Natural parasitism in fruit fly (Diptera: Tephritidae) and interaction with wild hosts surrounding apple orchards adjacent to Atlantic Forest fragments in Paraná State, Brazil / Parasitismo natural de mosca-das-frutas (Diptera: Tephritidae) e interação com hospedeiros silvestres em pomares de macieira adjacentes a fragmentos de Mata Atlântica no Estado do Paraná, Brasil

The South American fruit fly, Anastrepha fraterculus (Wiedemann, 1830) (Diptera: Tephritidae), is an important pest in the subtropical region of Brazil. This insect has tritrophic relation between wild fruits and parasitoids and is associated with apple (Malus domestica Borkh.) orchards adjacent to the Atlantic Forest in Paraná. We thus investigated the degree of infestation of the fruit fly and natural parasitism in wild and cultivated fruits surrounding apple orchards. For this purpose, we collected fruits of Acca sellowiana (Berg.) Burret, Campomanesia xanthocarpa (Mart), Eugenia uniflora L., Eugenia pyriformis Cambessèdes, Psidium cattleianum Sabine, Psidium guajava (L.), Annona neosericea Rainer and Eriobotrya japonica (Thumb) in apple orchards adjacent to the Atlantic Forest located in Campo do Tenente, Lapa and Porto Amazonas counties. In total, we collected 18,289 fruits during four growing years. The occurrence of A. fraterculus depends on the susceptible period of apple fruits. A. sellowiana and P. cattleianum were considered primary fruit fly multipliers and P. guajava was secondary, all occurring after the apple harvest (IS period). The group of parasitoids with A. fraterculus was Aganaspis pelleranoi (Brèthes, 1924) (Hymenoptera: Figitidae), Opius bellus (Gahan, 1930), Doryctobracon areolatus (Szépligeti, 1911) and Doryctobracon brasiliensis (Szépligeti, 1911) (Hymenoptera: Braconidae) all of which are first records in the Atlantic Forest in Paraná. First record of O. bellus occurring in the State of Paraná, as well as, first record of the tritrophic association between host plant A. neosericea, parasitoids D. areolatus and O. bellus and fruit fly A. fraterculus. The host P. cattleianum stood out among the Myrtaceae species in regard to the high diversity of parasitoid species (81% of parasitoids). The total number of Figitidae species (76.5%) was higher [...].

Mosca-das-frutas sul-americana, Anastrepha fraterculus (Wiedemann, 1830) (Diptera: Tephritidae), é uma importante praga da região subtropical do Brasil. Este inseto tem relação tritrófico entre frutos silvestres e parasitoides e está associado a pomares de macieiras (Malus domestica Borkh.) adjacentes à Mata Atlântica no Paraná. Assim, investigamos o grau de infestação da mosca-das-frutas e o parasitismo natural em frutas silvestres e cultivadas ao redor de pomares de maçã. Para tanto, foram coletados frutos de Acca sellowiana (Berg.) Burret, Campomanesia xanthocarpa (Mart), Eugenia uniflora L., Eugenia pyriformis Cambessèdes, Psidium cattleianum Sabine, Psidium guajava (L.), Annona neosericea Rainer e Eriobotrya japonica (Thumb) em pomares de maçã adjacentes à Mata Atlântica localizados nos municípios de Campo do Tenente, Lapa e Porto Amazonas. No total, coletamos 18.289 frutos durante quatro anos de cultivo. A ocorrência de A. fraterculus depende do período de suscetibilidade dos frutos da maçã. A. sellowiana e P. cattleianum foram considerados multiplicadores primários de mosca-das-frutas e P. guajava foi secundário, todos ocorrendo após a colheita da maçã (período IS). Os parasitóides a associados a A. fraterculus foram Aganaspis pelleranoi (Brèthes, 1924) (Hymenoptera: Figitidae), Opius bellus (Gahan, 1930), Doryctobracon areolatus (Szépligeti, 1911) e Doryctobracon brasiliensis (Szépligeti, 1911) (Hymenoptera: Braconidae), todos os quais são primeiros registros na Mata Atlântica no Paraná. Primeiro registro de O. bellus ocorrendo no Estado do Paraná, assim como, primeiro registro da associação tritrófica entre o hospedeiro A. neosericea, parasitoides D. areolatus e O. bellus e mosca-das-frutas A. fraterculus. O hospedeiro P. cattleianum se destacou entre as espécies de Myrtaceae pela alta diversidade de parasitóides associados (81% dos parasitóides). O número total de espécies de Figitidae (76,5%) foi superior [...].

