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.     

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.     

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.     

Reproductive rates of tsetse flies in the field in Zimbabwe

Abstract. Tsetse flies Glossina morsitans morsitans Westwood and G. pallidipes Austen were marked and released within 12 h of emergence at Rekomitjie Research Station, Zambezi Valley, Zimbabwe, and on Redcliff Island, Lake Kariba. Ovarian dissections were performed on recaptured flies and on wild collected samples. At Rekomitjie >90% of female G. m. morsitans were inseminated by age 4 days and G. pallidipes by 7 days. For both species at both sites the length of the largest oocyte, for flies in ovarian category zero, increased approximately linearly for about the first 6 days and was ovulated at c. 6–8 days. The largest oocyte grew significantly more slowly in later cycles. For G. m. morsitans , but not for G. pallidipes , the rate increased with temperature; the rates were always higher than observed in the laboratory. At Rekomitjie, for both species and at a mean screen temperature of 22C, the first larva was produced at c. 18 days and subsequent larvae at 11–day intervals; the intervals decreased with temperature by c. 0.5 days/C. On Redcliff Island the intervals for both species were 2 days shorter than at Rekomitjie at any given screen temperature and were sometimes as short as 7 days. The length of the larva in utero increased exponentially during pregnancy.     

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.     

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.     

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

Abstract Tsetse are vectors of trypanosomes that cause diseases both in humans and livestock. Traditional tsetse surveys, using sampling methods such as Epsilon traps and black screen fly rounds, are often logistically difficult, costly and time-consuming. The distribution of tsetse, as revealed by such survey methods, is strongly influenced by environmental conditions, such as climate and vegetation cover, which may be readily mapped using satellite data. These data may be used to make predictions of the probable distribution of tsetse in unsurveyed areas by determining the environmental characteristics of areas of tsetse presence and absence in surveyed areas. The same methods may also be used to characterize differences between tsetse species and subspecies. In this paper we analyse the distribution of Glossina morsitans centralis, Glossina morsitans morsitans and Glossina pallidipes in southern Africa with respect to single environmental variables. For G.m.centralis the best predictions were made using the average NDVI (75% correct predictions; range > 0.37) and the average of the maximum temperature (70% correct predictions; 27.0–29.2°C). For G.m.morsitans the best prediction was given by the maximum of the minimum temperature (84% correct predictions; range > 18.8°C), and for G.pallidipes , also by the maximum of the minimum temperature (86% correct predictions; range > 19.6 °C). The following paper compares a range of multivariate techniques for making predictions about the distribution of these species in the same region.     

Feeding frequency in relation to reproduction in Glossina morsitans morsitans and G.pallidipes

Abstract Measurements of residual haematin in males of Glossina morsitans morsitans Westwood reared in the laboratory at 25^oC suggest that blood meal digestion is completed in 4 or 5 days after feeding. However, a high proportion of haematin is present as faecal matter 2 days after feeding and it is concluded that digestion is completed sooner than indicated by the regression of logio haematin on time. Therefore, low levels of residual haematin in field-caught tsetse provide no indication of the frequency with which they feed. For this reason the effects of feeding frequency upon various reproductive parameters in the laboratory have been examined. It is concluded that the best performance is achieved by G.m. morsitans females which ingest four blood meals per inter-larval period and that for a similar performance in G.pallidipes five blood meals are required. The extent to which such feeding frequencies are a reflection of feeding activity in the field are discussed in terms of the biochemical requirements to maintain a reproductive adult female tsetse in positive energy balance.     

Characterization of microsatellite markers in the tsetse fly, Glossina pallidipes (Diptera: Glossinidae)

Glossina pallidipes is a vector of African trypanosomiasis. Here we characterize eight new polymorphic microsatellite loci in 288 G. pallidipes sampled from 12 Kenya populations. The number of alleles per locus ranged from four to 36 with a mean of 20.5 +/- 10.1. Expected single locus heterozygosities varied from 0.044 to 0.829. Heterozygosity averaged 0.616 +/- 0.246. No linkage disequilibrium was found. We also report results in eight other tsetse species estimated by using the primers developed in G. pallidipes. The primers worked best in G. swynnertoni and G. austeni and worst in G. m. morsitans and G. m. submorsitans.     

