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Michael P Parrella - One of the best experts on this subject based on the ideXlab platform.

  • improving the biological control of leafminers diptera agromyzidae using the sterile insect technique
    Journal of Economic Entomology, 2006
    Co-Authors: Roy Kaspi, Michael P Parrella
    Abstract:

    The leafminer Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) is a worldwide pest of ornamental and vegetable crops. The most promising nonchemical approach for controlling Liriomyza leafminers in greenhouses is regular releases of the parasitoid Diglyphus isaea (Walker) (Hymenoptera: Eulophidae). In the current study, we examine the hypothesis that the use of D. isaea for biological control of leafminers in greenhouse crops may be more practical and efficient when supplemented with additional control strategies, such as the sterile insect technique (SIT). In small cages, our SIT experiments suggest that release of sterile L. trifolii males in three sterile-to-fertile male ratios (3:1, 5:1, and 10:1) can significantly reduce the numbers of the pest offspring. In large cage experiments, when both parasitoids and sterile males were released weekly, the combined methods significantly reduced mine production and the adult leafminer population size. Moreover, a synergistic interaction effect between these two methods was found, and a model based on our observed data predicts that because of this effect, only the use of both methods can eradicate the pest population. Our study indicates that an integrated pest management approach that combines the augmentative release of the parasitoid D. isaea together with sterile leafminer males is more efficient than the use of either method alone. In addition, our results validate previous theoretical models and demonstrate synergistic control with releases of parasitoids and sterile insects.

  • abamectin compatibility with the leafminer parasitoid Diglyphus isaea
    Biological Control, 2005
    Co-Authors: Roy Kaspi, Michael P Parrella
    Abstract:

    Abstract The most promising non-chemical approach for controlling Liriomyza leafminers in greenhouses is the augmentative/inoculative releases of the parasitoid Diglyphus isaea. However, the insecticide/miticide abamectin is still an important chemical control method against Liriomyza leafminers and the mite complex that attacks numerous greenhouse crops. It is not uncommon for abamectin to be applied where Diglyphus is being used for leafminer control. Consequently, there is a need to develop compatibility data for this material and Diglyphus. The effect of abamectin on D. isaea (adults and larvae) was investigated using laboratory and greenhouse experiments. Direct application and uptake of abamectin had a significant negative effect on the survival of D. isaea adult (both sexes). Abamectin residue on chrysanthemum leaves had a significant negative effect on adult D. isaea female longevity up to 5 days after application. Abamectin was lethal for D. isaea larvae when applied directly to larvae or when contaminated leafminer larvae were consumed by parasitoid larvae. However, the percent emergence of D. isaea was not affected by abamectin treatments when applied to chrysanthemum plants that contained parasitoid larvae. Moreover, the longevity of these emerged adults was not affected by abamectin applications. In light of these data, abamectin compatibility with D. isaea for leafminer control in IPM programs for greenhouses is discussed.

Komivi Senyo Akutse - One of the best experts on this subject based on the ideXlab platform.

  • interaction between two leafminer parasitoids halticoptera arduine hymenoptera pteromalidae and Diglyphus isaea hymenoptera eulophidae in the management of liriomyza huidobrensis diptera agromyzidae
    Environmental Entomology, 2018
    Co-Authors: Komivi Senyo Akutse, Samuel K Muchemi, Claus P. W. Zebitz, Christian Borgemeister, Caroline N Foba, Sunday Ekesi, Komi K M Fiaboe
    Abstract:

