Dissecting insect responses to climate warming: overwintering and post‐diapause performance in the southern green stink bug,Nezara viridula,under simulated climate‐change conditions

The effect of simulated climate change on overwintering and post‐diapause reproductive performance is studied in Nezara viridula (L.) (Heteroptera: Pentatomidae) close to the species' northern range limit in Japan. Insects are reared from October to June under quasi‐natural (i.e. ambient outdoor) conditions and in a transparent incubator, in which climate warming is simulated by adding 2.5°C to the ambient temperatures. Despite the earlier assumption that females of N. viridula overwinter in diapause, whereas males do so in quiescence, regular dissections show that the two sexes overwinter in a state of true diapause. During winter, both sexes are dark‐coloured and have undeveloped reproductive organs. Resumption of development does not start until late March. During winter, the effect of simulated warming on the dynamics and timing of physiological processes appears to be limited. However, the warming significantly enhances winter survival (from 27–31% to 47–70%), which is a key factor in range expansion of N. viridula. In spring, the effect of simulated warming is complex. It advances the post‐diapause colour change and transition from dormancy to reproduction. The earlier resumption of development is more pronounced in females: in April, significantly more females are already in a reproductive state under the simulated warming than under quasi‐natural conditions. In males, the tendency is similar, although the difference is not significant. Warming significantly enhances spring survival and percentage of copulating adults, although not the percentage of ovipositing females and fecundity. The results suggest that, under the expected climate‐warming conditions, N. viridula will likely benefit mostly as a result of increased winter and spring survival and advanced post‐diapause reproduction. Further warming is likely to allow more adults to survive the critical cold season and contribute (both numerically and by increasing heterogeneity) to the post‐overwintering population growth, thus promoting the establishment of this species in newly‐colonized areas.     

Surviving winter: diapause syndrome in the southern green stink bug Nezara viridula in the laboratory,in the field,and under climate change conditions

The southern green stink bug Nezara viridula (L.) (Heteroptera: Pentatomidae) has long attracted attention not only as a serious pest of numerous agricultural crops, but also as a species expanding its range in many parts of the world. Nezara viridula has also been widely used as a model in different experimental studies. The present review focuses on reproductive (i.e. adult) winter diapause, which is the pivotal element of the species' seasonal cycle. Results from numerous field experiments and observations, as well as laboratory ecophysiological investigations conducted during the few last decades, are analyzed and interpreted. Experimental findings are used to describe in detail the dynamics of physiological changes during overwintering. Reproductive diapause in N. viridula is controlled in both sexes by photoperiodic conditions. The induction of diapause is associated with a reversible change of body colour from green or yellow to russet (or brown). The proper timing of adult emergence and the induction of diapause, as well as the size of adults, is vitally important for successful overwintering. Nezara viridula has been shown to respond strongly to the current trend in climate change by shifting the limit of its northern range, particularly in central Japan. Analysis of historic climate data suggests that the environmental conditions during the last few decades have become more favourable for the overwintering survival of N. viridula in many locations in central Japan. This has likely promoted the northward spread of the species. The relationships between reproductive diapause, reversible body colour change, overwintering success and the recent range expansion are analyzed. Perspectives of the range dynamics of the species are discussed in light of further predicted climate change.     

Climate change trade-offs in the side-blotched lizard (Uta stansburiana): effects of growing-season length and mild temperatures on winter survival

An expanding body of literature has demonstrated that global climate change continues to adversely affect many populations, species, and ecosystems. However, life-history theory also predicts possible benefits from longer growing seasons and less severe winters, particularly for ectotherms. To test the idea that climate change will have benefits as well as costs, I studied the impacts of growing-season length on growth and overwintering conditions on survival time using side-blotched lizards (Uta stansburiana). Experiments in replicate field enclosures revealed that fall growing-season length has a direct effect on overwintering body size. Laboratory experiments revealed that both size and overwintering temperature have direct effects on winter survival time. Larger lizards are more likely to survive longer regardless of winter temperature. Furthermore, animals in colder (but still mild) winter microenvironments are more likely to survive longer than those in warmer winter environments. These results indicate that warmer winters caused by global climate change have the potential to negatively affect ectotherm populations. However, longer growing seasons may offset losses by allowing additional growth and energy storage. Thus, environmental alterations associated with climate change may be simultaneously beneficial and detrimental, and the long-term persistence of certain organisms may depend on the relative strength of their effects.     

