Micro-scale differences in thermal habitat quality and a possible case of evolutionary flexibility in the thermal physiology of lacertid lizards

We studied the thermal ecology of the lacertid lizards Lacerta oxycephala and Podarcis melisellensis on the Adriatic island of Vis (Croatia) during summer. These species obviously differ in microhabitat use: L. oxycephala climbs on rocks and stone walls, whereas P. melisellensis is mainly ground-dwelling in vegetation. Since theoretical considerations predict a difference in thermal quality between the species' microhabitats, this system seems to present a good opportunity to test the influence of thermal microhabitat quality on body temperature, thermoregulatory behaviour and evolution of thermal characteristics. Data on thermoregulatory behaviour, body temperatures (T_b) and habitat quality were collected in the field and selected temperatures (T_sel) were estimated in a laboratory thermogradient. Accuracy and effectiveness of thermoregulation were quantified. Thermoregulatory behaviour consisted of timing of activity, selection of places in full sun and near sun-shade transitions, and basking. As predicted, L. oxycephala occupied the microhabitat with the lower thermal quality and had on average a lower T_b. However, L. oxycephala also selected lower temperatures in the experimental thermogradient. Thus, if T_sel can be regarded as the thermoregulatory target, both species proved to be accurate and effective thermoregulators. These results corroborate the "labile view" on the evolution of thermal physiology: both L. oxcephala and P. melisellensis appear to be adapted to their respective thermal microhabitat. This is a surprising conclusion, since earlier studies have found the thermal characteristics of Lacertidae to be evolutionarily rigid.     

Diurnal and Seasonal Changes in Thermal Preference of Single,Isolated Bees and Small Groups of Bees (Apis mellifera L.)

The aim of the present study was: to compare thermoregulatory behaviour of single honeybee workers and groups of 3–15 bees over their annual activity period and to check out whether the annual fluctuations of ambient temperature selection are correlated with phases of the colony development. Thermal behaviour of both single workers and groups of bees was recorded, using a video camera, in a thermal gradient system. Thermal preferences of the insects were tested seasonally in spring (May/June), summer (July/August) and autumn (September–November). Both single bees and small groups of bees changed their thermal behaviour in daily cycle. The season of the year had distinct effect on temperature preferences of both single honeybee workers or small groups of bees. In single honeybee workers the lowest ambient temperatures were preferred in late spring (the swarming phase) while the highest temperatures were selected during the summer (the colony growing phase). There were significant seasonal changes in ambient temperature selected by groups of honeybee workers. Groups of honeybee workers tended to prefer the lowest temperatures in late spring and the highest temperatures were selected during the summer. The day-night differences exhibited by small groups of bees in our experiments are likely to represent behavioural responses of the honeybee colony. In our experiments we proved an influence of the season of the year on the honeybees’ thermal behaviour, which might be connected with seasonal shifts of temperature regulated by the honeybee colony.     

Thermoregulatory behaviour explains countergradient variation in the upper thermal limit of a rainforest skink

The vulnerability of a terrestrial ectotherm to high environmental temperatures depends on the animal's thermal physiology and thermoregulatory behaviour. These variables – environment, physiology, and behaviour – interact with each other, complicating assessment of species vulnerability to global warming. We previously uncovered a counterintuitive pattern in rainforest sunskinks Lampropholis coggeri: a negative relationship between their critical thermal maximum (CT_max) and the temperature of their environment. Could this result be explained by a three‐way interaction between environment, physiology, and behaviour? Here we find that sunskink thermal preference is correlated positively with CT_max, but, importantly, skinks from hotter environments prefer lower temperatures than conspecifics from cooler environments. In an acclimation experiment, we find that CT_max is plastic and shifts in alignment with acclimation temperature. We also found heritable variation in this trait in a common garden study, but this variation was small relative to the plastic shifts observed in CT_max. Thus, our previous observation of a negative correlation between field CT_max and temperature is explained, at least in part, by the lizard's thermoregulatory behaviour: lizards from hot environments preferentially choose cool microenvironments, and their physiology acclimates to these cooler experienced temperatures. Our results suggest that behavioural adjustments to the environment can produce countergradient variation in physiological traits. More broadly, our work underscores the importance of interactions between environment, behaviour, and physiology in ectotherms. Understanding these interactions will be crucial in assessing vulnerability to climate change.     

