Variation in the phenology of photosynthesis among eastern white pine provenances in response to warming

In higher‐latitude trees, temperature and photoperiod control the beginning and end of the photosynthetically active season. Elevated temperature (ET) has advanced spring warming and delayed autumn cooling while photoperiod remains unchanged. We assessed the effects of warming on the length of the photosynthetically active season of three provenances of Pinus strobus L. seedlings from different latitudes, and evaluated the accuracy of the photochemical reflectance index (PRI) and the chlorophyll/carotenoid index (CCI) for tracking the predicted variation in spring and autumn phenology of photosynthesis among provenances. Seedlings from northern, local and southern P. strobus provenances were planted in a temperature‐free‐air‐controlled enhancement (T‐FACE) experiment and exposed to ET (+1.5/3°C; day/night). Over 18 months, we assessed photosynthetic phenology by measuring chlorophyll fluorescence, gas exchange, leaf spectral reflectance and pigment content. During autumn, all seedlings regardless of provenance followed the same sequence of phenological events with the initial downregulation of photosynthesis, followed by the modulation of non‐photochemical quenching and associated adjustments of zeaxanthin pool sizes. However, the timing of autumn downregulation differed between provenances, with delayed onset in the southern provenance (SP) and earlier onset in the northern relative to the local provenance, indicating that photoperiod at the provenance origin is a dominant factor controlling autumn phenology. Experimental warming further delayed the downregulation of photosynthesis during autumn in the SP. A provenance effect during spring was also observed but was generally not significant. The vegetation indices PRI and CCI were both effective at tracking the seasonal variations of energy partitioning in needles and the differences of carotenoid pigments indicative of the stress status of needles. These results demonstrate that PRI and CCI can be useful tools for monitoring conifer phenology and for the remote monitoring of the length of the photosynthetically active season of conifers in a changing climate.     

Larval diapause of Monochamus urussovi and photoperiodic effects on larval development in tree bolts

To determine the larval diapause and the effect of photoperiod on development in Monochamus urussovi (Coleoptera: Cerambycidae), larvae were reared on Abies sachalinensis and Picea jezoensis logs and bolts. Larvae stopped developing in the final instar at 25°C and 16L : 8D (16 h light and 8 h dark) whereas an exposure to 5°C in the dark (134 days) following acclimation at 12°C under natural daylength led to adult emergence. When larvae were reared under 8L : 16D or 16L : 8D at 25°C with an intervening period of chilling at 5°C in the dark (112 days), a photoperiod of 8L : 16D induced a shorter time required for adult emergence after being returned to 25°C, and smaller adult body size than 16L : 8D.     

Effects of temperature and other environmental factors on the post‐diapause development of walnut husk fly Rhagoletis completa (Diptera: Tephritidae)

To more effectively manage walnut husk fly Rhagoletis completa (Diptera: Tephritidae), in California walnut orchards, it is important to understand the factors that affect the timing of adult emergence. In the present study, we examine the effects of incubation temperature, pre‐chill and chill durations, latitude, cultivar and size on the post‐diapause development of R. completa puparia. The lower developmental threshold, upper developmental threshold and optimal temperature for puparial development are estimated to be 4, 34 and 26.6 °C, respectively. The thermal requirement for adult emergence after 120 days of chilling is estimated to be 2024 degree days. Percentage adult emergence declines at both higher and lower incubation temperatures. Chill duration at 5 °C for diapausing puparia has a nonlinear negative effect on the thermal requirement but no effect on percentage emergence. Insufficient chilling leads to poor synchronization of adult emergence. Greater pre‐chill duration at room temperature increases the thermal requirement and slightly decreased percentage emergence. Latitude has a negative effect on the thermal requirement. Puparia from northern California black walnut (Juglans hindsii) have a slightly greater thermal requirement than puparia from cultivated walnut (Julgans regia). There is no significant difference in puparial fresh weight or mean thermal requirement between males and females, although the positive correlation between thermal requirement and puparial fresh weight is stronger for females than males. The effects of temperature and other environmental factors on the post‐diapause development of R. completa are discussed in relation to observations from other Rhagoletis species.     

