Prepupal diapause and instar IV developmental rates of the spruce beetle, Dendroctonus rufipennis (Coleoptera: Curculionidae, Scolytinae)

The spruce beetle, Dendroctonus rufipennis (Kirby), is an important mortality agent of native spruces throughout North America. The life-cycle duration of this species varies from 1 to 3 years depending temperature. The univoltine cycle (one generation per year) is thought to maximize outbreak risk and accelerate host mortality in established outbreaks. Prepupal diapause is associated with the semivoltine cycle (one generation per 2 years) and we investigated thermal conditions that result in diapause induction. Preliminary experiments used respirometry in an attempt to distinguish the diapause state of experimental insects but the technique was apparently confounded by low respiration before and during pupation, regardless of diapause status. Therefore, diapause induction was deduced using developmental delays. The observed developmental response was not a “switch”, with developmental delay either present or absent, but instead varied continuously. We found that temperatures <15 °C from instar III through mid-instar IV were associated with developmental delays beyond that expected from cool temperatures. Moreover, the duration of exposure to cool temperatures was important in determining the degree of developmental delay. Small, if any, delays were observed if the cumulative exposure to <15 °C was <20 d whereas >40 d cumulative exposure was associated with distinct developmental suppression. Intermediate exposure to cool temperatures resulted in minor developmental delays. We used our results to parameterize a maximum likelihood estimation model of temperature-dependent instar IV developmental rates, including the effect of diapause. This model can be included as part of a spruce beetle phenology model for predicting population dynamics.     

Diapause survival and post-diapause performance in bumblebee queens (Bombus terrestris)

Diapause survival and post-diapause performance (i.e., if a queen starts to lay eggs) of in total 2210 bumblebee queens (Bombus terrestris) were measured under different diapause regimes: 5 temperatures (–5, 0, 5, 10 and 15 °C) in combination with 5 durations of exposure (1, 2, 4, 6 and 8 months).The results show that weight at the start of diapause determines to a large extent whether a queen will be able to survive diapause. Queens with a wet weight below 0.6 g prior to diapause did not survive, but for those queens exceeding this threshold a higher pre- diapause weight did not increase their post-diapause performance.There was no effect of temperature on diapause survival; 76% of the variance in survival could be explained by the duration of the treatment. Neither temperature nor duration of exposure had an effect on post-diapause performance. The preoviposition period of the queens that laid eggs was also determined. The preoviposition period was affected by both temperature and duration of exposure: the preoviposition period decreased with decreasing temperature but also with increasing duration of the treatment.     

Egg development and diapause: ecophysiological and genetic basis of phenological polymorphism and adaptation to varied hosts in the green oak tortrix,Tortrix viridana L. (Lepidoptera: Tortricidae)

Experiments concerned 8 tortrix populations associated with varied oak species. They showed that the egg development included a phase of diapause. Completion of embryogenesis at 20 degrees C was used as a criterion for whether diapause was completed. Under semi-natural conditions diapause terminated in late autumn or early winter, then eggs developed continuously, without postdiapause winter quiescence, even in severe cold. The eggs from the populations associated with holm or cork oak completed diapause then hatched later than those from the populations associated with sessile or pubescent oak. Reciprocal crossbreedings confirmed that this phenological polymorphism was genetically determined. Under constant temperatures the physiological state of diapausing eggs, assessed by measuring their cold requirements to complete diapause by exposure to 8 degrees C, varied gradually with increasing age. This diapause development was strongly temperature-dependent. Cold requirements of diapausing eggs were much higher in a late-hatching than in an early-hatching population. This explains the phenological polymorphism of the tortrix: the more eggs need cold, the later they complete diapause in autumn, and the later they hatch in spring. Egg cold requirements varied widely within populations too, which resulted in large variations in the date of diapause end among individuals. The date of egg hatch was influenced by the temperatures occurring during diapause and postdiapause, but apparently not by photoperiod.     

