Feeding behavior ofRhagoletis pomonella flies (Diptera: Tephritidae): Effect of initial food quantity and quality on food foraging,handling costs,and bubbling

Under seminatural conditions feeding and postfeeding behaviors of individual apple maggot flies, Rhagoletis pomonella(Diptera: Tephritidae), were recorded after flies were presented with yeast hydrolysate or sucrose droplets, varying in either concentration, amount of food solute, or total droplet volume. The objectives were (a) to establish, at a constant level of previous food deprivation, food ingestion thresholds in relation to food quality and quantity and (b) to study the effect of initial food quantity and quality on food handling time and subsequent food foraging behavior. For both carbohydrate and protein substrates, fly foraging time after feeding on a tree branchlet was positively related to total amount of food solute previously encountered on a leaf surface, though largely independent of food volume or concentration. The volume and concentration of food presented, however, significantly affected food handling and processing time and therefore foraging time. In fact, total branchlet residence time was more closely linked to food handling and processing time than to foraging time. Less time was needed for uptake of liquid than dry food, the latter requiring liquification by salivary secretion and eliciting considerable intermittent cleaning of mouthparts by feeding flies. Similar to the situation in other fluid feeders, uptake time in R. pomonelladecreased with increasing dilution, although below a threshold of a 30% concentration of solute, the rate of nutrient intake decreased rapidly. When the level of dilution and total volume of food ingested were great enough, engorged flies entered extended quiescent postfeeding periods during which they extrude orally droplets of liquid crop contents (bubbling). After this they reinitiated feeding, followed by more bubbling and feeding bouts. Multivariate logistic regression analysis suggested that bubbling behavior is determined by liquid food volume and degree of dilution, hunger, and temperature. Although thresholds triggering bubbling decreased with increasing temperature, higher temperature by itself did not result in bubbling behavior. This suggests that bubbling is not primarily a mechanism to achieve evaporative cooling as has been suggested but, rather, a behavior to eliminate excess water, thereby enabling engorged flies to continue feeding on diluted food sources.     

Post-feed buzzing in the tsetse, Glossina morsitans morsitans, is an endothermic mechanism

ABSTRACT. Post-feed buzzing in Glossina morsitans morsitans Westw. causes a rise in thoracic temperature relative to the length of the buzz. As lift is proportional to the square of wing-beat frequency, which increases with temperature up to 32°C, buzzing results in an increase in the lift which the fly can produce. Heat generated by buzzing, in combination with the heat received from the host at the time of feeding, may well allow the fly to maximize lift generated in the immediate post-feeding period. Buzzing flies excrete excess water from the meal more rapidly than non-buzzing flies. It is argued that this is due to a rise in abdominal temperature. Maximized lift in the immediate post-feeding period and the rapid elimination of water from the very large blood meals taken by these flies may be expected to have strong selective advantages for the flies.     

Behavioural responses of dairy cattle to the stable fly, Stomoxys calcitrans, in an open field environment

Individual cows (25 in each of four herds) were monitored 8-10 times weekly for 12 weeks (stable fly season) on a southern California dairy, with 100 observations per cow. The numbers of biting stable flies, Stomoxys calcitrans (L.) (Diptera: Muscidae) on the front legs and the frequencies of four fly-repelling behaviours per 2-min observation period [head throws, front leg stamps, skin twitches (panniculus reflex) and tail flicks] were recorded. Fly numbers varied, peaking at 3.0-3.5 flies per leg in week 9 (late May). Weekly herd mean frequencies of fly-repelling behaviours were highly dependent on fly numbers, with a linear regression r(2) > 0.8. Head throws and stamps were less frequent than skin twitches and tail flicks. Individual cows differed in numbers of stable flies and behaviours. Behaviours were correlated with flies for individual cows, but at a lower level than were herd means (r = 0.3-0.7). Cows that stamped more within a herd tended to have lower fly counts; other fly-repelling behaviours were less effective. Cows maintained ranks within a herd with regard to fly numbers (r = 0.47), head throws (0.48), leg stamps (0.64), skin twitches (0.69) and tail flicks (0.64). Older cows tended to harbour higher fly numbers and to stamp less relative to younger adult cows. Ratios of leg stamps and head throws to fly numbers dropped significantly through time, suggesting habituation to pain associated with fly biting. Tail flicks were not effective for repelling Stomoxys, but were easiest to quantify and may help in monitoring pest intensity. At this low-moderate fly pressure, no consistent impacts on milk yield were detected, but methods incorporating cow behaviour are recommended for future studies of economic impact.     

