Upwind flight of tsetse (Glossina spp.) in response to natural and synthetic host odour in the field

Abstract. In Zimbabwe, studies were made of the flight responses of tsetse ( Glossina spp.) to synthetic and natural ox odour using arrangements of electric nets.Tsetse flying away from a target showed a significant upwind bias when a blend of carbon dioxide (2/1 min), acetone (500 mg/h), octenol (0.4 mg/h), 4-methylphenol (0.8 mg/h) and 3-n-propylphenol (0.1 ma) was dispensed 15 m upwind, with c. 35% flying upwind.Without carbon dioxide this percentage was significantly reduced to 15% which was not significantly different from that with no odour (8%).This pattern was not altered by reducing the doses of acetone, octenol and phenols by 10–100 times, to levels comparable to those produced by an ox.With natural ox odour or a synthetic equivalent of ox odour dispensed from a ventilated pit 8 m upwind of the target, c. 28% flew upwind.This was reduced significantly to 15% if carbon dioxide was removed.In studies using a 17 m line of nets arranged orthogonally across the prevailing wind line, c. 50% of the catch was caught on the downwind side in the absence of odour.This increased significantly to c.60% when acetone, octenol and phenols were dispensed 15 m upwind, with or without carbon dioxide.With a shorter line (9 m) or an incomplete one (16.5 m long with 5 times 1.5 m wide gaps along its length) there was no change in the proportion caught downwind.For all three lines, dispensing odour upwind increased the catch 2–5 times on both the up-and downwind sides of the nets.It is concluded that a stronger upwind response to host odour is elicited when carbon dioxide is present.It is suggested that in nature upwind flight is very imprecisely orientated, with tsetse making flights up and down an odour plume 'searching' for a host.     

The flight and landing of tsetse (Glossina) in response to components of host odour in the field

ABSTRACT. Studies were conducted in Zimbabwe of the responses of Glossina morsitans morsitans Westwood and Glossina pallidipes Austen to various host odours using either arrangements of electrocuting nets or visual observations. Tsetse flying upwind in a plume of carbon dioxide, acetone and octenol turned downwind upon flying into a plume of acetone or octenol, but did not turn upon flying into a plume of carbon dioxide. They also turned in response to a transient decline in odour concentration. Tsetse landed on the ground in the vicinity of a source of natural odour or artificial odour containing carbon dioxide but not at sources of acetone or octenol only. The proportion of female G.pallidipes caught at a source of natural odour (37%) was significantly different from that caught at a source of synthetic odour (17%). Resting tsetse stimulated by natural odour took off sooner than non-stimulated flies and had a strong upwind bias in the direction of take off. Tsetse stimulated with artificial odour did not take off sooner than non-stimulated flies. It is suggested that there is an unidentified components) of ox odour that activates resting tsetse.     

Rates of progress up odour plumes by tsetse flies: a mark-release video study of the timing of odour source location by Glossina pallidipes

Abstract. The arrival of individually marked Glossina pallidipes Austen at a host odour source after their video-timed release from 30–75 m downwind was measured in the field in Zimbabwe. In the absence of odour, the proportion recaptured was <2% (= - random expectation); when synthetic ox odour was released, the probability of recapture at the source increased with proximity of release, from 6% at 75 m to 21% at 30 m (about twice this number arrived within ∼2 m of the source). There were two distinct distributions of recaptures: a 'fast' cohort which found the source within 40 s, and a 'slow' cohort which took from one to >20 min, with ∼50% of the flies in each cohort. The fastest flies probably reached the source in a single, mainly straight flight from take-off, at an overall average (straight line) displacement speed of 2.8-4.5 ms^-1 (i.e. close to the preferred flight speed of ∼5 m s^-1). The flies apparently maintained their ground speed largely independent of the wind speed they headed into. The 'slow' cohort had a constant probability of arrival at the source, presumably after losing and re-contacting the plume, and after having stopped at least once on the way. There were no marked correlations with wind parameters, although the probability of recapture increased slightly with the directness of the wind from the source, and the probability of 'slow' flight increased slightly with wind speed. It is inferred that a repeated sequence of anemotactic 'aim-then-shoot' orientation at take-off plus optomotor-steered in-flight correction of direction is used as a form of biassed random walk to bring the flies close to the odour source, rather than the use of moth-type anemotactic zigzagging.     

