Estimating changes in mean population age using the death distributions of live‐captured medflies

1. A simple, low‐cost approach to estimating population ageing was introduced based on a modified version of the captive cohort method – a technique developed earlier (Carey et al., Aging Cell, 7 , 426–437, 2008) in which information on the remaining lifespans of live‐captured medflies of unknown age is used to estimate the overall population age structure. 2. To test this approach approximately 1200 medflies near Volos, Greece were live captured from daily sampling over a 3‐month field season. 3. This simplified method reported: (i) an extraordinary post‐capture longevity of wild medflies in early season (>200 days in longest lived); (ii) a decrease of 50–75 days in the mean longevity from early‐season to late‐season flies; (iii) seasonality of frailty as indicated by the shorter‐lived flies in late autumn; (iv) cessation of fly emergence in late season as indicated by the absence of long‐lived individuals (indicating newly emerged at capture) sampled in the autumn; and (v) increase in mean age from about 20 days in early season to approximately 60 days in late season. 4. The applications of this simplified captive cohort method are discussed, including its use in the analysis of insect vector populations, Drosophila ecology and ageing in the wild, demographic toxicology, and age bias in sampling.     

Relevance of Male Size to Female Mate Choice in Ceratitis capitata (Diptera: Tephritidae): Investigations with Wild and Laboratory-Reared Flies

We examined the influence of male size on the mating success of both wild and laboratory populations of the Mediterranean fruit fly (medfly) Ceratitis capitata. Laboratory females selected larger males even when smaller males were present in large proportions (5:1), although this phenomenon did not occur when the proportion was extended to 10:1. Wild females were observed to prefer wild males over laboratory-reared ones independent of their respective sizes, and wild males of different sizes were found to transfer similar quantities of sperm to the spermathecae of the females.     

Are degraded habitats from agricultural crops associated with elevated faecal glucocorticoids in a wild population of common vole (Microtus arvalis)?

The severe impact of agriculture on species’ abundance and diversity is widely recognized. However, its effects on the physiology of wild animal populations are poorly known. We analyzed faecal glucocorticoids levels in wild common voles (Microtus arvalis) living in a farmland landscape to test whether living in degraded habitats, such as crops, is correlated with increased glucocorticoids. Other factors such as sex, reproductive status, and population density were also considered. We captured voles with Sherman traps in crops and in their field margins which were comprised of semi-natural vegetation. We collected fresh faecal samples from captured individuals and quantified their levels of faecal corticosterone metabolites (FCM) in the laboratory. The quantification of FCM concentrations was performed by competitive enzyme immunoassay. Individuals captured within the crops had higher levels of FCM than those in field margins; females and breeding individuals exhibited higher FCM levels. In addition, FCM concentrations positively correlated with abundance of voles. Our results suggest that degraded habitats in agricultural landscapes are associated with increased glucocorticoid levels on common voles likely caused by a higher disturbance from agricultural practices and a lesser vegetation cover in crops compared with field margins.     

Genetic elimination of field-cage populations of Mediterranean fruit flies

The Mediterranean fruit fly (medfly, Ceratitis capitata Wiedemann) is a pest of over 300 fruits, vegetables and nuts. The sterile insect technique (SIT) is a control measure used to reduce the reproductive potential of populations through the mass release of sterilized male insects that mate with wild females. However, SIT flies can display poor field performance, due to the effects of mass-rearing and of the irradiation process used for sterilization. The development of female-lethal RIDL (release of insects carrying a dominant lethal) strains for medfly can overcome many of the problems of SIT associated with irradiation. Here, we present life-history characterizations for two medfly RIDL strains, OX3864A and OX3647Q. Our results show (i) full functionality of RIDL, (ii) equivalency of RIDL and wild-type strains for life-history characteristics, and (iii) a high level of sexual competitiveness against both wild-type and wild-derived males. We also present the first proof-of-principle experiment on the use of RIDL to eliminate medfly populations. Weekly releases of OX3864A males into stable populations of wild-type medfly caused a successive decline in numbers, leading to eradication. The results show that genetic control can provide an effective alternative to SIT for the control of pest insects.     