From observations to experiments in phenology research: investigating climate change impacts on trees and shrubs using dormant twigs

Background and Aims Climate change is advancing the leaf-out times of many plant species and mostly extending the growing season in temperate ecosystems. Laboratory experiments using twig cuttings from woody plant species present an affordable, easily replicated approach to investigate the relative importance of factors such as winter chilling, photoperiod, spring warming and frost tolerance on the leafing-out times of plant communities. This Viewpoint article demonstrates how the results of these experiments deepen our understanding beyond what is possible via analyses of remote sensing and field observation data, and can be used to improve climate change forecasts of shifts in phenology, ecosystem processes and ecological interactions.Scope The twig method involves cutting dormant twigs from trees, shrubs and vines on a single date or at intervals over the course of the winter and early spring, placing them in containers of water in controlled environments, and regularly recording leaf-out, flowering or other phenomena. Prior to or following leaf-out or flowering, twigs may be assigned to treatment groups for experiments involving temperature, photoperiod, frost, humidity and more. Recent studies using these methods have shown that winter chilling requirements and spring warming strongly affect leaf-out and flowering times of temperate trees and shrubs, whereas photoperiod requirements are less important than previously thought for most species. Invasive plant species have weaker winter chilling requirements than native species in temperate ecosystems, and species that leaf-out early in the season have greater frost tolerance than later leafing species.Conclusions This methodology could be extended to investigate additional drivers of leaf-out phenology, leaf senescence in the autumn, and other phenomena, and could be a useful tool for education and outreach. Additional ecosystems, such as boreal, southern hemisphere and sub-tropical forests, could also be investigated using dormant twigs to determine the drivers of leaf-out times and how these ecosystems will be affected by climate change.     

Use of RAPD patterns for clone verification and in studying provenance relationships in Norway spruce (Picea abies)

 We have used the RAPD technique to analyse samples of Picea abies obtained from an improvement forestry station. Two types of plant material were harvested, the first being clones and the second provenances from various regions. We first checked the clonal identity of elite tree cuttings and clones; some differences in the RAPD patterns resulting from mis-planting or mis-labelling of cuttings were found. We also established a reference library of RAPD fingerprints for 96 clones, which will serve as a reference source in cases of litigation concerning clone identity. The RAPD technique was also used to study the genetic relationship between nine European provenances of Norway spruce. A dendogram was obtained by individual pairwise comparison of 42 RAPD bands, which separated the nine provenances into two major groups, one containing the Nordic provenances (Sweden and Bielorussia) and another the Alpine provenances (France, Austria, Germany and Belgium). The Belgian provenance, which is not indigenous, is most closely related to the German provenance. We conclude that the RAPD technique is a useful tool for forestry stations in managing propagation operations. Received: 15 June 1996 / Accepted: 11 October 1996     

Phenotypic plasticity in Cardamine flexuosa: variation among populations in plastic response to chilling treatments and photoperiods

Variability in the plastic responses of seven life history traits to different chilling and photoperiod regimes was studied in four wild populations of Cardamine flexuosa. This species, a winter-green or year-long annual, showed a facultative long-day and chilling requirement for flowering. Considerable variation among populations was noted in plasticity of all traits. Differences in plasticity were greater among three paddy field populations from different climatic areas than between adjacent populations under different disturbance regimes. A paddy field population (OP) and an adjacent orchard population (OG) exhibited similar plasticity, in both amounts and patterns of response. TP, a Japan Sea coast population, was distinct from three other populations, especially in the small amounts of plasticity. Differences in amount of response were much more common than differences in pattern of response. Character expressions of five traits were significantly correlated with the number of days to flowering. Days to flowering and the numbers of inflorescences and siliques showed high negative correlations because the branching ability of meristems decreased with delay of flowering.     

Will phenotypic plasticity affecting flowering phenology keep pace with climate change?

Rising temperatures have begun to shift flowering time, but it is unclear whether phenotypic plasticity can accommodate projected temperature change for this century. Evaluating clines in phenological traits and the extent and variation in plasticity can provide key information on assessing risk of maladaptation and developing strategies to mitigate climate change. In this study, flower phenology was examined in 52 populations of big sagebrush (Artemisia tridentata) growing in three common gardens. Flowering date (anthesis) varied 91 days from late July to late November among gardens. Mixed‐effects modeling explained 79% of variation in flowering date, of which 46% could be assigned to plasticity and genetic variation in plasticity and 33% to genetics (conditional R² = 0.79, marginal R² = 0.33). Two environmental variables that explained the genetic variation were photoperiod and the onset of spring, the Julian date of accumulating degree‐days >5 °C reaching 100. The genetic variation was mapped for contemporary and future climates (decades 2060 and 2090), showing flower date change varies considerably across the landscape. Plasticity was estimated to accommodate, on average, a ±13‐day change in flowering date. However, the examination of genetic variation in plasticity suggests that the magnitude of plasticity could be affected by variation in the sensitivity to photoperiod and temperature. In a warmer common garden, lower‐latitude populations have greater plasticity (+16 days) compared to higher‐latitude populations (+10 days). Mapped climatypes of flowering date for contemporary and future climates illustrate the wide breadth of plasticity and large geographic overlap. Our research highlights the importance of integrating information on genetic variation, phenotypic plasticity and climatic niche modeling to evaluate plant responses and elucidate vulnerabilities to climate change.     