European field collections and Canadian releases ofCeranthia samarensis (Dipt.: Tachinidae), a parasitoid of the gypsy moth

A programme to collect, import and release into Canada the gypsy moth parasitoid,Ceranthia samarensis (Diptera: Tachinidae) is described. The parasitoid's potential for biological control in Canada is also discussed. The parasitoid was collected in Europe by exposing experimental gypsy moth larvae in areas where local gypsy moth populations were at low densities. Following field exposure, the host larvae were returned to the laboratory and parasitoids reared from them. This technique has shown thatC. samarensis is the suffers 7–16% hyperparasitism. From 83–90% of theC. samarensis typically enter diapause as pharate adults within the puparia. Laboratory tests of post-exposure host rearing conditions indicate that constant temperatures disrupt the normal parasitoid diapause and that this effect can not be offset by use of either static long or short photoperiods or natural daylengths. Shipping and cold-storage procedures for puparia are described. Post-storage time to emergence of adultC. samarensis decreased with longer cold storage periods and with higher post-storage incubation temperatures. Emergence requires 112 degree-days above a threshold of 8°C after a period of at least 8 months cold storage. Releases of adultC. samarensis into field cages at four locations in southern Ontario are documented. While dissection of host larvae from the field cages has failed so far to demonstrate evidence of parasitism, we remain hopeful that some establishment of the parasitoid has occurred.      

Development, survivorship, and reproduction of Helicoverpa armigera (Lepidoptera: Noctuidae) under constant and alternating temperatures

Laboratory studies were conducted to assess the effect of temperature on the survival, development, fecundity, and longevity of Helicoverpa armigera (Hübner) at 11 constant temperatures ranging from 12.5 to 40 degrees C, as well as at five alternating temperature regimes (25-10, 30-15, 32.5-17.5, 35-20, and 35-27.5 degrees C) and under a photoperiod of 16:8 (L:D) h. H. armigera reared at constant temperatures did not develop from egg to adult (emergence) outside the temperature range of 17.5-32.5 degrees C. The alternating conditions expanded this range from 10 to 35 degrees C. The lowest developmental thresholds of the immature stages were estimated by a linear model and ranged from 10.17 (pupal stage) to 11.95 degrees C (egg stage) at constant temperature regimes and from 1.1 to 5.5 degrees C, respectively at alternating temperatures. The values of developmental thresholds estimated using the nonlinear (Lactin-2) model were lower than those estimated by the linear model for constant and alternating temperature regimes except for larval and pupal stages at constant temperatures. Mean adult longevity fluctuated from 34.4 d at 15 degrees C to 7.6 d at 35 degrees C. Females reared under all alternating temperature regimes laid more eggs than females reared at any, except the 25 degrees C, constant temperature treatment. The intrinsic rate of increase was highest at 27.5 degrees C, at both the constant and the corresponding alternating temperature regimes (0.147 and 0.139, respectively). Extreme temperatures had a negative effect on life table parameters.     

Influence of temperature, photoperiod and humidity on oviposition and egg hatch of the root-feeding flea beetle Longitarsus bethae (Chrysomelidae: Alticinae), a natural enemy of the weed Lantana camara (Verbenaceae)

The root-feeding flea beetle Longitarsus bethae Savini & Escalona, was introduced into South Africa as a candidate biological control agent for the noxious and invasive weed, Lantana camara L. As part of the study to predict the beetles' survival in its new range, the influence of climatic conditions on its egg development and reproductive performance were investigated in the laboratory. The threshold temperature (T degrees) and degree-days (DD) required for egg hatch were determined after exposing the eggs to various constant temperatures (12, 17, 22, 27 and 32 degrees C) in separate growth chambers. The DD required for egg hatch was 178.6, and the temperature threshold required for egg hatch was 11.3 degrees C. Survival of eggs varied from 27 to 56% at 32 and 17 degrees C, respectively, and was optimum between 17 and 25 degrees C. Oviposition was examined under high and low relative humidity (RH) regimes while egg hatch was determined at six RH levels, each maintained in a separate controlled growth chamber set at a constant temperature (25 degrees C). Whilst RH had no influence on oviposition, eggs were highly susceptible to aridity, and continuous exposure to relative humidity below 63% for more than three days was wholly lethal at 25 degrees C. Optimum egg hatch occurred at RH between 85 and 95% for up to 12 days. The effect of day length on oviposition and subsequent egg hatch was investigated under two photoperiod regimes. Neither oviposition nor subsequent egg hatch was influenced by photoperiod. The knowledge obtained will be useful for mass rearing as well as field release programmes for L. bethae.     

