Thermal biology of the meadow grasshopper, Chorthippus parallelus, and the implications for resistance to disease

Abstract.  1. The thermal biology of the meadow grasshopper, Chorthippus parallelus , a common, habitat generalist acridid species found in the U.K., was characterised and the influence of thermoregulatory behaviour for resistance against a temperate ( Beauveria bassiana ) and tropical ( Metarhizium anisopliae var. acridum ) fungal pathogen was determined.
2.  Chorthippus parallelus was found to be an active behavioural thermoregulator, with a preferred temperature range of 32–35 °C.
3. Both pathogens proved lethal to fifth instar and adult grasshoppers. No evidence of behavioural fever in response to infection by either pathogen was found, but normal thermoregulation was found to reduce virulence and spore production of B. bassiana. Normal thermoregulation did not appear to affect M. anisopliae var. acridum .
4. These results suggest that the effects of temperature on host resistance depend on the thermal sensitivity of the pathogen and, in this case, derive from direct effects of temperature on pathogen growth rather than indirect effects mediated by host immune response.
5. The implications for possible risks of exotic pathogens and influence of climate change are discussed.     

Diapause induction and coloration in the Senegalese grasshopper, Oedaleus senegalensis

Abstract. Embryonic diapause induction in the Senegalese grasshopper, Oedaleus senegalensis Krauss (Orthoptera: Acrididae), is influenced both by the photoperiod and the temperature experienced by females. High temperatures (40°C) and long photoperiods (LD 14:10h), which characterize the beginning of the rainy season in the Sahel, cause non-diapausing eggs to be laid. Lower temperatures (25°C) and shorter photoperiods (LD 12:12h), which occur at the end of the rains, result in the production of diapausing eggs. At 30°C and constant photoperiods, O. senegalensis exhibited a long-day-short-day response with critical photoperiods of c. 13 h and c. 20 h, only the former value being of ecological significance. The photoperiodically sensitive stages to diapause induction in females occurred from the fifth stadium onwards. Temperature also affected the coloration of both nymphs and adults. Dark-black and pale-white individuals were produced by low (25°C) and high (40°C) temperatures respectively, whereas an intermediate temperature (30°C) produced individuals which were greyish brown. These results are discussed in relation to the ecology of O. senegalensis.     

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.     

Thermoregulation in four species of British grasshoppers (Orthoptera: Acrididae)

1. This study examines the extent to which thermal balance and thermoregulatory ability may contribute to habitat partitioning in insect herbivores.
2. The distribution of four species of grasshopper on a Breckland grass heath is described. Myrmeleotettix maculatus is restricted to short swards, Omocestus viridulus is restricted to long swards, and Chorthippus brunneus and Stenobothrus lineatus are found on swards of intermediate length.
3. Short swards are warmer on average than long swards, but lack cooler refuges on hot days.
4. Chorthippus brunneus and O. viridulus are better able to raise their body temperatures at low ambient temperatures than M. maculatus and S. lineatus . Omocestus viridulus is less able to reduce body temperature at high ambient temperature.
5. Myrmeleotettix maculatus may be precluded from inhabiting cooler long swards because of its inability to raise body temperatures at low ambient temperatures. Omocestus viridulus may avoid short swards because of the danger of overheating.
6. Thermoregulatory ability is a good predictor of the distribution of the grasshoppers in swards of differing length and microclimate.     

Thermal ecology of western tent caterpillars Malacosoma californicum pluviale and infection by nucleopolyhedrovirus

Abstract 1. Western tent caterpillars hatch in the early spring when temperatures are cool and variable. They compensate for sub-optimal air temperatures by basking in the sun.
2. Tent caterpillars have cyclic population dynamics and infection by nucleopolyhedrovirus (NPV) often occurs in populations at high density.
3. To determine whether climatic variation might influence viral infection, the environmental determinants of larval body temperature and the effects of temperature on growth and development rates and larval susceptibility to NPV were examined.
4. In the field, larval body temperature was determined by ambient temperature, irradiance, and larval stage. The relationship between larval body temperature and ambient temperature was curvilinear, a property consistent with, but not necessarily limited to, behaviourally thermoregulating organisms.
5. Larvae were reared at seven temperatures between 18 and 36 °C. Larval growth and development increased linearly with temperature to 30 °C, increased at a lower rate to 33 °C, then decreased to 36 °C. Pupal weights were highest for larvae reared between 27 and 30 °C.
6. The pathogenicity (LD₅₀) of NPV was not influenced by temperature, but the time to death of infected larvae declined asymptotically as temperature increased.
7. Taking into account larval growth, the theoretical yield of the virus increased significantly between 18 and 21 °C then decreased slightly as temperatures increased to 36 °C.
8. Control and infected larvae showed no difference in temperature preference on a thermal gradient. The modes of temperature preference were similar to those for optimal growth and asymptotic body temperatures measured in the field on sunny days.
9. Warmer temperatures attained by basking may increase the number of infection cycles in sunny springs but do not protect larvae from viral infection.     

