Sensory and behavioural responses of the stable fly Stomoxys calcitrans to rumen volatiles

Analysis of volatiles from rumen digesta by gas chromatography linked antennogram recordings from Stomoxys calcitrans (L) (Diptera: Muscidae) antennal receptor cells revealed about 30 electrophysiologically active constituents, the most important of which is dimethyl trisulphide with a sensory threshold in the femtogram range. The behavioural responses of S. calcitrans to five chemostimulants (dimethyl trisulphide, butanoic acid, p-cresol, oct-1-en-3-ol and skatole) were tested in a wind tunnel where activation and attraction of hungry flies to rumen volatiles were recorded. Dimethyl trisulphide, butanoic acid and p-cresol were found to attract S. calcitrans. This sensitivity to rumen volatile constituents, that also occur in animal wastes used for oviposition by Stomoxys spp., as well as in flowers used by stable flies as sources of nectar is discussed in the context of the behavioural ecology of these flies.     

A new morphological character for the identification of Stomoxys calcitrans and S. niger (Diptera: Muscidae). Comparison of La Réunion island populations

On La Reunion Island (France), two morphological closely related species of stable flies (Diptera: Muscidae), living in the same environment, Stomoxys calcitrans and S. niger, are involved in the transmission of blood parasites to the livestock. To facilitate a rapid identification of both species in the field conditions, we highlighted a diagnostic morphological character not yet described: the length of the maxillary palpus. The study of three populations of S. calcitrans and two populations of S. niger, collected at various elevations, showed that the maxillary palpi of S. niger were significantly longer than in S. calcitrans, independent of sex. This character, easily visible in the field with a simple magnifying glass, has been confirmed on individuals of both species from West Africa.     

Potential impacts of climate change on stable flies, investigated along an altitudinal gradient

Adult populations of stable flies were sampled along an altitudinal transect in Reunion Island to determine whether higher temperatures were associated with: (a) higher numbers of flies; (b) a longer season of infestation, and/or (c) different responses to warming in the cosmopolitan Stomoxys calcitrans (L) and the tropical Stomoxys niger niger Macquart (Diptera: Muscidae). Flies of both species were trapped at seven farms situated at four altitudes (100-1600 m a.s.l.) over a 90-week period. For both species, there were no relationships between the maximum or mean fly abundance and altitude. Only minimum abundance in winter was significantly higher at lower altitudes. Maximum and mean abundances differed significantly between nearby farms under similar climatic conditions. Seasonal fluctuations in fly abundance changed along the gradient. At lower altitudes, population growth started earlier after the winter but abundance declined earlier in summer, which resulted in a shift of the season of infestation. Seasonal fluctuations of both species were strongly related to climate variables at high altitude, mainly temperature. However, climate variables explained a decreasing proportion of the variations in abundance at lower altitudes. Stomoxys calcitrans was the most abundant species overall, but the proportion of S. n. niger increased significantly at lower altitudes and this species became predominant at 100 m a.s.l. It is concluded that stable fly infestations are unlikely to worsen in response to global warming. Maximum abundance is limited by local factors, possibly larval resources, which suggests that adequate husbandry practices could override the impact of climate change. Because S. n. niger tends to be the predominant pest at elevated temperatures, it is recommended that this species should not be introduced in areas where climate is changing.     

Nuisance flies on Australian cattle feedlots: immature populations

Species composition, seasonality and distribution of immature fly populations on a southern Queensland feedlot during 2001-2003 were determined. Similar data were collected on feedlots in central New South Wales and central Queensland. The fly species recovered in the highest numbers were Musca domestica L. (Diptera: Muscidae), Stomoxys calcitrans L. (Diptera: Muscidae) and Physiphora clausa Macquart (Diptera: Ulidiidae). Houseflies were the dominant species at all feedlots. Houseflies preferred the warmer months from October to June, but stable flies preferred the cooler months and peaked in spring (September-November) and autumn (March-May). Larval abundance ratings recorded in the feedlot and numbers of larvae extracted in the laboratory from corresponding samples followed similar trends. Larvae of M. domestica were most abundant in the hospital and induction area and least abundant in horse stables and yards. Pupae of M. domestica were abundant in the hospital and induction area and drains, but least abundant in horse stables and yards. Larvae of S. calcitrans were most abundant in drains and least abundant in horse stables and yards. Pupae of S. calcitrans were most numerous in drains and least numerous in old cattle pens. Feedlot design and management had little effect on fly reduction.     

