Incidence of frugivorous flies (Tephritidae and Lonchaeidae), fruit losses and the dispersal of flies through the transportation of fresh fruit

Frugivorous flies (Diptera: Tephritidae and Lonchaeidae) cause significant losses in horticultural crops. These losses are associated with the fruit damaged by frugivorous fly infestation occurring during all steps of the production and commercial sale. Pest monitoring is the main tool used to identify and avoid future losses in horticultural crops. In this study we monitor the population of frugivorous flies in a fruit orchard and a food supply centre (FSC) to assess fruit losses and examine the dispersal of flies. Six Anastrepha species, Ceratitis capitata (Wiedemann), five Neosilba species and Dasiops sp. were recovered from traps and fruit samples. The fruit losses were estimated for star fruit (26%), guava (17%), mango (4%), orange (1.9%), tangerine (0.9%) and apple (0.2%). The fruit collected at the FSC demonstrated that immature frugivorous flies can disperse for several kilometres through the transportation of fresh fruit. Due to the importance of frugivorous flies, this poses a threat to the production and commercial sale of fruit around the world. Our study presents information relevant to the pest management and risk mitigation for frugivorous flies.     

Molecular characterization of esterase E3 gene associated with organophosphorus insecticide resistance in the New World screwworm fly, Cochliomyia hominivorax

Abstract.  The New World screwworm, Cochliomyia hominivorax (Coquerel) (Diptera: Calliphoridae), is one of the most important myiasis-causing flies in South America. It is responsible for severe economic losses to livestock producers, mainly because it causes mortality in newborn calves and reductions in the quality of leather and in the production of milk and meat. The economic losses caused by myiasis, along with those caused by other internal and external parasites, are the main factors limiting meat production. In Brazil, C. hominivorax has been controlled by applying insecticides, particularly organophosphate (OP)-based compounds. However, the improper and continuous use of these chemicals can lead to the selection of OP-resistant strains. This, associated with the fast development of OP resistance in other myiasis-causing flies, shows the importance of investigating resistance in C. hominivorax. Based on the findings of previous studies, the objective of the current work was to isolate and sequence the E3 gene in C. hominivorax. Mutations at the positions (Gly137 and Trp251) responsible for conferring OP resistance in Lucilia cuprina and Musca domestica L. (Muscidae) were identified in C. hominivorax . In addition, the orthologous region in C. hominivorax contained motifs that are highly conserved among carboxyl/cholinesterases and contribute to the catalytic mechanism of the active site. The characterization of this gene in natural populations of New World screwworm can be an important tool for monitoring resistance to insecticides throughout its current geographic distribution. This will provide information for the selection and implementation of more effective pest management programmes.     

The Monetary Burden of Cystic Echinococcosis in Iran

Cystic echinococcosis (CE) is a globally distributed parasitic infection of humans and livestock. The disease is of significant medical and economic importance in many developing countries, including Iran. However, the socioeconomic impact of the disease, in most endemic countries, is not fully understood. The purpose of the present study was to determine the monetary burden of CE in Iran. Epidemiological data, including prevalence and incidence of CE in humans and animals, were obtained from regional hospitals, the scientific literature, and official government reports. Economic data relating to human and animal disease, including cost of treatment, productivity losses, and livestock production losses were obtained from official national and international datasets. Monte Carlo simulation methods were used to represent uncertainty in input parameters. Mean number of surgical CE cases per year for 2000–2009 was estimated at 1,295. The number of asymptomatic individuals living in the country was estimated at 635,232 (95% Credible Interval, CI 149,466–1,120,998). The overall annual cost of CE in Iran was estimated at US$232.3 million (95% CI US$103.1–397.8 million), including both direct and indirect costs. The cost associated with human CE was estimated at US$93.39 million (95% CI US$6.1–222.7 million) and the annual cost associated with CE in livestock was estimated at US$132 million (95% CI US$61.8–246.5 million). The cost per surgical human case was estimated at US$1,539. CE has a considerable economic impact on Iran, with the cost of the disease approximated at 0.03% of the country''s gross domestic product. Establishment of a CE surveillance system and implementation of a control program are necessary to reduce the economic burden of CE on the country. Cost-benefit analysis of different control programs is recommended, incorporating present knowledge of the economic losses due to CE in Iran.     