Temperature-dependent phenology of the parasitoid Fopius arisanus on the host Bactrocera dorsalis.

Fopius arisanus (Sonan), an egg-pupal parasitoid of numerous fruit fly species, was recently introduced into Africa for the control of the Oriental fruit fly, Bactrocera dorsalis (Hendel). In this study, life-table data of F. arisanus were generated under laboratory conditions at six constant temperatures (15, 20, 25, 30, 28 and 35 °C; 75% RH, L12:D12 photoperiod) and under variable conditions in a screenhouse on B. dorsalis. Several non-linear functions were fitted to model species development, mortality, longevity and oviposition using the Insect Life Cycle Modeling (ILCYM) software. The established phenology models were stochastically simulated at variable temperatures to estimate the life table parameters. Fopius arisanus completed development from egg to adult at all the temperatures tested except at 35 °C. Mean developmental time from egg to adult was inversely proportional to the temperature. The minimum temperature threshold (Tmin) from egg to adult was 8.15 °C, while the maximum temperature threshold (Tmax) was estimated at 34.2 °C. The optimal temperature for immature stages survival was predicted for 20-30 °C. The optimum fecundity estimated was 251 eggs/female at 22 °C. Following the stochastic simulations under natural conditions of the selected agro-ecological zones, it appears that the Humid Forest with Bimodal Rainfall provides a suitable thermal condition for potential population growth of F. arisanus. The present study shows the importance of temperature on the demographic parameters of F. arisanus. Implications of present findings on the biological control of B. dorsalis under climate change scenarios is discussed.

Determination of temperature thresholds for the parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae), using life cycle simulation modeling: Implications for effective field releases in classical biological control of fruit flies.

The braconid parasitoid Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) is one of the most important natural enemies in classical biological control programs against tephritid fruit flies worldwide. In light of the spread of the invasive fruit fly species, Bactrocera dorsalis in Africa and beyond, there is a need to implement classical biological control. The current study aimed to determine temperature thresholds for D. longicaudata reared on B. dorsalis, using life cycle simulation modeling to guide informed parasitoid releases in Africa. Simulated parameters included thermal requirements, population growth parameters at different temperature requirements, suitable areas for the establishment, and the number of generations per year under projected climatic conditions. The lower thermal threshold for the development was estimated at 10.0°C, with a thermal constant (k) of 333.3-degree days, while the maximum temperature threshold was estimated at 33.69°C. Fecundity was highest at 25°C, with 177.3 eggs per female. Temperature significantly affected the population growth parameters of D. longicaudata, and the maximum value of the intrinsic rate of increase (rm) was 0.145 at 27°C. Results indicate that D. longicaudata could successfully establish in tropical and sub-tropical regions under current and future climatic conditions. However, a slight change in the suitable areas is expected by the year 2050 due to a slight and gradual rise in temperature. Our findings provide important information for further release of this parasitoid in Africa as well as designing pest management strategies to limit the spread and reduce the impact of fruit flies sustainably.

Can Anastrepha fraterculus larval feeding influence chemotaxis and parasitism of Diachasmimorpha longicaudata and Aganaspis pelleranoi?

Anastrepha fraterculus (Diptera: Tephritidae) is a major barrier to fruit production and exportation. In Brazil, the native parasitoid Aganaspis pelleranoi (Hymenoptera: Figitidae) and the exotic parasitoid Diachasmimorpha longicaudata (Hymenoptera, Braconidae) stand out as biological control agents. Knowledge of the factors that affect interactions among parasitoids, A. fraterculus, and host fruits may enhance the use of these agents in biological control programmes. This study evaluated the chemotaxis and parasitism of A. pelleranoi and D. longicaudata females reared on A. fraterculus larvae and kept on an artificial diet, red guava (Psidium guajava) or apple (Malus domestica). Females of both parasitoid species that emerged from larvae raised on artificial diet, guava or apple, were tested to Y olfactometer choice tests. In the parasitism tests, both parasitoid species were made to choose between A. fraterculus larvae brushed with water, apple pulp or guava pulp. D. longicaudata females from artificial diet (control) did not distinguish between fruit odours; however, females of D. longicaudata from larvae kept in apple or guava directed to the odours of their original fruit. The greatest parasitism for D. longicaudata occurred in the units that contained the pulp in which the larvae grew. A. pelleranoi from artificial diet preferred guava odours, including the females kept in apple. Similar results were observed in the parasitism bioassays. Our results found that A. fraterculus larval feeding influenced search behaviour and parasitism of D. longicaudata, whereas A. pelleranoi rearing experience did not affect its host choices.

