Fly abundance and climate as determinants of sheep blowfly strike incidence in southwest England

The relationships between abundance of the blowfly Lucilia sericata (Meigen) (Diptera: Calliphoridae), climate, animal management procedures and the incidence of cutaneous myiasis (blowfly strike) in sheep were examined in three sheep pasture systems in southwest England during the summers of 2002 and 2003. In each year, flies were collected using liver-baited sticky targets, daily weather and routine husbandry practices were noted and the age-class of each animal infested and body position of each strike were recorded. On sites where no strike control was used, 5.8-12.1% of ewes and 5.7-15.8% of lambs were struck. Ewe strikes predominated at the beginning of the season. The incidence of ewe strikes was significantly associated with higher mean temperature, rainfall and shearing; shearing was associated with a 95% reduction in the risk of ewe strike. In lambs, the incidence of strike was significantly related to higher fly abundance, ewe shearing, treatment and mean ambient temperature. Lambs were 4.6 times more likely to be struck after the ewes had been shorn than before; however, the strongest relationship was with mean L. sericata abundance. Average minimum threshold temperatures of 9.5 degrees C for lamb strikes and 8.5 degrees C for all strikes were extrapolated, below which oviposition did not occur. Over 80% of ewe strikes occurred in the breech region in 2002, as did 100% in 2003. However, in lambs both body and breech strikes occurred in both years. The distribution of lamb strikes appeared to change over time, with breech strikes predominating in May, June and July and body strikes occurring increasingly later in the season. The incidence of lamb breech strikes was significantly associated with higher L. sericata abundance and ewe shearing but there was no relationship with weather conditions. By contrast, the risk of body strike in lambs was significantly associated with higher blowfly abundance, higher rainfall and higher maximum temperatures. The relationship between strike incidence and L. sericata abundance is important because it enhances our understanding of strike incidence patterns and management of this disease. Clearly, any factors that facilitate larger L. sericata populations, such as inappropriate carcass burial or increased average ambient temperatures, are likely to increase the incidence of strike.     

Development and validation of a simulation model for blowfly strike of sheep

A comprehensive simulation model for sheep blowfly strike due to Lucilia sericata (Meigen) (Diptera: Calliphoridae), which builds on previously published versions but also incorporates important new empirical data, is used to explain patterns of lamb and ewe strike recorded on 370 farms in south-west, south-east and central England and Wales. The model is able to explain a significant percentage of the variance in lamb strike incidence in all four regions, and ewe strike in three of the four regions. The model is able to predict the start of seasonal blowfly strike within one week in three of the four regions for both ewes and lambs, and within 3 weeks in the fourth region. It is concluded that the accuracy of the model will allow it to be used to assess the likely efficacy of new control techniques and the effects of changes in existing husbandry practices on strike incidence. The model could also be used to give sheep farmers advance warning of approaching strike problems. However, the ability to forecast future strike patterns is dependent on the accuracy of the weather projections; the more long-term the forecast, the more approximate the prediction is likely to be. When applied on a regional basis, model forecasts indicate expected average patterns of strike incidence and may not therefore be appropriate for individual farmers whose husbandry practices differ substantially from the average.     

Uptake and fate of specific antibody in feeding larvae of the sheep blowfly,Lucilia cuprina

Abstract. The quantity of specific antibody ingested by larvae of Lucilia cuprina and its fate after ingestion were studied in larvae grown on sheep and on an artificial diet. Larvae grown to late first or early second instar on sheep vaccinated with horse myoglobin contained 66% less specific antibody detected by enzyme linked immunosorbent assay than larvae grown to a similar stage on an artificial diet containing 75% serum from the same sheep. A similar result was obtained when larvae were grown to mid-third instar. Larvae grown on sheep to first or second instar contained approximately the same quantity of specific antibody per unit weight of larvae as those grown to third instar. Larvae grown on diet to third instar contained 22% less specific antibody per unit weight than those grown to first or second instar. In larvae grown on diet to late third instar, ingested diet retained 91 ± 12% of its original specific antibody activity in the crop, 50 ± 11% in the anterior midgut, 8 ± 2% in the posterior midgut and 13 ± 6% in the hindgut. The mean concentration of total immunoglobulin detectable in the haemolymph of individual third instar larvae grown on diet was 1.7 ± 2.8 ug/ml. Assays of specific antibody in the haemolymph of similarly reared larvae indicated that all or most of this immunoglobulin remained functional. The implications of the quantities and distribution of ingested functional antibody found in feeding larvae of L.cuprina are discussed in relation to the possibility of vaccinating sheep against these larvae and the selection of likely internal targets as sources of potential protective antigens.     

