Inflexible versus flexible: the influence of temperature and photoperiod on pre‐ and post‐eyespot development time in Libellulidae (Odonata)

Temperature and photoperiod are important environmental parameters for organisms. The present study tests the hypothesis that, during embryogenesis, temperature and photoperiod influence pre ‐ and post ‐ eyespot development time in dragonflies of the family Libellulidae differently. Eggs are used from eight species (five different genera, from Africa/Europe, and lentic/lotic habitat preferences). The eggs are reared under different constant or fluctuating temperature and light conditions. There are no general species‐specific degree‐days for pre ‐ or the post ‐ eyespot development in these species. In all study species, the variance within and between the treatments of the duration in days and the degree‐days of pre ‐ eyespot development is lower than that of post ‐ eyespot development. Pre ‐ eyespot development appears to be less flexible in its reaction to environmental influences. By contrast, post ‐ eyespot development appears to react more flexibly to environmental influences. All eight species show the same pattern. This indicates strongly that this flexibility is a general pattern in Libellulidae that might help the species within this family to cope successfully with variations in environmental conditions. Because eyespot development and katatrepsis occur close to each other, the above‐described pattern might also appear in other odonates and in other insect groups that exhibit katatrepsis. For all of them, it is essential for survival to match the time of hatching with adequate external temperature and photoperiodic conditions.     

Effects of photoperiod and temperature on diapause induction in Conogethes punctiferalis (Lepidoptera: Pyralidae)

The yellow peach moth, Conogethes punctiferalis (Guenée), a multivoltine species that overwinters as diapausing larvae, is one of the most serious insect pests on maize in China. Effect of photoperiod and temperature on larval diapause was examined under empirical laboratory conditions. Short‐day treatments caused larval diapause at 25°C, and the critical photoperiod was between 12 and 13 h (or 12 h 51 min) light per day. No sensitive instar was identified for diapause induction under alternated short‐ (L : D 11 : 13 h) and long‐day (L : D 14 : 10 h) treatments at different larval stages. However, accumulative treatment of three instars and 10 d under short‐day treatment was required for the induction of 50% larval diapause. All larvae entered diapause at 20°C, whereas less than 3% did so at 30°C, irrespective of the long‐ or short‐day treatment. Furthermore, under the short‐day treatment, more than 90% of larvae went into diapause with temperatures ≤ 25°C, but less than 17% did so at 28°C. In contrast, under the long‐day treatment, less than 19% of larvae went into diapause with temperatures ≥ 23°C. The forward shift (5°C) of critical temperature under the long‐day regime demonstrated the compensatory effect of temperature and photoperiod on diapause induction. In conclusion, C. punctiferalis had a temperature‐dependent type I photoperiodic diapause response; there was no sensitive instar for diapause determination, but the photoperiodic accumulation time countermeasures both of the short‐day cycles and the number of instars exposed, and the photoperiodic diapause response, was a temperature‐compensated phenomenon.     

Diapause induction as an interplay between seasonal token stimuli,and modifying and directly limiting factors: hibernation in Chymomyza costata

Larvae of wild type (WT) strain of Chymomyza costata Zetterstedt (Diptera: Drosophilidae) enter diapause (stop developing) in response to short‐day signal at a constant 18 °C, whereas larvae of a non‐photoperiodic‐diapause (NPD) strain do not respond to photoperiodic signalling and continue in larval development irrespective of daylength. The present study shows that WT larvae also respond reliably to thermoperiodic signalling (daily cycles of temperature) under constant darkness, whereas the NPD larvae do not, suggesting that the pathways transducing the environmental token stimuli (photoperiod and thermoperiod) onto the diapause developmental programme might merge functionally in the central biological clock system known to be mutated in NPD strain. Temperature and larval population density modify the output of token stimuli signalling. High temperatures (>24 °C) tend to avert, whereas low temperatures (<18 °C), especially in combination with constant darkness, stimulate diapause induction in WT strain. Overcrowding (>200 larvae per 5 g of larval diet) lengthens the duration of larval development and induces a ‘diapause‐like’ developmental arrest of relatively weak intensity in up to 60% of larvae of both strains. At high temperatures (>30 °C), all WT larvae continue direct development but subsequently die during the pupal stage. Low temperature exposure (<12 °C) causes quiescence in the majority of the larvae of both strains. Starvation blocks development and causes mortality when applied in larvae younger than day 3 of the third instar. Older larvae survive starvation and their photoperiodically‐induced developmental pre‐programming is not affected. Collectively, the results show that diapause induction in C. costata is a result of various interacting effects of multiple environmental factors.     

