Genetics of hybridization of Glossina swynnertoni with Glossina morsitans morsitans and Glossina morsitans centralis

Abstract Reciprocal crosses were performed with Glossina swynnertoni and Glossina morsitans morsitans and with G. swynnertoni and Glossina morsitans centralis , using strains that carried marker genes in all three linkage groups. Glossina swynnertoni males can inseminate, but not fertilize, G.m.morsitans; all other crosses produced some fertile females. Hybridization did not cause sex ratio distortion among F_{ flies. Most F and backcross females were fertile, but all F, males were sterile. Sterility among backcross males was also high (99% in BXj, 85% in Bx₂, and about 50% in Bx₃ to Bx₅). Chromosome transmission by hybrid females usually conformed to Mendelian expectations, but genetic recombination was lower than observed in G.m.morsitans. The reduction in fertility among backcross females was not associated with heterozygosity in any linkage group. Sterility among hybrid and backcross males was associated with heterozygosity of sex chromosomes and probably autosomes. The results support the systematic placement of G.swynnertoni closer to G.mxentralis than to G.m.morsitans.     

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.     

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.     

Effects of age, sex and hunger on the antennal olfactory sensitivity of tsetse flies

Abstract The effects of age on electroantennogram (EAG) responses were investigated in male and female Glossina morsitans morsitans and comparative studies on the effects of starvation and sex on the EAG in G.m. morsitans, G.austeni, G.tachinoides and G.fuscipes fuscipes were made. Stimuli were the vapours of l-octen-3-ol, 4-heptanone, 3-nonanone and acetone. EAG decreased with age in both sexes of G.m.morsitans , responses in 5-day-old flies already being significantly lower than those in 1-day-old flies. In G.m.morsitans and G.tachinoides , EAG responses of males were higher than those of females. In G.austeni and G.f.fuscipes , however, the reverse was found. With increasing starvation EAG sensitivity increased in both sexes of G.m.morsitans and G.tachinoides. In G.austeni and in G.f.fuscipes no clear effects of starvation were observed. Response spectra of the individual species to the four odour substances did not change with increasing hunger. It is concluded that receptor sensitivity may be modulated depending on the insect's needs. Possible mechanisms of regulation and significance of this modulation are discussed.     

The causes of mating abstention in male tsetse flies Glossina morsitans

ABSTRACT. The sexual response of a Glossina morsitans morsitans Westw. male towards a decoy consisting of a dead male dosed with synthetic sex pheromone, was depressed by the presence of the cuticular lipids of the decoy male. However, addition of a C24 paraffin, not normally found on or in the cuticle of male or female G.m.morsitans , to a decoy male from which cuticular lipids were removed had a similar effect. Addition of either the C24 paraffin or an extract of male cuticular lipids to decoy females which possess natural sex pheromone in their cuticles, again depressed the sexual responses of males towards the decoys. The n-hexane solvent used in the application of compounds to decoys, itself depressed responses towards natural pheromone on a dead female at volumes in excess of 100μl, presumably because it mixed the surface waxes which masked the pheromone. Occlusion experiments implied that sex pheromone receptors are present on both the tarsi and tibiae of the male. No evidence was found for the existence of a specific mating deterrent on or in the cuticle of male G.m. morsitans.     

Persistent courtship reduces male and female longevity in captive tsetse flies Glossina morsitans morsitans Westwood (Diptera: Glossinidae)

Where males can increase their mating success by harassing femalesuntil they accept copulation, harassing tactics can be expectedto evolve to a point where they have costs to the longevityof both sexes. By experimentally manipulating the sex ratioin captive groups of tsetse flies Glossina morsitans morsitans,we demonstrated that the longevity of females declines wheresex ratios are biased toward males, while the longevity of malesdeclines where the sex ratio is biased toward females. Neitherirradiation of males nor prevention of copulation by blockingor damaging the external male genitalia increased the longevityof females caged with them, suggesting that female longevitywas reduced by the physical aspects of male harassment ratherthan by components of the ejaculate     

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

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

A comparison of phototactic responses to red and green light in Glossina morsitans morsitans and Musca domestica

ABSTRACT. Experiments were performed on male Glossina morsitans morsitans Westw. and female Musca domestica L. to investigate their phototactic responses at long wavelengths. In both species, green light was less attractive than red (relative to a 400 nm standard). In M. domestica this can be adequately explained by an intensity-dependent phototactic reversal at wavelengths above 500 nm; red light could be matched in attractiveness by a low intensity green light, implying that no true red-green discriminating ability exists, but that red is perceived as a dim green. Although a partial phototactic reversal was also observed in G. m. morsitans at 525–575 nm, no tested intensity of green matched red in attractiveness. It appears therefore that G. m. morsitans may be an unusual dipteran in possessing a method of discriminating red and green light independently of intensity. Both species also resolved near-UV from green.     

The roles of vision and olfaction in mate location by males of the tsetse fly Glossina morsitans morsitans

The roles of visual and/or olfactory stimuli in eliciting mating responses from male Glossina morsitans morsitans Westwood were examined, using a system for automatically recording the number and duration of mating strikes made towards decoys, under controlled conditions. The results confirm that there is no olfactory component of the female sex recognition pheromone sensed by the male antennae, and the attraction of males to females appears to be visual. The absence of male-male mating strikes was the result of the absence of female sex-pheromone, rather than the presence of a repellent mating deterrent in the male cuticle. Experiments with coloured, artificial, sex-pheromone-dosed, cotton decoys showed that colour had only weak effects on attractiveness and number of encounters with decoys, and that no colour caused significant enhancement of mating responses over those shown to decoy females.     