    Liriomyza spp., leafminer flies (Mik; Diptera: Agromyzidae), are economically important quarantine pests that puncture and mine leaves and fruits of various horticultural crops worldwide, affecting yield and trade. Halticoptera arduine (Walker; Hymenoptera: Pteromalidae), a key parasitoid from the pests' areas of origin in South America, was introduced as a potential alternative management strategy. Prior to H. arduine release, its potential interactions with the dominant local ectoparasitoid, Diglyphus isaea (Walker; Hymenoptera: Eulophidae), were assessed. Halticoptera arduine and D. isaea were released in single, sequential and simultaneous combinations on Liriomyza huidobrensis (Blanchard; Diptera: Agromyzidae) to evaluate possible effect on the parasitism rate, reproduction and host mortality. The combination of both parasitoids did not significantly affect the specific parasitism rates of either of them, an indication that H. arduine and D. isaea can coexist. Parasitism rates of the exotic H. arduine were significantly superior to the indigenous D. isaea in all release combinations except when both species were released simultaneously. While 50 individuals of D. isaea resulted only in 21.23 ± 2.1% parasitism, 50 parasitoids composed of 25 H. arduine and 25 D. isaea caused 53.27 ± 4.99%. Both parasitoids further induced significant nonreproductive host mortalities. Both parasitoids' F1 progenies sex ratios were female-biased in all parasitoid release combinations except in single release of D. isaea with a balanced sex ratio. The improvement in D. isaea's sex ratio induced by the presence of H. arduine suggests a synergetic effect on D. isaea's reproductive performance. The introduction of H. arduine in horticulture production systems may therefore improve natural control of Liriomyza leafminers in East Africa.

  • interaction between chrysocharis flacilla and Diglyphus isaea hymenoptera eulophidae two parasitoids of liriomyza leafminers
    Journal of Economic Entomology, 2018
    Co-Authors: Komivi Senyo Akutse, Samuel K Muchemi, Claus P. W. Zebitz, Christian Borgemeister, Caroline N Foba, Sunday Ekesi, Komi K M Fiaboe
    Abstract:

    Agromyzid Liriomyza leafminer flies are a major threat to horticultural production in East Africa with low natural control reported. The endoparasitoid Chrysocharis flacilla (Walker; Hymenoptera: Eulophidae) was introduced from Peru into quarantine facilities at ICIPE in Kenya for a leafminer classical biological control program. Interaction assays with one of the dominant local parasitoids, Diglyphus isaea (Walker; Hymenoptera: Eulophidae), using Liriomyza huidobrensis (Blanchard; Diptera: Agromyzidae) was assessed through sole, simultaneous and sequential releases. C. flacilla resulted to superior host parasitism rates over D. isaea. When used separately, specific parasitism rates of D. isaea and C. flacilla were 26.33 ± 2.07% and 60.27 ± 2.53% respectively but, when simultaneously used, the total parasitism rose to 72.96 ± 4.12%. Presence of C. flacilla after D. isaea reduced significantly parasitism rate of D. isaea. Both parasitoids caused separately and simultaneously additionally significant nonreproductive host mortalities of between 48.33 ± 3.75% and 69.33 ± 3.92 for D. isaea and C. flacilla respectively. Sex ratios of C. flacilla and D. isaea F1 progenies were female biased and were not affected by interspecific interactions. Implications of these results for subsequent combined use of C. flacilla and D. isaea against Liriomyza leafminers in East Africa are discussed.

  • prospects of fungal endophytes in the control of liriomyza leafminer flies in common bean phaseolus vulgaris under field conditions
    Biocontrol, 2016
    Co-Authors: Komivi Senyo Akutse, Komi K M Fiaboe, Sunday Ekesi, Jane Wairimu Gathage, Zipporah Osiemo Lagat, Nguya K Maniania
    Abstract:

    Field trials were carried out for two seasons in two sites (Sagana and Naromoru, Central province of Kenya) to evaluate the prospects of endophyte isolates of Beauveria bassiana (Balsamo) Vuillemin G1LU3 and Hypocrea lixii Patouillard F3ST1 for the control of Liriomyza leafminer in common bean Phaseolus vulgaris L. crops through seeds inoculation. Autodissemination device (AD) treated with conidia of Metarhizium anisopliae (Metschnikoff) Sorokin ICIPE 20 was also added as a treatment. Leafminer infestation was not significantly different during the first season but was higher in the controls than in endophtyte treatments at both sites during the second season. Three key Liriomyza species [(L. huidobrensis Blanchard, L. sativae Blanchard and L. trifolii (Burgess)] and six parasitoid species [(Opius dissitus Muesebeck, Phaedrotoma scabriventris Nixon, Diglyphus isaea Walker, Neochrysocharis formosa Westwood, Hemiptarsenus varicornis Girault and Halticoptera arduine (Walker)] were identified during the trials. Leafminer infestation, number of pupae, leafminer flies and parasitoids emergence and yield were the parameters evaluated. Both isolates successfully colonized different parts of P. vulgaris plants, although the colonization was higher with H. lixii F3ST1 than B. bassiana G1LU3 at both sites. The mean number of pupae from the infested leaves varied between 141–252 and 331–416 in endophyte and control treatments, respectively, during the first season and from 110–223 to 366–523, respectively, in endophyte and control treatments during the second season. There were no significant differences among the treatments in the number of parasitoids that emerged from pupae. Higher yield of P. vulgaris seeds was obtained in endophyte than in control treatments. The inclusion of AD treatment did not have significant effect on all the parameters evaluated, except yield. Results of the present study suggest that both fungal isolates hold potential for pest management and could be considered for the control of leafminer flies. However, there is the need to confirm these results on large-scale trials.

