy both in shade and in sunshine, with SD of residuals = 5.7 mW and R2 = 0.89. Its prediction accuracy is similar throughout the main range of environmental conditions foragers usually experience, covering any combination of ambient air temperature of 14–38 °C and global radiation of 3–1000 W m−2. [ABSTRACT FROM AUTHOR]

Polistes paper wasps are a widespread taxon inhabiting various climates. They build nests in the open without a protective outer layer, which makes them vulnerable to changing temperatures. To better understand the options they have to react to environmental variation and climate change, we here compare the thermoregulatory behavior of Polistes biglumis from cool Alpine climate with Polistes gallicus from warm Mediterranean climate. Behavioral plasticity helps both of them to withstand environmental variation. P. biglumis builds the nests oriented toward east-south-east to gain solar heat of the morning sun. This increases the brood temperature considerably above the ambience, which speeds up brood development. P. gallicus, by contrast, mostly avoids nesting sites with direct insolation, which protects their brood from heat stress on hot days. To keep the brood temperature below 40–42 °C on warm days, the adults of the two species show differential use of their common cooling behaviors. While P. biglumis prefers fanning of cool ambient air onto the nest heated by the sun and additionally cools with water drops, P. gallicus prefers cooling with water drops because fanning of warm ambient air onto a warm nest would not cool it, and restricts fanning to nests heated by the sun. [ABSTRACT FROM AUTHOR]

The worldwide distribution of honeybees and their fast propagation to new areas rests on their ability to keep up optimal 'tropical conditions' in their brood nest both in the cold and in the heat. Honeybee colonies behave like 'superorganisms' where individuals work together to promote reproduction of the colony. Social cooperation has developed strongly in thermal homeostasis, which guarantees a fast and constant development of the brood. We here report on the cooperation of individuals in reaction to environmental variation to achieve thermal constancy of 34–36 °C. The measurement of body temperature together with bee density and in-hive microclimate showed that behaviours for hive heating or cooling are strongly interlaced and differ in their start values. When environmental temperature changes, heat production is adjusted both by regulation of bee density due to migration activity and by the degree of endothermy. Overheating of the brood is prevented by cooling with water droplets and increased fanning, which start already at moderate temperatures where heat production and bee density are still at an increased level. This interlaced change and onset of different thermoregulatory behaviours guarantees a graded adaptation of individual behaviour to stabilise the temperature of the brood. [ABSTRACT FROM AUTHOR]

Simple Summary: Ambient temperature is a main external parameter in the life of ectothermic insects. It affects egg and larval development as well as adults' survival, thriving and propagation, and successful overwintering. We conducted temperature measurements in Central Europe in the habitat and in the microhabitats of Pyrrhocoris apterus, a herbivorous bug species almost ubiquitous in Eurasia, with a high invasive potential (USA, Central America, India and Australia). and set them against freely available climate data commonly used to characterize habitat climate. Our temperature measurements were also compared to the bug species' thermal limits (critical thermal minima and maxima). Ambient temperatures outside the thermal boundaries of P. apterus can and do occur in the habitat. Microhabitat measurement showed that in summer, individuals simply moved from hot areas to cooler ones, and in winter they sought areas with bearable temperatures for hibernation. Temperatures in these particular areas are not (always) represented in large-scale climate tables, leading to possible misinterpretation of the future dispersal behavior, e.g., of invasive species. Ambient temperature is a main parameter that determines the thriving and propagation of ectothermic insects. It affects egg and larval development as well as adults' survival and successful overwintering. Pyrrhocoris apterus is a herbivorous bug species almost ubiquitous in Eurasia. Its distribution extends from the Atlantic Coast to Siberia, Northwest China and Mongolia. After introduction, it established successfully in the USA, Central America, India and Australia, which indicates a high invasive potential of this species. We determined the climatic conditions in Central Europe in a habitat where P. apterus has been continuously observed for decades. We conducted temperature measurements in the habitat and in the microhabitats where individuals could be found during the year and set them against freely available climate data commonly used to characterize habitat climate. Our temperature measurements were also compared to thermal limits (critical thermal minima and maxima). Although ambient temperatures outside the thermal boundaries of P. apterus can and do occur in the habitat, the bugs thrive and propagate. Microhabitat measurement in winter showed that individuals sought areas with favorable temperatures for hibernation. In particular, these areas are not (always) represented in large-scale climate tables, leading to possible misinterpretation of future patterns of spread of invasive species spread. [ABSTRACT FROM AUTHOR]