Influence of ambient and thoracic temperatures upon sexual behaviour of the gypsy moth, Lymantria dispar

ABSTRACT. In an ambient temperature ( T _a) range of 18–28°C, thoracic temperatures ( T _th) of individual male Lymantria dispar (L.), caught at flight in the field, ranged from 21 to 36.5°C, with a correlation coefficient of 0.63 between T _th and ambient temperature ( T _a). Ambient temperature (and insolation) altered the insect's body temperature and the probabilities, latencies, and durations of preflight responses to pheromone. In a wind tunnel at 16 and 20°C, quiescent males exposed to pheromone raised their T _th by sustained wing fanning from 17 and 21°C, respectively, to c. 24°C before takeoff. At 24 and 28°C ambient, T _th rose by takeoff to 28 and 31°C, respectively. The latencies of male wing fanning in response to pheromone decreased from 1.44 min at 16°C ambient, to 0.58 min at 20°C, to 0.26 min at 24°C, and to 0.16min at 28°C. The components of behaviour (antennal twitch, body jerk, step and wing tremor) that occurred between quiescence and wing fanning were more frequent at ambients of 16 and 20°C than at 24 and 28°C.     

Rate of metabolism, temperature regulations, and evaporative water loss in the lesser gymnure Hylomys suillus (Insectivora, Mammalia)

Rate of metabolism, body temperature, wet thermal conductance, and evaporative water loss were measured at different ambient temperatures in four lesser gymnures Hylomys suillus. Gymnures responded as typical endothermic homoiotherms to changes in ambient temperature. Below the lower critical temperature of 32°C, they maintained a body temperature of 37.3± 0.3°C by an increased rate of metabolism. Minimum wet thermal conductance was 111% of that expected on the basis of body mass. Average basal rate of metabolism was 1.04 ml O₂ g⁻¹ h⁻¹, which represents 106% of the expected value. Within and above the thermoneutral zone, heat loss by evaporation did not account for more than 30% of the heat produced. As a consequence, the body temperature of gymnures was maintained 4°C above ambient temperature. These metabolic and thermoregulatory patterns differ strikingly from those of other members of the family Erinaceidac and can be interpreted as a result of physiological adaptation to a different ecology. Being smaller than hedgehogs and inhabiting montane tropical rainforests, lesser gymnures lack the physiological traits which enable many hedgehogs to invade hot, arid and/or strongly seasonal environments.     

Thermoregulation and daily activity patterns in a black desert grasshopper,Taeniopoda eques

Behavioural thermoregulation was studied in the western horse lubber grasshopper Taeniopoda eques (Burmeister), a native of the Chihuahuan Desert of North America. The grasshoppers regulated their temperature through a series of daily cyclical vertical movements between vegetation and the soil, and by the adoption of four thermoregulatory postures: flanking, crouching, stilting and stem-shading. At dawn, the grasshoppers moved from their nocturnal roost-plants to the ground, returned to bushes during the middle of the day, moved back to the open ground in the afternoon, then reascended vegetation at dusk. The occurrences of the four thermoregulatory postures were synchronized with these microhabitat shifts. During the cooler mornings and afternoons, the insects maximized heat gain by flanking and crouching, achieving thoracic temperatures of up to 16°C above ambient. Throughout the hot middle of the day the insects stilted and shaded, minimizing heat gain. These behaviours effectively kept the grasshoppers' body temperatures near the preferred temperature (36·2°C), but lower than the maximum voluntarily tolerated temperature (41·9°C), critical thermal maximum (45·2°C) and instantaneous lethal maximum (46·5°C). The body size of flanking insects influenced heating and cooling rates, wind effects and temperature excess at equilibrium. Both infrared and visible radiation appeared to elicit flanking. The need and ability to thermoregulate are influenced by this insect's reliance on chemical deterrents for defence.     