  • differential effects of pesticide applications on liriomyza huidobrensis diptera agromyzidae and its parasitoids on pea in central kenya
    Journal of Economic Entomology, 2015
    Co-Authors: Daisy Salifu, Komivi Senyo Akutse, M M Guantai, C P K O Ogol, J M Kasina, Komi K M Fiaboe
    Abstract:

    Three Liriomyza species [Liriomyza huidobrensis (Blanchard), Liriomyza trifolii (Burgess), and Liriomyza sativae Blanchard] have been reported as the most important leafminer pests in vegetable production systems in Africa. In Kenya, farmers rely on indiscriminate synthetic insecticides use. On-farm field investigations were set up at three different locations (Sagana, Kabaru, and Naromoru) in central Kenya to determine the effect of pesticide application on the abundance of leafminers and their parasitoids under three management practices, namely: farmer practice (FP), reduced pesticide use (RP), and a control with no use of pesticides (CO). In addition, laboratory experiments were designed to test the effect of commonly used pesticides in pea production systems in central Kenya--Dimethoate, Dynamec, Thunder, Cyclone, Bestox, Folicur, Milraz, and Bulldock--on L. huidobrensis and two of its parasitoids, Diglyphus isaea Walker and Phaedrotoma scabriventris Nixon. The mean numbers of leafminer flies in control treatment were higher than in RP and FP in both first and second seasons across all sites, but RP and FP did not differ significantly. Parasitoid numbers were very low and there was no much variation between treatments at each location in both first and second seasons. No significant differences were observed between the three management practices with regards to the yield measurements. In the laboratory, the estimated LD50 values for L. huidobrensis larvae were all more than two times higher than the recommended dosages, while the LD50 of adults were below the recommended dosages. The estimated LD50 values for the parasitoids were much lower than recommended dosages for all pesticides except Thunder. This study, therefore, demonstrates that the pesticides currently used do not control the Liriomyza leafminer larvae that constitute the most destructive stage of the pest, but are rather detrimental to their parasitoids. In addition, the current low level of parasitoids recorded under field conditions even where no pesticide was used during this study, warrants consideration of classical biological control programs.

  • interactions between phaedrotoma scabriventris nixon hymenoptera braconidae and Diglyphus isaea walker hymenoptera eulophidae parasitoids of liriomyza huidobrensis blanchard diptera agromyzidae
    Biological Control, 2015
    Co-Authors: Komivi Senyo Akutse, Komi K M Fiaboe, Nguya K Maniania, J Van Den Berg, Sunday Ekesi
    Abstract:

    Abstract Liriomyza leafminer flies represent a serious threat to horticultural production in East Africa. Total field parasitism rates recorded in Kenya are below 5%, with the indigenous ectoparasitoid Diglyphus isaea Walker being one of the key parasitoid species. The International Centre of Insect Physiology and Ecology ( icipe ), in collaboration with the International Potato Centre (CIP), imported into Kenya the endoparasitoid Phaedrotoma scabriventris Nixon to improve natural control of leafminers. The objective of this study was to investigate the interactions between D. isaea and P. scabriventris when used together for the biological control of Liriomyza species. These interactions were studied under laboratory conditions, using treatments that involved single, simultaneous and sequential releases of the different parasitoid species onto plants infested by L. huidobrensis larvae. While used separately, parasitism rates of D. isaea and P. scabriventris were 30.4 ± 10.9% and 63.6 ± 7.7% respectively. However, when used simultaneously, the total parasitism rate increased to 77.0 ± 5.3%. Although P. scabriventris had no effect on D. isaea , the presence of D. isaea reduced the specific parasitism rate of P. scabriventris . In addition, both parasitoids induced leafminer mortality through larval-feeding and stinging. However, feeding and stinging mortality induced by D. isaea (41.9 ± 9.1%) was significantly higher compared to P. scabriventris (11.9 ± 8.7). Similarly, pupal mortality due to feeding and stinging activity was 49.1 ± 6.5% and 21.6 ± 1.9% when exposed to D. isaea and P. scabriventris respectively. The implication for simultaneous use of both parasitoids in East Africa is discussed.

