Age, sex, adult and larval diet shape starvation resistance in the Mediterranean fruit fly: an ecological and gerontological perspective.

The ability of an animal to withstand periods of food deprivation is a key driver of invasion success (biodiversity), adaptation to new conditions, and a crucial determinant of senescence in populations. Starvation resistance (SR) is a highly plastic trait and varies in relation to environmental and genetic variables. However, beyond Drosophila, SR has been studied poorly. Exploiting an interesting model species in invasion and ageing studies-the Mediterranean fruit fly (Ceratitis capitata)- we investigated how age, food and gender, shape SR in this species. We measured SR in adults feeding in rich and poor dietary conditions, which had been reared either on natural hosts or artificial larval diet, for every single day across their lifespan. We defined which factor is the most significant determinant of SR and we explored potential links between SR and ageing. We found that SR declines with age, and that age-specific patterns are shaped in relation to adult and larval diet. Females exhibited higher SR than males. Age and adult diet were the most significant determinants of SR, followed by gender and the larval diet. Starvation resistance proved to be a weak predictor of functional ageing. Possible underlying mechanisms, ecological and gerontological significance and potential applied benefits are discussed

Effect of Sex, Age and Temperature on the Functional Response of <i>Macrolophus pygmaeus</i> Ramber and <i>Nesidiocoris tenuis</i> Reuter (Heteroptera: Miridae) on Eggs of <i>Tuta absoluta</i>

The predatory mirids Macrolophus pygmaeus Ramber (Heteroptera: Miridae) and Nesidiocoris tenuis Reuter (Heteroptera: Miridae) are used for the biological control of Tuta absoluta Meyrick (Lepidoptera: Gelechiidae) and other pests in tomato greenhouses. The functional response of 1-day-old (young) and 10-day-old (old) adult females and males of M. pygmaeus and N. tenuis on eggs of T. absoluta was determined on tomato at two temperatures (20 °C and 25 °C) and LD 16:8. Females of M. pygmaeus exhibited higher predation efficiency than males at both tested temperatures. Young M. pygmaeus females had a higher efficiency than old ones, whereas males had a low efficiency irrespective of age. The predation efficiency of N. tenuis was high (but lower than M. pygmaeus) in both young females and males, although old females had a higher efficiency than the respective males. Our results show that the two predatory species have different functional response characteristics to their prey depending mainly on sex and age, which may affect their role as biological control agents

Artificial diet alters activity and rest patterns in the olive fruit fly

Olive fruit flies, Bactrocera oleae (Diptera: Tephritidae) reared in the laboratory on an artificial diet are essential for the genetic control techniques against this pest. However, the colony's laboratory adaptation can affect the quality of the reared flies. We used the Locomotor Activity Monitor to track the activity and rest patterns of adult olive fruit flies reared as immatures in olives (F2-F3 generation) and in artificial diet (&gt;300 generations). Counts of beam breaks caused by the adult fly activity were used as an estimation of its locomotor activity levels during the light and dark period. Bouts of inactivity with duration longer than five minutes were considered a rest episode. Locomotor activity and rest parameters were found to be dependent on sex, mating status and rearing history. In virgin flies reared on olives, males were more active than females and increased their locomotor activity towards the end of the light period. Mating decreased the locomotor activity levels of males, but not of female olive-reared flies. Laboratory flies reared on artificial diet had lower locomotor activity levels during the light period and more rest episodes of shorter duration during the dark period compared to flies reared on olives. We describe the diurnal locomotor activity patterns of B. oleae adults reared on olive fruit and on artificial diet. We discuss how locomotor activity and rest pattern differences may affect the laboratory flies' ability to compete with wild males in the field

Artificial diet alters activity and rest patterns in the olive fruit fly.

