A trial to control the tsetse fly,Glossina morsitans centralis,with low densities of odour-baited targets in west Zambia

Abstract. A large-scale trial investigated the possibility of eradicating G.m.centralis from a traditional cattle rearing area using odour-baited targets at a reduced overall target density from 4 to 0.5-2.3 per km², thus cutting down initial material costs by about 50%. Only the periphery of what was thought to be prime tsetse habitat (dense woodland) was treated with targets. These were all black or blue/black cloth (1.8 × 1m), sprayed with deltamethrin suspension concentrate and baited with butanone and/or acetone (40-130mg/h) and l-octen-3-ol (0.5mg/h). Although fly catches from traps and flyrounds initially dropped by approximately 3% per day and trypanosomiasis cases declined by 99% within a year, eradication was not achieved, so that more targets were deployed at a later stage. Although initially cheaper, the option of using reduced target densities proved financially unattractive because of prolonged periods of target maintenance prior to eradication. Revised strategies for tsetse control with odour-baited, insecticide-impregnated targets in west Zambia are presented.     

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.     

Performance of the tsetse fly Glossina pallidipes reared under simple laboratory conditions

A simple grass-thatched hut was used to rear Glossina pallidipes Austen. Constructed to allow free flow of air, the temperature and humidity inside correspond to that of the outside environment. Pupae used to start the colony were obtained from pregnant females from Lambwe Valley, Western Kenya. Nine day-old females were paired with 12 day-old males for 7 days after which they were separated.Colony performance was evaluated on the basis of female survival, number and weight of puparia produced, and mortality rates. The emergence rate of puparia weighing more than 30 mg was above 80%. Data on the performance of the colony between June 1985 and June 1986 are presented. We suggest that the simple nature of the rearing facility and its direct contact with the outside environment have facilitated the colonization of the flies.

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Résumé Une cage très simple, conçue pour assurer l'arrivé d'air, et le maintien de l'humidité et de loa température extérieure, a été employée pour conserver et élever G. pallidipes. Les pupes utilisées pour constituer la souche provenaient de Lambwe valley (ouest du Kénya). Des couples de femelles vieilles de 9 jours et de mâles de 12 jours ont été consitués, les sexes ont ensiute été séparés. La performance de la souche a été estimée en fonction de la survie des femelles et du nombre et du poids des pupes produites, et des taux de mortalité. Plus de 80% des pupes ont pesé 30 mg et plus, et le taux d'émergence a été de 80%. Les résultats fournis concernent les performances observées entre juin 1985 et juin 1986. Nous estimons que la simplicité du dispositif d'élevage et le contact direct avec les conditions écologiques externes ont facilité l'éstablissement de la souche.

     

Factors affecting density-independent survival of an island population of tsetse flies in Zimbabwe

Analysis is presented of the factors affecting survival probability in populations of tsetse flies Glossina morsitans morsitans Westwood and G. pallidipes Austen (Diptera: Glossinidae) on Antelope Island, Lake Kariba, Zimbabwe. For mature male and female adult G. m. morsitans mean temperature (T _bar) accounted for 70 and 50%, respectively, of the variance in mark-recapture estimates of survival when the flies were not subjected to trapping. Saturation deficit (SD) only accounted for 36 and 33%, respectively. Maximum temperature (T_max) and SD accounted for 36–42% of the variance in male and female G. pallidipes. For the corresponding Moran curve estimates of the survival over all developmental stages, SD lagged by three weeks accounted for 61 and 41% of the variance for male and female G. m. morsitans, respectively, and 64 and 56% for G. pallidipes. The corresponding figures for plots against T_max were 44, 23, 23, and 21%, respectively. The same patterns were seen in the whole data set once allowance was made for the effect of trapping on survival and for an effect of season, correlated with an index of photosynthetic activity. For male G. m. morsitans there was a significant effect of saturation deficit, but not temperature, on immature survival. Decreased adult survival at high temperatures results from the need to feed more frequently and hence to take more risks per unit time. High saturation deficits result directly in reduced emergence of healthy flies from pupae.     

