Geographic variation in diapause induction and termination of the cotton bollworm, Helicoverpa armigera Hübner (Lepidoptera: Noctuidae)

Overwintering diapause in Helicoverpa armigera, a multivoltine species, is controlled by response to photoperiod and temperature. Photoperiodic responses from 5 different geographical populations showed that the variation in critical photoperiod for diapause induction was positively related to the latitudinal origin of the populations at 20, 22 and 25 °C. Diapause response to photoperiod and temperature was quite different between northern and southern populations, being highly sensitive to photoperiod in northern populations and temperature dependence in southern populations. Diapause pupae from southern population showed a significantly shorter diapause duration than from northern-most populations when they were cultured at 20, 22, 25, 28 and 31 °C; by contrast, overwintering pupae from southern populations emerged significantly later than from northern populations when they were maintained in natural conditions, showing a clinal latitudinal variation in diapause termination. Diapause-inducing temperature had a significant effect on diapause duration, but with a significant difference between southern and northern populations. The higher rearing temperature of 22 °C evoked a more intense diapause than did 20 °C in northern populations; but a less intense diapause in southern population. Cold exposure (chilling) is not necessary to break the pupal diapause. The higher the temperature, the quicker the diapause terminated. Response of diapause termination to chilling showed that northern populations were more sensitive to chilling than southern population.     

Repeated cycles of chilling and warming effectively terminate prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis (Coleoptera: Curculionidae) requires one or more years to complete its life cycle, owing to prolonged larval diapause. To compare the effects of temperature cycles and total periods of chilling on the termination of prolonged diapause, larvae were subjected to different chilling (5 degrees C) and warming (20 degrees C) cycles ranging from 30 to 720 days, and all cycles were repeated until the sum of chilling and warming periods reached 720 days. The prolonged diapause of C. sikkimensis was more effectively terminated by repeated cycles of chilling and warming than by prolonging the continuous chilling period. However, extremely short temperature cycles were not highly effective in enhancing diapause termination, even when such cycles were repeated many times. To examine the role of warming periods on diapause termination, diapause larvae were subjected to a sequence of chilling (120 days at 5 degrees C) and warming (240 days at 20 degrees C) with a warming period (0-120 days at 20 degrees C) inserted in the chilling period. Diapause larvae that were not reactivated in the first chilling period required exposure to a certain period of warming before they were able to complete diapause development in the subsequent chilling. Thus, C. sikkimensis appears to spread its reactivation times over several years in response to seasonal temperature cycles.     

Diapause termination, post-diapause development and reproduction in the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae)

Effects of photoperiod and cold exposure on diapause termination, post-diapause development and reproduction in Loxostege sticticalis were examined. Larvae were reared at diapause inducing condition (22 °C, L:D 12:12) consistently or transferred to long day photoperiod (L:D 16:8) and darkness (L:D 0:24) respectively, after entering into diapause. Diapause was terminated in approximately 40% of the larvae after 36 days, and no significant differences were observed between photoperiods, suggesting larval diapause was terminated spontaneously without being induced by photoperiods. Cold exposure significantly hastened diapause termination. The diapause termination incidence increased significantly with peaks of 98% at both 5 °C and 0 °C exposure for 30 days, as compared to 42% in controls not exposed to cold, while the mortality and number of days required for diapause termination decreased dramatically. The optimal low temperature exposure periods under 5 °C or 0 °C were 20 days and 30 days, showing a higher termination incidence and shorter time for diapause termination. This suggests that the low temperatures in winter play an important role in diapause termination under natural conditions. The threshold temperatures for post-diapause development in prepupae and pupae were 9.13 °C and 10.60 °C respectively, with corresponding accumulations of 125 and 200 degree-days. Adults that experienced larval diapause significantly delayed their first oviposition, oviposition period was prolonged, and the lifetime number of eggs laid was decreased, however both males and females have significantly longer longevity. The field validation of diapause termination, the degree-days model, and the relationship between diapause and migration in L. sticticalis were also discussed.     