Bacterially enriched diet improves sexual performance of sterile male Mediterranean fruit flies

The Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), (medfly) is a polyphagous and cosmopolitan agricultural pest, targeted in many areas for control by the Sterile Insect Technique (SIT). Our objective in the present study was to test the hypothesis that a bacterially enriched diet provided to sterile males will improve their sexual performance in competitive settings that emulate natural conditions. Specifically we determined how feeding on diets enriched with Klebsiella oxytoca affected the ability of sterile males to compete for wild females against wild males, their ability to inhibit female receptivity, and their survival. We found that enriching the sterile male diet with K.oxytoca significantly improved mating competitiveness in the laboratory and in field cages. In addition, bacterially enriched sterile males inhibited female receptivity more efficiently than sugar fed males and survived longer duration of starvation. We conclude that inoculating mass reared sterile flies with bacteria prior to their release is a valid approach to improve the efficacy of SIT.     

Life table assay of field-caught Mediterranean fruit flies, Ceratitis capitata, reveals age bias

Though traps are used widely to sample phytophagous insects for research or management purposes, and recently in aging research, possible bias stemming from differential response of individuals of various ages to traps has never been examined. In this paper, we tested the response of Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) males and females of four ages (spanning from 1 to 40 days) to McPhail-type traps baited with a synthetic food attractant in field cages and found that the probability of trapping was significantly influenced by age. The type of food on which flies were maintained before testing (sugar or protein) also had a strong effect and interacted with age. In another experiment, we collected wild C. capitata adults of unknown age using 1–3 methods and then reared them in the laboratory until death. The survival schedules of these flies were subsequently used in a life table assay to infer their age at the time of capture. Results showed that on a single sampling date, males captured in traps baited with a food attractant were younger compared with males aspirated from fruiting host trees, or males captured in traps baited with a sex attractant. Likewise, females captured in food-baited traps were younger compared with aspirated females. In addition to providing the first evidence of age-dependent sampling bias for a phytophagous insect species, this paper also provides a novel approach to estimate the differences in the age composition of samples collected with different techniques. These findings are of utmost importance for several categories of insects, medically important groups notwithstanding.     

Effect of male and female age on the mating success of the obliquebanded leafrollerChoristoneura rosaceana (Lepidoptera: Tortricidae) under different ecological conditions

The present study was undertaken to test the hypothesis that male and female age, as well as temperature, can affect the mating success ofChoristoneura rosaceana, given the role that these two factors play in the pheromone biology of this species. In the laboratory, the mating success of females generally declined linearly with age, whereas in males, it increased during the first 3 days and then decreased. The decline in female mating success was more pronounced under warm than cool thermocycles, while the changes observed in males were greater under fluctuating than constant temperature regimes. The onset time of mating was unaffected by male age, however, older females always mated earlier than younger ones, with the advance being more pronounced at cool than warm temperatures. Similar results were also obtained under field conditions. When a single 3-day-old male was provided with 0-, 3-, and 5-day-old females simultaneously, older females obtained mates significantly more often than younger individuals at all temperatures, indicating that calling earlier may afford a reproductive advantage to older females. At both constant temperatures, the time spent mating was longer in older than in younger females, but not under fluctuating thermocycles. Very young and very old males generally spent more time in copula than middle-aged individuals both at constant temperatures and under the warm thermocycle. Under the cool thermocycle, the duration of mating was considerably prolonged at all ages, which could increase the risk of predation.     

