Estimation of the effects of buffalo fly (Haematobia irritans exigua) on the milk production of dairy cattle based on a meta-analysis of literature data

Published reports on the effect of buffalo fly Haematobia irritans exigua De Meijere (Diptera: Muscidae) and the closely related horn fly (H. irritans) were examined and analysed using non-linear weighted regression techniques in an attempt to establish the relationship between daily production loss (D), average number of parasites (n) and the average damage per parasite per day (d), and to provide estimates of expected losses in milk yield (MYD) and live-weight gain (LWG) in dairy cattle. A Mitscherlich three-parameter model was used to explain the relationship between the total loss of production attributable to buffalo flies and the average number of flies associated with cattle. This model was significant (P<0.01), with R2 = 20.2% and predicted a threshold number of flies (n = 30) below which no adverse effects would be noted. At a moderate level of infestation (n = 200) dMYD was 2.6 ml/fly/day and dLWG was 0.14 g/fly/day, resulting in estimated daily losses in milk yield (D(MYD)) and live-weight gain (D(LWG)) of 520 ml and 28 g, respectively.     

Skin lesions and cattle hide damage from Haematobia irritans infestations

The horn fly Haematobia irritans L. (Diptera: Muscidae) has recently spread to Argentina and Uruguay and is believed to cause damage to cattle hides. Four groups of ten Holstein steers each were maintained for 58 weeks under different infestation levels with H. irritans to determine if it was the cause of this problem. Hides (chrome tanned) from steers maintained under minimum infestation level had 4.7 +/- 3.8% of the area damaged. Maintaining the steers under low H. irritans level for the last 44 days of the trial using insecticidal ear-tags, resulted in 29.5 +/- 15.8% of hide area being damaged. Steers that were treated with 5% cypermethrin pour-on, when the H. irritans population was close to 50 flies, showed that 31.3 +/- 16.6% of hide area was injured, and 46.6 +/- 12.8% of damaged hide area was found in hides from non-treated steers. Significant differences were found between mean hide damage from steers maintained continuously under low H. irritans infestation levels and all other groups. Hyperaemia was significantly lower in the skin of steers under low H. irritans infestation level than in the skins of non-treated steers and steers maintained under low-level infestations for the final 44 days. Eosinophil and mononuclear cell infiltration was significantly lower when the population of H. irritans was less than six per steer than when the population was more than 100 flies per steer. Low numbers of Stomoxys calcitrans were found in all groups, but most hide damage was presumed due to H. irritans.     

Summer mastitis experimentally induced by Hydrotaea irritans exposed to bacteria

Abstract. Summer mastitis is an acute suppurative bacterial infection of the udder in heifers and dry cows. To ascertain the possible role of flies in the transmission of the disease, experimental exposures of recipient heifers to Hydrotaea irritans previously exposed to bacteria were carried out. Flies were allowed to feed on secretions from clinical cases of summer mastitis. The pathogens present were the bacteria Staphylococcus aureus, Streptococcus dysgalactiae, Actinomyces pyogenes , Stuart-Schwan cocci, Peptococcus indolicus, Fusobacterium necrophorum and Bacterioides species. The teats of eight heifers were exposed to flies with verified pathogen content. Two teats of each animal were deliberately damaged before fly exposure. One teat was cut, another pricked with insect needles to mimic insect bites. Two of the heifers developed summer mastitis in the quarters where teats had been cut. The bacterial species isolated from these quarters corresponded to those that had previously been fed to the flies. For the first time, it is now demonstrated that H.irritans is capable of transmitting summer mastitis pathogens and so causing summer mastitis in recipient heifers. Lesions on the teat orifice may be a predisposing factor in the development of the disease.     

