Mapping tsetse habitat suitability in the common fly belt of Southern Africa using multivariate analysis of climate and remotely sensed vegetation data

Abstract The distribution of Glossina morsitans centralis, Glossina morsitans morsitans and Glossina pallidipes are described in part of southern Africa, using a range of multivariate techniques applied to climate and remotely sensed vegetation data. Linear discriminant analysis is limited in its predictive power by the assumption of common co-variances in the classes within multivariate environment space. Maximum likelihood classification is one of a variety of alternative methods that do not have this constraint, and produce a better prediction, particularly when a priori probabilities of presence and absence are taken into account. The best predictions are obtained when the habitat is subdivided, prior to classification, on the basis of a bimodality detected on the third component axis of a principal component analysis. The results of the predictions were good, particularly for G.m.centralis and G.m.morsitans , which gave overall correct predictions of 92.8% and 85.1 %, with a Kappa index of agreement between the predion and the training data of 0.7305 and 0.641 respectively. For G.pallidipes , 91.7% of predictions were correct but the value of Kappa was only 0.549. Very clear differences are demonstrated between the habitats of the two subspecies Gmxentralis and G.m.morsitans.     

Comparative study on the susceptibility of different laboratory strains of Glossina species to Trypanosoma simiae

Abstract. Teneral tsetse of four Glossina species from laboratory-reared colonies were fed on four Large White pigs infected with three different stocks of Trypanosoma simiae isolated in Coast Province, Kenya. Thereafter the tsetse were maintained on goats and dissected on day 28 to determine the trypanosome infection rates. Glossina brevipalpis was as susceptible as G.pallidipes whilst G.palpalis gambiensis was not susceptible to T.simiae CP 11 a stock causing acute infection, which was isolated from a wild G.austeni. Glossina brevipalpis was as susceptible as G.pallidipes to another stock causing acute infection, T.simiae CP 813 isolated from a wild G.pallidipes. Glossina morsitans centralis was also as susceptible as G.brevipalpis and G.pallidipes whilst G.p.gambiensis was not susceptible to this T.simiae stock. Glossina m.centralis showed very low susceptibility to a stock causing chronic infection, T.simiae CP 1896 isolated from a bushpig, whilst G.brevipalpis, G.p.gambiensis and G.pallidipes could not be infected by this T.simiae stock. Male Glossina were generally more susceptible than females to the three T.simiae stocks.     

The effects of temperature, body mass and feeding on metabolic rate in the tsetse fly Glossina morsitans centralis

Abstract.  Metabolic rate variation with temperature, body mass, gender and feeding status is documented for Glossina morsitans centralis . Metabolic rate [mean ± SE; VCO₂= 19.78 ± 3.11 μL CO₂ h⁻¹ in males (mean mass = 22.72 ± 1.41 mg) and 27.34 ± 3.86 μL CO₂ h⁻¹ in females (mean mass = 29.28 ± 1.96 mg) at 24 °C in fasted individuals] is strongly influenced by temperature, body mass and feeding status, but not by gender once the effects of body mass have been accounted for. A significant interaction between gender and feeding status is seen, similar to patterns of metabolic rate variation documented in Glossina morsitans morsitans . Synthesis of metabolic rate-temperature relationships in G. m. centralis , G. m. morsitans and Glossina pallidipes indicate that biting frequency as well as mortality risks associated with foraging will probably increase with temperature as a consequence of increasing metabolic demands, although there is little evidence for variation among species at present. Furthermore, metabolic rate–body mass relationships appear to be similarly invariant among these species. These data provide important physiological information for bottom-up modelling of tsetse fly population dynamics.     

Genetic Variation at Structural Loci in the Glossina morsitans Species Group

Gene diversity was investigated in four taxa of tsetse flies (Diptera: Glossinidae) including Glossina morsitans morsitans, G. m. centralis, G. swynnertoni, and G. pallidipes. Histochemical tests were performed for 35–46 isozymes. Polymorphic loci were 20% in G. morsitans morsitans, 32% in G. m. centralis, 17.6% in G. swynnertoni, and 26% in G. pallidipes. Mean heterozygosities among all loci were 6.6% in G. morsitans morsitans, 6.0% in G. m. centralis, 7.1% in G. swynnertoni, and 6.8% in G. pallidipes. Allozyme gene diversities were considerably less than those reported for many Diptera. The low gene diversities are probably related to small effective population sizes.     

