The origin of winter populations of infective larvae of nematode parasites of cattle in a Mediterranean-type climatic environment

An experiment to determine the origin of populations of infective larvae of cattle nematode parasites on pasture during winter was conducted in south-west Western Australia. Six pasture plots were contaminated with worm eggs by grazing worm-infected cattle for periods of a month during summer and autumn. Each plot was contaminated at a different time from the rest. The levels of infective larvae were determined by counting the worm burdens of tracer calves which test-grazed the plots the following winter.Tracer calves which grazed the plots contaminated during summer acquired few worms, whereas those that grazed the plots contaminated during autumn acquired many worms. It was concluded that the hot, dry conditions prevailing during summer and early autumn prevented the development of eggs or survival of larvae in dung pats or free on pasture. In this environment, a programme of worm control which relied on administration of anthelmintic to grazing cattle to prevent autumn contamination of pasture would be most likely to succeed if the first treatment was given in early autumn.     

Response of young cattle to anthelmintic treatment during autumn in a Mediterranean-type climatic environment

A worm control programme in which heifers were treated with anthelmintic on three occasions during autumn, was tested in the Mediterranean-type climatic environment of south-west Western Australia. The experiment aimed to determine if the treatments would prevent the heifers contaminating their pastures with worm eggs during autumn, thereby improving their growth performances the following winter. An attempt was made to measure the availability of infective larvae of abomasal worms on the heifers' pastures during early winter by counting the worms in steers, previously of low worm status, that grazed with the heifers from late autumn until the start of mating in mid-winter.

The anthelmintic treatments reduced the contamination of pasture for most of autumn. The treated heifers that grazed these pastures grew faster, and by the start of mating two months after the last treatment were about 22 kg heavier, than untreated heifers grazing contaminated pasture. At the end of mating six weeks later the difference was 45 kg in favour of the treated heifers. At this time half the heifers grazing contaminated pasture were treated with anthelmintic. The following month these heifers grew faster than those left untreated, but by late November they had not attained the wieght of the heifers grazing uncontaminated pasture.

The heifers that grazed uncontaminated pasture produced more calves the following autumn than did those grazing contaminated pasture. The abomasal worm counts of the steers, with a mean of about 46 000 worms, failed to reveal any difference between treatments in the availability of larvae of abomasal worms on pasture. However, it was concluded that the treatments probably exerted their effect on growth rates by reducing the number of infective larvae ingested by heifers grazing the uncontaminated pasture during winter.     

Nematodirus battus infection in calves

In studies on the control of parasitic gastroenteritis in calves and sheep, involving an annual rotation of pastures grazed by these host species, it was shown that young cattle could play an important role in the epidemiology of Nematodirus battus, a species usually regarded as a parasite of lambs. Thus, young cattle readily acquired heavy burdens of N battus in spring and the contamination of pastures with eggs from these infections resulted in significant populations of larvae on the herbage, which were infective to both calves and lambs grazed on these pastures in the following year. Although the majority of the N battus eggs hatched in the spring, some hatched in the autumn. The calves developed a strong immunity to N battus during the grazing season as demonstrated by the absence of worms at necropsy in the autumn, despite the presence of infective larvae on the pasture.     

Dynamics of pasture contamination by gastrointestinal nematodes of cattle under extensive management systems: proposal for strategic control.

An epidemiological study of gastrointestinal nematode parasitism in beef cattle in mountainous areas of Spain was performed. The dynamics of contamination with gastrointestinal nematode larvae of Pyrenean pastures was studied over four years at five areas at different altitudes (900 m to 2100 m), grazed by animals according to traditional systems of beef cattle in mountainous areas. Grass samples were taken every two weeks and larval differentiation was performed. Worm egg counts of grazing animals were assessed in cows, heifers and calves. A consistent seasonal pattern of infective larvae on pasture through the study was observed. In hay meadows, located below 1000 m, infective larvae were found from the end of October until June of the following year. At higher altitudes (1200-2100 m), a bimodal pattern of pasture larvae contamination was observed with increases in late spring (March-June) and in late autumn (September-November). Ostertagia spp., Cooperia spp., Trichostrongylus spp., Oesophagostomum spp., and Nematodirus spp. were found, with Ostertagia spp. being the most frequently found, followed by Cooperia spp. The highest increase of larval contamination in autumn coincided with the grazing of animals in hay meadows. This elevated autumn larval population had a very important epidemiological role because these larvae remained as overwintered larvae until the following grazing season, starting the cycle of contamination of the animals.     

