Delayed Egg Hatching and Semivoltinism in the Nearctic Stonefly Megarcys signata (Plecoptera: Perlodidae)

Delayed egg hatching can influence the survival of aquatic insects by reducing exposure to competitors, predators, parasites, or unfavorable environmental conditions. We examined egg development in a Colorado population of the stonefly, Megarcys signata (Plecoptera: Perlodidae), whose larvae inhabit high altitude streams in the Rocky Mountains of western North America. Five-thousand fertilized eggs were collected from 40 different females and incubated in the East River, Gunnison County, Colorado, until late fall, at which time unhatched eggs were transported to incubation chambers in Ithaca, New York. We used three different combinations of photoperiod and temperature (10L:14D, 4°C; 10L:14D, 8°C; 8L:16D, 8°C) in an attempt to induce hatching. Eggs in the treatments with temperatures elevated above normal winter temperatures in the East River (8°C) began developing after 6 months, whereas eggs in the treatment most closely simulating natural stream conditions (4°C) did not hatch after 10 months of incubation. Our data indicate that this population of Megarcys signata has an extended egg stage that persists for almost a year, and that it is semivoltine at these elevations of 2885 and 2895 meters.     

Variable egg development of Dinocras spp. (Plecoptera, Perlidae) and the stonefly seed bank theory

1. Temperature dependence of egg development of Dinocras cephalotes (Curtis) (three German and one Norwegian population) and Slovenian D. megacephala (Klapálek) was studied under a constant 14 : 10 light : dark photoperiod and constant temperature ranges of 4–24 °C and 4–18 °C, respectively. D. cephalotes was also incubated under seasonal field conditions; natural daylength and fluctuating temperatures had no modifying effect. 2. Both species have very similar lower threshold temperatures (4 and 3.5 °C, respectively), thermal demand for development (c. 600 degree days) and high dependence of mean incubation period on temperature (exponents of regressions near 1.5). Present data on D. cephalotes agree with the literature on British and Norwegian material of the same species. 3. Development occurs only at cue temperatures above the lower threshold. Cue temperatures range from 6 °C (some D. megacephala) to 14 °C (some D. cephalotes) and vary strongly within and between egg masses of D. cephalotes. Variation is not random, but seems to be genetically determined. 4. The variable temperature response renders study of effects of particular experimental regimes, and comparisons between local populations, difficult. 5. A latitudinal gradient in cue temperatures for development from 6 °C at c. 46 °N to 12 or even 14 °C at c. 61 °N seems to reflect reduced diversity at high latitudes. 6. Average success of spontaneous hatching exceeded 90% between 12 and 20 °C, but declined towards higher and lower temperatures. 7. Unhatched eggs were not dead but in parapause; development at other, higher or lower, temperatures was induced. Spontaneous plus induced hatching success approached 90%. Developing eggs rarely died; most dead eggs were apparently unfertilized. 8. Dormant plecopteran eggs are proposed to form a seed bank in stream bed sediments. Highly successful development after up to 220 days of dormancy was ascertained in Dinocras, and survival for up to 3 years is reported for other Perloidea. 9. Only systellognathan egg morphology provides options for long dormancy; the other plecopteran superfamilies, notably Nemouroidea, follow different strategies.     

Temperature dependence for development of the whitefly predator <Emphasis Type="Italic">Clitostethus arcuatus</Emphasis> (Rossi)

