The apple sawfly, Hoplocampa testudinea: egg development and forecasting of egg hatch

The effect of temperature on egg development of the apple sawfly Hoplocampa testudinea Klug was studied under controlled conditions. Based on a linear model describing the relationship between temperature and developmental rates a thermal threshold of 6.9 °C and a thermal constant of 85 day-degrees were established. The experimental results were used to expand and parameterise an existing phenology model to include egg development and hatching of first instar larvae. The model thus became a valuable forecasting tool not only for adult emergence and the optimal monitoring period but also for egg hatch and the best timing of control measures against first instar larvae. The model was validated by comparing predictions with independent observations on egg-laying and subsequent hatching of first instar larvae under semi-field conditions.     

Development of Apheliuus mali an endoparasitoid of woolly apple aphid, Eriosoma lanigerum at different temperatures

Developmental times for both sexes of Aphelinus mali (Haldeman) (Hymenoptera: Aphelinidae) an endoparasitoid of woolly apple aphid, Eriosoma lanigerum (Hausmann) (Hemiptera: Pemphigidae) were studied at 13, 15, 18, 20, 25 and 30°C and compared with those of E. lanigerum. Mean developmental times ranged from 11.7–53.3 days for males, 11.8–55.4 days for females and 11.8–54.4 days for both sexes combined and were significantly inversely related to temperature. A good linear model fit (r²>0.99) between developmental rate and temperature in the range 13–30°C was observed. Results indicate significant differences between developmental times of males and females at 13, 18, 20, and 25°C but no differences at 15 and 30°C. The notional developmental threshold was 8.3°C for both males and females. Compared with its host, A. mali has a higher lower developmental threshold. An average of 252.8, 256.7 and 254.8 degree-days (DD) above the lower threshold were required by A. mali to complete development from time of oviposition to adult emergence for males, females and both sexes combined, respectively. Field experimentation also indicated that the developmental time of A. mali lags significantly behind that of its aphid host throughout the year. These findings are discussed in relation to its status as a biological control agent for E. lanigerum.     

The apple sawfly, Hoplocampa testudinea: a temperature driven model for spring emergence of adults

The effect of temperature on post-diapause development of the apple sawfly Hoplocampa testudinea Klug was studied under controlled conditions. Survival was found to decrease at temperatures higher than 15°C, the lethal temperature being close to 25°C. The lower thermal threshold was 4.5°C; the values for males and females did not differ significantly. However, different values were established for unprotected individuals, in vermiculite and in potting compost respectively. Based on the overall mean and variance of the developmental time and the overall thermal threshold a simple soil temperature driven phenology model was constructed using a time- varying distributed delay. The validation with independent field and semi-field data revealed the model to be a reliable tool to predict apple sawfly phenology and hence to determine the optimal time for the installation of white sticky traps or for control measures such as insecticide treatments.     

Relationship between cumulative heat units and fruit-body emergence of ectomycorrhizal fungusTricholoma bakamatsutake in the fields

The relationship between cumulative heat units and fruit-body emergence ofTricholoma bakamatsutake was examined by using the method used for estimating dates of adult emergence of insects. Fruit-body numbers at a study site at Tateyama, Chiba Prefecture and records at a weather station near the site from 1987 to 1993 were used. For the seven years, daily average temperatures above the developmental threshold were cumulated for the days following induction of fruit-body development. The developmental threshold was taken to be 0°C, and the day when the daily minimum temperature fell to near 20°C in each year was taken as the day of induction of fruit-body development. A linear relationship between fruit-body numbers and daily average temperatures was observed as follows: Y=0.00376X + 1.873 (r=0.810), where Y is the probit of percentage of cumulative fruit-bodies, and X is the heat units above 0°C cumulated for the days following induction of fruit-body development. X for Y=5, which represents the cumulative emergence of 50% of fruit-bodies, was estimated to be 832 day-degrees.     

The life-cycle of Apatania fimbriata Pictet in the Breitenbach

The life-cycle of Apatania fimbriata (Pictet, 1843) (Trichoptera: Limnephilidae) was studied in the Breitenbach (Hesse, Federal Republic of Germany) and in the laboratory at 6 °C, 10 °C and 14 °C at 14 h day length. Embryogenesis was mainly temperature dependent in the 4 °C–18 °C range. At a constant 22 °C embryos developed to well-differentiated larvae which did not hatch. In the laboratory experiments, developmental rates increased with increasing temperature (6 °C, 10 °C and 14 °C); larvae went through a dormancy in the third and fourth larval instars; in the field this occurred between November and February. The dormancy synchronized the larval development. Laboratory studies led to the conclusion that the dormancy was not temperature dependent but endogenously determined. The emergence pattern of adults showed a clear diurnal pattern, with a maximum emergence approximately two hours after sunset. A comparison of several years emergence studies showed a decreasing number of females with increasing distance from the spring. A comparison of the dry weight of females between two traps at the stream showed a significantly higher weight from the downstream site. For males no between-site differences were observed.     

