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The selection of suitable host plants for oviposition is critical for herbivorous insects to maximise survival of their offspring. Olfaction plays an important role in this process. However, little is known about how olfaction shapes the interaction between the fall armyworm (Spodoptera frugiperda) and host plants. In this study, we tested the hypothesis that olfaction guides the host selection process in the fall armyworm using oviposition and wind tunnel bioassays.In no-choice and dual-choice assays, female moths oviposited on all seven host plants that were tested (maize, sorghum, wheat, bean, cowpea, tomato and cabbage). However, in multiple-choice assays, no eggs were deposited on cowpea and cabbage. We found that maize, sorghum and wheat were most preferred for oviposition, whereas cowpea was least preferred. Wind tunnel assays confirmed these divergent oviposition preferences, with maize, sorghum and wheat odours being the most attractive. Gas chromatography-mass spectrometry analysis followed by random forest classification identified terpenes as the potential host-plant attractants.Our results improve our understanding of the chemical ecology of the fall armyworm and suggest that some of these host plants could offer potential for use in an intercropping strategy to manage S. frugiperda. © 2022 Society of Chemical Industry.

IntroductionA changing environment can select on life-history traits and trade-offs in a myriad of ways. For example, global warming may shift phenology and thus the availability of host-plants. This may alter selection on survival and fertility schedules in herbivorous insects. If selection on life-histories changes, this may in turn select for altered nutrient intake, because the blend of nutrients organisms consume helps determine the expression of life-history traits. However, we lack empirical work testing whether shifts in the timing of oviposition alter nutrient intake and life-history strategies.MethodsWe tested in the marula fruit fly, Ceratitis cosyra, how upward-selection on the age of female oviposition, in comparison with laboratory adapted control flies, affects the sex-specific relationship between protein and carbohydrate intake and life-history traits including lifespan, female lifetime egg production and daily egg production. We then determined the macronutrient ratio consumed when flies from each selection line and sex were allowed to self-regulate their intake.ResultsLifespan, lifetime egg production and daily egg production were optimised at similar protein to carbohydrate (P:C) ratios in flies from both selection lines. Likewise, females and males of both lines actively defended similar nutrient intake ratios (control =1:3.6 P:C; upward-selected = 1:3.2 P:C).DiscussionOur results are comparable to those in non-selected C. cosyra, where the optima for each trait and the self-selected protein to carbohydrate ratio observed were nearly identical. The nutrient blend that needs to be ingested for optimal expression of a given trait appeared to be well conserved across laboratory adapted and experimentally selected populations. These results suggest that in C. cosyra, nutritional requirements do not respond to a temporal change in oviposition substrate availability.

IntroductionTemperature fluctuations are important for the distribution and survival of insects. Rapid hardening, a type of phenotypic plasticity, is an adaptation that can help individuals better tolerate lethal temperatures because of earlier exposure to a sublethal but stressful temperature. Nutrition and sex are also known to influence a species ability to tolerate thermal stress. This study determined the effects of larval diet, adult diet, sex and hardening on the thermal tolerance of Ceratitis cosyra (Walker) (Diptera: Tephritidae) at lower and upper lethal temperatures.MethodsLarvae were raised on either an 8% torula yeast (high) or a 1% torula yeast (low) larval diet and then introduced to one of three dietary regimes as adults for thermal tolerance and hardening assays: no adult diet, sugar only, or sugar and hydrolysed yeast diet. Flies of known weight were then either heat- or cold-hardened for 2 hours before being exposed to a potentially lethal high or low temperature, respectively.ResultsBoth nutrition and hardening as well as their interaction affected C. cosyra tolerance of stressful temperatures. However, this interaction was dependent on the type of stress, with nutrient restriction and possible adult dietary compensation resulting in improved cold temperature resistance only.DiscussionThe ability of the insect to both compensate for a low protein larval diet and undergo rapid cold hardening after a brief exposure to sublethal cold temperatures even when both the larva and the subsequent adult fed on low protein diets indicates that C. cosyra have a better chance of survival in environments with extreme temperature variability, particularly at low temperatures. However, there appears to be limitations to the ability of C. cosyra to cold harden and the species may be more at risk from long term chronic effects than from any exposure to acute thermal stress.

Bioconversion is the process whereby nutrients are recovered from organic waste products, often by flies, to produce value-added products such as protein for animal feed and lipids for biodiesel production. Currently, research and industry focus on a select few fly species for use in bioconversion that are generalists in their feeding behaviour, the black soldier fly, Hermetia illucens L., and the housefly, Musca domestica L. More investigation is needed on species of flies that are more suited to underutilised waste streams, including meat processing waste. Species of flies that breed in carrion, such as blowflies (Calliphoridae) and flesh flies (Sarcophagidae) can be used to reduce meat processing waste and produce a valuable source of protein. In this review, we propose more investigation and use of a wider range of fly species for bioconversion of organic waste. Four blowfly species are recommended for use in the bioconversion of meat processing waste or a mixture of manure waste and meat processing waste. Chrysomya chloropyga (Wiedemann) is a large mammal carcass specialist and has been found to be effective at recovering nutrients from abattoir waste within four days and producing large larvae in the process. Chrysomya putoria (Wiedemann) and Chrysomya megacephala (Wiedemann) naturally breed in carrion and faeces, are associated with pit latrines and respond well under mass rearing conditions, with high egg production. These species would be recommended for a large-scale bioconversion facility that receives mixed waste streams including manure and animal remains. Lucilia sericata is known to produce antimicrobial compounds that assist in wound healing and has been frequently bred and studied and responds well to lab and mass rearing conditions. We recognise the potential obstacles to using alternative species in industrial-scale bioconversion facilities and pose future directions for research to overcome these challenges.