Speed over efficiency: locusts select body temperatures that favour growth rate over efficient nutrient utilization

Ectotherms have evolved preferences for particular body temperatures, but the nutritional and life-history consequences of such temperature preferences are not well understood. We measured thermal preferences in Locusta migratoria (migratory locusts) and used a multi-factorial experimental design to investigate relationships between growth/development and macronutrient utilization (conversion of ingesta to body mass) as a function of temperature. A range of macronutrient intake values for insects at 26, 32 and 38°C was achieved by offering individuals high-protein diets, high-carbohydrate diets or a choice between both. Locusts placed in a thermal gradient selected temperatures near 38°C, maximizing rates of weight gain; however, this enhanced growth rate came at the cost of poor protein and carbohydrate utilization. Protein and carbohydrate were equally digested across temperature treatments, but once digested both macronutrients were converted to growth most efficiently at the intermediate temperature (32°C). Body temperature preference thus yielded maximal growth rates at the expense of efficient nutrient utilization.     

The effect of carbon dioxide enrichment on apparent stem respiration from <Emphasis Type="Italic">Pinus taeda</Emphasis> L. is confounded by high levels of soil carbon dioxide

Huey and Slatkin’s (Q Rev Biol 51:363–384, 1976) cost–benefit model of lizard thermoregulation predicts variation in thermoregulatory strategies (from active thermoregulation to thermoconformity) with respect to the costs and benefits of the thermoregulatory behaviour and the thermal quality of the environment. Although this framework has been widely employed in correlative field studies, experimental tests aiming to evaluate the model are scarce. We conducted laboratory experiments to see whether the common lizard Zootoca vivipara, an active and effective thermoregulator in the field, can alter its thermoregulatory behaviour in response to differences in perceived predation risk and food supply in a constant thermal environment. Predation risk and food supply were represented by chemical cues of a sympatric snake predator and the lizards’ food in the laboratory, respectively. We also compared males and postpartum females, which have different preferred or “target” body temperatures. Both sexes thermoregulated actively in all treatments. We detected sex-specific differences in the way lizards adjusted their accuracy of thermoregulation to the treatments: males were less accurate in the predation treatment, while no such effects were detected in females. Neither sex reacted to the food treatment. With regard to the two main types of thermoregulatory behaviour (activity and microhabitat selection), the treatments had no significant effects. However, postpartum females were more active than males in all treatments. Our results further stress that increasing physiological performance by active thermoregulation has high priority in lizard behaviour, but also shows that lizards can indeed shift their accuracy of thermoregulation in response to costs with possible immediate negative fitness effects (i.e. predation-caused mortality).     

Correlates of active body temperatures and microhabitat occupation in nine species of central Australian agamid lizards

Body temperatures of active lizards and their correspondence with microhabitat occupation were studied for nine species of agamid lizards in the central Australian arid zone. Thermoregulatory behaviour was also documented using several measures, such as the use of shade and perch height. The effects of thermal environment on lizard habitat occupation were hypothesized to be significant, because desert regions experience daily and seasonal extremes of temperature that are well in excess of a lizard’s preferred temperature range. All species, except Ctenophorus isolepis and Diporiphora winneckei, were found to have body temperatures that corresponded closely to ground and surface temperatures. Thermoregulatory behaviour was also found to be important throughout a lizard’s daily activity; all study species, other than Ctenophorus isolepis, were found to increase their perch height in the middle of the day. Ctenophorus isolepis was shown to be a strictly terrestrial species that uses the shade of spinifex in its thermoregulatory behaviour. Species exhibited a non‐random selection of microhabitats and a preference for a particular set of thermal and structural factors. In this study, it was shown that structural factors were particularly important in microhabitat occupation. Thermal factors accounted for a smaller proportion of variance in microhabitat occupation, but still played a considerable role in the microhabitat use in central Australian agamids.     