Vespine wasps ( Vespula sp.) are endowed with a pronounced ability of endothermic heat production. To show how they balance energetics and thermoregulation under variable environmental conditions, we measured the body temperature and respiration of sucrose foragers (1.5 M, unlimited flow) under variable ambient temperature ( T = 20-35 °C) and solar radiation (20-570 W m). Results revealed a graduated balancing of metabolic efforts with thermoregulatory needs. The thoracic temperature in the shade depended on ambient temperature, increasing from ~37 to 39 °C. However, wasps used solar heat gain to regulate their thorax temperature at a rather high level at low T (mean T ~ 39 °C). Only at high T they used solar heat to reduce their metabolic rate remarkably. A high body temperature accelerated the suction speed and shortened foraging time. As the costs of foraging strongly depended on duration, the efficiency could be significantly increased with a high body temperature. Heat gain from solar radiation enabled the wasps to enhance foraging efficiency at high ambient temperature ( T = 30 °C) by up to 63 %. The well-balanced change of economic strategies in response to environmental conditions minimized costs of foraging and optimized energetic efficiency. [ABSTRACT FROM AUTHOR]

Heterothermic insects like honeybees, foraging in a variable environment, face the challenge of keeping their body temperature high to enable immediate flight and to promote fast exploitation of resources. Because of their small size they have to cope with an enormous heat loss and, therefore, high costs of thermoregulation. This calls for energetic optimisation which may be achieved by different strategies. An ‘economizing’ strategy would be to reduce energetic investment whenever possible, for example by using external heat from the sun for thermoregulation. An ‘investment-guided’ strategy, by contrast, would be to invest additional heat production or external heat gain to optimize physiological parameters like body temperature which promise increased energetic returns. Here we show how honeybees balance these strategies in response to changes of their local microclimate. In a novel approach of simultaneous measurement of respiration and body temperature foragers displayed a flexible strategy of thermoregulatory and energetic management. While foraging in shade on an artificial flower they did not save energy with increasing ambient temperature as expected but acted according to an ‘investment-guided’ strategy, keeping the energy turnover at a high level (∼56–69 mW). This increased thorax temperature and speeded up foraging as ambient temperature increased. Solar heat was invested to increase thorax temperature at low ambient temperature (‘investment-guided’ strategy) but to save energy at high temperature (‘economizing’ strategy), leading to energy savings per stay of ∼18–76% in sunshine. This flexible economic strategy minimized costs of foraging, and optimized energetic efficiency in response to broad variation of environmental conditions. [ABSTRACT FROM AUTHOR]

Abstract: Endothermic insects like honeybees and some wasps have to cope with an enormous heat loss during foraging because of their small body size in comparison to endotherms like mammals and birds. The enormous costs of thermoregulation call for optimisation. Honeybees and wasps differ in their critical thermal maximum, which enables the bees to kill the wasps by heat. We here demonstrate the benefits of a combined use of body temperature measurement with infrared thermography, and respiratory measurements of energy turnover (O2 consumption or CO2 production via flow-through respirometry) to answer questions of insect ecophysiological research, and we describe calibrations to receive accurate results. To assess the question of what foraging honeybees optimise, their body temperature was compared with their energy turnover. Honeybees foraging from an artificial flower with unlimited sucrose flow increased body surface temperature and energy turnover with profitability of foraging (sucrose content of the food; 0.5 or 1.5mol/L). Costs of thermoregulation, however, were rather independent of ambient temperature (13–30°C). External heat gain by solar radiation was used to increase body temperature. This optimised foraging energetics by increasing suction speed. In determinations of insect respiratory critical thermal limits, the combined use of respiratory measurements and thermography made possible a more conclusive interpretation of respiratory traces. [Copyright &y& Elsevier]

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 ( Tth) was regulated at a high and rather constant level over a broad range of ambient temperatures ( Tth = 33.7-35.7°C, Ta = 10-27°C). However, at a certain Ta, Tth showed a strong variation, depending on the plants from which the bees were foraging. At warmer conditions ( Ta = 27-32°C) the Tth increased nearly linearly with Ta to a maximal average level of 42.6 °C. The thorax temperature excess decreased strongly with increasing Ta ( Tth− Ta = 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 Tth in the spring than in the summer. An anova revealed that season had the greatest effect on Tth, followed by Ta and radiation. 4. It was presumed the foragers' motivational status to be the main factor responsible for the variation of Tth between seasons and different plants. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

Honeybee larvae and pupae are extremely stenothermic, i.e. they strongly depend on accurate regulation of brood nest temperature for proper development (33-36°C). Here we study the mechanisms of social thermoregulation of honeybee colonies under changing environmental temperatures concerning the contribution of individuals to colony temperature homeostasis. Beside migration activity within the nest, the main active process is ''endothermy on demand'' of adults. An increase of cold stress (cooling of the colony) increases the intensity of heat production with thoracic flight muscles and the number of endothermic individuals, especially in the brood nest. As endothermy means hard work for bees, this eases much burden of nestmates which can stay ectothermic. Concerning the active reaction to cold stress by endothermy, age polyethism is reduced to only two physiologically predetermined task divisions, 0 to ∼2 days and older. Endothermic heat production is the job of bees older than about two days. They are all similarly engaged in active heat production both in intensity and frequency. Their active heat production has an important reinforcement effect on passive heat production of the many ectothermic bees and of the brood. Ectothermy is most frequent in young bees (<∼2 days) both outside and inside of brood nest cells. We suggest young bees visit warm brood nest cells not only to clean them but also to speed up flight muscle development for proper endothermy and foraging later in their life. Young bees inside brood nest cells mostly receive heat from the surrounding cell wall during cold stress, whereas older bees predominantly transfer heat from the thorax to the cell wall. Endothermic bees regulate brood comb temperature more accurately than local air temperature. They apply the heat as close to the brood as possible: workers heating cells from within have a higher probability of endothermy than those on the comb surface. The findings show that thermal homeostasis of honeybee colonies is achieved by a combination of active and passive processes. The differential individual endothermic and behavioral reactions sum up to an integrated action of the honeybee colony as a superorganism. [ABSTRACT FROM AUTHOR]