Ancestral populations perform better in a novel environment: domestication of medfly populations from five global regions

Geographically isolated populations of a species may differ in several aspects of life-history, morphology, behavior, and genetic structure as a result of adaptation in ecologically diverse habitats. We used a global invasive species, the Mediterranean fruit fly to investigate, whether adaptation to a novel environment differs among geographically isolated populations that vary in major life history components such as life span and reproduction. We used wild populations from five global regions (Kenya, Hawaii, Guatemala, Portugal, and Greece). Adult demographic traits were monitored in F(2), F(5), F(7) and F(9) generations in captivity. Although domestication in constant laboratory conditions had a different effect on the mortality and reproductive rates of the different populations, a general trend of decreasing life span and age of first reproduction was observed for most medfly populations tested. However, taking into account longevity of both sexes, age-specific reproductive schedules, and average reproductive rates we found that the ancestral Kenyan population kept the above life history traits stable during domestication compared to the other populations tested. These findings provide important insights in the life-history evolution of this model species, and suggest that ancestral medfly populations perform better than the derived - invasive ones in a novel environment.     

Sex‐ and age‐biased mortality in wild Grey Partridge Perdix perdix populations

Differing reproductive effort, individual qualities and local environmental conditions can lead to uneven mortality risk among individuals within populations and may result in survival differences according to age and sex. Identification of factors contributing to unequal operational sex ratios has been important for understanding population dynamics and conservation management. In this study, sex‐ and age‐specific mortality was estimated in three wild Grey Partridge populations from analysis of year‐round radiotracking data from 168 individuals. Survival days were counted in three periods defined individually for each bird: the pairing period (covey break‐up to laying of the first egg); the nesting period (between clutch initiation date and failure of the last nesting attempt, or the date when chicks were 14 days old); and the covey period (the end of the nesting period or joining a group until covey break‐up). Predation was the main cause of mortality. A significant effect of age on survival was found during the pairing period, when older individuals paired off faster and survived better. The highest mortality risk overall was found during the nesting period. Furthermore, significantly higher mortality of females was recorded during the nesting period, suggesting that greater investments in reproduction, behaviour at the nest or the quality of nesting habitats can decrease survival of females and cause a male‐skewed sex ratio. No significant effect of age or sex was found during the covey period, or for the year as a whole, but there was a significant difference in annual mortality rates between the three study populations. Our results confirm age‐ and sex‐specific variation of adult mortality in a ground‐nesting bird with biparental care during the annual cycle, documenting differing sensitivities of various population cohorts to predation.     

Sources of variability in copula duration of Mediterranean Fruit Flies

Previous studies have reported considerable variation in copula duration in the Mediterranean Fruit Fly (medfly), Ceratitis capitata Weidemann (Diptera: Tephritidae). Presently, however, this variation is largely unexplained. In this study we measure copula duration in two medfly strains used in applied research in Israel and examine how much of the variation in copula duration is due to the strain, age, the time of day the mating begins and the relative size of the mating pair. We found the following: flies of a strain reared in the laboratory since 1976 (Vienna) copulated for longer than flies of a strain collected in 1969 and periodically refreshed with wild males (Sade); older flies copulated for longer than young flies; and pairs in which males were smaller than females copulated for longer than pairs in which males were larger or the sexes were more evenly size-matched. The time of day the mating commenced (morning vs afternoon) had no effect. We discuss potential mechanisms underlying these variations in medfly copula duration.     

Copula duration and sperm storage in Mediterranean fruit flies from a wild population

In the Mediterranean fruit fly (Ceratitis capitata Weidemann, ‘medfly’), a lekking tephritid, evidence from laboratory studies of flies from laboratory strains suggests that copulation is shorter, and sperm storage more abundant, if males are large or protein‐fed, and that copulation is longer when females are large. In addition, sperm tend to be stored asymmetrically between the female’s two spermathecae and this asymmetry declines with abundance of stored sperm. The primary objective of this study was to investigate whether these trends persist in other experimental contexts that bear closer resemblance to nature. Accordingly, we carried out experiments in a field‐cage using males derived as adults from a wild population and virgin females reared from naturally infested fruit. The results of this study were consistent with laboratory studies in that copula duration increased with female size, that sperm were stored asymmetrically between the females’ spermathecae, and that this asymmetry declined with number of sperm stored. However, we also found some previously unreported effects of female size; large females stored more sperm and stored sperm more asymmetrically between their two spermathecae than did small females. Unlike the laboratory studies, copula duration and sperm storage patterns were unaffected by male size and diet. This may be due to overwhelming variation from other sources in the wild‐collected males used, as well as environmental variability in the semi‐natural setting.     