High levels of pyrethroid resistance in horn flies (Diptera: Muscidae) selected with cyhalothrin.

Lambdacyhalothrin cattle ear tags controlled horn fly, Haematobia irritans (L.), for 14 wks or longer during 1986-1988 in Georgia, USA. In 1989 and 1990, control of < 50 horn flies per side of cow was achieved for < or = 4 wk because of high levels of pyrethroid resistance in horn flies selected with lambdacyhalothrin. The highest resistance ratios (RRs) were seen in 1989. These were 498 for lambdacyhalothrin; 92,000 for fenvalerate; and 54 for permethrin. RRs for cypermethrin as high as 8,800 were estimated in 1990 when the RR for fenvalerate was only 1,060. No cross-resistance to diazinon was detected. These high levels of pyrethroid resistance seem to have a large component of metabolic resistance. Synergistic coefficients as high as 3,600 were determined by addition of nonlethal amounts of piperonyl butoxide. Resistance development in a no-pyrethroid-use area indicates movements of > or = 3km by sufficient numbers of horn flies can significantly change the RR.     

Winter feeding sites of hay in round bales as major developmental sites of Stomoxys calcitrans (Diptera: Muscidae) in pastures in spring and summer

The stable fly, Stomoxys calcitrans (L.), historically has been a pest of livestock in confined operations but seldom of animals on pastures or rangelands. In the past two decades, however, S. calcitrans has become a major pest of cattle and horses on pastures in the midwestern United States. Although there usually is an overabundance of diverse stable fly and house fly, Musca domestica L., larval habitats in confined livestock operations, no larval habitat for stable flies has been clearly identified in the pasture-range environment. Because the winter feeding of hay in round bales results in significant amounts of hay wastage that when mixed with manure, might develop into suitable larval habitats, this study evaluated these areas as developmental sites for the abundant stable flies in pastures. There was a trend for fly traps placed in the vicinity of hay feeding sites to catch more stable flies than those placed distant from these sites. Estimates of stable flies emerging from these sites ranged from 102 to 1225 flies per core sample (25 by 25 cm). The mean number of adult stable flies during May and June 2001 through 2004 correlated negatively with the average minimum temperatures during the preceding winter (November-February) but not with rainfall or temperatures during the spring. These results support the hypothesis that winter feeding sites of hay in round bales are the main source of stable flies in pastures.     

New approaches to the chemical control of arthropod pests of livestock

Chemicals are important tools in our effort to control livestock ectoparasites, and they are likely to remain so in the foreseeable future. Chemical control is the most widely used and in most cases the only method available to the producer. Therefore, it is imperative that we continue to seek new chemicals as well as new systems for delivery of those chemicals in an effective, efficient, and safe manner.New approaches to chemical control involve both new chemicals and new delivery systems in a complementary and often a synergistic relationship. Three relatively new classes of chemicals, pyrethroids, insect growth regulators, and avermectins, have provided special opportunities for development of new delivery systems.The development of the insecticidal ear tag has been perhaps the most significant new approach to chemical control of livestock ectoparasites in recent years. Although originally developed for control of ear ticks, the tag has had its greatest impact on control of the horn fly, a major pest of cattle in the U.S. The pyrethroid tags provided season-long control of horn flies in most climates and at the same time dramatically reduced the use of pesticide. Unfortunately, the problem of resistance to pyrethroids in the tags has been encountered. Unless effective techniques for resistance management can be developed, the producer may lose this valuable new tool.Sustained-release boluses provide a means of delivering IGR's, such as methoprene and diflubenzuron, for control of immature stages of the horn fly and face fly in manure of cattle. These devices are designed to lodge in the reticulum of cattle and release the chemical to the digestive tract over an extended period. Bolus formulations may in fact provide a tool for management of horn fly resistance to pyrethroids.Ivermectin, a potent new chemical from a new class of agricultural pesticides, provides an important opportunity for developing innovative delivery systems for control of livestock pests. Its systemic activity against ticks and blood-sucking flies at extremely low dosages provides potential for development of long-lasting subcutaneous implants. Its ability to control immature stages of dung-breeding flies, such as the horn fly and face fly, offers the potential for development of sustained-release boluses for larvicidal treatments.These three selected examples of new approaches to chemical control of livestock ectoparasites point to two obvious trends: 1) reduction in quantity of a chemical to be applied without sacrifice of efficacy and 2) use of controlled-release technology to prolong effectiveness of the chemical. Although the examples presented are focused on applications to cattle, similar approaches have application for other livestock. New approaches such as these have important implications in implementation of either area-wide or IPM control programs. Such programs will need convenient and effective chemical delivery systems to be successful if in fact chemicals are to be a part of the programs.     