Attempted mechanical transmission of lumpy skin disease virus by biting insects

The mosquitoes Anopheles stephensi Liston and Culex quinquefasciatus Say (Diptera: Culicidae), the stable fly Stomoxys calcitrans Linnaeus (Diptera: Muscidae) and the biting midge Culicoides nubeculosus Meigen (Diptera: Ceratopogonidae) were allowed to feed on either lumpy skin disease (LSD) infected animals or through a membrane on a bloodmeal containing lumpy skin disease virus (LSDV). These arthropods were then allowed to refeed on susceptible cattle at various intervals after the infective feed. Virus was detected in the insects by polymerase chain reaction immediately after feeding and at sufficiently high titre to enable transmission to occur. However, no transmission of virus from infected to susceptible animals by An. stephensi, S. calcitrans, C. nubeculosus and Cx. quinquefasciatus was observed.     

First survey of seasonal abundance and daily activity of Stomoxys spp. (Diptera: Muscidae) in Kamphaengsaen Campus, Nakornpathom province, Thailand

The seasonal changes and the daily activity of Stomoxyine species (Diptera: Muscidae) were examined, using Vavoua traps, in a dairy and a beef cattle farm in Nakhonpathom province, Thailand during July 2004 to June 2005. Over this period, Stomoxys calcitrans was the most commonly trapped species, followed by S. sitiens and S. indica. For the later species, this is the first report of its presence in Thailand. A total of 80 % of flies were captured during the rainy season from May to October and 20 % during the dry season from November to April. No major difference of fly density was observed between the dairy and the beef cattle farm. The activity pattern of S. calcitrans was diurnal with a peak between 08:00 am to 10:00 am and another less marked one in the afternoon. The activity pattern of S. sitiens and S. indica was mainly crepuscular with 2 peaks, early in the morning (06:00 a.m.) and late in the afternoon (6:00 p.m.). Those species are important pests of livestock in Thailand, where they are known as a mechanical vector of trypanosomes. A better knowledge of their ecology is a prerequisite for more efficient control measures.     

Retention and attempted mechanical transmission of Ehrlichia risticii by Stomoxys calcitrans

Abstract. The ability of adult Stomoxys calcitrans (L.) (Diptera: Muscidae) to retain viable Ehrlichia risticii (Rickettsiaceae), the aetiologic agent of Potomac horse fever (PHF), and mechanically transmit the pathogen from citrated bovine blood artificially infected with E.risticii to susceptible mice was studied. Viable E. risticii were found in the digestive tract of S.calcitrans 3h after the flies had engorged to repletion on infected blood; however, no E. risticii were detected in flies ≥2 days after feeding. Subsequently, groups of S.calcitrans were fed for 20 s on infected blood, then fed to repletion on mice 30, 60,130, 180 or 220 min after having feeding interrupted. Mice displayed no clinical signs of PHF and did not produce anti- E. risticii antibodies when assayed 30 days after S. calcitrans had fed. Although S.calcitrans were able to harbour viable E.risticii for at least 3h, transmission of the disease agent to susceptible mice during interrupted feeding was not demonstrated under these experimental conditions.     

Feeding response of the stable fly, Stomoxys calcitrant L., to blood fractions and adenine nucleotides

Abstract Feeding responses of the stable fly ( Stomoxys calcitrans L., Diptera: Muscidae) to blood fractions and saline-based diets were studied in an artificial feeding device. Flies fed equally on whole blood, plasma and erythrocyte fractions while resuspended platelets or the buffy layer did not stimulate feeding. Evidence from filtration studies indicates that plasma contains both less-than-5000D and greater-than-100,000D phagostimulatory components. ATP, ADP and AMP in saline stimulate feeding in this insect in a dose-dependent manner. cAMP also stimulates feeding in the range of concentrations tested. It is argued that the less-than-5000D phagostimulant may be one or more of these adenine nucleotides.     