Maternal thermal and photoperiodic effects on the progeny diapause in Trichogramma telengai Sorokina (Hymenoptera: Trichogrammatidae)

Although maternal photoperiodic and maternal thermal effects on the progeny diapause have been demonstrated in a number of insect species, their interaction has been rarely studied. We investigated this interaction in Trichogramma telengai. In a series of experiments, maternal females were reared at day lengths of 12–18 h and at temperatures of 17, 20, 25 and 30°C. Their progeny developed under day length of 12 h and temperatures of 13, 14 and 15°C. The experiments showed that both short day and low temperature experienced by the maternal generation significantly increased the proportion of diapausing progeny. In particular, the threshold of the maternal photoperiodic response decreased with temperature. Under combinations of photoperiod with daily thermoperiod, the role of the “night” temperature in the induction of diapause in the progeny was much more important than that of the “day” temperature. We conclude that the interaction pattern between the photoperiodic and thermal maternal effects in T. telengai is generally the same as that between the photoperiodic and thermal responses directly influencing diapause induction in other long‐day insects. The threshold temperature of the maternal thermal response of T. telengai was about 25–27°C, while diapause can be induced if larvae develop at temperatures not higher than 15–16°C. This suggests that, at least in the studied Trichogramma species, the maternal thermal effect has no ecological value. In the practice of biocontrol, however, rearing of Trichogramma wasps at high temperature can drastically reduce the proportion of diapausing progeny.     

Photoperiodic and temperature effects on the intensity of larval diapause in Sesamia nonagrioides

Abstract. The intensity of larval diapause in Sesamia nonagrioides Lef (Lepidoptera: Noctuidae) was investigated under laboratory conditions. Newly hatched larvae were exposed to different stationary photoperiods (from LD 7 : 17 h to LD 14 : 10 h), at a constant temperature of 25 °C. Diapause incidence was higher when larvae were exposed to daylengths shorter than the critical value (LD 12 : 12 h), whereas the within‐treatment variation in the larval period appeared to be significantly correlated with the photoperiod applied. The incidences of diapause and the duration of larval development were also measured after exposing larvae to short photoperiods (LD 8 : 16 h, LD 10 : 14 h or LD 12 : 12 h) in combination with various temperatures (20, 22.5 or 25 °C). Although an increase in the incidence of diapause appeared with the lowering of the temperature, no statistical differences were observed in the time needed for pupation within the photoperiodic treatments at the temperatures of 20 and 22.5 °C. Furthermore, when diapausing larvae were transferred to the long photoperiod of LD 16 : 8 h, they immediately proceeded to pupation, regardless of the photoperiod or the temperature to which they had been previously exposed, indicating that there were no differences in the intensity of diapause. Photoperiodic changes from LD 10 : 14 h to LD 12 : 12 h or to LD 14 : 10 h at different larval ages reduced the intensity of diapause with (a) early age of transfer and (b) increase of daylength. By contrast, when larvae were transferred from the long photoperiod of LD 14 : 10 h to shorter, such as LD 10 : 14 h or LD 12 : 12 h, a small increase in the intensity of diapause with the shortening of the daylength was apparent. These results support the hypothesis that insects may compare the duration of the photoperiod and could classify them as either longer or shorter in relation to the critical value.     