Behavioural responses of stable flies to cattle manure slurry associated odourants

Stable flies (Stomoxys calcitrans [Diptera: Muscidae] L.) are blood‐feeding synanthropic pests, which cause significant economic losses in livestock. Stable fly antennae contain olfactory sensilla responsive to host and host environment‐associated odours. Field observation indicated that the abundance of stable flies increased significantly in grasslands or crop fields when cattle manure slurry was applied. Major volatile compounds emanating from manure slurry were collected and identified. Behavioural responses of stable flies to those compounds were investigated in laboratory bioassays and field‐trapping studies. Results from olfactometer assays revealed that phenol, p‐cresol and m‐cresol were attractive to adult stable flies. When tested individually, attraction was higher with lower dosages. Stable flies were most attracted to blends of phenol and m‐cresol or p‐cresol. Traps with binary blend lures caught more stable flies in field trials as well.     

The role of the frontal ganglion and corpora cardiaca/corpora allata complex in post-feeding weight loss in adult Heliothis zea

Abstract A post-feeding diuretic response has been observed in adult Heliothis zea (Boddie) (Lepidoptera: Noctuidae). 2-day-old starved adults which were allowed to feed to repletion on a 10% (w/v) sucrose solution lost 49.1% (females) and 85.8% (males) of the weight of the ingested meal during the first hour following feeding. Ligation between head and thorax or frontal ganglionectomy, when performed immediately following feeding, each resulted in a significant and permanent reduction in this normal weight-loss. Injection of homogenates of the corpora cardiaca/corpora allata (CC/CA) complex into non-ligated insects immediately after feeding also reduced significantly the post-feeding weight-loss, but this inhibition was transient and disappeared after 1 h. Dissection and weighing of the crop from either ligated, frontal ganglionectomized, or CC/CA-injected insects confirmed the crop as the predominant site of fluid retention in each case. We suggest that a soluble antidiuretic factor from the CC/CA acts in conjunction with the frontal ganglion to control the rate of crop emptying and subsequent diuresis by regulating the volume of ingested fluid that is passed into the haemolymph from the crop/midgut.     

Flies on vacation: evidence for the migration of Australian Syrphidae (Diptera)

1. Hover flies (Syrphidae: Diptera) are a cosmopolitan group of insects that provide important ecosystem services including pollination and pest control. The seasonal migration of hover flies is probably best known in Europe, but it remains unstudied in many other parts of the world. 2. Australia is believed to be home to around 160 hover fly species, some of which are common in urban and agricultural environments. The current evidence for hover fly migration in Australia is scarce and anecdotal, yet migration may be critical to the success of pollination and the biological control of aphids. 3. In this study, species occurrence records from an online biodiversity database (Atlas of Living Australia) were used to look for evidence of migratory behaviours in all Australian hover flies with more than 200 occurrence records. 4. Four of the 10 species displayed seasonal changes in their distribution consistent with migration, including Australia's two most abundant species: Melangyna viridiceps and Simosyrphus grandicornis. This work is an important first step in understanding the prevalence of migration in Australian hover flies. However, confirmation of our findings requires additional evidence to rule out other plausible explanations for the observed patterns. 5. Based on changes in summer and winter latitudinal distribution, it is estimated that some Australian hover flies may make annual migrations of 400–1800 km. 6. This work suggests that the management of beneficial insects requires consideration of factors at both local and continental scales, as landscape use changes may have an impact on ecosystem services delivered hundreds of kilometres away.     

Aggressive behaviours of female Drosophila melanogaster are influenced by their social experience and food resources

Abstract Aggressive behaviours of female Drosophila melanogaster (Meigen) (Diptera: Drosophilidae) were studied in the laboratory. These behaviours included 'approach', a seemingly intentional movement of a fly to another fly, 'lunge', a dash at another fly, and 'wings erect', a brief erection of wings toward another fly. These behaviours were influenced by social experience and food resources. First, event frequencies of aggressive behaviours were higher in flies that had been held in isolation than in those that had been held in crowded conditions. This difference was the result of the social experience of attacking flies and not that of attacked flies. Second, flies showed aggressive behaviours more frequently when they were on food containing yeast colonies than when they were on food without yeast colonies. Third, flies conditioned to a food solution containing live yeast prior to observation showed lower frequency of aggressive behaviours than flies conditioned to a boiled food solution. Development of aggressive behaviours became apparent one day after eclosion.     