Autumn photosynthetic decline and growth cessation in seedlings of white spruce are decoupled under warming and photoperiod manipulations

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high‐latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down‐regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change.     

Control of diapause in codling moth larvae

Diapause in a New Zealand strain of codling moth (Cydia pomonella Linnaeus [Lepidoptera: Olethreutidae]) was induced in larvae by photoperiods of 15 h or less. Once diapause had been initiated, it could not be terminated by any combination of conditions tested for at least 20 days after cocooning. In diapausing larvae a low rate of pupation occurred at 25 °C under a long day (18 h) photoperiod. A high rate of pupation was achieved under a long day regime when larvae were decocooned, and provided with apple as nourishment. Diapause could be terminated predictably in 94–100% of larvae by 1) conditioning at 15 °C and constant darkness for periods of 40–100 days, then 2) chilling at 2±2 °C and constant darkness for 20–50 days followed by 3) any post-chill condition periods at 25 °C, 18 h photoperiod. Complete diapause termination was achieved when 100 days conditioning was followed by 30 days or 50 days post-chill period. Under these conditions, 76% termination occurred in the post-chill period after 10 days, and 93% after 25 days.To terminate diapause in codling moth larvae, we recommend that a 100 days conditioning followed by 30 days chilling and 50 days post chilling periods be used.     

Inflorescence commitment and subsequent development differ in their responses to temperature and photoperiod in Osteospermum jucundum

Although Osteospermum is a species which is known to require a period of chilling to induce flowering, the precise form of the relationships between temperature and photoperiod on the phases of flowering has not been quantified. This study aimed to investigate the effects of temperature and photoperiod on time to inflorescence commitment and on the rate of subsequent floral development in Osteospermum jucundum cv. Zulu. To assess how temperature and photoperiod affected the number of days needed for inflorescence commitment, plants were transferred from a range of photothermal environments to a non‐inductive environment. The effect of temperature and photoperiod on subsequent inflorescence development was examined by transferring plants with initiated inflorescences to a range of photothermal environments. Inflorescence commitment occurred first in plants grown at a low average diurnal temperature (10.6°C), but no evidence was found to suggest that photoperiod affected the duration of this phase. Once initiated, high temperatures and long days hastened inflorescence development. The rate of progress to flowering from initiation increased linearly with photoperiod and temperature (up to an optimum of 23.5°C).     

Latitudinal variation in life-cycle characteristics of Potamogeton pectinatus L.: vegetative growth and asexual reproduction

Across latitudinal gradients, environmental conditions that influenceplant growth and reproduction largely change. Here we study clonal variation inlife-cycle characteristics of the cosmopolitan water plantPotamogetonpectinatus L. across a broad latitudinal range.Two consecutive experiments were performed under standardised laboratoryconditions (photoperiod, irradiance and temperature). In the first experimentweinvestigated asexual reproduction among fifteen clones, obtained from latitudesranging from 24 to 68° N. After 90 days of growth, high-latitude clonesproduced more but smaller tubers, while the aboveground biomass was lower ascompared to the clones obtained from low latitudes.In a second experiment we studied inherent differences in early growth,morphology and photosynthesis for eleven clones (obtained from the samelatitudinal range as in experiment 1). We found high among clonal variation formost measured variables, but the number of latitude-correlated traits waslimited. The only trait that correlated with latitude was the number of leavesper plant, which increased in clones from higher latitudes.Our results agree with the hypothesis of a latitude-correlated switch inlife-cycle strategy for this species. For northern clones this results in ashort life-cycle, with an early and high investment in tuber biomass, while forlow latitude clones the length of the life-cycle is prolonged, with a delayedreproduction and increased total plant biomass.     