Effects of temperature,photoperiod and soil humidity on induction of pseudopupal diapause in the bean blister beetle Epicauta gorhami

Larvae of the bean blister beetle Epicauta gorhami Marseul (Coleoptera: Meloidae) feed on grasshopper eggs in soil and undergo hypermetamorphosis. This beetle undergoes larval diapause in the fifth instar as a pseudopupa, a form characteristic of hypermetamorphosis in meloid beetles. The effects of temperature, photoperiod and soil humidity on larval development of E. gorhami are examined in a population in Miyazaki, Japan, using egg pods of Locusta migratoria L. as food. At lower temperatures (20 and 22.5 °C), all larvae become pseudopupae, regardless of the photoperiod. By contrast, at higher temperatures (27.5 and 30 °C), almost all larvae pupate at the end of the fourth instar, again regardless of the photoperiod. A long‐day photoperiodic response occurs only at an intermediate temperature (25 °C): under an LD 12 : 12 h photocycle, all larvae enter diapause, although the diapause incidence tends to decrease as the day length becomes longer. Pseudopupae are immobile and remain in diapause for ≥120 days when they are kept under the same conditions, except that diapause terminates within a relatively short time at 30 °C. Although lower soil humidity retards post‐feeding development, soil humidity has no effect on the diapause incidence. On the basis of the short developmental period and diapause avoidance under summer conditions, it is suggested that this beetle partially produces two generations a year in southwestern Japan.     

Photoperiod-temperature interaction-a new form of seasonal control of growth and development in insects and in particular a Carabid Beetle, <Emphasis Type="Italic">Amara communis</Emphasis> (Coleoptera: Carabidae)

Amara communis larvae were found to develop significantly faster and to have higher growth rate at short-day (12 h) as compared to long-day (22 h) photoperiods at all used temperatures (16, 18, 20, and 22°C). The coefficient of linear regression of larval development rate on temperature was significantly higher at the short day than at the long day. The thermal developmental thresholds appeared similar at both photoperiods. Body weight of young beetles reared under different photoperiods was almost the same. Thus, photoperiod does not simply accelerate or decelerate insect development, but modifies the thermal reaction norm. At short days, larval development becomes faster and more temperature-dependent, which provides a timely completion of development at the end of summer. The analysis of literature data has allowed us to find the photoperiodic modification of thermal requirements for development in 5 insect orders: Orthoptera, Heteroptera, Coleoptera, Lepidoptera, and Diptera. Modification may result in significant changes in the slope of the regression line, and hence the sum of degree-days, and in the thermal developmental threshold. Consequently, the thermal requirements for development in many insects gradually vary during summer under the effect of changing day-length, which may have adaptive significance. Thus, the photoperiodic modification of thermal reaction norms acts as a specific form of seasonal control of insect development.     

Effects of temperature experienced during embryonic development on biomass and C and N composition at hatching in Palaemon serratus (Pennant, 1777)

In decapod crustaceans, the conditions experienced during embryonic development trigger phenotypic plasticity of the larvae at hatching. The objective of this study was to test the effects of temperature during embryonic development of Palaemon serratus on the phenotypic plasticity of hatching larvae. We incubated egg-bearing females from eggs laying to hatching at four temperatures (10, 15, 18 and 20°C). Weight, carbon and nitrogen contents were measured on newly laid eggs and on freshly hatched larvae. The duration of embryonic development was negatively correlated with incubation temperature. At 20°C, all females abandoned their eggs during development. Incubation temperature had no effect on the weight and the percentage of N of the larvae at hatching, while it did affect their percentage of C and their C/N ratio. Embryos incubated at 10°C seemed to produce larvae with fewer lipid reserves than those incubated at 15 and 18°C. They probably overconsumed their lipid reserves to compensate for the metabolic losses due to the low temperature. These results provide information on the link between maternal investment per egg and larval development in P. serratus.     