Effects of adults body size and larvae diet on the fecundity and percent fertility of eggs laid by Xylotrechus arvicola (Coleoptera: Cerambycidae) females,insect pest in Spanish vineyards

Xylotrechus arvicola is a pest of grape in some vine‐producing regions of the Iberian Peninsula. Biological parameters and relationships (fecundity and percent fertility of eggs in relationship to body size) of females obtained in the laboratory and captured in vineyards were studied. In laboratory conditions, the mean developmental time of larvae ranged from 384 to 392 days and pupal stage varied between 12 to 14 days. Body size (BS) of X. arvicola females was significantly bigger than males. Fecundity was greater in the laboratory (147 eggs) than in the field (50 eggs) females, but the percent fertility of the laboratory eggs was lower (16 eggs). Laboratory females showed a bigger relationship between the production of eggs and BS than females captured in vineyards. Wild females (PDO Ribera del Duero and Tierra de León) had a positive relationship between the percent fertility of eggs and the BS. No correlation between the percent fertility of eggs and the BS was displayed by females captured in PDO Toro, but these females had a higher percent fertility (53 eggs) than the others PDO's. These biological parameters and relationships studied suggest that the artificial diet may lack certain essential nutrients that vine varieties can provide that favor the fertility of eggs. This explains why wild females have the potential to become a problem pest in the Tempranillo grape variety, with bilateral cordon and bush vines training systems that have the highest incidence of this cerambycid.     

Improving male mating competitiveness and survival in the field for medfly,Ceratitis capitata (Diptera: Tephritidae) SIT programs

The success of the sterile insect technique (SIT) depends critically upon mating between released sterilized males and wild females. In Hawaii, improvements in the efficiency of sterile males were attempted on two separate fronts – mating enhancement and survival improvement. In the former, two methods have been investigated – selective breeding and aromatherapy. In the latter, flies which survived in field cages for several days were selected and bred to produce progeny with enhanced survival ability compared to control flies. Regarding mating selection, standard laboratory-reared males that successfully mated with wild females in field cages were allowed to breed. F1 offspring were inbred, then the selection procedure was repeated for four additional cycles. In the aromatherapy procedure, laboratory-reared males were exposed to ginger root oil for several hours 1 day prior to testing in field cages. Compared to controls, the selected flies improved the mating competitiveness of male flies ca. 3-fold, irradiation reduced this increase to ca. 2.5-fold. Exposing the selected, hybrid strain raised the fitness of the lab males to ca. 9-fold that of wild males. In the ongoing survival selection study, we have obtained lines in which the selected males survived ca. 2-fold better than laboratory control males over several days in an outdoor field cage, with food and water provided. The goal is to combine the traits of higher survival and mating ability into a single strain for SIT release.     

Protein electrophoresis in Andean condors (Vultur gryphus): Reference values and differences between wild and rehabilitating individuals

The study of wildlife health greatly contributes to understanding population dynamics and detecting conservation threats. The determination of the different fractions of plasma proteins (proteinogram) is an important laboratory tool to study wildlife health. The aim of this study was to characterize protein electrophoresis in wild Andean condors (Vultur gryphus) from north‐western Patagonia and to evaluate differences according to age and sex classes. Once reference values of wild, apparently healthy individuals, were established, we compared these values to those of individuals received at the Buenos Aires Zoo in Argentina for rehabilitation due to various health problems. Reference proteinograms from wild Andean condors differed only in the α 1 and β 2‐fractions between sex categories. Males showed higher concentrations of these protein fractions than females. We found clear differences between wild birds and rehabilitating individuals. Total proteins, globulins, α 1‐globulins, total α‐globulins, β 2‐globulins, total β‐globulins, and γ‐globulins were significantly higher in rehabilitating than in wild individuals, whereas albumin, α 2, and β1‐globulins were similar between these groups. The albumin/globulin ratio, as a general indicator of health, was significantly lower in rehabilitating than in wild individuals. The results indicate the effects on different protein fractions of pathologic processes occurring in individuals undergoing rehabilitation. Our results provide useful insights, contributing to improving diagnoses and prognoses in this species. This information may also be useful to assess the health status of Andean condors in studies of wild populations and for comparisons with other bird species.     