Thermal biology and locomotor performance of the Andean lizard Liolaemus fitzgeraldi (Liolaemidae) in Argentina

Ectotherms thermoregulate to maintain their body temperature within the optimal range needed for performing vital functions. The effect of climate change on lizards has been studied as regards the sensitivity of locomotor performance to environmental temperatures. We studied thermoregulatory efficiency and locomotor performance for Liolaemus fitzgeraldi in the Central Andes of Argentina. We determined body temperature, micro-environmental temperatures and operative temperatures in the field. In the laboratory, we measured preferred temperatures and calculated the index of thermoregulatory efficiency. We estimated the thermal sensitivity of locomotion by measuring sprint speed (initial velocity and long sprint) and endurance at five different body temperatures. Body temperature was not associated with either micro-environmental temperature, nor did it show differences with preferred temperatures. Thermoregulatory efficiency was moderate (0.61). Initial velocity and long sprint trials showed differences at different temperatures; however, endurance did not. Moreover, the optimal temperatures for the performance trials showed no significant differences among themselves. We conclude that Liolaemus fitzgeraldi has thermal sensitivity in locomotor performance with respect to body temperature and that it is an eurythermic lizard that experiences a large variation in body temperature and that has thermal flexibility in the cold.     

Body Temperature and Thermoregulatory Behaviour of the Lizard Ameiva ameiva in Central Amazonian Forests

Body temperatures and thermoregulatory behaviour of the teiid lizard Ameiva ameiva inhabiting the edge and the understory were studied in Central Amazonian forests. Despite of differences in the thermal profile of the habitats, the mean body temperature was the same for active lizards observed at the edge or inside the forest, where only slight peculiarities in thermoregulatory behaviour were observed. A. ameiva is capable of maintaining body temperature significantly above microhabitat temperature.     

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.     

Thermoregulatory abilities of Alaskan bees: effects of size, phylogeny and ecology

1. The thermoregulatory capabilities of 18 species of Alaskan bees spanning nearly two orders of magnitude of body mass were measured. Thoracic temperature, measured across the temperature range at which each species forages, was regressed against operative (environmental) temperature to determine bees' abilities to maintain relatively constant thoracic temperatures across a range of operative temperatures (thermoregulatory performance).
2. Previous studies on insect thermoregulation have compared thoracic temperature with ambient air temperature. Operative temperature, which integrates air temperature, solar radiation and effects of wind, was estimated by measuring the temperature of a fresh, dead bee in the field environment. It is suggested that this is a more accurate measure of the thermal environment experienced by the insect and also allows direct comparisons of insects under different microclimate conditions, such as in sun and shade.
3. Simple regression analysis of species and family means, and analysis of phylogenetically based independent contrasts showed thermoregulatory capability, ability to elevate thoracic temperature, and minimum thoracic temperature necessary for initiating flight all increased with body size.
4. Bumble-bees were better thermoregulators than solitary bees primarily as a consequence of their larger body size. However, their thermoregulatory abilities were slightly, but significantly, better than predicted from body size alone, suggesting an added role of pelage and/or physiology. Large solitary bees were better thermoregulators than small solitary bees apparently as a result of body-size differences, with small bees acting as thermal conformers.     

Direct effects of incubation temperature on morphology,thermoregulatory behaviour and locomotor performance in jacky dragons (Amphibolurus muricatus)

Incubation temperature is one of the most studied factors driving phenotypic plasticity in oviparous reptiles. We examined how incubation temperature influenced hatchling morphology, thermal preference and temperature-dependent running speed in the small Australian agamid lizard Amphibolurus muricatus. Hatchlings incubated at 32 °C grew more slowly than those incubated at 25 and 28 °C during their first month after hatching, and tended to be smaller at one month. These differences were no longer significant by three months of age due to selective mortality of the smallest hatchlings. The cooler incubation treatments (25 °C and 28 °C) produced lizards that had deeper and wider heads. Hatchlings from 28 °C had cooler and more stable temperature preferences, and also had lower body temperatures during a 2-h thermoregulatory behaviour trial. Locomotor performance was enhanced at higher body temperatures, but incubation temperature had no measurable effect either independently or in interaction with body temperature. Our study demonstrates that incubation temperature has direct effects on morphology and thermoregulatory behaviour that appears to be independent of any size-dependent effects. We postulate a mechanistic link between these two effects.     