Biological characteristics of a non‐photoperiodic‐diapause strain of the cabbage beetle Colaphellus bowringi (Coleoptera: Chrysomelidae)

Comparison of biological characteristics between diapausing and non‐diapausing strains of insects provides some insights into the mechanisms regulating diapause. In this study, biological characteristics, especially diapause characteristics and life‐history traits, of a non‐photoperiodic‐diapause (NPD) strain of the cabbage beetle Colaphellus bowringi were compared with those of a normal, high‐diapause (HD) strain that enters diapause in response to either long day length or low temperature. The NPD strain did not enter diapause at any photoperiod at 22°C or higher, but still had a capacity to enter diapause at low temperatures. Although diapause could be induced in both strains by exposure to 20°C, the proportion of individuals having shorter diapause duration was greater in the NPD strain compared to the HD strain. The NPD strain had significantly lower hatching and larval survival rates, longer developmental duration of immature stages, smaller body size and lower longevity and female fecundity compared to the HD strain. The NPD strain of this species is a promising model for investigating diapause regulation in insects in general.     

Temperature and Photoperiodic Control of Development in the Green Shield Bug Palomena prasina (L.) (Heteroptera,Pentatomidae) in Leningrad Province

Thermal reaction norms for development (the lower temperature threshold, temperature sensitivity, and sum of degree-days) can show phenotypic plasticity in response to a combination of ecological factors. The goal of this study was to evaluate the degree of plasticity of the thermal reaction norms for development under different photoperiodic conditions in the green shield bug Palomena prasina. Experiments were conducted in 2015 and 2016. In 2015, two photoperiodic regimens (12 and 22 h of light per day) and five constant temperatures (20, 22, 24, 26, and 28°C) were used; a lower temperature of 16°C was added in 2016. There were no differences in the egg developmental time between the two photoperiodic conditions and across the two experiments. Under the short-day photoperiodic regimen, nymphal development was faster at all the temperatures and was characterized by greater thermal sensitivity and a higher temperature threshold than under the long-day photoperiod. Besides, the relationship between the developmental rate and temperature deviated from linearity at 26 and 28°C under short-day conditions. The adaptive nature of the observed nymphal response to photoperiodic conditions was confirmed by our phenological observations and an outdoor cage experiment. The adult body mass slightly increased with rising temperature under short-day conditions but did not depend on the temperature under long-day ones. Females were larger than males, and both sexes had a greater body mass under long-day conditions than under short-day ones. In 2015, the eggs for experiments were collected before mid-July, almost a month later than in 2016. Nymphs that hatched from the later eggs (in 2015) had significantly higher relative growth rates than the early-season nymphs (in 2016) at 20, 22, and 24°C under both photoperiods. This discrepancy between years was probably related to the maternal effect, namely, the difference in the female physiological age.

     

Environmental induction of larval diapause and life-history consequences of post-diapause development in the Large Copper butterfly,Lycaena dispar (Lepidoptera: Lycaenidae)

Many insects in temperate zones withstand the adverse conditions of winter through entering diapause and the two most important environmental stimuli that induce diapause are photoperiod and ambient temperature. The Large Copper butterfly, Lycaena dispar Haworth (Lepidoptera: Lycaenidae), is a Palearctic butterfly that hibernates as larvae. Since this butterfly is a near threatened species in some regions, there has been a growing need for a standardized protocol for mass rearing of this butterfly based on the adequate knowledge of its ecology. In the present study, we first identified that L. dispar larvae were sensitive to the photoperiodic induction of diapause during their first larval instar. We then investigated to what extent the diapause-inducing effects of photoperiod could be modified by ambient temperatures in L. dispar larvae by exposing them to the range of day-lengths (L:D 14:10, 12:12, 10:14 and 8:16) at three different temperatures (15, 20 and 25 °C). All larvae were induced to enter diapause at low ambient temperature (15 °C) regardless of photoperiod, whereas most of them (86 %) exhibited direct development when temperature was high (25 °C). The photoperiodic induction of diapause was evident when day-length was shorter than 14 h at intermediate temperature (20 °C). Pre-diapause development was prolonged at low temperatures. Finally, we found that post-diapause development of L. dispar larvae was determined by both the chilling temperature experienced by diapausing larvae and the duration of larval diapause. Adult emergence was enhanced when larvae were chilled at 8 °C and when they had been under the state of diapause for 20 days before they were treated to terminate diapause.     

