Means of the formation of gonotrophic relations in blood-sucking Diptera

Gonotrophic relations in low and higher bloodsucking Diptera are fundamentally different that results from an initial type of feeding and trophic behaviour. Trophic behaviour of low dipterous hematophages and gonotrophic harmony peculiar to them can be traced from entomophagy, that is from the predatory mode of life on account of small insects. By the type of the trophic behaviour low bloodsucking Diptera are predators with a typical moment contact with the prey. More primitive is a type of gonotrophic harmony characteristic of hunters for diffusely spread prey (incomplete blood portion provides the maturation of incomplete portion of eggs). Hunting for diffusely spread prey is characteristic of entomophages too. The appearance of gregarious ruminants facilitates the possibility of repeated contacts with prey and blood satiation threshold increases. This is a higher type of gonotrophic harmony providing a maximum realization of potential fecundity. The initial saprophagy of higher Diptera is associated with another type of trophic behaviour (long contact with food substratum) that is a prerequisite for quite a different way of evolution of host-parasite relationships in higher Diptera. This leads to more close connections with the host and excludes gonotrophic harmony. Females were the first to begin the exploitation of vertebrate animals. This is connected with the peculiarities of their behaviour during egg laying such as the stay near animals for laying eggs into fresh dung. Autogeneity, nectarophagy and aphagia are homologous phenomena which reflect the loss of an animal component of food or both components at the level of non-specialized saprophagy rather than secondary loss of bloodsucking. The scheme of gonotrophic relations is given.     

Endophagy of biting midges attacking cavity-nesting birds

Feeding behaviour, host preferences and the spectrum of available hosts determine the role of vectors in pathogen transmission. Feeding preferences of blood-feeding Diptera depend on, among others factors, the willingness of flies to attack their hosts either in the open (exophagy) or in enclosed places (endophagy). As far as ornithophilic blood-feeding Diptera are concerned, the biting midges (Diptera: Ceratopogonidae) and blackflies (Diptera: Simuliidae) are generally considered to be strictly exophagous. We determined which blood-sucking Diptera enter nest cavities and feed on birds by placing sticky foil traps inside artificial nest boxes. A total of 667 females of eight species of biting midges of the genus Culicoides (Latreille, 1809) were captured on traps during 2006-2007, with Culicoides truncorum (Edwards, 1939) being the dominant species. DNA blood analyses of blood-engorged females proved that midges actually fed on birds nesting in the boxes. Three species were identified as endophagous: Culicoides truncorum, Culicoides pictipennis (Staeger, 1839), and Culicoides minutissimus (Zetterstedt, 1855). Our study represents the first evidence that ornithophilic biting midges are endophagous. The fact that we caught no blackflies in the bird boxes supports the exophagy of blackflies. We believe that our findings are important for surveillance programmes focusing on Diptera that transmit various bird pathogens.     

A comparison of the feeding behaviour of tsetse and stable flies

In Zimbabwe, observations were made of the behaviour of individual stable flies (Stomoxys spp.) (Diptera: Muscidae) and tsetse (Glossina spp.) (Diptera: Glossinidae) feeding on cattle during the wet (Stomoxys and tsetse) and dry (tsetse only) seasons. For Stomoxys landing on adult cattle, only 27% took a full meal (mean feeding time = 147 s). Most Stomoxys left the host before completing their meal, largely due to disturbance by the host's defensive behaviour (24%, mean time = 59 s) or other flies (44%, 71 s). The probability of a Stomoxys leaving the host progressively increased with time. Simultaneous observations of tsetse showed that, compared to Stomoxys, their feeding success was lower (15%), feeding was interrupted earlier (33 s) and the time taken to complete a meal was shorter (109 s). Further studies of tsetse across different seasons and hosts showed that feeding success varied according to host age (adult = 7%; calf = 3%) and was negatively correlated with the frequency of host defensive behaviour and the relative abundance of non-biting Diptera. Disturbances were more often caused by host behaviour (69%) than other flies (31%) and the probability of tsetse leaving decreased with time on the host. Overall, these results suggest that tsetse and Stomoxys have different feeding strategies. In particular, tsetse appear to be more responsive to host defensive behaviour, which reduces their feeding success relative to Stomoxys. These behavioural differences are consistent with the respective life-history characteristics of Stomoxys and tsetse.     