Goats and sheep as hosts for some common cattle trichostrongylids

Recently weaned kid goats, lambs and calves, raised under worm-free conditions, were dosed with infective larvae of Ostertagia spp. and Cooperia spp. derived from naturally infected cattle. Each animal received a similar mixed dose of 60 000 infective larvae. Postmortem examination four weeks after infection revealed that the overall establishment of Ostertagia spp. in the kids was markedly less than in either the calves or the lambs, while the establishment in the lambs was marginally less than in the calves. Species composition of the Ostertagia burdens at necropsy differed widely between the host species. In the calves, O. ostertagi was the dominant species (57.8–73.4%), whereas in the lambs it was less successful (12.3–51.4%), and in the goats it was present in only very small numbers (0.0–3.8%). The proportion of O. crimensis present in the Ostertagia burden of each of the lost species was inversely related to the level of O. ostertagi present. A similar level of establishment of Cooperia spp. occurred in both lambs and calves, but no Cooperia established in the goats.     

SEASONAL AVAILABILITY OF NEMATODE LARVAE ON PASTURES GRAZED BY CATTLE IN NEW SOUTH WALES

Trends in the numbers of infective nematode larvae on pasture plots contaminated by cattle at different seasons of the year were defined in 3 different climatic regions. The main nematodes were Ostertagia ostertagi, Trichostrongylus spp, Haemonchus spp and Cooperia spp.
On the North Coast of New South Wales with a sub-tropical climate, the numbers of infective larvae of all 4 nematodes rose rapidly to peak levels soon after each seasonal period of contamination began, then fell quickly within a few months. On the Central Coast of New South Wales, the trends were similar to those on the North Coast, except that the larvae persisted on the pasture for a much longer time. On the Northern Tablelands of New South Wales, where temperatures were much colder than on the coast, larval development was slower and major peaks of larval availability did not occur until early spring. These different seasonal trends in each region were considered to be related to the climatic differences between the regions.
On pastures which were contaminated continuously, larval numbers reached maximum levels in mid-winter on the Central Coast and in early spring on the Northern Tablelands. It was concluded that the majority of these larvae were derived from the contamination of pastures in autumn and winter. Subsequently in summer, a rapid dying out of larvae was observed in all the regions, probably due to the effect of hotter weather.
The studies suggest that a reduction in the contamination of pasture with nematode eggs in autumn and winter could result in pastures carrying fewer larvae and thus form the basis of effective worm control programs for cattle.     

Evaluation of the morantel sustained release bolus in cows and calves

The efficacy of the morantel sustained release bolus (MSRB) in controlling gastrointestinal parasitism in beef cattle was assessed during the 1982 spring-autumn grazing season. Forty-eight cows and their calves were allotted to three equal groups. One group (T-1) served as a nonmedicated control group. One MSRB was administered to each calf of the T-2 group, and to each cow and calf of the T-3 group at the beginning of the study. The efficacy of the bolus was assessed by comparison of weight gain performance and parasitological data (fecal worm egg counts, herbage larval counts, worm counts from tracer and principal trial calves, and plasma pepsinogen level determinations). Though not statistically significant, treated calves from Group T-2 had a numerical mean weight gain advantage of 2.6 kg, and those from Group T-3 of 4.7 kg, over control calves. Average daily gains (ADG) for the three groups of calves were 0.69, 0.72, and 0.73 kg, respectively. Untreated cows from Group T-2 and treated cows from Group T-3 outperformed the control cows by 12.3 and 7.5 kg, respectively. Fecal worm egg counts from both groups of treated calves were significantly (P less than 0.01) lower than counts from control calves during the entire 169-day trial; notably, egg counts were reduced by 99% 28 days after MSRB administration to both groups of calves. There were no significant differences in the number of eggs counted from the three groups of cows, probably because of the very low numbers of eggs encountered. Mean total worm burdens of principal calves (six per group) necropsied at trial termination indicated a 91% (P less than 0.01) reduction in Group T-2 and an 87% reduction (P less than 0.01) in Group T-3. Worm-free tracer calves were introduced onto pastures every 28 days to monitor availability of infective larvae. The mean number of worms recovered at necropsy from tracer calves that grazed with control cattle increased as the season progressed. However, the numbers of parasites recovered each month from mid-August through mid-October from tracers that grazed pastures with treated cattle were lower (P less than 0.05) than those levels displayed at trial initiation. In addition, the mean numbers of worms from treated group tracers were lower than from the controls for each necropsy period.(ABSTRACT TRUNCATED AT 400 WORDS)     