A comparison of African buffalo, N''Dama and Boran cattle as reservoirs of Trypanosoma congolense for different Glossina species

Teneral Glossina morsitans centralis Machado were fed on the flanks of the African buffalo (Syncerus caffer Sparrman), N'Dama (Bos taurus L.) or Boran (Bos indicus L.) cattle infected with Trypanosoma congolense Broden. The infected tsetse were maintained on rabbits and on day 30 after the infected feed, the surviving tsetse were dissected to determine the infection rates. The mean infection rates (% +/- SE) in the midgut of tsetse fed on buffalo, N'Damas and Borans were 23.5 +/- 3.3, 31.6 +/- 2.7 and 33.7 +/- 4.6, respectively. The differences were not significant. However, the mean mature infection rate in tsetse fed on the buffalo (13.2 +/- 2.1%) was significantly lower compared to the rates in tsetse fed on the N'Dama (20.4 +/- 1.4) or the Boran cattle (21.4 +/- 1.1). When groups of teneral G.m.centralis, G.pallidipes Austen, G.p.gambiensis Vanderplank, G.f.fuscipes Newstead, G.brevipalpis Newstead and G.longipennis Corti were fed simultaneously on either an infected buffalo, an N'Dama or a Boran steer, the mature infection rates ranged from 0 to 16.1%. Irrespective of the host species used, the T.congolense infection rate was highest in G.m.centralis, lowest in the palpalis and fusca group tsetse, with G.pallidipes being intermediate. Nevertheless, the trypanoresistant African buffalo and N'Dama may serve as reservoirs of T.congolense as can trypanosusceptible Boran cattle.     

Behaviour of tsetse flies (Glossina) in host odour plumes in the field

ABSTRACT. In Zimbabwe, studies were made of the responses of Glossina pallidipes Austen and G.morsitans morsitans Westwood to artificial host odour using an incomplete ring of electrocuting nets. In a plume of synthetic host odour tsetse flew generally upwind, with 50–60% flying within 35^o of due upwind. More than 80% of tsetse flew at < 50 cm above ground level. Upon losing contact with odour they executed a reverse turn within about 2 m, and upon regaining contact they turned upwind. There were no clear differences in the responses of G.m.morsitans and G.pallidipes. Using electrocuting nets lying horizontally on the ground it was found that tsetse landed in the vicinity of the odour source, the propensity to land being greater for G.pallidipes than for G.m, morsitans , greater for immature than mature flies, and greater for males than females.     

Feeding behaviour of Glossina pallidipes and g. morsitans centralis on Boran cattle infected with trypanosoma congolense or T. vivax under laboratory conditions

In field studies, tsetse flies (Diptera: Glossinidae) feed more successfully on cattle infected with Trypanosoma congolense Broden (Kinetoplastida: Trypanosomatidae) than on cattle infected with T. vivax Ziemann or uninfected cattle. Here we describe the first laboratory investigation of this phenomenon. In the first experiment, caged Glossina pallidipes Austen were fed for 1 and 5 min on a Boran steer infected with T. congolense clone IL 1180 and on an uninfected steer. Feeding success was recorded in this way five times over several weeks. The same protocol was subsequently used in three additional experiments with the following combinations: G. pallidipes and a steer infected with T. vivax stock IL 3913, G. morsitans centralis Machado and a steer infected with T. congolense, and G. morsitans centralis and a steer infected with T. vivax. The four experiments were replicated once, making eight experiments in total. In three experiments there was increased tsetse feeding success, measured at 1 min, after a steer became infected (T. congolense, two experiments and T. vivax, one experiment). Analysis of all data combined found no significant differences in tsetse feeding success on the different groups of cattle prior to infection, but after infection tsetse feeding success was significantly greater on the infected cattle (P< 0.001). Trypanosoma congolense infection led to a greater increase in tsetse feeding success than T. vivax infection. The increase in feeding success was not related to changes in the level of anaemia, skin surface temperature or parasitaemia. A possible explanation is the effects of trypanosome infection on cutaneous vasodilation and/or blood clotting in infected cattle. When allowed to feed for 5 min, nearly all tsetse engorged successfully and effects of cattle infection on feeding success were not found.     