Diapause induction and clock mechanism in the cabbage butterfly Pieris melete Ménétriés

Photoperiodic control of diapause induction was systematically investigated in the cabbage butterfly, Pieris melete, which enters summer and winter diapause as a pupa. Summer and winter diapause are induced principally by short and long scotophases, respectively; the intermediate scotophases (11-12 h) permit pupae to develop without diapause. Photoperiodic responses under 24-h light-dark cycles at 16.9, 18, 20 and 22 °C showed that the hibernation response was temperature compensated, whereas aestivation response was strongly temperature-dependent. The incidence of diapause for both aestivation and hibernation showed a decline at the ultra-short and ultra-long scotophases. Experiments using non-24-h light-dark cycles showed that the length of the scotophase played an essential role in the determination of diapause. The highest photosensitivity differed under hibernation and aestivation conditions. With a 3 × LD 12:12 interruption, a maximal inhibition of aestivation occurred in the L3/2 stage, and of hibernation it occurred in the L4/0 stage. A long-night of LD 10:14 induced hibernation diapause but inhibited aestivation diapause and, conversely, a short-night of LD 14:10 inhibited hibernation diapause but induced aestivation diapause. With a 1-h light pulse at LD 11:13, a maximal inhibition of hibernation occurred 3 h before lights-on (late scotophase), whereas, with a 1-h light pulse at LD 12.5:11.5, a maximal induction of aestivation occurred 2-3 h after the onset of darkness (early scotophase). Nanda-Hamner and Bünsow experiments failed to reveal the involvement of a circadian system, suggesting that the photoperiodic time measurement for diapause induction in this butterfly resembles an hourglass-like timer or a damped circadian oscillator.     

Combined effects of overwintering temperature and habitat degradation on the survival of boreal wood ant

The survival of insects during diapause may be affected by overwintering temperature and other environmental stress, such as anthropogenic habitat degradation. We experimentally studied the effects of overwintering temperature (+1 and +7°C) and commercial forest clear-cutting on the overwintering survival of the forest-dwelling wood ant Formica aquilonia. We found that both the higher overwintering temperature and clear-cutting lowered the overwintering survival and body fat resources of Formica aquilonia. The survival and body fat resources were highest in lower temperature treatment forest nests and lowest in higher temperature treatment clear-cut nests. The overall survival of ants increased with higher body fat resources. It is possible that both forest clear-cutting and higher winter temperature due to possible climate warming may increase the wintertime mortality of wood ants and other forest-dwelling ants.     

Timing of diapause induction and its life‐history consequences in Nezara viridula: is it costly to expand the distribution range?

Abstract. 1. From the early 1960s to 2000 Nezara viridula (Heteroptera: Pentatomidae) expanded its range northwards in Japan and reached Osaka following climate warming recorded in the region. 2. The timing of diapause induction and its effect on life‐history traits were studied under quasi‐natural conditions in Osaka. Egg masses were placed outdoors in six series in July–November 1999. Developmental events were monitored until September 2000. 3. Adult diapause was induced in September–October in agreement with the photoperiodic response obtained under laboratory conditions. Induction of diapause in early October ensured the highest winter survival. Nymphs that hatched after late September died by December–April showing that the species cannot survive winter in the nymphal stage. 4. Life‐history traits varied between the early (non‐diapause reproduction) and late (post‐diapause reproduction) series. Thus, non‐diapause females produced significantly fewer egg masses than did females that reproduced only after diapause. The timing of diapause induction strongly affected overwintering success and post‐diapause performance: females that became adults and entered diapause in October lived longer, had a longer period of oviposition, and produced more eggs in larger egg masses than females that attained adulthood and entered diapause in September. 5. Females from the early series reproduced until late November, although progeny from the late September eggs were destined to die during the winter. Pre‐winter reproduction of adults that emerged in mid‐September or later was a result of the imperfect timing of diapause induction. It is an ineffective allocation of resources and may be considered the ecological cost of range expansion. 6. To establish in the region, N. viridula will probably evolve a lengthening of the critical photoperiod of the diapause induction response. This will allow the species to enter diapause earlier and, thus, avoid maladaptive late‐autumn reproduction but, perhaps, increase the cost of diapause because of a possible adverse impact of pre‐winter high temperature conditions on overwintering.     