Effects of growth temperature and winter duration on leaf phenology of a spring ephemeral (Gagea lutea) and a summergreen forb (Maianthemum dilatatum)

Effects of growth temperature and winter duration on leaf longevity were compared between a spring ephemeral, Gagea lutea, and a forest summergreen forb, Maianthemum dilatatum. The plants were grown at day/night temperatures of 25/20°C and 15/10°C after a chilling treatment for variable periods at 2°C. The temperature regime of 25/20°C was much higher than the mean air temperatures for both species in their native habitats. Warm temperature of 25/20°C and/or long chilling treatment shortened leaf longevity in G. lutea, but not in M. dilatatum. The response of G. lutea was consistent with that reported for other spring ephemerals. Air temperature increases as the vegetative season progresses. The decrease in leaf longevity in G. lutea under warm temperature condition ensures leaf senescence in summer, an unfavorable season for its growth. This also implies that early leaf senescence could occur in years with early summers. Warm spring temperatures have been shown to accelerate the leafing-out of forest trees. The decrease in leaf longevity due to warm temperature helps synchronize the period of leaf senescence roughly with the time of the forest canopy leaf-out. Prolonged winter due to late snowmelt has been shown to shorten the vegetative period for spring ephemerals. The decrease in leaf longevity due to long chilling treatment would correspond with this shortened vegetative period.     

Chilling and forcing temperatures interact to predict the onset of wood formation in Northern Hemisphere conifers

The phenology of wood formation is a critical process to consider for predicting how trees from the temperate and boreal zones may react to climate change. Compared to leaf phenology, however, the determinism of wood phenology is still poorly known. Here, we compared for the first time three alternative ecophysiological model classes (threshold models, heat‐sum models and chilling‐influenced heat‐sum models) and an empirical model in their ability to predict the starting date of xylem cell enlargement in spring, for four major Northern Hemisphere conifers (Larix decidua, Pinus sylvestris, Picea abies and Picea mariana). We fitted models with Bayesian inference to wood phenological data collected for 220 site‐years over Europe and Canada. The chilling‐influenced heat‐sum model received most support for all the four studied species, predicting validation data with a 7.7‐day error, which is within one day of the observed data resolution. We conclude that both chilling and forcing temperatures determine the onset of wood formation in Northern Hemisphere conifers. Importantly, the chilling‐influenced heat‐sum model showed virtually no spatial bias whichever the species, despite the large environmental gradients considered. This suggests that the spring onset of wood formation is far less affected by local adaptation than by environmentally driven plasticity. In a context of climate change, we therefore expect rising winter–spring temperature to exert ambivalent effects on the spring onset of wood formation, tending to hasten it through the accumulation of forcing temperature, but imposing a higher forcing temperature requirement through the lower accumulation of chilling.     

Chilling outweighs photoperiod in preventing precocious spring development

It is well known that increased spring temperatures cause earlier onset dates of leaf unfolding and flowering. However, a temperature increase in winter may be associated with delayed development when species' chilling requirements are not fulfilled. Furthermore, photosensitivity is supposed to interfere with temperature triggers. To date, neither the relative importance nor possible interactions of these three factors have been elucidated. In this study, we present a multispecies climate chamber experiment to test the effects of chilling and photoperiod on the spring phenology of 36 woody species. Several hypotheses regarding their variation with species traits (successional strategy, floristic status, climate of their native range) were tested. Long photoperiods advanced budburst for one‐third of the studied species, but magnitudes of these effects were generally minor. In contrast to prior hypotheses, photosensitive responses were not restricted to climax or oceanic species. Increased chilling length advanced budburst for almost all species; its effect greatly exceeding that of photoperiod. Moreover, we suggest that photosensitivity and chilling effects have to be rigorously disentangled, as the response to photoperiod was restricted to individuals that had not been fully chilled. The results indicate that temperature requirements and successional strategy are linked, with climax species having higher chilling and forcing requirements than pioneer species. Temperature requirements of invasive species closely matched those of native species, suggesting that high phenological concordance is a prerequisite for successful establishment. Lack of chilling not only led to a considerable delay in budburst but also caused substantial changes in the chronological order of species' budburst. The results reveal that increased winter temperatures might impact forest ecosystems more than formerly assumed. Species with lower chilling requirements, such as pioneer or invasive species, might profit from warming winters, if late spring frost events would in parallel occur earlier.     