Effect of temperature on the termination of prolonged larval diapause in the chestnut weevil Curculio sikkimensis (Coleoptera: Curculionidae)

Adults of the chestnut weevil Curculio sikkimensis emerged over a 3-year period under laboratory and quasi-field conditions due to a prolonged diapause that occurred at the mature larval stage. Variable proportions of the larvae remained in diapause after a single cold (5 degrees C) treatment of 120 days. Extension of the chilling period to as long as 540 days did not increase the percentage of diapause termination, and excessively long chilling actually reduced the percentage. Chilling was not indispensable to the termination of larval diapause. Diapause intensity was very high and variable, and more than 1000 days at 20 degrees C was necessary to reactivate all diapause larvae. When the diapause larvae were exposed to cycles of low (5 degrees C for 120 days) and high (20 degrees C for 240 days) temperatures, the percentage of diapause termination reached 100% after two or three such cycles. Thus, the prolonged diapause of C. sikkimensis has characteristics similar to the common short winter diapause in other insects, but has unique characteristics that ensure polymodal reactivation over several years.     

Evidence for reversible change in intensity of prolonged diapause in the chestnut weevil Curculio sikkimensis

The chestnut weevil Curculio sikkimensis undergoes a prolonged larval diapause that is completed by repeated exposure to chilling and warming. We examined the possible reversibility of diapause intensity in response to temperature changes. All larvae were subjected to an initial chilling followed by incubation at 20°C to force pupation of the 1-year-type larvae that require only one winter for diapause completion. We then exposed the larvae remaining in prolonged diapause to a second chilling at 5°C for different lengths of time, preceded or not preceded by incubation at 20°C (moderately high) and/or 25°C (high) and followed by a final post-chilling reincubation at 20°C. Many of the prolonged-diapausing larvae subjected only to a brief second chilling were re-activated upon reincubation. However, short exposure to 25°C before this second chilling dramatically decreased the percentage of larvae completing diapause. When larvae were exposed to 25°C for a short period, then incubated at 20°C and subjected to the brief second chilling, many were re-activated during reincubation. The chilling time required for most of the larvae to complete diapause decreased after pre-chilling incubation at 20°C and increased after incubation at 25°C. These results demonstrate that diapause intensity in C. sikkimensis changes reversibly in response to changes in ambient temperature.     

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.     

Diapause avoidance induced by low temperature in the yellow-spotted longicorn beetle, Psacothea hilaris

The diapause-averting effect of low temperature on pre-diapause larvae was examined in the yellow-spotted longicorn beetle, Psacothea hilaris. Larvae that had been reared under diapause-inducing conditions (25 °C , L12:D12) were temporarily exposed to 10 °C for various periods, and returned to the initial condition. Diapause was not averted by chilling for 15 days irrespective of the age of the larvae at chilling. After a 30-day chilling treatment, all of the 40- and 60-day-old larvae averted diapause, while diapause was averted in only one-third of the 10- and 20-day-old larvae. None of the pre-diapause larvae chilled for 60 days entered diapause irrespective of the age at chilling. With diapause avoidance, larvae that overwintered in earlier instars can start growing in earliest spring without any arrest; this phenomenon probably subserves the synchronization of larval development in a population.     

Effects of temperature during chilling and pre-chilling periods on diapause and post-diapause development in a katydid, Eobiana engelhardti subtropica

In Eobiana engelhardti subtropica, early laid eggs reach the diapause stage in early autumn. For long periods before winter, the eggs are exposed to temperatures higher than their theoretical lower threshold for development. In contrast, late-laid eggs cannot reach their diapause stage before winter. Our study showed that E. e. subtropica copes with these difficulties via the thermal response involving embryonic diapause. In this katydid, the almost fully developed embryo undergoes an obligatory diapause. When diapause eggs were maintained at a temperature of 20 degrees C or higher, diapause persisted for a long time. Diapause was effectively terminated by temperatures ranging from 1 to 11 degrees C, and hatching occurred successfully at temperatures from 11 to 15 degrees C. In addition to the chilling temperature, pre-chilling temperature modified diapause intensity and hatching time. Diapause eggs hatched earlier after chilling when the pre-chilling temperature was lower, within a range of 14.5-25 degrees C. Thus, the low-temperature requirement for diapause termination prevents early laid eggs from untimely hatching in autumn, and low temperatures before and during winter decrease diapause intensity and shorten the hatching time in the following spring. When eggs were chilled before diapause, they tolerated chilling and averted diapause. Thus, even if eggs encounter low temperatures before diapause, they can hatch in the following spring.     