An assessment of direct and indirect effects of climate change for populations of the Rocky Mountain Apollo butterfly (Parnassius smintheus Doubleday)

Abstract Climate change is occurring and insects are responding. Current challenges for ecologists and managers are predicting how organisms will respond to continuing climate change and determining how to mitigate potential negative effects. In contrast to broad scale predictions for climate change involving the distribution of species, in this article we highlight the many ways in which local populations of the Rocky Mountain Apollo butterfly (Parnassius smintheus Doubleday) are predicted to respond to climate change. Using experimental and observational data collected over the past 15 years, we detail both direct and indirect effects. In addition, we identify limitations in our knowledge restricting the ability to predict how populations will respond to climate change. Some changes, such as warmer winter temperatures, may have beneficial effects; however, most of the effects of climate change will be detrimental. Variability in snow cover during the overwintering period and habitat loss due to forest encroachment have the largest potential negative effects.     

Supercooling point plasticity during cold storage in the freeze‐tolerant sugarbeet root maggot Tetanops myopaeformis

Abstract The sugarbeet root maggot Tetanops myopaeformis (Röder) overwinters as a freeze‐tolerant third‐instar larva. Although most larvae are considered to overwinter for only 1 year, some may exhibit prolonged diapause in the field. In the laboratory, they can live for over 5 years using a combination of diapause and post‐diapause quiescence. In the present study, the cold survival strategies of these larvae during storage is investigated by measuring their supercooling points in combination with survival data. Supercooling points (SCPs) change significantly during storage, highlighted by a marked increase in the range of SCPs recorded, although the ability to tolerate freezing is not affected. Additionally, a freezing event ‘re‐focuses’ the SCPs of aged larvae to levels similar to those seen at diapause initiation. This change in SCPs is dependant not only on the initial freezing event, but also on the parameters of the incubation period between freezing events. Finally, the temperatures of larval overwintering microhabitats are monitored during the 2007–2008 boreal winter. The results indicate that, although overwintering larva are physiologically freeze‐tolerant, they may essentially be freeze avoidant during overwintering via microhabitat selection.     

Climate and land-use interactively shape butterfly diversity in tropical rainforest and savanna ecosystems of southwestern China

Human-induced habitat conversion and degradation, along with accelerating climatic change, have resulted in considerable global biodiversity loss. Nevertheless, how local ecological assemblages respond to the interplay between climate and land-use change remains poorly understood. Here, we examined the effects of climate and land-use interactions on butterfly diversity in different ecosystems of southwestern China. Specifically, we investigated variation in the alpha and beta diversities of butterflies in different landscapes along human-modified and climate gradients. We found that increasing land-use intensity not only caused a dramatic decrease in butterfly alpha diversity but also significantly simplified butterfly species composition in tropical rainforest and savanna ecosystems. These findings suggest that habitat modification by agricultural activities increases the importance of deterministic processes and leads to biotic homogenization. The land-use intensity model best explained species richness variation in the tropical rainforest, whereas the climate and land-use intensity interaction model best explained species richness variation in the savanna. These results indicate that climate modulates the effects of land-use intensity on butterfly alpha diversity in the savanna ecosystem. We also found that the response of species composition to climate varied between sites: specifically, species composition was strongly correlated with climatic distance in the tropical rainforest but not in the savanna. Taken together, our long-term butterfly monitoring data reveal that interactions between human-modified habitat change and climate change have shaped butterfly diversity in tropical rainforest and savanna. These findings also have important implications for biodiversity conservation under the current era of rapid human-induced habitat loss and climate change.     

Dispersion of flightless adults of the Asian lady beetle, Harmonia axyridis, in greenhouses containing cucumbers infested with the aphid Aphis gossypii: effect of the presence of conspecific larvae

Nezara viridula (L.) (Heteroptera: Pentatomidae) recently expanded its distribution range in Japan and reached Osaka. In the southern temperate zone, the species overwinters in the adult stage and reproductive diapause is associated with a body colour change from green to russet. In Osaka, the reproductive diapause is only induced in September–October and nymphs from late egg masses are destined to die during winter. However, the fate of adults emerging late in the season remained unknown. Survival, body colour change, and post‐diapause reproduction were studied under quasi‐natural conditions in Osaka in 1999–2000 in those adults that attained adulthood as late as in November. Two experimental cohorts were used: in the Outdoor cohort, insects were reared outdoors starting in their second instar (28 September); in the Laboratory cohort, nymphs and subsequently adults were reared from the same day under diapause‐inducing conditions (L10:D14 at 25 °C), then acclimatized (5 days at 20 °C and 5 days at 15 °C; L10:D14) and transferred outdoors on 1 December. Adults in both cohorts did not reproduce in autumn and survived the winter with a low mortality. More than 20% of adults in the Outdoor cohort failed to change body colour from green to russet during winter, apparently because of the low ambient temperature, suggesting that the environmental conditions required for colour change do not completely coincide with those required for diapause induction, and that the colour of the adults is not always a reliable indicator of diapause in this species. After overwintering, females from the Outdoor cohort produced significantly fewer egg masses and eggs and had a significantly shorter period of oviposition than females that entered diapause under short‐day conditions in the Laboratory cohort. Thus, if progeny from the late egg masses attain adulthood late in the autumn, these adults have high chances of successful overwintering, but their reproductive output after the winter diapause is significantly reduced.     