Pheromone perception in the tsetse fly Glossina morsitans morsitans

The perception of cuticular female sex pheromone (15, 19, 23-trimethylheptatriacontane=morsilure) in Glossina m. morsitans was studied electrophysiologically and behaviourally. Electrophysiological studies indicated no sensitivity of the tibial sensilla or the contact chemoreceptive hairs on the legs of males to the pheromone. However, electroantennograms were recorded during stimulation of antennae from male flies with the odour of morsilure, which indicated that the pheromone is detected by the insect via olfactory receptors on the antennae. This was further confirmed by behavioural experiments in which the antennal movements of intact males stimulated with the odour of both synthetic pheromone and female decoys were studied. Behavioural responsiveness to morsilure increased with increasing starvation and was not due to a general sensitivity to paraffins. These studies indicate that tsetse flies may be able to recognize conspecifics at close range using their sense of olfaction.

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Résumé La perception de la phéromone sexuelle des femelles de Glossina m.morsitans, la morsilure (15, 19, 23-triméthylheptatriacontane) a été examinée par électrophysiologie et par le comportement. L'électrophysiologie n'a pas révélé de sensibilité des sensilles tibiales, ni des poils chimioréceptives de contact sur les pattes des mâles à la phéromone. Cependant, les électroantennogrammes recueillis pendant la stimulation des antennes des mâles par l'odeur de morsilure, ont montré que la phéromone est décelée par l'insecte grâce aux récepteurs olfactifs des antennes. Ceci a été confirmé ultérieurement par des expériences de comportement au cours desquelles ont été examinés les mouvements antennaires de mâles intacts stimulés par l'odeur de phéromone synthétique et de femelles.La réponse comportementale à la morsilure a augmenté avec le jeûne, elle n'était pas due à une sensibilité générale aux paraffines.Ces études montrent que les mouches tsé-tsé peuvent être capables de reconnaître leurs congénères à faible distance grâce à l'olfaction.

     

The response of tsetse flies to artificial baits in relation to age, nutritional and reproductive state

In various vegetation types in Zimbabwe, the catches of Glossina pallidipes Austen and G. morsitans morsitans Westw. (Diptera: Glossinidae) at a target baited with odour (acetone, 1-octen-3-ol and two phenols) were positively correlated with catches of the same species at an unbaited net. No correlation existed between target catches and hand net catches of tsetse flies sitting on the vegetation. G. pallidipes females caught at a target and at an unbaited net were older than those caught from vegetation. Of the female G. pallidipes caught at the target, 46% were in the first 3 days of pregnancy. Of those caught at the unbaited net, significantly fewer, 21%, were in this stage. G. pallidipes males caught from vegetation contained more fat (3.07±0.333 mg) than those caught at the unbaited net (2.06±0.339 mg) or at the target (2.19±0.218 mg). It is inferred that target catches consisted predominantly of tsetse which were already in flight when they sensed the stimuli from the target, and that target catches were biased towards female G. pallidipes in the first 3 days of pregnancy.     

A general model for mortality in adult tsetse (Glossina spp.)

Tsetse exhibit a U-shaped age-mortality curve, with high losses after eclosion and a well-marked ageing process, which is particularly dramatic in males. A three-parameter (k(1) -k(3) ) model for age-dependent adult instantaneous mortality rates was constructed using mark-recapture data for the tsetse fly Glossina morsitans morsitans Westwood (Diptera: Glossinidae). Mortality changed linearly with k(1) over all ages; k(2) affected only losses in roughly the first week of adult life, and k(3) controlled the ageing rate. Mortality pooled over age was twice as sensitive to changes in k(3) as in k(1) . Population growth rate was, however, similarly affected by these two parameters, reflecting the disproportionate effect of k(3) on mortality in the oldest flies that contribute least to the growth rate. Pooled-age mortality and growth rate were insensitive to changes in k(2) . The same model also provided good fits to data for laboratory colonies of female G. m. morsitans and Glossina austeni Newstead and should be applicable to all tsetse of both sexes. The new model for tsetse mortality should be incorporated into models of tsetse and trypanosome population dynamics; it will also inform the estimation of adult female mortality from ovarian dissection data.     