This work aimed to study the biology of Clitostethus arcuatus (Rossi) (Coleoptera: Coccinellidae) under different temperatures and evaluate the optimum temperature for its mass rearing. Studies were carried out in the laboratory at four constant temperatures (15°C, 20°C, 25°C and 30°C), 75 ± 5% relative humidity and a photoperiod of 16 h light:8 h dark, in which C. arcuatus was fed ad libitum with nymphs of all instar of Aleyrodes proletella L. (Homoptera: Aleyrodidae) on Brassicae oleracea L. (var. Costata). The following biological parameters were evaluated: development time and survival rates of pre-imaginal stages, adult longevity (female and male), length of the pre-oviposition and oviposition periods, fecundity, fertility and percentage of egg hatching. Population growth parameters, the lower development threshold and the sum of effective temperatures were estimated. Temperatures ranging from 20°C to 30°C were suitable for the development of C. arcuatus, suggesting that this species is well adapted to the temperatures usually found inside greenhouses or in open fields in temperate regions. Although the intrinsic rate of natural increase and doubling time were similar at 25°C and 30°C, the temperature of 25°C was shown to be the most suitable for mass rearing and development of populations under field conditions, since the percentage of egg hatching and the accumulated survival rates of the pre-imaginal stages were the highest. Considering the estimated lower threshold for pre-imaginal development (7.9°C) and the sum of effective temperatures [293.6 degree-days (°D)], it is predicted for Ponta Delgada (Azores, Portugal) that the first adults of C. arcuatus should emerge by the first fortnight of February and that up to 12 generations per year can occur.     

Control of preimaginal development by photoperiod and temperature in the dock leaf beetle <Emphasis Type="Italic">Gastrophysa viridula</Emphasis> (De Geer) (Coleoptera,Chrysomelidae)

Development times of preimaginal stages and immature adult body mass of the dock leaf beetle Gastrophysa viridula from Bryansk (53°N, 34°E) were determined at six constant temperatures ranging from 16 to 26°C and at two photoperiods, 12L: 12D or 22L: 2D. The lower developmental threshold (LDT) and sum of degreedays (SDD) were calculated for eggs, larvae, and pupae; the values for the total preimaginal development were 7.9°C and 274.8 °C×d, respectively. The thermal reaction norms for preimaginal development were not affected by photoperiod, although the adult body mass was on average greater under short-day conditions. Our results confirm the previously published data on G. viridula concerning diapause induction by shorter daylengths, greater body mass under lower temperatures, and narrowing of the thermal optimum range during preimaginal development. The thermal reaction norms for larval development were shown to be the same in individuals that died during the pupal stage, and in those that successfully emerged as adults. Our results are more precise than the previous data on this species due to a larger sample size, careful control of rearing conditions, and the use of an alternative method for calculating LDT and SDD. In particular, a more accurate estimation of LDTs allowed us to demonstrate that rate isomorphy was violated in this species, contrary to what had been stated by Honěk and co-authors (2003).     

Fecundity and feeding of <Emphasis Type="Italic">Galerucella calmariensis</Emphasis> and <Emphasis Type="Italic">G. pusilla</Emphasis> on <Emphasis Type="Italic">Lythrum salicaria</Emphasis>

Fecundity and feeding of two introduced sibling biological control species, Galerucella calmariensis and G. pusilla (Coleoptera: Chrysomelidae) on purple loosestrife, Lythrum salicaria L. (Lythraceae) were compared at constant temperatures of 12.5, 15, 20, 25, and 27.5 °C. Larval feeding was also carried out at 30 °C, but at this temperature, larvae developed only to the L2 stage and none pupated. Thus, data for this temperature were not used in the analysis. There were significant species × temperature interactions in fecundity. Of the two species, Galerucella pusilla laid more eggs. Although egg production of both species was lowest at 12.5 °C and increased to 20 °C, at higher temperatures, the two species reacted differently. From 25 to 27.5 °C, egg production decreased for G. pusilla, but G. calmariensis fecundity peaked at 27.5 °C. Significant temperature × species × life-stage interactions were also observed in feeding. For each species, the amount of feeding varied with temperature and stage of development. Galerucella pusilla adults consumed more foliage at 15, 20, and 27.5 °C. However, at 12.5 °C G. calmariensis adults fed more than G. pusilla. G. pusilla larvae consumed an average of 25% less foliage than G. calmariensis. The lower larval consumption of G. pusilla suggests that when food is limited, G. pusilla larvae may have a higher survival rate because of its ability to complete larval development with less food and produce more progeny due to its greater fecundity. When food is not limited neither species would have a competitive advantage and both species could coexist temporally and spatially. However, since G. calmariensis larvae consumed more leaf material, the larval stage of this species would have a greater impact on purple loosestrife than G. pusilla.     

