Cuticular ridge patterns applied for identifying adult female worms of Haemonchus species with various vulvar morphological types in infected native goats in Laos.

Species of the blood sucking nematode Haemonchus are a main problem in the small ruminant industry worldwide. Haemonchus worms were taken from 68 infected native goats slaughtered in three provinces of Laos in June and July 2019. Cuticular ridge patterns were used for the first time to identify adult female Haemonchus spp. and their vulvar morphs were characterized. The results showed that the variations in vulvar morphology of female Haemonchus spp. presented a knobbed morph as the dominant morphotype and predominant linguiform B subtype was also detected. In total, 270 selected female worms from each vulvar morph type were examined based on their cuticular ridge patterns in cross sections at positions of the esophageal-intestinal junction (EI), the 4 mm region from the anterior end (4 mm), and the mid-body (MB). Only Haemonchus contortus was identified and most worms had constant numbers of ridges at EI, 4 mm, and MB, namely 30, 26, and 22 ridges, respectively, accounting for 99.26%, 97.41%, and 97.04%, respectively, of worms detected, while the lowest variation in the number of ridges was at region EI which is recommended as the single best position. Based on synlophe and ITS2 sequence analysis, it was assumed that H. contortus might dominate in the sample areas with the possible numbers of ridges of H. contortus females in the ranges 29-30, 25-27, and 21-23 for positions EI, 4 mm, and MB, respectively. The cuticular ridge pattern was a useful character for identifying female Haemonchus species in this study and could be utilized as an affordable alternative method for epidemiological studies and as part of parasite control management in native goats of Laos.

Bacterial profiling of Haemonchus contortus gut microbiome infecting Dohne Merino sheep in South Africa.

A metagenomic approach was used to study the gut microbiome of Haemonchus contortus field strains and that of its predilection site, the abomasum of Dohne Merino sheep. The abomasum contents and H. contortus were collected from 10 naturally infected Dohne Merino sheep. The H. contortus specimens were classified and sexually differentiated using morphometric characters and was further confirmed through molecular identification. We investigated differences and similarities between the bacterial composition of the adult male and female H. contortus gut microbiomes, which were both dominated by bacteria from the Escherichia, Shigella, Vibrio and Halomonas genera. Major abundance variations were identified between the shared adult male and female H. contortus microbiomes. The results also revealed that Succiniclasticum, Rikenellaceae RC9 gut group and Candidatus Saccharimonas were the predominant genera in the Dohne Merino abomasum. This study provides insight into the highly diverse bacterial composition of the H. contortus gut microbiome and the Dohne Merino abomasum which needs to be studied further to explore the complex interactions of different gastrointestinal nematode microbiomes with the host.

Ovine haemonchosis: a review.

Sheep farming is the backbone of a rural economy in developing countries, and haemonchosis is a major impediment in the way of its progress. Haemonchus contortus (H. contortus) infection persists all over the world particularly in the tropical and sub-tropical regions. Various review articles have been published to substantially cover one or more aspects of its morphology, prevalence, pathogenesis, symptoms, diagnosis, immune response, drug resistance, treatment, and control measure. The objective of this paper is to briefly review past and present information available in the aforementioned areas in one place to enable the readers to fully understand the problem from a broader perspective. H. contortus parasite harbours in abomasum of affected animal and feeds on its blood, producing mild to severe symptoms and even death in acute form. The parasite thus inflicts heavy production losses and is of economic importance. H. contortus has developed diverse characters over the years leading to limited success in the production of vaccines. Indiscriminate use of the anthelmintics has produced drug resistance against almost all conventional products. Efficacy of medicinal plants and non-conventional chemicals has been reported under controlled experiments; however, research on their adverse effects on growth and fertility is yet to be studied. Research on molecular tools for identification and introduction of resistant genes into the flock is also underway but still a long journey to find its field application. Crossbreeding may compromise the production traits of the existing flock. In given circumstances, a targeted selective treatment approach along with selective breeding, culling of more susceptible animals, and maintaining a good body condition score through the provision of a balanced diet remains a workable strategy to control haemonchosis in sheep.

Morphological identification of adult male Haemonchus species in goats from Thailand and Lao PDR.

