Parasitism of Aganaspis daci against Ceratitis capitata under Mediterranean climate conditions

The effect of environmental factors is essential to the success of parasitoids as biological control agents, as it determines their foraging activity, development, and survival. The larval‐pupal parasitoid wasp Aganaspis daci (Weld) (Hymenoptera: Figitidae) is known to have a very low fertility (i.e., offspring production) in the field in certain Mediterranean areas, probably due to its inability to efficiently oviposit under such climatic conditions. In this study, the percentage of parasitism and induced mortality (mortality of host pupae attributed to parasitoids, from which adults do not emerge) caused by this wasp to the Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), was assessed under field conditions across 1 year, using medfly‐infested apples and parasitoid‐confined release in a lemon orchard of southeastern Spain. As A. daci is known to have very few emergences in the field, fertility was assessed in the laboratory from parasitized pupae recovered from the field. We found average parasitism rates of 27% and high induced mortality rates of 66% under field conditions. Consequently, medfly population reduction (total mortality of C. capitata caused by A. daci, i.e., induced mortality + % parasitism) was, on average, 87%. Parasitism and induced mortality varied throughout the year, depending on the average temperature and relative humidity. The interaction of these factors resulted in the highest parasitism rates at low mean temperature and humidity values; likewise, the highest percentages of induced mortality were obtained with a combination of high mean temperature and low mean humidity values. In conclusion, A. daci may exert a strong impact on medfly populations, being a good candidate for inundative field releases for the management of C. capitata in the Mediterranean Basin.     

Field parasitism levels of Ceratitis capitata larvae (Diptera: Tephritidae) by Aganaspis daci on different host fruit species in the coastal region of Tartous,Syria

The Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), is a key pest of fruit trees in Syria and is attacked by the larval-pupal parasitoid, Aganaspis daci (Weld) (Hymenoptera: Figitidae). The seasonal population dynamics and distribution of A. daci were studied in surveys of orchards [Citrus reticulatum (clementine), Citrus sinensis (maourdi blood orange), Citrus paradise (grapefruit), C. sinensis (sweet orange), Eriobotrya japonica (loquat), Prunus persica (peach), Prunus domestica (plum), Malus domestica (apple), Figus carica (fig) and Psidium guajava (guava)] at eight locations along the coastal region of Syria, between November 2013 and November 2014. A total of 69.4?kg of infested fruits were sampled, yielding 4274 puparia. From these, 3173 emerged as C. capitata adults, 351 as hymenopteran parasitoids and 750 did not eclose. The parasitoids were identified as A. daci (Hymenoptera: Figitidae) and Psyttalia sp. (Szépligeti) (Hymenoptera: Braconidae). The predominant species was A. daci comprising 348 (99.1%) of the total number of parasitoids collected. Populations of A. daci were present from May 2014 to September 2014, and numbers increased with increasing temperature to peak at 32.05% and 34.01% parasitism in June and August 2014, respectively. Parasitoids were only retrieved from larvae reared from loquat, grapefruit, peach and guava with parasitism levels of 1.68%, 30.76%, 18.28% and 16.15%, respectively. These results are important because they suggest that populations of A. daci have potential to be used for biological control in integrated pest management programmes for C. capitata in the coastal region of Syria.     

Response of Ceratitis capitata to citrus chemicals under semi-natural conditions

Responses of Mediterranean fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) adults to chemicals from citrus fruits were quantified in choice tests conducted in large field cages housing naturally planted orange trees, on the Greek island of Chios. Both sexes responded strongly to chemicals released from artificial cuts made in the pulp of peeled oranges and also to natural or commercial orange juice applied to the surface of yellow 7.0 cm diameter spheres. Males also responded strongly to chemicals released from cuts made in the oily region of the orange peel, or flavedo and less to sour orange and lemon. In contrast, females did not respond to flavedo chemicals. Male response to peel chemicals occurred throughout the day and increased as the number of cuts per fruit increased from 2 to 8. Contact of male mouthparts with sap from cuts, and mating attempts between males on or near the cuts were frequently observed. The males also responded to commercial essential orange oils applied to yellow spheres, but less to essential mandarin oils. The role of citrus chemicals in host finding behaviour of C. capitata and their potential use for the development of effective traps for monitoring and controlling the fly are discussed. The experimental method followed in this study can be used as a standard technique for evaluation of the quality of flies to be used in sterile insect technique programmes.     

