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The biology of key citrus pests Thaumatotibia leucotreta Meyrick (Lepidoptera: Tortricidae), Ceratitis capitata Wiedemann (Diptera: Tephritidae) and Ceratitis rosa Karsch (Diptera: Tephritidae) includes their dropping from host plants to pupate in the soil below citrus trees. Since most EP fungi are soil-borne microorganisms, the development and formulation of alternative control strategies using these fungi as subterranean control agents, targeted at larvae and pupae in the soil, can potentially benefit existing IPM management of citrus in South Africa. Thus, a survey of occurrence of entomopathogenic fungi was undertaken on soils from citrus orchards and natural vegetation (refugia) on conventionally and organically managed farms in the Eastern Cape Province in South Africa. A method for baiting soil samples with citrus pest T. leucotreta and C. capitata larvae, as well as with the standard bait insect, Galleria mellonella Linnaeus (Lepidoptera: Pyralidae), was implemented. Sixty-two potentially useful entomopathogenic fungal isolates belonging to four genera were collected from 288 soil samples, an occurrence frequency of 21.53%. The most frequently isolated entomopathogenic fungal species was Beauveria bassiana (Balsamo) Vuillemin (15.63%), followed by Metarhizium anisopliae var. anisopliae (Metschnikoff) Sorokin (3.82%). Galleria mellonella was the most effective insect used to isolate fungal species (χ2=40.13, df=2, P≤ 0.005), with a total of 45 isolates obtained, followed by C. capitata with 11 isolates, and T. leucotreta with six isolates recovered. There was a significantly (χ2=11.65, df=1, P≤ 0.005) higher occurrence of entomopathogenic fungi in soil samples taken from refugia compared to cultivated orchards of both organically and conventionally managed farms. No significant differences were observed in the recovery of fungal isolates when soil samples from both farming systems were compared. The physiological effects and host range of 21 indigenous fungal isolates obtained in the Eastern Cape were investigated in the laboratory to establish whether these isolates could be effectively used as biological control agents against the subterranean life stages of C. rosa, C. capitata and T. leucotreta. When these pests were treated with a fungal concentration of 1 x 10⁷ conidia ml⁻¹, the percentage of T. leucotreta adults which emerged in fungal treated sand ranged from 5 to 60% (F=33.295; df=21; P=0.0001) depending on fungal isolate and the percentage of pupae with visible signs of mycosis ranged from 21 to 93% (F= 96.436; df=21; P=0.0001). Based on fungal isolates, the percentage adult survival in C. rosa and C. capitata ranged from 30 to 90% and 55 to 86% respectively. The percentage of C. rosa and C. capitata puparia with visible signs of mycosis ranged from 1 to 14% and 1 to 11% respectively. Deferred mortality due to mycosis in C. rosa and C. capitata adult flies ranged from 1 to 58% and 1 to 33% respectively, depending on fungal isolate. Entomopathogenic fungal isolates had a significantly greater effect on the adults of C. rosa and C. capitata than they did on the puparia of these two fruit fly species. Further, C. rosa and C. capitata did not differ significantly in their response to entomopathogenic fungi when adult survival or adult and pupal mycosis were considered. The relative potency of the four most virulent Beauveria isolates as well as the commercially available Beauveria bassiana product, Bb Plus® (Biological Control Products, South Africa), were compared against one another as log-probit regressions of mortality against C. rosa, C. capitata and T. leucotreta which all exhibited a dose-dependent response. Against fruit flies the estimated LC50 values of all five Beauveria isolates ranged from 5.5 x 10¹¹ to 2.8 x 10¹² conidia/ml⁻¹. There were no significant differences between the relative potencies of these five fungal isolates. When T. leucotreta was considered, isolates: G Moss R10 and G 14 2 B5 and Bb Plus® were significantly more pathogenic than G B Ar 23 B3 and FCM 10 13 L1. The estimated LC₅₀ values of the three most pathogenic isolates ranged from 6.8 x 10⁵ to 2.1 x 10⁶ conidia/ml⁻¹, while those of the least pathogenic ranged from 1.6 x 10⁷ to 3.7 