Electroantennogram responses of tsetse flies (Glossina pallidipes) to host odours in an open field and riverine woodland

The present study was initiated to gain insight into the way in which tsetse flies ( Glossina spp.) sense odours at different locations in odour plumes in both an open field and a wooded area.
We recorded the antennal responses (EAGs) from stationary living female G. pallidipes 15 m upwind and at various (60, 40, 20, 10, 5 and 1 m) distances downwind from a synthetic host odour source (containing 1-octen-3-ol, acetone and two phenols), in the natural habitat of the fly (Zimbabwe) using a portable electrophysiological device. Experiments were performed in a flat open area (an airstrip) and in riverine woodland. Differences between responses in different environments were determined by comparing various parameters of the EAGs (intermittency, frequency, amplitude, duration and rate of depolarization).
We found that a fly senses odours as puffs that, further downwind, contain less odour and pass less frequently. In an open field downwind from the source, tsetse perceive more olfactory information than upwind for only 10–20 m, whereas in woodland, olfactory responses remain higher and more frequent than upwind up to at least 60 m. In an open field, olfactory information rapidly increases when approaching the odour source from 20 m and in woodland from 5 m onwards.
It is proposed that averaging odour information over time may be of minor importance in long-range location of odour sources. The results suggest that tsetse may smell odour-baited targets from at least 60 m downwind and that the number of flies responding to and being caught by these baits may be higher in woodland than in an open field.     

The effects of host physiology on the attraction of tsetse (Diptera: Glossinidae) and Stomoxys (Diptera: Muscidae) to cattle

In Zimbabwe, studies were made of the numbers of tsetse (Glossina spp.) and stable flies (Stomoxys spp.) attracted to cattle of different nutritional status, age and sex. Host odours were analysed to determine the physiological basis of these differences and improved methods are described for measuring rates of production of kairomones. Seasonal fluctuations in host weight, related to changes in pasture quality, had no significant effect on attraction of tsetse or Stomoxys. However, both attraction to different individuals and carbon dioxide production by these individuals were strongly correlated with weight, suggesting a possible link. Attraction to the odour from different types of cattle decreased in the order ox>cow>heifer>calf, and oxen were twice as attractive as calves of less than 12 months old. Lactation did not alter the relative attractiveness of cows. Calves less than six months old produced lower levels of carbon dioxide, acetone, octenol and phenols than oxen, but for older calves and cows, levels of production of known kairomones and repellents were similar to those of an ox. Carbon dioxide produced by cattle varied according to time of day and the animal's weight; cattle weighing 500 kg produced carbon dioxide at a mean rate of 2.0 l min(-1) in the morning and 2.8 l min(-1) in the afternoon compared to respective rates of 1.1 and 1.9 l min(-1) for cattle weighing 250 kg. Artificially adjusting the doses of carbon dioxide produced by individual cattle to make them equivalent did not remove significant differences in attractiveness for tsetse but did for Stomoxys. Increasing the dose of carbon dioxide from 1 to 4 l min(-1) in a synthetic blend of identified kairomones simulating those produced by a single ox, increased attractiveness to tsetse but not to the level of an ox. The results suggest that the main sources of differences in the attractiveness of individual cattle are likely to be variation in the production of carbon dioxide and, for tsetse, other unidentified kairomone(s). The biological and practical implications of these findings are discussed.     

The activation of resting tsetse flies (Glossina) in response to visual and olfactory stimuli in the field

ABSTRACT Studies were conducted in Zimbabwe of the responses of Glossina morsitans morsitans Westwood and Glossina pallidipes Austen resting in a refuge to various host stimuli. Tsetse took off in response to 100% ox odour, 0.08% carbon dioxide or a visual stimulus consisting of a 0.75 × 0.75 m black target placed c . 5 m from the refuge moving at 4^o s^-1, but the level of response was low with only 35%, 19% and 29% responding, respectively. Tsetse did not take off in response to any one of 25% ox odour, 0.8% carbon dioxide, acetone (3 μg 1^-1) or octenol (0.03 μg 1^-1). In the absence of any host stimuli, flies emerged from the refuge later on hotter days (35–37^oC) than on cooler days (32–34.5^oC). Male G.pallidipes emerging later in the afternoon contained significantly more haematin than those emerging relatively earlier. There were no significant differences between the responses of G.m. morsitans and G.pallidipes. It is suggested that the initial activation of resting flies is primarily mediated through endogenous, rather than host, stimuli.     