Relationships between temperature, development and survival of different life stages of the tomato fruit fly, Neoceratitis cyanescens

The development and survival of female Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae) from egg to complete ovarian maturation were studied in the laboratory at five different constant temperatures: 15, 20, 25, 30, and 35 °C. The aim of this study was to get information on the influence of temperature on pre-mature stages, as a prerequisite to optimise rearing procedures and to understand temporal and geographical patterns of fruit fly occurrence. The developmental rate of the different life stages increased linearly with increasing temperatures up to 30 °C. The fastest development of pre-mature stages was recorded at 30 °C (22±1 days) and the slowest at 15 °C (98±3 days). The day-degrees requirements (K) to complete total development were 432.6 day-degrees. Lower temperature thresholds were 11.4, 11.9, 10.0, and 11.1 °C for egg, larval, pupal stages and ovarian maturation, respectively. The number of adults obtained from an initial batch of 100 eggs reached a maximum (64) at 25 °C. At 35 °C, no adults emerged. Larval developmental time was significantly shorter in green tomato fruits than in potato tubers at 15, 20, and 25 °C. Mortality rate of larvae was higher in green tomato fruits than in potato tubers at 25 and 30 °C.     

Effect of temperature and sperm supply on the reproductive potential ofTetranychus evansi (Acari: Tetranychidae)

The biology ofTetranychus evansi Baker and Pritchard was studied at five different temperatures on excised leaves ofSolanum douglasii Dunal. The theoretical lower threshold temperature of development was ca. 13°C. The combined immature stages required ca. 148 day-degrees for completion. Progressively later initiation of mating resulted in higher peak proportions of females produced, up to an 8-day period between emergence and mating at 30°C. The proportion of male offspring always increased towards the end of the oviposition period, apparently because of sperm depletion. The calculated intrinsic rate of increase was highest (0.432) at 35°C, the highest temperature tested. At that same temperature, daily oviposition rate and net reproductive rate were maximum (9.5 and 111.77, respectively), and mean generation time and doubling time were minimum (10.92 and 1.6 days, respectively).     

Life history study of Thrips setosus

Development, reproduction and population growth of Thrips setosus Moulton (Thysanoptera, Thripidae), reared on a leaf of kidney bean, was studied under six different constant temperatures, and the effect on reproduction of short photoperiod during immature stages was examined. Survival rates from hatch to adult were more than 67.5% at temperatures between 17.5 and 27.5 °C, but less than 55% at 30 °C. Developmental rates increased linearly as rearing temperature increased. A total of 181.1 degree-days, above a developmental zero of 12.5 °C, were required to complete development from egg to adult oviposition. These data were related to records of field temperatures in Kurashiki in western Japan, and an estimate produced that, under outdoor conditions, a maximum of between seven and 12 generations could have developed annually between 1990 and 1999. There were no significant differences in mean adult longevity and mean fecundity among three temperatures (20, 22.5 and 25 °C). The intrinsic rate of natural increase (r _m) was 0.1997 at 25 °C. Reproductive diapause was induced by a photoperiod less than 12 h at 20 °C.     

The effects of temperature and host density on the development and survival of Copidosoma koehleri

Developmental rates for Copidosoma koehleri Blanchard (Hymenoptera: Encyrtidae) and its host, Phthorimaea operculella (Zeller), were determined at 10, 15, 23, 27, 29, 32, 34 and 35°C from host egg to adult. The developmental rates determined for both species showed good fit to mathematical models of insect development. At 15°C mean emergence of adult C. koehleri was 15 days after P. operculella adults emerged. At 29°C mean emergence of C. koehleri was only 5 days after that of P. operculella. P. operculella developed at 35°C, but parasitized larvae, and therefore C. koehleri, did not tolerate temperatures above 32°C. In the presence of C. koehleri, host survival was low (3%) at low host egg densities, but greater (20%) at higher host densities. Parasitized larvae of P. operculella were less able to compete for food resources, as measured by adult emergence.     