Understanding the dynamics of larval habitat utilization by mosquito communities is crucial for the design of efficient environmental control strategies. The authors investigated the structure of mosquito communities found at hotel compounds in Zanzibar, networks of mosquito interactions with larval habitats and robustness of mosquito communities to elimination of larval habitats. A total of 23 698 mosquitoes comprising 26 species in six genera were found. Aedes aegypti (n = 16 207), Aedes bromeliae/Aedes lillie (n = 1340), Culex quinquefasciatus (n = 1300) and Eretmapodites quinquevitattus (n = 659) were the most dominant species. Ecological network analyses revealed the presence of dominant, larval habitat generalist species (e.g., A. aegypti), exploiting virtually all types of water holding containers and few larval habitat specialist species (e.g., Aedes natalensis, Orthopodomyia spp). Simulations of mosquito community robustness to systematic elimination of larval habitats indicate that mosquito populations are highly sensitive to elimination of larval habitats sustaining higher mosquito species diversity. This study provides insights on potential foci of future mosquito-borne arboviral disease outbreaks in Zanzibar and underscores the need for detailed knowledge on the ecological function of larval habitats for effective mosquito control by larval sources management.

BACKGROUND The sterile insect technique (SIT) is used in Australia to contain and eradicate outbreaks of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) in fruit-fly-free regions, and to manage populations in some endemic regions. To assess the likely impact of SIT on wild B. tryoni populations it is important to assess the colocation and synchrony of male calling between sterile and wild flies. We observed the location and timing of calling behaviours of marked mass-reared (fertile and sterile) and wild B. tryoni males in walk-in field cages. RESULTS We found that wild males called further from the canopy centre than mass-reared (fertile or sterile) males. Mass-reared (fertile or sterile) males called earlier in the evening than wild males and, consequently, mass-reared males called when temperature and light intensity were higher than when wild males called. CONCLUSION Male calling is a prerequisite to mating among dacine fruit flies. Therefore, our observations of spatio-temporal divergence in male calling behaviour may lead to assortative mating between mass-reared and wild B. tryoni in SIT applications. The importance of these spatio-temporal differences warrants further inquiry, with particular focus on how environmental conditions modify calling behaviour and avenues to ameliorate differences between sterile and wild flies. © 2021 Society of Chemical Industry.

Females and males have divergent strategies of energy investment, so the thermal preference of each sex in insects may differ because energetic conversion of metabolic reserves is dependent on temperature. We determined the thermal preference of virgin, sexually mature Mediterranean fruit flies, Ceratitis capitata, and found that males preferred a significantly higher temperature (23.8 ± 0.3 °C) than that of females (22.1 ± 0.3 °C). We then tested predictions for the difference in thermal preference related to the energetic demands of reproduction over a range of temperatures. The frequency and duration of calling bouts by male C. capitata were optimal at 26 °C. Mating propensity and latency, and copula duration, were optimal over the range of 22-28 °C. When mating occurred, temperature had little effect on the incidence of sperm storage by females, but there was a notable decline in the number stored at 28 °C. Female lifespan was highest at 18 °C, but lifetime egg production was optimal at 24 °C. These results illustrate temperature-related differences in the reproductive fitness of the sexes in C. capitata, although the optima for male traits align best with their thermal preference. They also support the theoretical prediction that insect thermal preference should be lower than the optimum for fitness.

Citrus Research International; Hortgro Science; South African Table Grape Industry and National Research Foundation.

The South African National Research Foundation (SFH150718127604), AgriProtein Technologies, South Africa and a University of Pretoria Research Development Programme grant.

The expression of life-history traits, such as lifespan or reproductive effort, is tightly correlated with the amount and blend of macronutrients that individuals consume. In a range of herbivorous insects, consuming high protein to carbohydrate ratios (P:C) decreases lifespan but increases female fecundity. In other words, females face a resource-based trade-off between lifespan and fecundity. Redox metabolism may help mediate this trade-off, if oxidative damage is elevated by reproductive investment and if this damage, in turn, reduces lifespan. Here, we test how diets varying in P:C ratio affect oxidative damage and antioxidant protection in female and male of the marula fly

Background A field survey was performed to investigate local environmental factors promoting occurrence and abundance of Aedes aegypti and Ae. bromeliae mosquitoes at hotel compounds in the south-east coastal region of Zanzibar Island. Methods The potential risk factors were determined using generalized linear mixed models. Aedes (Stegomyia) spp. indices such as container index (CI) and pupae per container (PPC) index were also estimated. Results Aedes aegypti and Ae. bromeliae were the most abundant vector species, accounting for 70.8% of all Aedes mosquitoes collected. The highest CI was observed for plastic containers irrespective of the season, whereas the highest PPC was observed for coconut shells and aluminium containers in the rainy and dry seasons, respectively. The risk of Aedes mosquito occurrence and abundance were significantly associated with presence of plastic containers, coconut shells, used tyres and steel containers. These were discarded in shaded places, in the open and gardens, or found in plant nurseries. Conclusion This study shows that Aedes species of global health significance occur at hotel compounds on this part of Zanzibar Island. The occurrence and abundance are sustained by the presence of abundant and poorly managed solid wastes and containers used for gardening tasks. This highlights an urgent need for the adoption of area-wide environmentally sustainable Aedes mosquito management interventions that also integrate solid waste management and ornamental plant production practices for reducing the risk of arboviral disease epidemics. Graphical Abstract