Diapause pupal color diphenism induced by temperature and humidity conditions in Byasa alcinous (Lepidoptera: Papilionidae)

We investigated whether diapause pupae of Byasa alcinous exhibit pupal color diphenism (or polyphenism) similar to the diapause pupal color polyphenism shown by Papilio xuthus. All diapause pupae of B. alcinous observed in the field during winter showed pupal coloration of a dark-brown type. When larvae were reared and allowed to reach pupation under short-day conditions at 18 °C under a 60 ± 5% relative humidity, diapause pupae exhibited pupal color types of brown (33%), light-brown (25%), yellowish-brown (21%), diapause light-yellow (14%) and diapause yellow (7%). When mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C and 25 °C under a 60 ± 5% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 91.2, 8.8 and 0.0% at 10 °C, and 12.2, 48.8 and 39.0% at 25 °C, respectively. On the other hand, when mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C, 18 °C and 25 °C under an over 90% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 79.8, 16.9 and 3.3% at 10 °C, 14.5, 26.9 and 58.6% at 18 °C, and 8.3, 21.2 and 70.5% at 25 °C, respectively. These results indicate that diapause pupae of brown types are induced by lower temperature and humidity conditions, whereas yellow types are induced by higher temperature and humidity conditions. The findings of this study show that diapause pupae of B. alcinous exhibit pupal color diphenism comprising brown and diapause yellow types, and suggest that temperature and humidity experienced after a gut purge are the main factors that affect the diapause pupal coloration of B. alcinous as environmental cues.     

Reversible change in embryonic diapause intensity by mild temperature in the Chinese rice grasshopper, Oxya chinensis

The effects of temperature on maintenance and termination of embryonic diapause were investigated in Jining (35.4°N, 116.6°E) and Sihong (33.5°N, 118.2°E) strains of the Chinese rice grasshopper, Oxya chinensis Thunberg (Orthoptera: Catantopidae). Eggs of both strains entered diapause when incubated at 30, 25, or 20 °C. Chilling at 8 °C had an evident effect on diapause termination and almost all eggs chilled for 60 days ended diapause development. Chilling of eggs at 8 °C for only 20 days failed to result in any hatching at 20 °C, suggesting that such level of chilling was not enough to induce diapause termination. However, the treatment combining incubation of eggs at 30 °C for varying lengths of time with subsequent incubation to 20 °C had a distinct effect on the completion of diapause of the eggs. The results indicate that there were two temperature optima, that is, low temperature (chilling) and high temperature, for diapause development in this grasshopper species. Incubation of chilled eggs at 20 °C for 5–15 days followed by further incubation at 25 °C reduced termination of diapause significantly compared with the eggs only chilled at 8 °C. Exposure of eggs chilled at 8 °C to a pulse of 25 °C from 1 to 7 days, separated by a 20-day interval at 8 °C, resulted in a decrease in the percentage of successfully hatched eggs as the length of the pulse of 25 °C increased. The results suggest that diapause intensity may be restored at moderately high temperatures. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Thermal response and reversibility of diapause in the eggs of Locusta migratoria