Induction of adult diapause in Pyrrhocoris apterus L. by short cold exposure

Summary The ripening of ovaries in the females of Pyrrhocoris apterus can be inhibited not only by short-day photoperiod but also by cold shocks during the first week of the adult life.Ovaries in a part of the females are prevented from ripening by transferring the bugs from 25° C and long-day photoperiod to substantially lower temperatures for 2 (or 4) days. The oviposition is inhibited at least for 3 weeks after the return to original conditions favourable for reproduction. This effect is achieved when cold treatment is combined either with darkness or with long day. A difference between 2 and 4 days of chilling was not recorded. The effect of cold exposure (evaluated in our experiments by the proportion of mature and immature females) depends on the temperature of the cold exposure and on the age of females at the time of transfer to cold. The incidence of immature females was higher when lower temperatures (+2 to +6° C) were used and when females were transferred to cold early (0–2 days) after the adult moult.     

Passerines may be sufficiently plastic to track temperature‐mediated shifts in optimum lay date

Projecting the fates of populations under climate change is one of global change biology's foremost challenges. Here, we seek to identify the contributions that temperature‐mediated local adaptation and plasticity make to spatial variation in nesting phenology, a phenotypic trait showing strong responses to warming. We apply a mixed modeling framework to a Britain‐wide spatiotemporal dataset comprising >100 000 records of first egg dates from four single‐brooded passerine bird species. The average temperature during a specific time period (sliding window) strongly predicts spatiotemporal variation in lay date. All four species exhibit phenological plasticity, advancing lay date by 2–5 days °C^?1. The initiation of this sliding window is delayed further north, which may be a response to a photoperiod threshold. Using clinal trends in phenology and temperature, we are able to estimate the temperature sensitivity of selection on lay date (B), but our estimates are highly sensitive to the temporal position of the sliding window. If the sliding window is of fixed duration with a start date determined by photoperiod, we find B is tracked by phenotypic plasticity. If, instead, we allow the start and duration of the sliding window to change with latitude, we find plasticity does not track B, although in this case, at odds with theoretical expectations, our estimates of B differ across latitude vs. longitude. We argue that a model combining photoperiod and mean temperature is most consistent with current understanding of phenological cues in passerines, the results from which suggest that each species could respond to projected increases in spring temperatures through plasticity alone. However, our estimates of B require further validation.     

Dormancy, Turion Formation, and Germination by Different Clones of Spirodela polyrrhiza

Some clones of Spirodela polyrrhiza form dormant bodies called turions which require several weeks of chilling treatment before they proceed to renew growth and develop into vegetative fronds. The individual fronds of Spirodela are less than 5 mm long and can be grown aseptically in liquid culture. Turion formation and germination can serve as a bioassay for the various compounds involved in dormancy development.

Turion formation can be induced by manipulation of light intensity during the day, photoperiod, night temperature, day temperature, and concentration of nitrate in the culture medium. Different clones of Spirodela from northeastern United States, Puerto Rico, and Argentina had different requirements for turion formation. The clones from Argentina and Puerto Rico did not form turions under any of the experimental conditions imposed. Turions of some clones required chilling treatments for renewed vegetative growth while others did not. Both gibberellic acid and long photoperiods were required to bypass the chilling requirements of some clones, but not others.

     

STAMINATE BLOOM DATE AND TEMPERATURE RESPONSES OF POLLEN GERMINATION AND TUBE GROWTH IN TWO WALNUT (JUGLANS) SPECIES

Pollen was collected from ten cultivars of Juglans regia and three cultivars of the later-blooming species, J. nigra. Extensive phenological data were available for these cultivars. Cultivars were chosen on the basis of staminate bloom date to include the earliest and latest blooming individuals available and a representative range throughout the bloom season. Mean staminate bloom dates for the cultivars examined covered a period of 46 days over which time mean daily temperatures rose 6 C. In order to determine if adaptations to temperature were expressed by the gametophyte generation, pollen was subjected to controlled temperatures from 5 to 40 C in 2 to 4 C increments and analyzed for germination percentages and pollen tube elongation. A positive relationship was found for pollen germination percentage and mean staminate bloom date such that earlier blooming individuals showed lower minimum temperature thresholds for germination, and optimum temperature for pollen germination was positively correlated with mean staminate bloom date. Differences in pollen tube growth, determined separately from hydration and germination responses, were less clear. Most J. regia clones had lower minium temperature thresholds for growth than the J. nigra clones, but there was no clear relationship to earliness of bloom within the species. No differences were discerned in optimum temperatures for pollen tube growth either between the two species or within species.     

Influences of daylength and temperature on the period of diapause and its ending process in dormant larvae of burnet moths (Lepidoptera,Zygaenidae)

The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3–10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of 5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30–180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120–180 days, molting to the next instar occurred after 6–10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6–10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30–90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10–25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10–40 days and had molted after the 6–10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.