Abstract: 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. [Copyright &y& Elsevier]

Abstract: The relation between the respiratory activity of resting honeybees and ambient temperature (T a) was investigated in the range of 5–40°C. Bees were kept in a temperature controlled flow through respirometer chamber where their locomotor and endothermic activity, as well as abdominal ventilatory movements was recorded by infrared thermography. Surprisingly, true resting bees were often weakly endothermic (thorax surface up to 2.8°C warmer than abdomen) at a T a of 14–30°C. Above 33°C many bees cooled their body via evaporation from their mouthparts. A novel mathematical model allows description of the relationship of resting (standard) metabolic rate and temperature across the entire functional temperature range of bees. In chill coma (<11°C) bees were ectothermic and CO2 release was mostly continuous. CO2 release rate (nls−1) decreased from 9.3 at 9.7°C to 5.4 at 5°C. At a T a of >11°C CO2 was released discontinuously. In the bees’ active temperature range mean CO2 production rate (nls−1) increased sigmoidally (10.6 at 14.1°C, 24.1 at 26.5°C, and 55.2 at 38.1°C), coming to a halt towards the upper lethal temperature. This was primarily accomplished by an exponential increase in gas exchange frequency (0.54 and 3.1breathsmin−1 at 14.1 and 38.1°C) but not in released CO2 volume per respiratory cycle (1487 and 1083nlcycle−1 at 14.1 and 38.1°C). Emission of CO2 bursts was mostly (98%) accompanied by abdominal ventilation movements even in small CO2 bursts. Larger bursts coincided with a longer duration of active ventilation. An increased amount of CO2 expelled per unit time of ventilation indicates a higher efficiency of ventilation at high ambient temperatures. [Copyright &y& Elsevier]

We report here on the interrelationship of aggressive behaviour and thermoregulation in honeybees. Body temperature measurements were carried out without behavioural disturbance by infrared thermography. Guard bees, foragers, drones, and queens involved in aggressive interactions were always endothermic, i.e. had their flight muscles activated. Guards made differential use of their endothermic capacity. Mean thorax temperature was 34.2–35.1°C during examination of bees but higher during fights with wasps (37°C) or attack of humans (38.6°C). They usually cooled down when examining bees whereas examinees often heated up during prolonged interceptions (maximum >47°C). Guards neither adjusted their thorax temperature (and thus flight muscle function and agility) to that of examined workers, nor to that of drones, which were 2–7°C warmer. Guards examined cool bees (<33°C) longer than warmer ones, supporting the hypothesis that heating of examinees facilitates odour identification by guards, probably because of vapour pressure increase of semiochemicals with temperature. Guards in the core of aggressive balls clinged to the attacked insects to fix them and kill them by heat (maximum 46.5°C). Bees in the outer cluster layers resembled normal guards behaviourally and thermally. They served as active core insulators by heating up to 43.9°C. While balled wasps were cooler (maximum 42.5°C) than clinging guards balled bees behaved like examinees with maximum temperatures of 46.6°C, which further supports the hypothesis that the examinees heat up to facilitate odour identification. [ABSTRACT FROM AUTHOR]

The CO2 production of individual larvae ofApis mellifera carnica, which were incubated within their cells at a natural air humidity of 60-80%, was determined by an open-flow gas analyzer in relation to larval age and ambient temperature. In larvae incubated at 34 °C the amount of CO2 produced appeared to fall only moderately from 3.89±1.57 µl mg-1 h-1 in 0.5-day-old larvae to 2.98±0.57 µl mg-1 h-1 in 3.5-day-old larvae. The decline was steeper up to an age of 5.5 days (0.95±1.15 µl mg-1 h-1). Our measurements show that the respiration and energy turnover of larvae younger than about 80 h is considerably lower (up to 35%) than expected from extrapolations of data determined in older larvae. The temperature dependency of CO2 production was determined in 3.5-day-old larvae, which were incubated at temperatures varying from 18 to 38 °C in steps of 4 °C. The larvae generated 0.48±0.03 µl mg-1 h-1 CO2 at 18 °C, and 3.97±0.50 µl mg-1 h-1 CO2 at 38 °C. The temperature-dependent respiration rate was fitted to a logistic curve. We found that the inflection point of this curve (32.5 °C) is below the normal brood nest temperature (33-36 °C). The averageQ10 was 3.13, which is higher than in freshly emerged resting honeybees but similar to adult bees. This strong temperature dependency enables the bees to speed up brood development by achieving high temperatures. On the other hand, the results suggest that the strong temperature dependency forces the bees to maintain thermal homeostasis of the brood nest to avoid delayed brood development during periods of low temperature. [ABSTRACT FROM AUTHOR]