Comparative study on the infection rates of different laboratory strains of Glossina species by Trypanosoma congolense

Teneral Glossina morsitans centralis Machado, G.austeni Newstead, G.palpalis palpalis Robineau-Desvoidy, G.p.gambiensis Vanderplank, G.fuscipes fuscipes Newstead, G.tachinoides Westwood and G.brevipalpis Newstead, from laboratory-bred colonies, were fed at the same time on the flanks of ten goats infected with Trypanosoma congolense Broden isolated in Tanzania or in Nigeria. The seven tsetse species were infected over the range 0.3-49.2%. Survival of both T.congolense isolates was best in G.m.centralis, poorest in G.austeni and the four palpalis group tsetse, with G.brevipalpis intermediate. It is suggested that there are differences in the gut of different laboratory-bred cultures of Glossina Westwood species and subspecies such that T.congolense parasites can survive better in the gut of some than in others and undergo cyclical development to metacyclics in the hypopharynx.     

Responses of tsetse flies, Glossina morsitans morsitans and Glossina pallidipes, to baits of various size

Recent studies of Palpalis group tsetse [Glossina fuscipes fuscipes (Diptera: Glossinidae) in Kenya] suggest that small (0.25 × 0.25 m) insecticide-treated targets will be more cost-effective than the larger (≥1.0 × 1.0 m) designs currently used to control tsetse. Studies were undertaken in Zimbabwe to assess whether small targets are also more cost-effective for the Morsitans group tsetse, Glossina morsitans morsitans and Glossina pallidipes. Numbers of tsetse contacting targets of 0.25 × 0.25 m or 1.0 × 1.0 m, respectively, were estimated using arrangements of electrocuting grids which killed or stunned tsetse as they contacted the target. Catches of G. pallidipes and G. m. morsitans at small (0.25 × 0.25 m) targets were, respectively, ～1% and ～6% of catches at large (1.0 × 1.0 m) targets. Hence, the tsetse killed per unit area of target was greater for the larger than the smaller target, suggesting that small targets are not cost-effective for use against Morsitans group species. The results suggest that there is a fundamental difference in the host-orientated behaviour of Morsitans and Palpalis group tsetse and that the former are more responsive to host odours, whereas the latter seem highly responsive to visual stimuli.     

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.     

Detection and characterization of <Emphasis Type="Italic">Wolbachia</Emphasis> infections in laboratory and natural populations of different species of tsetse flies (genus <Emphasis Type="Italic">Glossina</Emphasis>)

BACKGROUND: Wolbachia is a genus of endosymbiotic α-Proteobacteria infecting a wide range of arthropods and filarial nematodes. Wolbachia is able to induce reproductive abnormalities such as cytoplasmic incompatibility (CI), thelytokous parthenogenesis, feminization and male killing, thus affecting biology, ecology and evolution of its hosts. The bacterial group has prompted research regarding its potential for the control of agricultural and medical disease vectors, including Glossina spp., which transmits African trypanosomes, the causative agents of sleeping sickness in humans and nagana in animals. RESULTS: In the present study, we employed a Wolbachia specific 16S rRNA PCR assay to investigate the presence of Wolbachia in six different laboratory stocks as well as in natural populations of nine different Glossina species originating from 10 African countries. Wolbachia was prevalent in Glossina morsitans morsitans, G. morsitans centralis and G. austeni populations. It was also detected in G. brevipalpis, and, for the first time, in G. pallidipes and G. palpalis gambiensis. On the other hand, Wolbachia was not found in G. p. palpalis, G. fuscipes fuscipes and G. tachinoides. Wolbachia infections of different laboratory and natural populations of Glossina species were characterized using 16S rRNA, the wsp (Wolbachia Surface Protein) gene and MLST (Multi Locus Sequence Typing) gene markers. This analysis led to the detection of horizontal gene transfer events, in which Wobachia genes were inserted into the tsetse flies fly nuclear genome. CONCLUSIONS: Wolbachia infections were detected in both laboratory and natural populations of several different Glossina species. The characterization of these Wolbachia strains promises to lead to a deeper insight in tsetse flies-Wolbachia interactions, which is essential for the development and use of Wolbachia-based biological control methods.     