Pupation site and emergence time influence the mating success of bagworm females,Oiketicus kirbyi

In commercial oil palm plantations in Costa Rica, we tested the hypotheses that pupation site and emergence time affect the mating success of protogynous female bagworms,Oiketicus kirbyi (Guilding) (Lepidoptera: Psychidae). Greater proportions of female than male pupae on upper leaves of oil palms and greater proportions of mated females in the upper rather than lower crown strata support the hypothesis that selection of pupation site by female larvae influences the mating success of adults. Increasing captures of males with increasing trap height further suggest that enhanced mating success of females in tree tops may be attributed either to most effective dissemination of sex pheromone on higher sites, or to males foraging predominantly in the upper strata of oil palms. As the majority of females pupated in the middle rather than upper crown of oil palms, selection of pupation site by females may be affected by additional as yet unknown factors. Emergence of females significntly preceded emergence of males. Increasing proportions of mated females throughout the emergence seasons probably resulted from an increased ‘availability’ of males. In tropical rainforests with local variations inO. kirbyi developmental time and stage, protogyny may represent an evolutionary strategy that furthers outbreeding.     

Seasonal size distribution of developmental stages of sub-antarctic copepod

Drepanopus pectinatus, the most numerous species of Copepod in the Kerguelen Archipelago completes 4 generations a year. The succession of generations follows the hydrological seasons. Each generation may be characterized by the cephalothorax length of the developmental stages C1 to C6. The mean lengths of stages C2 to C6 are shown to be significantly different (P > 0.05) between seasons (i.e. between generations). The size-classes have a unimodal distribution from stages C1 to C4. For copepodite C5 and adults, bimodal histograms reflect the differentiation of males and females.A reverse trend of the development pattern of lengths is demonstrated in this subantarctic species: a decrease with decreasing temperatures (6° to 2°) from April to September, and an increase with the rising temperatures (2° to 7°) from October to February.Two populations of females are distinguishable in spring and summer, by their small (0.950–1.299 mm length of cephalothorax, 55%) or large (1.300–1.800 mm, 45%) size, respectively.The differences in mean length cannot be explained by their relation to the mean temperature alone. Pigment content is suspected to play a role.     

Development, survival and reproduction of Euseius finlandicus (Acari: Phytoseiidae) at different constant temperatures

Development, survival and reproduction of Euseius finlandicus Oudemans were studied at seven constant temperatures (15, 20, 25, 27, 30, 32 and 34°C) in the laboratory. Within the temperature range tested, developmental period from egg to adult varied from 148 to 360.5h and 133.7 to 336.5h for females and males, respectively. The lower thermal threshold for immature development for females and males was 8.9 and 6.4°C, respectively. Survival during immature development exceeded 90% at all the temperatures from 15 to 32°C, but at 34°C an abrupt decline was recorded. Female longevity decreased gradually from 82.7d at 15°C to 12.2 d at 34°C. The mean generation time ranged from 44.3d at 15°C to 15.9d at 32°C. The highest r _m value (0.2817) was obtained at 30°C and the lowest at 15°C (0.0976). Temperatures above 30°C had an adverse effect on population increase.     

Plasticity in life-history traits of the bark beetle Ips calligraphus as influenced by phloem thickness

Summary Reproduction and development of Ips calligraphus (Germar) (Coleoptera: Scolytidae) occur primarily in the phloem (inner bark) tissue of its pine hosts. In slash pine (Pinus elliottii Engelm. var. elliottii), phloem thickness can vary widely from tree to tree. We compared adult residence time and fecundity and progeny development time, body size, and sex ratio of this beetle when reared in slash pine bolts and bark slabs with phloem thicker or thinner than average adult body width. Most studies were conducted at 20°, 25°, and 30° C. Residence time of parent males and females decreased with increasing temperature but it was not affected by phloem thickness. Residence time of parent males was about 2 days shorter than that of parent females at each temperature. Reproductive fitness was greater in thick phloem at all temperatures as measured by faster larval development, earlier emergence of progeny (F ₁ ) adults, and emergence of more progeny adults per parent female. Progeny survival was similar between phloem those from thick phloem, and their sex ratio was female biased (1:2 vs 1:1, male:female). When allowed to inhabit thick phloem, the progeny (F ₂) of adults from thin phloem were just as large and of similar sex ratio (1:1) as were progeny of F ₁ adults from thick phloem. The high degree of plasticity in the life-history traits of this beetle may enable it to successfully colonize pines that vary widely in phloem thickness.     