The trade-off between lifespan and reproduction is central to our understanding of life-history evolution. Laboratory selection experiments have been a powerful tool for quantifying this trade-off, but these tend to be restricted in taxonomic scope, which may limit our understanding. In addition, research often focuses on the trade-off between lifespan and reproductive effort in females, and far less data test how lifespan trades off with different aspects of male reproduction (e.g. pre- and post-copulatory reproductive investment). Here, we examined the trade-off between lifespan and reproduction in females and males of the marula fruit fly, Ceratitis cosyra (Walker) (Diptera: Tephritidae). To do so, we selected downward or upward on age of peak female egg laying in C. cosyra for twenty generations. In multiple generations, we measured female and male lifespan and body size, female daily and lifetime fecundity, male courtship and mating success, as well as the number of sperm transferred at different ages and sperm storage asymmetry in spermathecae. Our selection regime appeared to achieve its aim; egg laying peaked earlier in females from downward selected lines than upward selected lines. The number of sperm transferred by males decreased in the upward selected flies, but other male reproductive traits remained the same across selection regimes. In contrast, with the wider literature, upward selection did not extend the lifespan of females or males after ten generations of selection. While lifespan in both sexes responded to selection on female egg laying schedules, it did not do so in a straightforward way. Moreover, male investment in reproductive traits was largely independent of selection regime. These counter-intuitive findings highlight the importance of working with a broad range of species and of considering the trade-off between reproduction and lifespan in both sexes. The trade-off between lifespan and reproduction has been extensively studied in model species using various types of laboratory selection. A limited number of species have been considered using this approach, and the majority of the studies have focused on female, rather than male, reproductive effort. Here, we selected downwards and upwards on age of female reproduction in the marula fruit fly and measured survival, female fecundity, reproductive schedule, as well as male sperm transfer, sperm storage asymmetry, mating and calling success. We found a moderate trade-off between lifespan and early fecundity in downward selected flies, whereas no obvious trade-off was observed in upward selected lines. Regardless of the selection regime, reproductive scheduling was affected in females but not in males, while lifespan was affected in both sexes. Our results show that the timing of reproduction can evolve independently across the sexes, highlighting the importance of studying both females and males.

Insects tend to live within well-defined habitats, and at smaller scales can have distinct microhabitat preferences. These preferences are important, but often overlooked, in applications of the sterile insect technique. Different microhabitat preferences of sterile and wild insects may reflect differences in environmental tolerance and may lead to spatial separation in the field, both of which may reduce the control program efficiency. In this study, we compared the diurnal microhabitat distributions of mass-reared (fertile and sterile) and wild Queensland fruit flies, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae). Flies were individually tagged and released into field cages containing citrus trees. We recorded their locations in the canopies (height from ground, distance from canopy center), behavior (resting, grooming, walking, feeding), and the abiotic conditions on occupied leaves (temperature, humidity, light intensity) throughout the day. Flies from all groups moved lower in the canopy when temperature and light intensity were high, and humidity was low; lower canopy regions provided shelter from these conditions. Fertile and sterile mass-reared flies of both sexes were generally lower in the canopies than wild flies. Flies generally fed from the top sides of leaves that were lower in the canopy, suggesting food sources in these locations. Our observations suggest that mass-reared and wild B. tryoni occupy different locations in tree canopies, which could indicate different tolerances to environmental extremes and may result in spatial separation of sterile and wild flies when assessed at a landscape scale.

1. Hopkins' host selection principle (HHSP) proposes that, in holometabolous insects, gravid females prefer to oviposit on their developmental substrate. This hypothesis is widely explored in phytophagous insects, but few studies have considered blood‐feeding insects. In this study, the HHSP was tested using the haematophagous stable fly, Stomoxys calcitrans (Diptera: Muscidae). Using no‐choice and two‐choice tests, the study first sought to demonstrate whether S. calcitrans larvae recognise their rearing substrate. 2. It was found that when a rearing substrate is offered to S. calcitrans larvae singly (no‐choice) or associated with a non‐rearing substrate (two‐choice), they were able to recognise the substrate in which they developed. This ability disappeared when larvae were transferred to another substrate for 5 h. 3. Next, using oviposition bioassays, it was investigated whether information gathered by pre‐imaginal stages during their development can persist into the adult stages after metamorphosis. It was hypothesised that gravid females emerging from clean and uncleaned pupae reared on sheep and camel dung would still prefer to lay on their rearing substrate. The results revealed that gravid females did not prefer to oviposit on their developmental substrate. In addition, failure to recognise and prefer their developmental substrate was found in gravid females that had emerged from puparia that were cleaned as well as those contaminated with sheep dung. 4. It is concluded that S. calcitrans adult preference is not influenced by their offspring's experience. Overall, the HHSP was not supported by our study, suggesting that this principle may not apply to generalist insects characterised by high plasticity in oviposition substrate acceptance.

