Survival ability of Mexican fruit fly males from different strains in presence of the predatory orb-weaving spider Argiope argentata (Araneae: Araneidae).

The sterile insect technique (SIT) is a key element for the integrated management of pest populations of the Mexican fruit fly, Anastrepha ludens, in Mexico. Its success depends on the survival of mass-reared sterile males and their ability to mate with wild females. However, colonization and mass-rearing conditions can adversely affect their ability to avoid predators. To test if colony management strategies could contribute to improve field survival abilities of mass-reared flies, we compared the survival of males exposed to the orb-weaver spider Argiope argentata. Males compared originated from three strains with different colonization strategies: (a) a colony started from field-collected wild flies (replacement), (b) a colony started by hybridizing wild males with mass-reared adapted females (hybrid) and (c) a colony started with mass-reared males selected on the basis of their survival ability and mating competitiveness in field cages (selected). Mass-reared males and wild males were used as controls. Males were exposed to spiders under laboratory cage conditions. Overall, wild males showed better survival ability than mass-reared males. Regarding the colonization approach, wild males survived better than a hybrid, replaced and selected males. We conclude that mass-rearing conditions have a strong negative effect on the ability of males to escape spiders. The colonization systems evaluated did not counter this effect. The lower survival of males from the selected colony suggests that the selection over one generation did not contribute to improve males' predator avoidance and escape abilities and probably needs to be modified. Possible explanations for this and implications on colonization and colony management for SIT purpose are discussed.

A comparison of sexual competitiveness and demographic traits of Anastrepha obliqua (Macquart) (Diptera: Tephritidae) among fruit-associated populations.

The control of Anastrepha obliqua includes the sterilization of mass-reared insects grown in isolation in a constantly controlled environment. Through time, laboratory mass-reared colonies may produce flies with lower field performance. To recover the genetic variation and aptitude of mass-reared populations, wild insects are introduced into mass-reared colonies. Our aim in this study was to determine whether the host species from two localities influence the life history traits of A. obliqua. We collected flies as larvae from infested fruits of Spondias purpurea, S. mombin, Mangifera indica cv. 'piña', and M. indica cv. 'coche' from two localities in Chiapas, Mexico. There were significant differences in the mating competitiveness of males collected from mango cv. 'coche' compared with mass-reared males. There were no differences in the mating propensity between flies from the two localities, even in the number of matings, when weight was considered as a covariable. The mass-reared strain showed the earliest age at first oviposition. The locality affected the longevity and oviposition period, and these influenced the birth rate, intrinsic rate of increase, finite rate of population increase, mean generation time, and doubling time. According to the demographic parameters, the population of S. mombin would allow artificial colonization in less time, considering that it has a high reproduction rate starting at an early age. Even in the propensity test, it had the highest number of matings. However, males with greater sexual competitiveness and longevity for colonization corresponded to those collected from S. purpurea.

Efficiency of two larval diets for mass-rearing of the mosquito Aedes aegypti.

Aedes aegypti is a major vector of arboviruses that may be controlled on an area-wide basis using the sterile insect technique (SIT). Larval diet is a major factor in mass-rearing for SIT programs. We compared dietary effects on immature development and adult fitness-related characteristics for an International Atomic Energy Agency (IAEA) diet, developed for rearing Ae. albopictus, and a standardized laboratory rodent diet (LRD), under a 14:10 h (light:dark) photoperiod ("light" treatment) or continuous darkness during larval rearing. Larval development was generally fastest in the IAEA diet, likely reflecting the high protein and lipid content of this diet. The proportion of larvae that survived to pupation or to adult emergence did not differ significantly between diets or light treatments. Insects from the LRD-dark treatment produced the highest proportion of male pupae (93% at 24 h after the beginning of pupation) whereas adult sex ratio from the IAEA diet tended to be more male-biased than that of the LRD diet. Adult longevity did not differ significantly with larval diet or light conditions, irrespective of sex. In other aspects the LRD diet generally performed best. Adult males from the LRD diet were significantly larger than those from the IAEA diet, irrespective of light treatment. Females from the LRD diet had ~25% higher fecundity and ~8% higher egg fertility compared to those from the IAEA diet. Adult flight ability did not differ between larval diets, and males had a similar number of copulations with wild females, irrespective of larval diet. The LRD diet had lower protein and fat content but a higher carbohydrate and energetic content than the IAEA diet. We conclude that the LRD diet is a low-cost standardized diet that is likely to be suitable for mass-rearing of Ae. aegypti for area-wide SIT-based vector control.

