Insecticide-treated cattle against tsetse (Diptera: Glossinidae): what governs success?

The distributions of insecticide-treated cattle from sites in Tanzania and Zimbabwe were assessed from interviews with livestock owners, analysis of secondary livestock data and mapping technologies. The time-course of tsetse control operations at these sites were then simulated using a mathematical model that assumed diffusive movement and logistic growth in fly populations. A simulation of a tsetse control operation in Mudzi district, north-east Zimbabwe, was in accord with observations that the use of insecticide-treated cattle was unable to prevent substantial re-invasion of tsetse from Mozambique, consequent on the patchy distribution of cattle. The simulation was also consistent with the observed efficacy of a 10-km wide barrier of insecticide-treated targets deployed evenly at 4 km/(-2). Simulation of a control operation on Mkwaja Ranch in Tanzania was in accord with the observation that the use of insecticide-treated cattle reduced the tsetse population on the ranch by c. 90%. Insecticide-treated cattle were used to better effect in the Kagera Region of Tanzania. Simulation of this operation predicts that the deployment of 35,000 treated cattle in the area would result in > 99% control of the tsetse population, consistent with the observed decline, by 1-2 orders of magnitude, in cases of trypanosomiasis in the region. The greater success of the Kagera operation was due to the size and shape of the treated area and, particularly, to the restriction of re-invasion to 20% of the perimeter, compared with > 80% on Mkwaja. Simulation was used to assess how tsetse control could have been improved at Mkwaja. The results suggest that splitting herds into smaller, more numerous, units could have achieved some improvement but, in general, the disease problem would not have been solved by the use of insecticide-treated cattle alone. Only by deploying odour-baited targets in ungrazed areas, or in a 1-3-km barrier around the ranch, could substantially better control (99-99.9%) have been achieved. Sensitivity analyses of the Mkwaja simulation showed that the general conclusions were robust to assumptions regarding cattle distribution and the rates of fly movement and growth. Properly managed and appropriately applied insecticide-treated baits are powerful weapons for tsetse control but should not be used without regard to potential levels of re-invasion, consequent largely on considerations of the size and shape of the treatment area and the density and distribution of the baits.     

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.     

Selective use of odour-baited, insecticide-treated targets to control tsetse flies Glossina austeni and G. brevipalpis in South Africa

The effectiveness of odour-baited targets treated with 0.8% deltamethrin in controlling Glossina austeni Newstead and G. brevipalpis Newstead (Diptera: Glossinidae) was evaluated in Zululand, South Africa. Targets were initially deployed in the three habitat types (grassland, woodland and forest) of two adjacent areas at a density of four targets per km(2). One area functioned as the treatment block (c. 35 km(2)) and included the focus of the target deployment, and the second area functioned as a barrier block (c. 40 km(2)) against tsetse fly re-invasion from the untreated area to the south. After 8 months, targets were removed from open grassland in both areas and target density in wooded habitats and sand forest was increased to eight per km(2). Twelve months later, all targets were removed from the barrier block and used to increase target density in the wooded and sand forest habitats of the treatment block to 12 per km(2). This target density was maintained for 14 months. In the treatment area, a 99% reduction in G. austeni females occurred after 13 months at a target density of eight per km(2) in wooded habitat; this was maintained for 22 months. Reduction in G. brevipalpis was less marked. The relatively poor reduction in G. brevipalpis is attributed to the high mobility of this species and its distribution throughout less wooded and more open habitats.     