Regulation of adult diapause inCoccinella septempunctata septempunctata andC. septempunctata brucki from two regions of Japan (A Minireview)

In similar climatic conditions of central/northern Europe and Hokkaido, Japan, both subspecies ofCoccinella septempunctata L. have similar life-cycles: long-day photoperiodic response ensures the induction of winter diapause. In the mild climate of Honshu, Japan,C. s. brucki Mulsant shows a different life cycle: short-day photoperiodic response leads to the induction of summer diapause while overwintering is quiescence. All populations show an important variation in photoperiodic response.     

Seasonal development and variation in abundance among four annual flight periods in a butterfly: a 20‐year study of the speckled wood (Pararge aegeria)

Insects typically spend the winter in a species‐specific diapause stage. The speckled wood butterfly, Pararge aegeria, is unique in having two alternative diapause stages, hibernating as larvae or pupae. In southern Sweden this creates a seasonal flight pattern with four annual adult flight periods: the first in May (pupal diapause), the second in June (larval diapause), and the third and fourth directly developing offspring generations in July and August, respectively. We address the raison d'être of the two diapause pathways by (1) outdoor rearing of cohorts, and (2) performing transect censuses throughout the season for 20 years. We contend that an early start of next season provides a benefit accruing to pupal diapause; conversely, a large proportion of the offspring from adults of the fourth flight peak are unable to reach the pupal stage before winter, providing a benefit accruing to larval winter diapause. The results obtained show that the two hibernation pathways are unlikely to be genetically distinct because of a strong overlap between the two offspring generations, and because sibling offspring from the third and fourth flight periods are likely to choose either of the two hibernation pathways, thereby resulting in a genetic mixing of the pathways. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 635–649.     

Insects in a warmer world: ecological, physiological and life-history responses of true bugs (Heteroptera) to climate change

Focusing on the southern green stink bug, Nezara viridula (Pentatomidae), in central Japan the effects of climate change on true bugs (Insecta: Heteroptera) are reviewed. In the early 1960s, the northern edge of the species's distribution was in Wakayama Prefecture (34.1°N) and distribution was limited by the +5°C coldest month (January) mean temperature isothermal line. By 2000, N. viridula was recorded 70 km further north (in Osaka, 34.7°N). Historical climate data were used to reveal possible causes of the northward range expansion. The increase of mean and lowest winter month temperatures by 1–2°C in Osaka from the 1950s to the 1990s improved potential overwintering conditions for N. viridula. This promoted northward range expansion of the species. In Osaka, adult diapause in N. viridula is induced after mid‐September, much later than in other local seed‐feeding heteropterans. This late diapause induction results in late‐season ineffective reproduction: some females start oviposition in autumn when the progeny have no chance of attaining adulthood and surviving winter. Both reproductive adults and the progeny die. A period from mid‐September to early November represents a phenological mismatch: diapause is not yet induced in all adults, but it is already too late to start reproduction. Females that do not start reproduction but enter diapause in September have reduced postdiapause reproductive performance: they live for a shorter period, have a shorter period of oviposition and produce fewer eggs in smaller egg masses compared with females that emerge and enter diapause later in autumn. To some extent, N. viridula remains maladapted to Osaka environmental conditions. Ecological perspectives on establishment in recently colonized areas are discussed. A review of available data suggests that terrestrial and aquatic Heteroptera species respond to climate change by shifting their distribution ranges, changing abundance, phenology, voltinism, physiology, behaviour, and community structure. Expected responses of Heteroptera to further climate warming are discussed under scenarios of slight (<2°C) and substantial (>2°C) temperature increase.     