Performance and feeding behavior of Ceratitis capitata: comparison of a wild population and a laboratory population

Experiments were carried out to test the performance and some aspects of feeding behavior in two populations of Ceratitis capitata (a population reared in the laboratory for 16 years, i.e., approximately 160 generations, and a wild one obtained from infested coffee, Coffea arabica grains). Two types of food were used in the experiment: an artificial yeast diet used for laboratory rearing and papaya (Carica papaya), a natural host of the fly. The performance parameters tested were percent emergence, time to emergence, adult female size, and egg production during the pre-oviposition phase (first five days of adult life). The behavioral aspects tested were food preference by newly hatched larvae, induction, estimated ingestion of the two diets, whether the larvae placed on one diet stayed there or moved to the other diet, and acceptance of food for oviposition. The results indicated that the performance of the wild population was superior when the flies fed on papaya, whereas the performance of the laboratory population was similar with the two diets; the wild population showed a strong preference for papaya in all choice experiments, whereas the laboratory population showed no diet preference; the females of the wild population only oviposited on pieces of papaya that had not been peeled, and did not oviposit in the artificial diet; the females of the laboratory population oviposited indiscriminately on unpeeled and peeled papaya and on the artificial diet.     

The effect of four fruit species on the parasitization rate of Anastrepha fraterculus (Diptera: Tephritidae,Trypetinae) by Diachasmimorpha longicaudata (Hymenoptera: Braconidae,Opiinae) under laboratory rearing conditions

In the laboratory, the effect of host fruit species Citrus paradisi, C. aurantium, Prunus persica, and Psidium guajava on A. fraterculus parasitism by Diachasmimorpha longicaudata was studied. The number of ovipositor-probing events and the probing-time were documented to evaluate the role of fruit chemistry, and epicarp and mesocarp thickness, respectively. The relationship between the parasitization rate and fruit size in particular plant species was analyzed by applying a simple regression. Results showed that guava and peach yielded significantly more parasitoids than both Citrus spp. Probably, the parasitization rate of D. longicaudata on A. fraterculus would be influenced in part by chemical and physical factors from fruit species.     

Pupal and larval-pupal parasitoids (Hymenoptera) obtained fromAnastrepha Spp. andCeratitis capitata (Dipt.: tephritidae) pupae collected in four localities of Tucuman province,Argentina

Pupal and larval-pupal parasitoids were obtained from 5 % of the 1,413 tephritid puparia collected in four localities of the Tucumán province, Argentina, from April, 1991 to April, 1993.Ceratitis capitata (Wiedemann) was attacked byPachycrepoideus vindemmiae (Rondani) (Pteromalidae), a pupal parasitoid, andAganaspis pelleranoi (Brèthes) (Eucoilidae), a larval parasitoid.Anastrepha spp. were attacked byDoryctobracon areolatus (Szépligeti) (Braconidae), a larval parasitoid, and also byA. pelleranoi. Information about parasitism, percentage of emergence of tephritid species and pupal viability in different localities is provided.     

Biological attributes of three introduced parasitoids as natural enemies of fruit flies,genus Anastrepha (Diptera: Tephritidae)