Exploration for olive fruit fly parasitoids across Africa reveals regional distributions and dominance of closely associated parasitoids.

The olive fruit fly, Bactrocera oleae, has been a key pest of olives in Europe and North America. We conducted the largest exploration for parasitoids associated with the fly across Sub-Saharan Africa (Kenya, Namibia, and South Africa) including some of the fly's adjoining regions (Canary Islands, Morocco, Réunion Island and Tunisia). From Sub-Saharan regions, four braconids were collected: Bracon celer, Psytallia humilis, P. lounsburyi, and Utetes africanus. Results showed that their regional dominance was related to climate niches, with P. humilis dominant in hot semi-arid areas of Namibia, P. lounsburyi dominant in more tropical areas of Kenya, and U. africanus prevalent in Mediterranean climates of South Africa. Psytallia concolor was found in the Canary Islands, Morocco and Tunisian, and the Afrotropical braconid Diachasmimorpha sp. near fullawayi on Réunion Island. Furthermore, we monitored the seasonal dynamics of the fly and parasitoids in Cape Province of South Africa. Results showed that fruit maturity, seasonal variations in climates and interspecific interactions shape the local parasitoid diversity that contribute to the low fly populations. The results are discussed with regard to ecological adaptations of closely associated parasitoids, and how their adaptations impact biocontrol.

Host discrimination in the fruit fly parasitoid Diachasmimorpha longicaudata: evidence from virgin female behaviour and egg distribution patterns.

Many parasitoid species discriminate already parasitized hosts, thus avoiding larval competition. However, females incur in superparasitism under certain circumstances. Superparasitism is commonly observed in the artificial rearing of the parasitoid Diachasmimorpha longicaudata, yet host discrimination has been previously suggested in this species. Here, we addressed host discrimination in virgin D. longicaudata females in a comprehensive way by means of direct and indirect methods, using Ceratitis capitata and Anastrepha fraterculus which are major fruit fly pests in South America. Direct methods relied on the description of the foraging behaviour of females in arenas with parasitized and non-parasitized host larvae. In the indirect methods, healthy larvae were offered to single females and the egg distributions were compared to a random distribution. We found that D. longicaudata was able to recognize parasitized host from both host species, taking 24 h since a first parasitization for A. fraterculus and 48 h for C. capitata. Indirect methods showed females with different behaviours for both host species: complete discrimination, non-random (with superparasitism), and random distributions. A larger percentage of females reared and tested on A. fraterculus incurred in superparasitism, probably associated with higher fecundity. In sum, we found strong evidence of host discrimination in D. longicaudata, detecting behavioural variability associated with the host species, the time since the first parasitization and the fecundity of the females.

Irradiation of early immature Anastrepha ludens stages for the rearing of Doryctobracon areolatus (Hymenoptera: Braconidae), a fruit fly parasitoid.

Doryctobracon areolatus is a native parasitoid of the Neotropical region that presents the highest percentages of natural parasitism of fruit flies of the genus Anastrepha. In the Moscafrut Program SADER-SENASICA, located in Metapa de Domínguez, Chiapas, Mexico, a laboratory colony of this species is maintained on Anastrepha ludens, the Mexican fruit fly, with the aim to scale the production of the parasitoid up to massive levels. In order to eliminate unwanted emergence of adult flies during the rearing process, this study evaluated the effect of irradiation (at doses of 20, 30, 40, and 50 Gy) applied to eggs, and first and second instar larvae of A. ludens; all irradiated stages were subsequently exposed as second instar larvae to adult females of D. areolatus. Irradiation did not affect the eclosion of A. ludens eggs but, at doses of 40 and 50 Gy, it did cause delayed larval development and pupation, as well as lower larval weight. Adult fly emergence was suppressed at all doses, except in eggs irradiated at 20 Gy. Doses of 20 and 30 Gy applied to the eggs and larvae did not affect the emergence, survival, fecundity or flight ability of the emerged parasitoids, but the second instar larvae were easily handled during the rearing process. Our results suggest that D. areolatus can be successfully produced in second instar larvae of A. ludens irradiated at 30 Gy.