Estivation and hibernation of Pieris brassicae (L.) in southern Spain: synchronization of two complex behavioral patterns

 The large white butterfly, Pieris brassicae, has an unusually complex life-history in its southernmost range in Western Europe. This complexity results (1) from two developmental rests, a short-day induced hibernation and a long-day induced estivation response, and (2) from the exceptionally early appearance of the first adult generation in January/February and a subsequent winter diapause in some of their progeny. It was found that in spring and autumn, different generations are faced with critical photophases which induce hibernation or estivation, with the consequence that in five out of six generations per year, only some develop directly whereas the others enter a dormancy phase. To assess the implications of this high number of optional responses on the generation succession, the development time was studied at various photoperiods and temperatures. The results showed that a threshold response determines the duration of estivation. With unchanged summer conditions (daylengths ≥15 h) estivation lasts on average 18–19 weeks, while with autumn conditions (daylengths ≤14 h) it lasts only 7 weeks. A change of photophases from ≥15 h to ≤14 h terminates estivation within about 3–5 weeks, slightly depending on the pupal exposure time in summer conditions. The duration of estivation is not affected by temperature or by the photophases experienced by the caterpillars. The winter diapause lasts 18–19 weeks on average with winter conditions (12°C/10.30 h light), but only 8–10 weeks with late spring conditions (21°C/15 h light). These results were used to assess the effects on the population phenology, with the finding that despite the different developmental pathways, a desynchronization of the generation succession is largely prevented. Estivation, hibernation, and direct development at different seasons are well adjusted to a common phenological pattern of a continuously reproducing population. This pattern of activity covers a cryptic dormant subpopulation, and could not have been deduced by field observations. Received: October 3, 2001 / Accepted: October 3, 2002     

Neonicotinoids override a parasite exposure impact on hibernation success of a key bumblebee pollinator

1. Seasonal adaptations enabling the bridging of periodic challenging abiotic conditions are taxonomically widespread. However, sensitivity to other environmental stresses can be heightened during these periods. 2. Several temperate insects with over‐wintering strategies play key ecosystem and economic roles, including wild bee pollinators. For example, hibernation survival in temperate bumblebees is crucial, as only new queens of future social colonies over‐winter. These bees are also faced with other abiotic and biotic stressors, some of which have been linked to recent pollinator declines, such as exposure to pesticides and parasites. 3. Using a fully crossed experiment, the influence of dietary exposure to neonicotinoid insecticides (thiamethoxam and clothianidin) and/or the prevalent bumblebee parasite Crithidia bombi on hibernation survival and hibernation weight change of Bombus terrestris bumblebee queens was investigated. 4. Both neonicotinoid and C. bombi exposures reduced hibernation success individually, but no additive or synergistic effects between the stressors were found. Further, effects were asynchronous, with early neonicotinoid effects on hibernation mortality overriding later parasite effects under combined exposures. Neonicotinoid exposure also increased hibernation weight loss of surviving queens. 5. Diapause periods, employed by numerous temperate organisms, are likely to be especially vulnerable to environmental stresses, besides the seasonal challenge for which these periods are an adaptation. Thus, diapause requires inclusion during the consideration of the impacts of such stresses. Accordingly, it is demonstrated here that naturally relevant exposures of pesticides and parasites have important detrimental effects on bumblebees during a critical hibernation period, with potential consequences for populations of these key wild pollinators.     

小菜蛾在温带地区越冬研究进展

小菜蛾是世界性重要害虫。在热带和亚热带地区冬季的十字花科植物上能正常发育繁殖,可见到各种虫态。但在温带冬季十字花科植物不能生长的地区,小菜蛾的越冬成为一个重要生态学问题。综述了亚洲、北美洲和欧洲小菜蛾越冬的研究进展。小菜蛾在日本的北海道、本州岛的北陆和东北大部分地区不能越冬,越冬北限相当于冬季积雪覆盖时间为60d的区域;在中国,小菜蛾不能在寒冷的东北地区越冬,在长江中下游以南地区冬季可见各虫态,但越冬北限尚不清楚;小菜蛾在冬天气候温和的韩国以及澳大利亚东南部继续发生;北美洲的加拿大西部和安大略地区大量的试验证明小菜蛾不能成功越冬,在美国南部小菜蛾冬季可正常发生,北部小菜蛾的越冬尚未见系统研究报道,但确认春季从南部运输的受小菜蛾感染的甘蓝等种苗是美国北部的重要虫源。小菜蛾在欧洲各地越冬的系统研究未见报道,没有证据表明在英国小菜蛾会发生有显著意义的越冬。目前小菜蛾越冬研究主要采用冬季直接试验观察和基于耐寒性试验的越冬预测两种方法。冬季直接试验观察法包括:(1)利用人工饲养的小菜蛾在田间各种潜在的越冬场所的越冬试验;(2)在秋播、野生或残留的十字花科植物上进行冬季种群的系统抽样调查;(3)越冬前后在前茬为十字花科植物的田块广泛搜寻普查小菜蛾的存活个体。基于耐寒性试验的越冬预测法:在获取小菜蛾越冬场所温度的基础上设计低温处理模式,试验低温处理后小菜蛾的存活率及后续发育和生殖。将试验数据和各地气温或小气候相结合,对小菜蛾在的越冬可能性进行推断。     