The biological attributes of three introduced species of parasitoids which attack the fruit fly Anastrepha ludens were evaluated. Larvae and eggs of A. ludens were exposed to larval parasitoids Diachasmimorpha longicaudata and D. tryoni and the egg parasitoid Fopius arisanus. Parasitoid longevity and fecundity were determined using larvae and eggs of A. ludens. Likewise, the parasitism rates of these parasitoid species in infested host fruits were recorded. The intrinsic rate of increase for F. arisanus was 0.1019 followed by D. tryoni with a rate of 0.1641 and D. longicaudata with the highest rate of 0.2233. Although F. arisanus females had the highest levels of fecundity, only 50% of them remained alive until reproductive age. These results in combination with the longer generation time (in comparison with D. longicaudata and D. tryoni), can be considered as the most important factors explaining F. arisanus reduced rate of increase. However, we note that oviposition activity caused egg mortality which reduced Anastrepha egg hatch by ca. 20%. This result suggests that F. arisanus has a high potential as a natural enemy of A. ludens, in accordance with our research efforts to develop a new F. arisanus strain specialized for development in Anastrepha eggs. The results show that D. tryoni is not a good candidate for biological control of Anastrepha. Meanwhile, D. longicaudata continues to be the most important exotic parasitoid for suppression of Anastrepha fruit fly populations.     

Competitive interactions and partial displacement of Anastrepha obliqua by Ceratitis capitata in the occupation of host mangoes (Mangifera indica)

1. We examined the competitive interactions between a native fruit fly species (Anastrepha obliqua Macquart) and the invasive medfly (Ceratitis capitata Wiedemann) when these co-occur on a shared mango fruit host (Mangifera indica L.).
2. Using mango fruits of distinct levels of ripeness, we investigated both competition among larvae and among adult females for oviposition. We quantified competition by the numbers of eggs laid and the intensity of agonistic interactions between adult females.
3. Interactions between immature fruit flies led to reduced size and number of emerged adults of both species. These impacts were felt more acutely in the native species.
4. Interspecific competition between females led to fewer eggs laid on semi-ripe fruit by both species, which may be the result of niche overlap associated with oviposition.
5. Intraspecific interactions between A. obliqua individuals led to intense agonistic behaviour, with a concurrent decrease in number of landings on these host fruits.
6. These results suggest that the native species undergoes a partial niche displacement when facing the invasive species. A portion of the fundamental niche of A. obliqua remained unoccupied by the invading C. capitata, which may allow their coexistence under natural conditions.

     

广东南岭国家级自然保护区实蝇多样性研究

2011年对南岭国家级自然保护区实蝇科昆虫进行了诱集采集,得到实蝇科昆虫标本1640号,隶属于2亚科11属19种。不同海拔高度实蝇科昆虫群落组成不同,在6个采样地中海拔最高点实蝇科昆虫种类和数量发生都较少。不同时间实蝇科群落组成及数量也呈现一定的差异。调查发现,7、8月份诱集的数量最多,分别为525头和537头。不同采样地和不同时间实蝇科昆虫的总体优势度指数（D）、多样性指数（H′）、物种丰富度指数（E）、均匀度指数（J）随时间的变化趋势基本一致。     

Hosts of Mediterranean fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) from Broome and the Broome Peninsula, Western Australia

Abstract  In the Kimberley region of Western Australia, Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) is only established at Broome. From 1998 to 2001 extensive fruit sampling was carried out at Broome during a trial investigating the eradication of medfly from the Kimberley using the sterile insect technique. Fruit was collected from 82 native and introduced plants growing in and surrounding the town of Broome and held for emergence of medfly. Eighteen plants, including the native tree Terminalia petiolaris , were found to be medfly hosts . On the basis of abundance, fruiting phenology and host suitability, eight species (kumquat Fortunella japonica , mango Magnifera indica , Barbados cherry Malpighia glabra , orange jessamine Murraya paniculata , guava Psidium guajava , Pacific almond Terminalia catapa , blackberry tree Terminalia petiolaris and yellow oleander Thevetia peruviana ) were the most important hosts ensuring medfly survival and population growth. Despite medfly being reared from T. petiolaris , it is likely that medfly can only maintain populations in areas close to human habitation, and eradication from these areas would lead to eradication from the whole Kimberley region.     