Reproductive abnormalities in tsetse flies in Zimbabwe

Between November 1988 and July 1995 five technicians carried out ovarian dissections on 16,013 Glossina morsitans morsitans Westwood and 123,848 G. pallidipes Austen captured at Rekomitjie Research Station, Zambezi Valley, Zimbabwe. The ovarian age and uterine content were recorded, as were the lengths (l₁, l₂ and l_u) of the largest and second largest oocyte, and of any uterine inclusion. Major abnormalities and abnormal spermathecal contents were found in <0.1% of all flies dissected. Apparent abortions rates varied significantly between dissectors and occurred at frequencies of 0.8–4.5% in G. m. morsitans and 0.3–2.8% in G. pallidipes. The lowest estimates give the best picture of the field situation. Abortion rates were higher in flies caught on electric nets than in trapped flies where the rate was only 0.15%, indicating that reproductive losses are negligible for most of the year at Rekomitjie. The rates did, however, increase to >2% when mean temperatures exceeded 27°C and flies were captured in artificial refuges. There was little effect of ovarian age on the abortion rate, but the frequency of empty uteri declined markedly with age – with a suggestion, however, that it might increase again in the oldest flies. A knowledge of the rates of reproductive loss is important for the construction of realistic models of the dynamics of tsetse populations.     

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.     

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.     

Activation of three species of tsetse (Glossina spp.) in response to host derived stimuli

Recordings were made of the activation of hungry Glossina morsitans morsitans Westwood, G. pallidipes Austen, and G. austeni Newstead in response to odours from ox breath and ox urine, and a moving visual stimulus, in a wind tunnel. The spontaneous activity of G.m.morsitans was very low (less than 4% of males and 2% of females active per min during control periods). That of G.austeni and G.pallidipes was in the region of 20% except for G.pallidipes females when in excess of 40% were active during control periods. Addition of ox urine odours to the airstream had no effect on activity in any of the species investigated but addition of ox breath odours to the airstream significantly increased activity of G.pallidipes and of G.m.morsitans, although for the latter only approximately 12% of flies were active. For G.austeni the addition of ox breath odours resulted in a significant increase in activity of females but not of males. The moving visual stimulus resulted in a significant increase in the activity of both sexes of G.austeni and G.m.morsitans but no change in the activity of G.pallidipes. The low level of spontaneous activity and the low response to ox breath odours in a strain of G.m.morsitans maintained in the laboratory since 1969 was compared with a new colony of this species which originated from puparia collected in Zimbabwe in 1991. No differences in either spontaneous activity or the response to ox breath odour was recorded, but females from the new colony were significantly more responsive to a moving visual stimulus.(ABSTRACT TRUNCATED AT 250 WORDS)     

The influence of sex and fly species on the development of trypanosomes in tsetse flies

Unlike other dipteran disease vectors, tsetse flies of both sexes feed on blood and transmit pathogenic African trypanosomes. During transmission, Trypanosoma brucei undergoes a complex cycle of proliferation and development inside the tsetse vector, culminating in production of infective forms in the saliva. The insect manifests robust immune defences throughout the alimentary tract, which eliminate many trypanosome infections. Previous work has shown that fly sex influences susceptibility to trypanosome infection as males show higher rates of salivary gland (SG) infection with T. brucei than females. To investigate sex-linked differences in the progression of infection, we compared midgut (MG), proventriculus, foregut and SG infections in male and female Glossina morsitans morsitans. Initially, infections developed in the same way in both sexes: no difference was observed in numbers of MG or proventriculus infections, or in the number and type of developmental forms produced. Female flies tended to produce foregut migratory forms later than males, but this had no detectable impact on the number of SG infections. The sex difference was not apparent until the final stage of SG invasion and colonisation, showing that the SG environment differs between male and female flies. Comparison of G. m. morsitans with G. pallidipes showed a similar, though less pronounced, sex difference in susceptibility, but additionally revealed very different levels of trypanosome resistance in the MG and SG. While G. pallidipes was more refractory to MG infection, a very high proportion of MG infections led to SG infection in both sexes. It appears that the two fly species use different strategies to block trypanosome infection: G. pallidipes heavily defends against initial establishment in the MG, while G. m. morsitans has additional measures to prevent trypanosomes colonising the SG, particularly in female flies. We conclude that the tsetse-trypanosome interface works differently in G. m. morsitans and G. pallidipes.     

Responses of tsetse flies (Glossina spp.) to compounds on the skin surface of an ox: a laboratory study

The behaviour of male Glossina morsitans morsitans Westwood and Glossina pallidipes Austen (Diptera: Glossinidae) alighting on targets with or without ox sebum was compared. The presence of ox sebum did not increase significantly the number of flies alighting on the target in either species. However, after contact with the sebum coated target, both species showed an increase in flight activity, and G. m.morsitans showed a greater tendency to return to the target. This behaviour resulted in a number of short flights which may reflect the search for a feeding site on a host. The duration of each visit to the target was significantly reduced when sebum was present for G. m. morsitans but not for G. pallidipes. This is explained by documented differences in the resting behaviour of the two species which shows that G. m. morsitans normally rests for longer periods on the surface of an untreated black target than does G. pallidipes. Other experiments showed that the presence of sebum elicited a probing response in G. m. morsitans and G. pallidipes. The results are discussed with reference to the possible use of host sebum to improve trap catches in the field.     