Variation in the load of the horn fly, Haematobia irritans, in cattle herds is determined by the presence or absence of individual heifers

The distribution of horn flies, Haematobia irritans (L.) (Diptera: Muscidae), in herds of Danish Holstein-Friesian cattle was investigated in two studies conducted during two field seasons. In the first study, highly significant differences in fly distribution between the most and the least fly-susceptible heifers were observed. In one herd, the mean difference between the most fly-susceptible and the most fly-resistant heifers was 268 Ha. irritans specimens. The highest ratio between upper and lower mean fly number was 64.1:1, whereas the lowest was 3.1:1. In the second year, it was demonstrated that the heifers kept their rank in fly attraction over time. The trial clearly demonstrated that some heifers were attracting flies, whereas others, even in the same herd, only carried a few. In the second study, heifers were moved in and out of herds in an attempt to manipulate fly loads in the herds. In year 1, one herd (herd A) received four fly-resistant heifers from another herd (herd B), resulting in a drop in the mean number of flies, whereas herd B received four fly-susceptible heifers from herd A, resulting in an elevation of the mean number of flies. In year 2, a similar pattern emerged using herds C and D, and when the cattle were later returned to their original herds, the fly loads returned to their original distribution. The data presented here show unequivocally that, for horn flies, there can be considerable differences in fly loads for individual heifers within the Holstein-Friesian breed. Furthermore, the overall fly load within herds can be manipulated, and can be reversed. Thus, the distribution in the number of flies within a herd appears to depend on the number of fly-resistant or fly-susceptible heifers. The possible role of chemical factors emitted by heifers, i.e. volatile semiochemicals, in determining differences in fly loads is discussed, whereby attractants are emitted by fly-susceptible heifers and enable flies to locate their host, and repellents are emitted by fly-resistant heifers such that the flies are actively repelled from the herd.     

Dynamics of cypermethrin resistance in the field in the horn fly,Haematobia irritans

The toxicity of cypermethrin to the horn fly Haematobia irritans (L.) (Diptera: Muscidae) was determined for samples collected from untreated herds at a farm in central Argentina from October 1997 to May 2001. Field tests of the efficacy of cypermethrin against horn flies were first carried out at this farm in 1993, when the fly was shown to be susceptible to pyrethroids. Subsequently the horn fly populations on this farm were shown to have become resistant and, since 1997, the use of cypermethrin has been restricted to experimental purposes. In this study, fly samples collected in 1999, 2000 and 2001 were subjected to a polymerase chain reaction (PCR) to detect the presence of a specific nucleotide substitution in the sodium channel gene sequence, which has been associated with target site insensitivity to pyrethroids. This analysis showed that the level of cypermethrin resistance had diminished between 1997 and 2001. However, this was not sufficient to restore the efficacy of this pyrethroid to the level found prior to the onset of resistance. Heterozygous and homozygous resistant flies were detected in all samples of flies subjected to PCR diagnosis of alleles conferring target site resistance.     

The prevalence of Stephanofilaria sp. in buffalo fly, Haematobia irritans exigua, in Central Queensland

Abstract  Buffalo fly ( Haematobia irritans exigua ) infestations of cattle are associated with characteristic lesions, the initial cause of which has been attributed to a filarial nematode of the genus Stephanofilaria , for which the fly acts as a vector. Survey work in the 1980s estimated the prevalence of microfilaria in female buffalo fly in Queensland at 2.91%. Since then no information has been published and the current prevalence of microfilarial infection in buffalo fly is not known. Buffalo fly were collected from four geographically distinct sites in Central Queensland in mid-summer 2004 and were dissected to estimate Stephanofilaria sp. infection rates . Larval stages of the nematodes were recovered from female flies from all four sites and the percentage of female flies from which nematodes were recovered ranged from 29% to 57%. The average number of larvae recovered from infected female flies ranged from 1.25 to 1.75. Whereas no infected male flies were found from Sites 2–4, larvae were recovered from 43% of male flies collected at Site 1. This high prevalence of filarial infection in buffalo flies implies a correspondingly high level of transmission to cattle in Central Queensland.     