In order to survive cold northern winters, honeybees crowd tightly together in a winter cluster. Present models of winter cluster thermoregulation consider the insulation by the tightly packed mantle bees as the decisive factor for survival at low temperatures, mostly ignoring the possibility of endothermic heat production. We provide here direct evidence of endothermic heat production by 'shivering' thermogenesis. The abundance of endothermic bees is highest in the core and decreases towards the surface. This shows that core bees play an active role in thermal control of winter clusters. We conclude that regulation of both the insulation by the mantle bees and endothermic heat production by the inner bees is necessary to achieve thermal stability in a winter cluster. [ABSTRACT FROM AUTHOR]

In honeybee colonies, guards protect their nest from various robbers including bees from other colonies. Infrared thermography showed that the guards and the bees examined by them (examinees) differ considerably in their thermal behaviour according to their particular role in the nestmate recognition process. The thorax surface temperature was on average higher and more variable in the examinees (36.1°C, S.D. = 4.14, N = 1545, 303 bees) than in the guards (34.0°C, s.n. = 2.00, N = 1681, 772 bees). During thorough examinations lasting longer than 30s, more than 60% of the examinees showed phases of intense thoracic heating of more than 2°C (maximum temperature 48.5°C), whereas most guards cooled down. Our data suggest that these examinees heat up their surface to enhance chemical signalling during examinations. [ABSTRACT FROM AUTHOR]

The temporal dynamics and spatial distribution of heat production by inflorescences of Arum maculatum L. were investigated by infra-red thermography. Two centres of heat production, the appendix and the male flowers, and three thermogenic phases, two of the male flowers and one of the appendix, could be observed. On the first day of flowering, when the spathe was still firmly closed, the male flowers became thermogenic ( = first thermogenic phase of the inflorescence) and reached surface temperatures of 4.1-8.0 °C above ambient temperatures. Afterwards the spathe unfolded and the appendix started hearing to temperatures of 5.0-14.0 °C above ambient temperature (second thermogenic phase of the inflorescence). On the second day of flowering the male flowers revealed a second temperature maximum of 0.5-6.l °C above ambient temperature (third thermogenic phase of the inflorescence) which was followed by the release of the pollen grains. When the male flowers started heating, a fruit-like and pleasant scent became evident and continued until the heat production in the male flowers stopped. Only during the heating phase of the appendix was thus fruit-like scent overlaid by the dung-like odour typical of many aroid species. The surface of the spadix and the inner surface of the basal bulb (floral chamber) as investigated by scanning electron microscopy were characterized by smooth epidermal cells of papillate shape. These cells were turgid before the onset of flowering and heat production and collapsed afterwards. Distinct intercellular spaces creating a lacunose epidermis ('Lückenepidermis') were observed between the papillate epidermal cells of the inner surface of the basal bulb (floral chamber). This coincided with a high density of stomata on its outer surface. [ABSTRACT FROM AUTHOR]

Simple Summary: The heat-loving paper wasp Polistes dominula builds small nests in sheltered places in quite differing habitats, providing a favorable microclimate for brood development. Their careful choice of the nesting site enables them to keep the nest and brood temperature above the ambient air temperature. In temperate Austrian (Central European) climates, the cooling efforts of the adult wasps keep the mean comb temperature from exceeding approximately 39 °C, which seems to be a threshold for optimal development. Measurement of the nest and the body temperature of the wasps at typical nesting sites revealed a close relationship with the air temperature at the nests but a suboptimal correlation with climate data from a local standard weather station or model-generated macroclimate data. The comparison emphasizes the importance of microclimate measurements if it is to judge the vulnerability of insects to climate change. The paper wasp Polistes dominula is a thermophilic species originating from the Mediterranean climate, but is now widely spread in Europe. They live in quite differing habitats; and as synanthropic species, they have been established in human settlement areas. They build a single small comb at protected places with a favorable microclimate. We measured the temperature of the wasps, the nests and their environment at typical nesting sides in Austria (Europe) in the temperate climate, in order to reveal relationships between nest and body temperature and the habitats' microclimate. The temperatures of the comb and of the wasps' body were in a wide range (~20–37 °C) above the ambient air temperature at the nest. This is an advantage as higher temperatures accelerate the development speed of the brood. However, the mean comb temperature did not exceed approximately 38.6 °C. This was managed by cooling efforts of the adult wasps. The ambient air temperature near the nest (~1–2 cm) was always clearly elevated above the ambient air temperature at a local standard weather station in the habitat. A comparison with climate-model-generated macroclimate data revealed the necessity of measuring microclimate data for a reliable description of the insects' thermal environment. [ABSTRACT FROM AUTHOR]