Haemonchus spp. or barber's pole worms are one of the most highly pathogenic nematodes of ruminants causing economic losses in livestock worldwide. The current study was a first attempt to identify Haemonchus spp. from goats in Thailand and Lao PDR. Utilizing the inexpensive tools of the discriminant function (DF) combined with synlophe patterns is fundamental for understanding their epidemiological aspects. In total, 255 randomly chosen adult male Haemonchus worms from goats in various areas in each country were identified individually. For both these countries, about 94% based on the DF values, and 99%, 98%, and 97% based on synlophe patterns in the region of the esophageal intestinal junction (EI), 4 mm from the anterior end, and at both these positions, respectively, were identified as H. contortus. Other identified specimens defined as H. placei and hybrids as well as unclassified species based on synlophe patterns were proved using polymerase chain reaction (PCR); this also included some randomly chosen H. contortus by DF and synlophe patterns. All those specimens were confirmed as H. contortus being strongly supported by some genetic evidences and UPGMA analysis. Thus, it was assumed that all specimens in the current study were H. contortus. The morphological differences of this predominant species (H. contortus) in goats between the two countries were: body length, gubernaculum length, and left spicule barb length, while almost all characters of male worms individually measured appeared to overlap, mostly in H. contortus and H. placei, which may lead to misclassification. Therefore, using the DF along with synlophe patterns can assist in increasing the accuracy of Haemonchus spp. identification from goats in some areas where funding is limited, particularly in Lao PDR. The present results revealed that synlophe patterns in the EI region seemed to be promising for the identification of Haemonchus spp., while molecular techniques are also required to address ambiguous identification with some specimens.

Investigating the function and possible biological role of an acetylcholine-gated chloride channel subunit (ACC-1) from the parasitic nematode Haemonchus contortus.

The cys-loop superfamily of ligand-gated ion channels are well recognized as important drug targets for many invertebrate specific compounds. With the rise in resistance seen worldwide to existing anthelmintics, novel drug targets must be identified so new treatments can be developed. The acetylcholine-gated chloride channel (ACC) family is a unique family of cholinergic receptors that have been shown, using Caenorhabditis elegans as a model, to have potential as anti-parasitic drug targets. However, there is little known about the function of these receptors in parasitic nematodes. Here, we have identified an acc gene (hco-acc-1) from the sheep parasitic nematode Haemonchus contortus. While similar in sequence to the previously characterized C. elegans ACC-1 receptor, Hco-ACC-1 does not form a functional homomeric channel in Xenopus oocytes. Instead, co-expression of Hco-ACC-1 with a previously characterized subunit Hco-ACC-2 produced a functional heteromeric channel which was 3x more sensitive to acetylcholine compared to the Hco-ACC-2 homomeric channel. We have also found that Hco-ACC-1 can be functionally expressed in C. elegans. Overexpression of both cel-acc-1 and hco-acc-1 in both C. elegans N2 and acc-1 null mutants decreased the time for worms to initiate reversal avoidance to octanol. Moreover, antibodies were generated against the Hco-ACC-1 protein for use in immunolocalization studies. Hco-ACC-1 consistently localized to the anterior half of the pharynx, specifically in pharyngeal muscle tissue in H. contortus. On the other hand, expression of Hco-ACC-1 in C. elegans was restricted to neuronal tissue. Overall, this research has provided new insight into the potential role of ACC receptors in parasitic nematodes.

Anthelmintic resistance in goat herds-In vivo versus in vitro detection methods.

Anthelmintic resistance (AR) is a serious threat to animal health and has a major economic impact worldwide due to production and financial losses. The aim of this study was to determine the occurrence of AR on 30 goat farms in Slovakia during the pasturing seasons and to compare three widely used in vitro and in vivo methods for detecting AR in field conditions. A three-year survey was conducted during the pasturing seasons of 2014-2016. Goats on each farm were split into treated and control groups and were treated by recommended (5 mg/kg body weight) and double doses (10 mg/kg b.w.) of albendazole. Comparisons between percent reduction in a faecal egg count reduction test (FECRT) and an egg hatch test (EHT) and the presence of L3 larvae in a larval development test (LDT) using resistant concentrations of benzimidazole (BZ) were monitored after treatment. The FECRT indicated percent reductions of 69.2-86.2% for the single dose and of 36.3-45.4% for the double dose. The EHT indicated that all farms had BZ-resistant nematodes. Low (<15% hatching) and high (>15% hatching) levels of resistance were detected on 13 and 17 farms, respectively. The LDT failed to detect resistant larvae on seven farms but detected low and high levels of resistance on seven and 14 farms, respectively. The data indicate a moderate correlation between in vitro and in vivo tests for detecting BZ resistance among the 30 goat farms. The hatching detected by the EHT and the presence of L3 larvae by the LDT at resistant BZ concentrations provided reasonable identification of low levels of resistance in the parasite populations, but the use of a double dose for a treatment may underestimate the real occurrence of low levels of resistant parasites on goat farms.