On the Natural Seeding of Marshland Pastures with Bovine Gastrointestinal Parasites

By placing parasite-free calves in paddocks grazed by infected animals for 18 day periods at various times during the previous season it was shown that eggs of Ostertagia ostertagi, Cooperia oncophora and Nematodirus helvetianus deposited on pastures from early July to October of one year were able to survive in the Maritime area of Canada over winter either as eggs and/or larvae and contribute to residual infections on these pastures the following spring. The greatest deposition and/or survival of those eggs that were shed on pasture occurred in August and September for Cooperia and in September and October for Ostertagia. Greatest deposition of Nematodirus occurred in July and August and relatively few Nematodirus eggs shed in late September or early October were infective early in the next season.

The number of generations of worms per year was low, ranging from one to two or perhaps three per year depending on the species. There was a delay in the maturation of many worm eggs.

Residual overwintering infections play a significant role in the establishment of initial infections each summer in susceptible stock. These animals recontaminate the pastures leading to the subsequent development of large numbers of infective larvae by late summer and autumn.

     

Strategic use of anthelmintics to prevent parasitic gastroenteritis in cow-calf herds in California

On the basis of the hypothesis that the peak numbers of infective nematode third-stage larvae (L3) on herbage in winter months results from fall contamination of pastures, 2 methods to reduce fall contamination were tested. In trial 1, morantal sustained-release boluses were administered to 15 fall-calving cows on Sept 7, 1982. Fifteen untreated cows (controls) were placed on separate pastures. Numbers of L3 on herbage during the winter and spring were assessed by use of worm-free tracer calves. In trial 2, 19 cattle due to calve in the fall were administered 200 micrograms of invermectin/kg of body weight, SC, on Sept 2, 1983. Also, 17 cattle similarly were given a placebo injection and served as control animals. Treated cattle were placed on the pasture used by control cattle in trial 1 and control cattle on the pasture used by treated cattle in trial 1. Worm-free tracer calves were again used to assess numbers of L3 on herbage. In trial 1, tracer calves grazing the control animal pasture from January 14 to 28 acquired 37 times as many nematodes as did those grazing the treated animal pasture. In trial 2, the greatest difference observed was a 10-fold increase of nematodes in calves grazing control animal pastures, compared with worm numbers in tracer calves grazing the treated animal pasture.     

Observations on the epidemiology of Dictyocaulus viviparus in north west England

A five year ley pasture was used as a source of natural infection with Dictyocaulus viviparus for cattle in anthelmintic trials. Pasture larval counts, faecal larval counts of permanently grazing calves and lungworm burdens harboured by tracer calves were monitored in three grazing seasons to assess the pattern of infection. Carrier calves were introduced at the beginning of the grazing season in the first two years of the study but not in the third. In the fourth year the pasture was subdivided into two paddocks where overwintered infection with and without carrier infection were compared. A control paddock exposed to carrier infection but no overwintered infection was also monitored. Pasture larvae survived the winter but carrier infection appeared to make a larger contribution to pasture larval counts and the onset of parasitic bronchitis in susceptible calves. In the absence of grazing cattle at the end of the grazing season the concentration of D viviparus larvae on the herbage fell rapidly to undetectable levels. Discrepancies between contamination of herbage by infective D viviparus larvae and infectivity of pasture for susceptible cattle occurred in all years but were particularly marked on the third year when natural immunity appeared to influence the number of lungworms accumulating in tracer calves. Failure to recover lung worms from tracer calves cannot be regarded as an accurate indication of lungworm free pasture. In the first three years the proportion of the lungworm population which was inhibited in tracer calves was higher early and late in the grazing season and negligible in mid season. This suggests that a predisposition to inhibition in larvae which have overwintered on pasture may influence the time of onset of parasitic bronchitis in the next grazing season, but results from the fourth year did not support this hypothesis.     