How climate change affects the seasonal ecology of insect parasitoids

1. In the context of global change, modifications in winter conditions may disrupt the seasonal phenology patterns of organisms, modify the synchrony of closely interacting species and lead to unpredictable outcomes at different ecological scales. 2. Parasites are present in almost every food web and their interactions with hosts greatly contribute to ecosystem functioning. Among upper trophic levels of terrestrial ecosystems, insect parasitoids are key components in terms of functioning and species richness. Parasitoids respond to climate change in similar ways to other insects, but their close relationship with their hosts and their particular life cycle – alternating between parasitic and free-living forms – make them special cases. 3. This article reviews of the mechanisms likely to undergo plastic or evolutionary adjustments when exposed to climate change that could modify insect seasonal strategies. Different scenarios are then proposed for the evolution of parasitoid insect seasonal ecology by exploring three anticipated outcomes of climate change: (i) decreased severity of winter cold; (ii) decreased winter duration; and (iii) increased extreme seasonal climatic events and environmental stochasticity. 4. The capacities of insects to adapt to new environmental conditions, either through plasticity or genetic evolution, are highlighted. They may reduce diapause expression, adapt to changing cues to initiate or terminate diapause, increase voltinism, or develop overwintering bet-hedging strategies, but parasitoids' responses will be highly constrained by those of their hosts. 5. Changes in the seasonal ecology of parasitoids may have consequences on host–parasitoid synchrony and population cycles, food-web functioning, and ecosystem services such as biological pest control.     

Are aphid parasitoids from mild winter climates losing their winter diapause?

Temperature is both a selective pressure and a modulator of the diapause expression in insects from temperate regions. Thus, with climate warming, an alteration of the response to seasonal changes is expected, either through genetic adaptations to novel climatic conditions or phenotypic plasticity. Since the 1980s in western France, the winter guild of aphid parasitoids (Hymenoptera: Braconidae) in cereal fields has been made up of two species: Aphidius rhopalosiphi and Aphidius matricariae. The recent activity of two other species, Aphidius avenae and Aphidius ervi, during the winter months suggests that a modification of aphid parasitoid overwintering strategies has taken place within the guild. In this study, we first performed a field survey in the winter of 2014/15 to assess levels of parasitoid diapause incidence in agrosystems. Then, we compared the capacity of the four parasitoid species to enter winter diapause under nine different photoperiods and temperature conditions in the laboratory. As predicted, historically winter-active species (A. rhopalosiphi and A. matricariae) never entered diapause, whereas the species more recently active during winter (A. avenae and A. ervi) did enter diapause but at a low proportion (maximum of 13.4 and 11.2%, respectively). These results suggest rapid shifts over the last three decades in the overwintering strategies of aphid parasitoids in Western France, probably due to climate warming. This implies that diapause can be replaced by active adult overwintering, with potential consequences for species interactions, insect community composition, ecosystem functioning, and natural pest control.     

Role of natural day-length and temperature in determination of summer and winter diapause in Pieris melete (Lepidoptera: Pieridae)

Under field conditions, the cabbage butterfly, Pieris melete, displays a pupal summer diapause in response to relatively low daily temperatures and gradually increasing day-length during spring and a pupal winter diapause in response to the progressively shorter day-length. To determine whether photoperiod is 'more' important than temperature in the determination of summer and winter diapause, or vice versa, the effects of naturally changing day-length and temperature on the initiation of summer and winter diapause were systematically investigated under field conditions for five successive years. Field results showed that the incidence of summer diapause significantly declined with the naturally increasing temperature in spring and summer generations. Path coefficient analysis showed that the effect of temperature was much greater than photoperiod in the determination of summer diapause. In autumn, the incidence of diapause was extremely low when larvae developed under gradually shortening day-length and high temperatures. The incidence of winter diapause increased to 60-90% or higher with gradually shortening day-length combined with temperatures between 20.0°C and 22.0°C. Decreasing day-length played a more important role in the determination of winter diapause induction than temperature. The eco-adaptive significance of changing day-length and temperature in the determination of summer and winter diapause was discussed.