Hydroprene prolongs developmental time and increases mortality of Indianmeal moth (Lepidoptera: Pyralidae) eggs

Eggs of the Indianmeal moth, Plodia interpunctella (Hübner), were exposed to the labeled rate of hydroprene (1.9 x 10(-3) mg [AI]/cm2) sprayed on concreted petri dishes. These eggs were exposed for 1, 3, 6, 12, and 18 h and until hatching (continuous exposure) at temperatures of 16, 20, 24, 28, and 32 degrees C and 57% RH until the emergence of first instars. The developmental time and egg mortality were significantly influenced by temperature and exposure periods. At 16 degrees C, hydroprene did not cause differences in developmental time when eggs were exposed for different periods. At temperatures >16 degrees C, both exposure period and temperature influenced developmental time. The maximum developmental time (15.0 +/- 0.2 d) occurred at 16 degrees C, and the minimum developmental time (3.2 +/- 0.3 d) occurred at 32 degrees C. Mortality increased when eggs were exposed to hydroprene for longer periods at all of the five tested temperatures. The greatest mortality (81.6 +/- 2.1%) occurred when eggs were continuously exposed on treated surfaces at 32 degrees C. We used developmental time instead of rate (1/ developmental time) to fit simple linear or polynomial regression models to the development data. Appropriate models for developmental time and mortality were chosen based upon lack-of-fit tests. The regression models can be used in predictive simulation models for the population dynamics of Indianmeal moth to aid in optimizing use of hydroprene for insect management.     

Thermal and thermoperiodic effects on larval development and diapause in the European corn borer, Ostrinia nubilalis

Effects of temperature and thermoperiod on larval development and the induction of diapause were investigated in the European corn borer, Ostrinia nubilalis. Developmental threshold temperatures characteristic of the first four larval instars were estimated under both thermoperiods and constant temperatures. Threshold values were similar under the two conditions, but the 4th-larval instar was shown to display a significantly lower developmental threshold temperature than was characteristic of the earlier instars. Although developmental times (days per instar) were not greatly affected by fluctuating temperature regimes, the quantity of growth (weight, head width) during the 4th and 5th (last) stadia was found to be much greater under thermoperiodic regimes than under comparable constant temperature conditions. By means of thermoperiodic regimes having identical mean temperatures but different cryophase durations, it was demonstrated that the induction of diapause is dependent on the duration of the cryophase rather than on the mean temperature of the thermoperiod. To be effective, the cryophase must be colder than the insect's thermoperiodic response threshold. It was also demonstrated that thermoperiodic responses that are readily produced under continuously dark rearing conditions are not manifested under continuous light.     

Developmental arrest during embryonic development of the common chameleon (Chamaeleo chamaeleon) in Spain

Embryonic development of the common chameleon, Chamaeleo chamaeleon, was monitored from oviposition to hatching at a field site in southwestern Spain and in the laboratory under five experimental temperature regimes. Embryos were diapausing gastrulae at the time of oviposition; developmental arrest in the field continued as cold torpor during winter. Postarrest development in the field commenced in April, and hatching occurred in August, for a total incubation period of 10.5 mo. In the laboratory, one group of eggs was incubated at a constant warm (26 degrees C) temperature. The remaining treatments simulated field conditions and consisted of initial periods of warm temperature of 0, 27, 46, and 71 d, a subsequent 4-mo period of cold winter (16 degrees C) temperature, and a final period of warm (26 degrees C) temperature. Embryos in the constant warm temperature treatment were in diapause an average of 3 mo, with clutch means ranging from 2 to 4 mo. Hatching among clutches occurred over 2 mo. In contrast, for field and experimental eggs that experienced cold winter conditions, hatching within treatments occurred over 2-14 d; "winter" conditions synchronized development. The length of time between the end of cold conditions and hatching did not differ among treatments; development thus resumed as soon as temperature was suitable regardless of the initial period of warm temperature. Diapause in nature thus insures that embryos remain gastrulae after oviposition despite nest temperatures that may be warm enough to support development.     

Effects of constant and changing temperature conditions on diapause induction in Helicoverpa armigera (Lepidoptera: Noctuidae)

The effects of photoperiod and temperature on the induction and termination of facultative pupal diapause in Helicoverpa armigera (Lepidoptera: Noctuidae) were investigated under laboratory conditions. Exposing H. armigera larvae to both constant and fluctuating temperature regimes with a mean of 25°C and 20°C resulted in a type-III photoperiodic response curve of a short-long day insect. The long-day critical daylengths for diapause induction were ten hours and 12 hours at the constant temperatures of 25°C and 20°C, respectively. Higher incidences of diapause and higher values both for the longer and the shorter critical photoperiods for diapause induction were observed at fluctuating regimes compared with the corresponding constant ones. At alternating temperatures, the incidence of diapause ranged from 4.2% to 33.3% and was determined by the temperature amplitude of the thermoperiod and by the interaction of cryophase or thermophase with the photoperiod. Helicoverpa armigera larvae seem to respond to photoperiodic stimuli at temperatures >15°C and <30°C; all insects entered diapause at a constant temperature of 15°C, whereas none did so at a constant temperature of 30°C under all the photoperiodic regimes examined. Although chilling was not a prerequisite for diapause termination, exposure of diapausing pupae to chilling conditions significantly accelerated diapause development and the time of adult emergence. Therefore, temperature may be the primary factor controlling the termination of diapause in H. armigera.     