Roy Kaspi - One of the best experts on this subject based on the ideXlab platform.

  • improving the biological control of leafminers diptera agromyzidae using the sterile insect technique
    Journal of Economic Entomology, 2006
    Co-Authors: Roy Kaspi, Michael P Parrella
    Abstract:

    The leafminer Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) is a worldwide pest of ornamental and vegetable crops. The most promising nonchemical approach for controlling Liriomyza leafminers in greenhouses is regular releases of the parasitoid Diglyphus isaea (Walker) (Hymenoptera: Eulophidae). In the current study, we examine the hypothesis that the use of D. isaea for biological control of leafminers in greenhouse crops may be more practical and efficient when supplemented with additional control strategies, such as the sterile insect technique (SIT). In small cages, our SIT experiments suggest that release of sterile L. trifolii males in three sterile-to-fertile male ratios (3:1, 5:1, and 10:1) can significantly reduce the numbers of the pest offspring. In large cage experiments, when both parasitoids and sterile males were released weekly, the combined methods significantly reduced mine production and the adult leafminer population size. Moreover, a synergistic interaction effect between these two methods was found, and a model based on our observed data predicts that because of this effect, only the use of both methods can eradicate the pest population. Our study indicates that an integrated pest management approach that combines the augmentative release of the parasitoid D. isaea together with sterile leafminer males is more efficient than the use of either method alone. In addition, our results validate previous theoretical models and demonstrate synergistic control with releases of parasitoids and sterile insects.

  • abamectin compatibility with the leafminer parasitoid Diglyphus isaea
    Biological Control, 2005
    Co-Authors: Roy Kaspi, Michael P Parrella
    Abstract:

    Abstract The most promising non-chemical approach for controlling Liriomyza leafminers in greenhouses is the augmentative/inoculative releases of the parasitoid Diglyphus isaea. However, the insecticide/miticide abamectin is still an important chemical control method against Liriomyza leafminers and the mite complex that attacks numerous greenhouse crops. It is not uncommon for abamectin to be applied where Diglyphus is being used for leafminer control. Consequently, there is a need to develop compatibility data for this material and Diglyphus. The effect of abamectin on D. isaea (adults and larvae) was investigated using laboratory and greenhouse experiments. Direct application and uptake of abamectin had a significant negative effect on the survival of D. isaea adult (both sexes). Abamectin residue on chrysanthemum leaves had a significant negative effect on adult D. isaea female longevity up to 5 days after application. Abamectin was lethal for D. isaea larvae when applied directly to larvae or when contaminated leafminer larvae were consumed by parasitoid larvae. However, the percent emergence of D. isaea was not affected by abamectin treatments when applied to chrysanthemum plants that contained parasitoid larvae. Moreover, the longevity of these emerged adults was not affected by abamectin applications. In light of these data, abamectin compatibility with D. isaea for leafminer control in IPM programs for greenhouses is discussed.

Fanghao Wan - One of the best experts on this subject based on the ideXlab platform.

  • interactions between Diglyphus isaea and neochrysocharis formosa hymenoptera eulophidae two parasitoids of agromyzid leafminers
    Biological Control, 2018
    Co-Authors: Jingli Xuan, Wanxue Liu, Jianyang Guo, Yibo Zhang, Xiaoqin Cheng, Fanghao Wan
    Abstract:

    Abstract When two parasitoids exploit a common host simultaneously, competitive interactions between them usually occur, and are more complex among host-killing species. Diglyphus isaea (Walker) and Neochrysocharis formosa (Westwood) are larval parasitoids of agromyzid leafminers. They have widespread distributions and share the same host resources when sympatric. In this study, interactions between these two parasitoids and different parasitoid release modes were investigated. Parasitoids were exposed to leaves infested with late second to early third instar Liriomyza sativae at two different host densities in Petri dishes. The total death rate of N. formosa released alone was density-dependent. And release modes also impacted the total death rate of parasitoids at a high host density. However, the oviposition number of D. isaea was significantly greater than that of N. formosa when they coexisted. In addition, host-feeding proportions were greatest among three host-killing behaviors: parasitism, host feeding and host stinging. These results contribute to a further understanding of parasitoids’ host-killing strategies.