Ecological and Evolutionary Stochasticity Shape Natural Selection.

The distribution of biodiversity depends on the combined and interactive effects of ecological and evolutionary processes. The joint contribution of these processes has focused almost exclusively on deterministic effects, even though mechanisms that increase the importance of random ecological processes are expected to also increase the importance of random evolutionary processes. Here we manipulate the sizes of old field fragments to generate correlated sampling effects for a focal population (a gall maker) and its enemy community. Traits and communities were more variable in smaller patches. However, because of the preference of some enemies for some trait values (gall sizes), random variation in population mean trait values exacerbated differences in community composition. The random distribution of traits and interactions created predictable but highly variable patterns of natural selection. Our study highlights how stochastic processes can affect ecological and evolutionary processes structuring the strength and direction of selection locally and at larger scales.

Insights into the Interaction between the Monophagous Tephritid Fly Anastrepha acris and its Highly Toxic Host Hippomane mancinella (Euphorbiaceae).

Despite their enormous economic importance and the fact that there are almost 5000 tephritid (Diptera) species, fruit fly - host plant interactions are poorly understood from a chemical perspective. We analyzed the interactions among Anastrepha acris (a little studied monophagous tephritid) and its highly toxic host plant Hippomane mancinella from chemical, ecological and experimental perspectives, and also searched for toxicants from H. mancinella in the larval-pupal endoparasitoid Doryctobracon areolatus. We identified 18 phenolic compounds from H. mancinella pulp belonging to different chemical groups including phenylpropanoids, flavonoids, chalcones and coumarins. No traces of Hippomanin A were detected in larvae, pupae or A. acris adults, or in D. areolatus adults, implying that A. acris larvae can metabolize this toxicant, that as a result does not reach the third trophic level. We tested the "behavioral preference - lack of larval specialization-hypothesis" via feeding experiments with a larval rearing medium containing H. mancinella fruit (skin + pulp or pulp alone). The high toxicity of H. mancinella was confirmed as only two (out of 2520 in three experiments) A. ludens larvae (a polyphagous pest species that preferentially feeds on plants within the Rutaceae) survived without reaching the adult stage when fed on media containing H. mancinella, whereas A. acris larvae developed well and produced healthy adults. Together, these findings open a window of opportunity to study the detoxification mechanisms used by tephritid fruit flies.

Interaction Between Beauveria bassiana (Hypocreales: Cordycipitaceae) and Coptera haywardi (Hymenoptera: Diapriidae) for the Management of Anastrepha obliqua (Diptera: Tephritidae).

The interaction between the entomopathogenic fungus Beauveria bassiana (Balsamo) and the parasitoid Coptera haywardi (Oglobin), as potential biological control agents for Anastrepha obliqua (Macquart) fruit flies, was evaluated under laboratory and semi-protected field cage conditions. The effects of the parasitoids and fungus were individually and jointly assessed in Plexiglas cages. Application of B. bassiana dry conidia to soil produced 40% mortality in A. obliqua adults. However, mortality was lower (21.2%) on evaluation under field cage conditions. According to the multiple decrement life table analysis, the probability of death of A. obliqua was 88% when C. haywardi parasitoids and B. bassiana conidia were used in conjunction, 89% when only C. haywardi parasitoids were released and 23% when only B. bassiana conidia were applied. These results demonstrate that no synergistic, additive or antagonistic interaction took place with the simultaneous use of these natural enemies, since the presence of B. bassiana had no effect on the C. haywardi parasitism. These results indicate that the parasitoid is a better natural enemy for the control of A. obliqua, and show that, although the two biological control agents can be used simultaneously, their joint application will not produce increased control.

Effect of Density and Lethargy Duration in Prerelease Packaging of the Fruit Fly (Diptera: Tephritidae) Parasitoid, Diachasmimorpha longicaudata (Hymenoptera: Braconidae).