Exposure of <Emphasis Type="Italic">Eocanthecona furcellata</Emphasis> (Hemiptera: Pentatomidae) nymphs and adults to high temperatures induces an aestivo-hibernal egg diapause: a strategy for surviving hot summers

The predatory stink bug Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae), which is widely distributed in the subtropics and tropics, is a potential biological control agent for lepidopteran, coleopteran, and hemipteran pests. We investigated development of eggs oviposited by E. furcellata females subjected to various photoperiods (i.e., 8L:16D, 12L:12D, and 16L:8D) and temperature regimes (i.e., 26:26, 30:28, 32:30, 34:32, 35:33, and 36:34 °C; temperatures at light and dark phases, respectively) at different developmental stages. Exposure of E. furcellata nymphs and adults to different photoperiods and temperature regimes affected the development of eggs they subsequently oviposited. Temperatures greater than 30 °C induced diapause in a portion of the eggs, as evidenced by the delayed appearance of the black circle and the red spot, characteristics of early and late stage of embryonic development, respectively. The proportion of diapausing eggs generally increased with increasing temperature exposure, whereas it decreased with advancing developmental stage/age at which E. furcellata was subjected to the temperature regime. Although viability of diapausing eggs was lower than that of non-diapausing eggs, diapausing eggs survived longer under high temperatures. The trade-off between survivability and viability may be a strategy to survive unpredictability of summer temperatures.     

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.     

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.     

Effect of thermoperiod on diapause intensity in Pyrrhocoris apterus (Heteroptera Pyrrhocoridae)

The intensity of adult diapause in Pyrrhocoris apterus was measured in two series of experiments as the duration of pre-oviposition period at a constant temperature of 25 degrees C after transfer from short (12L:12D) to long day conditions (18L:6D). Higher diapause intensity was induced with a thermoperiod than at constant temperatures. After the induction throughout larval instars 3-5 and during 4 weeks of adult life at short days and a thermoperiod of 25/15 degrees C the pre-oviposition period was 30+/-4 and 26+/-3 days. After induction at constant 25 degrees C the pre-oviposition period was 22+/-3 and 23+/-4 days, while after induction at constant 20 degrees C it was 17+/-4 and 19+/-4 days. Induction at a lower constant temperature of 20 degrees C was thus followed by a less intense diapause than the induction at a higher constant temperature of 25 degrees C. These counterintuitive results are discussed. The oxygen consumption rate measured at experimental temperatures prior to transfer from short to long days was higher at thermoperiodic conditions than at constant temperatures and it was similar at constant 20 and 25 degrees C. Thus, the oxygen consumption rate measured prior to the transfer was highest (indication of the least intense diapause) in the insects that showed later, after the transfer to long days, the longest pre-oviposition period (indication of the most intense diapause). Within the first two days after transfer to constant 25 degrees C, oxygen consumption rate measured at 25 degrees C decreased in the thermoperiodic insects, while it transiently increased in insects from constant 20 degrees C. Two days and later after the transfer, oxygen consumption rate was similar in all groups. Cold hardiness was not correlated with diapause intensity. The low lethal temperature in diapausing insects was correlated with the night temperature during diapause induction.     

Abiotic factors influencing embryonic development, egg hatching, and larval orientation in the reindeer warble fly, Hypoderma tarandi

Wild-caught, tethered females of the reindeer warble fly, Hypoderma tarandi (L.) (= Oedemagena tarandi (L.)), (Diptera, Oestridae) were stimulated to oviposit on hairs of a reindeer hide. Newly laid eggs incubated at constant temperatures and relative humidities hatched within 3 days to 2 weeks, depending on the experimental conditions. Over a range of 7-40 degrees C, hatching only occurred between 20 and 37 degrees C. Eggs held at 100% relative humidity had lower hatchability and longer time to hatch relative to eggs held at 77% relative humidity. The average number of degree-days for hatching was 50.35. Between 20 and 33 degrees C there was a temperature-dependent linear trend in developmental rate, and the proportion of eggs hatching was highest, and least variable, at the mid-temperature ranges. The temperature range found in the natural host micro-habitat where H. tarandi commonly affix their eggs (close to the skin at the base of a host hair) was consistent with the experimental temperature treatments that produced the highest hatching rate. Newly emerged larvae displayed positive thermotaxis, while showing no phototaxic or geotaxic behaviour. Results indicate that constraints of the host environment, coupled with temperature-dependent hatching success, may impose a selective pressure on oviposition behaviour.     