The oxidative damage caused to cells by reactive oxygen species (ROS) is one of several factors implicated in causing ageing. Oxidative damage may also be a proximate cost of reproductive effort that mediates the trade-off often observed between reproduction and survival. However, how the balance between oxidative damage and antioxidant protection affects life-history strategies is not fully understood. To improve our understanding, we selected on female reproductive age in the marula fruit fly, Ceratitis cosyra, and quantified the impact of selection on female and male age-dependent mortality, female fecundity, male sperm transfer, calling and mating. Against expectations, upward-selected lines lived shorter lives and experienced some reductions in reproductive performance. Selection affected oxidative damage to lipids and total antioxidant protection, but not in the direction predicted; longer lives were associated with elevated oxidative damage, arguing against the idea that accumulated oxidative damage reduces lifespan. Greater reproductive effort was also associated with elevated oxidative damage, suggesting that oxidative damage may be a cost of reproduction, although one that did not affect survival. Our results add to a body of data showing that the relationship between lifespan, reproduction and oxidative damage is more complex than predicted by existing theories.

In herbivorous insects, the degree of host specialisation may be one ecological factor that shapes lifespan. Because host specialists can only exploit a limited number of plants, their lifecycle should be synchronised with host phenology to allow reproduction when suitable hosts are available. For species not undergoing diapause or dormancy, one strategy to achieve this could be evolving long lifespans. From a physiological perspective, oxidative stress could explain how lifespan is related to degree of host specialisation. Oxidative stress caused by Reactive Oxygen Species (ROS) might help underpin ageing (the Free Radical Theory of Aging (FRTA)) and mediate differences in lifespan. Here, we investigated how lifespan is shaped by the degree of host specialisation, phylogeny, oxidative damage accumulation and antioxidant protection in eight species of true fruit flies (Diptera: Tephritidae). We found that lifespan was not constrained by species relatedness or oxidative damage (arguing against the FRTA); nevertheless, average lifespan was positively associated with antioxidant protection. There was no lifespan difference between generalist and specialist species, but most of the tephritids studied had long lifespans in comparison with other dipterans. Long lifespan may be a trait under selection in fruit-feeding insects that do not use diapause.

Start-up funds were provided to CWW by the University of Pre-toria in the form of a Research Development Programme grant. Further research funding, as well as a Grant Holder-Linked Masters Scholarship to PDR, was also provided by the South African National Research Foundation through a Competitive Programme for Rated Researchers grant (Grant No: 93686) to CWW and CRA.

Markets importing citrus fruit including lemons, Citrus limon (L.) Burman f., from South Africa require that the fruit be free of fruit fly pests (Diptera: Tephritidae). Historically there has been no fruit fly infestation recorded on lemons destined for export from South Africa. In this study, we assessed the host status of commercial export grade Eureka lemons, Citrus limon (L.) Burmanf. cv. Eureka, for four fruit fly pest species of economic importance in South Africa: Ceratitis capitata (Wiedemann), Ceratitis rosa Karsch, Ceratitis quilicii De Meyer, Mwatawala & Virgilio, and Bactrocera dorsalis (Hendel). Trapping was conducted in 10 Eureka lemon orchards in two major citrus production regions over two citrus seasons between 2016 and 2017 to determine the level of fruit fly abundance in the sampled orchards. Lemons were collected at harvest over the two seasons in the same orchards where trapping was conducted. Fruit fly infestation of the sampled lemons was determined by dissection. Additionally, infestation of lemons was determined under forced exposure to mature mated females of C. capitata and B. dorsalis. Trapping data showed the presence of adults of all four fruit fly species in the sampled lemon orchards. No fruit fly infestation was detected in 43 222 Eureka lemons sampled at harvest. There was also no infestation of lemons under forced exposure conditions. The results of this study provide evidence with 99.99 % efficacy and a 99%confidence level that SouthAfrican commercial export grade Eureka lemon fruit is not a host for C. capitata, C. rosa, C. quilicii or B. dorsalis.

The highly invasive oriental fruit fly, Bactrocera dorsalis (Hendel), has been declared present in the northern parts of South Africa since 2013. A study was thus initiated in July 2014 to determine the host range and field population of the pest species in the region. Fruit were collected from commercial fruit production, interface (smaller commercial blocks surrounded by natural savannah vegetation) and natural areas (savannah vegetation) throughout Limpopo and Mpumalanga provinces, South Africa. Field sites consisted of five commercial fruit production sites, two interface sites and two natural areas. Fruit samples from the tree and the ground were collected and incubated separately to determine infesting fruit fly species and the degree of infestation. Adult B. dorsalis populations were monitored at each field site using three methyl eugenol-baited bucket traps to estimate population pressure and to determine with the use of time series analysis if monthly trap captures were correlated with fruit infestation. Bactrocera dorsalis was reared from seven plant species: two from commercial orchards (Mangifera indica cv. [Tommy Atkins, Sensation], Citrus sinensis cv. [Valencia]), and five from other plant species (Psidium guajava, Anacardium occidentale, Solanum mauritianum, Xylotheca kraussiana, Vangueria infausta). Fruit utilized by B. dorsalis was also infested or damaged by other species, which may indicate opportunism by the pest, and the potential for competitive interactions. Time series analyses show adult population increased 2 months after an increase in mean temperature in all sites, 4 months after rainfall in natural and interface sites, and 1 month and 3 months after fruit infestation in commercial and natural and interface sites, respectively. This study shows B. dorsalis utilizing a limited range of hosts in South Africa. However, the host range of B. dorsalis may expand as it may not yet have encountered all potential hosts.