Simple Summary: During overwintering diapause, the gynes of paper wasps (Polistes sp.) are mainly dormant in sheltered hibernacles, protecting them against predators and adverse weather conditions but hardly against low temperatures. By measuring the temperature inside hibernacles occupied by species from both Mediterranean (Italian; P. dominula, P. gallicus) and temperate (Austrian; P. dominula) climates (mean hibernacle temperatures: 8.5 °C and 3.2 °C, respectively), we were able to calculate the energetic demand of overwintering. The cumulative energetic costs differed between the populations. Costs were lowest for the P. dominula population from the cooler Austrian winter climate and significantly higher in P. dominula and P. gallicus from the warmer Italian climate. The lower costs of the temperate species were a result of the lower winter temperature and physiological acclimation processes. Energetic calculations with an assumed temperature increase of up to 3 °C due to climate change predict a dramatic increase of up to 40% in overwintering costs in all species. Gynes of paper wasps (Polistes sp.) spend the cold season in sheltered hibernacles. These hibernacles protect against predators and adverse weather conditions but offer only limited protection against low temperatures. During overwintering diapause, wasps live on the energy they store. We investigated the hibernacles' microclimate conditions of species from the Mediterranean (Italy, P. dominula, P. gallicus) and temperate (Austria, P. dominula) climates in order to describe the environmental conditions and calculate the energetic demand of overwintering according to standard metabolic rate functions. The temperatures at the hibernacles differed significantly between the Mediterranean and temperate habitats (average in Austria: 3.2 ± 5.71 °C, in Italy: 8.5 ± 5.29 °C). In both habitats, the hibernacle temperatures showed variance, but the mean hibernacle temperature corresponded closely to the meteorological climate data. Cumulative mass-specific energetic costs over the studied period were the lowest for the temperate P. dominula population compared with both Mediterranean species. The lower costs of the temperate species were a result of the lower hibernacle temperature and acclimation to lower environmental temperatures. Model calculations with an increased mean temperature of up to 3 °C due to climate change indicate a dramatic increase of up to 40% in additional costs. [ABSTRACT FROM AUTHOR]

Honeybees need water for different purposes, to maintain the osmotic homeostasis in adults as well as to dilute stored honey and prepare liquid food for the brood. Water is also used for cooling of the hive. Foraging in endothermic insects is energy-intensive and the question arises how much energy bees invest in a resource without any metabolically usable energy. We investigated the energy demand of water collecting bees under natural conditions. The thermoregulation and energetic effort was measured simultaneously in a broad range of experimental ambient temperatures (Ta = 12-40 °C). The thorax temperature as well as the energetic turnover showed a great variability. The mean Tthorax was ranging from ~ 35.7 °C at 12 °C to nearly 42.5 °C at 40 °C. The energy turnover calculated from CO2-release was highest at a Ta of 20 °C with about 60 mW and lowest at 40 °C with about 22 mW per bee. The total costs during collection decreased from 10.4 J at 12 °C to 0.5 J at 40 °C. The energetic effort of the water collectors was comparable with that of 0.5 M sucrose foraging bees. Our investigation strongly supports the hypothesis that the bees’ motivational status determines the energetic performance during foraging. [ABSTRACT FROM AUTHOR]

Sexual displays of acoustically signalling insects are used in the context of mate attraction and mate choice. While energetic investment in sound production can increase the reproductive success of the sender, this entails metabolic costs. Resource allocation to sexually selected, reproductive traits can trade off against allocation to naturally selected traits (e.g. growth, immunity) when individuals' energy budgets are limited. Estimating the magnitude of the costs invested in acoustic signalling is necessary to understand this trade-off and its influence on fitness and life history. To compare the costs associated with acoustic signalling for two ensiferan species, we simultaneously took respiratory measurements to record the rate of CO2 production and used infrared thermography to measure the increase in thorax temperature. Furthermore, to identify what combinations of acoustic parameters were energetically costly for the sender, we recorded the calling songs of 22 different cricket and katydid species for a comparative analysis and measured their thorax temperature while they sang. Acoustic signalling was energetically costly for Mecopoda sp. and Anurogryllus muticus, requiring a 12- and 16-fold increase over resting levels in the CO2 production rate. Moreover, calling increased thorax temperature, on average by 7.6 and 5.8°C, respectively. We found that the song intensity and effective calling rate, not simply the chirp/trill duty cycle or the pulse rate alone, were good predictors for the thorax temperature increase in males. [ABSTRACT FROM AUTHOR]