Identification of stable fly attractant compounds in vinasse,a byproduct of sugarcane–ethanol distillation

The stable fly, Stomoxys calcitrans (Diptera: Muscidae), is a worldwide pest of livestock. Recent outbreaks of stable flies in sugarcane fields in Brazil have become a serious problem for livestock producers. Larvae and pupae found inside sugarcane stems after harvesting may indicate that stable flies use these stems as potential oviposition or larval development sites. Field observations suggest that outbreaks of stable flies are associated with the vinasse and filter cake derived from biomass distillation in sugarcane ethanol production that are used as fertilizers in sugarcane fields. Adult stable flies are attracted to vinasse, which appears to present an ideal larval development site. The primary goal of the present study is to demonstrate the role of vinasse in influencing the sensory physiological and behavioural responses of stable flies, and to identify its associated volatile attractant compounds. Both laboratory and field studies showed that vinasse is extremely attractive to adult stable flies. Chemical analyses of volatiles collected revealed a wide range of carboxylic acids, alcohols, phenols and aldehydes as potential attractant compounds. These newly identified attractants could be used to develop a tool for the attractant‐baited mass trapping of stable flies in order to reduce infestations.     

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%.     

Luring houseflies (Musca domestica) to traps: do cuticular hydrocarbons and visual cues increase catch?

Abstract.  Houseflies ( Musca domestica L.) are a major pest species of livestock units and landfill sites. Insecticide resistance has resulted in an increased emphasis on lure-and-kill control methods, but the success of this approach relies on the effective attraction of houseflies with olfactory or visual stimuli. This study examined the efficacy of olfactory (cuticular hydrocarbons) or visual (colours and groups of flies) attractants in a commercial poultry unit. Despite simulating the cuticular hydrocarbon profiles of male and female houseflies, we found no significant increase in the number of individuals lured to traps and no sex-specific responses were evident. The use of target colours selected to match the three peaks in housefly visual spectral sensitivity yielded no significant increase in the catch rate of traps to which they were applied. This study also demonstrated that male and female flies possess significantly different spectral reflectance (males are brighter at 320–470 nm; females are brighter at 470–670 nm). An experiment incorporating groups of recently killed flies from which cuticular hydrocarbons were either removed by solvent or left intact also failed to show any evidence of olfactory or visual attraction for houseflies of either sex. This study concluded that variations of the most commonly applied methods of luring houseflies to traps in commercial livestock units fail to significantly increase capture rates. These results support commonly observed inconsistencies associated with using olfactory or visual stimuli in lure-and-kill systems, possibly because field conditions lessen the attractant properties observed in laboratory experiments.     