Head-body thermal control, thermal preferenda, and voluntary maxima in the taipan, Oxyuranus scutellatus (Serpentes: Elapidae)

Head-body temperature differences of up to 5.7° and 6.1° C were observed during solar and radiant heating, respectively. Head temperature was more precisely regulated than body temperature and within preferred limits head temperature was significantly lower than body temperature. Higher temperatures were tolerated by the body than the head. Preferred temperatures for both the head and body are higher in the taipan than in other ophidians which have been reported upon, and approximate those temperatures reported for the voluntary maxima in other snakes.     

The relative importance of temperature and diet to larval development and adult size of the winter stonefly, Soyedina carolinensis (Piecoptera: Nemouridae)

SUMMARY. 1. Soyedina carolinensis Claassen, a leaf shredding stonefly, was reared in a series of three laboratory experiments from early instar to adult on different species of deciduous leaves and at various constant and fluctuating temperature regimes.
2. Experiment 1, which involved rearing larvae on fourteen different leaf diets at ambient stream temperatures, showed that diet significantly affected larval growth and adult size but did not affect overall developmental time.
3. Experiment 2, which involved rearing larvae on five different leaf diets at each of three fluctuating temperature regimes (viz ambient White Clay Creek (WCC), ambient WCC+3°C, and ambient WCC+6°C), showed that: (i) adding 6°C to the normal temperature regime of WCC was lethal to 99% of the larvae regardless of diet; and (ii) warming WCC by 3°C did not affect developmental time but did significantly reduce adult size relative to adults reared at WCC temperatures on certain diets.
4. Experiment 3, which involved rearing larvae on five different leaf diets at each of five constant temperatures (viz 5, 10, 15, 20, 25°C), showed that: (i) temperature significantly affected the mortality, growth, and development time of larvae whereas diet only affected larval growth and mortality; (ii) temperatures at or near 10°C yielded maximum larval growth and survival for most diets; (iii) at 5°C, larval mortality was high and growth was low resulting in a few small adults for most diets; (iv) larval mortality was at or near 100% at 15°C regardless of diet; and (v) no larvae survived at 20 and 25°C.     

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.     

Synchronization of larval emergence in winter moth (Operophtera brumata L.) and budburst in pedunculate oak (Quercus robur L.) under simulated climate change

1. The hypothesis that a 3 °C elevation in temperature and doubled CO₂ concentration would have no effect on the synchronization of winter moth egg hatch with budburst in oak was tested by comparing the separate and interactive effects of ambient and elevated (+ 3 °C) temperature and ambient and elevated (doubled to 340 p.p.m.) CO₂ in eight experimental Solardomes. In addition, an outdoor control was compared with the ambient temperature/CO₂ treatment combination.
2. Elevated temperature accelerated darkening (preceding egg hatch by about 5–10 days) and hatching of eggs developing off the trees; elevated CO₂ had no effect. The same effects were observed in eggs developing on the trees.
3. Within treatments, date of egg hatch was the same on trees with early or late budburst.
4. Egg darkening and budburst were closely synchronized at both ambient and elevated temperatures.
5. Both eggs and trees required fewer cumulative heat units (day degrees > 4 °C), for hatching and budburst, respectively, at ambient than elevated temperatures. The requirements in the outdoor control treatment were similar to those in the ambient Solardome treatment.
6. Egg hatch between 10 and 25 °C, on a temperature gradient in the laboratory, required a constant number of heat units; fewer were required below 10 °C.
7. Elevated temperatures, in the Solardomes and the field, delayed adult emergence from the pupae.
8. The results suggest that a general increase in temperature with climatic change would not affect the closeness of the synchronization between egg hatch of winter moth and budburst of oak.     

Thermal ecology of the mudskippers, Periophthalmus koelreuteri (Pallas) and Boleophthalmus boddarti (Pallas) of Kuwait Bay

The annual range of body temperatures (14–35°C) of emergent mudskippers are substantially less than that of air temperatures (10–42°C) as a result of behavioural thermoregulation. In winter, low surface temperatures are avoided by remaining in burrows. Newly emerged mudskippers then bask until body temperatures rise above 14°C before they move onto the mud. In summer, body temperatures are kept lower than ambient by selecting areas where evaporative cooling is high. Body temperatures generally match those of wet mud, which can be 7°C lower than air shade temperatures. The smaller, more terrestrial, Periophthalmus koelreuteri have body temperatures which are mainly lower in summer and higher in winter than Boleophthalmus boddarti .     