The metabolism of ectotherms is temperature-dependent, and the exponential nature of the metabolic rate- temperature relationship means that warmer temperatures are disproportionately affecting the energy balance. T SB a sb (standard met.) temperature deviation from microclimate ambient nest temperature was -3.6 °C in I P. dominula i , -2.4 °C in I P. gallicus i , and -3.3 °C in I P. biglumis i ). The energetic calculations were conducted using temperature-dependent metabolic rate models of the three I Polistes i species, I P. dominula i , I P. gallicus i and I P. biglumis i [[17], [29]], and adequate body temperature models [[4], [17]]. The nest ambient air temperatures in the three habitats were clearly elevated above the meteorological standard ambient temperature recorded by the nearest weather stations (temperature difference, I P. dominula i : 3.6 °C, I P. gallicus i : 2.4 °C, I P. biglumis i : 3.3 °C; Figure 2). [Extracted from the article]

The Western honeybees Apis mellifera carnica and A. m. ligustica are closely related subspecies living in neighbouring regions. Metabolism and the upper lethal thermal limits are crucial physiological traits, adapted in the evolutionary process to environment and climate conditions. We investigated whether samples from these two ecotypes differ in these traits. The standard metabolic rate was higher in the A. m. ligustica population only at a high temperature ( T ~ 40 °C; dVCO = 12 nl s; P < 0.05), probably due to a higher body temperature (dT = 1.5 °C; P < 0.01). The critical thermal maximum of activity and respiration was similar (difference activity CT = 0.8 °C, respiratory CT = 1.1 °C). The lethal temperature (LT) revealed higher tolerance and survival rates of the Ligustica bees (Carnica 50.3 °C; Ligustica 51.7 °C; P < 0.02). Results reveal the adaptation of the two subspecies to their historic climate conditions, possibly favouring Ligustica in a warming environment. [ABSTRACT FROM AUTHOR]

Winter in climatic regions with low temperatures is a challenge for overwintering insects. They are exposed to temperature extremes, which directly cause mortality or lead to energy depletion. The winter hibernaculum of paper wasp gynes protects from predators and rain, but only poorly from ambient temperature. In order to detect physiological adaptations to differing climates, we compared the respiratory metabolism of overwintering gynes of two polistine species from the Mediterranean climate in Italy (Polistes dominula and Polistes gallicus), and of one species from the temperate climate in Austria (Polistes dominula). The wasps' CO2 emission was measured with stop‐flow respirometry in a temperature range from 2.5 to 20 °C. The mass‐specific standard (resting) metabolic rate, the main type of metabolism of the dormant insects during overwintering, increased exponentially with ambient temperature but was suppressed in comparison to individuals measured in the summer, which conserves the energy stores. In addition, it was lower in the Mediterranean species (P. dominula and P. gallicus) in comparison to the temperate species (P. dominula), especially at higher temperatures. The active metabolic rate was suppressed to a similar amount. The suppressed metabolism of the overwintering Mediterranean gynes could be an adaptation to the higher winter temperatures to prevent a premature depletion of the energy resources. [ABSTRACT FROM AUTHOR]

We investigated the respiration patterns of wasps (Vespula sp.) in their viable temperature range (2.9–42.4°C) by measuring CO2 production and locomotor and endothermic activity. Wasps showed cycles of an interburst–burst type at low ambient temperatures (T a <5°C) or typical discontinuous gas exchange patterns with closed, flutter and open phases. At high T a of >31°C, CO2 emission became cyclic. With rising T a they enhanced CO2-emission primarily by an exponential increase in respiration frequency, from 2.6mHz at 4.7°C to 74mHz at 39.7°C. In the same range of T a CO2 release per cycle decreased from 38.9 to 26.4μlg−1 cycle−1. A comparison of wasps with other insects showed that they are among the insects with a low respiratory frequency at a given resting metabolic rate (RMR), and a relatively flat increase of respiratory frequency with RMR. CO2 emission was always accompanied by abdominal respiration movements in all open phases and in 71.4% of the flutter phases, often accompanied by body movements. Results suggest that resting wasps gain their highly efficient gas exchange to a considerable extent via the length and type of respiration movements. [ABSTRACT FROM AUTHOR]

Abstract: Vespine wasps are known for their high endothermic capacity. Endothermic activity is directly linked to respiration. However, knowledge on wasp respiration is sparse and almost nothing is known about their resting metabolism. We investigated the yellowjackets’ CO2 production in a flow-through respirometer chamber overnight. Endothermic and behavioral activity was observed by real-time infrared thermography. Most resting wasps were ectothermic or only slightly endothermic (thoracic temperature excess against abdomen <0.6°C). In the investigated temperature range (T a =2.9–42.4°C) mean CO2 production rate of resting wasps increased steeply according to an exponential function, from 5.658μlg−1 min−1 at 8.3°C to 8.504μlg−1 min−1 at 20.2°C, 58.686μlg−1 min−1 at 35.3°C and 102.84μlg−1 min−1 at 40°C. The wasps’ respiratory critical thermal maximum (CTmax), marking the upper edge of their viable temperature range, was 45.3°C. The respiratory CTmax did not differ significantly from the activity CTmax of 44.9°C. CTmax values were considerably below that of honeybees (48.9 and 49.0°C for respiration and activity, respectively). This allows honeybees to kill wasps by heat-balling. Comparison with other arthropods showed that vespine wasps are among the insects with the highest mass-specific resting metabolic rate and the steepest increase of metabolism with ambient temperature. [Copyright &y& Elsevier]