Mechanisms of heat inactivation in Salmonella serotype Typhimurium as affected by low water activity at different temperatures

AIMS: To determine the effect of reduced water activity (a(w)) on thermal inactivation of Salmonella serotype Typhimurium at different temperatures and its mechanism. METHODS AND RESULTS: D-value determinations at a range of different temperatures showed that heating at reduced a(w) (0.94, produced by addition of glucose or sodium chloride to nutrient broth) was protective at temperatures above 53-55 degrees C but sensitizing below this temperature. Using selective enumeration media to determine injury, it was shown that at lower heating temperatures cells survived at high a(w) with cytoplasmic injury whereas at low a(w) these cells were killed. At higher temperatures ribosome degradation was a more important cause of death and was inhibited by low a(w) heating media thereby providing greater heat resistance. CONCLUSIONS: The observed change in behaviour reflects the different reactions responsible for thermal death at different temperatures and their different response to reduced a(w). SIGNIFICANCE AND IMPACT OF THE STUDY: This work qualifies the previous assumption that reduced a(w) is protective and suggests that the efficacy of low temperature pasteurization regimes may be increased by reduced a(w).     

绿僵菌侵染后东亚飞蝗营养生理参数变化研究

本文采用饵剂饲喂方法研究东亚飞蝗Locusta migratoria manilensis(Meyen)营养生理参数变化,结果表明:绿僵菌(Metarhizium anisopliae)侵染后,5个温度梯度下蝻期总取食量分别为96.4、108.3、131.9、103.2、249.5mg;21、24、27、30℃4个温度...     

Compensatory dietary selection occurs in larval Locusta migratoria but not Spodoptera littoralis after a single deficient meal during ad libitum feeding

ABSTRACT Final instar nymphs of Locusta migratoria (L.) and larvae of Spodoptera littoralis (Boisduval) were given an artificial diet deficient in either protein or digestible carbohydrate for a single meal during ad libitum feeding, after which they were provided with a choice of two diets, one containing protein but no digestible carbohydrate and the other containing carbohydrate but no protein. Detailed analyses of feeding behaviour showed that locusts exhibited a degree of compensatory dietary selection following the single deficient meal. No such response was evident for the caterpillars, although previous work (Simpson et al. , 1988) has demonstrated that compensatory selection behaviour is marked in this species after periods of 4 h or more on the same deficient diets. These results show that locusts are able to respond extremely rapidly to the nutritional quality of their food by utilizing nutritional feedbacks. This capability may have important implications for the study of foraging strategies in other herbivorous insects.     

Calling behaviour under climate change: geographical and seasonal variation of calling temperatures in ectotherms

Calling behaviour is strongly temperature‐dependent and critical for sexual selection and reproduction in a variety of ectothermic taxa, including anuran amphibians, which are the most globally threatened vertebrates. However, few studies have explored how species respond to distinct thermal environments at time of displaying calling behaviour, and thus it is still unknown whether ongoing climate change might compromise the performance of calling activity in ectotherms. Here, we used new audio‐trapping techniques (automated sound recording and detection systems) between 2006 and 2009 to examine annual calling temperatures of five temperate anurans and their patterns of geographical and seasonal variation at the thermal extremes of species ranges, providing insights into the thermal breadths of calling activity of species, and the mechanisms that enable ectotherms to adjust to changing thermal environments. All species showed wide thermal breadths during calling behaviour (above 15 °C) and increases in calling temperatures in extremely warm populations and seasons. Thereby, calling temperatures differed both geographically and seasonally, both in terrestrial and aquatic species, and were 8–22 °C below the specific upper critical thermal limits (CT_max) and strongly associated with the potential temperatures of each thermal environment (operative temperatures during the potential period of breeding). This suggests that calling behaviour in ectotherms may take place at population‐specific thermal ranges, diverging when species are subjected to distinct thermal environments, and might imply plasticity of thermal adjustment mechanisms (seasonal and developmental acclimation) that supply species with means of coping with climate change. Furthermore, the thermal thresholds of calling at the onset of the breeding season were dissimilar between conspecific populations, suggesting that other factors besides temperature are needed to trigger the onset of reproduction. Our findings imply that global warming would not directly inhibit calling behaviour in the study species, although might affect other temperature‐dependent features of their acoustic communication system.     