Different densities prerelease packing and times of lethargy in the fruit fly parasitoids Diachasmimorpha longicaudata (Ashmead) were evaluated in order to standardize the process of chilled insect technique for this species. Adults were kept at densities of 0.048, 0.072, 0.096, 0.120, and 0.144 parasitoids/cm2 before release in a México tower, where thermal lethargy was induced at a temperature of 2 ± 2°C for 45 min. Samples of parasitoids were collected to evaluate mortality, survival, fecundity, and flight capacity. All densities showed a similar mortality, both for males (ca. >10%) and females (ca. <7). There was no effect of density on survival and flight capacity in both sexes. On the other hand, fecundity increased with density, 1.66 sons/♀/day, similar to the control. We conclude that a density of 30,000 pupae per cage (0.144 parasitoids/cm2) is adequate for the massive prerelease packaging of the parasitoid D. longicaudata. Regarding the thermal lethargy period, 180 min under 2 ± 2°C conditions, considered as time for management, does not affect the survival, fecundity, and flight capacity of adults. The results obtained are of great utility to establish prerelease packaging parameters for D. longicaudata used in the biological control of Tephritidae fruit fly populations.

Individual and Population Differences Shape Species Interactions and Natural Selection.

Trait variation is central to our understanding of species interactions, and trait variation arising within species is increasingly recognized as an important component of community ecology. Ecologists generally consider intraspecific variation either among or within populations, yet these differences can interact to create patterns of species interactions. These differences can also affect species interactions by altering processes occurring at distinct scales. Specifically, intraspecific variation may shape species interactions simply by shifting a population's position along a trait-function map or by shifting the relationship between traits and their ecological function. I test these ideas by manipulating within- and among-population intraspecific variation in wild populations of a gall-forming insect before quantifying species interactions and phenotypic selection. Within- and among-population differences in gall size interact to affect attack rates by an enemy community, but among-population differences were far more consequential. Intraspecific differences shaped species interactions by both shifting the position of populations along the trait-function map and altering the relationship between traits and their function, with ultimate consequences for patterns of natural selection. I suggest that intraspecific variation can affect communities and natural selection by acting through individual- and population-level mechanisms.

The Effect of Winter Length on Duration of Dormancy and Survival of Rhagoletis completa (Diptera: Tephritidae) and Associated Parasitoids From Northeastern Mexico.

The walnut husk fly Rhagoletis completa (Cresson), native to the Midwestern United States and Mexico, is invasive in California and Europe. It is one of the most important pests of walnuts in areas gathering 30% of the world production. Knowledge of life-history regulation is important for the design of management strategies. Research on dormancy has been performed on invasive populations, and not on populations at the southern extreme of its native range. Here, we examined the effect of winter length on fly and parasitoid emergence, survival, and duration of dormancy. Percent emergence was higher for chill periods at 5°C ranging from 8 to 20 wk. No or insufficient chill resulted in low emergence and a significant proportion of individuals in prolonged dormancy (>1 yr). Duration of dormancy was longer for pupae at constant temperatures and a 4-wk chill period than longer winter durations. Dormancy was longer for Mexican than that reported for U.S. populations, suggesting the existence of a latitudinal cline where populations at southern latitudes have evolved slower metabolic rates. Three parasitoid species were found associated with R. completa (Aganaspis alujai (Wharton and Ovruski) (Hymenoptera: Figitidae), Diachasmimorpha juglandis Muesebeck, and Diachasmimorpha mellea Gahan) (Hymenoptera: Braconidae). Results suggest that rearing of R. completa is possible by subjecting pupae to chill periods between 8 and 20 wk. Overwintering mortality of flies and A. alujai could be further reduced above 5°C. Our findings can contribute for the accurate development of predictive models on invasion potential, development, fly and parasitoid rearing, and biological control.

Effect of fruit and host fly species on the associative learning by Fopius arisanus.