Insecticide-treated cattle for tsetse control: the power and the problems

Trypanosomiasis control increasingly involves financial input from livestock owners and their active participation. If control is carried out on smaller scales than in the past, methods such as aerial and ground spraying and sterile insect techniques will have reduced application. There will be increased reliance on trypanocidal drugs, and bait methods of tsetse control--where flies are attracted to point sources and killed. If drug resistance develops, cheap and simple bait methods offer the only means of disease control that might be applied, and paid for, by stockowners themselves. The methods have been effective in some circumstances, but not in others, and it is important to understand the reasons for the successes and the failures. Analysis is presented of the results of two Tanzanian tsetse control campaigns involving the use of insecticide-treated cattle. Between 1991 and 1996, following the introduction of widespread dipping in the Kagera Region, trypanosomiasis declined from >19000 cases to <2400 and deaths from >4000 to 29. On four ranches in the region, tsetse have been almost eliminated and trypanosomiasis prophylaxis is no longer used. Similarly aggressive use of pyrethroids on Mkwaja Ranch in Tanga Region has not had such dramatic effects. Tsetse and trypanosomiasis are still common, despite high levels of prophylaxis and the deployment of approximately 200 odour-baited targets. The difference in the results is attributed to a combination of the much smaller area covered by treated animals at Mkwaja, a greater susceptibility to re-invasion and a more suitable habitat for the flies. A better understanding of the dynamics of the use of insecticide-treated cattle is needed before we can predict confidently the outcome of particular control operations.     

The use of discriminant analysis for the estimation of sampling biases for female tsetse

ABSTRACT.

• 1 In this paper we investigate whether the technique of discriminant analysis can be used to estimate sampling biases for female tsetse.
• 2 Discriminant analysis was first applied to laboratory samples of female tsetse, Glossina morshans morsitans Westwood, to test whether flies of known history could be assigned to the correct day of the pregnancy cycle on the basis of their fat, haematin and corrected residual dry weight.
• 3 Following the satisfactory results from the laboratory samples, the same technique was applied to field samples of G.m. centralis Machado captured by electric traps and hand nets in Zambia and of G.palpalis palpalis (Robineau-Desvoidy) captured in biconical traps at five sites in Ivory Coast. The results show that flies on day 1 of the pregnancy cycle were most likely to be caught, with a second peak of day-6 and day-7 flies, while very few day-8 or day-9 flies were caught.
• 4 These major peaks in fly trappability coincide with the known feeding habits of female tsetse, and indicate synchrony of feeding by many members of the population immediately after larviposition and again as the larva in utero moults from the second to third instar. G.palpalis is relatively more available at this later stage of its pregnancy cycle to the capture methods used than is G.morsitans. A third feed may be taken at a more variable point in the pregnancy cycle.
• 5 This method of estimating the sampling biases of female tsetse could allow an estimate of total population size, as long as the absolute sampling efficiency of flies on any one day of the pregnancy cycle could be established by, for example, mark-release-recapture experiments.

     

Control of tsetse and trypanosomiasis transmission in Uganda by applications of lambda-cyhalothrin

The pyrethroid insecticide lambda-cyhalothrin was evaluated in field trials against Glossina f.fuscipes and sleeping sickness transmission in Iyolwa sub-county, Tororo District, Uganda. The insecticide was applied selectively to the resting-sites of tsetse, by bush-spraying, using 10% wettable powder (10WP) formulation at an application rate of 11.6 g a.i./ha over an area of 28 km2, or by a 2% Electrodyn formulation (2ED) applied at 0.9 g a.i./ha over 30 km2. In a third trial area of 32 km2, 215 pyramidal traps treated with lambda-cyhalothrin 100 mg/m2 were set. The best impact was obtained with 10WP lambda-cyhalothrin which eliminated tsetse within 1-2 months, whereas G.f.fuscipes persisted at very low density in part of the area treated with 2ED lambda-cyhalothrin. In both treated areas the numbers of human sleeping sickness cases fell to no more than one per month, compared with four to twelve per month previously. The overall rate of cattle trypanosomiasis (T.brucei and T.vivax) was also reduced slightly. Insecticide-treated traps remained fully effective for at least 6 months under field conditions and catches were reduced 20-90-fold. These results in the control of tsetse and trypanosomiasis transmission lead us to recommend lambda-cyhalothrin for tsetse control operations.     