x 10⁷ conidia/ml⁻¹. Thaumatotibia leucotreta final instar larvae were exposed to two conidial concentrations, at four different exposure times (12, 48, 72 and 96 hrs) and showed an exposure time-dependant relationship (F=5.43; df=3; P=0.001). At 1 x 10⁷conidia/ml⁻¹ two Beauveria isolates: G Moss R10 and G 14 2 B5 were able to elicit a response in 50% of test insects at 72 hrs (3 days) exposure. Although a limited amount of mycosis was observed in the puparia of both fruit fly species, deferred adult mortality due to mycosis was high. The increased incidence of adult mortality suggests that post emergence mycosis in adult fruit flies may play a more significant role in field suppression than the control of fruit flies at the pupal stage. The increased incidence of pupal mortality, as well as the relatively low concentrations of conidia required to elicit meaningful responses in T. leucotreta pupae may suggest that pre-emergent control of false codling moth will play a more significant role in field suppression than the control of adult life stages using indigenous isolates of entomopathogenic fungi. Various entomopathogenic fungal application techniques targeted at key insect pests within integrated pest management (IPM) systems of citrus are discussed.

This study investigated understorey management in Canterbury, New Zealand, apple orchards for the enhancement of populations of Dolichogenidea tasmanica (Cameron) (Braconidae) for leafroller (Lepidoptera: Tortricidae) biological control. The first objective was to determine the influence of understorey plants on the abundance of D. tasmanica and leafroller parasitism, and to investigate the mechanisms behind this influence. The second was to determine the most suitable understorey plants in terms of their ability to enhance parasitoid abundance, leafroller parasitism, parasitoid longevity, parasitoid fecundity and its ability to not benefit leafroller. Results from three consecutive field trials showed that buckwheat (Fagopyrum esculentum Moench), coriander (Coriandrum sativum L.), alyssum (Lobularia maritima (L.) Desv), and, to a lesser extent, broad bean (Vicia faba L.), enhanced parasitoid abundance and leafroller parasitism. The mechanisms behind the effects of understorey plants had previously been unexplored. However, results here showed that it was the flowers or the buckwheat that 'attracted' the parasitoid to the plant and not the shelter, aphids or microclimate that the plant may also provide. Providing flowering plants in the orchard understorey also increased immigration of parasitoids and enhanced parasitoids and enhanced parasitoid longevity and fecundity in the laboratory. In contrast, the understorey plants had no influence on the female:male ratio of D. tasmanica. Although coriander enhanced leafroller parasitism three-fold in field experiments compared with controls, it failed to enhance the longevity of both sexes of D. tasmanica in the laboratory compared with water-only. Broad bean significantly enhanced parasitoid abundance three-fold and significantly increased parasitism from 0% to 75% compared with the controls on one leafroller release date. However, laboratory trials showed that of male D. tasmancia but it did not enhance female longevity. Also, female D. tasmanica foraging on broad bean produced a total of only three parasitoid cocoons, but this result was based on an overall 6.5% survival of larvae to pupae or to parasitoid cocoon. Furthermore, results suggested that extrafloral nectar secretion decreased as the plants matured. Phacelia (Phacelia tanacetifolia Benth.) did not significantly enhance parasitism rate in the field compared with controls, and numbers of D. tasmanica captured by suction sampling were significantly lower in phacelia treatments compared with alyssum, buckwheat and control plots. Also, laboratory experiments showed that survival of D. tasmanica on phacelia flowers was equivalent to that on water-only and significantly lower than on buckwheat. These results suggest that phacelia does not provide nectar to D. tasmanica, only pollen, and therefore is not a suitable understorey plant for D. tasmanica enhancement in orchards. Buckwheat and alyssum showed the most potential as understorey plants for the enhancement of natural