Distant olfactory response of the cabbage seed weevil, Ceutorhynchus assimilis, to oilseed rape odour in the field

Abstract. The olfactory orientation of adult cabbage seed weevils (Ceutorhynchus assimilis Payk.) from a distance of 20 m to extracts of oilseed rape (Brassica napus L.) was studied in the field using mark–release–recapture experiments. Male and female weevils dispersed upwind from the centre of a circular array of traps baited with Oilseed rape extracts. In the absence of host-plant odour, female weevils dispersed randomly, and male weevils downwind. Percentage recapture of weevils was significantly greater in traps baited with rape flower than rape leaf extracts. Significantly fewer weevils were recaptured in unbaited traps compared to traps baited with rape flower or leaf extracts. We suggest that seed weevils can use odour-mediated upwind anemotaxis to locate their host-plants from distances of at least 20 m.     

Dispersal of phlebotomine sand flies (Diptera: Psychodidae) in a Colombian focus of Leishmania (Viannia) braziliensis.

Dispersal of five species of phlebotomine sand flies was studied in a coffee plantation near Arboledas, Colombia, by mark-release-recapture studies using fluorescent powders. The estimated recapture rate for males of Lutzomyia shannoni marked and released during the day was 28.1%, significantly higher than that for all other species (p < 0.05). Recapture rate of female Lu. shannoni was 9.5%, and no females of the other four species were recovered. This suggests either that Lu. shannoni is a more sedentary species than the others, or that the large trees on which these insects were captured and recaptured function as foci of lekking behaviour in males. The high recapture rates of females of this species may indicate that oviposition occurs in close proximity to the bases of these trees. Although most marked sand flies were recaptured within 200 m of their release point, a single female Lu. gomezi was recovered 960 m away 36 h after release. This suggests that the dispersal capacity of Lutzomyia species may be greater than has been thought, an important consideration in future control programs directed against these insects in Leishmania-endemic areas.     

Effects of seawater-acclimatization and release sites on survival of hatchery–reared brown trout Salmo trutta

To test whether seawater–acclimatization of hatchery–reared anadromous and freshwater resident brown trout before release increased the survival of adults, smolts were retained 0, 2, 4 and 8 weeks in sea water before release. Total recapture rate increased for smolts retained 4 and 8 weeks in sea water before release relative to the controls. This trend was more pronounced for Freshwater resident than for anadromous stocks, Offspring of anadromous fish stayed longer at sea than offspring of freshwater resident fish. Recapture rates in fresh water were higher for brown trout released in the river than in the fjord in the R, Drammen area, but not in the R. Imsa. In both cases, most fish were recaptured in the sea. Moving into the R. Imsa (relative to other rivers) appeared higher for fish released at the mouth of the river (93%) than in the fjord (47%). Judged from the recapture rates, sea survival appeared to be the same whether released in the fjord or at the mouth of the river.     

Flight responses of tsetse flies (Glossina) to octenol and acetone vapour in a wind-tunnel

Abstract. Female Glossina morsitans morsitans Westwood were video-recorded in a wind-tunnel as they entered, in crosswind flight, a broad plume of either octenol or acetone (two components of ox odour). Both odours produced upwind turning responses (in-flight anemotaxis) to a range of concentrations, with thresholds at around 10^-8mg1^-l for octenol and 10^-6mg1^-1 for acetone. Kinetic responses were unaffected by octenol at low concentrations, but flight speed was significantly reduced and sinuosity (^om^-1) and angular velocity (^os^-1) significantly increased by concentrations at or above those in ox breath; for acetone, these effects were apparent but inconsistently related to concentration. It is concluded that octenol and acetone vapour are used by tsetse flies to locate hosts by upwind anemotaxis, probably combined with kinetic responses. The behavioural basis for the 'repellency' of high octenol concentrations in the field is discussed in the context of the virtual loss of upwind anemotaxis to octenol at the highest concentration tested in the tunnel (30 × ox breath).     