Phenotypic responses of hatchlings to constant versus fluctuating incubation temperatures in the multi-banded krait, Bungarus multicintus (Elapidae)

Most studies on egg incubation in reptiles have relied on constant temperature incubation in the laboratory rather than on simulations of thermal regimes in natural nests. The thermal effects on embryos in constant-temperature studies often do not realistically reflect what occurs in nature. Recent studies have increasingly recognized the importance of simulating natural nest temperatures rather than applying constant-temperature regimes. We incubated Bungarus multicintus eggs under three constant and one fluctuating-temperature regimes to evaluate the effects of constant versus fluctuating incubation temperatures on hatching success and hatchling phenotypes. Hatching success did not differ among the four treatments, and incubation temperature did not affect the sexual phenotype of hatchlings. Incubation length decreased as incubation temperature increased, but eggs incubated at fluctuating temperatures did not differ from eggs incubated at constant temperatures with approximately the same mean in incubation length. Of the hatchling phenotypes examined, residual yolk, fat bodies and locomotor performance were more likely affected by incubation temperature. The maximal locomotor speed was fastest in the fluctuating-temperature and 30 degrees C treatments and slowest in the 24 degrees C treatment, with the 27 degrees C treatment in between. The maximal locomotor length was longest in the fluctuating-temperature treatment and shortest in the 24 degrees C and 27 degrees C treatments, with the 30 degrees C treatment in between. Our results show that fluctuating incubation temperatures do not influence hatching success and hatchling size and morphology any differently than constant temperatures with approximately the same mean, but have a positive effect on locomotor performance of hatchlings.     

Molecular characterization of the insect immune protein hemolin and its high induction during embryonic diapause in the gypsy moth,Lymantria dispar

During the embryonic (pharate first instar) diapause of the gypsy moth, Lymantria dispar, a 55 kDa protein is highly up-regulated in the gut. We now identify that protein as hemolin, an immune protein in the immunoglobulin superfamily. We isolated a gypsy moth hemolin cDNA and demonstrated a high degree of similarity with hemolins from three other moth species. Hemolin mRNA levels increased at the time of diapause initiation and remained high throughout the mandatory period of chilling required to terminate diapause in this species, and then dropped in late diapause. This mRNA pattern reflects the pattern of protein synthesis. These results suggest that hemolin is developmentally up-regulated in the gut during diapause. Diapause in this species can be prevented using KK-42, an imidazole derivative known to inhibit ecdysteroid biosynthesis, and gypsy moths treated in this manner failed to elevate hemolin mRNA. Conversely, this diapause appears to be initiated and maintained by the steroid hormone, 20-hydroxyecdysone, and the addition of 20-hydroxyecdysone to the culture medium elevated hemolin mRNA in the gut. Our results thus indicate a role for 20-hydroxyecdysone in the elevation of hemolin mRNA during diapause. Presumably, hemolin functions to protect the gypsy moth from microbial infection during its long, overwintering diapause.     

Effects of photoperiod and temperature on pupal diapause induction of grape berry moth Lobesia botrana

Abstract. The grape berry moth, Lobesia botrana Denn. & Sciff. (Lepidoptera: Tortricidae), one of the most injurious pest of grape berries in Greece, is a multivoltine species that overwinters as diapausing pupae. The effects of several diel and non-diel photoperiods and of temperature, experienced by eggs and larvae, on pupal diapause induction were investigated. The diapause response curve was of Type I (long day type) and the determining factor was the duration of scotophase (> 11 h), regardless of the duration of photophase. However, at very short (< 4 h) photoperiods, the incidence of diapause was also high. Diapause was positively and significantly correlated with the egg-larval developmental time, pupal mortality and the duration of the pupal stage. Eggs and larvae reared under LD 12 : 12 h photoperiod and various temperatures (from 12 to 30 °C) produced diapausing pupae (almost 100%), but the duration of the pupal stage (intensity of diapause) increased with increasing temperature. Under continuous darkness, however, the percentage diapause decreased with increasing temperature. Single and double 1-h light pulses were applied systematically at various times during the scotophase of six diapause-inducing diel photoperiods. Two photosensitive points in time (called A and B) were revealed, during which illumination resulted in a significant decrease of diapause induction. The decrease was much greater during the first sensitive period (early in scotophase) rather than in the second (late in the scotophase).