Altitudinal variation in behavioural thermoregulation: local adaptation vs. plasticity in California grasshoppers

We investigated the adaptive significance of behavioural thermoregulation in univoltine populations of the grasshopper Melanoplus sanguinipes along an altitudinal gradient in California using laboratory tests of animals raised under different temperatures. Trials consisted of continuous body temperature measurements with semi-implanted microprobes in a test arena, and observation and simultaneous recording of behavioural responses. These responses included mobility, basking and orientation of the body axes (aspect angle) towards a radiation source. Mobility and basking are determined by the altitudinal origin of the parental generation and not by the temperature treatments. With increasing altitude, individuals tend increasingly to raise body temperatures via mobility and increased basking. In contrast, body orientation towards the radiation source is influenced by the temperature treatments but not by the altitude of origin. Individuals experiencing higher temperatures during rearing show a lower tendency to lateral flanking. We conclude that body orientation responses are not adapted locally. In contrast other components of the behavioural syndrome that increase body temperature, such as mobility and basking, are adaptive in response to local selection pressure. The thermoregulatory syndrome of these grasshoppers is an important contribution to life-history adaptations that appropriately match season lengths.     

Nest thermoregulation in Vespa simillima, V.tropica and V.analis

Abstract. 1. Nest thermoregulation follows a similar pattern in Vespa simillima Smith and V. tropica L. There is a gradual decline in the daily fluctuations of nest temperature to a constant steady state which is maintained during the production of the first sexuals, followed by a sudden loss of stability at the end of the colony cycle.
2. The larvae are not major producers of heat, as they are unable to raise their body temperature by more than 1–2°C above ambient. However, they act as heat reservoirs and providers of food in the form of larval secretions. This feeding may allow workers to raise their body temperature during non-foraging periods.
3. The adults are capable of raising their body temperature many degrees above ambient and the presence of even one adult, in this case a V.analis F. mother queen, was able to raise the nest temperature.
4. Supplementary carbohydrate food promotes thermogenesis and enhances colony development.     

Overwintering survivorship of pupae of the mimosa web worm, Homadaula anisocentra (Lepidoptera: Plutellidae), in an urban landscape

ABSTRACT. 1. Overwintering survivorship of pupae of the mimosa webworm, Homadaula anisocentra Meyrick (Lepidoptera: Plutellidae), was examined in several urban habitats in central Iowa during the winters of 1981–82, 1982–83 and 1983–84.
2. Survivorship and supercooling point temperatures were determined throughout the winters. Corroborative laboratory studies were conducted during the winter of 1982–83.
3. Minimum ambient temperatures that equalled or were below the supercooling point of the insect, at any time, were lethal.
4. Prolonged cold exposure below 0°C and above the supercooling point resulted in high mortality levels. To quantify this relationship, a concept of minimum-temperature exposure was developed by tabulating the number of degrees that the daily minimum temperature was below 0°C for a given sampling period.
5. Some mimosa webworm pupae were found to overwinter in highly protected sites (2.5–5.0°C warmer than the ambient air temperatures) in the urban environment, resulting in less minimum-temperature exposure and reducing the probability of reaching the lethal supercooling point temperature.     

Diapause duration, survival in relation to desiccation and egg-pod morphology of the Senegalese grasshopper, Oedaleus senegalensis

Abstract. The Senegalese grasshopper, Oedaleus senegalensis (Krauss) (Orthoptera: Acrididae), is periodically a devastating pest of subsistence crops in the sahelian zone of West Africa.Egg diapause has been suggested as an important component of the mechanism acting to generate outbreaks and therefore diapause duration, egg survival in relation to desiccation and egg-pod morphology were investigated in this study.
Three months after oviposition, 50% of initially diapausing egg pods maintained in the laboratory exhibited 'partial emergence' and produced hoppers.No difference in diapause duration was evident between egg pods maintained at a constant 30 ± 1°C and those maintained under alternating temperature conditions of 23 ± 1°C and 35 ± 1°C.Laboratory maintained eggs showed significant variation in diapause duration with diapause lasting between 2 months and 4 years.Diapause duration was shorter in the field with only c. 1.4% of eggs remaining in diapause after a single dry season (6–7 months).Survival of eggs in relation to desiccation was high in the field and low in the laboratory, and field eggs lost only c. 20% of their body weight over the dry season.Survival of diapausing and non-diapausing egg pods in the laboratory was similar, indicating that diapause in this species is primarily a mechanism for preventing emergence at an inappropriate time of year, rather than conferring any additional resistance to desiccation.Egg pods oviposited under diapause-inducing conditions (LD 14:10 h, 25°C) were significantly shorter than those laid under non-diapausing conditions (LD 10:14 h, 40°C).These results are discussed both in relation to possible O.senegalensis 'bet-hedging' strategies and to the probable mechanism operating behind major outbreaks of this species.     