Pre-adult mortality in the blowfly Lucilia sericata

Populations of the blowfly, Lucilia sericata (Meigen) (Diptera: Calliphoridae), have a considerable potential for rapid increase; the lifetime reproductive output of each adult female has been estimated to be between 130 and 172 eggs. Nevertheless, in the field, absolute population densities of this species are relatively low. To account for this difference, the levels of mortality affecting the eggs, feeding and wandering larvae and pupae of L. sericata were assessed in the field and laboratory. Percentage egg hatch was dependent on relative humidity with no egg eclosion at humidities below 50%; there was no significant effect of temperature on egg hatch. On infested sheep, the mean mortality of feeding larvae was 53%, but this ranged widely from 0% to 96%. There was no effect of atmospheric temperature or humidity on the mortality of feeding larvae in vivo. In the laboratory, only 10% of wandering larvae pupariated at 10 degrees C. At above 20 degrees C pupariation was consistently almost 100%. Percentage emergence increased from 0% at 10 degrees C to about 80% between 20 and 30 degrees C. The upper lethal temperature for pupae was approached at 35 degrees C. Analysis of the predation of pupae in the field revealed a weak, but significant curvilinear relationship between temperature and proportionate mortality and a median mortality of 0.49% per 24 h exposure (interquartile range = 5.2%). There was no evidence of density dependence in pupal predation. Overall, it is estimated that pre-adult mortality accounts for losses of approximately 97% of each generation, but this figure is subject to considerable variation depending on factors such as climate, time of year and host susceptibility.     

Population suppression for control of the blowfly Lucilia sericata and sheep blowfly strike

Abstract.

• 1 The control of ovine myiasis by suppression of populations of the blowfly Lucifia sericata was investigated experimentally on three farms in the south-west of England in 1992 and 1993.
• 2 In blind trials, sheep on one farm (control) were given two doses of placebo, on a second two doses of the larvicide cyromazine (Vetrazin®, CibaGeigy), and on a third cyromazine and a subsequent dose of placebo.
• 3 The first treatment was given shortly before the predicted spring emergence of L.sericata and the second shortly before the predicted emergence of the second generation. Previous simulation analysis had identified strategic early-season treatment as the optimum for blowfly population suppression.
• 4 On both treatment farms significantly smaller L.sericata populations were recorded throughout 1992 and the incidence of strike was significantly lower than on the control farm. The results show that appropriate early-season timing of sheep treatment can suppress populations of L.sericata and could be used by farmers to reduce the incidence of blowfly strike.
• 5 The results suggest, however, that the effectiveness of population suppression and strike incidence may have been influenced by immigration into the control areas and by adverse weather, the latter changing the susceptibility of sheep to strike and resulting in rising strike incidence even when L.sericata population densities were low. In practice, therefore, blowfly population suppression should be employed as a component of an integrated strike management programme.

     

Suppression of the blowfly Lucilia sericata using odour-baited triflumuron-impregnated targets

Abstract. Field trials were carried out in 1995 and 1996 on farms in the south-west of England to assess the extent to which odour-baited targets could be used to suppress populations of the ectoparasitic blowfly, Lucilia sericata , in sheep pastures. Targets were constructed from 41 × 41 cm squares of aluminium sheet, covered by white cloth which had been dipped in a mixture of sucrose solution (50% w/v) and the chitin synthesis inhibitor triflumuron (10% suspension concentrate). Each target was baited with ≈ 300 g of liver and sodium sulphide solution (10%). Three matched sheep farms were used in the trials. In 1995, triflumuron-impregnated targets were placed around the periphery of sheep pastures at one of the farms in late June, at approximately one target per hectare. In 1996, triflumuron-impregnated targets were placed around the periphery of sheep pastures of a second of the farms in early May, at approximately five targets per hectare. Each year, five sticky targets, used to monitor the L. sericata populations, were also placed in fields at the experimental and the other two farms, which acted as controls. In 1995, the results provided some, although inconclusive, evidence that the triflumuron-impregnated targets had reduced the numbers of L. sericata relative to the populations on the two control farms. In 1996, however, the density of L. sericata on the experimental farm was reduced to almost zero and remained significantly lower than on two control farms throughout the period during which the triflumuron-impregnated targets remained in the field. The results are discussed in relation to the use of triflumuron-treated targets as a practical means of controlling L. sericata and sheep blowfly strike.     

Hypersensitivity responses and repeated infections with Lucilia cuprina, the sheep blowfly.