Olive fruit flies, Bactrocera oleae (Diptera: Tephritidae) reared in the laboratory on an artificial diet are essential for the genetic control techniques against this pest. However, the colony's laboratory adaptation can affect the quality of the reared flies. We used the Locomotor Activity Monitor to track the activity and rest patterns of adult olive fruit flies reared as immatures in olives (F2-F3 generation) and in artificial diet (>300 generations). Counts of beam breaks caused by the adult fly activity were used as an estimation of its locomotor activity levels during the light and dark period. Bouts of inactivity with duration longer than five minutes were considered a rest episode. Locomotor activity and rest parameters were found to be dependent on sex, mating status and rearing history. In virgin flies reared on olives, males were more active than females and increased their locomotor activity towards the end of the light period. Mating decreased the locomotor activity levels of males, but not of female olive-reared flies. Laboratory flies reared on artificial diet had lower locomotor activity levels during the light period and more rest episodes of shorter duration during the dark period compared to flies reared on olives. We describe the diurnal locomotor activity patterns of B. oleae adults reared on olive fruit and on artificial diet. We discuss how locomotor activity and rest pattern differences may affect the laboratory flies' ability to compete with wild males in the field

Supine behaviour predicts the time to death in male Mediterranean fruitflies (Ceratitis capitata).

Over 97% of the 203 male medflies monitored in a lifetime study of their behaviour exhibited what we term supine behaviour (temporary upside-down orientation) starting an average of 16.1 days prior to their death (mean lifespan of 61.7 days). Supine onset increased the mortality risk by 39.5-fold and a unit increase in supine level increased mortality by 26.3%. The discovery that behavioural traits in insects can be used as biomarkers of their health and to predict their time to death has important implications regarding research on morbidity dynamics, behavioural neuroethology and gerontology, and the interpretation of longevity extension in model organisms

Females of <i>Halyomorpha halys</i> (Hemiptera: Pentatomidae) Experience a Facultative Reproductive Diapause in Northern Greece

Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) is a native pest of East Asia that overwinters as an adult in natural and human-made structures. Adult emergence from overwintering sites starts in spring, whereas females produce offspring in early summer on host plants, where most feeding occurs. In this study, we investigated the reproductive physiology of overwintering females of H. halys in Northern Greece, by determining the duration of the preoviposition period and fecundity of individuals that were left to overwinter in natural conditions and were subsequently transferred to chambers with standard conditions monthly, from December 2020 to March 2021. According to our results, overwintering H. halys females do not initiate egg laying once they emerge from overwintering sites, but rather need some additional time to exit diapause and mature reproductively. The mean preoviposition period of overwintering females that were transferred from their overwintering sites to the chambers in December 2020 was 29.0 days, which was significantly longer by 8.3 days than that of females that overwintered until March 2021, and by 13.2 days than the control (26 °C, 60% RH and a 16:8 h light: dark photoperiod). No significant difference among the average number of eggs per egg mass laid by overwintering individuals brought in the chambers in different time intervals and the laboratory colony was observed. However, females that were left to overwinter until March laid a significantly higher number of eggs in total, compared to the ones whose overwintering was disrupted in February. Based on our findings, overwintering females of H. halys experience a facultative reproductive diapause in Northern Greece. Our study was the first to determine the occurrence of diapause of H. halys in N. Greece and our findings could be very valuable for assessing the damage of this pest to early-season crops and designing successful management practices

Remating in wild females of the Mediterranean fruit fly, Ceratitis capitata

Female remating is widespread among animal taxa, and shows high interspecific diversity in frequency. While there is no consensus on general explanations for why females mate multiply, it is clear that remating rate has important implications for many evolutionary processes such as sperm competition, cryptic female choice and sexual conflict. We investigated the refractory period and the effect of sex ratio on remating in wild-derived individuals of the Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) (Diptera, Tephritidae), an insect pest species of major worldwide economic importance. In addition, we determined the remating rates of wild females throughout the flying season. Experiments with medflies that were the offspring of wild flies showed that remating was frequent and was observed in 3.3–13.3% of females on each of the 4 days after the first mating. Many matings thus induce only a short refractory period. Again using the offspring of wild flies, we found that female remating increased significantly under a highly male-biased sex ratio. Microsatellite analysis of offspring from wild-collected females showed that remating is common (4–28% of offspring arrays showed multiple paternity) and occurred throughout the flying season of these insects. These results have important implications for insect pest management using the sterile insect technique

The Roles of Mating, Age, and Diet in Starvation Resistance in <i>Bactrocera oleae</i> (Olive Fruit Fly)