Fattening strategies of wintering great tits support the optimal body mass hypothesis under conditions of extremely low ambient temperature

1.  In a seasonal environment, subcutaneous energy reserves of resident animals often increase in winter and decline again in summer reflecting gradual seasonal changes in their fattening strategies. We studied changes in body reserves of wintering great tits in relation to their dominance status under two contrasting temperature regimes to see whether individuals are capable of optimizing their body mass even under extreme environmental conditions.
2.  We predicted that dominant individuals will carry a lesser amount of body reserves than subordinate great tits under mild conditions and that the body reserves of the same dominant individuals will increase and exceed the amount of reserves of subordinates under conditions of extremely low ambient temperatures, when ambient temperature dropped down to −37 °C.
3.  The results confirmed the predictions showing that dominant great tits responded to the rising risk of starvation under low temperatures by increasing their body reserves and this was done at the expense of their safety.
4.  Removal experiments revealed that lower body reserves of subordinate flock members are due to the increased intraspecific competition for food under low ambient temperatures.
5.  Our results also showed that fattening strategies of great tits may change much quicker than previously considered, reflecting an adaptive role of winter fattening which is sensitive to changes in ambient temperatures.     

Influence of pregnancy on the thermal biology of the lizard, Sceloporus jarrovi: why do pregnant females exhibit low body temperatures?

1. Selected body temperatures of female lizards, Sceloporus jarrovi , were measured on a photothermal gradient during late pregnancy and again when postpartum, and pregnant females were subjected to one of three fluctuating temperature regimes that simulated body temperatures of (1) pregnant females, (2) postpartum females or (3) allowed normal thermoregulation.
2. Overall, females selected lower body temperatures when pregnant (mean = 32·0°C) than when postpartum (mean = 33·5°C).
3. Females regulated body temperature more precisely when pregnant than when postpartum as judged by their smaller variances in body temperature throughout the day.
4. When pregnant, females selected a lower mean maximum body temperature (mean: pregnant = 32·8°C; postpartum = 34·5°C) than when postpartum, but selected mean minimum body temperatures did not differ.
5. None of the experimental temperature treatments was detrimental to pregnant females. Female body length increased during pregnancy but the rate of increase did not differ among treatments. Moreover, length-adjusted body mass of postpartum females did not differ among treatments.
6. Pregnant females that experienced postpartum body temperatures produced neonates that were smaller in body mass and length than pregnant females that experienced pregnant body temperatures and females that were allowed to thermoregulate.
7. For neonates resulting from the postpartum body temperature treatment, the disparity in the body length, but not mass, was still observed at 9 days of age, although survival and growth of neonates was high and did not differ among treatments.
8. The results demonstrate that pregnant females could maintain higher postpartum body temperatures without compromising their physical condition, but select relatively low body temperatures, presumably to avoid decrements in offspring fitness.     

Thermoregulation by an Australian murine rodent, the ash-grey mouse (Pseudomys albocinereus)

We examine here the thermal physiology of the ash-grey mouse, as there is a paucity of data to explain how Australian rodents meet thermoregulatory demands. Most ash-grey mice remained normothermic over a range of ambient temperatures (10°C to 30°C), although they became hyperthermic at high ambient temperatures. One individual entered torpor at ambient temperatures of 20°C and 25°C, with minimal body temperatures of 24.5°C and 28.4°C respectively, before spontaneously arousing. This is the first evidence of torpor use by an Australian murine rodent. Our data suggest that although ash-grey mice have the physiological ability to use torpor, it is used rarely, presumably due to other behavioural and physiological adaptations. Their higher-than-expected basal metabolic rate (1.56±0.25mLO(2)g(-1)h(-1)) indicates that ash-grey mice do not have a frugal approach to energy expenditure. Other standard physiological variables were typical of a generalised rodent. A readily-available omnivorous diet, nocturnal activity, semi-fossorial habit and social behaviour presumably allow a high energy lifestyle. A reluctance to use torpor, despite an apparent physiological ability to do so, supports the idea that the use of torpor reflects a net balance between the costs and benefits of a heterothermic thermoregulatory strategy.