Effect of puparia incubation temperature: increased infection rates of Trypanosoma congolense in Glossina morsitans centralis, G.fuscipes fuscipes and G.brevipalpis

Puparia of Glossina morsitans centralis (Machado), G.fuscipes fuscipes (Newstead) and G.brevipalpis (Newstead) were incubated at 25 +/- 1 degrees C, 28 +/- 1:25 +/- 1 degrees C, day:night or 29 +/- 1 degrees C throughout the puparial period, and maintained at 70-80% relative humidity. Puparial mortality was higher at 29 than at 25 degrees C (optimum temperature) in all three species, particularly in G.f.fuscipes and G.brevipalpis. Adults of G.m.centralis from puparia incubated at 29 degrees C, and those of this subspecies, G.f.fuscipes and G.brevipalpis from puparia incubated at 28:25 degrees C, day:night or 25 degrees C throughout, were infected as tenerals (27 h old) by feeding them at the same time on goats infected with Trypanosoma congolense (Broden) IL 1180 after the parasites were detected in the wet blood film. Infection rates on day 25 post-infected feed were higher in G.m.centralis from puparia incubated at 29 degrees C and in adults of the three different tsetse species from puparia incubated at 28:25 degrees C, day:night, than in those from puparia incubated at 25 degrees C. However, in G.f.fuscipes the labral and hypopharyngeal infection rates were not significantly different from those of the tsetse produced by puparia kept at 25 degrees C.     

Relationships between host blood factors and proteases in Glossina morsitans subspecies infected with Trypanosoma congolense

Abstract. Host blood effects on Trypanosoma congolense establishment in Glossina morsitans morsitans and Glossina morsitans centralis were investigated using goat, rabbit, cow and rhinoceros blood. Meals containing goat erythrocytes facilitated infection in G. m. morsitans , whereas meals containing goat plasma facilitated infection in G. m. centralis. Goat blood effects were not observed in the presence of complementary rabbit blood components. N-acetyl-glucosamine (a midguMectin inhibitor) increased infection rates in some, but not all, blood manipulations. Cholesterol increased infection rates in G. m. centralis only. Both compounds together added to cow blood produced superinfection in G. m. centralis , but not in G. m. morsitans. Midgut protease levels did not differ 6 days post-infection in flies maintaining infections versus flies clearing infections. Protease levels were weakly correlated with patterns of infection, but only in G. m. morsitans. These results suggest that physiological mechanisms responsible for variation in infection rates are only superficially similar in these closely-related tsetse.     

Tsetse mating behaviour: effects of age and hunger in Glossina morsitans morsitans and G.pallidipes

ABSTRACT. The effects of age and hunger on the responses of male Glossina morsitans morsitans Westwood and G.pallidipes Austen to freeze-killed female decoys, were examined in the laboratory. In both species, activity, estimated as the total number of interactions between males and decoys, increased with both age and hunger. Interactions were divided into short-stay (<60 s) and long-stay, full copulatory responses. In both species, young, unfed males were significantly less likely to attempt to copulate with a decoy after encounter than were fed males. Among fed males the proportion of interactions that proceeded to full copulatory attempts did not change with increasing age, but decreased consistently with increasing hunger. At all ages and hunger levels, G.pallidipes were more active than G.m.morsitans. However, after encountering a decoy, G.pallidipes were less likely to attempt to copulate than G.m.morsitans. In both species the duration of copulatory attempts did not change with age, but declined with increasing hunger. Copulatory attempts by G.pallidipes were significantly shorter than those of G.m.morsitans. The results are discussed in relation to the behaviour of tsetse in response to control devices such as traps and targets.     