Efficacy of the morantel sustained-release bolus in grazing cattle in North America

The efficacy of using a bolus containing morantel in a sustained-release preparation for controlling naturally acquired gastrointestinal parasitic infections in weaned calves and yearling cattle was investigated during the 1982 grazing season at selected sites in the United States and Canada. According to a common trial design under various climatic and management conditions, 10 field trials were conducted with the bolus. At the time of spring turnout, a bolus was administered to each calf or yearling in the treated group. Then, treated and control cattle grazed separate but equal areas of divided pasture(s). The epidemiologic pattern of parasitic gastroenteritis in control animals and the effect of treatment on this pattern was determined in each trial. Safety and practicality of use of the bolus also were established. When compared with untreated cattle (control), those given the bolus deposited significantly (P less than 0.05) fewer worm eggs (89% reduction) during the first 90 days of the grazing season, as well as significantly fewer (P less than 0.05) worm eggs (84% reduction) during the entire grazing season. Consequently, during the second half of the grazing season, larval populations on treated pastures remained significantly (P less than 0.05) lower (66% reduction), compared with numbers of larvae found on control pastures. For pastures grazed by treated and control cattle at trial initiation, mean worm counts recovered from tracer calves were equal, indicating comparable pasture contamination at the beginning of the grazing season.(ABSTRACT TRUNCATED AT 250 WORDS)     

Proceedings in life sciences: Fish diseases : Editor W. Ahne, Third Cooperative Programme of Research on Aquaculture session, 1980, 120 figs., x + 252 pp., DM 89.00 (cloth bound), approx. US $52.60, Springer-Verlag, Berlin-Heidelberg-New York, ISBN 3-540-10406-2

The effect of repeated intradermal injections of small quantities of adult worm extract on the resistance of calves to Cooperia spp. was investigated.A group of five calves injected intradermally was less resistant than a similar group inoculated twice with infective larvae, but more resistant than a group of five calves kept as worm-free controls. The resistance was reflected by worm numbers, worm lengths, inhibition percentage of L4 larvae, number of eggs per female worm, egg production and by antibody titres.     

SEASONAL CHANGES IN THE STRUCTURE OF NEMATODE POPULATIONS OF CATTLE IN NEW SOUTH WALES IN RELATION TO INHIBITED LARVAL DEVELOPMENT

Pasture plots in 3 climatic regions were contaminated with worm eggs of Ostertagia ostertagi, Trichostrongylus axel, Haemonchus spp and Cooperia spp in the autumn, winter and spring. Successive pairs of parasite-free calves were grazed on the plots for 7 to 10 days at 4-week intervals and then killed for worm counts 14 days after their removal from pasture.
On the Northern Tablelands of New South Wales, irrespective of the season of pasture contamination, the degree of inhibition of O. ostertagi was low in winter and highest in spring. T. axei showed similar trends while Cooperia spp showed negligible inhibition.
On the North Coast of New South Wales, inhibited larvae accounted for a very small proportion of the O. ostertagi burdens, while in comparison T. axei showed a much greater degree of inhibition. Larval inhibition of Haemonchus spp occurred in autumn and early winter after which it did not occur. There was negligible inhibition in Cooperia spp.
On the Central Coast of New South Wales, there was little inhibition of O. ostertagi and none in T. axel . For Haemonchus spp, inhibited larvae were found mainly in autumn and winter. The numbers of inhibited Cooperia larvae were also highest in autumn and winter and were associated with large worm burdens.
The marked difference between the tablelands and coastal regions in the seasonal trends of inhibition of O. ostertagi was considered to be due to a difference in strains between the geographical regions. The possible effect of climatic factors on the inhibition-proneness of infective larvae on pasture is discussed for Ostertagia and other nematodes. The roles of host resistance and density-dependence are also discussed.     

Experimental Ostertagia spp. infection of sheep: Populations and lesions in previously worm-free and previously exposed animals

Sheep raised and maintained under worm-free conditions were subjected to one heavy exposure and several low-level exposures of infective Ostertagia spp. larvae. The infections were terminated by treatment with thiabendazole. These sheep and not previously exposed controls were then challenged with 400 000 or 20 000 Ostertagia spp. larvae and killed after a further 8 days. In the previously exposed sheep, worm numbers, rate of worm development, and pathological effect, were greatly reduced. No significant differences were seen between sheep aged 6 and 18 months at the time of challenge.     