enemies. Buckwheat not only increased numbers of D. tasmanica seven-fold, but also increased numbers of beneficial lacewings (Micromus tasmaniae (Walker)) and hover flies (Syrphidae) captured on yellow sticky traps compared with the controls. It significantly increased leafroller parasitism by D. tasmanica from 0% to 86% compared with the controls (on one date only), and in the laboratory enhanced D. tasmanica longevity and increased fecundity compared with water-only. Similarly, alyssum significantly increased parasitism rate compared with controls, and two-fold more D. tasmanica were suction sampled in these plots compared with controls. It also enhanced longevity of both sexes of D. tasmanica compared with water, and showed the most favourable characteristics in terms of being of no benefit to leafrollers. This is because it was not preferred over apple by leafroller larvae and when they were forced to feed on it, it caused high mortality (94.3%) and low pupal weight (15 mg). Furthermore, alyssum did not enhance the number of fertile eggs produced by adult leafrollers compared with water only. However, further research is required to address the overall effect of buckwheat and alyssum on crop production and orchard management, including effects on fruit yield and quality, frost risk, disease incidence, soil quality, weeds and other pests. Also, research into the ability of these plants to survive in the orchard with little maintenance, and into the optimal sowing rates, would be useful. Sampling natural populations of leafroller within each treatment showed that damage from leafrollers and the number of leafroller larvae were respectively 20.3% and 29.3% lower in the flowering treatments compared with the controls. Furthermore, field trials showed up to a six-fold increase in leafroller pupae in controls compared with buckwheat and alyssum. This suggests that increasing leafroller parasitism rate from understorey management in orchards will translate into lower pest populations, although neither larval numbers/damage nor pupal numbers differed significantly between treatments. Trapping D. tasmanica at a gradient of distances showed that this parasitoid travels into rows adjacent to buckwheat plots, indicating that growers may be able to sow flowering plants in every second or third row of the orchard, and still enhance leafroller biocontrol while minimising the adverse effects of a cover crop. Sowing buckwheat and alyssum in orchard understoreys may enhance biological control of apple pests in organic apple production and reduce the number of insect growth regulators applied in IFP programmes. However, the challenge still remains to investigate whether conservation biological control can reduce leafroller populations below economic thresholds.

This study investigated understorey management in Canterbury, New Zealand, apple orchards for the enhancement of populations of Dolichogenidea tasmanica (Cameron) (Braconidae) for leafroller (Lepidoptera: Tortricidae) biological control. The first objective was to determine the influence of understorey plants on the abundance of D. tasmanica and leafroller parasitism, and to investigate the mechanisms behind this influence. The second was to determine the most suitable understorey plants in terms of their ability to enhance parasitoid abundance, leafroller parasitism, parasitoid longevity, parasitoid fecundity and its ability to not benefit leafroller. Results from three consecutive field trials showed that buckwheat (Fagopyrum esculentum Moench), coriander (Coriandrum sativum L.), alyssum (Lobularia maritima (L.) Desv), and, to a lesser extent, broad bean (Vicia faba L.), enhanced parasitoid abundance and leafroller parasitism. The mechanisms behind the effects of understorey plants had previously been unexplored. However, results here showed that it was the flowers or the buckwheat that 'attracted' the parasitoid to the plant and not the shelter, aphids or microclimate that the plant may also provide. Providing flowering plants in the orchard understorey also increased immigration of parasitoids and enhanced parasitoids and enhanced parasitoid longevity and fecundity in the laboratory. In contrast, the understorey plants had no influence on the female:male ratio of D. tasmanica. Although