User-friendly models of the costs and efficacy of tsetse control: application to sterilizing and insecticidal techniques

An interactive programme, incorporating a deterministic model of tsetse (Diptera: Glossinidae) populations, was developed to predict the cost and effect of different control techniques applied singly or together. Its value was exemplified by using it to compare: (i) the sterile insect technique (SIT), involving weekly releases optimized at three sterile males for each wild male, and (ii) insecticide-treated cattle (ITC) at 3.5/km(2). The isolated pre-treatment population of adults was 2500 males and 5000 females/km(2); if the population was reduced by 90%, its growth potential was 8.4 times per year. However, the population expired naturally when it was reduced to 0.1 wild males/km(2), due to difficulties in finding mates, so that control measures then stopped. This took 187 days with ITC and 609 days with SIT. If ITC was used for 87 days to suppress the population by 99%, subsequent control by SIT alone took 406 days; the female population increased by 48% following the withdrawal of ITC and remained above the immediate post-suppression level for 155 days; the vectorial capacity initially increased seven times and remained above the immediate post-suppression level for 300 days. Combining SIT and ITC after suppression was a little faster than ITC alone, provided the population had not been suppressed by more than 99.7%. Even when SIT was applied under favourable conditions, the most optimistic cost estimate was 20-40 times greater than for ITC. Modelling non-isolated unsuppressed populations showed that tsetse invaded approximately 8 km into the ITC area compared to approximately 18 km for SIT. There was no material improvement by using a 3-km barrier of ITC to protect the SIT area. In general, tsetse control by increasing deaths is more appropriate than reducing births, and SIT is particularly inappropriate. User-friendly models can assist the understanding and planning of tsetse control. The model, freely available via http://www.tsetse.org, allows further exploration of control strategies with user-specified assumptions.     

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.     

Evaluations of permethrin-impregnated clothing and three topical repellent formulations of deet against tsetse flies in Zambia

Permethrin-impregnated clothing and three topical repellent formulations of deet (diethyltoluamide) were field tested against natural populations of tsetse flies, mostly Glossina morsitans centralis Machado, in central Zambia. Volunteers wore different combinations of clothing impregnated with permethrin 0.125 mg ai/cm2 and repellents while riding in a vehicle that was driven slowly (4-6 km/h), with the windows and rear door open, through fly-infested areas. The mean rate of tsetse bites was about twenty per 75 min for unprotected people. The treatment combination of permethrin-impregnated clothing (blue cotton coveralls) and either of two controlled-release deet formulations on exposed skin of face and arms provided 91% mean protection, but this was not significantly better (P greater than 0.05) than wearing deet repellent alone (76-87% protection). No significant differences of protection were observed between the three repellent treatments, although the two controlled-release formulations (intended to be more persistent) were applied at approximately half the dosage of the standard 75% deet. Wearing permethrin-impregnated coveralls alone provided relatively poor protection (34%) for the untreated and exposed skin of head and hands. However, olive drab mesh jackets treated with permethrin reduced the tsetse biting rate by 75%.     

Glossina austeni (Diptera: Glossinidae) eradicated on the island of Unguja, Zanzibar, using the sterile insect technique

An area-wide integrated tsetse eradication project was initiated in Zanzibar in 1994 by the International Atomic Energy Agency and the governments of Tanzania and Zanzibar, to eradicate Glossina austeni Newstead from Unguja Island (Zanzibar) using the sterile insect technique. Suppression of the tsetse population on Unguja was initiated in 1988 by applying residual pyrethroids as a pour-on formulation to livestock and by the deployment of insecticide impregnated screens in some of the forested areas. This was followed by sequential releases of gamma-sterilized male flies by light aircraft. The flies, packaged in carton release containers, were dispersed twice a week along specific flight lines separated by a distance of 1-2 km. More than 8.5 million sterile male flies were released by air from August 1994 to December 1997. A sterile to indigenous male ratio of >50:1 was obtained in mid-1995 and it increased to >100:1 by the end of 1995. As a consequence the proportion of sampled young females (1-2 ovulations), with an egg in utero in embryonic arrest or an uterus empty as a result of expulsion of a dead embryo, increased from <25% in the 1st quarter to >70% in the last quarter of 1995. In addition, the age structure of the female population became significantly distorted in favor of old flies (> or = 4 ovulations) by the end of 1995. The apparent density of the indigenous fly population declined rapidly in the last quarter of 1995, followed by a population crash in the beginning of 1996. The last trapped indigenous male and female flies were found in weeks 32 and 36, 1996, respectively. Time for 6 fly generations elapsed between the last catch of an indigenous fly and the end of the sterile male releases in December 1997.     