Abstract. Eggs of a local population of Locusta migratoria L. (Orthoptera: Acrididae) collected near Hirosaki (40.5^oN) entered diapause when incubated at temperatures between 20 and 35^oC. For diapause development the optimum temperature was 10^oC. The lower thermal threshold for post-diapause development was 14.7^oC. After chilling at 10^oC for 20 days, the rate of hatching varied with incubation temperature, being 0, 61% and 81% at 20, 25 and 30^oC, respectively. After chilling for 40 days or more, however, almost all eggs hatched at 20–30^oC. Diapause with a reduced intensity seemed to be eliminated easily by a high temperature of 25 or 30^oC.
When eggs chilled at 10^oC for 20 days were kept at 20^oC for 7 days or more before incubation at 25^oC, almost all eggs maintained diapause. Most eggs chilled at 10^oC for three 10 day periods separated by 3 days of warming at 25^oC failed to terminate diapause. Daily alternations of 10^oC (18 h) and 25^oC (6h) decreased the diapause-terminating effect of chilling. These facts suggest that diapause intensity can be restored if eggs chilled insufficiently are exposed to a moderately high temperature. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Diapause induction, maintenance and termination in the rice stem borer Chilo suppressalis (Walker)

The rice stem borer, Chilo suppressalis, enters facultative diapause as fully grown larvae in response to short-day conditions during the autumn. Our results showed that the critical night length for diapause induction in C. suppressalis was between 10 h 22 min and 10 h 45 min at 22, 25 and 28 °C, 11 h 18 min at 31 °C, and between 10 h 5 min and 10 h 20 min under field conditions (average temperature ranged from 27.2 to 30.7 °C). The diapause incidence declined in ultra-long nights (18-22 h scotophases) and DD, and increased in ultra-short nights (2-6 h scotophases) and LL. Moreover, we found that the third instar was the stage most sensitive to the photoperiod, and night length played an essential role in the initiation of diapause. Night-interruption experiments with a 1-h light pulse at LD 12:12 (light 12:dark 12) exhibited two troughs of diapause inhibition, with one occurring in early scotophase and the other in late scotophase. Field observations for six years showed that most larvae entered winter diapause in August in response to declining day lengths, despite the high temperatures prevailing during August. By periodically transferring the field-collected overwintering larvae to different photoperiods and temperatures, the results showed that photoperiod had a significant influence on diapause development during the early phase of diapause, while high temperature significantly accelerated the termination of larval diapause.     

Effect of anoxia on diapause termination in eggs of the false melon beetle, Atrachya menetriesi

The effect of anoxia on diapause development in the leaf beetle Atrachya menetriesi was investigated to elucidate the role of oxygen in regulation of egg diapause. While anoxia alone had no effect on diapause termination, it decreased diapause intensity before chilling. Such an effect reached a maximum level when anoxia lasted for about 10 days. Anoxia applied during the pre-diapause stage also reduced diapause intensity. On the other hand, anoxia terminated diapause when the diapause intensity had been lowered by sufficient duration of chilling (50 days at 7.5 degrees C). The effect of anoxia was temperature dependent; the larger effect was elicited when anoxia was combined with a higher temperature. A 50-day chilling caused more than 20% of eggs to terminate diapause upon transfer to warm conditions. However, when this chilling period was interrupted on the 20th day by a 5-day exposure to a high temperature of 20, 25 or 30 degrees C, the effect of the former chilling was cancelled partially or completely, suggesting that warming reversed diapause development. This reversing effect of a high temperature, however, was not manifested when the warming was combined with anoxia. The results suggest that anoxia inhibits diapause reversal and facilitates a certain process of diapause development. The sequence of exposure to anoxia and chilling is not important.     

Effects of low temperature on diapause termination and body colour change in adults of a stink bug, Plautia stali

Abstract. Diapause adults of Plautia stali Scott maintained at 20°C under short day conditions (LD 12:12 h) were exposed to four temperatures of 5–20°C to examine the effect on diapause development which was assessed in terms of oviposition. Diapause adults kept at 20°C under short day conditions changed their body colour gradually from brown to green and started egg laying after a prolonged preoviposition period. Those transferred to either 10 or 15°C also showed colour change but did not lay eggs. Bugs exposed to 5°C underwent neither body colour change nor oviposition and died more rapidly than those kept at higher temperatures. When 30-day-old diapause adults were chilled at 5, 10 or 15°C for 30 or 60 days and returned to 20°C and long day conditions (LD 16:8 h), the preoviposition period varied primarily depending on the chilling, but not on the temperature. On the other hand, when 60day-old diapause adults chilled for 30 days were observed at 20°C and long day conditions, their preoviposition period tended to be longer as the chilling temperature was lower In this case, a temperature of 10°C appeared to intensify diapause. Therefore, the effect of chilling on diapause development varied depending on the age at which insects were chilled. When chilled bugs were transferred to short day conditions at 20°C, most females failed to lay any eggs and some turned green, then after a while, some green bugs changed to brown again. These results indicate that bugs remained sensitive to short day conditions even after a 60-day chilling at 10 or 15°C.     