Courtship behaviour in the sandfly Lutzomyia longipalpis, the New World vector of visceral leishmaniasis

Courtship behaviour in Lutzomyia longipalpis Lutz and Neiva (Diptera: Psychodidae) was examined to determine the sequence of behaviours that occur prior to copulation. Courtship consisted of a series of male and female touching and wing-flapping behaviours, with males performing a greater variety of wing-flapping behaviours than previously described. Occurrence of male approach-flapping, semi-circling and female stationary-flapping were all predictors of eventual copulation, and may coincide with the dispersal of pheromones or the production of auditory signals important to courtship. Touching occurred in the majority of observations, with contact most often made with the tips of the legs and antennae. This behaviour, not previously described in sandflies, was initiated by males and females, and may indicate the use of contact pheromones, a form of communication previously overlooked in L. longipalpis. Future studies are required to separate the auditory and chemical signals associated with wing-flapping, and to confirm whether L. longipalpis possesses cuticular hydrocarbons capable of inducing behavioural responses. The identification of signals that inhibit sexual behaviour during courtship may be particularly relevant to developing mating disruption techniques against L. longipalpis.     

Candidate sex pheromones of the New World screwworm Cochliomyia hominivorax

Five novel homologous acetate derivatives of long-chain secondary alcohols and a related ketone were tested for their efficacy as contact mating stimulants for Cochliomyia hominivorax Coquerel (Diptera: Calliphoridae). Full copulatory behaviour at a high percentage was found in tests with racemic 6-acetoxy-19-methylnonacosane at 2.5-20 microg using fertile males from three strains. Males of two strains responded nearly as well to 7-acetoxy-15-methylnonacosane, but an older strain first colonized in 1992 did not respond to this compound. Few or no copulatory responses were obtained to the other secondary alcohol acetates and a related ketone. These two acetate derivatives are the first sex pheromones identified in a calliphorid fly. The threshold of response was also tested, but could not be pinpointed.     

Hunger‐dependent and sex‐specific antipredator behaviour of larvae of a size‐dimorphic mosquito

1. Modification of behaviours in the presence of predators or predation cues is widespread among animals. The costs of a behavioural change in the presence of predators or predation cues depend on fitness effects of lost feeding opportunities and, especially when organisms are sexually dimorphic in size or timing of maturation, these costs are expected to differ between the sexes. 2. Larval Aedes triseriatus (Say) (Diptera: Culicidae) were used to test the hypothesis that behavioural responses of the sexes to predation cues have been selected differently due to different energy demands. 3. Even in the absence of water‐borne predation cues, hungry females (the larger sex) spent more time browsing than did males, indicating a difference in energy needs. 4. In the presence of predation cues, well‐fed larvae of both sexes reduced their activity more than did hungry larvae, and males shifted away from high‐risk behaviours to a greater degree than did females, providing the first evidence of sex‐specific antipredator behaviour in foraging mosquito larvae. 5. Because sexual size dimorphism is common across taxa, and energetic demands are probably correlated with size dimorphism, this research demonstrates the importance of investigating sex‐specific behaviour and behavioural responses to enemies, and cautions against generalising results between sexes.     

Blood-feeding in mosquitoes: probing time and salivary gland anti-haemostatic activities in representatives of three genera (Aedes, Anopheles, Culex)

Mosquitoes (Diptera: Culicidae) face their hosts' haemostatic mechanisms when attempting to feed on blood. Accordingly, they antagonize haemostasis by salivary agents that include anti-clotting, anti-platelet and vasodilatory compounds. Because haemostasis is a complex and redundant physiological response that varies between vertebrates, it is to be expected that haematophagous animals have a salivary armoury that most efficiently counteracts their preferred hosts. The mosquito Culex quinquefasciatus Say, which has a strong tendency to ornithophagy, appears to have only recently adapted to mammals and may not have evolved efficient mechanisms to counteract mammalian platelet responses, while birds only have relatively inefficient thrombocytes. Accordingly, we compared the probing behaviour of Cx. quinquefasciatus with two other mosquito species from different backgrounds: Aedes aegypti (L.) and Anopheles albimanus Weidemann, that have apparently had a longer evolutionary association with mammals. Culex takes much more time to find blood on a mammalian host (human or mouse) when compared to the two other mosquito species, but does not differ in probing behaviour when feeding on a chicken. Salivary anti-haemostatic components were also measured in those three species of mosquito and results are discussed in context with the probing behaviour.     