, , and 1992. Hypersensitivity responses and repeated infections with Lucilia cuprina, the sheep blowfly. International Journal for Parasitology 22: 1175–1177. Sheep repeatedly infected with L. cuprina at 2- but not 4-week intervals developed partial resistance to infection after five infections, as measured by larval recovery. However, resistance did not persist for more than three infections. Skin weal responses were measured after injection of larval products simultaneously with each infection. The only correlation between weal size and larval recoveries occurred at infection 1 and indicated a relationship between skin sensitivity and innate rather than acquired resistance. The results suggest that resistance to L. cuprina can develop after repeated infections but that it is short lived and requires frequent larval exposure. A role for hypersensitivity responses was not confirmed by the weal responses but was suggested by the size of wound developed per larva recovered.     

Blowfly species composition in sheep myiasis in Scotland

Abstract Samples of dipteran larvae were collected from live sheep throughout Scotland, reared in the laboratory, and identified once adult flies emerged. Lucilia sericata was found in 77% of samples, and other species in 49%. The most common alternative species were L.caesar , which occurred in 31 % of samples, and Protophormia terraenovae , which occurred in 18%. Three other calliphorid species, Calliphora vomitoria, C.vicina and Lillustris , and the muscid Muscina pabulorum were also found. The proportion of samples containing alternative species was significantly lower in eastern Scotland than in western Scotland. Significantly higher proportions of samples containing alternative species were collected at altitudes of 200 m and above; from sheep of hill breeds; from rough grazing conditions and moorland; in the absence of trees; and in the presence of bracken.     

Seasonal and spatial changes in blowfly production from small and large carcasses at Durham in lowland northeast England

Colonization by blowflies (Diptera: Calliphoridae) of mouse carcasses exposed in open agricultural land near Durham (54 45'N) changed from early spring monopolization by Calliphora vicina R.-D. to a summer pattern of multiple species exploitation by this species together with Lucilia caesar L., L. illustris Mg., L. silvarum Mg., L. sericata Mg. and L. richardsi Collin. In a garden at the edge of Durham, mouse carcasses were dominated by C. vicina from spring to autumn. Difference in mouse colonization between the agricultural and garden sites seemed to reflect differences in the blowfly species present, as measured by baited trap catches at the sites. In sets of C. vicina reared from mice under conditions of competition for larval food, it was found that resulting females were significantly larger than males, size being measured as mean wing length. Blowfly production from three sheep carcasses exposed successively at the agricultural site was dominated by C. vomitoria L. and L. caesar, but also produced other Lucilia species in small numbers, including L. sericata. These L. sericata females from sheep that had died from causes other than myiasis included full-sized specimens, in contrast to those produced from mouse carcasses that were all undersized individuals. As L. sericata females trapped on sheep pastures are predominantly full-sized, this suggests that large carcasses may, in part, be a source of the L. sericata population that attacks sheep as a myiasis agent. The nature of large carcasses as possible sources of L. sericata in lowland Britain is discussed.     

Mitochondrial genomes of the sheep blowfly, Lucilia sericata, and the secondary blowfly, Chrysomya megacephala

This paper presents complete mitochondrial genomes for the sheep blowfly, Lucilia sericata (Meigen), and the secondary blowfly, Chrysomya megacephala (Fabricius). Both L. sericata and C. megacephala had standard dipteran-type mitochondrial genome architectures and lengths of 15 945 bp and 15 831 bp, respectively. Additionally, C. megacephala possessed a tRNA duplication either side of the D-loop, as previously reported in another Chrysomya species, C. putoria; this duplication appears to be synapomorphic for the genus Chrysomya. As in other insect mitochondrial genomes, base compositions had a high AT content, with both genomes more than 76% AT-rich.     

The patterning of compensatory sugar feeding in the Australian sheep blowfly

Abstract. The pattern of feeding is described for males of the Australian sheep blowfly, Lucilia cuprina , with ad libitum access to either 0.1 M or 1.0 M glucose solution. Flies given the 0.1 M solution ingested nearly 3 times the volume taken by flies given the 1.0 M solution by eating meals of, on average, twice the size about 1.5 times as frequently. Flies were usually relatively inactive following a meal, with the extent of this post-prandial quiescence being a function both of meal size and concentration of sugar. Quiescence lasted only about 20% of the average intermeal interval, however, and there was no correlation between meal size and time to the next meal. The crop emptied more slowly when it contained 1.0 M rather than 0.1 M glucose solution and the crop was, on average, fuller at the beginning of a meal on the higher concentration. The volume of solution imbibed during a meal was positively correlated with time since the end of the preceding meal. The average crop volume at the end of a meal was similar in flies feeding on 0.1 M and 1.0 M solutions. The results are considered in relation to published information on control of feeding and compensation in the blowfly Phormia regina.     