Mitigating thermal effect of behaviour and microhabitat on the intertidal snail Littorina saxatilis (Olivi) over summer

High shore intertidal ectotherms must withstand temperatures which are already close, at or beyond their upper physiological thermal tolerance. Their behaviour can provide a relief under heat stress, and increase their survival through thermoregulation. Here, we used infrared imaging to reveal the thermoregulatory behavioural strategies used by the snail Littorina saxatilis (Olivi) on different microhabitats of a high shore boulder field in Finistère (western France) in summer. On our study site, substrate temperature is frequently greater than L. saxatilis upper physiological thermal limits, especially on sun exposed microhabitats. To maintain body temperatures within their thermal tolerance window, withdrawn snails adopted a flat posture, or elevated their shells and kept appended to the rock on the outer lip of their aperture with dried mucous (standing posture). These thermal regulatory behaviours lowered snail body temperatures on average by 1–2 °C. Aggregation behaviour had no thermoregulatory effect on L. saxatilis in the present study. The occupation of biogenic microhabitats (barnacles) was associated with a 1 °C decrease in body temperatures. Barnacles and microhabitats that experienced low sun exposure, low thermal fluctuations and low thermal maxima, could buffer the heat extremes encountered at high shore level especially on sun exposed microhabitats.     

Defence tactics cycle with diel microhabitat choice and body temperature in the desert locust,Schistocerca gregaria

We examined environmental factors influencing plasticity in antipredator defences of adult gregarious desert locusts, Schistocerca gregaria, including daily cycles in temperature, light, microhabitat occupied and predator threat. In the Sahara Desert in Mauritania, West Africa, daily temperature fluctuated widely from below locusts’ cold thermal limits for jump‐ and flight‐defence, to above their preferred body temperature. Locusts changed microhabitats throughout the 24‐hr period in synchrony with the daily thermo‐photocycle. They roosted in tall trees and large bushes at night, moved to the ground in the morning, shaded under or in small bushes and annuals at midday, moved back to the ground in the afternoon and then returned to night roosting sites around dusk. Locust antipredator defences varied throughout the 24‐hr period, and these changes were correlated with temperature, photocycle and habitat. Flight escape was associated with daytime, high temperatures and the ground habitat. Dropping escape (= releasing hold of vegetation and dropping to the ground or into vegetation) was associated with cool temperatures and low‐to‐medium sized bushes. Stationary behaviour was associated with the tree microhabitat and height off the ground. Roosting (a primary defence) was associated with cool temperatures at night and early morning, tree habitats and nocturnal ground‐foraging times of endothermic mammals. In summary, we propose that temperature is the key factor in determining both changing microhabitat choice and changing antipredator defence, due to thermal constraints on locust muscle for this ectothermic insect. The thermocycle also influences temporal predator loads, which influence the evolution of locust diel defence strategies. These various environmental factors not only influence one another, they also interact to influence antipredator defence expression in locusts. Overall, our study suggests that plasticity in locust antipredator defences is a complicated matter mediated by the interactions of multiple environmental factors and physiological and ecological constraints and trade‐offs.     