Population Fluctuation of Ceratitis capitata (Diptera: Tephritidae) as a Function of Altitude in Eastern Guatemala.

Population fluctuations of Ceratitis capitata (Wiedemann) were evaluated over a period of 12 mo in four altitudinal strata (400-750, 750-1,100, 1,100-1,450, and 1,450-1,800 meters above sea level, masl) in Eastern Guatemala. Within each altitudinal range, sampling plots were established in coffee plantations and adjacent areas, in which Jackson traps were set and baited with Trimedlure. Coffee berries and other host fruits were collected. Population density was lowest at the 400-750 masl stratum and highest at 1,450-1,800 masl. At every altitudinal range, the fluctuations of the pest were associated mainly with the availability of ripe coffee berries as a primary host. From 750-1,450 masl, the pest was also associated with the availability of sweet orange and mandarins in commercial and backyard orchards. The highest densities of the pest were recorded in the dry season. Citrus were the main alternate host where ripe coffee berries were not available. This knowledge on population dynamics of C. capitata will contribute to develop more effective area-wide pest management strategies including the use of sterile insects, natural enemies, and bait sprays.

Selection by mating competitiveness improves the performance of Anastrepha ludens males of the genetic sexing strain Tapachula-7.

The sexual performance of Anastrepha ludens males of the Tapachula-7 genetic sexing strain, produced via selection based on mating success, was compared with that of males produced without selection in competition with wild males. Mating competition, development time, survival, mass-rearing quality parameters and pheromone production were compared. The results showed that selection based on mating competitiveness significantly improved the sexual performance of offspring. Development time, survival of larvae, pupae and adults, and weights of larvae and pupae increased with each selection cycle. Differences in the relative quantity of the pheromone compounds (Z)-3-nonenol and anastrephin were observed when comparing the parental males with the F4 and wild males. The implications of this colony management method on the sterile insect technique are discussed.

Foraging Allocation in the Honey Bee, Apis mellifera L. (Hymenoptera, Apidae), Tuned by the Presence of the Spinosad-Based Pesticide GF-120.

Agroecosystem management commonly involves the use of pesticides. As a result, a heterogeneous landscape is created, in which suitable and unsuitable spaces are defined by the absence/presence of pesticides. In this study, we explored how foragers of the honey bee, Apis mellifera L., adapt to such context. We specifically evaluated the effect of GF-120, a spinosad-based fruit fly toxic bait, on the allocation of foragers between food sources under the hypothesis that foragers will move from food sources with GF-120 to food sources without it. We thus carried out three experiments: in experiment 1, a group of foragers was trained to collect honey solution from a feeder; next, this feeder offered a GF-120/honey solution. A minority of foragers continued collecting the GF-120/honey solution. In experiment 2, we trained two groups of foragers from a colony to two food sources equally rewarding. Next, GF-120 was added to one of the food sources. We found that the majority of foragers moved from the GF-120-treated feeder to the feeder without GF-120 and that the minority that continued visiting the GF-120-treated feeder did not collect the GF-120/honey solution. In a third experiment, we wanted to know if foragers in an experimental setup as in experiment 1 would perform waggle dances: none of the foragers that collected GF-120/honey were observed dancing. Our results emphasize the importance of "food refuges" for non-target species, since they minimize the impact of agrochemicals upon them.

Effects of size, sex and teneral resources on the resistance to hydric stress in the tephritid fruit fly Anastrepha ludens.

Water availability is recognized as one of the most important factors in the distribution and activity of terrestrial organisms. In the case of insects, hydric stress imposes a major challenge for survival because of the small surface-area-to-volume ratio they exhibit. In general, stress resistance is expected to co-vary positively with size; however, this pattern can become obscured in insects that exhibit sexual size dimorphism, as sexes differ in size and/or shape and have dissimilar resource allocations. In the present study, we use an allometric-based approach to (i) assess the desiccation and starvation stress resistance of teneral Anastrepha ludens flies, (ii) disentangle the relationships between resistance, size and sex and (iii) examine the adult fly body differences in water and lipid contents before and after exposure to stress. After controlling for sexual size dimorphism, an allometric increase of resistance with overall size was observed for all stress-based treatments. The scaling exponents that define the proportion of increase resistance varied with size traits and with type and degree of hydric stress. In this allometric relationship, and also in the relationships between mass and wing length and between size and teneral resources, the sexes maintained similar scaling exponents but differed in the intercepts. Males were more resistant to stress than females; this competitive advantage is probably linked to greater amounts of teneral lipids and more water use during stress.