Effect of temperature on the development, fecundity, progeny sex ratio and life-table of Campoletis chlorideae, an endolarval parasitoid of the pod borer, Helicoverpa armigera

Development, survival, fecundity, progeny sex ratio (PSR) and age-specific life-table parameters of the parasitoid Campoletis chlorideae Uchida (Hymenoptera: Ichneumonidae) were examined at six different constant temperatures (12, 17, 22, 27, 32 and 37°C) in the laboratory [70 ± 10% RH and 10:14 h (light:dark) photoperiod]. Second instar larvae of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) were reared on chickpea (Cicer arietinum L.) and used as the host. Development times shortened as the temperature increased from 12 to 37°C. The estimated lower developmental threshold (tL) was 3.4°C. The thermal summation for total immature stages was 379.97 degree-days. A reciprocal relationship between temperature and longevity was observed in the range of 12–17°C. The maximum mortality of pupae (71.8%) occurred at 37°C. At 22°C, the yield of a female parasitoid averaged 137.3 ± 14.7 (mean ± SD) progeny, of which 89.6 ± 7.6 were daughters. The number of daughters produced decreased when the females were kept either above or below 22°C, although the PSR was female biased in the range of 17–27°C. The analyses of life-table parameters, developmental rates, reproduction, mortality and PSR suggest that maximum population growth (r _m ) is near 27°C. There was little variation observed in most of the desired qualities of C. chlorideae in the range of 17–27°C, and it appears that the parasitoid is adapted to a wide range of temperatures. We suggest that for maximum production the parasitoid should be reared at 22 ± 4°C and be released in areas where the temperature ranges between 17° and 27°C, as in the plains of northern India.     

Role of photoperiod and temperature in the induction of overwintering pupal diapause in the spotted tentiform leafminer,Phyllonorycter blancardella

The role of photoperiod and temperature in the induction of overwintering diapause inPhyllonorycter blancardella (F.) (Lepidoptera: Gracillariidae) was examined in the laboratory and field using leafminers from commercial apple orchards in Ontario, Canada.P. blancardella exhibited a long-day response to photoperiod: long daylengths resulted in uninterrupted development whereas short daylengths induced diapause. The estimated critical photoperiod for diapause induction was L14.25∶D9.75. The larvae of leafminers destined to enter diapause took ca. 3× longer to complete development than the larvae of non-diapausing leafminers. The development prolonging effect of photoperiod decreased with decreasing daylength. Temperature modified the diapause inducing effect of photoperiod. At L14.25∶D9.75, diapause incidence was similar at 15 and 20°C but was lower at 25°C. Photoperiod also altered the normal relationship between development rate and temperature. At L14.25∶D9.75, the duration of larval development of diapausing leafminers was similar at 15, 20 and 25°C. Temperature alone is unlikely to have a role in the induction of diapause because leafminers exposed to natural late summer and fall temperature regimes and L16∶D8 did not enter diapause.     

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.     

On the effects of diet and photoperiod on <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas) (Coleoptera,Coccinellidae) larval development

It is known that the light day length and diet could act as cue factors inducing adult (reproductive) diapause in aphidophagous coccinellids. However, the rate of coccinellid larval development could also depend on photoperiod and diet. The present study examines the effects of photoperiod (L: D = 12: 12 or 16: 8) and diet (the green peach aphid Myzus persicae or eggs of the Angoumois grain moth Sitotroga cerealella) on preimaginal development of the multicolored Asian lady beetle Harmonia axyridis. At 20°C, a short light day and feeding on aphids shortened larval development by 1.4 and 4.6 days, respectively. In combination, both factors caused a simple sum of separate impacts indicating the absence of their interaction. The duration of the pupal stage was independent of photoperiod, but decreased by 0.3 days in individuals fed on aphids. The proportion of the larvae that successfully completed preimaginal development was also independent of the light day length, but significantly depended on diet (84 and 60% in individuals fed on green peach aphids and on grain moth eggs, respectively). In sum, these results suggest that in the case studied, diet does not play the role of cue factor, but directly influences the larval growth and development of H. axyridis.     