Diversity of Stomoxys spp. (Diptera: Muscidae) and diurnal variations of activity of Stomoxys indicus and S. calcitrans in a farm, in Wang Nam Khiao District, Nakhon Ratchasima Province, Thailand

A study of species diversity of Stomoxys spp. and diurnal variations of activity of the most abundant was performed during a one year period at a local dairy cattle farm in Wang Nam Khiao District, Nakhon Ratchasima Province, Thailand. Four species of stomoxyine flies were morphologically identified, including Stomoxys indicus Picard 1908, S. calcitrans (Linnaeus 1758), S. sitiens Rondani 1873 and S. uruma Shinonaga and Kanao 1966. The most common species were S. indicus (50.2%) and S. calcitrans (49.5%). S. sitiens and S. uruma were found in small proportions (< 1%). The number of flies captured was significantly different among the three seasons with the greatest number in the rainy season (mean = 66%; df = 2, P < 0.05). The variations of diurnal activity were observed during different period of times (06:00 to 18:00) during three seasons. Both sexes of S. indicus and males of S. calcitrans showed unimodal activity pattern in cool and summer seasons. But a bimodal activity pattern was recorded in rainy season. For females S. calcitrans, a unimodal peak of activity was observed in cool season and a constant variation of activity all along the day in summer and rainy seasons, with an increase from the morning to the evening. A better understanding of stomoxyine fly behavior, especially the daily flight activity, can assist in prioritization and design of appropriate vector prevention and control strategies.     

Purinoceptors in blood feeding behaviour in the stable fly, Stomoxys calcitrans

Abstract Both sexes of the stable fly, Stomoxys calcitrans L. (Diptera: Muscidae), have receptors on their mouthparts that mediate blood feeding. The potency ranking of the adenine nucleotides (ATP > ADP > AMP > adenosine) in eliciting feeding and suppressing the NaCl-sensitive cell may indicate the involvement of a P₂-type receptor. This is supported by the lack of effect on feeding by methyl xanthines. Feeding-related behavioural and electrophysiological results demonstrate that the potency of CH₃-S-ATP is not greater than that of ATP. These ATP-mediated responses are antagonized by ANAPP₃. Results support the conclusion that the putative 'ATP receptor' involved in stable fly phagostimulation resembles the P_2x purinoceptor of vertebrates.     

Tsetse and other biting fly responses to Nzi traps baited with octenol, phenols and acetone