The epidemiology of nematode and fluke infections in cattle in the Red River Delta in Vietnam

Over a period of 13 months, faecal samples were collected monthly from approximately 45 cattle over 3 months of age. Additionally, 74 calves of 1-2 months were sampled to determine the presence of Toxocara vitulorum eggs. Individual egg counts and infective strongyle larvae from pooled faecal samples were examined. Post-mortem worm counts were carried out on six groups of tracer calves (n=12) that had been kept for 4 weeks on pasture in and around the village studied. The following helminths were identified: T. vitulorum, Cooperia punctata, C. pectinata, C. oncophora, Oesophagostomum radiatum, Trichostrongylus axei, T. colubriformis, Haemonchus spp., Fasciola spp. and Paramphistomum spp. In 8% of the samples collected from young calves, individual egg counts for T. vitulorum were found indicative for pathogenic worm burdens. Strongyle egg counts and worm counts indicated that transmission is low without a distinct seasonality. In animals of 3-9 months old, a strongyle egg count peak can be demonstrated which at a higher age steadily and significantly decreased. In faecal cultures Cooperia spp. were most prominent in all age groups throughout the year with the exception of the period September-November when Haemonchus spp. and Oesophagostomum spp. were most prevalent. Fasciola spp. eggs were found in 22% of the collected faecal samples and the egg counts were low indicating that the intensity of Fasciola spp. infection is mild. Based on the present data, regular anthelmintic treatments seem not to be justified, except for a single treatment at the age of 2 weeks against toxocariosis.     

Relative importance of winter survival of larval nematodes in pasture and infected carrier calves in a study of parasitic gastroenteritis in calves.

A study was done in Maine to determine the relative importance of winter survival of nematode larvae in pasture and infected carrier animals as sources of infection for susceptible calves. Under the conditions of the experiment, it appeared that winter survivals in pasture of the infective stages of the genera Ostertagia, Cooperia, Nematodirus, and Trichostrongylus were of greater importance than carrier animals as sources of infection for susceptible calves. While animals in plots infected the previous summer and simultaneously allowed to graze alongside infected carrier animals did show more worms than those grazed only in infected plots, these differences were not statistically significant. Both groups had significantly (P greater than 0.01) more worms than did calves grazed only with carrier animals for the period of the experiment (8 weeks). It was also observed that carrier calves with low fecal egg counts (less than 200 eggs/g of feces) introduced in early spring to uncontaminated pasture could produce enough parasitic contamination by early fall to cause fulminating infections in susceptible calves grazing the pasture at the same time. Infected animals that survived clinical disease during their 1st summer developed a strong immunity which limited their acquisition of further infections when they were exposed to severe pasture contamination the following year.     

Dynamics of Ostertagia spp. and Cooperia oncophora in field-grazed cattle from weaning to 2 years old in New Zealand, with particular reference to arrested development

Gastrointestinal nematode parasite burdens were monitored in a herd of field-grazed cattle from weaning to 2 years old to allow observations to be made on the dynamics of burdens of arrested Ostertagia spp. and Cooperia oncophora. Arrested Ostertagia spp. accumulated in the herd over their first autumn and winter to reach a peak in late winter/spring (August-October). From October until late January they declined at a rate comparable to that at which they had accumulated. Numbers continued to decline at a slower rate until the following autumn (May) when they began to increase again. Observations on the decline of worm burdens in a group of animals which had been moved from pasture to worm-free conditions in October indicated that intake of infective larvae over late spring/summer contributed little to the number of arrested Ostertagia spp. in the herd. No clinical Type II ostertagiosis was observed in the field-grazed animals over the time of decline of the arrested worm burden, nor was there an increase in the number of adult Ostertagia spp. present. However, two of the animals removed from pasture to worm-free conditions developed Type II ostertagiosis approximately a month after the move. Worm burdens of successive groups of "tracers" grazed with the herd every 2 months indicated that there was a small seasonal increase in the propensity of Ostertagia spp. for arrested development. However, estimated accumulation rates of arrested larvae calculated from tracer worm burdens, suggest that this seasonal increase in propensity was not adequate in itself to account for the accumulation rate observed in "resident" animals of the herd. No comparable accumulation of arrested C. oncophora took place in resident animals over the first autumn and winter of the trial despite the fact that tracer worm burdens over this period indicated a marked seasonal increase in the propensity for arrested development by this species.     