Ashworthius sidemi Schulz, 1933 and Haemonchus contortus (Rudolphi, 1803) in cervids in France: integrative approach for species identification.

Among gastro-intestinal nematodes, the blood-sucking worms belonging to the subfamily of Haemonchinae are considered to be of pathogenic and economic great importance, particularly in small ruminants. Haemonchus contortus, primary found in domestic ruminants and wild bovines (Mouflon, Chamois), is probably the most studied, but occurrence of Ashworthius sidemi has gradually increased over recent years, especially in Cervids and free roaming wild bovid as the European bison in eastern Europe, and some cases of co-infestation were recently observed on five Roe deer (Capreolus capreolus) and one Red deer (Cervus elaphus) in France. If the diagnosis is possible on the morphological features for adult worms for helminthologists, the identification on some stages (female, subadult, eggs and larvae) is difficult or impossible. Sequencing ND4 domain from the mitochondrial DNA of H. contortus and A. sidemi worms, we observed clearly two distinct clades, with an inter-specific divergence of 28.1%. Basing on this specific domain, a multiplex PCR-based method was developed: new primers were designed and used pooled in one mix PCR, producing amplicons of 454bp for H. contortus and 330bp for A. sidemi, allowing a trivial and an inexpensive taxonomic affiliation after migration. This multiplex PCR-based method was developed here to distinguish H. contortus and A. sidemi regardless their developmental stage, easy to use for highlighting co-infestation cases in both wild and domestic ruminants. It is a non-invasive approach appearing as a good diagnostic tool relevant to coprological cultures.

Explaining premunition with Kin selection using Haemonchus contortus.

Premunition is the state in a disease where an existing infection protects the host from reinfection with the same species. The cause of premunition is not clearly understood. In this study, we hypothesized that kin-selection might be a contributing factor in premunition. To test this theory, sheep were infected either once with a linguiform or smooth vulval morphotype of Haemonchos contortus, twice with the same morphotype or twice with different morphotypes. All infections resulted in a similar number of adult parasites. However, there were differences in the morphotypes recovered providing potential evidence of kin selection. Negative interference competition might also contribute to the reduction of the incoming population. Allelopathic or physical interactions between the parasites may be the mechanism behind the observed phenomena.

Host-specificity and morphometrics of female Haemonchus contortus, H. placei and H. similis (Nematoda: Trichostrongylidae) in cattle and sheep from shared pastures in São Paulo State, Brazil.

To better evaluate the usefulness of female Haemonchus specimens for specific identification, we undertook a detailed analysis of the morphology of a collection of worms obtained from cattle and sheep in shared pastures. Based on the results, we also more precisely evaluated the host-specificity of Haemonchus contortus, H. placei and H. similis occurring sympatrically in a farm located in the western region of São Paulo State, Brazil. A synlophe analysis was employed to identify the three species of Haemonchus. In cattle, the predominant species was H. similis (90.9%), followed by H. placei (9.1%). With the exception of one H. placei specimen, only H. contortus was found in sheep. The longest body length was found for H. placei specimens, followed by H. contortus and then H. similis. It was possible to distinguish H. similis females from H. contortus and H. placei on the basis of vulval structure. The synlophe analysis proved to be very useful for identification of H. contortus, H. placei and H. similis in epidemiological studies involving different species of ruminants in the same pastures. The finding that H. placei and H. similis were adapted to cattle and that H. contortus was adapted to sheep also confirmed the high host-specificity of the three nematodes species.

Differentiation of Haemonchus placei from Haemonchus contortus by PCR and by morphometrics of adult parasites and third stage larvae.