Octenol (1-octen-3-ol), acetone, 4-methylphenol, 3-n-propylphenol, and other potential attractants (human urine, stable fly faeces), as well as guiacol, creosol (potential repellents), were tested as baits for biting flies in North America using standard phthalogen blue IF3GM cotton Nzi traps, or similar commercial polyester traps. Baits were tested during the summers of 2001-04 at a residence in Canada and during January-August 2001 at a dairy in the U.S.A. Behaviour in the presence of octenol was also studied by intercepting flies approaching a trap through the use of transparent adhesive film. Analogous bait and/or trap comparisons were conducted in natural settings in June 1996 in Kenya and in September-December 1997 in Ethiopia. In Canada, catches of five of six common tabanids (Tabanus similis Macquart, Tabanus quinquevittatus Wiedemann, Hybomitra lasiophthalma [Macquart], Chrysops univittatus Macquart, Chrysops aberrans Philip) and the stable fly Stomoxys calcitrans L. were increased significantly by 1.2-2.1 times with octenol (1.5 mg/h). Catches of T. quinquevittatus and S. calcitrans were 3.5-3.6 times higher on a sticky enclosure surrounding a trap baited with octenol. No other baits or bait combinations had an effect on trap catches in North America. In Ethiopia, standard Nzi traps baited with a combination of acetone, octenol and cattle urine caught 1.8-9.9 times as many Stomoxys as similarly baited epsilon, pyramidal, NG2G, S3, biconical and canopy traps, in order of decreasing catch. When baits were compared, catches in Nzi traps of six stable fly species, including S. calcitrans, were not affected by octenol (released at approximately 1 mg/h), or cattle urine (140 mg/h), used alone or in combination with acetone (890 mg/h). Acetone alone, however, significantly increased the catches of common Stomoxys such as Stomoxys niger niger Macquart, Stomoxys taeniatus Bigot, and S. calcitrans by 2.4, 1.6 and 1.9 times, respectively. Catches of Glossina pallidipes Austen were increased significantly in traps baited with acetone, urine or octenol, or any combination, relative to those in unbaited traps (1.4-3.6x). Catches of Glossina morsitans submorsitans Newstead were increased significantly by 1.5-1.7 times, but only when baits were used individually. Unlike other studies with East African tsetse, catches of both tsetse species with the complete bait combination (acetone, urine and octenol) did not differ from those in unbaited traps. Experiments with an incomplete ring of electric nets surrounding a Nzi trap, and a new approach using a sticky enclosure made from transparent adhesive film, revealed diverse responses to artificial objects and baits among biting flies. In Kenya, daily trap efficiency estimates for traps baited with either carbon dioxide (6 L/min) or a combination of acetone, cattle urine and octenol were 21-27% for G. pallidipes, 7-36% for Glossina longipennis Corti, 27-33% for S. n. niger, and 19-33% for Stomoxys niger bilineatus Grünberg, assuming 100% electrocution efficiency. Actual trap efficiencies may have been lower, given observed outside : inside electric net catch ratios of 0.6 : 1.6. Observed ratios averaged 54% of expected values, with 10 of 15 possible ratios less than the minimum possible value of 1.0.     

Larvicidal activity of endectocides against pest flies in the dung of treated cattle

Cattle were treated with topical formulations of endectocides to assess the larvicidal activity of faecal residues against horn fly, Haematobia irritans (L.), house fly, Musca domestica L., and stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae). In laboratory bioassays, doramectin, eprinomectin and ivermectin suppressed horn fly in dung of cattle treated at least 4 weeks previously and suppressed house fly and stable fly in dung of cattle treated 1-5 weeks previously. Moxidectin suppressed horn fly in dung from cattle treated no more than one week previously and did not suppress house fly and stable fly. Results combined for the three species across two experiments suggested that, ranked in descending order of larvicidal activity, doramectin > ivermectin approximately = eprinomectin > moxidectin.     

Relationships between chemical properties of larval media and development of two Stomoxys species (Diptera: Muscidae) from Reunion Island

The development of two cattle pests, Stomoxys calcitrans L. and Stomoxys niger niger Macquart (Diptera: Muscidae), was studied in the laboratory using seven potential larval media from a dairy farm on Reunion Island. The media were six types of cattle feed and an old manure medium. Egg-to-adult survival, duration of development, and adult live weight at emergence were determined for both fly species on each medium. The media were analyzed for pH, nitrogen, organic matter, and structural compounds (cellulose, hemicellulose, lignin). For S. calcitrans, immature survival was significantly higher on sugarcane leaves, Rhodes grass, and elephant grass; for S. niger, survival was significantly higher on the same substrates plus sugarcane tops. These substrates were characterized by slightly acid pH values (range, 5.4-6.0). In both species, there were significant bell-shape relationships between immature survival and substrate pH. The developmental time of both fly species was significantly shorter on Rhodes grass, Rhodes grass hay, and elephant grass. These substrates were characterized by high cellulose contents and low soluble organic fractions. In both species, there were significant linear relationships between developmental time and cellulose content of substrates. Similarly, there were significant linear relationships between adult live weight and cellulose content of substrates. The C:N ratio of the most favorable substrates was highly variable. Although the relationships revealed in this study do not establish causation, it is suggested that pH and cellulose content may have direct and indirect effects on Stomoxys development.     