The effects of selection for size in cattle on horn fly population density

Abstract. Statistically significant differences were observed in the population density of the horn fly, Haematobia irritans irritans L., on Angus cows having significantly different frame sizes. Angus cows, averaging <112.5 cm in height at the hip, had significantly lower numbers of horn flies than Angus cows that measured 112.5–117.5 cm, 117.5–120 cm, 120–126 cm and >126 cm in height at the hip. The Angus I cows(126 cm). The estimated heritability (h²) of horn fly resistance was 0.43 ± 0.07 and 0.95 ± 0.31 for 1989 and 1990, respectively. Horn fly counts on the Angus I herd (<112.5 cm in height) was 118.1 (probable breeding value, PBV = -20.69) to 165 horn flies per cow (PBV = 26.9 flies per cow in 1989) and from 75.9 (PBV = -29.1) to 134.5 (PBV = 29.5) flies per cow in 1990. Angus I bulls had PBV = -23.7 to 13.4 and from-26.5 to 14.75 in 1989 and 1990, respectively. The Angus II cows had horn fly counts that ranged from 159.6 (PBV of-23.5) to 208.1 (PBV of 25) per cow in 1989 and from 232.3 (PBV of-56.2) to 378.7 (PBV of 90) per cow in 1990. Angus II bulls had PBVs that ranged from-17.1 to 18.9 in 1989 and from -28.1 to 48.8 in 1990. The Angus I cows had significantly (P < 0.0001) lower numbers of hom flies (mean of 63.8 horn flies per m²) than the small, medium or large Angus II cows (mean of 129.4, 149.6 and 145.5 hom flies per m², respectively). The data indicated that some specific factor(s) associated with cow size contribute(s) to innate resistance of cattle to the horn fly.     

Insecticide resistance in the horn fly: alternative control strategies

Abstract.  The horn fly, Haematobia irritans (Linnaeus 1758) (Diptera: Muscidae) is one of the most widespread and economically important pests of cattle. Although insecticides have been used for fly control, success has been limited because of the development of insecticide resistance in all countries where the horn fly is found. This problem, along with public pressure for insecticide-free food and the prohibitive cost of developing new classes of compounds, has driven the investigation of alternative control methods that minimize or avoid the use of insecticides. This review provides details of the economic impact of horn flies, existing insecticides used for horn fly control and resistance mechanisms. Current research on new methods of horn fly control based on resistant cattle selection, semiochemicals, biological control and vaccines is also discussed.     

Studies on the possible role of cattle nuisance flies, especially Hydrotaea irritans, in the transmission of summer mastitis in Denmark

The summer mastitis pathogens Actinomyces pyogenes, Peptococcus indolicus, Bacteroides melaninogenicus ss. levii, Fusobacterium necrophorum and Streptococcus dysgalactiae were isolated from the polyphagous symbovine dipterans Hydrotaea irritans (Fallén) and Morellia sp. caught around dairy heifers on pasture, but not from the haematophagous species Haematobia irritans (L.), Haematobosca stimulans (Meigen), Culicoides sp. and Simulium sp. Secretions from clinical cases of summer mastitis proved to be sources of summer mastitis bacteria for more than 3 weeks despite antibiotic treatment and teat amputation. Taking into account the seasonal activity pattern of Hydrotaea irritans and its topographical distribution on grazing cattle, it appears evident that this fly may play a central role in the establishment and maintenance of the bacterial contamination with summer mastitis pathogens on the teats of healthy cattle. In the present study the survival of A.pyogenes and P.indolicus for 7 days in experimentally infected Hydrotaea irritans, as demonstrated by the recovery of these microorganisms from agar plates exposed to live infected flies, is described. However, experimental transmission of summer mastitis from sick to healthy heifers by Hydrotaea irritans proved unsuccessful.     