Anastrepha egg deposition induces volatiles in fruits that attract the parasitoid Fopius arisanus.

Fopius arisanus is a solitary egg-pupal endoparasitoid that attacks several species of tephritid fruit flies, particularly Bactrocera spp. This species, indigenous from the Indo-Australian region, was introduced into Mexico for biological control purposes. From the standpoint of the 'new associations' concept this parasitoid has been evaluated against fruit flies in the Anastrepha complex. We investigated the specificity of F. arisanus responses to fruits infested with two species of Anastrepha. We examined whether fruit volatiles attractive to this parasitoid are induced as a result of fruit fly oviposition. We also investigated whether F. arisanus females are able to discriminate between the oviposition-induced volatiles from host eggs parasitised by conspecifics and volatiles from unparasitised eggs. All experiments were performed in a wind tunnel. Results showed that mango fruits infested with A. ludens eggs (2-3 days after egg deposition) were significantly more attractive to naïve F. arisanus females compared with non-infested fruits or fruits infested with larvae. In addition, guava fruits harbouring A. striata eggs were significantly more attractive to the parasitoid than non-infested fruits or fruits infested with larvae. Thus, the parasitoid was attracted to fruits with eggs, but fruit and fly species did not influence the parasitoid attraction. We also found that F. arisanus females were more attracted to fruits exposed to fertile A. ludens females (i.e. fruits with eggs inside) compared with fruits exposed to sterile females (i.e. fruits with no eggs inside) or fruits with mechanical damage. Parasitoid females were not attracted to A. ludens eggs. The results suggest that the presence of eggs induces volatiles that attract parasitoids. Finally, we found that F. arisanus was able to discriminate between fruits with unparasitised eggs vs. eggs parasitised by conspecifics, indicating that host discrimination could be mediated by olfactory cues.

Effect of the natural pesticide spinosad (GF-120 formulation) on the foraging behavior of Plebeia moureana (Hymenoptera: Apidae).

In this study we evaluated the effects of the biorational pesticide, Spinosad (GF-120 formulation), on foraging behavior in the stingless bee Plebeia moureana (Ayala). Several foragers were individually trained to collect an unscented 1.0 M sucrose solution (31% sucrose wt:wt) from a blue plate in one arm of a Y-tube maze. The other arm offered plain water on a yellow plate. After 20-30 visits to the setup, the sucrose solution was exchanged for a sucrose solution mixed with one of five concentrations of GF-120 and 30 consecutive choices of each bee were recorded. Interestingly, the foragers collected the sucrose solution with GF-120 at all concentrations. Our results show that: 1) the GF-120 formulation, when applied at the recommended concentration and mixed with food, does not discourage engaged foragers and, 2) foraging behavior over time is not significantly impaired by the continuous collection of GF-120.

Beauveria bassiana as a pathogen of the Mexican fruit fly (Diptera: Tephritidae) under laboratory conditions.

Bioassays were carried out under controlled conditions (27 +/- 2 degrees C, 80 +/- 5% RH, and a photoperiod of 12:12 [L:D] h) to evaluate the effect of eight strains of the entomopathogenic fungus Beauveria bassiana upon larvae, pupae, and adult females of the Mexican fruit fly, Anastrepha ludens (Loew). Mortality of the immature stages was low, 2-8% in larvae and 0% in pupae. However, very high levels of mortality were obtained for adult flies, with values of 100, 98, and 98% for the strains Bb16, Bb24, and Bb26, respectively. LC50 values for these three strains ranged from 3.12 x 10(6) to 9.07 x 10(6) conidia/ml. Lethal time 50 (LT50) was 2.8, 3.7, and 4.2 d for Bb16, Bb26, and Bb24 strains, respectively, with an average LT50 of 4.4 d across all strains. The fungal mycelium emerged through the soft parts of the exoskeleton, such as the wing bases, mouth, intersegmental regions of the legs, and membranous regions of the abdomen, coxae, and neck. Maximum percentage sporulation ranged from 66.4 to 74.7% for the three most virulent strains.

Reproductive potential predicts longevity of female Mediterranean fruitflies.

