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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.

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 (T a  = 12-40 °C). The thorax temperature as well as the energetic turnover showed a great variability. The mean T thorax was ranging from ~ 35.7 °C at 12 °C to nearly 42.5 °C at 40 °C. The energy turnover calculated from CO 2 -release was highest at a T a 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.

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.

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]

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]

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]