The potential for control of the blowfly Lucilia sericata using odour-baited targets

Abstract A deterministic simulation model is used to explore the potential for control of the sheep blowfly, Lucilia sericata , using odour-baited targets. The simulations show that the ambient temperatures of 15^oC or 20^oC, 13% and 24% of the females present in a population would need to be killed per day, respectively, to prevent the population growing from its initial spring density during the summer blowfly season. In contrast, if both sexes could be sterilized, so that sterilized males were also able to disrupt the fertility of unsterilized females, only 6.5–13% of both sexes would need to be attracted and sterilized at 15^oC or 20^oC respectively. To examine whether these numbers of individuals could be attracted by targets baited with liver and sodium sulphide, mark-release-recapture studies in sheep pastures were used to quantify sticky target efficiencies. The percentage of the Lsericata population caught on any one day was shown to be positively related to average ambient temperature. At 15^oC each sticky target caught approximately 13% and at 20^oC, 41% of the females within a 20 m radius. The results indicate that, at ambient temperatures between 15 and 20^oC, if distributed at a density of approximately 5/ha, targets baited with liver and sodium sulphide could effectively suppress a population of L.sericata. Lower densities of targets would be required if they were able to sterilize both sexes rather than kill or if more attractive synthetic semiochemical baits could be developed.     

The seasonal pattern of sheep blowfly strike in England and Wales

Abstract. Information on 4049 cases of sheep blowfly strike on 495 farms in England and Wales was gathered in a longitudinal survey conducted in 1991. The breech was the most commonly infested area, accounting for 70.9% of all strikes recorded. Body strikes accounted for 19.7% and foot strikes 11.4% of all strikes. Early in the season the incidences of both breech and body strikes were higher in ewes than in lambs, but from June to October lamb strikes predominated. The incidence of foot strike was greater in ewes than lambs throughout the year, with a peak incidence in September. Regional differences in strike incidence were evident; the highest overall incidences were recorded in the southern regions of England where peak incidence occurred earlier in the year. The highest monthly incidence risk was recorded in August in the south-east of England where there were 14.2 strikes per 1000 lambs and 4.0 strikes per 1000 ewes at risk. The importance of faecal soiling and ambient temperature are discussed.     

Sheep blowfly strike risk and management in Great Britain: a survey of current practice

Abstract.  The methods used for the control of sheep blowfly strike (ovine cutaneous myiasis) and the farm management factors associated with strike prevalence were examined using data from questionnaire survey returns provided by 966 sheep farmers in Great Britain, based on the period between March 2003 and February 2004. Overall, 91% of participants treated prophylactically with insecticides against blowfly strike; 39% treated twice and 11% treated more than three times in the year. Insect growth regulators (IGRs) were the most commonly chosen product (40%), especially the IGR cyromazine. Only 12% of farmers opted to dip their sheep in organophosphate insecticide against fly strike and 2% of farmers reported applying inappropriate products against strike to their sheep, such as ivermectin or 'drenches'. Farmers worming their ewes more often were 0.8 times less likely to report blowfly strike, but those who wormed their lambs more often were 1.2 times more likely to report strike. Pure-breed flocks were 0.7 times less likely to record an outbreak of blowfly strike than cross-breed flocks. Strike was less likely in ewe flocks grazed at higher altitude; however, this relationship with altitude was not seen in lambs. The results show that insecticides remain the primary tool used by almost all farmers to prevent strike and that the type of insecticides used and means of application have altered dramatically over the past 15 years. However, the prevalence of strike has remained almost unchanged over this period. Clearly careful attention to the type and timing of insecticide application, in association with a detailed understanding of the husbandry factors that predispose sheep to higher strike risk, is essential to allow the optimal management of strike problems.     