coriander enhanced leafroller parasitism three-fold in field experiments compared with controls, it failed to enhance the longevity of both sexes of D. tasmanica in the laboratory compared with water-only. Broad bean significantly enhanced parasitoid abundance three-fold and significantly increased parasitism from 0% to 75% compared with the controls on one leafroller release date. However, laboratory trials showed that of male D. tasmancia but it did not enhance female longevity. Also, female D. tasmanica foraging on broad bean produced a total of only three parasitoid cocoons, but this result was based on an overall 6.5% survival of larvae to pupae or to parasitoid cocoon. Furthermore, results suggested that extrafloral nectar secretion decreased as the plants matured. Phacelia (Phacelia tanacetifolia Benth.) did not significantly enhance parasitism rate in the field compared with controls, and numbers of D. tasmanica captured by suction sampling were significantly lower in phacelia treatments compared with alyssum, buckwheat and control plots. Also, laboratory experiments showed that survival of D. tasmanica on phacelia flowers was equivalent to that on water-only and significantly lower than on buckwheat. These results suggest that phacelia does not provide nectar to D. tasmanica, only pollen, and therefore is not a suitable understorey plant for D. tasmanica enhancement in orchards. Buckwheat and alyssum showed the most potential as understorey plants for the enhancement of natural enemies. Buckwheat not only increased numbers of D. tasmanica seven-fold, but also increased numbers of beneficial lacewings (Micromus tasmaniae (Walker)) and hover flies (Syrphidae) captured on yellow sticky traps compared with the controls. It significantly increased leafroller parasitism by D. tasmanica from 0% to 86% compared with the controls (on one date only), and in the laboratory enhanced D. tasmanica longevity and increased fecundity compared with water-only. Similarly, alyssum significantly increased parasitism rate compared with controls, and two-fold more D. tasmanica were suction sampled in these plots compared with controls. It also enhanced longevity of both sexes of D. tasmanica compared with water, and showed the most favourable characteristics in terms of being of no benefit to leafrollers. This is because it was not preferred over apple by leafroller larvae and when they were forced to feed on it, it caused high mortality (94.3%) and low pupal weight (15 mg). Furthermore, alyssum did not enhance the number of fertile eggs produced by adult leafrollers compared with water only. However, further research is required to address the overall effect of buckwheat and alyssum on crop production and orchard management, including effects on fruit yield and quality, frost risk, disease incidence, soil quality, weeds and other pests. Also, research into the ability of these plants to survive in the orchard with little maintenance, and into the optimal sowing rates, would be useful. Sampling natural populations of leafroller within each treatment showed that damage from leafrollers and the number of leafroller larvae were respectively 20.3% and 29.3% lower in the flowering treatments compared with the controls. Furthermore, field trials showed up to a six-fold increase in leafroller pupae in controls compared with buckwheat and alyssum. This suggests that increasing leafroller parasitism rate from understorey management in orchards will translate into lower pest populations, although neither larval numbers/damage nor pupal numbers differed significantly between treatments. Trapping D. tasmanica at a gradient of distances showed that this parasitoid travels into rows adjacent to buckwheat plots, indicating that growers may be able to sow flowering plants in every second or third row of the orchard, and still enhance leafroller biocontrol while minimising the adverse effects of a cover crop. Sowing buckwheat and alyssum in orchard understoreys may enhance biological control of apple pests in organic apple production and reduce the number of insect growth regulators applied in IFP programmes. However, the challenge still remains to investigate whether conservation biological control can reduce leafroller populations below economic thresholds.