A framework for studying transient dynamics of population projection matrix models

Empirical models are central to effective conservation and population management, and should be predictive of real-world dynamics. Available modelling methods are diverse, but analysis usually focuses on long-term dynamics that are unable to describe the complicated short-term time series that can arise even from simple models following ecological disturbances or perturbations. Recent interest in such transient dynamics has led to diverse methodologies for their quantification in density-independent, time-invariant population projection matrix (PPM) models, but the fragmented nature of this literature has stifled the widespread analysis of transients. We review the literature on transient analyses of linear PPM models and synthesise a coherent framework. We promote the use of standardised indices, and categorise indices according to their focus on either convergence times or transient population density, and on either transient bounds or case-specific transient dynamics. We use a large database of empirical PPM models to explore relationships between indices of transient dynamics. This analysis promotes the use of population inertia as a simple, versatile and informative predictor of transient population density, but criticises the utility of established indices of convergence times. Our findings should guide further development of analyses of transient population dynamics using PPMs or other empirical modelling techniques.     

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.     

Effects of tsetse targets on mammals and birds in Kasungu National Park, Malawi

Possible effects on wildlife of targets baited with an attractive odour (acetone), impregnated with deltamethrin and used to control tsetse flies (Glossina morsitans morsitans), were investigated in the Kasungu National Park, Malawi. Mammals and birds were censused simultaneously along transects in 'Miombo' woodland or 'Dambo' grassland/mixed woodland, with and without targets. Mammals were also monitored by surveys of their spoor on experimentally cleared plots (5 m in diameter) and a target relocation experiment was conducted when the targets were removed from the test transects and transferred to the controls. Significantly fewer small antelopes (e.g. common Duiker Sylvicapra grimmia) were detected in plots along transects with targets (tests) than along control transects. The presence of targets affected the frequency of occurrence of antelopes, suids and large herbivores in the experimental plots, but small carnivores, monkeys, rodents and hares were unaffected. A conclusion of the relocation experiment was that the deltamethrin-impregnated cloth was responsible for the observed effects and not the acetone. A total number of 23 species of birds meeting a criterion for their abundance in the areas surveyed were selected for detailed analyses. Lower indices in the test areas than in the controls were recorded for 15 of these 23 species in the Dambos and for 10 of 21 species in the Miombo woodland. Matched paired comparisons revealed significantly lower numbers in the test areas than in the controls in both habitats only for black-headed Oriole Oriolus larvatus, but for Grey Lourie Corythaixoides concolor, little bee-eater Merops pusillus, fork-tailed Drongo Dicrurus adsimilis and combined data on three species of sunbirds in the Dambos. The black-eyed bulbul Pycnonotus barbatus had significantly lower numbers in the tests than in the controls in Miombo woodland. The numbers of three species of dove were higher in the tests than in the controls in both habitats, significantly so for the Cape turtle dove Streptopelia capicola, and numbers of the Flappet Lark Mirafra rufocinnamomea were significantly higher in the tests than in the controls in Dambos. The results are discussed in the light of previous studies on environmental effects of tsetse control, including effects of tsetse targets on pollinators especially non-target horseflies.     

Integral projection models perform better for small demographic data sets than matrix population models: a case study of two perennial herbs

1. Matrix population models are widely used to describe population dynamics, conduct population viability analyses and derive management recommendations for plant populations. For endangered or invasive species, management decisions are often based on small demographic data sets. Hence, there is a need for population models which accurately assess population performance from such small data sets.
2. We used demographic data on two perennial herbs with different life histories to compare the accuracy and precision of the traditional matrix population model and the recently developed integral projection model (IPM) in relation to the amount of data.
3. For large data sets both matrix models and IPMs produced identical estimates of population growth rate (λ). However, for small data sets containing fewer than 300 individuals, IPMs often produced smaller bias and variance for λ than matrix models despite different matrix structures and sampling techniques used to construct the matrix population models.
4. Synthesis and applications . Our results suggest that the smaller bias and variance of λ estimates make IPMs preferable to matrix population models for small demographic data sets with a few hundred individuals. These results are likely to be applicable to a wide range of herbaceous, perennial plant species where demographic fate can be modelled as a function of a continuous state variable such as size. We recommend the use of IPMs to assess population performance and management strategies particularly for endangered or invasive perennial herbs where little demographic data are available.     