  • comparing immature development and life history traits in two coexisting host feeding parasitoids Diglyphus isaea and neochrysocharis formosa hymenoptera eulophidae
    Journal of Integrative Agriculture, 2014
    Co-Authors: Yibo Zhang, Wanxue Liu, Wei Wang, L U Shulong, Wenxia Wang, Fanghao Wan
    Abstract:

    Abstract Coexisting natural enemies that share a common host resource in the same guild usually exhibit variation in their life history traits, due to their need to share a similar ecological niche. In this study, we compared the immature development times and adult life history traits of two coexisting, host-feeding parasitoids, Diglyphus isaea Walker and Neochrysocharis formosa Westwood (Hymenoptera: Eulophidae), of which both attack larvae of the same agromyzid leafminers. These two species are both synovigenic, idiobiont parasitoids, whose adults consume host fluids (“host feeding”) and lay anhydropic eggs. Of the two, D. isaea has a larger body but little or no initial egg load, and engages in similar lifetime host-feeding events. However, it achieves higher fecundity, longer adult longevity, and higher host suppression ability than N. formosa, which has a smaller body and higher initial egg load. Although D. isaea engages in similar lifetime host-feeding events with N. formosa , all of its gains in life history traits per host-feeding event of D. isaea were larger than those of N. formosa. The age-specific fecundity and host mortality curves of N. formosa were more skewed in early life than those of D. isaea. In addition, the ovigeny index of N. formosa was negatively correlated to body size. Our results confirmed that two coexisting parasitoids, which share the same host resource, show different immature development patterns and life history traits, suggesting that different resource allocation mode could be a general rule of coexisting species sharing the same habitat or host.

  • contrasting patterns of ovarian development and oogenesis in two sympatric host feeding parasitoids Diglyphus isaea and neochrysocharis formosa hymenoptera eulophidae
    Applied Entomology and Zoology, 2014
    Co-Authors: Wanxue Liu, Wei Wang, Lisheng Cheng, Jianyang Guo, Fanghao Wan
    Abstract:

    Diglyphus isaea (Walker) and Neochrysocharis formosa (Westwood) (Hymenoptera: Eulophidae) are common idiobiont parasitoids of leafminers attacking vegetable crops. They exhibit differing levels of synovigeny, and host feeding enhances their fecundity and longevity. The reproductive systems of these two parasitoids are typical of hymenopteran eulophids, consisting of two ovaries, each usually comprising three polytrophic meroistic ovarioles. Diglyphus isaea possesses two obvious oviduct accessory glands, which are absent in N. formosa. Both parasitoids underwent oosorption when starved, while feeding on host larvae promoted oogenesis and egg maturation. In both, oogenesis and vitellogenesis commenced on the first day of the pupal stage rather than after eclosion. Formation of ovarioles in D. isaea commenced 1 day earlier than in N. formosa. Mature eggs were rarely observed in ovaries of newly emerged D. isaea, but usually a few were present in N. formosa. When hosts (second–third instar Liriomyza sativae larvae) were provided, the number of mature eggs in D. isaea ovaries initially increased and then stabilized, while in N. formosa, the number first increased and then decreased. Diglyphus isaea had fewer but larger eggs than N. formosa did. Thus, synovigenic divergence begins at the pupal stage and may result in different life-history traits of adults.

  • lifetime gains and patterns of accumulation and mobilization of nutrients in females of the synovigenic parasitoid Diglyphus isaea walker hymenoptera eulophidae as a function of diet
    Journal of Insect Physiology, 2011
    Co-Authors: Yibo Zhang, Wanxue Liu, Wei Wang, Fanghao Wan
    Abstract:

