Mediterranean fruit fly population phenological patterns are strongly affected by elevation and host presence.

The Mediterranean fruit fly (medfly) (Ceratitis capitata, Diptera: Tephritidae), is an extremely polyphagous pest that threatens the fruit production and trading industry worldwide. Monitoring C. capitata populations and analysing its dynamics and phenology is considered of outmost importance for designing and implementing sound management approaches. The aim of this study was to investigate the factors regulating the population dynamics of the C. capitata in a coastal and semi-mountainous area. We focused on effects of topography (e.g. elevation), host presence and seasonal patterns of ripening on the phenological patterns considering data collected in 2008. The experimental area is characterized by mixed fruit orchards, and Mediterranean climate with mild winters. Two trap types were used for population monitoring. The female targeted McPhail type and the male targeted Jackson type. Traps were placed in farms located at different elevations and landscape morphology (coastal and semi-mountainous areas). The main crops included citrus, apples, peaches, plums, pears, figs, quinces and apricots. Adult captures were first recorded in May, peaked in mid-summer and mid-autumn and almost ceased at the end of the season (January 2008). Captures in the coastal areas preceded that of highlands by 15 days. Most of the adults detected during the fruit ripening of late stone fruit cultivars (first peak) and citrus (second peak). The probability of capturing the first adults preceded almost three weeks the peak of adult captures either considering the elevation or host focus analyses. The results provide valuable information on the seasonal population trend of C. capitata in mixed fruit Mediterranean orchards and can support the set-up of IPM systems in areas with various landscapes and different hosts throughout the fruit growing season.

Latitudinal variation in survival and immature development of Ceratitis capitata populations reared in two key overwintering hosts.

Ceratitis capitata, a major agricultural pest, is currently expanding its geographic distribution to northern, temperate areas of Europe. Its seasonal biology and invasion success depend on temperature, humidity and host availability. In coastal warmer Mediterranean regions and cooler temperature areas, bitter oranges and apples serve as overwintering hosts during the larval stage. We assessed the overwintering capacity of C. capitata populations obtained from different areas of the northern hemisphere by studying the survival and development rates of immature stages in both fruits under laboratory conditions. Eggs from each population were artificially inserted in the flesh of the two hosts and kept at 15, 20, or 25 °C until pupation and adult emergence. Climatic analysis of the area of the population origin showed combined effects of latitude, host and macroclimatic variables on immature survival and development rates. Egg to adult survival rates and developmental duration were longer in apples than in bitter oranges. For populations originated from southern-warmer areas, egg to adult developmental duration was prolonged and adult emergence reduced at 15 °C compared to those populations obtained from northern regions. Our findings reveal varying plastic responses of medfly populations to different overwintering hosts and temperatures highlighting the differential overwintering potential as larvae within fruits. This study contributes towards better understanding the medfly invasion dynamics in temperate areas of Northern Europe and other parts of the globe with similar climates.

Acute cold stress and supercooling capacity of Mediterranean fruit fly populations across the Northern Hemisphere (Middle East and Europe).

The Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae), holds an impressive record of successful invasion events promoted by globalization in fruit trade and human mobility. In addition, C. capitata is gradually expanding its geographic distribution to cooler temperate areas of the Northern Hemisphere. Cold tolerance of C. capitata seems to be a crucial feature that promotes population establishment and hence invasion success. To elucidate the interplay between the invasion process in the northern hemisphere and cold tolerance of geographically isolated populations of C. capitata, we determined (a) the response to acute cold stress survival of adults, and (b) the supercooling capacity (SCP) of immature stages and adults. To assess the phenotypic plasticity in these populations, the effect of acclimation to low temperatures on acute cold stress survival in adults was also examined. The results revealed that survival after acute cold stress was positively related to low temperature acclimation, except for females originating from Thessaloniki (northern Greece). Adults from the warmer environment of South Arava (Israel) were less tolerant after acute cold stress compared with those from Heraklion (Crete, Greece) and Thessaloniki. Plastic responses to cold acclimation were population specific, with the South Arava population being more plastic compared to the two Greek populations. For SCP, the results revealed that there is little to no correlation between SCP and climate variables of the areas where C. capitata populations originated. SCP was much lower than the lowest temperature individuals are likely to experience in their respective habitats. These results set the stage for asking questions regarding the evolutionary adaptive processes that facilitate range expansions of C. capitata into cooler temperate areas of Europe.

Fitness cost of Rhagoletis cerasi (Diptera: Tephritidae) adults emerged from pupae with different dormancy regimes: the case of prolonged chilling.

The European cherry fruit fly, Rhagoletis cerasi (Diptera: Tephritidae), is a univoltine species that overwinters at pupal stage. Under optimum overwintering conditions pupae will develop into adults the next spring. Shorter or longer than optimum chilling periods induce prolonged pupae dormancy. Pupae that enter prolonged dormancy due to a short chilling period exhibit high emergence rates after a second cycle of cold/warm periods. Adults found to be larger and less fecund compared to their counterparts from pupae with annual diapause. On the other hand, extreme long chilling periods at pupal stage results in high mortality rates. However, for one Greek population, a substantial number of adults emerged following prolonged chilling of pupae (ca. 18 consecutive months). In this study, we used three R. cerasi populations in order to address possible geographical variation in fitness cost of adults from pupae with prolonged dormancy. In addition, the fitness traits of these adults emerging after prolonged pupae chilling were compared with that of their counterparts from pupae with annual diapause or prolonged dormancy. Our results reveal no population-specific variation in fitness cost of adults from pupae with prolonged dormancy. Within a population, lifetime fecundity did not differ between adults emerged from pupae with prolonged dormancy and those emerged after prolonged pupae chilling. Adults emerged from pupae exposed to prolonged chilling suffer an additional reduction in adult longevity compared to adults from pupae with prolonged dormancy. Hence, fitness of R. cerasi adults is regulated by diapause regimes of pupae.