Daylight fluorescent pigment powders are frequently used to self-mark tephritid flies that are released in sterile insect technique programmes and for studies on their population ecology, movement and behaviour. This study was conducted to determine the effects of pigment colour and dose in marking the Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae). Six pigment colours (Astral Pink 1, Blaze 5, Stellar Green 8, Lunar Yellow 27, Comet Blue 60 and Invisible Blue 70) were applied to pupae at doses of 0, 2, 4, or 6 g/l. Under laboratory conditions, pigment colour had a small but significant effect on the number of partially emerged and deformed adults; the fewest of these were observed when flies were marked with Astral Pink 1. Pigment concentration, on the other hand, had no effect on adult emergence, partial emergence, deformed adults and mortality on the last day of eclosion. There was no significant effect of pigment colour on adult survival under laboratory and semi-field conditions. Under laboratory conditions, however, there was an effect of pigment concentration on adult survival depending on pigment colour. Visibility under an ultraviolet light and persistence of marks was significantly affected by pigment colour and concentration when observed under laboratory conditions, but not under semi-field conditions. Regardless of colour or dose, pigments used in the study were visible for at least 14 days, but began to fade by 21 days after adult eclosion. To mark B. dorsalis under temperate, warmsummer African conditions, all pigment colours tested in this study may be applied at 2–4 g/l pupae. Recaptures of marked and released flies may be underestimated as the flies age.

Blowfly larvae aggregate on exposed carcasses and corpses and pass through three instars before wandering from the carcass and pupating. The developmental landmarks in this process can be used by forensic entomologists to estimate the time since the insects colonised the carcass, which sets a minimum post mortem interval. Large aggregations of feeding larvae generate a microclimate with temperatures up to 15°C above ambient conditions, which may accelerate larval development and affect forensic estimates of post-mortem intervals. This study investigated the effects of heat accumulated by maggot masses of Lucilia cuprina at aggregations of 20, 50 and 100 larvae, each at incubation temperatures of 18°C, 24°C and 30°C, using body length and life stage as developmental indicators. Aggregation temperatures reached up to 18.7°C above ambient temperature, with significant effects of both size and temperature of the aggregation on the development time of its larvae. Survivorship was highest for all life stages at 24°C, which is near the developmental optimum of L. cuprina. The results of this study provide a broadly applicable method of quantifying heat accumulation by aggregations of a wide range of species of forensic importance, and the results obtained from such studies will demonstrate that ambient temperature cannot be considered the only source of heat that blowfly larvae experience when they develop on a carcass. Neglect of temperatures within larval aggregations will result in an overestimation of post-mortem intervals and thus have far-reaching medicolegal consequences.

Background Oviposition site selection by gravid female insects is an important determinant in species distribution, abundance, and population dynamics. Females may assess the suitability of a potential oviposition substrate by using cues from conspecific or heterospecific individuals already present. Here, we assessed whether the presence of conspecific, heterospecific larvae and parasites influenced oviposition decisions by the stable fly, Stomoxys calcitrans (Linneaus). Methods and Results Using dual and multiple-choice oviposition bioassays, we found that gravid female S. calcitrans avoided substrates with conspecific larvae, the larvae of house flies, Musca domestica (Linneaus), and the mite Macrocheles muscaedomesticae (Scopoli). Avoidance of conspecific and heterospecific larvae persisted in the dark, suggesting that this behaviour is mediated by olfactory cues. When we reared S. calcitrans in the presence of conspecific larvae and the larvae of house flies at different densities we found that this negatively affected emergence time, larval weight, larval survival, pupal weight, pupal survival, and adult weight. We also demonstrated that individuals of S. calcitrans developed in the presence of mites exhibited low egg hatchability, and poor larval and adult survival. Conclusion Our study provides additional support for the “preference-performance” hypothesis in S. calcitrans, with gravid females preferring to lay eggs on a substrate that will enhance offspring fitness. We recommend that the olfactory cues involved in avoidance by gravid female S. calcitrans of substrates with conspecific and heterospecific larvae should be elucidated. This could lead to the discovery of repellent chemicals important for S. calcitrans management.

Selection of oviposition substrate is critical in holometabolous insects. Female stable flies, Stomoxys calcitrans, locate and select vertebrate herbivore dung in which they lay their eggs. However, the preference for vertebrate herbivore dung by S. calcitrans females, its fitness consequences for offspring, and the semiochemicals used to locate and select oviposition substrates remain unclear. Using oviposition choice tests and life table bioassays we found that gravid female S. calcitrans prefer to oviposit on donkey and sheep dung, which also improves the performance of their offspring. GC-MS analysis followed by random forest classification identified β-citronellene and carvone as the most important predictive volatile organic compounds of donkey and sheep dung, respectively. In multiple choice oviposition bioassays, S. calcitrans laid more eggs in wet sand containing β-citronellene and carvone than in other treatments. The attractiveness of these compounds was confirmed in a field trial, with traps baited with β-citronellene and carvone catching more S. calcitrans. We conclude that gravid female S. calcitrans use semiochemical cues to choose oviposition substrates that maximise offspring fitness.