The pupal parasitoid Pimpla turionellae (L.) uses self-produced vibrations transmitted on the plant substrate, so-called vibrational sounding, to locate immobile concealed pupal hosts. The wasps are able to use vibrational sounding reliably over a broad range of ambient temperatures and even show an increased signal frequency and intensity at low temperatures. The present study investigates how control of body temperature in the wasps by endothermic mechanisms may facilitate host location under changing thermal environments. Insect body temperature is measured with real-time IR thermography on plant-stem models at temperature treatments of 10, 18, 26 and 30 °C, whereas behaviour is recorded with respect to vibrational host location. The results reveal a low-level endothermy that likely interferes with vibrational sound production because it occurs only in nonsearching females. At the lowest temperature of 10 °C, the thoracic temperature is 1.15 °C warmer than the ambient surface temperature whereas, at the high temperatures of 26 and 30 ° C, the wasps cool down their thorax by 0.29 and 0.47 °C, respectively, and their head by 0.45 and 0.61 °C below ambient surface temperature. By contrast, regardless of ambient temperature, searching females always have a slightly elevated body temperature of at most 0.30 °C above the ambient surface temperature. Behavioural observations indicate that searching females interrupt host location more frequently at suboptimal temperatures, presumably due to the requirements of thermoregulation. It is assumed that both mechanisms, producing vibrations for host location and low-level endothermy, are located in the thorax. Endothermy by thoracic muscle work probably disturbs signal structure of vibrational sounding, so the processes cannot be used at the same time. [ABSTRACT FROM AUTHOR]

European Polistine wasps inhabit mainly temperate and warm climate regions. However, the paper wasp Polistes biglumis represents an exception; it resides in mountainous areas, e.g., in the Alps and in the Apennines. In these habitats, the wasps are exposed to a broad temperature range during their lifetime. We investigated whether they developed adaptations in their metabolism to their special climate conditions by measuring their CO2 production. The standard or resting metabolic rate and the metabolism of active wasps was measured in the temperature range which they are exposed to in their habitat in summer. The standard metabolic rate increased in a typical exponential progression with ambient temperature, like in other wasps. The active metabolism also increased with temperature, but not in a simple exponential course. Some exceptionally high values were presumed to originate from endothermy. The simultaneous measurement of body temperature and metabolic rate revealed a strong correlation between these two parameters. The comparison of the standard metabolic rate of Polistes biglumis with that of Polistes dominula revealed a significantly lower metabolism of the alpine wasps. This energy saving metabolic strategy could be an adaptation to the harsh climate conditions, which restricts foraging flights and energy recruitment. [ABSTRACT FROM AUTHOR]

Polistine wasps collect nectar for their energetic demand and for the provision of the brood. They are mainly ectothermic during different behavioral tasks. We investigated the body temperature of two species living in differing habitats and climate regions, in order to reveal the environmental influence on their thermoregulatory behavior. The species were Polistes dominula in the temperate climate of Central Europe, and Polistes gallicus in the warm Mediterranean climate of Southern Europe. The wasp's body temperature was measured during foraging on lovage (Levisticum officinale) and fennel (Foeniculum vulgare) by infrared thermography in the entire ambient temperature range they are usually exposed to (Ta ~ 20–40 °C). The temperature of all body parts increased nearly linearly with ambient temperature, with the thorax as the warmest part. To achieve optimal foraging temperatures, they preferably use solar radiation. An "operative temperature model" enabled the evaluation of the endothermic effort. Polistes dominula foraging on lovage exhibited no endothermic activity. However, while foraging on fennel they had a weak and almost constant endothermic performance of about 1 °C. Polistes gallicus, by contrast, exhibited mostly no or only minor endothermy during foraging. Both wasps avoid a high energetic effort and this way reduce their foraging costs. [ABSTRACT FROM AUTHOR]

The two paper wasps, Polistes dominula and Polistes gallicus, are related species with strongly differing distribution ranges. We investigated thermal tolerance traits (critical thermal limits and metabolic response to temperature) to gain knowledge about physiological adaptations to their local climate conditions and to get evidence for the reasons of P. dominula's successful dispersion. Body and ambient temperature measurements at the nests revealed behavioural adaptations to microclimate. The species differed clearly in critical thermal minimum ( P. dominula −1.4 °C, P. gallicus −0.4 °C), but not significantly in critical thermal maximum of activity ( P. dominula 47.1 °C, P. gallicus 47.6 °C). The metabolic response did not reveal clear adaptations to climate conditions. At low and high temperatures, the metabolic rate of P. dominula was higher, and at intermediate temperatures, we determined higher values in P. gallicus. However, the species exhibited remarkably differing thermoregulatory behaviour at the nest. On average, P. gallicus tolerated a thoracic temperature up to ~41 °C, whereas P. dominula already tried at ~37 °C to keep the thorax below ambient temperature. We suggest this to be an adaptation to the higher mean ambient temperature we measured at the nest during a breeding season. Although we determined for P. dominula a 0.5 °C larger thermal tolerance range, we do not presume this parameter to be solely responsible for the successful distribution of P. dominula. Additional factors, such as the thermal tolerance of the queens could limit the overwintering success of P. gallicus in a harsher climate. [ABSTRACT FROM AUTHOR]