Chill coma recovery of Ceratitis capitata adults across the Northern Hemisphere.

The Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae), is an invasive pest, that is currently expanding its geographic distribution from the Mediterranean coasts to more temperate areas of Europe. Given that low temperature is a primary determinant of insect species' range boundaries especially in the Northern Hemisphere with pronounced seasonality, we used chill coma recovery time for assessing latitudinal clines in basal chill tolerance of C. capitata adults. We selected six populations obtained from areas with broad climatic variability based on the main bioclimatic variables of temperature and precipitation, spanning a latitudinal range of about 19° from Middle East to Central Europe. Adults were exposed to 0 °C for 4 h, and time to regain the typical standing position of a fly at 25 °C were recorded. The post-stress survival after a period of 8 days was also recorded. Results revealed that adults from Israel and Austria were less chill tolerant than those from Greece, resulting in curvilinear trends with latitude. Analysis of macroclimatic conditions revealed combined effects of latitude (as a proxy of photoperiod) and macroclimatic conditions on chill coma recovery time. Nonetheless, there was not a deleterious effect on post-recovery survival, except for flies obtained from the northern most point (Vienna, Austria). Overall, it seems that evolutionary patterns of basal chill coma recovery time of C. capitata adults are driven mainly by local climatic variability.

Effects of photoperiod and relative humidity on diapause termination and post-winter development of Rhagoletis cerasi pupae.

The European cherry fruit fly, Rhagoletis cerasi (Diptera: Tephritidae), is a univoltine species that undergoes obligatory summer-winter diapause at pupal stage in the soil (2-5 cm) beneath host trees. To study the effects of photoperiod and relative humidity on diapause termination and post-winter developmental duration of R. cerasi, pupae collected from Dossenheim (Germany) were exposed to different photoperiod or relative humidity regimes during a chilling period ranging from 2 to 8.5 months. Specifically, pupae were exposed to four photoperiod regimes: (a) light conditions (24L:00D), (b) dark conditions (00L:24D), (c) short photoperiod (08L:16D) and (d) long photoperiod (16L:08D), as well as to three relative humidity regimes: (a) low (40% RH), (b) medium (60% RH) and (c) high (70-80% RH). Data revealed that relative humidity is not a significant predictor of diapause termination, but it affects the post-winter developmental period. Higher relative humidity promotes post-winter pupae development. On the other hand, photoperiod significantly affected both diapause termination and post-winter development of R. cerasi pupae. Light conditions (24L:00D) accelerate adult emergence, particularly for females. Regardless of the photoperiod (24L:00D, 00L:24D, 08L:16D), rates of adult emergence were high (>75%) for chilling intervals longer than 6.5 months. Nonetheless, exposure to a long day photoperiod (16L:08D), during chilling, dramatically reduced the proportion of adult emergence following 6 months exposure to chilling. Our findings broaden the understanding of factors regulating diapause responses in European cherry fruit fly, local adaptation and synchronization of adult emergence with the ripening period of major hosts.

Microsatellite and Wolbachia analysis in Rhagoletis cerasi natural populations: population structuring and multiple infections.

Rhagoletis cerasi (Diptera: Tephritidae) is a major pest of sweet and sour cherries in Europe and parts of Asia. Despite its economic significance, there is a lack of studies on the genetic structure of R. cerasi populations. Elucidating the genetic structure of insects of economic importance is crucial for developing phenological-predictive models and environmental friendly control methods. All natural populations of R. cerasi have been found to harbor the endosymbiont Wolbachia pipientis, which widely affects multiple biological traits contributing to the evolution of its hosts, and has been suggested as a tool for the biological control of insect pests and disease vectors. In the current study, the analysis of 18 R. cerasi populations collected in Greece, Germany, and Russia using 13 microsatellite markers revealed structuring of R. cerasi natural populations, even at close geographic range. We also analyzed the Wolbachia infection status of these populations using 16S rRNA-, MLST- and wsp-based approaches. All 244 individuals screened were positive for Wolbachia. Our results suggest the fixation of the wCer1 strain in Greece while wCer2, wCer4, wCer5, and probably other uncharacterized strains were also detected in multiply infected individuals. The role of Wolbachia and its potential extended phenotypes needs a thorough investigation in R. cerasi. Our data suggest an involvement of this symbiont in the observed restriction in the gene flow in addition to a number of different ecological factors.

Prolonged pupal dormancy is associated with significant fitness cost for adults of Rhagoletis cerasi (Diptera: Tephritidae).

In temperate areas, dormancy (diapause and/or quiescence) enables herbivorous insect species to persist and thrive by synchronizing growth and reproduction with the seasonal phenology of their host plants. Within-population variability in dormancy increases survival chances under unpredictable environmental changes. However, prolonged dormancy may be costly, incurring trade-offs in important adult fitness traits such as life span and reproduction. We used the European cherry fruit fly, Rhagoletis cerasi, a stenophagous, univoltine species that overwinters in the pupal stage for usually one or more years to test the hypotheses that prolonged dormancy of pupae has trade-offs with body size, survival and reproduction of the resulting adults. We used two geographically isolated populations of R. cerasi to compare the demographic traits of adults obtained from pupae subjected to one or two cycles of warm-cold periods (annual and prolonged dormancy respectively). Regardless of population, adults from pupae that experienced prolonged dormancy were larger than counterparts emerging within 1year. Prolonged dormancy did not affect adult longevity but both lifetime fecundity and oviposition were significantly decreased. Extension of the life cycle of some individuals in R. cerasi populations in association with prolonged dormancy is likely a bet-hedging strategy.