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.     

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.     

Interactions between cattle and biting flies: effects on the feeding rate of tsetse

In Zimbabwe, studies were made of the effect of host behaviour on the feeding success of Glossina pallidipes Austen and G. morsitans morsitans Westwood (Diptera: Glossinidae) attracted to cattle of different age and sex. The mean feeding rates for male and female G. pallidipes attracted to oxen were 60% and 58%, respectively, compared to 33% and 53% for male and female G. m. morsitans. The feeding rate of G. pallidipes varied between oxen and was inversely correlated with a host's rate of defensive leg movements, which, in turn, was positively correlated with the density of Stomoxys spp. (Diptera: Muscidae) caught in the vicinity of the host. Tsetse were significantly less successful in feeding from young cattle. For G. pallidipes, the feeding rate on calves (<6 months) was 11%, whereas for male and female G. m. morsitans the rates were 12% and 20%, respectively. Significantly lower feeding rates were apparent for cattle aged up to 2 years, when the feeding rate for G. pallidipes (31%) was still significantly less than that on mature oxen (68%). Feeding rates for G. pallidipes on adult female cattle were lower than those on oxen (45% vs. 61%). The lower feeding rates in young animals were attributed to higher rates of defensive movements. The results suggest that higher rates of defensive activities by young cattle reduce the risk of them contracting trypanosomiasis.     

Evaluation of insecticide-treated cattle as a barrier to re-invasion of tsetse to cleared areas in northeastern Zimbabwe

A field trial in Zimbabwe investigated the efficacy of insecticide-treated cattle as a barrier to prevent the re-invasion of tsetse, Glossina morsitans and G. pallidipes (Diptera: Glossinidae), into cleared areas. The original tsetse barrier consisted of insecticide-treated odour-baited targets, at an operational density of four to five targets per km2, supported by insecticide-treatments of cattle with either deltamethrin dip (Decatix, Coopers) at two-weekly intervals, or deltamethrin pouron (Spoton, Coopers) at monthly intervals, in a band approximately 20 km wide from the re-invasion front. Tsetse catch, and trypanosomiasis incidence in nine sentinel herds was recorded for 7-8 months, respectively, before the targets were removed, leaving only the insecticide treatment of the local cattle to stem the re-invasion of tsetse. After the removal of the target barrier, the tsetse readily invaded the trial area and the incidence of trypanosomiasis in sentinel herds increased, while their PCVs decreased. After seven months without the targets in place, trypanosomiasis prevalence in the local stock had reached alarmingly high levels; the trial was terminated prematurely and the target barrier re-deployed. Immediately after the re-deployment of the target barrier, the tsetse catch in the trial area reverted to acceptable levels along the re-invasion front, and trypanosomiasis incidence in the sentinel cattle decreased. It is concluded that, under the conditions of the field trial, the insecticidal treatment of local cattle did not in itself form an effective barrier to tsetse re-invasion. By contrast, the target barrier performed as was predicted by mathematical and experimental analysis, and readily cleared the tsetse infestation and reduced trypanosomosis incidence in the trial area.     

The host-orientated behaviour of tsetse flies (Glossina): the interaction of visual and olfactory stimuli

ABSTRACT. Studies were made in Zimbabwe of the propensity of Glossina pallidipes Austen and G. morsitans morsitans Westwood to divert from flying upwind in plumes of host odour to various visual features (termed targets). Using various arrangements of electrocuting nets with targets placed downwind of an odour source it was found that 45% diverted to a square target, c. 30% diverted to a black vertical oblong and there was no significant diversion to a bark-coloured vertical oblong that simulated the bole of a tree. The relative propensity of tsetse to divert to variously coloured targets decreased in the order: black = blue > red > yellow; for different shapes it decreased in the order: circle > square > horizontal oblong = vertical oblong. Changes in the composition or concentration of the odour, or loss of contact with it, did not markedly affect the percentage that diverted. Tsetse that diverted to a target and subsequently flew away from it showed an upwind bias in the presence of odour. In the absence of odour there was a slight crosswind bias. If these crosswind fliers then flew into a plume of host odour they turned c. 50 upwind.