Experimental Manipulation of Melanism Demonstrates the Plasticity of Preferred Temperature in an Agricultural Pest (Phaulacridium vittatum)

Phenotypic plasticity is a key trait of successful pest species, and may increase the ability to cope with higher, more variable temperatures under climate change. We investigate the plasticity of preferred temperature in a widespread agricultural pest, the wingless grasshopper (Phaulacridium vittatum). Preferred temperature is a measure of thermoregulatory behaviour through habitat selection. It is influenced by melanism, which affects body temperature by determining the amount of radiation absorbed by the body. First we demonstrate that body temperature and preferred temperature in P. vittatum is influenced by melanism, by comparing the preferred temperature of the colour morphs in laboratory thermal gradients and field body temperatures in natural populations. We then test whether preferred temperature changes in response to changes in body temperature, by determining preferred temperature before and after manipulation of melanism by painting. When melanism was manipulated experimentally in live grasshoppers, preferred temperature changed to reflect the thermal qualities of the new colour. The preferred temperature of light grasshoppers increased after they were painted black, and decreased after being painted white. Similarly, dark individuals that were painted white behaved like a light individual, maintaining a lower body temperature. Preferred temperature in P.vittatum is a plastic thermoregulatory response to ambient temperature, mediated by the influence of melanism on body temperature.     

Too cold is better than too hot: Preferred temperatures and basking behaviour in a tropical freshwater turtle

Thermoregulation is critical to the survival of animals. Tropical environments can be particularly thermally challenging as they reach very high, even lethal, temperatures. The thermoregulatory responses of tropical freshwater turtles to these challenges are poorly known. One common thermoregulatory behaviour is diurnal basking, which, for many species, facilitates heat gain. Recently, however, a north-eastern Australian population of Krefft's river turtles (Emydura macquarii krefftii) has been observed basking nocturnally, possibly to allow cooling. To test this, we determined the thermal preference (central 50% of temperatures selected) of E. m. krefftii in an aquatic thermal gradient in the laboratory. We then conducted a manipulative experiment to test the effects of water temperatures, both lower and higher than preferred temperature, on diurnal and nocturnal basking. The preferred temperature range fell between 25.3°C (±SD: 1.5) and 27.6°C (±1.4) during the day, and 25.3°C (±2.4) and 26.8°C (±2.5) at night. Based on this, we exposed turtles to three 24 h water temperature treatments (‘cool’ [23°C], ‘preferred’ [26°C] and ‘warm’ [29°C]) while air temperature remained constant at 26°C. Turtles basked more frequently and for longer periods during both the day and night when water temperatures were above their preferred range (the ‘warm’ treatment). This population frequently encounters aquatic temperatures above the preferred thermal range, and our results support the hypothesis that nocturnal basking is a mechanism for escaping unfavourably warm water. Targeted field studies would be a valuable next step in understanding the seasonal scope of this behaviour in a natural environment.     

Physiological and behavioural responses to seasonal changes in environmental temperature in the Australian spiny crayfish Euastacus sulcatus

The strategies used by ectotherms to minimise the detrimental effects of suboptimal thermal environments on physiological performance are often related to whether they inhabit a terrestrial or aquatic environment. Most terrestrial ectotherms use thermoregulatory strategies to maintain body temperature within an optimal range, while many aquatic ectotherms utilise thermal acclimation to maintain performance over varying seasonal temperatures. This study aimed to elucidate the relative contributions of acclimation and behavioural thermoregulation to maintaining whole-animal performance over varying seasonal temperatures in the semi-terrestrial Lamington spiny crayfish (Euastacus sulcatus). Crayfish activity and surface temperatures were determined by means of radio tracking and behavioural observations. Field studies demonstrated that E. sulcatus is exposed to stable daily temperatures, varying only between seasons from 10°C in late winter to over 20°C in summer. Also, terrestrial behaviour corresponded to a small portion of crayfish time (1.13%), much lower than predicted, indicating that E. sulcatus has limited opportunity for behavioural thermoregulation. We also tested the effect of acclimation to either 10 or 20°C on chela strength and stamina. We found acclimation had a more marked effect on chela stamina than maximum strength measures; however, overall acclimatory capacity was limited in E. sulcatus. Thus, we found that the semi-terrestrial crayfish E. sulcatus used neither thermoregulatory behaviours nor physiological strategies to deal with seasonal changes in environmental temperature.