Effect of food deprivation on attractiveness of food sources, containing natural and artificial sugar and protein, to three African fruit flies: Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis capitata

The effects of food deprivation, age, and mating status on the responses of three fruit fly species, Ceratitis cosyra (Walker), Ceratitits fasciventris (Bezzi), and Ceratitits capitata (Wiedemann) (Diptera: Tephritidae) to natural and artificial sugar and protein food sources were investigated. Natural food sources included guava [Psidium guajava L. (Myrtaceae)] juice (a common host fruit for all three fruit fly species) and bird faeces (farm chicken). Artificial food sources included molasses (obtained from a local sugar factory) and a locally produced protein bait (the International Centre of Insect Physiology and Ecology yeast). In all species studied, sugar deprivation of immature (1–2‐day‐old) male and female flies increased their response to food odours, although it did not change their preference for the type of odour (protein or sugar). Protein deprivation of mature (14–17‐day‐old) male and female flies also increased their response to food odours compared to protein‐fed flies. Protein‐deprived females were highly attracted to odours from protein sources in particular. Odours from natural food sources, guava juice, and chicken faeces, were more attractive to food‐deprived flies than were odours from artificial sugar and protein sources. Attraction to food odours increased significantly with increasing age for protein‐deprived females of all species. For males and females of all species, nutritional state was a more important factor than mating status in influencing responses of flies to food odours. Practical implications of these findings are discussed in terms of strategies for fruit fly control using food baits.     

Seasonal trends in Ceratitis capitata reproductive potential derived from live‐caught females in Greece

Reproductive data of individual insects are extremely hard to collect under natural conditions, thus the study of research questions related to oviposition has not advanced. Patterns of oviposition are often inferred only indirectly, through monitoring of host infestation, whereas the influence of age structure and several other factors on oviposition remains unknown. Using a new approach, in this article, we live‐trapped wild Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) females on the Greek island of Chios during two field seasons. For their remaining lifetime, these females were placed individually in small cages and their daily oviposition was monitored. Reproduction rates between cohorts from different collection dates were then compared. The results showed that in the different captive cohorts the average remaining lifetime and reproduction were highly variable within and between seasons. Multivariate regression analysis showed that the month of capture had a significant effect on captive life span, average daily reproduction, and patterns of egg laying. The effect of year was significant on reproduction, but not on captive life span. These differences between sampling periods probably reflect differences in the availability of hosts and other factors that vary during the season and affect age structure and reproduction. Using a non‐parametric generalized additive model, we found a statistically significant correlation between the captive life span and the average daily reproduction. These findings and the experimental approach have several important implications.     

Effect of ant predation and abiotic factors on the mortality of medfly larvae,Ceratitis capitata,in the Argan forest of Western Morocco

The Argan Argania spinosa (L.) Skeels (Ericales: Sapotaceae) is an endemic and emblematic tree growing in the centre and the southwest of Morocco. Argan fruits are frequently infested by the Mediterranean fruit fly Ceratitis capitata Wied (Diptera: Tephritidae). To date, no control program has been undertaken in the Argan forest. This research is the first investigation of natural mortality factors of C. capitata larvae in the Argan forest. The study was conducted at two localities near the city of Essaouira with different climatic conditions. Observations were conducted using 260 late instar larvae which were individually deposited on the ground, under fruiting host trees, where they naturally fall to find a suitable pupation site. Overall, 42% of the larvae died before being able to burrow into the ground, irrespectively of the location. Larval survival and their likelihood of burrowing decreased with increasing ground temperature, as 53% of all the larvae deaths occurred at temperatures over 48°C. However, at lower ground temperature, predation by ants and spiders accounted for 47% of all mortality. Four species of ants carried out 94% of the captures. Monomorium subopacum Mayr (Hymenoptera: Formicidae), which frequently forages under Argan trees, was by far the most efficient predator, due to a powerful sting that rapidly immobilizes the prey. Our results suggest that the burrowing performance of larvae is constrained by high thermal conditions and the presence of ant predators at lower temperatures.     