Observations on the orientation of tsetse flies (Glossina pallidipes) to wind-borne odours

ABSTRACT. Observations of the upwind flight of Glossina pallidipes Austen near a source of host odour show that in the absence of a visual target the insects tend to overshoot the odour source in fast, low flight. There is no sign of the crosswind 'casting' flight which characterizes the behaviour of moths under similar circumstances, except that a 180 turn is executed to bring the tsetse flies back to the vicinity of the odour source in downwind flight. This may be followed by a second overshoot and another 180 turn before the insects alight within a metre or so of the source. The results indicate that the orientation of tsetse flies to host odour may involve a step-wise approach to the odour source, providing an opportunity for assessment of wind direction when the insects are at rest between successive bursts of flight.     

Dose responses of tsetse flies (Glossina) to carbon dioxide, acetone and octenol in the field

ABSTRACT Studies were conducted in Zimbabwe of the catch of Glossina pallidipes Austen from an electric net plus target baited with mixtures of acetone plus carbon dioxide or 1-octen-3-ol (octenol) plus carbon dioxide. For acetone dispensed alone at 5–50, 000 mg h^-1, ten-fold increments in the dose increased the catch 1.7 times. For carbon dioxide dispensed alone, dose increments from 12 to 1201 h^-1 doubled the catch, but the catch was not further increased by dispensing carbon dioxide at 600–1200 1 h^-1. For mixtures of these two odours, ten-fold increments in the dose of carbon dioxide between 12 and 12, 0001 h^-1 increased the catch c . 2.5 times if acetone was also dispensed at >50 mg h^-1; changes in the dose of acetone between 50 and 50 000 mg h^-1 did not affect the catch. The addition of octenol (0.05 mg h^-1) to carbon dioxide (12–12001 h^-1) doubled the catch. Ten-fold increments in the dose of octenol between 0.05 and 5 mg h^-1 did not increase the catch significantly and the catch was independent of changes in the dose of carbon dioxide between 120 and 12001 h^-1. The behavioural basis of the dose-response curves was investigated using an incomplete ring of electric nets to assess the flight orientation of tsetse in different odours. Upwind flight was not elicited by acetone or octenol alone, or by carbon dioxide unless it was at very high doses, however, mixtures of carbon dioxide with acetone or octenol elicited upwind flight. It is suggested that the attractiveness of mixtures of acetone and carbon dioxide is a function of the region of overlap of these two odours at above threshold concentration. Acetone and octenol on their own appear to increase the responsiveness of flies to visual cues.     

Orientation of tsetse flies to wind, within and outside host odour plumes in the field

Abstract Free-flying wild tsetse flies ( Glossina pallidipes Aust. and G. m. morsitans Westw.) were video recorded in Zimbabwe as they flew within an artificial host odour plume at 3, 7 or 15 m from the source, or in no odour, with and without a 0.75 m² vertical, black visual target present aligned with the wind. With no visual target present, flights in odour were strongly biased upwind, and in the absence of odour strongly biased downwind. With the target present, between 16% and 40% of the upwind approaching flies responded visually as they passed the target, by circling it, in proportion to the proximity of the source (taken to be proportional to the mean odour concentration). Crosswind approaching flies (for whom the target will have been visible for some metres away) circled more frequently (34–56%), but without obvious correlation with the odour concentration. Circling flies also responded orthokinetically, by slowing down as they passed the target. The departure directions relative to the wind of flies leaving the target were significantly affected by the odour concentration. At 3 m they left the target in all directions, except possibly avoiding due upwind. At 7 m they left with an obliquely upwind bias, but at 15 m and also in no odour, they left with a strong crosswind bias.     

Distant olfactory response of the onion fly, Delia antiqua, to host-plant odour in the field