Climate change and livestock parasites: integrated management of sheep blowfly strike in a warmer environment

The incidence of parasite‐mediated livestock disease is the result of a complex interaction of factors such as parasite and host abundance, host susceptibility, climate and, critically, farmer husbandry and intervention strategies, all of which change seasonally in space and time. Given the complexity of the interacting factors, the effects of environment changes on disease incidence are hard to predict, as accordingly are the optimal husbandry responses required to ameliorate any effects. Here a model system is used to explore these issues. Cutaneous myiasis (blowfly strike) is common disease of livestock and would be expected to be highly sensitive to even small changes in climate; it provides a good model for highlighting the problems inherent in attempting to predict the effect of climate change on livestock disease incidence. For this, a stochastic simulation model is used to examine the changes in the seasonal incidence of ovine cutaneous myiasis on farms in the United Kingdom and the likely effects of changes in husbandry and control strategies. The simulations show that the range of elevated temperatures predicted by current climate change scenarios result in an elongated blowfly season with earlier spring emergence and a higher cumulative incidence of strike. Overall, higher temperatures increased strike incidence disproportionately in ewes in early summer, but had relatively less direct effect on the pattern of lamb strike incidence; a +3 °C increase in average temperature approximately doubles the cumulative incidence of strike in lambs but results in four times more strikes in ewes. A range of strike management options is examined and the models show that changes in husbandry practices are also likely to have an important effect in reducing early season ewe strike incidences. The simulations suggest that integrated changes in husbandry practices are likely to be able to manage expected increases in strike, given the range of climate changes currently predicted.     

Circadian control of spontaneous flight activity in the blowfly, Lucilia cuprina

ABSTRACT. Under laboratory light: dark cycles, the flight activity of adult Lucilia cuprina (Wied.) was low during darkness and uniformity high during light. This pattern persisted as a rhythm both in constant darkness and in constant light of intensity up to 1lx, with a period of approximately 22 h in each. Light pulses of 15 min at l00lx applied to the free-running rhythm in constant darkness generated phase shifts of up to 60°, 12-h light pulses of the same intensity generated maximal (180°) phase shifts. The phase response curves had shapes similar to those of a number of other insect rhythms. When exposed to light periods (70 lx) of greater than 12 h followed by constant darkness, the rhythm reinitiated at the light-dark transition from a constant phase equivalent to that at the time of the light-dark transition in the LD 12:12 cycle.     

Mortality rate, reproductive output, and trap response bias in populations of the blowfly Lucilia sericata

1. The ages of 877 females of the blowfly Lucilia sericata, collected from two farms in south-west England between May and September 1996, were determined using a combination of ovarian dissection and wing fray analysis. 2. Using survivorship analysis, the mortality rates over the entire field season were estimated to be 2.2 and 1.9% per day-degree at the two farms, respectively. These gave a mean life expectancy of 46 and 53 day-degrees, and a lifetime reproductive output of 130 and 172 eggs per female. 3. The mortality rate remained relatively constant over time, with one exception, the first sample of the season when no older flies were present, indicating that underlying changes in demography (emergence and senescence of generations) over the field season were not sufficient to affect the rates calculated by survivorship analysis. 4. The response of L. sericata to liver-baited traps was affected strongly by age and reproductive status, with gravid females and females in the early stages of ovarian development being most markedly over-represented relative to other age categories.     

Effects of targets impregnated with the chitin synthesis inhibitor triflumuron on the blowfly Lucilia sericata

The effects of exposure of adults of the blowfly Lucilia sericata (Meigen) for 24 h to cloth targets impregnated with the chitin synthesis inhibitor triflumuron plus sucrose or sucrose only, were considered. When both male and female L. sericata were exposed to targets treated with sucrose only, the percentage egg hatch varied between 60–75%. When both sexes were exposed to targets treated with 10% triflumuron plus sucrose the percentage egg hatch varied between 3–7%. When females alone were exposed to triflumuron, prior to the introduction of unexposed males, egg hatch varied between 4–14%. The exposure of males alone to 10% triflumuron treated targets, prior to the introduction of unexposed virgin females, did not give any consistent significant decrease in the mean percentage egg hatch. However, repetition of this trial with males exposed to targets treated with 15 or 20% triflumuron, resulted in a percentage egg hatch of 3.1% and 10.3%, respectively. These results suggest that at relatively high target concentrations, males can transfer effective doses of triflumuron to unexposed virgin females.The exposure of adults to triflumuron treated targets did not significantly increase the mortality of either male or female L. sericata. Furthermore, bioassays showed that there was no significant decline in the ovicidal effectiveness of targets treated with 10% triflumuron and sucrose solution following exposure to sun and rain for up to 8 weeks in 1995 and 1996.The results suggest that targets treated with triflumuron suspension concentrate and sucrose may form the basis of an effective system for the control of L. sericata in the field. However, the relatively high concentrations of triflumuron required and the low efficiency with which males are able to transfer effective doses to unexposed females may diminish one of the main theoretical advantages of using a chitin synthesis inhibitor, such as triflumuron, over a conventional insecticide.