Toxicity of baited spinosad formulations to Ceratitis capitata: from the laboratory to the application

GF‐120, a fruit fly bait designed to attract and kill adult fruit flies, was tested in the laboratory and outdoors to determine effects of pre‐treatment diet and bait aging on mortality of Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Two spinosad‐based compounds, GF‐120 and Tracer^® Ultra, had generated two distinctive dose–mortality responds, with LC⁸⁰, LC⁹⁰, and LC⁹⁹ values of 2.4, 2.8, and 4.1 p.p.m., and 255, 479, and 1 143 p.p.m., respectively. The residues of GF‐120 drops, after feeding to the flies, generated 14.3% mortality. The droplet size of the baited spray plays an important role. The toxicity of large drops lasted more than that of small droplets. In the field, exposure to the sun further deteriorates the compound, which lost 50% of its toxicity within 6 days. Disappearance of the compound in the field, due to consumption by various insects, also played a role as 50% of the GF‐120 drops disappeared within 7 days. As mortality was directly related to the amount of insecticide eaten, the effect of GF‐120 depended on the feeding status of the flies: well‐fed flies were almost unaffected compared with starved ones.     

Ecological niches and potential geographical distributions of Mediterranean fruit fly (Ceratitis capitata) and Natal fruit fly (Ceratitis rosa)

Aim To predict and compare potential geographical distributions of the Mediterranean fruit fly (Ceratitis capitata) and Natal fruit fly (Ceratitis rosa). Location Africa, southern Europe, and worldwide. Methods Two correlative ecological niche modelling techniques, genetic algorithm for rule‐set prediction (GARP) and a technique based on principal components analysis (PCA), were used to predict distributions of the two fly species using distribution records and a set of environmental predictor variables. Results The two species appear to have broadly similar potential ranges in Africa and southern Europe, with much of sub‐Saharan Africa and Madagascar predicted as highly suitable. The drier regions of Africa (central and western regions of southern Africa and Sahelian zone) were identified as being less suitable for C. rosa than for C. capitata. Overall, the proportion of the region predicted to be highly suitable is larger for C. capitata than for C. rosa under both techniques, suggesting that C. capitata may be tolerant of a wider range of climatic conditions than C. rosa. Worldwide, tropical and subtropical regions are highlighted as highly suitable for both species. Differences in overlap of predictions from the two models for these species were observed. An evaluation using independent records from the adventive range for C. capitata and comparison with other predictions suggest that GARP models offer more accurate predictions than PCA models. Main conclusions This study suggests that these species have broadly similar potential distributions worldwide (based on climate), although the potential distribution appears to be broader for C. capitata than for C. rosa. Ceratitis capitata has become invasive throughout the world, whereas C. rosa has not, despite both species having broadly similar potential distributions. Further research into the biology of these species and their ability to overcome barriers is necessary to explain this difference, and to better understand invasion risk.     

Performance of controlled-release formulations of trimedlure to attract the Mediterranean fruit fly, Ceratitis capitata

Mediterranean fruit flies (Ceratitis capitata Wied.) are detected by traps baited with the attractant, trimedlure (TML), deposited on the standard cotton dental roll dispenser. Since this dispenser is effective for only 2–4 weeks, controlled-release dispensers may extend the field life of TML-baited traps. A polymeric plug containing 70% by weight TML was highly effective for 8–12 weeks. Other formulations were less effective in the tests with released flies. A release rate of ca. 17 mg of standard TML/day or higher gave insect captures which were generally lower but statistically equivalent to those with the reference dispenser which released 61–63 mg/day. Normalized insect catches increased approximately as the square root of the release rate; the curvilinear regression equation suggested that TML release rates of ca. 52 and 15 mg/day give insect captures equal to ca. 100% and 50%, respectively, of the reference catches. The most attractive TML isomer (TML-C) was a highly effective attractant at a release rate of ca. 4 mg/day.