Effects of attachment substrate,larval diet,and temperature on development and survival of immature Coquillettidia linealis (Skuse) and Coquillettidia xanthogaster (Edwards)

A sustainable colony allowed investigations into attachment substrates, diet and temperature in the development of the immature stages of Cq. linealis and its local congener Cq. xanthogaster. As immatures, these mosquitoes attach to submerged plants for respiration, and various field‐collected and laboratory‐raised plants were compared with non‐living substrates. Hydroponically‐grown wheat plant seedlings provided the most suitable attachment substrate. Development and survival of immatures to eclosion were investigated with three types and three quantities of a standardized diet. Development and survival at rearing temperatures between 10° and 30° C were also investigated. Diet type appeared to have little influence on development. However, at the lowest diet quantity, development to pupation was significantly reduced while the highest diet quantity impacted on the eclosion of pupae. Mean duration of immature development for Cq. linealis increased as temperature decreased. Time from 1^st instar to median eclosion did not differ significantly between 23° C (6.0 weeks) and 25° C (5.3 weeks), although it was 10 weeks at 20° C, and 30° C was lethal. For Cq. xanthogaster, similarly, mean duration of immature development increased as temperature decreased, with development time from 1^st instar to median eclosion significantly longer at 20° C (7.5 weeks) compared to 23° C (5.0 weeks) and 30° C (4.0 weeks).     

Effect of photoperiod on slow and fast developing individuals in aphidophagous ladybirds,Menochilus sexmaculatus and Propylea dissecta (Coleoptera: Coccinellidae)

The effects of environmental parameters on insect development have been studied extensively. But the reasons for 2 differential developmental rates within same cohort under varying environmental factors have not been explored. For the purpose, in this study the existence and stability of slow and fast development under 5 photoperiods (i.e., 8L: 16D, 10L : 14D, 12L : 12D, 14L : 10D and 16L : 8D; light and dark hours per day) and its effect on body mass and reproductive attributes in 2 aphidophagous ladybirds, Menochilus sexmaculatus (Fabricius) and Propylea dissecta (Mulsant) was examined on Aphis craccivora Koch at 27 ± 1 °C temperature. A clear bimodal (2 peaks, where the first peak represented the fast developing and the 2nd peak slow developing individuals) pattern of distribution at each photoperiod was found. The proportion of slow and fast developing individuals in a cohort differed with photoperiods. The slow developing individuals were more in numbers at 8L : 16D, in equal numbers at 14L : 10D and in less numbers at 16L: 8D, indicating that the variation in emergence was owing to exogenous cues influenced differential rates of mortality. Slow developing individuals had female biased sex ratio, higher longevity and lower body mass than fast developing individuals. Fast developing females laid higher numbers of eggs with higher egg viability than slow developing females. Study of such variations in development at different photoperiods is helpful to understand its role in the development of insects particularly ladybirds and permits the selection of fast developing bioagents for their use in biocontrol of pest species.     

Life history parameters of the cattle long‐nosed sucking louse,Linognathus vituli

Cattle sucking lice, Linognathus vituli (L.) (Phthiraptera: Linognathidae), were obtained from naturally infected cattle and maintained within ‘arenas’ affixed to the backs of cattle confined in controlled environment chambers maintained at a constant temperature of 15 °C. Temperatures measured within the arenas at an ambient temperature of 15 °C were constant at about 34 °C and only slightly above the temperature on nearby skin. The effect of temperature on egg development was determined using a gradient of temperatures between 25 °C and 41 °C. Eggs did not develop at temperatures of < 26 °C or > 39 °C. Survival of eggs was highest at temperatures of 30 °C and 35 °C. The earliest hatch was observed at 5 days post‐oviposition (at 33–35 °C). Development was extended to as long as 13 days at the lower temperatures. Kaplan–Meier survival probabilities were compared for lice kept at two densities in the arenas and showed there to be no effect of density on louse survival. Similarly, the mean number of eggs/louse/day over an 8‐day period was not influenced by louse density.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Induction and termination of diapause inOrius predatory bugs