An order-specific monoclonal antibody to Diptera reveals the impact of alternative prey on spider feeding behavior in a complex food web

Generalist predators have the capacity to restrict pest population growth, especially early in the season before densities increase. However, their polyphagous feeding habits sometimes translate into reduced pest consumption when they target alternative prey. An order-specific monoclonal antibody was developed to examine the strength of trophic connections between Diptera, a major category of non-pest prey, and linyphiid spiders in alfalfa. We report the development and characterization of a monoclonal antibody with order-level specificity to Diptera. This antibody elicited strong absorbance to 22 Diptera from 13 families, no false-positive reactivity to non-dipteran invertebrates, and antigen detection periods following prey consumption that were comparable between spiders. Over 900 field-collected females of the linyphiid spiders Erigone autumnalis and Bathyphantes pallidus were screened for Diptera antigen. Significantly more B. pallidus screened positive for Diptera (40%) compared to E. autumnalis (16%), indicating differential reliance on these prey. In parallel with the collection of spiders for gut-content analysis, prey availability was estimated at web sites. The two spiders exhibited different feeding responses to prey availability. Consumption of Diptera by B. pallidus was strongly correlated with Diptera abundance whilst the availability of other potential prey did not influence predation rates. Conversely, E. autumnalis did not prey upon Diptera in proportion to availability, but increased Collembola activity-density reduced dipteran consumption. Integration of molecular gut-content analysis with precise sampling of prey demonstrated how two closely related linyphiid spiders exhibit different feeding responses to the availability of prey under natural field conditions. Elucidating the feeding preferences of natural enemies is critical to effective incorporation of biological control by generalist predators in the management of agricultural pests.     

Aggregated oviposition in the Simulium damnosum complex is mediated by eggs in a laboratory bioassay

Abstract. . The phenomenon of aggregated oviposition in blackflies (Diptera: Simuliidae) was investigated under controlled laboratory conditions, using wild-caught females of the Simulium damnosum complex in Sierra Leone. A method was developed for inducing Simulium females to lay eggs, and used as a bioassay to measure the responses of gravid S. damnosum s.l. to freshly laid eggs of the same species complex. In a series of two-choice tests, significantly more ovipositing flies chose substrates already containing eggs over control substrates ( P = 0.004). The time from introduction of flies into the oviposition system to the onset of egg-laying was significantly less when eggs were already present ( P = 0.049). Flies responded more quickly when more eggs were present and the relationship between egg-batch number and the time of this response was curvilinear ( P = 0.012). Ecological advantages and disadvantages of such aggregation behaviour and the possible role of semiochemicals in its mediation are discussed.     

Responses of male Mediterranean fruit flies, Ceratitis capitata, to trimediure in a wind tunnel of novel design

ABSTRACT. The responses of male Ceratitis capitata (Wiedemann) (Diptera; Tephritidae) to trimediure were investigated in a wind tunnel of novel design with disposable plastic walls. Trimediure was presented to the flies either in the form of a discrete 'plume' or uniformly dispersed in the air passing through the tunnel. The flies showed an increase in flight activity in response to the two forms of stimulation and performed a zig-zag anemotactic flight in response to the plume, reminiscent of the behaviour of moths responding to a sex pheromone. They performed a straight, upwind, anemotactic response to the uniformly dispersed odour. The design of wind tunnel used in these tests was found to have several advantages: air flow and turbulence can be easily varied and the plastic walls can be changed to eliminate contamination problems.     