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Résumé C. capitata était attiré par des pièges alimentés avec de la trimedlure (TML), attractif déposé sur un distributeur commercial de coton dentaire en rouleau. Puisque ce distributeur ne fonctionne que 2 à 4 semaines, des distributeurs à libération contrôlée peuvent augmenter le temps de fonctionnement de pièges fournis en TML. Un bouchon en polymère contenant en poids 70% de TML a été très efficace pendant 8 à 12 semaines. D'autres formules ont été moins efficaces avec des mouches lâchées. Un taux de libération, de 17 mg ou plus de TML moyen par jour, capture moins, mais sans différence statistique significative, qu'un distributeur témoin qui libérait 61 à 63 mg/j. Les captures d'insectes ont augmenté à peu près comme le carré du taux de libération; l'équation de régression curviligne a conduit à penser que la libération de TML à des taux de 52 et 15 mg/j, provoque des captures égales à 100% et 50% des captures de référence. L'isomère le plus attractif (TML-C) a été un attractif très efficace au taux de libération de 4 mg/j.

     

Electroantennogram responses of the Mediterranean fruit fly, Ceratitis capitata, to the volatile constituents of nectarines

Electroantennograms (EAGs) were recorded from laboratory-reared, male and female Ceratitis capitata (medfly) in response to individual volatiles identified from ripe nectarines. Uniformity in EAG response between the sexes was observed for most test compounds. Only 10 volatiles, of the 44 nectarine volatiles tested, elicited significantly different EAG responses between the sexes. No correlation was observed between the magnitude of antennal responsiveness and the concentration of a particular volatile in the headspace odor of ripe nectarines. The most abundant (major) nectarine volatiles were among the least EAG stimulatory compounds tested. And certain minor and trace volatiles were the most potent compounds in eliciting EAGs. Moreover, the magnitude of antennal response to a nectarine volatile was related to the functional-group, chain-length, and unsaturation of the compound. The degree of potency of the compounds was as follows: six-carbon unsaturated aldehydes and alcoholsmethyl and ethyl hexanoates and octanoateshexenyl acetates and monoterpenes>shorter chain-length acetates and alcohols>lactones. Unsaturated aldehydes, alcohols, and acetates generally elicited larger EAGs than their saturated analogs, with the (E)-2-isomers being the most potent isomeric configurations. In addition, medfly antennae exhibited long recovery periods (i.e., >10 sec.) for the EAG tracings to return to baseline potential after stimulations with certain classes of compounds, e.g., C₆ to C₈ acid esters, monoterpenes, and hexen-1-ols. The potential adaptiveness to medflies for selective sensitivity to these volatiles is discussed.     

Function of multiple sperm storage organs in female Mediterranean fruit flies (Ceratitis capitata, Diptera: Tephritidae)

Sperm storage organs allow females to temporally separate insemination from fertilization, manipulate ejaculates and control fertilization. In the reproductive tract of female fruit flies (Diptera: Tephritidae), sperm are found in two different organs--a pair or triplet of spermathecae, and a "fertilization chamber". In order to understand the specific function of each of these organs, we tested the following hypotheses: (1) Sperm are distributed equally amongst the various sperm storage organs; (2) Both organ types maintain sperm viability; and (3) Sperm used in fertilization come from the fertilization chamber. We counted sperm in spermathecae and fertilization chamber of Mediterranean fruit flies (Ceratitis capitata) every 3 days for 18 days following insemination, and used a live/dead staining technique to determine the viability of sperm in these organs. Finally, by extirpating spermathecae from inseminated females and allowing them to oviposit, we were able to identify the fertilization chamber as the source of fertilizing sperm. Numbers of sperm in the spermathecae declined from an average of 3575 on the day of copulation to 649, 18 days later. Conversely, the fertilization chamber maintained a fairly constant level of sperms, ranging between an average of 207 cells on day 3 to 115 sperms on day 18. Throughout the period we monitored, we found high levels of sperm viability in both organs (> 80%). Sperm viability was similarly high in the fertilization chambers of females without spermathecae. However, fertility of eggs laid by these females declined rapidly, as did the number of sperm in the fertilization chamber. We conclude that both the spermathecae and the fertilization chamber are active sperm storage organs, with separate functions: the spermathecae for long-term storage and the fertilization chamber, periodically filled by the spermathecae, a staging point for fertilizing sperm. We suggest that the use of both organs by females results in sperm economy, which adaptively prolongs the intervals between copulations.