Tolerance and antibiosis resistance to cabbage root fly in vegetable Brassica species

Four accessions of the wild species Brassica fruticulosa Cyrillo (Brassicaceae) were studied in order to identify its tolerance and antibiosis resistance to the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae), in comparison to a widely cultivated cauliflower cultivar and a rapid cycling Brassica oleracea L. line. Antibiosis was prominent, as the insects reared on resistant accessions showed reduced individual pupal weight, total pupal weight, adult dry weight, and the longest average fly eclosion time. Host plant resistance, however, did not affect the sex ratio of adult flies. A study of the root architecture of plants with and without root fly inoculation revealed differences in the structure within B. oleracea accessions. A long main root and a high number of lateral roots appeared to be important characteristics for a Brassica type, with a higher tolerance level to cabbage root fly attack.     

X-ray diffraction analysis of particles ingested by fieter-feeding animals

The size and relative abundance of mineral particles ingested by two filter-feeding aquatic insects, Simulium vittatum Zett. (Diptera: Simuliidae) and Aedes triseriatus (Say) (Diptera: Culicidae) were determined by X-ray diffraction methods. Different minerals representing different particle size categories were supplied larval black flies and mosquitoes. Since minerals possess characteristic diffraction properties, their presence and relative abundance can be determined. Early instars of the black fly ingested and retained particles of three different size ranges; however, more coarse particles relative to fine particles were retained by smaller instars as compared with the larger instars. With mosquito larvae, there was a proportional increase in coarser material ingested with increasing age and size of larvae. Other applications of this method are discussed.

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Résumé La dimension et l'abondance relative des particules minérates, ingerées par deux larves d'insectes aquatiques se nourrissant par filtration, Simulium vittatum (Diptera: Simuliidae) et Aedes triseriatus (Diptera: Culicidae) ont été déterminées par diffraction aux rayons X. Divers minéraux sous forme de particules de différentes dimensions ont été fournis aux larves de Simulies et de moustiques. Puisque les minéraux sont caractérisés par leurs propriétés de diffraction, on peut donc vérifier leur présence et déterminer leur relative abondance. Les larves de Simulies des premiers stades retiennent et ingèrent des particules de trois rangs de taille, cependant les particules plus grossières sont mieux retenues dans l'appareil filtreur des jeunes larves, par comparaison avec les larves plus grandes. Avec les larves de moustiques il y a une augmentation de l'ingestion du matériel le plus grossier proportionnellement à l'accroissement d'âge et de taille des larves. D'autres applications de cette méthode sont discutées.

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This research was partially supported by National Science Foundation Grant DEB76-20122, Black Fly Regional Research Project NE-118, and the Michigan State University Agricultural Experiment Station, Journal Article No. 8318.     

Pasture dragging fails to reliably suppress the emergence of horn flies (Haematobia irritans) and face flies (Musca autumnalis) from dung pats in a Mid-Atlantic North American climate

The beef industry endures major economic losses from a complex of flies that feed on bovine blood and mucus. For cattle on pasture, the most important of these pests are horn flies (Haematobia irritans [L.] [Diptera: Muscidae]) and face flies (Musca autumnalis [Diptera: Muscidae] De Geer). Pasture dragging to spread manure pats has been promoted as a management tactic for these species because their larvae inhabit bovine manure pats, but the efficacy of this practice has not been empirically validated. Spreading pats might promote fly mortality through desiccation or overheating, but these processes are weather-dependent and warrant testing in disparate climates. We evaluated pasture dragging effects while monitoring for weather interactions throughout nine experiment rounds in summers of 2018 and 2020 in Pennsylvania, USA. The manure spreading treatments increased pat surface area up to 300% but failed to significantly reduce emergence of horn flies and face flies as compared to controls. In contrast, precipitation and temperature were significant predictors in fly emergence models. Surprisingly, face fly emergence was significantly elevated in dragged pats twice in 2020. These data call for a reevaluation of pasture dragging as a management technique for horn flies and face flies across a range of climates.     