Behavioural responses of the carrot fly larva, Psila rosae, to carrot root volatiles

ABSTRACT. The behaviour of the carrot fly larva, Psila rosae F. (Diptera, Psilidae), was investigated in the presence of six concentrations of thirty-four volatiles associated with carrot root and of trans -methyl-iso-eugenol, a constituent of carrot herb oil. Bomyl acetate, 2,4-dimethyl styrene, α-ionone, SbT-ionone and biphenyl were the most consistently preferred compounds. Klinotaxis and klinokinesis were the orientation responses near the stimulus source. Trans -2-nonenal was the most consistently avoided compound. Extracts derived by steam distillation of carrot roots and by concentration of vapour from roots were resolved, by gas liquid chromatography, into forty-five and twenty-two major constituents, respectively. The five preferred compounds are micro-components of these extracts, collectively representing 1% and 0.6% of root extract oil and vapour, respectively. It is suggested that these secondary plant constituents are specific host finding cues for the larva, and that frara-2-nonenal is a carrot defence compound.     

Extraordinary feeding behaviour in Diptera Sciomyzidae, snail-killing flies

In the laboratory, the larva of Sepedonella nana, Diptera Sciomyzidae, were only able to develop at the expense of small aquatic oligochets collected in adult habitats. Up till now, larval cycles have been elucidated for 38% of the Sciomyzidae; all these larvae proved to be strict consumers of aquatic or terrestrial molluscs (including slugs). This specific predation was considered as a major distinguishing characteristic of Sciomyzidae among Diptera. The unusual nutritional behaviour of S. nana reported in the present note is in contradiction with well-established prior knowledge. The species presented all the characteristics of the family, particularly the ventral arch sclerite present on the larva cephalopharyngeal skeleton. It was considered as a derived species; this behaviour, probably of secondary type, revealed new nutritional types for Afrotropical Sciomyzidae for which the biology of only five species was elucidated out of about 60 species described.     

The interaction of taste and heat on the biting response of the tsetse fly Glossina fuscipes fuscipes

Abstract.Tsetse flies probe more on a heated surface with a trace of uric acid than on one without. Uric acid is one of the components of human sweat and it elicits spike responses from taste hairs on the flies' legs. In this paper it is examined how heat from the surface and taste interact to affect the biting behaviour of Glossina fuscipes fuscipes Newstead 1910 (Diptera: Glossinidae) over successive days of food deprivation. The biting behaviour consists of bouts of probing, both ambulatory and stationary, intercalated with short hops of flight. The number of bouts increases over successive days, whereas the average bout duration does not. Although uric acid alone could not induce the flies to probe, in combination with surface heat it affected the flies greatly. Average bout duration was two‐fold that on a heated surface not treated with uric acid. The frequency of bouts was not affected by uric acid. These experiments and auxiliary ones on mechanoreceptive input and odours lead to the insight that the factors which affect biting behaviour can be viewed as a hierarchy. The hierarchy extends from those that affect the onset of biting to those that affect its course.     

Spatial feeding preferences of ornithophilic mosquitoes,blackflies and biting midges

The section of habitat used by particular bloodsucking insects when seeking bloodmeals may influence the spectrum of hosts to which they have access and consequently the diseases they transmit. The vertical distribution of ornithophilic bloodsucking Diptera (Culicidae, Simuliidae and Ceratopogonidae) was studied using bird‐baited traps set at both ground and tree canopy levels. In total, 1240 mosquito females of eight species, 1201 biting midge females of 11 species, and 218 blackfly females of two species were captured during 2003–2005. Culex pipiens (L.) (Diptera: Culicidae) was found to prefer ground‐level habitats, whereas Anopheles plumbeus (Stephens) (Diptera: Culicidae), biting midges [Culicoides spp. (Diptera: Ceratopogonidae)] and Eusimulium angustipes (Edwards) (Diptera: Simuliidae) preferred the canopy. The results of this study with regard to Cx. pipiens behaviour differ from those of most previous studies and may indicate different spatial feeding preferences in geographically separate populations. The occurrence of E. angustipes in the canopy is concordant with its role in the transmission of avian trypanosomes. These findings may be important for surveillance programmes focusing on ornithophilic Diptera which transmit various pathogenic agents.     

Description of the larva and pupa of Nasiternella regia Riedel, 1914 (Diptera: Pediciidae) from Slovakia, with notes on ecology and behaviour

The previously unknown larvae and pupae of Nasiternella regia Riedel, 1914 (Diptera, Pediciidae) are described and illustrated from specimens collected in water-filled tree holes in deciduous forests in Slovakia. Brief comments on their ecology and behaviour are provided. Comparisons are made to the larvae of Nasiternella varinervis (Zetterstedt, 1851) as described by Krivosheina (2009).     