Molecular and morphological methods were evaluated to distinguish between Haemonchus contortus and Haemonchus placei species. A total of 141 H. contortus and 89 H. placei male adult specimens collected from artificially infected lambs were identified individually by PCR analysis, using a species-specific primer pair. These PCR results were used as gold standard for Haemonchus spp. identification. Haemonchus placei presented higher mean spicule and barb lengths than H. contortus (P<0.05). However, some measurements overlapped. For this reason, a discriminate function did not allow the correct identification of 13 H. contortus and one H. placei specimen. The sheath tail length of the third stage larvae (L3), which comprises the distance between the tip of the larval tail and the end of the sheath tail, were measured. Only three of the 485 H. placei larvae (0.619%) had a sheath tail shorter than 85 µm, while only four of the 500 H. contortus larvae (0.8%) presented a sheath tail longer than 85 µm. The results indicated that 6.09% of the male adult specimens would be misclassified based on the discriminate function, while only 0.71% of infective larvae would be misclassified. Therefore, identification of L3 can be used as the first method to indicate the presence of H. placei and/or H. contortus in a population of domestic ruminants.

Haemonchus longistipes Railliet & Henry, 1909 (Nematoda, Trichostrongylidae) from the Egyptian dromedary, Camelus dromedarius (Artiodactyla: Camelidae), first identification on the basis of light and ultrastructural data.

Haemonchus longistipes is a gastrointestinal abomasal nematode which is one of the most prevalent and pathogenic parasites infesting the stomach of ruminants. On the basis of light and ultrastructural data, the objective of the present study was to introduce a first identification of the cameline haemonchosis caused by H. longistipes. Abomasa of 42 Egyptian camels Camelus dromedarius (Artiodactyla: Camelidae) were collected monthly from September 2013 to April 2014 from the main slaughter house of Cairo, Egypt. Adult male and female nematode worms were recovered from 26 (62%) specimens of the examined abomasa. The parasites were of yellow color; the body was filiform (slender) tapered towards the anterior end in male and towards both ends in female. Buccal capsules absent, the buccal cavity was small with a conspicuous dorsal lancet extended from dorsal wall. The cervical papillae were prominent and spine-like. The body length of the female worm was 16.6-20.5 (18.5 ± 0.3) mm. The anterior end to the cervical papillae was 3.19-4.30 (4.12 ± 0.5) mm. The vulva of the female had a linguiform process or flap, the tail is without a spine, and the anal pore at the posterior end of the body had a simple dorsal rim. The body of male was 10.4-14.7 (13.9 ± 2.0) mm in length. The male bursa had elongated lobes supported by long, slender rays. The small dorsal lobe was asymmetrical with Y-shaped dorsal rays. The spicules were long with a length of 0.52-0.54 (0.53 ± 0.05) mm, each provided with a small barb and pore near its extremity. Synlophe was bilaterally and dorsoventrally symmetrical; it extended from cephalic expansion over anterior 50% of prebursal or prevulvar body and consisted of a maximum of 42 ridges. The described species herein was compared with the three morphologically similar species Haemonchus mitchelli, Haemonchus okapiae, and H. longistipes with their synlophes consist of 42 ridges distributed over the anterior half of the body. These species can be separated by unique structural characteristics of their synlophes, spicules, and copulatory bursa. The most morphologically similar species to the recovered worm was H. longistipes. Also, some of the parameters with regard to morphology and morphometry of this parasite were described for the first time.

Morphological variation between isolates of the nematode Haemonchus contortus from sheep and goat populations in Malaysia and Yemen.

Haemonchus contortus is a highly pathogenic nematode parasite of sheep and goats. This work was conducted to investigate the population and host variations of the parasitic nematode H. contortus of sheep and goats from Malaysia and Yemen. Eight morphological characters were investigated, namely the total body length, cervical papillae, right spicule, left spicule, right barb, left barb, gubernaculum and cuticular ridge (synlophe) pattern. Statistical analysis showed the presence of morphological variation between populations of H. contortus from Malaysia and Yemen, with minor variation in the synlophe pattern of these isolates. Isolates from each country were grouped together in the scatterplots with no host isolation. Body, cervical papillae and spicule lengths were the most important characters that distinguished between populations of the two countries. This variation between Malaysia and Yemen may be attributed to geographical isolation and the possible presence of a different isolate of this worm in each country.