Induction of larval diapause in the blowfly, Calliphora vicina R.-D. (Diptera, Calliphoridae) under field and laboratory conditions

Field experiments conducted in the environs of St. Petersburg (Russia) with a local population of Calliphora vicina showed that induction of larval diapause under natural conditions was significantly dependent on day lengths and temperature. The maternal photoperiodic response had a distinct threshold: the first diapausing larvae hatched from the eggs laid in the middle of August when the day length was 16 h; at shorter photoperiods, the fraction of diapausing larvae depended only on temperature. At the mean temperature of 16°C, larval diapause was rarely recorded; at 12–13°C, about 50% of the larvae entered diapause; at 7–9°C, nearly all the larvae entered diapause. These results of the field experiments agree well with the parameters of photoperiodic and thermal responses studied in the laboratory at constant temperatures and photoperiods.     

Climate change, diapause termination and zooplankton population dynamics: an experimental and modelling approach

1. Earlier spring warming as predicted for climate change will alter combinations of water temperature and photoperiod that act as emergence cues for zooplankton resting stages. As a result, water temperature cue thresholds will be experienced at shorter photoperiods, a variable independent of weather variations. Also, light intensity, another potentially important cue for zooplankton emergence, could decrease in many lakes if symptoms of climate change resemble those of eutrophication. 2. We designed a laboratory experiment to test the effects of three factors, temperature (6, 9 and 12 °C), photoperiod (13L : 11D and 16L : 8D) and light intensity (20 and 35 μE m⁻² s⁻¹) on hatchling abundance and timing of hatching of daphniids (Daphnia ambigua) and rotifers (Keratella spp. and Synchaeta pectinata) from resting eggs. Further, we investigated the implications of potential changes in hatching dynamics, following variations in hatching cues, on zooplankton spring population development using predator–prey simulation models. 3. For hatchling abundance and timing of hatching, photoperiod had a significant effect for D. ambigua but not rotifers. Daphnia ambigua hatchling abundance decreased by 50% when incubated at conditions mimicking early spring (12 °C + 13‐h photoperiod) compared to a later spring (12 °C + 16‐h photoperiod). Light intensity has a significant effect only for S. pectinata, producing greater hatchling abundance at lower light intensity. 4. Simulation models suggest that in contrast to a later spring, an early warming produces a shift in spring zooplankton community composition, from daphniid to rotifer dominance. These patterns are primarily driven by differential zooplankton emergence response with variations in temperature–photoperiod cues. 5. Overall, our laboratory experiments and simulation models suggest that lakes with strong dependence on the ‘resting egg‐bank’, characteristic of many shallow north‐temperate lakes or in years with low winter survivorship of adult zooplankton, may be most susceptible to climate change. Further, fewer large grazers such as daphniids with an earlier spring may result in less control of cyanobacterial blooms in eutrophic lakes.     

Thermal reaction norms in two Coenagrion damselfly species: contrasting embryonic and larval life-history traits

1. We studied the temperature‐dependence of important life‐history traits both at the embryonic (egg hatching success, embryonic development time and hatchling size) and the larval stage (larval growth rate, larval survival and larval size after 100 days) using full‐sib families of two congeneric damselflies, Coenagrion hastulatum and Coenagrion puella, that differ in latitudinal distribution. Larvae were reared in the laboratory from the egg stage at four temperatures (12, 17, 22 and 27 °C). 2. The observed patterns of thermal plasticity in embryonic traits showed that the northern species was more successful than the southern species at lower temperatures, in line with the pattern of temperature adaptation in thermal reaction norms. 3. At the larval stage, we found no consistent pattern of latitudinal compensation. The thermal family reaction norms indicate, however, the potential for latitudinal compensation to evolve. We observed an ontogenetic shift in thermal optima for larval growth rate, with a higher optimal temperature for growth rate during the first 2 weeks of the larval stage. 4. This is the first indication of the existence of latitudinal compensation at the interspecific level in an invertebrate; it is stage‐specific, being present only in the embryonic stage. We argue that compensation in the embryonic stage may be much more likely than in the larvae and stress the importance of including more then one life‐history stage when drawing conclusions about the adaptiveness of patterns in thermal reaction norms.     