Adult holometabolous insects may derive metabolic resources from either larval or adult feeding, but little is known of whether adult diets can compensate for deficiencies in the larval diet in terms of stress resistance. We investigated how stress resistance is affected and compensated for by diet across life stages in the marula fruit fly, Ceratitis cosyra (Walker) (Diptera: Tephritidae). Larvae were fed diets containing either 8% torula yeast, the standard diet used to rear this species, or 1% yeast (low protein content similar to known host fruit). At emergence, adults from each larval diet were tested for initial mass, water content, body composition, and desiccation and starvation resistance or they were allocated to one of two adult diet treatments: sucrose only, or sucrose and yeast hydrolysate. The same assays were then repeated after 10 days of adult feeding. Development on a low protein larval diet led to lower body mass and improved desiccation and starvation resistance in newly emerged adults, even though adults from the high protein larval diet had the highest water content. Adult feeding decreased desiccation or starvation resistance, regardless of the diet provided. Irrespective of larval diet history, newly emerged, unfed adults had significantly higher dehydration tolerance than those that were fed. Lipid reserves played a role in starvation resistance. There was no evidence for metabolic water from stored nutrients extending desiccation resistance. Our findings show the possibility of a nutrient-poor larval environment leading to correlated improvement in adult performance, at least in the short term.

The five-spotted fruit fly, Ceratitis quinaria (Bezzi) (Diptera: Tephritidae), is distributed in parts of eastern and western Africa, northern Africa and southern Africa (including South Africa). The species also occurs outside of Africa, in Yemen. The host range of C. quinaria is narrow, with Mangifera indica L. being the main host of commercial importance. Ceratitis quinaria was claimed to be associated with citrus in Sudan although this claim was never substantiated and there has never been any confirmed reared record of C. quinaria on citrus despite numerous surveys of citrus across Africa. In order to verify the type of association that C. quinaria has with citrus, field surveys were carried out in South Africa to determine the distribution and seasonal occurrence of C. quinaria in commercial citrus orchards and the possible natural infestation of citrus by this species. Surveys on distribution of C. quinaria across South Africa were carried out in 1999 and 2000 by trapping with Ceratitislure (containing protein hydrolysate and β-caryophyllene). The seasonal occurrence of C. quinaria in the northern areas of South Africa was determined over two years between 2015 and 2017 by trapping with Enriched Ginger Oil (EGO), a male lure containing α-copaene. In the same trapping period, citrus and other fruit were sampled to determine infestation by C. quinaria. Additionally between 2009 and 2018, citrus fruit was sampled from the trees and ground in other commercial and non-commercial areas in the north of South Africa. All fruit samples collected were incubated for at least five weeks to allow rearing of flies to the pupal and adult stages. Trapping surveys conducted between 1999 and 2000 showed the presence of C. quinaria only in the northern areas of South Africa. Catches of C. quinaria males in EGO-baited traps were low in commercial citrus orchards (peak of catches being lower than 0.05 flies/trap/day). Catches of C. quinaria were mainly recorded outside of the citrus ripening period. No C. quinaria was reared from any of the citrus fruit sampled, even in those areas where the presence of the species was demonstrated by catches in EGO-baited traps. The fruit surveys therefore demonstrated the absence of natural infestation of citrus with C. quinaria in South Africa and supported existing biological information that citrus is not a host for C. quinaria.

Adult holometabolous insects may derive metabolic resources from either larval or adult feeding, but little is known of whether adult diets can compensate for deficiencies in the larval diet in terms of stress resistance. We investigated how stress resistance is affected and compensated for by diet across life stages in the marula fruit fly

The blowfly Lucilia cuprina is a primary colonizer of decaying vertebrate carrion, and its development provides a temperature-dependent clock that may be used to estimate the post-mortem interval of corpses and carcasses in medicolegal forensic investigations. This study uses the development of L. cuprina raised on a substrate of chicken liver at six constant temperatures from 18 to 33 °C to calibrate a thermal accumulation model of development for forensic applications. Development was optimal near 24 °C; above this temperature, survival of post-feeding life stages was increasingly compromised, while below it, development was increasingly retarded. The lower developmental threshold (~12 °C) and thermal summation constants of L. cuprina are distinct from those reported for Lucilia sericata, verifying that it is essential to identify African Lucilia specimens accurately when using them to estimate post-mortem intervals.

Large numbers of flies are needed to produce the quantity of larvae required for insect bioconversion of waste. However, this 'mass-rearing' may negatively affect adult survival and reproductive output. This study assessed the suitability for mass-rearing of four blow fly species, Chrysomya chloropyga, Chrysomya chloropyga (Wiedemann), Chrysomya megacephala (F.), Chrysomya putoria (Wiedemann) and Lucilia sericata (Meigen). Flies were kept at densities of 20, 50, 100, 250, 500, and 1,000 flies per 30 × 30 × 30 cm cage with an even sex ratio. Time to 50% mortality (LT50) was recorded, and the effects of density, species, and sex on LT50, fecundity, and fertility were determined. Females survived longer than males across all species. There was evidence for a trade-off between survival and high fecundity in L. sericata and C. chloropyga at density 250. C. megacephala had low fecundity across all densities. At high densities, C. putoria had the lowest mortality and highest fecundity, making it the most suitable for mass-rearing.