Isolation and characterization of a trypsin-like enzyme from the buffalo fly, Haematobia irritans exigua

Abstract. The incorporation of soybean trypsin inhibitor (SBTI) into the diet of the buffalo fly, Haematobia irritans exigua (De Meijere), results in increased mortality and reduced fecundity. A trypsin-like enzyme which binds to SBTI was isolated by affinity chromatography on a Sepharose-SBTI column followed by ion-exchange chromatography. The enzyme was inhibited by benzamidine, phenylmethylsulfonyl fluoride, ovomucoid, leupeptin and a-2 macroglobulin. The enzyme was not inhibited by EDTA or p-chloromecuribenzoic acid and had a broad pH optimum of pH 7–9. Vaccination of sheep produced antibodies specific for the trypsin-like enzyme which inhibited enzyme activity in vitro but did not affect the survival of flies maintained in in vitro culture.     

Cattle breed-variation in infestation by the horn fly Haematobia irritans

A study was carried out to assess the resistance of pure and cross-bred groups of cattle to the horn fly Haematobia irritans (Linnaeus) (Diptera: Muscidae) in northern Argentina. Pure-bred cattle were Criolla, Iberian Bos taurus Linnaeus (Artiodactyla: Bovidae) and Nellore, Bos indicus Linnaeus (Artiodactyla: Bovidae). Cross-bred cattle were Hereford, British B. taurus (34%) X Nellore (66%) and Hereford (66%) X Nellore (34%). All were heifers and animals were maintained in two groups, each containing a mixture of pure and cross-breeds. The lowest fly numbers were found on Criolla heifers and the highest on Hereford X Nellore cross-breeds. However, it could not be determined from this study whether this was a consequence of breed and/or size, as Criolla heifers were lighter than the corresponding Hereford X Nellore heifers. Fly numbers on the heifers followed an approximately negative binomial distribution. However, the ranking of individual animals in their level of infestation within subgroups was not consistent. Hence, culling the most infested heifers on any given date would at best give only a small improvement in H. irritans control.     

Use of electroporation as an option to transform the horn fly,Haematobia irritans: a species recalcitrant to microinjection

The horn fly, Haematobia irritans, is a serious pest of cattle in North America. The control of horn flies has primarily relied on insecticides. However, the heavy use of insecticides has led to the development of insecticide resistance in horn flies. Novel methods to control horn flies are greatly needed. Transgenic technology is an effective tool to genetically modify insects and may lead to novel methods of pest control based on genomic approaches. Here we report a piggyBac‐mediated transformation of the horn fly via electroporation. Transformation with a DsRed fluorescent marker protein coding region was verified by PCR analysis of individual fly bodies and pupal cases and sequencing of PCR products. However, Southern blot analysis failed to indicate the DsRed gene was integrated into the horn fly genome. Thus, the electroporation protocol may have caused the DsRed gene to be integrated into bacterial symbionts of the horn fly.     

Accessibility of blood affects the attractiveness of cattle to horn flies

The burden of infestation of the horn fly, Haematobia irritans (Linnaeus) (Diptera: Muscidae), differs among bovines within the same herd. We hypothesized that these differences might be related to the epidermal thickness of the cattle and the blood intake capacity of the fly. Results showed that dark animals carried more flies and had a thinner epidermis than light‐coloured animals, which was consistent with the greater haemoglobin content found in flies caught on darker cattle. Similarly, epidermal thickness increased with body weight, whereas haemoglobin content decreased. Overall, we suggest that accessibility of blood is a factor that partially explains cattle attractiveness to flies.     