Cold storage of Trichogramma ostriniae reared on Sitotroga cerealella eggs

Efficient storage of the biological controlagent Trichogramma ostriniae couldimprove current parasitoid production methodsby making the system more flexible andefficient. Initial studies compared emergencerates of T. ostriniae reared onSitotroga cerealella eggs held at 6 °C, 9 °C, 12 °C,15 °C, and 24 °C for up to 8 weeks after parasitism.At 15 °C, emergence occurred in <2 weeks.Emeregence was >80% for parasitized eggsstored at 9 °C and 12 °C for 4 and 6 weeksrespectively. Storage at 6 °C caused asignificant decline in emergence after 2 weeks. Subsequent trials focused on fitness of storedT. ostriniae. Percentage of emergedfemales parasitizing eggs, female longevity,and fecundity were quantified after storage.The percentage of females successfullyparasitizing O. nubilalis eggs wasgreatest for the 9 °C four-week treatment(100%). Compared to 24 °C controls, storage at12 °C for 6 weeks or at 9 °C for 8 weeks reduced thepercentage of females parasitizing. Longevityof females held in cold storage was generallyless than that of controls. Rates of parasitismby stored Trichogramma was generallysimilar to controls after 2 to 4 weeks' storageat 9 °C and 12 °C but declined with storage longerthan 4 weeks. Emergence of progeny of cold-storedfemales was lower than controls for alltreatments. The percentage of female progenyfrom cold stored females was comparable tocontrols up to 4 weeks of storage.     

Rapid cold hardening in the olive fruit fly Bactrocera oleae under laboratory and field conditions

A rapid cold hardening response was studied in females and males of the olive fruit fly Bactrocera (Dacus) oleae. When laboratory-reared females and males were transferred and maintained from the rearing temperature of 24 °C for 2 h to –6.5 °C approximately 5% survived. However, conditioning of both females and males for 2 h at various temperatures from 0 to 10 °C before their exposure for 2 h to –6.5 °C increased survival to 80 to 92%. A similar rapid cold hardening response in both females and males was also induced through gradual cooling of the flies at a rate of approximately 0.4 °C per min. The rapid increase in cold tolerance after prior conditioning of the flies to low temperatures, was rapidly lost when they returned to a higher temperature of 24 °C. In the field, in late February and early March, females and males were capable of a rapid cold hardening response. After exposure to the critical temperature they suffered a high mortality when tested in the afternoon and low mortality early in the morning on consecutive days, probably because of differences in the prevailing field temperatures a few hours before testing. This plasticity of cold tolerance gained through rapid cold hardening may allow the flies to survive during periods of the year with great fluctuation in circadian temperatures.     

Pheromone-trap catch in relation to the phenology of codling moth (Cydia pomonella)

Relationships between a pheromone-trap catch, adult emergence and penetration of fruit by first-instar larvae of Cydia pomonella were investigated from 1975 to 1977 in an orchard in South West England. For the first generation the times of moth emergence and catch in the pheromone trap were not significantly different; nor were male and female emergence times. The catch of moths of the first generation in the pheromone trap anticipated the appearance of their larvae in the fruit by 140–169 day-degrees > 10 °C. Eggs hatched after 94 day-degrees in the laboratory but in the orchard, wind and sunshine modified the microclimate so that the number of day-degrees required for egg development, as measured by standard meteorological instruments, was affected by wind and sunshine but development lasted on average about 90 day-degrees. This indicated a lag of 50–80 day-degrees between the curves for trap catch and oviposition: the pre-oviposition period in the orchard was shorter than expected from laboratory studies. In 1975 and 1976, some larvae developed to produce a second generation of moths which gave rise to a second generation of eggs and larvae, after an interval of 161 day-degrees in 1975, but only 41 day-degrees in 1976, indicating that in 1976 some eggs were laid before moths of the second generation were trapped. These results indicate that the first insecticidal spray against first-generation larvae should be applied about 140 day-degrees after the start of the reference week in which five moths of the spring brood are caught per trap. A second spray, if required to maintain insecticidal cover, should be applied about 100 day-degrees later. A spray should be applied against second-generation larvae immediately after the reference week for second-generation moths.     

Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development

During direct development the butterfly Lycaena tityrus was previously found to display sex-related reaction norms in response to temperature. Based on selection for protandry in males and fecundity selection for larger females, males favoured early emergence over large size, leading to a dramatic weight loss at higher temperatures, whereas females maintained similar weights throughout. Because males were able to avoid a weight reduction relative to females in spite of their shorter development at lower temperatures, sexual size dimorphism existed at higher temperatures only. In the present paper we compare sexual differences in life-history traits in L. tityrus between direct and diapause development at 25 °C. We demonstrate that, regardless of developmental pathway, protandry persisted and relative sexual size dimorphism, with females being larger, remained unchanged. Although diapausing individuals were less time-constrained, allowing them to grow to considerably higher final weights in both sexes, males were not able to reduce their weight loss relative to females. This is explained by the pressure to gain a developmental advantage solely during post-diapause development, whereas direct developing males may spread the burden over the whole larval period. Our results highlight the importance of considering sexual differences in selective pressures, which may influence central life-history traits in manifold ways.     

The sublethal effects of tebufenozide on the precopulatory and copulatory activities of Choristoneura fumiferana and C. rosaceana

The sublethal effects of tebufenozide, an ecdysone agonist, on the reproductive biology of Choristoneura fumiferana (Clem) and of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), treated during the larval stage, were evaluated using two treatment methods: the force‐feeding method and the diet method. The percentage of mortality and the developmental time of survivors increased linearly with the concentration of tebufenozide used. This ecdysone analogue proved to be more toxic to C. fumiferana than to C. rosaceana. In C. rosaceana, the weight of males and females decreased proportionally with the dose ingested, but females were affected to a greater extent. This difference might be due to a greater consumption of the treated diet, or to a differential vulnerability to tebufenozide. Tebufenozide did not modify the pre‐copulatory activities associated with chemical communication in the females. However, the consumption of tebufenozide delayed ovarian maturation, causing a reduction in the fecundity of females. Treated males had smaller spermatophores and fewer eupyrene sperms in their bursa copulatrix and spermatheca, along with lower mating success. In C. fumiferana, tebufenozide delayed the females’ onset time of calling the first night after emergence, but did not affect the mean time spent calling or the production of the main component of the sex pheromone. The males showed significantly greater difficulty in executing oriented flight in a wind tunnel, although their mating success was not affected. We concluded that tebufenozide interferes with various aspects of the reproductive biology of males and females of C. fumiferana and C. rosaceana, including some pre‐copulatory behaviors associated with sex pheromone communication.     

Bionomics and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum, at constant temperatures

Development, survival, reproduction and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum (Hausmann) (Hemiptera: Aphididae) at constant temperatures of 10, 13, 15, 20, 25, 30 and 32°C are reported. The developmental times of all life stages were inversely related to temperature ranging from 10 to 30°C. Span of total development (time from birth to adulthood) decreased from 57.8 days at 10°C to 11.7 days at 30°C and increased to 16.8 days at 32°C. A good linear model fit (R²>0.96) between developmental rate and temperature in the range 10–25°C was observed for all life stages. The lower developmental threshold was estimated at 5.8°C for instar I, 4.8°C for instar II, 4.9° for instar III and 4.4°C for instar IV. The lower temperature threshold for total development was estimated at 5.2°C. The upper developmental limit was found to be 32°C. Mean degree-day accumulations required for completion of instars I, II, III, IV and total development were: 125.6, 51.0, 47.7, 50.7 and 267.6, respectively. Fecundity, larviposition period and adult longevity were reduced with increasing temperature. Net reproductive rate was greatest at 15°C whereas intrinsic rate of increase peaked at 25°C. Optimal performance, as measured by fecundity, survival and intrinsic rate of increase, ocurred in the range 13–25°C.     

Effects of different temperatures on the life history of Evania appendigaster L. (Hymenoptera: Evaniidae), a solitary oothecal parasitoid of Periplaneta americana L. (Dictyoptera: Blattidae)

The influence of temperatures on the life parameters of the solitary oothecal parasitoid Evania appendigaster, was investigated in the laboratory. Parasitized oothecae of Periplaneta americana were left to develop under seven constant temperatures: 15, 17, 20, 25, 30, 35, and 40 °C. At the end, we found that: (i) E. appendigaster was able to complete development within the temperature range of 17–34 °C; (ii) mean adult longevity decreased as temperature increased, with the temperature of 40 °C being fatal in a matter of hours; (iii) males lived longer than females between 15 and 30 °C; (iv) adult emergence rate was the highest at 25 °C, and (v) no wasps emerged at 15 or 40 °C. Non-emerged oothecae contained either unhatched eggs or dead larvae. We determined the theoretical lower developmental threshold and thermal constant for the complete development as 12.9 °C and 584.8 day-degrees for males, and 13.1 °C and 588.2 day-degrees for females, respectively. A good balance between faster development, maximum adult longevity and good egg viability was obtained between 25–30 °C, and that would be the best temperature range for rearing E. appendigaster.