The olive fruit fly (Bactrocera oleae (Rossi) (Diptera: Tephritidae)), although a pest of major economic importance for the olive industry, has not been sufficiently studied with respect to the factors affecting its survival resistance to food deprivation. In the present study, we examined the effect of the interaction between mating status (virgin/mated), age class (11–20/21–30/31–40/41–50), and diet quality (protein plus sugar or only sugar) on starvation resistance in B. oleae under constant laboratory conditions. We conducted a total of 16 treatments (2 × 4 × 2 = 16) for each gender. Our results showed that starvation resistance in B. oleae did not differ significantly between females and males. The main conclusions of our study regarding mating status, age, and diet indicated that mated adults showed much less starvation resistance compared to virgins, younger adults endured longer, and the adults fed a restricted diet endured longer than those fed a full diet. A three-way interaction between mating status, diet, and age class was also identified and was the same for both genders. The interaction between mating status, age class, and diet also had a significant influence on starvation resistance in both sexes

Age Related Assessment of Sugar and Protein Intake of Ceratitis capitata in ad libitum Conditions and Modeling Its Relation to Reproduction

In the inquiry on the age related dietary assessment of an organism, knowledge of the distributional patterns of food intake throughout the entire life span is very important, however, age related nutritional studies often lack robust feeding quantification methods due to their limitations in obtaining short-term food-intake measurements. In this study, we developed and standardized a capillary method allowing precise life-time measurements of food consumption by individual adult medflies, Ceratitis capitata (Diptera: Tephritidae), under laboratory conditions. Protein or sugar solutions were offered via capillaries to individual adults for a 5 h interval daily and their consumption was measured, while individuals had lifetime ad libitum access to sugar or protein, respectively, in solid form. Daily egg production was also measured. The multivariate data-set (i.e., the age-dependent variations in the amount of sugar and protein ingestion and their relation to egg production) was analyzed using event history charts and 3D interpolation models. Maximum sugar intake was recorded early in adult life; afterwards, ingestion progressively dropped. On the other hand, maximum levels of protein intake were observed at mid-ages; consumption during early and late adult ages was kept at constant levels. During the first 30 days of age, type of diet and sex significantly contributed to the observed difference in diet intake while number of laid eggs varied independently. Male and female adult longevity was differentially affected by diet: protein ingestion extended the lifespan, especially, of males. Smooth surface models revealed a significant relationship between the age dependent dietary intake and reproduction. Both sugar and protein related egg-production have a bell-shaped relationship, and the association between protein and egg-production is better described by a 3D Lorenzian function. Additionally, the proposed 3D interpolation models produced good estimates of egg production and diet intake as affected by age, providing us with a reliable multivariate analytical tool to model nutritional trends in insects, and other organisms, and their effect upon life history traits. The modeling also strengthened the knowledge that egg production is closely related to protein consumption, as suggested by the shape of the medfly reproduction-response function and its functional relationship to diet intake and age

Trapping of <i>Ceratitis capitata</i> Using the Low-Cost and Non-Toxic Attractant Biodelear

Trapping is considered a powerful tool in the monitoring and control of fruit flies of high economic importance such as the Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae). However, the cost of trapping and, in some cases, the safety of the chemicals used as baits are concerning for growers and the environment. Here we present a novel, low cost, environmentally friendly, female-specific bait for C. capitata, called Biodelear, that consists of a mixture of attracting compounds such us pyrazines, pyranones and amorphous nitrogen-based polymers. The new bait was compared to the commercially available attractant Biolure® (Suterra LLC, Bend, OR, USA) in Greece. McPhail-type traps were deployed in an orange orchard located in Athens. Five traps per treatment were used for several weeks during 2009 and four traps per treatment in 2010. Traps contained either 17 g of Biodelear or one Biolure Unipack dispenser. The results showed that both baits were highly efficient in attracting C. capitata females, and to a lesser extent, males. Although Biolure initially appeared to outperform Biodelear, later in the season the two attractants converged in efficacy. In both years, female captures were similar in traps baited with Biolure and Biodelear. However, male captures were higher in Biodelear-baited traps in 2010. In addition, Biodelear seemed to be longer lasting than Biolure, despite not being formulated into a slow-release dispersion system. The low cost of Biodelear and its strong, long-lasting effects render it suitable for mass trapping of the Mediterranean fruit fly