Photoperiodic induction of reproductive diapause at 18°C was investigated in fourOrius [Heteroptera: Anthocoridae] species.Orius insidiosus (Say) displayed a long-day response with a critical photoperiod between L11:D13 and L12:D12. Diapause in this species was terminated rapidly when the temperature and/or the daylength were increased.Orius majusculus (Reuter) also displayed a long-day response. The critical photoperiod fell between L14:D10 and L16:D8. Diapause in this species was not terminated within 14 days when both temperature and daylength were increased. InOrius albidipennis (Reuter) no diapause could be induced at photoperiods varying from L8:D16 to L16:D8. InOrius tristicolor (White) a high proportion of diapause was found at all photoperiods tested. The effect of temperature on photoperiodic induction of diapause was studied inO. insidiosus at L10:D14. Diapause occurred at 18°C, 21°C and 25°C, but not at 30°C. Again, diapause was terminated rapidly after transfer to 25°C/L16:D8. Exposing only the nymphal instars 1–5 to short daylength was not enough to induce diapause in the whole population ofO. majusculus. Orius predatory bugs are used as biocontrol agents against western flower thrips,Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidael, in greenhouses. The consequences of photoperiodic induction of diapause for the success of early season releases ofOrius are discussed.     

Reproduction and population parameters of the Nearctic predator Geocoris punctipes at constant and varying temperature regimes

The heteropteran predator Geocoris punctipes (Say) has been used in augmentative biological control since 2000 to control Lepidoptera. However, surprisingly, few data are available about the influence of temperature on its population development, which is of key importance to plan the number and moment of releases to obtain sufficient pest reduction. The objective of this study was to evaluate daily and total fecundity, longevity and life table parameters (m_x, l_x, r_m, R, λ, T and TD) of G. punctipes at constant (16.8°C, 21.5°C, 24.5°C and 28.3°C) and corresponding varying (day/night) (21/11°C, 24/18°C, 27/21°C and 30/26°C) temperatures. Pairs of adult predators aged 24 h and originating from nymphs exposed to the same temperature regimes were kept at the above‐mentioned temperature regimes in Petri dishes containing Anagasta kuehniella (Zeller) eggs and an oviposition substrate. Tests were conducted in climatic chambers at the different temperature regimes and a RH 70 ± 10% and a 14L: 10D photoperiod. Reproduction, longevity and life table parameters were significantly affected by temperature, with clear differences between treatments at low (16.8°C, 21/11°C, 21.5°C, 24/18°C) or a high (24.5°C, 27/21°C, 28.3°C, 30/26°C) temperature regimes. Highest reproduction and fastest population growth of G. punctipes took place at average temperatures ranging from 24.5°C to 30°C, and neither reproduction nor population growth was negatively influenced by varying temperatures at any of the temperature regimes.     

Biology of Lysiphlebus fabarum following cold storage of larvae and pupae

Cold storage is one means of preserving parasitoids prior to release in augmentation biological control programs. This study examined the feasibility of storing larval and pupal stages of a sexual population of Lysiphlebus fabarum Marshall (Hymenoptera: Braconidae: Aphidiinae) at 6 ± 1 and 8 ± 1 °C, 50–60% r.h., and L14:D10 photoperiod. These life stages were stored for periods of 1, 2, and 3 weeks under fluctuating thermal regimes (2 h daily at 21 ± 1 °C). Generally, pupae gave better results than larvae, and 6 °C was better than 8 °C, considering wasp survival, wasp size (tibial and antennal lengths), egg load, and egg size. The best results were obtained with pupae stored for 2 weeks under a fluctuating temperature regime at 6 °C. Females emerging from this treatment did not differ from controls (developing directly at 21 °C) in body size, egg size, or progeny sex ratio, and suffered less than 20% mortality. Egg loads were reduced in these wasps, but the reductions were substantially less than occurred in other 2‐week‐storage treatments. Wasps stored in this manner successfully parasitized similar numbers of aphids as controls and produced similar progeny sex ratios. These results reveal a suitable set of low‐temperature conditions that can be used to delay the development of L. fabarum for 2 weeks with minimal impact on wasp fitness.     