Reproduction exacts a price in terms of decreased survival. Our analysis of the interplay between age patterns of fecundity and mortality for individual female medflies (Ceratitis capitata) revealed that individual mortality is associated with the time-dynamics of the egg-laying trajectory. In a sample of 531 medflies, we found that each individual has a characteristic rate of decline in egg laying with age. This defines an individual's rate of reproductive exhaustion. This rate was shown to predict subsequent mortality The larger the remaining reproductive potential, the lower the subsequent mortality An increased mortality risk was seen in flies for which egg production declined rapidly early on, irrespective of the level of egg production. Thus, reproductive potential and lifetime are coupled in such a way that those flies which are able to profit most from an extended life span in terms of increased egg output are indeed likely to live longer.

Female sensitivity to diet and irradiation treatments underlies sex-mortality differentials in the Mediterranean fruit fly.

Large-scale experiments on medflies that were subjected to sterilizing doses of ionizing radiation (plus intact controls) and maintained on either sugar-only or full, protein-enriched diets revealed that, whereas the mortality trajectories of both intact and irradiated male cohorts maintained on both diets are similar, the mortality patterns of females are highly variable. Mean mortality rates at 35 days in male cohorts ranged from 0.2 to 0.3 but in female cohorts ranged from 0.09 to 0.35, depending on treatment. The study reports three main influences: (a) qualitative differences exist in the sex-mortality response of medflies subjected to dietary manipulations and irradiation; (b) the female mortality response is linked to increased vulnerability due to the nutritional demands of reproduction; and (c) female sensitivity to environmental changes underlies the dynamics of the sex-mortality differential.

Anticipation of oxidative damage decelerates aging in virgin female medflies: hypothesis tested by statistical modeling.

Empirical analysis of survival data obtained from large samples of Mediterranean fruit flies shows that the trajectory of the mortality rate for virgin females departs from that for females maintained in mixed sex cages. It increases, decelerates, reaches its maximum, declines and then increases again within the reproductive interval. Non-virgin females, however, display an early-age plateau instead of this dip. We assume that these deviations are produced by the interplay between changes in oxygen consumption associated with reproductive behavior and the antioxidant defense that acts against anticipated oxidative damage caused by reproduction. Since there are no data on antioxidant mechanisms in medflies available that explain the observed patterns of mortality, we develop a model of physiological aging based on oxidative stress theory, which describes age-related changes in oxygen consumption and in antioxidative capacity during the reproductive period. Using this model, we simulate virtual populations of 25,000 virgin and non-virgin flies, calculate the respective mortality rates and show that they practically coincide with those of experimental populations. We show that the hypothesis about the biological support of reproduction used in our model does not contradict experimental data. The model explains how the early-age dip and plateau might arise in the mortality rates of female medflies and why the male mortality pattern does not exhibit such deviations.

Toxicity of Bacillus thuringiensis beta-exotoxin to three species of fruit flies (Diptera: Tephritidae).

The current study describes toxic effects of the Bacillus thuringiensis beta-exotoxin toward 3rd instars of 3 fruit fly species: Anastrepha ludens (Loew), A. obliqua (Macquart), and A. serpentina (Wiedemann). The beta-exotoxin was highly toxic to all 3 species tested, with LC50 values calculated as 0.641, 0.512, and 0.408 microgram/cm2 of filter paper used to expose the larvae, for A. ludens, A. obliqua, and A. serpentina, respectively. Exposure to beta-exotoxin was associated with an increase in the incidence of deformed pupae. The adult survivors from beta-exotoxin treatments showed no negative effects in terms of their longevity, fecundity, or egg eclosion (fertility). We conclude that the beta-exotoxin may have potential as a control agent for fruit fly pests.

Mortality oscillations induced by periodic starvation alter sex-mortality differentials in Mediterranean fruit flies.

Sex-specific mortality rates of medflies were monitored in cages containing individuals of both sexes and with food (either sugar-only or full diet) removed every 2nd, 3rd, or 4th day (plus ad libitum controls). The general finding is that periodic starvation led to marked oscillations in raw mortality rates. The specific findings are as follows: (i) female medflies live longer than male medflies when they are subjected to periodic starvation; (ii) male medflies maintained on a full diet experience a catastrophic increase in mortality (40%) on the first day food is removed. This mortality surge was not observed for females on either diet or for males maintained on a sugar-only diet; (ii) life expectancy is inversely related to the amplitude of mortality oscillations caused by food deprivation; and (iv) the large perturbations in mortality at younger ages caused by periodic starvation has little effect on the amplitude of mortality at older ages. In general, our data shed new light on the complexity of the mortality response of medflies to both the type and availability of food and thus provide a complimentary perspective to findings from dietary restriction studies on both vertebrate and invertebrate systems.