Disturbance to a stream food web by a bacterial larvicide specific to black flies: feeding responses of predatory macroinvertebrates

1. A field study was conducted to: (i) assess feeding habit changes of two predatory stoneflies following the loss of larval black fly (Diptera: Simuliidae) prey from two streams; and (ii) determine the relative importance of black fly larvae as prey for these and other selected predatory benthic macroinvertebrates. 2. Acroneuria lycorias and Paragnetina media (Plecoptera: Perlidae) diets were monitored in response to local reductions in larval black fly populations caused by Bacillus thuringiensis var. israelensis (B.t.i.) in two Michigan streams. These predators were collected from B.t.i.-treated and control sections of the streams, and their foreguts inspected for prey. 3. Black flies were the major dietary component of both predators collected from the control sections, but the number of black flies ingested was significantly less for predators collected from B.t.i.-treated habitats. Total number of prey ingested significantly decreased for A. lycorias, but not for P. media, and non-black fly prey consumption significantly increased for P. media, but not for A, lycorias, following B.t.i. applications. 4. In prey choice trials conducted in experimental channels, A. lycorias and P. media showed no preference between prey types (black flies and mayflies). Body mass gain of individual A. lycorias nymphs was measured, and was similar for nymphs in black fly-rich and black fly-poor environments. Conversely, Isoperla signata and I. dicala (Plecoptera: Perlodidae) ingested significantly more Simulium vittatum (Diptera: Simuliidae) than Baetis flavistriga (Ephemeroptera: Baetidae) or Epeorus vitrea (Heptageniidae) prey. Boyeria vinosa (Odonata: Aeshnidae) ingested significantly more B. flavistriga than S. vittatum prey. 5. Reducing black fly densities in these streams, using B.t.i., indirectly and differentially affected predators. In black fly-poor environments, feeding habits of specialist predators were most affected, and generalist predators least affected because the latter consumed alternative prey. Predator—predator and predator-prey interactions, and prey community structure may be affected indirectly by disturbances such as B.t.i. applications by reducing food resources and forcing predation onto less preferred prey.     

The effect of diet on activity in male Calliphora vicina blowflies

ABSTRACT. The effect of diet on the activity of male Calliphora vicina R.D. (Diptera, Calliphoridae) was studied. Protein–fed males (PF) were significantly more active than males fed only on sugar and water (NPF); as a consequence they ingested more sugar. The phenomenon was probably due to PF insects spending more time in flight than NPF, as no significant differences were found between the wingbeat frequencies and flight speeds of tethered flies of the two treatments. Protein ingestion also had a physiological effect, as PF flies had heavier thoraces than NPF flies with increased titres of protein.     

Estimation of the effects of buffalo fly (Haematobia irritans exigua) on the milk production of dairy cattle based on a meta-analysis of literature data

Published reports on the effect of buffalo fly Haematobia irritans exigua De Meijere (Diptera: Muscidae) and the closely related horn fly (H. irritans) were examined and analysed using non-linear weighted regression techniques in an attempt to establish the relationship between daily production loss (D), average number of parasites (n) and the average damage per parasite per day (d), and to provide estimates of expected losses in milk yield (MYD) and live-weight gain (LWG) in dairy cattle. A Mitscherlich three-parameter model was used to explain the relationship between the total loss of production attributable to buffalo flies and the average number of flies associated with cattle. This model was significant (P<0.01), with R2 = 20.2% and predicted a threshold number of flies (n = 30) below which no adverse effects would be noted. At a moderate level of infestation (n = 200) dMYD was 2.6 ml/fly/day and dLWG was 0.14 g/fly/day, resulting in estimated daily losses in milk yield (D(MYD)) and live-weight gain (D(LWG)) of 520 ml and 28 g, respectively.     