Interactive response of the horn fly (Diptera: Muscidae) and selected breeds of beef cattle

Statistically significant differences were observed in the population density of the horn fly, Haematobia irritans irritans (L.), on different breeds of beef cattle. The European breed Chianina had a population density of horn flies generally less than or equal to 50% than that of the British cattle breeds (Angus, Hereford, Polled Hereford, and Red Poll) and another European breed (Charolais). Generally, no significant difference existed among numbers of horn flies on Hereford, Polled Hereford, and Red Poll cows in 1988 or among Angus, Hereford, Polled Hereford, and Red Poll cows in 1989. Factors other than color appeared to be involved in the selective process between the horn fly and its host. Population densities on two white European breeds (Charolais and Chianina) were significantly different on all weekly intervals except for 4 wk in both 1988 and 1989. No significant difference existed among Charolais and British breeds except during 4 wk in 1988 and 3 wk in 1989. When weaning weights of all calves were adjusted for the effects of age to 205 d, sex of calf, and age of dam, the indirect effect of the horn fly on weaning weight showed a significant linear regression. Each 100 flies per cow caused a reduction of 8.1 kg in calf weaning weight. Cows within each breed with low numbers of horn flies weaned significantly heavier calves than cows with higher numbers of horn flies.     

Managing the horn fly (Diptera: Muscidae) using an electric walk-through fly trap

An electric walk-through fly trap was evaluated for the management of the horn fly, Hematobia irritans (L.), on dairy cattle in North Carolina over 2 yr. The trap relies on black lights and electrocution grids to attract and kill flies that are brushed from the cattle passing through. During the first season, horn fly densities were reduced from >1,400 to <200 flies per animal. Horn fly density averaged 269.2 +/- 25.8 on cattle using the walk-through fly trap twice daily, and 400.2 +/- 43.5 on the control group during the first year. The second year, seasonal mean horn fly density was 177.3 +/- 10.8 on cattle using the walk-through fly trap compared with 321.1 +/- 15.8 on the control group. No insecticides were used to control horn flies during this 2-yr study.     

The effects of hair density of beef cattle on Haematobia irritans horn fly populations

Abstract We show the relationships that exist between the amount of hair and quantity of sebum on cattle skin and the population density of the horn fly, Haematobia irritans. Brahman and Chianina steers had means of 2390 and 1587 hairs per cm², respectively, significantly more than the mean number of hairs on Angus, Brahman x Angus Crossbred, Charolais, and Red Poll steers. The Chianina steers had > 30% more sebum present on their skin and hair (0.58g/929cm²) than the Angus, Charolais, and Red Poll steers at the Beef Cattle Research Station Savoy, Arkansas. The Brahman steers had a significantly greater amount of sebum present on the skin (1.51 g/ 929 cm²) than the Crossbred and purebred Angus steers (0.55 and 0.25g/929cm², respectively) at the South Central Family Farms Research Centre Booneville, Arkansas. The Brahman and Chianina steers had means of 61.9 and 17.0 horn flies per steer, respectively, during the fly season, whereas the Angus, Crossbred, Charolais and Red Poll steers had fly season means that ranged from 76.9 to 265.8 flies per steer. Regression analysis showed that an increase of 100 hairs per cm², was associated with a reduction of 11 horn flies in the Angus II, 5 in Angus I, 20 in Charolais, 37 in Red Poll, and 0.4 in Chianina steers at the Savoy Station and a reduction of 6.6 horn flies for the Angus, Brahman, and Crossbred steers at the Booneville Centre. Regardless of cattle breed, an increase of 1.0 g of sebum per 929 cm² output by the steer was associated with 478.5 additional hairs per cm² on the animal. Each increase of 0.25 g of sebum per 929 cm² resulted in a decrease of 9.2 horn flies per steer. We conclude that some of the factors responsible for fly-resistance in cattle are hair density and the corresponding amount of sebum present on cattle skin and hair.     