Field efficacy of a morantel sustained release bolus for control of gastrointestinal nematodes in yearling steers

The efficacy of a morantel sustained release bolus (MSRB) for control of gastrointestinal nematodes in yearling steers was evaluated over a 6-month grazing period commencing on 26 March 1982. Three groups of 15 steers were allotted to the following treatments: Group 1 -- one MSRB at start of trial; Group 2 -- one therapeutic dose of thiabendazole at start of trial; Group 3 -- untreated control. The treatment groups were grazed separately. Parasite egg counts (EPG), herbage larval counts, pepsinogen levels and weight gains were monitored. Every other month, sets of 2 parasite-free tracer calves were placed in the pasture grazed by each treatment group and allowed to graze for 3 weeks before being subsequently necropsied for worm counts. At the end of the trial, 6 animals from each group were also necropsied for worm counts. The MSRB treatment resulted in significantly lower egg counts, fewer infective larvae on pasture, lower pepsinogen levels and lower worm burdens in tracer calves than was the case for the untreated group, but generally the levels were not significantly different from those associated with the thiabendazole treatment. The mean weight gain for the MSRB treated steers showed a significant advantage (70.9 lb) over the untreated animals, but was not significantly different from those which received thiabendazole. Total worm counts at the end of the trial were not different from any treatment group.     

Survival of infective larvae of nematode parasites of cattle during drought

In a study originally designed to determine the seasonal origin of the high levels of availability of nematode larvae to cattle in winter and spring, plots were serially contaminated with eggs of Ostertagia ostertagi and Cooperia oncophora by naturally-infected calves at monthly intervals from February 1980 to September 1980. The availability of infective larvae was monitored by monthly pasture sampling and larval recovery. Because of the intervention of a 15 month drought, recoveries of larvae from the pastures were very low until March 1981 (autumn in Australia) when large numbers of larvae appeared on pastures contaminated in the preceding spring. Examination of dry dung pats at that time showed that significant numbers of larvae were present in pats deposited up to a year previously, and particularly in pats deposited in May, August and September. Following the resumption of normal rainfall in May 1981, larval numbers in pats rapidly declined and concentrations of larvae on the pastures increased to extremely high levels. It is suggested that survival of infective larvae in dry dung pats was enhanced by the drought, with implications for control of nematode infections of cattle, particularly in winter rainfall environments.     

The efficacy of eprinomectin extended-release injection against naturally acquired nematode parasites of cattle,with special regard to inhibited fourth-stage Ostertagia larvae

The efficacy of eprinomectin in an extended-release injection (ERI) formulation in the treatment of cattle harboring naturally acquired nematode populations (including inhibited nematodes) was evaluated. Five studies were conducted under a similar protocol in the USA, the UK, and in Germany. All study animals were infected by grazing naturally contaminated pastures. The adequacy of pasture infectivity was confirmed by examining tracer calves prior to allocation and treatment of the study animals. The cattle were of various breeds or crosses, weighing 79–491 kg, and aged approximately 6–15 months. In each study, 20 animals were infected by grazing, and then removed from pasture and housed in a manner to preclude further nematode infections for 8–16 days until treatment. Animals were blocked based on descending pre-treatment body weight and randomly allocated to one of two treatments: ERI vehicle (control) at 1 mL/50 kg body weight or eprinomectin 5% (w/v) ERI at 1 mL/50 kg body weight (1.0 mg eprinomectin/kg). Treatments were administered once on Day 0 by subcutaneous injection in front of the shoulder. For parasite recovery and count, all study animals were humanely euthanized 14/15 days after treatment. Cattle treated with eprinomectin ERI had significantly (p < 0.05) fewer of the following nematodes than the controls with overall reduction of parasite counts of ≥94%: adult Dictyocaulus viviparus, Capillaria spp., Cooperia oncophora, Cooperia pectinata, Cooperia punctata, Cooperia surnabada, Haemonchus placei, Nematodirus helvetianus, Oesophagostomum radiatum, Ostertagia lyrata, Ostertagia ostertagi, Trichostrongylus axei, Trichostrongylus colubriformis, Trichuris discolor, Trichuris skrjabini, and Trichuris spp.; developing fourth-stage larvae of Ostertagia spp. and Trichostrongylus spp.; and inhibited fourth-stage larvae of Cooperia spp., Haemonchus spp., Nematodirus spp., Oesophagostomum spp., Ostertagia spp., and Trichostrongylus spp.