ABSTRACT. Distant olfactory orientation of female adult Delia antiqua (Meigen) to the host-plant Allium volatile dipropyl-disulphide (DPDS) was examined in the field using mark-release-recapture experiments and observations of flight behaviour. Onion-reared, post-diapause, virgin females from a laboratory colony dispersed upwind when released in the centre of 25, 50 and 100 m radius circles of eight 50 μl UDPDS baits. Percentage recapture and dispersal directedness did not decrease as a function of increasing distance to baits. In all cases the mean flight direction of recaptured flies closely correlated with mean wind direction. However, modes of the circular distributions of recaptured flies were located further crosswind when odour-baits were more distant. When distance was held constant (25 m) and DPDS concentration serially reduced (500–0.05 μ/bait), flies dispersed randomly in the absence of DPDS, crosswind in response to 0.05 μl baits and upwind in response to all other baits. Percentage recaptures on DPDS-baited traps of all concentrations were significantly greater than unbaited traps. Results from markrecapture studies were corroborated by observations of flight behaviour downwind. Flies located 100 m downwind from 50 μl DPDS baits flew upwind on take off while take-off flights in the absence of DPDS were random. Our data indicate that Allium volatiles like DPDS are involved not only in the acceptance phase of host-selection, but also in the first and probably most important stage when onion flies are initiating search long distances downwind. We conclude that D. antiqua orients to host-plants using olfactory cues from distances that should be classified as long-range ( sensu Kennedy , 1977     

Persistence of sterile screwworm (Diptera: Calliphoridae) flies at a release site.

Studies to determine the persistence of released, sterile screwworms, Cochliomyia hominivorax (Coquerel), were conducted in Belize, Central America, in 1987-1988. A total of nine releases were made, each consisting of 4,000 females marked with a fluorescent dust. Previous similar studies with baited fly traps indicated that flies died or dispersed from the release site within 3-4 d. For this study, flies were recaptured at sentinel animals. Recapture rates varied greatly between releases, a result that was not correlated with any weather parameter. The pattern of recapture indicated a sharp drop in the released population after 9-10 d, with a few flies persisting in the release area for up to 3 wk. The results suggest that survival of released sterile flies is comparable with estimated survival rate and life span of wild, native flies.     

基于荧光标记的怒江流域桔小实蝇 (Bactrocera dorsalis)的迁移扩散

2005年7月在云南怒江流域地区,以荧光粉作为标记物,采用“标记-释放-回收”方法,对怒江流域桔小实蝇迁移扩散规律进行了研究。以释放点潞江坝为中心,东南西北4个方位设立桔小实蝇回收点,经过7d的回捕,从释放点沿潞江以北至六库沿线共回收到的标记桔小实蝇43头,最远在距释放点以北97km的地方回收到标记桔小实蝇5头。沿怒江在释放点以南,仅在小于29km的范围内回收到标记桔小实蝇17头。释放点东面和西面未能回收到标记桔小实蝇。分析释放点以北各回收点标记桔小实蝇与非标记桔小实蝇的相互关系揭示,非标记桔小实蝇与标记桔小实蝇在空间动态中具有相同的行为模式,由此推论非标记桔小实蝇与标记桔小实蝇可能来自同一种群。分析怒江流域区的自然地理发现,潞江坝桔小实蝇可借助怒江两边高大山脉形成的天然河谷通道,在适宜的气候条件下,在北上气流的携带下,实现由南向北的远距离迁移扩散;同时,由于高大山脉的阻隔,使潞江坝桔小实蝇没能向东西两个方向作远距离扩散迁移。研究首次揭示了在特定峡谷地区桔小实蝇扩散迁移现象,为桔小实蝇迁移扩散研究提供了新鲜实例,为在当地制定切实有效的桔小实蝇防治策略提供了基础信息。     

Visually-guided, upwind turning behaviour of free-flying tsetse flies in odour-laden wind: a wind-tunnel study

Abstract. To test the hypothesis that tsetse flies use visual input from the apparent movement of the ground to assess wind direction while in flight, Glossina morsitans morsitans Westwood females were video- recorded in a wind-tunnel as they entered, in cross-wind flight, a broad plume of simulated host odour (C0₂ at c. 0.05%). The tunnel (2.3 times 1.2 m wide) generated winds up to 0.25 m s^-1 and had a strongly patterned floor that could be moved upwind or downwind to increase or decrease the visual input due to wind drift. Flight tracks were analysed for speed, direction relative to the wind, and angle of turn. Mean groundspeeds were c. 1.8 m s^-1. In control measurements in still air (with or without odour) flies turned 50:50 'upwind': 'downwind'. With a 0.25 m s^-1 odour-perme- ated wind, 79% turned upwind, and c. 70% left view flying upwind. When the floor was moved at 0.25 m s^-1 upwind (to mimic the visual input from the ground due to a 0.5 m s_^-1 wind), the strength of this response increased. If instead the floor was moved downwind, faster than the wind speed (to mimic the visual input due to a wind from the opposite direction), 59% turned downwind and c. 70% left view flying downwind, and thus away from the source (though progressing 'upwind' in terms of the visual input from apparent ground pattern movement). Upwind turns were on average significantly larger than downwind turns. It is concluded that tsetse navigate up host odour plumes in flight by responding to the visual flow fields due to their movement over the ground (optomotor anemotaxis), even in weak winds blowing at a fraction of their groundspeed.     