Phylogenetic analysis of New World screwworm fly, Cochliomyia hominivorax, suggests genetic isolation of some Caribbean island populations following colonization from South America

Abstract.  Larval infestations of the New World screwworm (NWS) fly, Cochliomyia hominivorax , cause considerable economic losses through the direct mortality and reduced production of livestock. Since the 1950s, NWS populations in North and Central America have been the target of virtually continuous eradication attempts by sterile insect technique (SIT). Nevertheless, in some areas, such as Jamaica, SIT-based control programmes have failed. Reasons for the failure of SIT-based programmes in some locations are unknown, but it is hypothesized that failure may be related to the mating incompatibility between sterile and wild flies or to the existence of sexually incompatible cryptic species. Accordingly, the current research investigates intraspecific phylogenetic relationships and associated biogeographic patterns between NWS populations from the Caribbean and South America, which represent those populations involved in, or earmarked for, forthcoming SIT programmes. Uniquely, this study also includes analyses of two North American samples, collected in Texas in 1933 and 1953 prior to initiation of the SIT-based eradication programme. The study utilizes three nucleotide datasets: elongation factor-1α (nuclear); cytochrome oxidase subunit 1 (mitochondrial), and 12S rRNA (mitochondrial). Phylogenetic analysis of these data, representing populations from across the Caribbean, South America and Texas, indicates sub-structuring of fly populations on several of the larger Caribbean islands, suggesting a period of isolation and/or founder effects following colonization from South America; significantly, our findings do not support a North American origin for Cuban flies. The importance of these findings in the light of proposed SIT programmes in the region is discussed.     

Genetic Approaches for Studying Myiasis-causing Flies: Molecular Markers and Mitochondrial Genomics

“Myiasis-causing flies” is a generic term that includes species from numerous dipteran families, mainly Calliphoridae and Oestridae, of which blowflies, screwworm flies and botflies are among the most important. This group of flies is characterized by the ability of their larvae to develop in animal flesh. When the host is a live vertebrate, such parasitism by dipterous larvae is known as primary myiasis. Myiasis-causing flies can be classified as saprophagous (free-living species), facultative or obligate parasites. Many of these flies are of great medical and veterinary importance in Brazil because of their role as key livestock insect-pests and vectors of pathogens, in addition to being considered important legal evidence in forensic entomology. The characterization of myiasis-causing flies using molecular markers to study mtDNA (by RFLP) and nuclear DNA (by RAPD and microsatellite) has been used to identify the evolutionary mechanisms responsible for specific patterns of genetic variability. These approaches have been successfully used to analyze the population structures of the New World screwworm fly Cochliomyia hominivorax and the botfly Dermatobia hominis. In this review, various aspects of the organization, evolution and potential applications of the mitochondrial genome of myiasis-causing flies in Brazil, and the analysis of nuclear markers in genetic studies of populations, are discussed.     

Salivary gland hypertrophy virus of house flies in Denmark: prevalence, host range, and comparison with a Florida isolate

House flies (Musca domestica) infected with Musca domestica salivary gland hypertrophy virus (MdSGHV) were found in fly populations collected from 12 out of 18 Danish livestock farms that were surveyed in 2007 and 2008. Infection rates ranged from 0.5% to 5% and averaged 1.2%. None of the stable flies (Stomoxys calcitrans), rat-tail maggot flies (Eristalis tenax) or yellow dung flies (Scathophaga stercoraria) collected from MdSGHV-positive farms displayed characteristic salivary gland hypertrophy (SGH). In laboratory transmission tests, SGH symptoms were not observed in stable flies, flesh flies (Sarcophaga bullata), black dump flies (Hydrotaea aenescens), or face flies (Musca autumnalis) that were injected with MdSGHV from Danish house flies. However, in two species (stable fly and black dump fly), virus injection resulted in suppression of ovarian development similar to that observed in infected house flies, and injection of house flies with homogenates prepared from the salivary glands or ovaries of these species resulted in MdSGHV infection of the challenged house flies. Mortality of virus-injected stable flies was the highest among the five species tested. Virulence of Danish and Florida isolates of MdSGHV was similar with three virus delivery protocols, as a liquid food bait (in sucrose, milk, or blood), sprayed onto the flies in a Potter spray tower, or by immersiion in a crude homogenate of infected house flies. The most effective delivery system was immersion in a homogenate of ten infected flies/ml of water, resulting in 56.2% and 49.6% infection of the house flies challenged with the Danish and Florida strains, respectively.     