Is the Even Distribution of Insecticide-Treated Cattle Essential for Tsetse Control? Modelling the Impact of Baits in Heterogeneous Environments

Background

Eliminating Rhodesian sleeping sickness, the zoonotic form of Human African Trypanosomiasis, can be achieved only through interventions against the vectors, species of tsetse (Glossina). The use of insecticide-treated cattle is the most cost-effective method of controlling tsetse but its impact might be compromised by the patchy distribution of livestock. A deterministic simulation model was used to analyse the effects of spatial heterogeneities in habitat and baits (insecticide-treated cattle and targets) on the distribution and abundance of tsetse.

Methodology/Principal Findings

The simulated area comprised an operational block extending 32 km from an area of good habitat from which tsetse might invade. Within the operational block, habitat comprised good areas mixed with poor ones where survival probabilities and population densities were lower. In good habitat, the natural daily mortalities of adults averaged 6.14% for males and 3.07% for females; the population grew 8.4× in a year following a 90% reduction in densities of adults and pupae, but expired when the population density of males was reduced to <0.1/km²; daily movement of adults averaged 249 m for males and 367 m for females. Baits were placed throughout the operational area, or patchily to simulate uneven distributions of cattle and targets. Gaps of 2–3 km between baits were inconsequential provided the average imposed mortality per km² across the entire operational area was maintained. Leaving gaps 5–7 km wide inside an area where baits killed 10% per day delayed effective control by 4–11 years. Corrective measures that put a few baits within the gaps were more effective than deploying extra baits on the edges.

Conclusions/Significance

The uneven distribution of cattle within settled areas is unlikely to compromise the impact of insecticide-treated cattle on tsetse. However, where areas of >3 km wide are cattle-free then insecticide-treated targets should be deployed to compensate for the lack of cattle.     

Contrasting population structures of two vectors of African trypanosomoses in Burkina Faso: consequences for control

Background

African animal trypanosomosis is a major obstacle to the development of more efficient and sustainable livestock production systems in West Africa. Riverine tsetse species such as Glossina palpalis gambiensis Vanderplank and Glossina tachinoides Westwood are the major vectors. A wide variety of control tactics is available to manage these vectors, but their removal will in most cases only be sustainable if the control effort is targeting an entire tsetse population within a circumscribed area.

Methodology/Principal Findings

In the present study, genetic variation at microsatellite DNA loci was used to examine the population structure of G. p. gambiensis and G. tachinoides inhabiting four adjacent river basins in Burkina Faso, i.e. the Mouhoun, the Comoé, the Niger and the Sissili River Basins. Isolation by distance was significant for both species across river basins, and dispersal of G. tachinoides was ∼3 times higher than that of G. p. gambiensis. Thus, the data presented indicate that no strong barriers to gene flow exists between riverine tsetse populations in adjacent river basins, especially so for G. tachinoides.

Conclusions/Significance

Therefore, potential re-invasion of flies from adjacent river basins will have to be prevented by establishing buffer zones between the Mouhoun and the other river basin(s), in the framework of the PATTEC (Pan African Tsetse and Trypanosomosis Eradication Campaign) eradication project that is presently targeting the northern part of the Mouhoun River Basin. We argue that these genetic analyses should always be part of the baseline data collection before any tsetse control project is initiated.     

Control of Glossina longipennis (Diptera: Glossinidae) by insecticide-treated targets at Galana ranch, Kenya, and confirmation of the role of G. longipennis as a vector of cattle trypanosomiasis

Glossina longipennis Corti was studied in Galana Ranch, Kenya over a four year period, in two areas (Tank E and Lali) where the species was abundant and other species were absent or scarce. There was active transmission of trypanosomiasis to cattle in both areas, the parasite species being Trypanosoma vivax Ziemann and T. congolense Broden. Mean infection rates of the G. longipennis were 1.1% and 0. 55% for T. vivax and T. congolense respectively at Tank E, and 0.88% and 0.15% at Lali. Experimental transmission studies showed that cattle in fly-proof enclosures challenged with wild G. longipennis collected from Galana became infected with both trypanosome species. A tsetse control operation in one area (Tank E) using targets impregnated with deltamethrin in an oil formulation reduced the population of G. longipennis by 98% over one year, despite evidence of re-invasion. Populations of G. longipennis in the other area (Lali) were relatively stable over the whole study period. The effect of tsetse control on the incidence of cattle trypanosomiasis at Tank E was less clear than that on tsetse numbers, probably due to the lack of a sustained reduction in tsetse numbers. However, a significant relationship was demonstrated between fortnightly incidence measurements and electric net catches of G. longipennis at Tank E. A further significant predictor of incidence was rainfall in the previous four to seven weeks. This study confirms the importance of G. longipennis as a vector of bovine trypanosomiasis in areas where it is the predominant tsetse present.     