    In nature, adult parasitoids feed to obtain and use nutrients for supplementing and/or replenishing some of their existing array of nutrient reserves. When adults feed on host or non-host food, they can enhance fitness, typically by increasing egg production or longevity. In the present study, ovigeny index (OI) and impact of female fitness, as well as physiological state on the reproductive strategies, were investigated in the synovigenic parasitoid, Diglyphus isaea, fed on host food (2–3rd instars of Liriomyza sativae larvae), non-host foods (10% honey solution) and starved (distilled water, control). The results showed that D. isaea was a strongly synovigenic parasitoid, of which OI value was 0.002. Both types of food enhanced the fecundity and prolonged the longevity of the females. D. isaea females fed on non-host food showed higher levels of gut sugar, body sugar and glycogen than those fed on host food, but the levels of lipid were higher in the host-fed females. D. isaea females seemed to show lipogenesis, with low rates of lipid catabolism sufficient to satisfy the requirement of egg maturation. Females might absorb lipids directly from the haemolymph of paralyzed hosts.

Valiollah Baniameri - One of the best experts on this subject based on the ideXlab platform.

  • temperature dependent functional response of Diglyphus isaea and hemiptarsenus zilahisebessi parasitizing liriomyza sativae
    Journal of Asia-pacific Entomology, 2020
    Co-Authors: Yaghoub Fathipour, Mostafa Haghani, Ali Asghar Talebi, Abdoolnabi Bagheri, Valiollah Baniameri
    Abstract:

    Abstract This study was conducted to address the effect of different constant temperatures (15, 20, 25, 30 and 35 °C) on the functional response of the parasitoid wasps Diglyphus isaea Walker and Hemiptarsenus zilahisebessi Erdos to different densities of Liriomyza sativae Blanchard (2, 4, 8, 16, 32 and 64 larvae) under laboratory conditions. The results revealed the Type II functional response for both parasitoids at different temperatures. The highest searching efficiency for D. isaea and H. zilahisebessi occurred at 25 °C (0.926 ± 0.211 h−1) and 30 °C (1.012 ± 0.241 h−1), respectively. In addition, the shortest handling time for D. isaea and H. zilahisebessi were observed at 25 °C (0.063 ± 0.008 h) and 30 °C (0.058 ± 0.008 h), respectively. These results demonstrated that H. zilahisebessi is more efficient at higher temperatures than D. isaea. Both parasitoids had higher parasitism performance when temperature increased (4.67 parasitized hosts/day at 15 °C vs 15.87 parasitized hosts/day at 25 °C for D. isaea and 3.89 parasitized hosts/day at 15 °C vs 17.24 parasitized hosts/day at 30 °C for H. zilahisebessi). In addition, a quadratic regression was found between handling time and temperature as well as between number of the parasitized larvae and temperature in D. isaea and H. zilahisebessi at different densities of L. sativae. This study provided a preliminary information on the parasitic behavior of these parasitoids and that D. isaea and H. zilahisebessi can be used properly beside other non-chemical approaches to manage L. sativae damage at a temperature range of 25–30 °C, respectively.

  • Temperature-dependent development of Diglyphus isaea (Hymenoptera: Eulophidae) on Liriomyza sativae (Diptera: Agromyzidae) on cucumber
    Journal of Pest Science, 2007
    Co-Authors: Mostafa Haghani, Yaghoub Fathipour, Ali Asghar Talebi, Valiollah Baniameri
    Abstract:

    The development of Diglyphus isaea (Walker), a parasitoid of leafminers, was studied under laboratory conditions at seven constant temperatures (10, 15, 20, 25, 30, 35, and 40°C) on Liriomyza sativae Blanchard reared on cucumber ( Cucumis sativus L.). The total development period (oviposition to adult emergence) decreased with increasing temperature between 15 and 35°C. In 10 and 40°C no development rate was found in D. isaea , so that it may be claimed that these temperatures fall outside the temperature range for development. Linear regression was used to describe the relationship between development rate and temperature. For egg to adult development, males of D. isaea required 153.8 degree-days (DD) above the theoretical threshold of 9.2°C and females required 161.3 DD above 9.4°C. Data were fitted to four nonlinear temperature-dependent models. Evaluation of the models took place based on the following criteria: fit to data, number and biological value of the fitted coefficient, and accuracy on the estimation of the thresholds. It could be concluded that the Briere-1 and Briere-2 models are suitable for estimating the minimum, maximum and optimal temperature thresholds of D. isaea . Thermal requirements and temperature thresholds can be used to predict the occurrence, number of generations and population dynamics of D. isaea .