The European paper wasp, Polistes dominulus Christ, is an abundant wasp species in South and Central Europe which dispersed to the north in recent times. Polistes dominulus exhibits an energy-extensive mode of life, spending much time resting at the nest, which should be reflected in adaptations regarding gas exchange and standard metabolism. We analysed the resting metabolism (CO emission) of Polistes dominulus workers in the ambient temperature range an individual may be exposed to during a breeding season ( T = 2.4-40.6 °C) via flow through respirometry. Behaviour and endothermic activity were assessed by infrared thermography. With rising T, CO release followed an exponential increase from 27 to 149 and 802 nl g min at T = 3, 20 and 35 °C, respectively. Measurements of the thermal regime at the nest showed that resting P. dominulus are most of the time in the lower range of their standard metabolic curve. A comparison with a 'highly energetic' wasp like Vespula sp. revealed that Polistes dominulus not only optimises behaviour but also reduces metabolism to save energy. The CO emission patterns changed with ambient temperature, from discontinuous (≤25 °C) to cyclic (25-36 °C) and continuous gas exchange at higher temperatures. A pronounced break appeared in the data progression regarding cycle frequency and CO emission per gas exchange cycle between 15 and 10 °C. This striking change in gas exchange features indicates a physiological adaptation to special respiratory requirements at low temperatures. [ABSTRACT FROM AUTHOR]

Climate change is expected to produce shifts in species distributions as well as behavioural and physiological adaptations to find suitable conditions or to cope with the altered environment. The paper wasps Polistes dominula and Polistes gallicus are closely related species, native in the European Mediterranean region and North Africa. P. dominula has expanded its range to the relatively cooler climates of Northern and Eastern Europe, but P. gallicus remained in its original distribution area. In order to reveal their metabolic adaptation to the current climate conditions, and the impact on energy demand at future climate conditions, we investigated the respiratory metabolic rate (CO 2 production) of P. dominula from Austria and P. gallicus from Italy. In contrast to the metabolic cold adaptation hypothesis their standard metabolic rate was nearly the same and increased in a typical exponential course with increasing ambient temperature. The metabolic rate of active wasps was higher than the standard metabolic rate and increased with the wasps' activity. There was no obvious difference in the active metabolism between the two species, with the exception that some P. gallicus individuals showed some extraordinary high values. A simultaneous measurement of metabolic rate and body temperature revealed that increased CO 2 production was accompanied by endothermic activity. The two investigated populations of paper wasps are quite similar in their metabolic response to temperature, although they live in different climate regions. The spread of P. dominula into cooler regions did not have significant influence on their active and standard metabolic rate. Unlabelled Image • A comparison of the standard and active metabolic rate of Polistes dominula and P. gallicus from different climates revealed similarities. • The metabolic rate was nearly the same in both species. • Simultaneous measurement of metabolism and body temperature revealed that increased CO 2 production is accompanied by endothermic activity. [ABSTRACT FROM AUTHOR]

Heteroptera, or true bugs populate many climate zones, coping with different environmental conditions. The aim of this study was the evaluation of their thermal limits and derived traits, as well as climatological parameters which might influence their distribution. We assessed the thermal limits (critical thermal maxima, CTmax, and minima, CTmin) of eight seed bug species (Lygaeidae, Pyrrhocoridae) distributed over four Köppen–Geiger climate classification types (KCC), approximately 6° of latitude, and four European countries (Austria, Italy, Croatia, Bulgaria). In test tubes, a temperature ramp was driven down to −5 °C for CTmin and up to 50 °C for CTmax (0.25 °C/min) until the bugs' voluntary, coordinated movement stopped. In contrast to CTmin, CTmax depended significantly on KCC, species, and body mass. CTmax showed high correlation with bioclimatic parameters such as annual mean temperature and mean maximum temperature of warmest month (BIO5), as well as three parameters representing temperature variability. CTmin correlated with mean annual temperature, mean minimum temperature of coldest month (BIO6), and two parameters representing variability. Although the derived trait cold tolerance (TC = BIO6 − CTmin) depended on several bioclimatic variables, heat tolerance (TH = CTmax − BIO5) showed no correlation. Seed bugs seem to have potential for further range shifts in the face of global warming. [ABSTRACT FROM AUTHOR]