Use of deltamethrin 'pour-on' insecticide for the control of cattle trypanosomosis in the presence of high tsetse invasion

A deltamethrin 'pour-on' insecticide was applied monthly to over 2000 cattle exposed to a high challenge of drug-resistant trypanosomes and high tsetse re-invasion pressure in the Ghibe valley, south-west Ethiopia. Blood samples were taken monthly from an average of 760 cattle for determination of PCV and presence of trypanosomes. The area of the valley is approximately 350 km2 and the cattle grazed in roughly four locations covering about a quarter to half of the area. Two years before the trial commenced, Glossina morsitans submorsitans Newstead (Diptera: Glossinidae) began to invade the valley. Despite the use of the pour-on the mean apparent density of G. m. submorsitans continued to rise, and, during the 4 years of tsetse control, was more than three-fold higher than that recorded during the previous 18 months. Over the same period there was little change in the apparent density of Glossina pallidipes Austen (Diptera: Glossinidae). By contrast, the mean monthly prevalence of trypanosome infections in cattle over 36 months of age decreased from 38.3 to 29.0%, the incidence of new infections decreased from 26.6 to 16.0% (a reduction of 40%), and packed cell volume in cattle increased from 21.7 to 24.1%. Evidence of a change in apparent parasite transmission rate was demonstrated by regression of infection incidence in cattle on the logarithm of apparent density of G. m. submorsitans. Before the trial started the regression coefficient was 45.8 +/- 6.3 and this reduced to 9.2 +/- 2.5% incidence per log(e) (flies/trap/day) during the period of tsetse control. It was concluded that this indicated reductions in tsetse numbers in the immediate vicinities of cattle in a way that was not reflected in overall tsetse catches. Nevertheless, the comparatively high levels of trypanosome prevalence that persisted in the cattle demonstrates that, where invasion prevalence is high, treatment of small pockets of cattle will not eradicate tsetse. To achieve more significant reduction in trypanosome prevalence in cattle, integrated methods of control utilizing target barriers in the major routes of invasion will be needed.     

The prevalence of African animal trypanosomoses and tsetse presence in Western Senegal

In 2005, the Government of Senegal initiated a tsetse eradication campaign in the Niayes and La Petite C?te aiming at the removal of African Animal Trypanosomosis (AAT), which is one of the main constraints to the development of more effective cattle production systems. The target area has particular meteorological and ecological characteristics that provide great potential for animal production, but it is unfortunately still infested by the riverine tsetse species Glossina palpalis gambiensis Vanderplank (Diptera: Glossinidae). The tsetse project in Senegal has adopted an area-wide integrated pest management (AW-IPM) approach that targets the entire tsetse population within a delimited area. During the first phase of the programme, a feasibility study was conducted that included the collection of entomological, veterinary, population genetics, environmental and socioeconomic baseline data. This paper presents the parasitological and serological prevalence data of AAT in cattle residing inside and outside the tsetse-infested areas of the target zone prior to the control effort. At the herd level, a mean parasitological prevalence of 2.4% was observed, whereas a serological prevalence of 28.7%, 4.4%, and 0.3% was obtained for Trypanosoma vivax, T. congolense and T. brucei brucei, respectively. The observed infection risk was 3 times higher for T. congolense and T. vivax in the tsetse-infested than in the assumed tsetse-free areas. Moreover, AAT prevalence decreased significantly with distance from the nearest tsetse captured which indicated that cyclical transmission of the parasites by tsetse was predominant over mechanical transmission by numerous other biting flies present. The importance of these results for the development of a control strategy for the planned AW-IPM campaign is discussed.     