Dynamics of Haematobia irritans irritans (Diptera: Muscidae) infestation on Nelore cattle in the Pantanal, Brazil

From June 1993 to May 1995, horn fly counts were conducted twice a month on untreated Nelore cattle raised extensively in the Pantanal. Horn fly population showed a bimodal fluctuation and peaks were observed every year after the beginning (November/December) and at the end (May/June) of the rainy season, which coincided with mid-late spring and mid-late fall, respectively. Horn flies were present on cattle throughout the year in at least 64% of the animals. Mean horn fly numbers on animals did not exceed 85 flies/cow during peaks and were under 35 flies/cow in most of the remaining periods. The highest infestations (population peaks) were short and dropped suddenly within two weeks. Less than 15% of the animals in both herds could be considered as "fly-susceptible" - showing consistently higher infestations, or "fly-resistant" - showing consistently lower infestations.     

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.     

The role of HiTI, a serine protease inhibitor from Haematobia irritans irritans (Diptera: Muscidae) in the control of fly and bacterial proteases

Blood-sucking arthropods are vectors responsible for the transmission of several pathogens and parasites to vertebrate animals. The horn fly Haematobia irritans irritans (Diptera: Muscidae) and the tick Boophilus microplus are important hematophagous ectoparasites that cause losses in cattle production. A serine protease inhibitor from a thorax extract of the fly H. irritans irritans (HiTI) was previously isolated, characterized and cloned. In the present study we described the expression, purification, and characterization of the recombinant HiTI (rHiTI) and its possible role in the control of different endogenous and bacterial proteases. rHiTI was successfully expressed using the pPIC9 expression vector with a yield of 4.2 mg/L of active rHiTI. The recombinant HiTI purified by affinity chromatography on trypsin-Sepharose had a molecular mass of 6.53 kDa as determined by LS-ESI mass spectrometry and inhibition constants (Kis) similar to those of native HiTI for bovine trypsin and human neutrophil elastase of 0.4 and 1.0 nM, respectively. Purified rHiTI also showed inhibitory activity against the trypsin-like enzyme of H. i. irritans using its possible natural substrates, fibrinogen and hemoglobin; and also inhibited the OmpT endoprotease of Escherichia coli using fluorogenic substrates. The present results confirm that HiTI may play a role in the control of fly endogenous proteases but also suggest a role in the inhibition of pathogen proteases.     

Dynamics and mechanisms of permethrin resistance in a field population of the horn fly, Haematobia irritans irritans

A study was conducted at the Pressler ranch, near Kerrville, Texas, USA between 2002 and 2006 to determine the dynamics and mechanisms of resistance to permethrin in a field population of the horn fly, Haematobia irritans irritans (L.). Changes of resistance to pyrethroid insecticide associated with use of a pour-on formulation of cyfluthrin in 2002 and use of diazinon ear tags in subsequent years were studied using a filter paper bioassay technique and a polymerase chain reaction assay that detects two sodium channel mutations, kdr and super-kdr resistance alleles. A maximum of 294-fold resistance to permethrin was observed in the summer of 2002. A significant decrease in the resistance level was observed in spring 2003, and resistance continued to decline after animals were treated with diazinon ear tags. In response to pyrethroid treatments, the allelic kdr and super-kdr frequency increased from 56.3% to 93.8% and from 7.5% to 43.8%, respectively in 2002, and decreased significantly in 2003 when the pyrethroid insecticide was no longer used to treat animals. Females were found to have a higher allelic super-kdr frequency than males in 2002, while no difference was detected between males and females in the allelic kdr frequency. There was a significant positive correlation between frequencies of the sodium channel mutations and levels of permethrin resistance, suggesting that the sodium channel mutations, kdr and super-kdr , are the major mechanisms of resistance to pyrethroids in this horn fly population. Results of synergist bioassays also indicated possible contributions of two metabolic detoxification mechanisms, the mixed function oxidases (MFO) and glutathione S-transferases (GST). Compared to a horn fly infestation of an untreated herd, treatments with the pyrethroid pour-on formulation failed to control horn flies at the Pressler ranch in 2002. Sustained control of horn flies was achieved with the use of diazinon ear tags in 2003 and subsequent years.