Effects of temperature on hatching rate, embryonic development and early larval survival of the edible sea urchin, Tripneustes gratilla

The temperature tolerances of embryonic and early larval development stages of Tripneustes gratilla were investigated from 13-34°C under laboratory conditions. Zygotes showed unequal cleavage at 13°C, whereas cleavage did not occurred at 34°C. Hatching was observed between 16–31°C with maximum hatching rates observed at 22–29°C. The lower and higher temperature limits for embryonic development were approximately 22°C and 29°C, respectively. Outside of this temperature range, embryos showed abnormality at different incubation times. Early larvae of this species have the ability to survive the higher temperature limit for short periods of time. Prism and 2 arm pluteus larvae survived at temperatures between 30 and 33°C, whereas 4 arm pluteus larvae survived at temperatures between 30 and 36°C for 2 h. These results suggest that the larval temperature tolerance capability of T. gratilla is stage dependent. These findings are important for understanding the life history strategy of this sea urchin in the shallow open water environment.     

Maternal thermal effect on diapause in Trichogramma species (Hymenoptera: Trichogrammatidae)

Both direct thermal and maternal photoperiodic effects on diapause induction have been thoroughly investigated in many insect species, while maternal thermal effects have been infrequently studied. We studied the effect of temperature during development of maternal generation on the proportion of diapausing progeny in four species of the genus Trichogramma Westw., minute egg parasitoids, widely used for biological control of lepidopteran pests. The maternal generations were reared at day lengths of 12 and 18 h and temperatures of 17, 20, 25 and 30°C, and their progeny developed under day length of 12 h and temperatures of 13 and 14°C. In T. evanescens and T. piceum, the proportion of diapausing progeny decreased with increasing temperature under all tested photoperiods and thermal regimes of progeny development; the high temperature of 30°C totally averted diapause of progeny. In T. buesi and T. principium, low temperatures of 17 and 20°C resulted in relatively high proportion of diapausing progeny only when the maternal generation developed under short‐day conditions. The threshold of the maternal thermal response varied from 17–18 to 22–23°C. Under field conditions, Trichogramma females are exposed to such high temperatures only during summer, when diapause in their progeny is in any case prevented by the maternal photoperiodic response and by the thermal response of the larvae. We conclude that the maternal thermal effect on diapause induction, although to a different extent, is inherent to Trichogramma species but, at least as suggested by laboratory experiments, it does not play any role in the regulation of seasonal development under natural conditions. However, during mass rearing of Trichogramma wasps, it should be taken into account that high temperature, even when combined with short photoperiod, can avert diapause in the next generation.     

Historical and projected interactions between climate change and insect voltinism in a multivoltine species

Climate change can cause major changes to the dynamics of individual species and to those communities in which they interact. One effect of increasing temperatures is on insect voltinism, with the logical assumption that increases in surface temperatures would permit multivoltine species to increase the number of generations per year. Though insect development is primarily driven by temperature, most multivoltine insect species rely on photoperiodic cues, which do not change from year‐to‐year or in response to climate warming, to initiate diapause. Thus, the relationship between climate change and voltinism could be complex. We use a phenology model for grape berry moth, Paralobesia viteana (Clemens), which incorporates temperature‐dependent development and diapause termination, and photoperiod‐dependent diapause induction, to explore historical patterns in year‐to‐year voltinism fluctuations. We then extend this model to predict voltinism under varying scenarios of climate change to show the importance of both the quality and quantity of accumulated heat units. We also illustrate that increases in mean surface temperatures > 2 °C can have dramatic effects on insect voltinism by causing a shift in the ovipositional period that currently is subject to diapause‐inducing photoperiods.     