Distribution and Density of Tsetse Flies (Glossinidae: Diptera) at the Game/People/Livestock Interface of the Nkhotakota Game Reserve Human Sleeping Sickness Focus in Malawi

In large parts sub-Saharan Africa, tsetse flies, the vectors of African human or animal trypanosomiasis, are, or will in the foreseeable future, be confined to protected areas such as game or national parks. Challenge of people and livestock is likely to occur at the game/livestock/people interface of such infested areas. Since tsetse control in protected areas is difficult, management of trypanosomiasis in people and/or livestock requires a good understanding of tsetse population dynamics along such interfaces. The Nkhotakota Game Reserve, an important focus of human trypanosomiasis in Malawi, is a tsetse-infested protected area surrounded by a virtually tsetse-free zone. The abundance of tsetse (Glossina morsitans morsitans) along the interface, within and outside the game reserve, was monitored over 15 months using epsilon traps. A land cover map described the vegetation surrounding the traps. Few flies were captured outside the reserve. Inside, the abundance of tsetse at the interface was low but increased away from the boundary. This uneven distribution of tsetse inside the reserve is attributed to the uneven distribution of wildlife, the main host of tsetse, being concentrated deeper inside the reserve. Challenge of people and livestock at the interface is thus expected to be low, and cases of trypanosomiasis are likely due to people and/or livestock entering the reserve. Effective control of trypanosomiasis in people and livestock could be achieved by increasing the awareness among people of dangers associated with entering the reserve.     

A cost-benefit analysis of feeding in female tsetse

Abstract. Three models for feeding in female tsetse are considered. Model I: there is a prolonged non-feeding phase after each meal followed by feeding at a constant rate, with a constant probability of dying as a consequence of feeding. Model II: the feeding rate increases linearly after each meal. Model III: the feeding rate increases exponentially after each meal. In Models II and III the feeding hazard is a linear function of the probability of feeding. Production of viable female offspring is estimated under each model, making allowance for losses of adults due to starvation and to background and feeding mortality, losses of pupae due to predation and parasitization, and losses of young flies if their mothers take insufficient blood during pregnancy. Under Model I, if females require three meals to produce viable pupae in 9 days, then for a non-decreasing population with a background mortality of 1%/day, and 25% pupal losses due to predation and parasitism, the feeding risk must be ≤5%/feed. At this maximum level the non-feeding phase should be 2–2.5 days for optimal productivity, with a mean feeding interval of 60–72 h. If the background mortality is 2%/day, feeding losses cannot exceed 1%/feed for a non-decreasing population. If four or five meals are required for the production of fully viable pupae, the optimal values of the non-feeding phase and mean feeding interval tend towards 1 and 2 days respectively. Under Models II and HI the mean feeding interval is 50–60 h for optimal productivity (with variances 3 times as large as for Model I), in good agreement with estimates from recent models for feeding and digestion. Field evidence suggests that feeding tsetse take greater risks as their fat levels dwindle. This should result in feeding (and feeding mortality) rates which increase during the feeding phase - as assumed in Models II and III but not in Model I. These models allow greater flexibility than Model I, because flies can feed early in the hunger cycle, at low probability, as long as the feeding risk is also low.     

Thermal tolerance in a south-east African population of the tsetse fly Glossina pallidipes (Diptera, Glossinidae): implications for forecasting climate change impacts