Instars ofMicroplitis rufiventris [Hym.: Braconidae] and their relative developmental speed under different photoperiods

The internal parasiteMicroplitis rufiventris Kok. passes through 3 instars but moults 3 times within its host. The last moult occuring just at emergence time. The morphology of the egg and larval stages of the parasite are discussed. At 27°C and a photoperiod of 6 h (6L:18D) the endo-developmental cycle of the parasite can summarize as follows: Egg 18–24 h; instar 1,4 days (fighting phase 48 h; feeding phase 30–48 h); instar 2, 12–18 h and instar 3,3 days. The effect of different photoperiods on the relative speeds of the endo-developmental stages of the parasite at each of 30, 25, 20°C were carefully studied. At the first 2 temperatures, the short photoperiod (6L:18D) accelerated the development of larval instars, while both of 18L:6D or 0L:24D slowed down the development. Under the latter photoperiods some larvae failed to moult and had emergence problems. The influence of photoperiod is significantly noticeable at 20°C. The incubation period of the egg-stage was prolonged significantly at 18L:6D and the development of larval instars was significantly faster and refined at 6L:18D. The factor(s) inhibiting the development of the egg-stage perhaps differ from those affecting the larval development. The ventral area of the host mid-gut among malpighian tubes seems to be where the surplus parasite larvae are eliminated by physical attack. A physiologically suppressed parasite larva is able to attack its developed competitor of the same age. Teratocytes cells perhaps play a part in eliminating the surplus parasite larvae by physiological suppression.     

Temperature‐dependent developmental model of Neoseiulus californicus (McGregor) (Acari,Phytoseiidae)

The developmental time and survival of immature stages of Neoseiulus californicus were studied at nine constant temperatures (12, 16, 24, 24, 28 32, 36, 38 and 40°C), 60–70% RH, and a photoperiod of 16 : 8 (L : D) h. The total mortality of immature N. californicus was lowest at 24°C (4.5%) and highest at 38°C (15.2%). The total developmental time decreased with increasing temperature between 12°C (18.38 days) and 32°C (2.98 days), and increased beyond 32°C. The relationship between the developmental rate and temperature was fitted by five nonlinear developmental rate models (Logan 6, Lactin 1, 2 and Briere 1, 2). The nonlinear shape of temperature development was best described by the Lactin 1 model (r² = 0.98). The developmental variation of each stage was well described by the three‐parameter Weibull distribution model (r² = 0.91–0.93). The temperature‐dependent developmental models of N. californicus developed in this study could be used to determine optimal temperature conditions for its mass rearing, to predict its seasonal population dynamics in fruit tree orchards or greenhouse crops, or to develop a population dynamics model of N. californicus.     

Effect of temperature on the development of the immature stages ofBembidion lampros [Coleoptera: Carabidae]

The effects of temperature on survival and development of immature stages ofBembidion lampros were examined under controlled conditions in the laboratory. The duration of development was examined at 5°C, 12°C, 17°C, 19°C, 22°C, 25°C, 30°C and 32°C and found to be inversely related to temperature. Between 12°C and 30°C there was a significant difference in duration of egg development, but no significant difference in percentage of eggs hatched. The upper and lower lethal limits for egg development were estimated (c 31°C and 4°C respectively). There are 3 larval instars and one pupal stage. Development of larva plus pupa required a minimum of 20.9 days at 30°C. Between 22°C and 30°C survival from larva to the adult stage was 80%.