In pest management programmes that incorporate the sterile insect technique (SIT), the ability of mass-reared insects to tolerate dry conditions may influence their survival after release in the field. In the present study, desiccation resistance of adult mass-reared Queensland fruit flies,Bactrocera tryoni(Frogatt) (Diptera: Tephritidae), that are routinely released in SIT programmes was compared with that of wild flies at 1, 10 and 20 days after adult eclosion. Under dry conditions without access to food or water, longevity of mass-rearedB. tryoniwas significantly less than that of their wild counterparts. Desiccation resistance of mass-reared flies declined monotonically with age, but this was not the case for wild flies. The sharp decline in desiccation resistance of mass-reared flies as they aged was likely explained by decreased dehydration tolerance. As in an earlier study, desiccation resistance of females was significantly lower than that of males but this was particularly pronounced in mass-reared females. Female susceptibility to dry conditions corresponded with declining dehydration tolerance with age and associated patterns of reproductive development, which suggests that water content of their oocyte load is not available for survival during periods of water stress.

Seasonal, spatial, and habitat responses of carrion-associated Diptera assemblages can provide valuable information about the presence or absence of species and their relative abundance, and thereby enhance understanding of their responses to environmental variables and how this may have an impact on forensic investigations. Three different nature reserves (localities) within the Municipality of Tshwane, South Africa, were selected to determine whether species assemblages of carrion-feeding flies differ between seasons, localities, and habitat types. A total of 59,511 adult Diptera, identified to 35 species in eight different families, were collected using modified Redtop hanging traps, baited with liver and fish, during four seasons in three different habitat types. Species assemblages differed temporally, with season being the main factor determining species diversity and not locality or habitat. However, savanna and human-disturbed habitats supported a higher abundance and species richness than grassland habitats. Areas adjacent to the localities, such as large urban expanses in Dinokeng or agricultural holdings in Rietvlei, led to an increase in the abundance and mean species richness of carrion-associated Diptera, and in increased numbers of pest or invasive species such as Chrysomya megacephala (F.). Despite this, the overall species assemblages present in human-disturbed areas were very similar to those recorded in natural habitats.

Background Mosquito lifespan can influence the circulation of disease causing pathogens because it affects the time available for infection and transmission. The life-cycle of mosquitoes is determined by intrinsic and environmental factors, which can include the availability of hosts and suitable resting environments that shelter mosquitoes from extreme temperature and desiccating conditions. This study determined the parity rates (an indirect measure of survival) and plant resting preference of vectors of Rift Valley fever (RVF) in northeastern Kenya. Methods Resting mosquitoes were trapped during the rainy and the dry season using a Prokopack aspirator from vegetation, whereas general adult populations were trapped using CDC light traps. At each site, sampling was conducted within a 1 km2 area, subdivided into 500 × 500 m quadrants and four 250 × 250 m sub-quadrants from which two were randomly selected as sampling units. In each sampling unit, plants were randomly selected for aspiration of mosquitoes. Only Aedes mcintoshi and Ae. ochraceus were dissected to determine parity rates while all mosquito species were used to assess plant resting preference. Results Overall, 1124 (79 %, 95 % CI = 76.8–81.1 %) mosquitoes were parous. There was no significant difference in the number of parous Ae. mcintoshi and Ae. ochraceus. Parity was higher in the rainy season than in the dry season. Daily survival rate was estimated to be 0.93 and 0.92 among Ae. ochraceus and Ae. mcintoshi, respectively. Duosperma kilimandscharicum was the most preferred plant species with the highest average capture of primary (3.64) and secondary (5.83) vectors per plant, while Gisekia africana was least preferred. Conclusion Survival rate of each of the two primary vectors of RVF reported in this study may provide an indication that these mosquitoes can potentially play important roles in the circulation of diseases in northern Kenya. Resting preference of the mosquitoes in vegetation may influence their physiology and enhance longevity. Thus, areas with such vegetation may be associated with an increased risk of transmission of arboviruses to livestock and humans.

Acclimation in the thermal tolerance range of insects occurs when they are exposed to novel temperatures in the laboratory. In contrast to the large number of studies that have tested for the ability of insects to acclimate, relatively few have sought to determine the time-course for attainment and reversal of thermal acclimation. In this study the time required for the Mediterranean fruit fly, Ceratitis capitata Wiedemann, and the Natal fruit fly, Ceratitis rosa Karsch, to acclimate to a range of constant temperatures was tested by determining the chill-coma recovery time and heat knock-down time of flies that had been exposed to novel benign temperatures for different durations. The time required for reversal of acclimation for both Ceratitis species was also determined after flies had been returned to the control temperature. Acclimation to 31 °C for only one day significantly improved the heat knock-down time of C. capitata, but also led to slower recovery from chill-coma. Heat knock-down time indicated that acclimation was achieved after only one day in C. rosa, but it took three days for C. rosa to exhibit a significant acclimation response to a novel temperature of 33 °C when measured using chill-coma recovery time. Reversal of acclimation after return to initial temperature conditions was achieved after only one day in both C. capitata and C. rosa. Adult C. capitata held at 31.5 °C initially exhibited improved heat knock-down times but after 9 days the heat knock-down time of these flies had declined to levels not significantly different from that of control flies held at the baseline temperature of 24 °C. In both Ceratitis species, heat knock-down time declined with age whereas chill-coma recovery time increased with age, indicating an increased susceptibility to high and low temperatures, respectively.