For tsetse (Glossina spp.), the vectors of human and animal trypanosomiases, the physiological mechanisms linking variation in population dynamics with changing weather conditions have not been well established. Here, we investigate high- and low-temperature tolerance in terms of activity limits and survival in a natural population of adult Glossina pallidipes from eastern Zambia. Due to increased interest in chilling flies for handling and aerial dispersal in sterile insect technique control and eradication programmes, we also provide further detailed investigation of low-temperature responses. In wild-caught G. pallidipes, the probability of survival for 50% of the population at low-temperatures was at 3.7, 8.9 and 9.6 degrees C (95% CIs: +/-1.5 degrees C) for 1, 2 and 3 h treatments, respectively. At high temperatures, it was estimated that treatments at 37.9, 36.2 and 35.6 degrees C (95% CIs: +/-0.5 degrees C) would yield 50% population survival for 1, 2 and 3 h, respectively. Significant effects of time and temperature were detected at both temperature extremes (GLZ, p<0.05 in all cases) although a time-temperature interaction was only detected at high temperatures (p<0.0001). We synthesized data from four other Kenyan populations and found that upper critical thermal limits showed little variation among populations and laboratory treatments (range: 43.9-45.0 degrees C; 0.25 degrees C/min heating rate), although reduction to more ecologically relevant heating rates (0.06 degrees C/min) reduce these values significantly from approximately 44.4 to 40.6 degrees C, thereby providing a causal explanation for why tsetse distribution may be high-temperature limited. By contrast, low-temperature limits showed substantial variation among populations and acclimation treatments (range: 4.5-13.8 degrees C; 0.25 degrees C/min), indicating high levels of inter-population variability. Ecologically relevant cooling rates (0.06 degrees C/min) suggest tsetses are likely to experience chill coma temperatures under natural conditions (approximately 20-21 degrees C). The results from acute hardening experiments in the Zambian population demonstrate limited ability to improve low-temperature tolerance over short (hourly) timescales after non-lethal pre-treatments. In flies which survived chilling, recovery times were non-linear with plateaus between 2-6 and 8-12 degrees C. Survival times ranged between 4 and 36 h and did not vary between flies which had undergone chill coma by comparison with flies which had not, even after factoring body condition into the analyses (p>0.5 in all cases). However, flies with low chill coma values had the highest body water and fat content, indicating that when energy reserves are depleted, low-temperature tolerance may be compromised. Overall, these results suggest that physiological mechanisms may provide insight into tsetse population dynamics, hence distribution and abundance, and support a general prediction for reduced geographic distribution under future climate warming scenarios.     

A parity-structured matrix model for tsetse populations

A matrix model is used to describe the dynamics of a population of female tsetse flies structured by parity (i.e., by the number of larvae laid). For typical parameter values, the intrinsic growth rate of the population is zero when the adult daily survival rate is 0.970, corresponding to an adult life expectancy of 1/0.030 = 33.3 days. This value is plausible and consistent with results found earlier by others. The intrinsic growth rate is insensitive to the variance of the interlarval period. Temperature being a function of the time of the year, a known relationship between temperature and mean pupal and interlarval times was used to produce a time-varying version of the model which was fitted to temperature and (estimated) population data. With well-chosen parameter values, the modeled population replicated at least roughly the population data. This illustrates dynamically the abiotic effect of temperature on population growth. Given that tsetse flies are the vectors of trypanosomiasis ("sleeping sickness") the model provides a framework within which future transmission models can be developed in order to study the impact of altered temperatures on the spread of this deadly disease.     

Nutritional stress affects the tsetse fly's immune gene expression

Tsetse-transmitted trypanosomiasis poses a serious threat to human and animal health in sub-Saharan Africa. The majority of tsetse flies ( Glossina spp.) in a natural population will not develop a mature infection of either Trypanosoma congolense or Trypanosoma brucei sp. because of refractoriness, a phenomenon that is affected by different factors, including the tsetse fly's immune defence. Starvation of tsetse flies significantly increases their susceptibility to the establishment of a trypanosome infection. This paper reports the effects of nutritional stress (starvation) on (a) uninduced baseline levels of gene expression of the antimicrobial peptides attacin, defensin and cecropin in the tsetse fly, and (b) levels of expression induced in response to bacterial ( Escherichia coli ) or trypanosomal challenge. In newly emerged, unfed tsetse flies, starvation significantly lowers baseline levels of antimicrobial peptide gene expression, especially for attacin and cecropin. In response to trypanosome challenge, only non-starved older flies showed a significant increase in antimicrobial peptide gene expression within 5 days of ingestion of a trypanosome-containing bloodmeal, especially with T. brucei bloodstream forms. These data suggest that a decreased expression of immune genes in newly hatched flies or a lack of immune responsiveness to trypanosomes in older flies, both occurring as a result of fly starvation, may be among the factors contributing to the increased susceptibility of nutritionally stressed tsetse flies to trypanosome infection.