A behavioural bioassay to identify attractive odours for Glossinidae

1. A behavioural bioassay, based on antennal movement responses, was developed using Glossina morsitans morsitans Westwood for screening chemical attractancy to tsetse. 2. Chemicals found to be attractive to male tsetse were acetone, formaldehyde, methylethylketone, methylvinylketone, 1-octen-3-ol and pentanal but not acetophenone, hexanal, lactic acid or urea. 3. Female tsetse also responded to all these chemicals in a similar fashion. Overall responses of females were, however, less than those of males. 4. These laboratory bioassay findings agree with field observations on tsetse responses to certain chemical odours. Therefore this behavioural bioassay should be a useful laboratory test procedure for screening attractants.     

Determining the age of adult male and female Glossina morsitans morsitans using a new technique

Abstract. 1. Pteridines accumulated linearly with age up to at least 63 days in male and 140 days in female Glossina morsitans morsitans Westwood kept under laboratory conditions. Time is the major factor influencing total pteridine accumulation levels with the accumulation rate being modulated by temperature.
2. It is suggested that pteridine accumulation may be used to determine the age of male and female tsetse flies in the field.     

Formulation of pyriproxyfen, a juvenile hormone mimic, for tsetse control

A topical dose, in 1 microliter acetone, of 0.02 microgram-2-[1-methyl-2-(4-phenoxyphenoxy) ethoxy] pyridine, the juvenile hormone mimic pyriproxyfen (S-31183, Sumitomo Chemical Co.), caused an adult female tsetse, Glossina morsitans morsitans Westwood, to produce non-viable offspring for the whole of her life. Using 14C labelled pyriproxyfen it was determined that as little as 0.001 microgram transferred to the in utero larva was sufficient to arrest development in the pupal stage. A formulation in vegetable oil was prepared for treating black cotton cloth targets which caused females to pick up 0.1 microgram active ingredient (a.i.) by tarsal contact during 1 min of exposure. Males exposed similarly for between 1 and 5 min transferred up to 0.016 microgram a.i. to females if they mated immediately after treatment. Doses as low as 0.01 micrograms in 10 microliters oil cm-2 on black cotton cloth targets caused females to produce non-viable offspring for at least two reproductive cycles following exposure. However, a dose of 0.1 microgram in 10 microliters oil cm-2 was necessary for an exposed male to cause disruption of the reproductive potential of his mate. This juvenile hormone mimic has potential to induce sterility via both sexes of tsetse using treated targets or traps under field conditions.     

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

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

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.     

The role of cattle as hosts of Glossina longipennisat Galana Ranch, south-eastern Kenya

Abstract. Glossina longipennis were recorded visiting and engorging on cattle in an enclosure and on a single ox in a crush using transparent electrocuting nets in an incomplete ring. Of the total flies caught, 3–6% of males and 5–6% of females in the total catches were engorged (a feeding success rate of up to 16.6% and 12.6%, respectively, depending on assumptions made about the proportion which had an opportunity to feed). Direct observation of tsetse from an observation pit showed 57% landing on the front legs, 13% on the hind legs, and 11 % on the belly of the host. The largest number of bloodmeals was taken from the front legs, although only 14% of landings there terminated in feeding; a higher proportion of the flies alighting on the hind legs and flank succeeded in feeding (28% and 21% respectively). Glossina longipennis were attracted to targets baited with ox odour from an underground pit in a dose-dependent manner. Odour of humans was much less attractive to G.longipennis than that of oxen (for equivalent biomass). Analysis of bloodmeal samples from tsetse caught in two sites on die ranch showed that G.longipennis preferentially feeds on suids, bovids and hippopotamus.     

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)     

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.     

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

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

Homoeosis in Drosophila. II. a Genetic Analysis of Polycomb

Three dominant mutant alleles of the Polycomb locus of Drosophila melanogaster are associated with homoeotic transformations of meso- and metathoracic to prothoracic legs, a homoeotic transformation of antennae to legs, and abnormalities of wings and some thoracic bristles. Puro and Nygrén (1975) localized Polycomb in the proximal left arm of chromosome 3 within salivary gland chromosome interval 77E,F-80. In the present study, the location and dosage relationships of this locus were examined, using translocation-generated segmental aneuploidy. The results indicate that Polycomb lies within interval 78C,D-79D, and that the locus is haplo-insufficient. Males hypoploid for this interval show meso- and metathoracic leg transformations, and both males and females show wing abnormalities. In addition, the legs of hypoploids of both sexes are shorter than those of wild-type flies, and show aberrancies of segmentation, chaetal number and distribution, and other morphological characteristics. Hypoploid flies do not express a homoeotic antennal-leg transformation, but the deficiency is associated with a Minute phenotype that is known to suppress this transformation in Polycomb flies; thus it cannot be ascertained whether the antennal-leg transformation is a haplo-insufficient phenotype. It is suggested that the expression of non-homoeotic pleiotropic effects provides a criterion for identifying homoeotic mutations that do not function directly in the establishment of determined states, but rather cause homoeosis indirectly. Polycomb is interpreted in this fashion, and it is suggested that the mutant syndrome may result from localized cell death.