Post-teneral dietary supplements have been found to improve mating performance of male Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) (Q-fly) and show considerable promise for enhancement of sterile insect technique (SIT) programmes even when applied within the current 48 h pre-release holding period. However, Q-flies are released as a bisexual strain, and the positive effects of a diet including yeast hydrolysate for males may also boost reproductive development and sexual performance of females. Increased prevalence of mature sterile females can substantially dilute SIT efficacy as mating capacity of sterile males is largely depleted by sterile females rather than the relatively rare wild females. Here, we demonstrate that providing yeast hydrolysate for 48 h after adult emergence, emulating the current pre-release holding period of Q-fly SIT, leads to a significant increase in reproductive development and sexual performance in male Q-flies. In contrast, female Q-flies with access to yeast hydrolysate for 48 h had ovaries that were poorly developed and, particularly at younger ages, were less likely to mate and remate than females with continuous, ad libitum access to yeast hydrolysate. Our findings suggest that addition of yeast hydrolysate into the pre-release diet of Q-flies could be a cost-effective means of releasing a bisexual strain with competitive males but with sexually immature females, thereby rendering it operationally more similar to a unisexual strain.

Adult Queensland fruit flies, Bactrocera tryoni (Froggatt), require adequate post-teneral nutrition to complete reproductive development, to perform sexually and for maximum longevity. Recent research has focussed on nutritional requirements of adults released in sterile insect technique (SIT) programmes that are used to manage these pests. Several studies have suggested benefits of providing yeast hydrolysate (YH) in addition to sugar during the 24- to 48-h pre-release holding period. Current evidence suggests that provision of YH can induce faster development, increased mating probability, longer copulations, increased sperm storage by mates, higher levels of sexual inhibition in mated females and increased longevity. We here review research on adult B. tryoni nutritional requirements, assess the potential application of this information in the context of SIT programmes and highlight future research that will help to determine whether YH, or other supplements, should be included as pre-release treatment in B. tryoni SIT.

Diet has a profound influence on the fitness of adult tephritid flies. Mass-reared flies are provided yeast hydrolysate as a rich source of nutrition that supports rapid sexual development and mating success. In contrast, wild tephritid flies often live in environments where food may be hard to find, and these are the conditions that sexually immature mass-reared sterile males encounter when released into the field during sterile insect technique campaigns. The effect of natural food sources (bat guano, bird droppings, citrus pollen, and wheat pollen) on the sexual development of adult mass-reared fertile, mass-reared sterile, and wild male Queensland fruit flies, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), was determined by measuring ejaculatory apodeme size. Inclusion of yeast hydrolysate in the adult diet was associated with faster growth of the ejaculatory apodeme in comparison with all other diets. Effects of diet were far less pronounced in mass-reared males, which may indicate reduced nutritional requirements, whereas the ejaculatory apodeme of wild males fed on natural sources of food or sucrose alone did not increase in size over the first 20 days of adult life.

Desiccation resistance is important for the survival of adult insects, although this key physiological trait has rarely been studied in tephritid flies. In the present study, desiccation resistance of female and male adult Queensland fruit flies Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) is determined with respect to age after adult eclosion. Resistance to acute starvation is measured over the same period to disentangle the competing roles of water loss and food deprivation. Survival of adult B. tryoni subjected to conditions of low humidity and starvation is reduced considerably compared with adults that are subjected to starvation alone. Desiccation resistance of adult female B. tryoni is generally lower than that of adult males. Desiccation resistance of adult B. tryoni declines in a continuous and regular manner over the first 20 days after adult eclosion. The regular pattern of declining resistance to desiccation with age in B. tryoni indicates that this reduction is not associated with the onset of maturity and maintenance of reproductive structures, nor with sexual activity. By contrast, resistance to starvation is similar at 0 and 6 days after adult eclosion, and declines thereafter. Survival under starvation and water stress is not related to wing length, which is a standard measure of fly size.

The distribution, systematics and ecology of Bactrocera tryoni, the Queensland fruit fly, are reviewed. Bactrocera tryoni is a member of the B. tryoni complex of species, which currently includes four named species, viz. B. tryoni ssp., B. neohumeralis, B. melas and B. aquilonis. The species status of B. melas and B. aquilonis is unclear (they may be junior synonyms of B. tryoni) and their validity, or otherwise, needs to be confirmed as a matter of urgency. While Queensland fruit fly is regarded as a tropical species, it cannot be assumed that its distribution will spread further south under climate change scenarios. Increasing aridity and hot dry summers, as well as more complex, indirect interactions resulting from elevated CO2, make predicting the future distribution and abundance of B. tryoni difficult. The ecology of B. tryoni is reviewed with respect to current control approaches (with the exception of sterile insect technique (SIT) which is covered in a companion paper). We conclude that there are major gaps in the knowledge required to implement most noninsecticide-based management approaches. Priority areas for future research include host–plant interactions, protein and cue-lure foraging and use, spatial dynamics, development of new monitoring tools, investigating the use of natural enemies and better integration of fruit flies into general horticultural IPM systems.