Ivermectin residues in cattle dung impair insect-mediated dung removal but not organic matter decomposition

1. Veterinary parasiticide residues in livestock dung have been repeatedly shown to negatively affect the abundance and diversity of dung-associated insects. While these losses are concerning from a conservation perspective, they can also translate to impairment of ecosystem functions in agricultural landscapes (e.g. nutrient cycling, primary productivity and greenhouse gas mitigation). 2. Most research focusing on decomposition-related ecosystem functioning has focused on dung beetles (Coleoptera: Scarabaeoidea) while other insects, particularly flies, have received comparatively less attention. 3. Using mesocosms, this study manipulated the insect groups colonising cow dung (beetles only, flies only, beetles and flies together, and an insect-free control). Half of the insects were exposed to 1 mg kg⁻¹ ivermectin in dung, while the other half were exposed to ivermectin-free dung. Dung decomposition (mass of organic matter lost) and dung removal (change in the dry mass of the dung pat attributed to both dung decomposition and burial) were measured. 4. Comparison of beetles and flies in ivermectin-free dung showed that beetles removed nearly twice as much dung as did flies. Comparison of dung removal across all treatments showed that ivermectin residues significantly reduced dung removal provided by beetles by 47% and dung removal provided by beetles and flies together by 32%. 5. Organic matter decomposition was not significantly affected by insect colonists or by the presence of ivermectin, indicating that organic matter decomposition can occur independently of insect activity and chemical perturbations.     

Selection for resistance to imidacloprid in the house fly (Diptera: Muscidae)

The house fly, Musca domestica L. (Diptera: Muscidae), continues to be a primary pest of livestock facilities worldwide. This pest also has shown a propensity for pesticide resistance development when under high selection pressures. In this study the house fly strain FDm was created by a 20% contribution from each of five colonies collected from dairies in Florida with known imidacloprid resistance. The FDm strain was used to evaluate the level ofimidacloprid resistance after five selections near the LC70 value of each selected generation. Overall, the mean selection mortality was 72.7, with males being considerably more susceptible than females. The unselected (F0) FDm strain showed considerable susceptibility to imidacloprid after its creation, compared with the five parental strains. Between 9500 and 14,000 virgin house flies were used in each selection. After the fifth and final selection, a 331-fold increase in imidacloprid resistance at the LC70 was observed over the parental FDm strain. In parallel studies, the FDm strain showed increasing tolerance of the commercial imidacloprid product QuickBayt. These results suggest that livestock producers should use caution when choosing pesticides and consider rotating fly baits, as is encouraged with other pesticide treatment regimes on farms.     

Bacterial genome sequencing tracks the housefly-associated dispersal of fluoroquinolone- and cephalosporin-resistant Escherichia coli from a pig farm

The regular use of antimicrobials in livestock production selects for antimicrobial resistance. The potential impact of this practice on human health needs to be studied in more detail, including the role of the environment for the persistence and transmission of antimicrobial-resistant bacteria. During an investigation of a pig farm and its surroundings in Brandenburg, Germany, we detected abundant cephalosporin- and fluoroquinolone-resistant Escherichia coli in pig faeces, sedimented dust, and house flies (Musca domestica). Genome sequencing of E. coli isolates revealed large phylogenetic diversity and plasmid-borne extended-spectrum beta lactamase (ESBL) genes CTX-M-1 in multiple strains. [Correction added on 28 February 2023, after first online publication: In the preceding sentence, ‘and TEM-1’ was previously included but has been deleted in this version.] Close genomic relationships indicated frequent transmission of antimicrobial-resistant E. coli between pigs from different herds and across buildings of the farm and suggested dust and flies as vectors for dissemination of faecal pathogens. Strikingly, we repeatedly recovered E. coli from flies collected up to 2 km away from the source, whose genome sequences were identical or closely related to those from pig faeces isolates, indicating the fly-associated transport of diverse ESBL-producing E. coli from the pig farm into urban habitation areas. The observed proximity of contaminated flies to human households poses a risk of transmission of antimicrobial-resistant enteric pathogens from livestock to man.     