A meta-analysis of predation risk effects on pollinator behaviour

Flower-visiting animals are constantly under predation risk when foraging and hence might be expected to evolve behavioural adaptations to avoid predators. We reviewed the available published and unpublished data to assess the overall effects of predators on pollinator behaviour and to examine sources of variation in these effects. The results of our meta-analysis showed that predation risk significantly decreased flower visitation rates (by 36%) and time spent on flowers (by 51%) by pollinators. The strength of the predator effects depended neither on predator taxa and foraging mode (sit-and-wait or active hunters) nor on pollinator lifestyle (social vs. solitary). However, predator effects differed among pollinator taxa: predator presence reduced flower visitation rates and time spent on flowers by Squamata, Lepidoptera and Hymenoptera, but not by Diptera. Furthermore, larger pollinators showed weaker responses to predation risk, probably because they are more difficult to capture. Presence of live crab spiders on flowers had weaker effects on pollinator behaviour than presence of dead or artificial crab spiders or other objects (e.g. dead bees, spheres), suggesting that predator crypsis may be effective to some extent. These results add to a growing consensus on the importance of considering both predator and pollinator characteristics from a community perspective.     

An alternative mating system in small male insects

Abstract. 1. Small males of all midge (Diptera: Chironomidae and Chaoboridae) species thus far examined (one chaoborid and six chironomids) are rare in mating swarms. They are found instead in vegetation adjacent to the swarm.
2. We show that it is here that females aggregate prior to embarking on mate acquisition flights. In the vegetation females appear to be accessible to males staying behind.
3. Such behaviour in small males may exploit the mate-attracting activities of large males in the swarm and may, at the same time, reduce competition and conserve energy.
4. The evolution and maintenance of these size-related types of mating behaviour are discussed.     

Parasitic effect of Opius concolor (Spzl) (Hymenoptera, Braconidae) on an intermediate host Ceratitis capitata Wied (Diptera, Trypetidae)

The parasitic incidence of Opius concolor (Hymenoptera, Braconidae) on a replacement host, Ceratitis capitata (Diptera, Trypetidae) was studied using polyacrylamide gel electrophoresis and immunoelectrophoresis methods. A noticeable modification of C. capitata proteins was observed when parasited by O. concolor. But the most important phenomenon is the finding of common antigens between the host and the parasite. These results are discussed with regard to trophic and parasitic behaviour of the parasite.     

A laboratory investigation of mosquito larval predation by Toxorhynchites moctezuma on Aedes aegypti

1. Functional responses of predatory Toxorhynchites moctezuma (Dyar & Knab) larvae feeding on Aedes aegypti (L.) larvae (Diptera: Culicidae) were found to be type II of Holling (1959) and Rogers (1972). 2. Estimates of searching rate were generally higher for later instar predators. The search rate of second instar predators declined as prey instar increased, but fourth instar Tx. moctezuma had the highest search rate for second instar Ae. aegypti. 3. Prey handling times were higher for early instar predators and late instar prey. 4. When presented with mixtures of two instars of Ae. aegypti, second instar Tx. moctezuma showed frequency independent selectivity for the early instars, whereas fourth instar predators showed frequency independent selectivity for the late instars of Ae. aegypti. There was no evidence of frequency dependent predation. Preferences appeared to be transitive. 5. Extended random predator equations, using parameters derived from the functional responses, did not adequately describe the outcome of predation in the prey mixture experiment, even when the possibility of optimal switching behaviour was accounted for.     

Host deprivation influence on response of Rhagoletis pomonella to its oviposition deterring pheromone

ABSTRACT. The response of host-depibved Rhagoletis pomonella Walsh (Diptera, Tephritidae) females to host fruit ( Crataegus viridis L.) marked with R.pomonella oviposition deterring pheromone was measured in the laboratory. A positive correlation was found between length of host deprivation (5, 10, 20, 40 or 80 min) and probability of host acceptance (i.e. oviposition) by flies. The results are discussed in relation to current theories on physiological control of insect feeding and oviposition behaviour.