In this study, buckwheat (Fagopyrum esculentum Moench) and alyssum (Lobularia maritima (L.)) flowers were used to examine the effect of floral resources on the efficacy of the leafroller parasitoid Dolichogenidea tasmanica (Cameron) in vineyards. This was done by assessing the influence of these flowers on parasitoid abundance and parasitism rate, and by investigating the consequences of this for leafroller abundance. In laboratory experiments, alyssum flowers were used to investigate the effect of floral food on the longevity, fecundity and sex ratio of D. tasmanica. Dolichogenidea tasmanica comprised more than 95 % of parasitoids reared from field collected leafrollers in this study. The abundance of D. tasmanica during the 1999-2000 growing season was very low compared with previous studies, possibly due to the very low abundance of its leafroller hosts during the experiment. The number of males of this species on yellow sticky traps was increased (although not significantly) when buckwheat flowers were planted in a Marlborough vineyard; however, the number of female D. tasmanica on traps was no greater with flowers than without. The abundance of another leafroller parasitoid, Glyptapanteles demeter (Wilkinson)(Hymenoptera: Braconidae), on traps was also not significantly affected by the presence of buckwheat flowers, although females of this species were caught in greater numbers in the control than in buckwheat plots. Naturally-occurring leafrollers were collected from three vineyard sites in Marlborough, and one in Canterbury during the 2000-2001 season to assess the effect of buckwheat and alyssum flowers on parasitism rate. Parasitism rate more than doubled in the presence of buckwheat at one of the Marlborough vineyards, but alyssum had no effect on parasitism rate in Canterbury. A leafroller release/recover method, used when naturally-occurring leafrollers were too scarce to collect, was unable to detect any effect of buckwheat or alyssum on parasitism rate. Mean parasitism rates of approximately 20 % were common in Marlborough, although rates ranged from 0 % to 45 % across the three vineyard sites in that region. In Canterbury in April, mean parasitism rates were approximately 40 % (Chapter 4). Rates were higher on upper canopy leaves (40-60 %) compared with lower canopy leaves and bunches (0-25 %). Leafroller abundance was apparently not affected by the presence of buckwheat in Marlborough, or alyssum in Canterbury. Buckwheat did, however, significantly reduce the amount of leafroller evidence (webbed leafroller feeding sites on leaves or in bunches) in Marlborough, suggesting that the presence of these flowers may reduce leafroller populations. Leafrollers infested less than 0.1 % of Cabernet Sauvignon leaves throughout the 1999-2000 growing season, but increased in abundance in bunches to infest a maximum of 0.5 % of bunches in late March in Marlborough. In Pinot Noir vines in the 2000-2001 season, leafroller abundance was also low, although sampling was not conducted late in the season when abundance reaches a peak. In Riesling vines in Canterbury, between 1.5 % and 2.5 % of bunches were infested with leafrollers in April. In the laboratory, alyssum flowers significantly increased the longevity and lifetime fecundity of D. tasmanica compared with a no-flower treatment. However, daily fecundity was not increased by the availability of food, suggesting that the greater lifetime fecundity was related to increases in longevity. Parasitoids were also able to obtain nutrients from whitefly honeydew, which resulted in similar longevity and daily fecundity to those when alyssum flowers were present. The availability of food had a significant effect on the offspring sex ratio of D. tasmanica. Parasitoids reared from naturally-occurring leafrollers produced an equal sex ratio, assumed to be the evolutionarily stable strategy (ESS) for this species. In the laboratory, this ESS was observed only when parasitoids had access to alyssum flowers. Without food, or with honeydew only, sex ratios were strongly male-biased. In the field, floral resources affected the sex ratio of D. tasmanica only when this species was reared from leafrollers released and recovered in Marlborough. In that experiment, buckwheat shifted the sex ratio in favour of female production from the equal sex ratio found in control plots. No firm explanations can be given to account for these results, due to a lack of research in this area. Possible mechanisms for the changes in sex ratio with flowers are discussed. This study demonstrated that flowers are an important source of nutrients for D. tasmanica, influencing the longevity, fecundity and offspring sex ratio of this species. However, only some of the field experiments were able to show any positive effect of the provision of floral resources on parasitoid abundance or parasitism rate. More information is needed on the role these parasitoids, and other natural enemies, play in regulating leafroller populations in New Zealand vineyards, and on how they use floral resources in the field, before recommendations can be made regarding the adoption of this technology by growers.