Dispersal pattern of the sand fly Lutzomyia neivai (Diptera: Psychodidae) in a cutaneous leishmaniasis endemic rural area in Southeastern Brazil

The dispersal pattern of the sand fly Lutzomyia neivai was studied through mark-release-recapture experiments in an American cutaneous leishmaniasis endemic rural area in Southeastern Brazil. Over 6500 specimens were marked with fluorescent powder and released in forest edge and peridomicile habitats from August to November 1999, February and April 2000. Recapture attempts were made using Shannon and CDC traps up to eight successive nights after releases. A total of 493 (7.58%) specimens were recaptured. The number of recaptured males and females of L. neivai in CDC traps was not affected by the distance between the trap and the release points. Approximately 90% of males and females recaptured in CDC traps were caught up to 70 m from the release points. The maximum female flight range recorded was 128 m. The average flight range per day was less than 60 m for males and females. Of the flies released in forest edge, approximately 16% of the recaptured females were caught in Shannon traps in the peridomicile habitat. The results indicate that the movements of L. neivai are spatially focal and the possibility of dispersion from forest to peridomicile habitat may be an important way of contracting leishmaniasis in dwellings.     

基于荧光标记的怒江流域桔小实蝇（Bactrocera dorsalis）的迁移扩散

2005年7月在云南怒江流域地区,以荧光粉作为标记物,采用“标记-释放-回收”方法,对怒江流域桔小实蝇迁移扩散规律进行了研究。以释放点潞江坝为中心,东南西北4个方位设立桔小实蝇回收点,经过7d的回捕,从释放点沿潞江以北至六库沿线共回收到的标记桔小实蝇43头,最远在距释放点以北97 km 的地方回收到标记桔小实蝇5头。沿怒江在释放点以南,仅在小于29 km 的范围内回收到标记桔小实蝇17头。释放点东面和西面未能回收到标记桔小实蝇。分析释放点以北各回收点标记桔小实蝇与非标记桔小实蝇的相互关系揭示,非标记桔小实蝇与标记桔小实蝇在空间动态中具有相同的行为模式,由此推论非标记桔小实蝇与标记桔小实蝇可能来自同一种群。分析怒江流域区的自然地理发现,潞江坝桔小实蝇可借助怒江两边高大山脉形成的天然河谷通道,在适宜的气候条件下,在北上气流的携带下,实现由南向北的远距离迁移扩散;同时,由于高大山脉的阻隔,使潞江坝桔小实蝇没能向东西两个方向作远距离扩散迁移。研究首次揭示了在特定峡谷地区桔小实蝇扩散迁移现象,为桔小实蝇迁移扩散研究提供了新鲜实例,为在当地制定切实有效的桔小实蝇防治策略提供了基础信息。     

Development of a low-cost tsetse trap and odour baits for Glossina pallidipes and G.longipennis in Kenya

Experiments were carried out to improve the NG2B tsetse trap (Brightwell et al., 1987), baited with acetone and cow urine, for use by rural communities to control G.pallidipes Austen and G.longipennis Corti. Modifications included a lower dose rate of acetone, a new cage design and raising the trap about 15-20 cm. Research on different trap cone materials showed that the degree of light transmission of the netting, rather than its colour, was the crucial factor affecting the catch of G.pallidipes. Adding an additional metre of blue cloth to one side of the trap increased catches of females of both species by about 60%. Traps baited with synthetic phenols yielded similar numbers of G.pallidipes and significantly more G.longipennis than those baited with natural cow urine. The latter difference was not apparent when octenol was also used, so cow urine was retained as one of the odour baits in preference to the imported phenols. Although octenol increased catches of G.pallidipes by only about 30%, catches of G.longipennis were increased 2-4-fold, making it a very useful attractant for the latter species. The cost of the trap/odour-bait system was estimated to be US$8.5 per unit per annum. The economics of this method of tsetse control are discussed.