Effect of temperature regime on diapause intensity in an adult-wintering Hymenopteran with obligate diapause

Osmia lignaria is a solitary bee that over-winters as a fully eclosed, cocooned, unfed adult. Our objective is to understand the effect of wintering temperature on diapause maintenance and termination in this species. We measure respiration rates and weight loss in individuals exposed to various wintering temperatures (0, 4, 7, 22 °C, outdoors) and durations (28, 84, 140, 196, 252 days). We use time to emerge and respiration response (respiration rate measured at 22 °C) as indicators of diapause intensity. Adults spontaneously lower their respiration rates to ∼0.1 ml/g h within 1 month after adult eclosion, indicating obligatory diapause. Non-wintered individuals maintain low respiration rates, but lose weight rapidly and die by mid-winter. In wintered adults, two phases can be distinguished. First, respiration response undergoes a rapid increase and then reaches a plateau. This phase is similar in bees wintered at 0, 4 and 7 °C. In the second phase, respiration response undergoes an exponential increase, which is more pronounced at the warmer temperatures. Composite exponential functions provide a good fit to the observed respiration patterns. Adults whose respiration response has reached 0.45 ml/g h emerge promptly when exposed to 20 °C, indicating diapause completion. Individuals wintered for short periods do not reach such respiration levels. When exposed to 20 °C these individuals lower their metabolic rate, and their emergence time is extended. The relationship between respiration rates and emergence time follows a negative exponential function. We propose two alternative models of diapause termination to interpret these results.     

Pupal diapause of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) mediated by larval host plants: pupal weight is important

Facultative diapause, a strategy that allows insects to initiate additional generations when conditions are favorable or to enter diapause when they are not, has a profound effect on the ecology and evolution of species. Most previous studies have concentrated on the role of photoperiod and temperature in inducing facultative diapause in insects. In contrast, here we studied pupal diapause mediated by larval host plants in the cotton bollworm Helicoverpa armigera, and confirmed that pupal weight is a critical factor. Two groups of third instar H. armigera larvae, kept at 25 °C with L:D = 8:16 and 20 °C with photoperiod of L:D = 8:16, respectively, were fed on six host plants and on artificial diet (as a control) to determine how larval host plants affect diapause incidence and related traits (such as pupal weight and developmental duration). The data showed larval host plants affected diapause incidence significantly and the effects could be masked by low temperature. Further analysis showed that pupal size, not the length of the sensitive stage, affected the decision to enter diapause. In a further experiment, third-instar to final-stage larvae deprived of artificial diet for 2 days demonstrated a direct relationship between pupal weight and diapause incidence. These results suggest that larval host plants, by affecting pupal size, may influence diapause occurrence in H. armigera. This has important adaptive significance for both over-wintering survival and the possibility for completing an additional generation.     

Diapause in a tropical species,Cothonaspis boulardi (Parasitic hymenoptera)

Summary The strain of Cothonaspis boulardi originating from a field collection at Guadeloupe (16° lat. N-West Indies) exhibits a facultative diapause that occurs at the final larval instar. The diapause was induced by low temperature (17.5° C) and was inhibited at 25° C. Diapause was independent of photoperiod. The termination of diapause was hastened in about 10 days by exposing larvae to a temperature of 25° C. Although a succession of sufficient cold days for diapause occurs only rarely in the collection area (Petit-Bourg), at higher elevations within 20 km of Petit-Bourg conditions that could induce diapause occur annually.This study is supported by research grant of C.N.R.S. (ATP Ecophysiologie, n^o 51-3571)     

Diapause development in frozen larvae of the goldenrod gall fly, Eurosta solidaginis fitch (diptera: tephritidae)