Parasitoids, released in augmentative biological control programmes, which display a rapid host-location capacity, have a higher likelihood of successfully controlling target pest species. By learning to associate sensory cues to a suitable oviposition site, might parasitoids used as biological control agents, locate hosts more rapidly, and perhaps increase the efficacity of e.g. Tephritidae fruit fly management. We studied associative learning of Fopius arisanus (Hymenoptera: Braconidae) and tested its range of learning in natural and conditional hosts and host fruits, i.e. Bactrocera dorsalis, Zeugodacus cucurbitae, Ceratitis capitata and Ceratitis cosyra (Diptera: Tephritidae) and on fruits (papaya, tomato, banana). Naïve female F. arisanus were compared with experienced wasps, which had been offered infested and non-infested fruit, and been allowed to oviposit. Preferences for olfactory cues from infested fruits were thereafter assessed in a two-choice olfactometer. Naïve and trained parasitoids preference differed in general and non-responders to infested fruits were higher among naïve parasitoids. The trained wasps preferred the fruit infested in the training more than the control fruit, for all combination, except when C. cosyra infested the fruits, hence avoidance behavioural response was observed towards the odour of the infested fruit. Fopius arisanus was capable of behaviourally respond to the learned information, e.g. associative odour learning was achieved, yet limited depending on interaction level, fruit fly and fruit combination. To create F. arisanus preference of an associated odour, it might hence be needed to ensure oviposition in perceived suitable host and host fruit, for the parasitoid learning to become favourable in a biological control setup.

Does Beauveria bassiana (Hypocreales: Cordycipitaceae) Affect the Survival and Fecundity of the Parasitoid Coptera haywardi (Hymenoptera: Diapriidae)?

The effect of the entomopathogenic fungus Beauveria bassiana (Balsamo) Vuillemin on the development of immature stages, and survival and fecundity of Coptera haywardi (Oglobin) adults was studied under laboratory conditions. The fungus was applied as dry conidia on parasitized pupae of Anastrepha obliqua (Macquart) (Diptera: Tephritidae) and on parasitoid adults of two different age groups (1- to 4-d-old, and 5- to 10-d-old). The fungus caused no negative effects on the development of the immature stages, since there were no differences on the emergence of adults compared with the untreated control. Adults were susceptible to the fungus on both sexes and age groups. Males showed shorter lifespan than females, even in untreated individuals. Despite the increased adult mortality produced by the fungus there was no effect on fecundity during first 18 d of adult life, as the net fecundity was 26.7 and 26.3 parasitoids per female treated and untreated, respectively. Our results suggest that, given the low susceptibility of parasitized pupae and the no effect on fecundity during the first 18 d of adult life, it is possible to develop management strategies using these two natural enemies in the biological control against A. obliqua.

Host availability affects the interaction between pupal parasitoid Coptera haywardi (Hymenoptera: Diiapridae) and larval-pupal parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae).

The use of multiple species in biological control programmes is controversial when interactions among them are not fully understood. We determined the response of the pupal parasitoid Coptera haywardi (Oglobin) to different availability of Anastrepha ludens (Loew) pupae previously parasitized or not by larval-pupal Diachasmimorpha longicaudata (Ashmead). The two types of pupae were exposed at different ages and proportions to different numbers of C. haywardi females for 48 h. The performance of C. haywardi adults emerging from parasitized and unparasitized pupae was measured. Coptera haywardi prefers to attack unparasitized A. ludens pupae rather than pupae parasitized by D. longicaudata. However, when the availability of unparasitized pupae was low or the number of foraging females was high, C. haywardi competed against early immature stages of the D. longicaudata, or hyperparasitized, feeding directly on the advanced-immature developmental stages of the early acting species. Adults of C. haywardi emerging as hyperparasitoids were no different in size, fecundity and longevity from those emerging as primary parasitoids. Our data suggest that simultaneous use of these species in augmentative biological control projects may be feasible but should be carefully planned in order to avoid any detrimental effect of its interaction.

Best Timing to Determine Field Parasitism by Released Diachasmimorpha longicaudata (Hymenoptera: Braconidae) Against Anastrepha (Diptera: Tephritidae) Pest Populations.