Interplay between ecological, behavioural and historical factors in shaping the genetic structure of sympatric walleye populations (Sander vitreus)

Disentangling ecological, behavioural and evolutionary factors responsible for the presence of stable population structure within wild populations has long been challenging to population geneticists. This study primarily aimed at decoding population structure of wild walleye (Sander vitreus) populations of Mistassini Lake (Québec, Canada) in order to define source populations to be used for the study of spatial partitioning using individual-based multilocus assignment methods, and decipher the dynamics of individual dispersal and resulting patterns of spatial resource partitioning and connectivity among populations. A second objective was to elucidate the relationships between biological characteristics (sex, size, age and population of origin) and an individual's probability to migrate and/or disperse. To do so, a total of 780 spawning individuals caught on five distinct spawning sites, and 1165 postspawning individuals, captured over two sampling seasons (2002-2003) were analysed by means of eight microsatellite loci. Four temporally stable walleye populations associated with distinct reproductive grounds were detected. These populations were differentially distributed among lake sectors during their feeding migration and their spatial distribution was stable over the two sampling seasons. Dispersing individuals were identified (n=61); these revealed asymmetrical patterns of dispersal between populations, which was also confirmed by divergent admixture proportions. Regression models underlined population of origin as the only factor explaining differential dispersal of individuals among populations. An analysis of covariance (ancova) indicated that larger individuals tended to migrate from their river of origin further away in the lake relative to smaller fish. In summary, this study underlined the relevance of using individual-based assignment methods for deciphering dynamics of connectivity among wild populations, especially regarding behavioural mechanisms such as differential spatial partitioning and dispersal responsible for the maintenance of genetic population structure.     

Age-dependent insemination success of sterile Mediterranean fruit flies

The Sterile Insect Technique (SIT) is used in many regions worldwide to manage wild populations of the Mediterranean fruit fly (`medfly'), an important pest species. This technique relies on released sterile males outcompeting their wild counterparts in fertilizing ova of wild females. Numerous studies have investigated the ability of sterile males to secure copulations, an essential step toward overall success. Here we progress further along the mating sequence by studying reproductive barriers that may remain ahead of sterile males that manage to secure copulations in field cage experiments and whether ability to pass these barriers is influenced by a male's age, diet and size, or the size of his mate. Amongst those virgin males that succeeded in copulating, both the number of sperm stored by mates and the chances of having any sperm stored at all decreased with age. Sperm tended to be stored asymmetrically between the females' two spermathecae, and this tendency was more apparent when few sperm were stored. In accord with effects of male age on number of sperm stored, sperm of older males were stored more asymmetrically than that of young males. We found no evidence that male size, male diet or female size influenced copula duration, number of sperm stored or allocation of sperm between the female's two spermathecae. The decline in number of sperm stored as males aged was not accompanied by age-dependent changes in copula duration, indicating that copula duration and insemination success are not deterministically linked. We discuss these results in light of their relevance to SIT and the medfly mating system.     

Swarming and mating activity of Anopheles gambiae mosquitoes in semi‐field enclosures

Anopheles gambiae Giles sensu stricto (Diptera: Culicidae) is the major Afro‐tropical vector of malaria. Novel strategies proposed for the elimination and eradication of this mosquito vector are based on the use of genetic approaches, such as the sterile insect technique (SIT). These approaches rely on the ability of released males to mate with wild females, and depend on the application of effective protocols to assess the swarming and mating behaviours of laboratory‐reared insects prior to their release. The present study evaluated whether large semi‐field enclosures can be utilized to study the ability of males from a laboratory colony to respond to natural environmental stimuli and initiate normal mating behaviour. Laboratory‐reared males exhibited spatiotemporally consistent swarming behaviour within the study enclosures. Swarm initiation, peak and termination time closely tracked sunset. Comparable insemination rates were observed in females captured in copula in the semi‐field cages relative to females in small laboratory cages. Oviposition rates after blood feeding were also similar to those observed in laboratory settings. The data suggest that outdoor enclosures are suitable for studying swarming and mating in laboratory‐bred males in field‐like settings, providing an important reference for future studies aimed at assessing the comparative mating ability of strains for SIT and other vector control strategies.     