Seasonal changes in metabolic rate and the potential for morphological development demonstrated that third-instar larvae of the goldenrod gall fly, Eurosta solidaginis Fitch, exhibit a distinct winter diapause. Metabolic rate (CO2 production) was significantly lower from 15 October to 9 February than in early autumn (9 September) and spring (1 March) samples. The induction of diapause coincided with the development of cold-hardening, maximum larval mass, and gall senescence, but our experiments did not identify specific cues triggering diapause induction. We examined the influence of exposure to 0 degrees C and -20 degrees C on diapause development. Diapause development in larvae stored at 0 degrees C occurred at approximately the same rate as in nature. Until 15 December the larvae were in the refractory phase of diapause (incapable of morphological development, even at permissive temperatures), but afterward moved to the activated phase within which diapause intensity decreased until termination in February. Diapause development occurred in larvae collected during the winter and stored at -20 degrees C for periods of 1 week to 3 months. Diapause intensity decreased in frozen larvae through the winter but at a slower rate than in larvae stored at 0 degrees C.     

Termination of the facultative diapause in the codling moth, Laspeyresia pomonella (Lepidoptera, Tortricidae)

The abiotic factors regulating the termination of the facultative diapause of the mature larva of the codling moth, Laspeyresia pomonella (L.), are described. The termination of diapause under long-day conditions (LD) is influenced by: (i) the rearing temperature of the larvae during prediapause development, (ii) the duration of the preincubation period, i.e. the time for which the diapausing larvae remain under prediapause rearing conditions, (iii) the reactivating incubation, i.e. the period for which the larvae are chilled, and (iv) the complementary incubation (LD and, except in one experiment, 26°), i.e. the period after the chilling needed for the pupation of the insects. The complementary incubation is distinctly shorter if the prediapause development of the larvae takes place at rearing temperatures below 26°, i.e. 21° or 19°. The latter conditions led to 100% pupae and pupation could be accelerated by prolonging the preincubation period. In larvae reared at 26°, the prolongation of the preincubation period raised the rate of pupation to a maximum of only 56% and also caused higher mortality. On the other hand, diapause was terminated under short-day conditions if the temperature was raised to 26° after a rearing temperature of 19° and a preincubation period of 90 days.

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Beendigung der fakultativen diapause des apfelwicklers, Laspeyresia Pomonella (Lepidoptera, Tortricidae)

Zusammenfassung Es wurde untersucht, welche abiotischen Faktoren die Beendigung der fakultativen Diapause ausgewachsener Apfelwicklerlarven (Laspeyresia pomonella) steuern. Die Beendigung der Diapause unter Langtagbedingungen (LT) wird beeinflusst durch: 1. die Zuchttemperatur der Larven während der Prädiapauseentwicklung; 2. die Dauer der Präinkubation, d.h. die Zeit, während der diapausierende Larven unter Prädiapause-Zuchtbedingungen bleiben; 3. die Reaktivierungs-inkubation, d.h. eine Periode von 70 d, während der die Larven auf 4° gekühlt werden und 4. die Komplementärinkubation (LT und, ausser in einem Experiment, 26°), d.h. die nach der Kühlung bzw. dem Wechsel in der Photoperiode benötigte Zeitdauer bis zur Verpuppung. Die Komplementärinkubation ist deutlich kürzer, wenn die Prädiapauseentwicklung der Larven bei Temperaturen unter 26° stattfindet. Bei 19° verpuppten sich 100% der Larven, wobei der Zeitpunkt der Verpuppung durch die Verlängerung der Präinkubationszeit beschleunigt wurde. Bei Larven, die bei 26° gezüchtet wurden, erreichte die Verpuppungsrate lediglich 56% bei einer relativ langen Präinkubationsdauer, wobei deren Verlängerung auch die Larvenmortalität erhöhte. Nach einer Zuchttemperatur von 19° und einer Präinkubationsdauer von 90 Tagen konnte die Diapause unter Kurztagbedingungen beendet werden, wenn die Temperatur auf 26° erhöht wurde.