Photoperiodic and thermal regulation of antifreeze protein levels in the beetle Dendroides canadensis

The importance of photoperiod, temperature and their interaction in controlling the seasonal pattern of haemolymph antifreeze protein levels in larvae of the beetle Dendroides canadensis was investigated. A complete photoperiodic response curve for antifreeze protein production was generated at 20°C with larvae collected in early fall. Individuals exposed to a 10-h photoperiod or less, including constant darkness, had significantly elevated antifreeze levels over those maintained in an 11-h photoperiod or more, including constant light. The critical daylength resulting in 50% population response lies between LD 11:13 and LD 10:14. This photoperiodic response was masked at sufficiently low (threshold between 15 and 10°C) and high (threshold between 25 and 30°C) temperatures. Partial photoperiodic response curves (at 17 and 25°C) obtained within this specified temperature range indicate that the position of the critical photoperiod (between 10 and 11 h) is stable while the amplitude of the response curve is temperature dependent.Experiments investigating the mechanisms controlling the spring depletion of protein antifreeze levels suggest that both photoperiod and temperature are important.The dominant response of photoperiod in the fall along with the modifying effects of temperature are considered to provide the necessary precision to assure adequate cold tolerance early in the fall and the flexibility to protect the species from yearly variation in weather conditions.     

Moving south: effects of water temperatures on the larval development of invasive cane toads (Rhinella marina) in cool‐temperate Australia

The distributional limits of many ectothermic species are set by thermal tolerances of early‐developmental stages in the life history; embryos and larvae often are less able to buffer environmental variation than are conspecific adults. In pond‐breeding amphibians, for example, cold water may constrain viability of eggs and larvae, even if adults can find suitable thermal conditions in terrestrial niches. Invasive species provide robust model systems for exploring these questions, because we can quantify thermal challenges at the expanding range edge (from field surveys) and larval responses to thermal conditions (in the laboratory). Our studies on invasive cane toads (Rhinella marina) at the southern (cool‐climate) edge of their expanding range in Australia show that available ponds often average around 20°C during the breeding period, 10°C lower than in many areas of the toads’ native range, or in the Australian tropics. Our laboratory experiments showed that cane toad eggs and larvae cannot develop successfully at 16°C, but hatching success and larval survival rates were higher at 20°C than in warmer conditions. Lower temperatures slowed growth rates, increasing the duration of tadpole life, but also increased metamorph body mass. Water temperature also influenced metamorph body shape (high temperatures reduced relative limb length, head width, and body mass) and locomotor performance (increased speed from intermediate temperatures, longer hops from high temperatures). In combination with previous studies, our data suggest that lower water temperatures may enhance rather than reduce recruitment of cane toads, at least in areas where pond temperatures reach or exceed 20°C. That condition is fulfilled over a wide area of southern Australia, suggesting that the continuing expansion of this invasive species is unlikely to be curtailed by the impacts of relatively low water temperatures on the viability of early life‐history stages.     

Temperature and photoperiodic regulation of diapause of the southwestern corn borer,Diatraea grandiosella

The effect of temperature and photoperiod on the onset and termination of the mature larval diapause of the southwestern corn borer, Diatraea grandiosella, was examined. The results showed that diapause induction was an extremely temperature-dependent process. Larvae reared at 23°C under short days all entered diapause whereas 90 per cent of those reared at 27°C developed continuously. A photoperiodic response was only demonstrable at 25°C, when diapause was instituted following larval exposure to daily photophases ranging from 8 to 14 hr. An examination of the sensitivity of immature larvae to low temperatures revealed that all instars showed some susceptibility to low temperature induction of diapause. No intermediate instar was found in which the developmental programme could be switched entirely from a diapause to a non-diapause one, or vice versa.Diapause development was also found to be primarily temperature regulated. The rate of termination of diapause at 30°C was significantly higher than that at 25 or 23°C. Continuous exposure to light rather than a daily photoperiod produced the highest rate of diapause development. It was also shown that a period of chilling (5°C) did not accelerate diapause development. The significance of the results in relation to the evolution and geographical distribution of the species is discussed.