Pest management through tropical tree conservation

Tropical trees can provide various ecological services to adjacent agricultural environments, including maintaining and amplifying the numbers of beneficial insects. In Mexico, certain tree species harbor a diverse guild of hymenopteran parasitoids that attack pest fruit flies (Diptera: Tephritidae) and are at the same time sources of valuable hardwood timber. Indigenous trees and their associated fauna are slowly disappearing due to forest clearance and the expansion of crop monocultures. Here we explore the relationship among pest and non-pest fruit flies, their fruit-hosts and parasitoids in the context of mango orchards and surrounding patches of uncultivated vegetation and propose a novel mechanism to use these associations in favor of conservation purposes and pest management. Trees of conservation biological control interest are classified as: (1) parasitoid multiplier plants, species that serve as alternate hosts for key fruit fly pests when their commercial hosts are not available, but in which they are unusually vulnerable to parasitism; (2) parasitoid reservoir plants, native or introduced trees in whose fruits non-pest fruit flies serve as hosts to generalist parasitoids that are able to attack pest tephritids in other species of commercially grown fruit; and (3) pest-based parasitoid reservoir plants, native or introduced species that are not economically important locally, but which harbor fruit flies that would be pests in other circumstances and that serve as hosts for parasitoids of the important pests in the vicinity. Protection, multiplication and dissemination of such tree species has the potential to increase the number of naturally produced fruit fly parasitoids and could assist in the management of tephritid pests in areas where destruction of forests has impoverished the historical sources of fruit fly natural enemies. Tropical forest conservation may help resource-poor farmers reduce crop losses, increase biodiversity within fruit-growing regions and conserve native forests for both conservation purposes and commercial use of native hardwoods.     

Desiccation resistance of adult Queensland fruit flies Bactrocera tryoni decreases with age

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 phylogeny of a mutualism: evolution and coadaptation between Trollius and its seed-parasitic pollinators

Interactions between seed-parasitic pollinators and their hosts provide useful model systems for the analysis of evolution of mutualism and potential coevolution between plants and insects. Here I present the systematics, pollination ecology and evolution of one of these interactions. I have documented and analysed the phylogenetic and geographic associations between Trollius (Ranunculaceae: 18 spp.) and Chiastocheta (Diptera: Anthomyiidae; 17 spp.), a host-specific genus of seed-parasitic flies that pollinate their host plants to varying extent. Their interactions are usually facultative mutualisms, but in the specialized T. europaeus three fly species are obligate mutualists and a fourth species is an antagonist. The distribution patterns of fly species among Trollius species suggest that the flies evolved in associations with five highly derived Trollius species, and secondarily colonized four more primitive taxa in the parts of their ranges that overlapped with primary hosts. In general, host specificity is maintained primarily through allopatry, with colonization occurring in regions of overlap between parapatric taxa. Fly speciation has occurred in allopatry, both within and among host taxa. Cospeciation is not evident, but convergent evolution in Trollius flowers of several traits, viz. orange sepals, elongated staminodia and increased carpel number per flower, may be the result of mutualism with Chiastocheta.     

Accessibility of blood affects the attractiveness of cattle to horn flies

The burden of infestation of the horn fly, Haematobia irritans (Linnaeus) (Diptera: Muscidae), differs among bovines within the same herd. We hypothesized that these differences might be related to the epidermal thickness of the cattle and the blood intake capacity of the fly. Results showed that dark animals carried more flies and had a thinner epidermis than light‐coloured animals, which was consistent with the greater haemoglobin content found in flies caught on darker cattle. Similarly, epidermal thickness increased with body weight, whereas haemoglobin content decreased. Overall, we suggest that accessibility of blood is a factor that partially explains cattle attractiveness to flies.     

Nepetalactones from essential oil of Nepeta cataria represent a stable fly feeding and oviposition repellent

The stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae), is one of the most serious pests to livestock. It feeds mainly on cattle and causes significant economic losses in the cattle industry. Standard stable fly control involving insecticides and sanitation is usually costly and often has limited effectiveness. As we continue to evaluate and develop safer fly control strategies, the present study reports on the effectiveness of catnip (Nepeta cataria L.) oil and its constituent compounds, nepetalactones, as stable fly repellents. The essential oil of catnip reduced the feeding of stable flies by >96% in an in vitro bioassay system, compared with other sesquiterpene-rich plant oils (e.g. amyris and sandalwood). Catnip oil demonstrated strong repellency against stable flies relative to other chemicals for repelling biting insects, including isolongifolenone, 2-methylpiperidinyl-3-cyclohexen-1-carboxamide and (1S,2'S)-2-methylpiperidinyl-3-cyclohexen-1-carboxamide. The repellency against stable flies of the most commonly used mosquito repellent, DEET, was relatively low. In field trials, two formulations of catnip oil provided >95% protection and were effective for up to 6 h when tested on cattle. Catnip oil also acted as a strong oviposition repellent and reduced gravid stable fly oviposition by 98%.