Trypanosoma brucei 29-13 strain is inducible in but not permissive for the tsetse fly vector

Using green fluorescent protein as a reporter, we have shown that the strain 29-13 of Trypanosoma brucei, widely used for inducible down-regulation of mRNA, is inducible in, but not permissive for the tsetse flies Glossina palpalis gambiensis and Glossina morsitans morsitans. Within two weeks post-infection, 42% males and females of teneral and non-teneral tsetse flies harboured intestinal infections, yet not a single infection progressed into the salivary glands.     

一种模拟昆虫种群动态的改进的变维矩阵模型

提出了一种模拟昆虫种群动态的改进的变维矩阵模型,该模型以发有历期为维数,采用分解与合成的方法变维,并考虑了个体间的发育差异。经模拟检验,模型模拟结果略优于徐汝梅等（1981）变维矩阵模型的结果。     

A population model for the alkali fly at Mono Lake: depth distribution and changing habitat availability

The densities of alkali fly larvae and pupae were measured in relation to depth and substrate type at six locations around Mono Lake. Samples representing a mixture of different bottom features were taken to a depth of 10 m (33 ft) using SCUBA. This is at or near the depth limit of fly larvae and pupae. The biomass of larvae and pupae on hard substrate were maximum and approximately equal at depths of 0.5 m and 1 m, substantially lower at intermediate depths of 3 m and 5 m, and over an order of magnitude further reduced at 10 m. Densities of flies on hard or rocky substrates (mainly calcareous tufa deposits), were significantly greater than those found on soft substrates such as mud or sand, at all but the greatest depth surveyed.Bathymetric maps of the areas of hard and soft substrate occurring at different lake depths were used to estimate the fly population size over the whole lake, based on the density distribution of larvae and pupae with depth on different substrates. The mapped areas of soft and hard substrates were also calculated for different lake levels, and applying the same procedure, a population model comparing the abundance of flies at different lake levels was developed. This habitat-based population model predicts that the abundance of the alkali fly is maximized at 6380 ft (1945 m) lake surface elevation. Most of the tufa substrate submerged at this lake level will become exposed and unavailable as habitat as the lake declines to 6370 ft (1942 m). In late 1991, the lake level was just over 6374 ft (1943 + m).     

Ivermectin as a possible control agent for the tsetse fly,Glossina morsitans

Ivermectin produced 100% mortality in adult teneral males, mature males and fertile females ofGlossina morsitans morsitans following a single meal of defibrinated pig blood containing concentrations of 0.1, 1.6 or >1.6 g ml^–1 respectively. The lethal concentration was reduced to <0.04 g ml^–1 for teneral males when fed repeatedly on treated blood. When pregnant females were fed a single blood meal containing ivermectin (0.08 g ml^–1) on the day after their first larviposition, followed by normal blood meals, no offspring were produced in the subsequent reproductive cycle but full recovery occurred thereafter. A dose dependent decline in fecundity was measured and data were subjected to Probit analysis. Thus estimates were made of ivermectin concentrations in the peripheral blood of treated animals by measuring the reduction in fecundity induced in flies fed on such blood. Indications are that with subcutaneous injections at least, amounts greatly in excess of the recommended clinical dose would be required to achieve levels lethal to feeding flies following a single blood meal. Oral treatment of a horse with twice the anthelmintic dose of ivermectin (0.4 mg kg^–1) produced a maximum concentration in the blood of about 0.14 g ml^–1 within 24 h and this was adequate to reduce tsetse fecundity to zero following a single meal. Such levels in a single blood meal were also sufficient to shorten the life expectancy of teneral male flies. The half-life of ivermectin in the horse was approximately 5–6 days with a maximum of 2.4% of ingested material entering the peripheral circulation. A cow treated with injectable ivermectin (0.2 mg kg^–1) produced maximum blood levels of about 0.005 g ml^–1 after one week; this was only 0.17% of the administered dose and sufficient to reduce fecundity in female flies to 50% of normal following a single blood meal. Such levels in a single blood meal had no effect on the longevity of flies. However, at least half the maximum activity was present in the circulation between 3 and 14 days following injection. Repeated feeding on the blood of a treated animal reduced considerably the dose of ivermectin required to produce a given effect. The fecundity of female flies was reduced to zero by repeated feeding on blood taken from the horse 8 days after treatment, and even after 15 days the blood of the horse contained sufficient drug to reduce fly fecundity to 50% of normal. Thus where domestic animals constitute major hosts of tsetse, treatment with ivermectin can be expected to achieve some measure of fly population reduction.