Optimized simulation of the control of tsetse flies Glossina pallidipes and G. m. morsitans (Diptera: Glossinidae) using odour-baited targets in Zimbabwe

In 1984-1985 insecticide-treated targets were deployed in the 600-km2 Rifa Triangle, Zambezi Valley, Zimbabwe. Trap catches of Glossina pallidipes Austen were modelled using a function combining logistic growth with diffusive movement. A simulation routine was linked to a non-linear least-squares optimization programme and fits optimized with respect to population carrying capacities, rates of growth and movement, and to levels of imposed mortality. In March-September 1984, the overall additional mortality was 2% per day of adult female G. pallidipes, increasing thereafter to 8% per day, due to the deployment of more targets, the onset of the hot, dry season and the ground-spraying of the adjoining Zambezi escarpment with DDT. For G. m. morsitans Westwood the corresponding estimates were 1 and 2% per day. For both species, the deployment of four targets km(-2) in a closed population will ensure eradication. For G. m. morsitans a halving of target efficacy would reduce the killing rate to the point where eradication would be unlikely. Estimated daily displacements were c. 200 m for G. m. morsitans and 660 m for G. pallidipes. The lower rate for G. m. morsitans means that, while targets kill this species less effectively, re-invasion of cleared areas is slower. Targets do not markedly affect robust populations outside the deployment area. The Zambian tsetse population adjacent to the Rifa Triangle declined markedly during the experiment, however, suggesting that it is largely maintained by immigration. The methods developed here will be applied to data from other campaigns with the aim of improving the efficiency of tsetse control programmes.     

Analysis of the mating activity of male tsetse flies Glossina m. morsitans and G. pallidipes in the laboratory

ABSTRACT. The responses of male Glossina morsitans morsitans West-wood and Glossina pallidipes Austen to freeze-killed females were examined in the laboratory. Analyses were performed using a specially designed, automated, computer-based, recording system. G. pallidipes were more active than G. m. morsitans , interacting with the female decoys twice as often. Interactions with the decoys divided broadly into short-stay (<60 s) and long-stay, full copulatory attempts. For G. m. morsitans 90% of interactions resulted in full copulatory attempts, the mean duration of which was >1 h. For G. pallidipes only 40% of interactions resulted in full copulatory attempts, the mean duration of which was 35 min. The initiation of interactions showed a clear V-shaped activity pattern in G. m. morsitans but in G. pallidipes only a morning peak was observed. In neither species was there a tendency for full copulatory responses to be initiated in any specific period of the diurnal activity pattern. The results indicate that the two species have very different mating systems, and represent an initial step in the quantification of these differences.     

Less is more: restricted application of insecticide to cattle to improve the cost and efficacy of tsetse control

Studies were carried out in Zimbabwe of the responses of tsetse to cattle treated with deltamethrin applied to the parts of the body where most tsetse were shown to land. Large proportions of Glossina pallidipes Austen (Diptera: Glossinidae) landed on the belly ( approximately 25%) and legs ( approximately 70%), particularly the front legs ( approximately 50%). Substantial proportions of Glossina morsitans morsitans Westwood landed on the legs ( approximately 50%) and belly (25%), with the remainder landing on the torso, particularly the flanks ( approximately 15%). Studies were made of the knockdown rate of wild, female G. pallidipes exposed to cattle treated with a 1% pour-on or 0.005% suspension concentrate of deltamethrin applied to the (a) whole body, (b) belly and legs, (c) legs, (d) front legs, (e) middle and lower front legs, or (f) lower front legs. The restricted treatments used 20%, 10%, 5%, 2% or 1% of the active ingredient applied in the whole-body treatments. There was a marked seasonal effect on the performance of all treatments. With the whole-body treatment, the persistence period (knockdown > 50%) ranged from approximately 10 days during the hot, wet season (mean daily temperature > 30 degrees C) to approximately 20 days during the cool, dry season (< 22 degrees C). Restricting the application of insecticide reduced the seasonal persistence periods to approximately 10-15 days if only the legs and belly were treated, approximately 5-15 days if only the legs were treated and < 5 days for the more restricted treatments. The restricted application did not affect the landing distribution of tsetse or the duration of landing bouts (mean = 30 s). The results suggest that more cost-effective control of tsetse could be achieved by applying insecticide to the belly and legs of cattle at 2-week intervals, rather than using the current practice of treating the whole body of each animal at monthly intervals. This would cut the cost of insecticide by 40%, improve efficacy by 27% and reduce the threats to non-target organisms and the enzootic stability of tick-borne diseases.     