Our objective was to determine the timing of the highest parasitic activity by released Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) in areas with fruits of sour orange and hog plums infested by Anastrepha ludens (Loew) and Anastrepha obliqua (McQuart) (Diptera: Tephritidae), under shaded and sunny conditions. Percent parasitism along fruit sampling period was related to host availability, which was influenced by the fruit size rather than environmental conditions. The highest parasitism in sour orange was obtained just the first day after release, but in hog plums this was observed during the first 3 days without significant differences between them. The levels of fruit infestation and parasitism were higher in shaded trees in sour orange as in hog plums. The high availability of larvae and the small size of hog plums were decisive for obtaining high levels of parasitism and keeping parasitoids near the release points. By contrast, the size and thick rind of sour orange provided to the larvae a physical refuge that was associated with lower parasitism, causing that parasitoids spread out in search of hosts more accessible. In sour orange, parasitism was exclusively by D. longicaudata, while in hog plums, we additionally found the coexistence of four native parasitoid species. This information suggests that in sour orange, the sampling should be performed 1 day after release, while in hog plums, the samplings can be extended to within the first 3 days. Such sampling can serve to better estimate the effect of D. longicaudata releases against Anastrepha pest populations in different fruit types.

From laboratory to the field: consistent effects of experience on host location by the fruit fly parasitoid Diachasmimorpha kraussii (Hymenoptera: Braconidae).

Associative learning is well documented in Hymenopteran parasitoids, where it is thought to be an adaptive mechanism for increasing successful host location in complex environments. Based on this learning capacity, it has been suggested that providing prerelease training to parasitoids reared for inundative release may lead to a subsequent increase in their efficacy as biological control agents. Using the fruit fly parasitoid Diachasmimorpha krausii we tested this hypothesis in a series of associative learning experiments which involved the parasitoid, two host fruits (tomatoes and nectarine), and one host fly (Bactrocera tryoni). In sequential Y-tube olfactometer studies, large field-cage studies, and then open field studies, naïve wasps showed a consistent preference for nectarines over tomatoes. The preference for nectarines was retained, but not significantly increased, for wasps which had prior training exposure to nectarines. However, and again consistently at all three spatial scales, prior experience on tomatoes led to significantly increased attraction to this fruit by tomato-trained wasps, including those liberated freely in the environment. These results, showing consistency of learning at multiple spatial scales, gives confidence to the many laboratory-based learning studies which are extrapolated to the field without testing. The experiment also provides direct experimental support for the proposed practice of enhancing the quality of inundatively released parasitoids through associative learning.

Density-dependent distribution of parasitism risk among underground hosts.

Variation in parasitism risk among hosts can arise from between-patch and within-patch factors, but considerably less information is known about the latter. This study investigated how distributions of the oriental fruit fly Bactrocera dorsalis influenced its parasitism by the pupal parasitoid Dirhinus giffardii in the laboratory. Because B. dorsalis larvae pupate underground, pupation depth was considered as an important factor that affects the risk of parasitism. When the density of B. dorsalis larvae was varied (1, 10, and 100 larvae per arena), average pupation depth increased with the density. When the depth of pupae was manipulated, the rate of parasitism differed by depths. Parasitism at 0 cm differed from the random parasitoid model expectation, but parasitism at 1 cm was not different from the model expectation. Few pupae at 2 cm were parasitized. In another experiment, when pupae were simultaneously presented at 0 cm and 1 cm depths, parasitism at 1 cm was weakened by the presence of puape at 0 cm. These results imply that the density of the host influences pupation depth as well as the distribution of parasitism and plays an important role in host-parasitoid dynamics.

Best Host Age of Anastrepha obliqua (Diptera: Tephritidae) for Multiplication of Four Native Parasitoids from the Americas.

The success of the mass rearing of parasitoids is directly related to host quality, and it requires selecting the best biological host age to ensure the optimal performance of the parasitoids released into the field. The larval development of the parasitoids Utetes anastrephae (Viereck) (Hymenoptera: Braconidae) and Odontosema anastrephae Borgmeier (Hymenoptera: Eucoilidae) and the pupal development of the parasitoids Coptera haywardi (Ogloblin) (Hymenoptera: Diapriidae) and Dirhinus sp. (Hymenoptera: Chalcididae) on the native host Anastrepha obliqua (Macquart) (Diptera: Tephritidae) in different larvae and pupae ages were investigated under laboratory conditions. Not all parasitoid species developed with the same efficiency in immature individuals of A. obliqua; U. anastrephae and C. haywardi showed the higher parasitism rates. The emergence and parasitism of U. anastrephae were equal using larvae from 5 to 8 d, while C. haywardi reared in 1- to 8-d-old pupae showed higher averages of parasitism. These results suggest that native parasitoids can be used to strengthen the implementation of biological control projects against A. obliqua, a pest of economic importance in South America.