Dual modes of aging in Mediterranean fruit fly females.

The life history of medflies is characterized by two physiological modes with different demographic schedules of fertility and survival: a waiting mode in which both mortality and reproduction are low and a reproductive mode in which mortality is very low at the onset of egg laying but accelerates as eggs are laid. Medflies stay in waiting mode when they are fed only sugar. When fed protein, a scarce resource in the wild, medflies switch to reproductive mode. Medflies that switch from waiting to reproductive mode survive longer than medflies kept in either mode exclusively. An understanding of the physiological shift that occurs between the waiting and reproductive modes may yield information about the fundamental processes that determine longevity.

Biodemographic trajectories of longevity.

Old-age survival has increased substantially since 1950. Death rates decelerate with age for insects, worms, and yeast, as well as humans. This evidence of extended postreproductive survival is puzzling. Three biodemographic insights--concerning the correlation of death rates across age, individual differences in survival chances, and induced alterations in age patterns of fertility and mortality--offer clues and suggest research on the failure of complicated systems, on new demographic equations for evolutionary theory, and on fertility-longevity interactions. Nongenetic changes account for increases in human life-spans to date. Explication of these causes and the genetic license for extended survival, as well as discovery of genes and other survival attributes affecting longevity, will lead to even longer lives.

Relationship of age patterns of fecundity to mortality, longevity, and lifetime reproduction in a large cohort of Mediterranean fruit fly females.

Daily reproduction was monitored throughout the lives of 1000 individual female Mediterranean fruit flies (Ceratitis capitata). Inasmuch as the average female medfly lived 35.6 days and laid 740 eggs in her lifetime, the overall data set consisted of information on around 740,000 eggs distributed over 35,600 fly-days. Results described include the frequency distributions of eggs/day at young (< or =30 days), middle (31-60 days), and older (61-90 days) ages, the relationship between individual life span and lifetime reproduction, the distribution of deaths for non-egg layers versus egg layers, and density plots for daily egg production relative to both lifetime reproduction and life span. One of the more surprising results was the lack of correlation of the total number of eggs laid by females at younger ages (all ages under 30) and subsequent reproduction and life span. Technical and conceptual implications for analyses of reproductive data on other species and groups are briefly discussed.

Early mortality surge in protein-deprived females causes reversal of sex differential of life expectancy in Mediterranean fruit flies.

Experiments based on over 400,000 medflies revealed that females maintained on a normal diet (sucrose plus protein) have a higher life expectancy than males maintained on a normal diet, with a difference of 1.30 +/- 0.27 days in favor of females. However, this sex differential reverses under protein deprivation, with a difference of 2.24 +/- 0.18 days in favor of males. The reversal of the male-female life expectancy differential is caused by a sustained surge in early female mortality under protein deprivation that is tied to egg-laying and physiological processes. In contrast, male mortality and life expectancy are only mildly affected by protein deprivation. The surge in early mortality for female medfly cohorts is an instance of a vulnerable period. These vulnerable periods are linked with patterns in hazard rates.

Mortality dynamics of density in the Mediterranean fruit fly.

The effects on medfly age-specific mortality of three types of densities--initial, current, and cumulative--were examined using sex-specific data from two sets of studies: (1) previous research on mortality patterns in 1.2 million individuals maintained in 167 different cages (1992 Science 258, 457) and ii) density experiments using a total of 210,000 individuals contained in 49 cages and maintained at one of three initial densities--2500, 5000 and 10,000 flies/cage. A central death rate was computed for each of the 216 cages at specified numerical levels (e.g., 5000, 4000, 1000, 500, 100, and so forth), which was distributed over a range of ages. This yielded a series of mortality schedules at "equivalent current densities." Two main results are reported. First, the leveling off and decline in mortality at the most advanced ages as observed in the original study of 1.2 million medflies cannot be explained as an artifact of declining current densities at older ages. Second, increased initial density heightened the mortality level at each age but had essentially no effect on mortality pattern. The overall methodology and many of the results are believed to be general and thus both logistical and conceptual implications for gerontology and population biology are discussed.