Use of fluorescent pigments for the automatic marking of tsetse flies in traps

A means of contaminating tsetse flies in the field with fluorescent pigment powders has been developed, using pigment in open-ended plastic chambers at the cage position on traps. Glossina pallidipes Austen and G.morsitans morsitans Westwood passed rapidly through the chambers, and on exit were contaminated with consistent doses of powder: about 90 micrograms/fly when powder was presented on the chamber roof and about 28 micrograms/fly when powder was presented on the chamber floor. The technique automatically marks tsetse flies with pigment, cheaply, simply and with the minimum imposition of stress and is expected to be particularly useful in ecological studies. Its potential for marking other biting flies is discussed.     

Artificial larviposition sites for field collections of the puparia of tsetse flies Glossina pallidipes and G. m. morsitans (Diptera: Glossinidae)

Tsetse flies Glossina pallidipes Austen and G. morsitans morsitans Westwood deposit their larvae in warthog burrows, in August-November, at Rekomitjie Research Station, Zambezi Valley, Zimbabwe. Artificial burrows, made from 200-l steel drums, were used to sample these flies and to collect their puparia. Sand-filled plastic trays in the burrows served as a substrate for larval deposition. The sand was covered with c. 2 cm of leaf litter after it was shown that only 3% of larvae were deposited on bare sand if both substrates were available. Other burrow modifications - artificially shading the burrow entrance, increasing the relative humidity inside the burrow, or reducing the size of the burrow entrance - significantly decreased deposition rates. The use of burrows in the hot season results in a reduction in the temperature experienced by the puparium towards an assumed optimum level of 26 degrees C. Artificial burrows maintained a mean temperature of 28.5 degrees C during October-November 1998, c. 2.5 degrees C cooler than ambient; earlier work has shown that natural burrows can be c. 5 degrees C cooler than ambient at these times. This may explain why natural burrows in full sunlight were used for larviposition, whereas artificial burrows were used only when they were in deep shade, and why significantly higher proportions of G. pallidipes were found in natural (66%) than in artificial burrows (34%). Better-insulated artificial burrows might produce more puparia with higher proportions of G. pallidipes. Burrows become waterlogged during the rains and may be too cool for optimum puparial development during the rest of the year. The percentages of G. m. morsitans in catches of females from artificial burrows, refuges and odour-baited traps were 34, 26 and < 10% respectively. Traps are biased in favour of G. pallidipes; artificial burrows may show a bias in favour of G. m. morsitans that is a function of temperature. Artificial warthog burrows provide a convenient way of studying the puparial stage in tsetse and for the first time facilitate the capture of females as they deposit their larvae.     

The optics of tsetse fly eyes in relation to their behaviour and ecology

Abstract The visual acuity of two species of tsetse flies, Glossina morsitans morsitans Westw. and Glossina pallidipes Aust., was investigated. Male G. morsitans eyes have an acute zone in the forward region, with large hexagonal lenses (mean minimum diameter, D=33, SE±0.7 μm), relatively small interommatidial angle (Δ(φ=1.08^o) and angular receptive field of individual ommatidia (Δp) of not less than 1.14^o. A narrow band of square lenses, with intermediate diameter and Δφ, merges with smaller hexagonal lenses in the periphery (24±0.7 μm), with relatively large interommatidial angle (Δφ=3.7^o) and small angular receptive field (Δp = c. 1.6^o). G.pallidipes eyes are similar, except that the lenses in the acute zone are larger than those of G.morsitans , in proportion to their larger body size. Female eyes are not significantly different from male eyes, except that they have a narrower region of binocular overlap (maximum for males = 24^o, for females = 18^o). The eye parameter (p=DΔφ) in the acute zone of male G.morsitans = 0.62, and in the peripheral zone = 1.56. These relatively high values are consistent with fast flight, visual detection of drift due to low wind speeds, mating chases and discrimination of cryptic host animals at high light intensities. The extended region of binocular overlap in males may serve as an early warning system of the approach of potential females. From our estimates, tsetse flies ought to be able to detect small objects against the sky c. 30 min before sunrise and after sunset, and to use their peripheral vision perhaps 15 min earlier and later than this.