Survey of insect growth regulator (IGR) resistance in house flies (Musca domestica L.) from southwestern Turkey

Insect growth regulators (IGRs) are currently the fastest‐growing class of insecticides, and in Turkey these products represent a new approach to pest control. In recent years, several IGRs were also registered for the control of the house fly, Musca domestica L. (Diptera: Muscidae), in Turkey. A field survey was conducted in the summers of 2006 and 2007 to evaluate resistance to some agriculturally and medically used IGRs on house flies from livestock farms and garbage dumps in the greenhouse production areas (Merkez, Kumluca, Manavgat, and Serik) of Antalya province (Southwestern Turkey). The results of larval feeding assay with technical diflubenzuron, methoprene, novaluron, pyripoxyfen, and triflumuron indicate that low levels (RF<10‐fold) of resistance to the IGRs exist in the house fly populations from Antalya province. Exceptions, however, were two populations, Guzoren and Toptas, from the Kumluca area which showed moderate resistance to diflubenzuron with 11.8‐fold in 2006 and 13.2‐fold in 2007, respectively. We found substantial variation in susceptibility of field‐collected house fly populations from year to year and from product to product. We generally observed an increase in resistance at many localities sampled from 2006 to 2007. The implications of these results to the future use of IGRs for house fly control are discussed. It will be critically important to continue monitoring efforts so that appropriate steps can be taken if resistance levels start to increase.     

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.     

Distribution and Density of Tsetse Flies (Glossinidae: Diptera) at the Game/People/Livestock Interface of the Nkhotakota Game Reserve Human Sleeping Sickness Focus in Malawi

In large parts sub-Saharan Africa, tsetse flies, the vectors of African human or animal trypanosomiasis, are, or will in the foreseeable future, be confined to protected areas such as game or national parks. Challenge of people and livestock is likely to occur at the game/livestock/people interface of such infested areas. Since tsetse control in protected areas is difficult, management of trypanosomiasis in people and/or livestock requires a good understanding of tsetse population dynamics along such interfaces. The Nkhotakota Game Reserve, an important focus of human trypanosomiasis in Malawi, is a tsetse-infested protected area surrounded by a virtually tsetse-free zone. The abundance of tsetse (Glossina morsitans morsitans) along the interface, within and outside the game reserve, was monitored over 15 months using epsilon traps. A land cover map described the vegetation surrounding the traps. Few flies were captured outside the reserve. Inside, the abundance of tsetse at the interface was low but increased away from the boundary. This uneven distribution of tsetse inside the reserve is attributed to the uneven distribution of wildlife, the main host of tsetse, being concentrated deeper inside the reserve. Challenge of people and livestock at the interface is thus expected to be low, and cases of trypanosomiasis are likely due to people and/or livestock entering the reserve. Effective control of trypanosomiasis in people and livestock could be achieved by increasing the awareness among people of dangers associated with entering the reserve.     

Chromosomal regions controlling resistance to gastro-intestinal nematode infections in mice

Quantitative trait loci (QTL) associated with resistance to an intestinal worm in a well-defined murine model are described. These have been identified in an F(2) population derived from resistant (SWR) and susceptible (CBA) parental mouse strains infected with the gastro-intestinal nematode parasite Heligmosomoides polygyrus. Seven QTL located on six chromosomes are described, associated with components of the complex host response and the differential regulation of parasite survival and reproduction. The combined additive effects of the five significant QTL associated with worm survival (total worm count at necropsy) account for about 60% of the difference in worm count between the parental lines. The dominance effect for these five QTL are all in the direction of resistance, supporting the heterosis for resistance established from the mean worm count for the F(2) line relative to the parental lines. It is now possible to identify the comparative chromosomal regions of these QTL in livestock and humans and to consider the possibility of future improved control strategies. These may include breeding of resistant or tolerant livestock, development of vaccines, or identification of new anthelmintic drugs.