Life history evolution in a globally invading tephritid: patterns of survival and reproduction in medflies from six world regions

Comparisons among populations from different localities represent an important tool in the study of evolution. Medflies have colonized many temperate and tropical areas all over the world during the last few centuries. In a common garden environment, we examined whether medfly populations obtained from six global regions [Africa (Kenya), Pacific (Hawaii), Central America (Guatemala), South America (Brazil), Extra-Mediterranean (Portugal) and Mediterranean (Greece)] have evolved different survival and reproductive schedules. Whereas females were either short-lived [life expectancy at eclosion (e₀) 48–58 days; Kenya, Hawaii and Guatemala] or long-lived (e₀ 72–76 days; Greece, Portugal and Brazil], males with one exception (Guatemala) were generally long-lived (e₀ 106–122 days). Although males universally outlived females in all populations, the longevity gender gap was highly variable (20–58 days). Lifetime fecundity rates were similar among populations. However, large differences were observed in their age-specific reproductive patterns. Short-lived populations mature at earlier ages and allocate more of their resources to reproduction early in life compared with long-lived ones. In all populations, females experienced a post-reproductive lifespan, with this segment being significantly longer in Kenyan flies. Therefore, it seems plausible that medfly populations, inhabiting ecologically diverse habitats, have evolved different life history strategies to cope with local environmental conditions.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 106–117.     

Measuring the sperm competition successes of field males of the yellow dung fly

Abstract 1. Sperm competition is extremely common in many species but conclusions about the relative successes of males, measured as the proportion of a female's offspring fertilised by the second male to mate with it, are based largely on laboratory studies using two laboratory-bred males or on interrupted last copulations.
2. A study was conducted in which virgin female yellow dung flies Scathophaga stercoraria mated uninterrupted with two field-captured males.
3. The results confirmed two previous conclusions: the proportion of a female's offspring fertilised by the second male was related strongly to the copula duration of the second male, and the proportion of a female's offspring fertilised by the second male was lower in larger females, though this effect was weak.
4. The results indicate that previous experiments on laboratory populations of this species are likely to be relevant to wild situations, but researchers are urged to conduct studies that will improve the confidence with which laboratory measures of the proportion of a female's offspring fertilised by the second male can be assumed to reflect the sperm precedence of the last male to mate in natural situations.     

Ovulation, insemination and mating in the tsetse fly Glossina pallidipes

ABSTRACT. Analyses of the ovulation, insemination and mating of female Glossina pallidipes Austen, were carried out in the field and laboratory. In unmated females, ovulation was delayed until at least the fifteenth day of adult life. The predictable pattern of growth and development of eggs was used to age nulliparous females to the nearest day in the field. Inseminated females first appeared at 5–6 days old and most females became inseminated at 8–10 days of age. There were no significant differences in the age at insemination in Zimbabwe or Kenya. Ovulation had occurred in the field by 10–11 days old. Mating therefore occurs at or just before ovulation. Females were able to mate at up to 12 days old and still produce a viable larva from their first egg. In laboratory flies originating from Uganda, factors which reduced and minimized the effects of disturbance, maximized insemination rate, so that under the best conditions the age at mating was identical to that found in the field. Nevertheless, the insemination rate of Fl generation Zimbabwe flies reared from wild-caught females was negligible. The differences in the laboratory mating behaviour of the Zimbabwe flies and the flies from the long-established colony of Uganda origin, are considered to be due to wild G. pallidipes being easily disturbed under laboratory conditions, whereas colony breeding rapidly selects for passivity. The concept of gross differences in G. pallidipes mating behaviour between geographic areas in thus rejected.