A comparison of the feeding behaviour of tsetse and stable flies

In Zimbabwe, observations were made of the behaviour of individual stable flies (Stomoxys spp.) (Diptera: Muscidae) and tsetse (Glossina spp.) (Diptera: Glossinidae) feeding on cattle during the wet (Stomoxys and tsetse) and dry (tsetse only) seasons. For Stomoxys landing on adult cattle, only 27% took a full meal (mean feeding time = 147 s). Most Stomoxys left the host before completing their meal, largely due to disturbance by the host's defensive behaviour (24%, mean time = 59 s) or other flies (44%, 71 s). The probability of a Stomoxys leaving the host progressively increased with time. Simultaneous observations of tsetse showed that, compared to Stomoxys, their feeding success was lower (15%), feeding was interrupted earlier (33 s) and the time taken to complete a meal was shorter (109 s). Further studies of tsetse across different seasons and hosts showed that feeding success varied according to host age (adult = 7%; calf = 3%) and was negatively correlated with the frequency of host defensive behaviour and the relative abundance of non-biting Diptera. Disturbances were more often caused by host behaviour (69%) than other flies (31%) and the probability of tsetse leaving decreased with time on the host. Overall, these results suggest that tsetse and Stomoxys have different feeding strategies. In particular, tsetse appear to be more responsive to host defensive behaviour, which reduces their feeding success relative to Stomoxys. These behavioural differences are consistent with the respective life-history characteristics of Stomoxys and tsetse.     

Mechanical transmission of Trypanosoma evansi and T. congolense by Stomoxys niger and S. taeniatus in a laboratory mouse model

Abstract .Mechanical transmission of Trypanosoma evansi (South American origin) and T. congolense of Kilifi DNA type (Kenyan origin) was studied in laboratory mice using the African stable flies Stomoxys niger niger and S. taeniatus . Altogether, 355 flies were interrupted after feeding on infected blood and then transferred immediately to an uninfected mouse to complete feeding. Microscopy and subinoculation of triturated flies into uninfected mice demonstrated the survival of T. congolense in Stomoxys for up to 210 min and T. evansi for up to 480 min. Parasites survived for much longer periods in the digestive tract than inside or on the mouthparts. Trypanosoma congolense was transmitted only by S. n. niger , and only at low rates of 3, 8 and 10% using flies of different feeding histories: fed on blood the previous day, freshly caught, and teneral. Trypanosoma evansi was transmitted by both Stomoxys species at higher rates: S. taeniatus range 13–18%; S. n. niger range 17–35%. The highest transmission rate occurred with the combination of teneral S. n. niger and T. evansi.     

Behavioural responses of stable flies to cattle manure slurry associated odourants

Stable flies (Stomoxys calcitrans [Diptera: Muscidae] L.) are blood‐feeding synanthropic pests, which cause significant economic losses in livestock. Stable fly antennae contain olfactory sensilla responsive to host and host environment‐associated odours. Field observation indicated that the abundance of stable flies increased significantly in grasslands or crop fields when cattle manure slurry was applied. Major volatile compounds emanating from manure slurry were collected and identified. Behavioural responses of stable flies to those compounds were investigated in laboratory bioassays and field‐trapping studies. Results from olfactometer assays revealed that phenol, p‐cresol and m‐cresol were attractive to adult stable flies. When tested individually, attraction was higher with lower dosages. Stable flies were most attracted to blends of phenol and m‐cresol or p‐cresol. Traps with binary blend lures caught more stable flies in field trials as well.     