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L'invermectine comme moyen de lutte utilisable contre la mouche tsé-tsé,Glossina morsitans

Résumé Après un repas unique de sang de porc défibrillé contenant 0,1; 1,6 ou plus de 1,6 g ml^–1 d'ivermectine, tous les mâles jeunes non encore alimentes, tous les mâles adultes et toutes les femelles fécondes deGlossina morsitans morsitans sont tués. Les concentrations léthales ont été réduites à moins de 0,04 g ml^–1 pour les jeunes mâles quand on les a alimentés régulièrement sur du sang traité. Quand des femelles en gestation ont été alimentées, le jour après leur première parturition, avec un seul repas de sang contenant 0,08 g ml^–1 d'ivermectine, et ensuite avec des repas de sang normal, il n'y a pas eu production de descendants pendant le cycle suivant, bien qu'une restauration totale ait eu lieu par la suite. Une diminution de la fécondité en relation avec la dose a été enregistrée, et les données soumisses à un test Probit. Ainsi des estimations de la concentration en ivermectine du sang périphérique des animaux traités ont été obtenues en mesurant la réduction de la fécondité induite chez les mouches ayant consommé ce sang. Ceci montre qu'une dose absorbée de 4 mg kg^–1, ou une injection souscutanée de 16 mg kg^–1, seraient nécessaires pour obtenir le seuil létal chez des mouches alimentées après un repas de sang unique, c'est-à-dire 20 fois la dose absorbée et 80 fois la dose subcutanée nécessaires contre les nématodes gastrointestinaux. Le traitement par voie buccale d'un cheval avec 2 fois la dose vermifuge d'ivermectine (0,4 mg kg^–1) provoque dans les 24 heures des taux sanguins suffisants pour réduire la fécondité jusqu'à zéro après un seul repas de sang. La demi-vie dans le cheval a été approximativement de 5 à 6 jours avec une pénétration dans la circulation périphérique d'une quantité maximale de 2,4% de l'ivermectine absorbée. Une vache traitée avec de l'ivermectine injectable (0,2 mg kg^–1) atteint la teneur sanguine maximale au bout d'une semain; celle-ci, correspondant seulement à 0,17% de la quantité administrée, était suffisante pour réduire de 50% la fécondité des mouches après un repas unique de sang. Cependant, entre les 3ème et 14ème jours suivant l'injection, la circulation sanguine présente au moins la moitié de l'activité maximale. Des repas répétés sur le sang des animaux traités réduisent considérablement la dose d'ivermectine nécessaire pour produire un effet donné. La fécondité des mouches devient nulle après des repas répétés sur le sang d'un cheval 8 jours après le traitement; et même après 15 jours, le sang de ce cheval contient suffisamment de produits pour abaisser la fécondité des mouches de 50%. Ainsi, à où les animaux domestiques constituent les principaux hôtes de la mouche tsé-tsé, avec le traitement à l'ivermectine, on peut espérer réduire d'une façon efficace la population de mouches.

     

Estimating tsetse population parameters: application of a mathematical model with density-dependence

A density-dependent model is used to describe the dynamics of an open population of tsetse flies (Diptera: Glossinidae). Immigration (or emigration) takes place when the total population is below (or above) a biologically determined threshold value. The population is also subjected to birth and death rates, as well as to the risk of being trapped (continuously or intermittently). During trapping the population decreases toward a 'low' equilibrium population and when trapping ceases the population starts recovering and increases toward a 'high' equilibrium. The model is fitted using data collected on trapped flies in four experiments. The first one was conducted with 'intermittent trapping' (i.e. several trapping-recovery cycles) on Glossina fuscipes fuscipes Newstead in the Central African Republic (Bangui area). In the other experiments, trapping data on Glossina palpalis palpalis (Robineau-Desvoidy) was collected in 'aggregate' form over several days at a time. Two of these were in Congo-Brazzaville (Bouenza area) and one in the Ivory Coast (Vavoua focus). Estimates are derived for the low and high equilibrium values as well as the trapping rate. The estimated effect of sustained trapping is to reduce the population to low equilibrium values that are 85-87% lower than the levels without trapping. The effects of the natural intrinsic growth and of the migration flows cannot be estimated separately because in the model they are mathematically indistinguishable.