Abundance of stable flies on heifers treated for control of horn flies with organophosphate impregnated ear tags

Ear tags containing 40% organophosphate insecticides (diazinon or diazinon plus chlorpyrifos-ethyl) were applied to control Haematobia irritans (L.) (Diptera: Muscidae) in treated (TG01 and TG02) and untreated (UG01 and UG02) groups of Holstein heifers born in 2001 and 2002, respectively. Control and treated groups were assessed for the abundance of Stomoxys calcitrans (L.) (Diptera: Muscidae) from August 2001 to April 2002 and again from August 2002 to April 2003. The treatment had a high efficacy for control of horn flies (maximum median number per heifer of TG01 and TG02 = 5) but a low effect on the abundance of stable flies. The total numbers of S. calcitrans were 1251 (42.9% of the total) and 1668 (57.1%) for TG01 and UG01, and 1423 (48.8%) and 1494 (51.2%) in TG02 and UG02, respectively. No significant difference in stable fly burden was found in 55 of the 76 weeks evaluated. A unimodal peak of abundance in the spring was found during the first fly season, and a bimodal abundance, with peaks in the spring and autumn, during the second season. No strong associations between horn fly and stable fly burdens was found in individuals of the CG01 (correlation coefficient = 0.13, P > 0.05) or CG02 (correlation coefficient = 0.538, P < 0.05, determination coefficient = 0.289).     

Effect of host behaviour on host preference in Stomoxys calcitrans

Field observations suggest that, in the U.K., cattle are the preferred host of Stomoxys calcitrans (L.), followed by horses. Differences were observed in the numbers of flies feeding on individual animals both in the field and under controlled conditions. Analysis of the behaviour of four Friesian calves under attack from S. calcitrans in controlled conditions revealed that the differences in the levels of attack between individual hosts are dependent on the reactions of the host when under attack. Those hosts which respond vigorously by tail flicks, foot stamps and head-swings suffer less from attack by S. calcitrans than their more placid contemporaries.     

Nepetalactones from essential oil of Nepeta cataria represent a stable fly feeding and oviposition repellent

The stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae), is one of the most serious pests to livestock. It feeds mainly on cattle and causes significant economic losses in the cattle industry. Standard stable fly control involving insecticides and sanitation is usually costly and often has limited effectiveness. As we continue to evaluate and develop safer fly control strategies, the present study reports on the effectiveness of catnip (Nepeta cataria L.) oil and its constituent compounds, nepetalactones, as stable fly repellents. The essential oil of catnip reduced the feeding of stable flies by >96% in an in vitro bioassay system, compared with other sesquiterpene-rich plant oils (e.g. amyris and sandalwood). Catnip oil demonstrated strong repellency against stable flies relative to other chemicals for repelling biting insects, including isolongifolenone, 2-methylpiperidinyl-3-cyclohexen-1-carboxamide and (1S,2'S)-2-methylpiperidinyl-3-cyclohexen-1-carboxamide. The repellency against stable flies of the most commonly used mosquito repellent, DEET, was relatively low. In field trials, two formulations of catnip oil provided >95% protection and were effective for up to 6 h when tested on cattle. Catnip oil also acted as a strong oviposition repellent and reduced gravid stable fly oviposition by 98%.     

Ecology of Stomoxys flies (Diptera: Muscidae) in Gabon. I. First survey in different ecological areas

The stomoxyine flies are hematophagous diptera and potential vectors of various pathogenic agents. Like those of the Afrotropical Region, the stomoxyine flies of Gabon remain nearly unknown. For these reasons, an entomological survey was conducted in a transverse way in eight localities representative of the various ecological zones of Gabon. The survey was based on the use of Vavoua traps. Various environmental factors able to influence the captures were noticed and included into a canonical correspondence analysis. In total, 15,966 Stomoxys spp., belonging to seven species or subspecies, were captured. The apparent densities (DAP) expressed as the number of flies per trap and per day, were highest in Franceville (41), Bakoumba (40), Makokou (25) and Mouila (21). The most abundant species were S. n niger (33.4%), S. transvittatus (33%), then S. calcitrans (17%). The principal factors that could explain the variability of the captures were the degree of anthropisation, the botanical facies (savanna or forest), the presence of wild and domestic fauna and the nature of the vegetal cover of the ground. S. calcitrans, S. niger niger were abundant in the areas where human presence was manifest. S. xanthomelas was present in forest belts. S. transvittatus, S. omega, S. inornatus were ubiquitous species. S. niger bilineatus was found in savannas areas.