Morphological identification of parasitic nematode infective larvae of small ruminants and cattle: a practical lab guide.

In 2004, a new concept was introduced for simplifying identification of larvae of the common nematodes of cattle, sheep and goats that comprises estimates of the lengths of the sheath tail extensions of infective third-stage larvae (L3) of each genus and/or species to that of Trichostrongylus spp., instead of having to be dependent only on measurements in micrometre. For example, if the mean length of the sheath tail extension (the extension of the sheath caudad, beyond the caudal tip of the larva) of Trichostrongylus colubriformis and Trichostrongylus axei is assumed to be 'X', then that of Haemonchus contortus is 2.0-2.7 'X' - a difference that is not difficult to estimate. An additional new approach suggested now, particularly for L3 of species and/or genera difficult to differentiate (such as Chabertia ovina and Oesophagostomum columbianum), is to estimate the proportion of the larval sheath tail extension comprising a terminal thin, whip-like filament. For the experienced person, it is seldom necessary to measure more than one or two sheath tail extensions of L3 in a mixed culture, because the identity of most of the remaining L3 can thereafter be estimated in relation to those measured, without having to take further measurements. The aim of this article was to present the novel approach in the form of a working guide for routine use in the laboratory. To facilitate identification, figures and a separate organogram for each of small ruminants and cattle have been added to illustrate the distinguishing features of the common L3.

Why is it important to correctly identify Haemonchus species?

Parasitic gastroenteritis caused by Haemonchus spp. is a major cause of economic losses in the livestock industry because it impairs weight gain and increases mortality in cattle and small ruminants, especially in tropical and subtropical areas. The proper identification of the various species, as well as knowledge regarding the epidemiology of parasitic gastroenteritis, is essential for the establishment of sustainable strategies of parasite control. This review focuses on the use of easily applied, low-cost parasitological methods of identifying Haemonchus species on the basis of their morphology. In most studies carried out in Brazil, the distinctions between Haemonchus contortus and Haemonchus placei have not been considered. Many reports of H. contortus, particularly in cattle, might actually represent H. placei. The appropriate identification of species is therefore indispensable. In addition to the measurement of male spicules, new morphological characteristics, such as the synlophe, should be evaluated in order to differentiate between and among species. Measurements of infective larvae in fecal cultures can also indicate the identity of Haemonchus species. This approach can be quite useful in studies that do not involve animal sacrifice, such as studies of anthelmintic resistance based on the fecal egg count reduction test.

Why is it important to correctly identify Haemonchus species? / Por que é importante a identificação correta das espécies de Haemonchus?

Parasitic gastroenteritis caused by Haemonchus spp. is a major cause of economic losses in the livestock industry because it impairs weight gain and increases mortality in cattle and small ruminants, especially in tropical and subtropical areas. The proper identification of the various species, as well as knowledge regarding the epidemiology of parasitic gastroenteritis, is essential for the establishment of sustainable strategies of parasite control. This review focuses on the use of easily applied, low-cost parasitological methods of identifying Haemonchus species on the basis of their morphology. In most studies carried out in Brazil, the distinctions between Haemonchus contortus and Haemonchus placei have not been considered. Many reports of H. contortus, particularly in cattle, might actually represent H. placei. The appropriate identification of species is therefore indispensable. In addition to the measurement of male spicules, new morphological characteristics, such as the synlophe, should be evaluated in order to differentiate between and among species. Measurements of infective larvae in fecal cultures can also indicate the identity of Haemonchus species. This approach can be quite useful in studies that do not involve animal sacrifice, such as studies of anthelmintic resistance based on the fecal egg count reduction test.

Infecções por Haemonchus spp. são uma das principais causas de perda econômica nas criações de ruminantes devido à redução no ganho de peso e mortalidade de bovinos e pequenos ruminantes, especialmente em regiões com clima tropical e subtropical. A identificação precisa das diferentes espécies, bem como o conhecimento sobre a epidemiologia das gastroenterites parasitárias, são fundamentais para a elaboração de estratégias sustentáveis de profilaxia das parasitoses. Essa revisão tem por objetivo central, abordar os principais métodos parasitológicos utilizados na identificação morfológica das espécies, os quais se caracterizam pela facilidade e baixo custo. Na maioria dos estudos realizados no Brasil, a distinção entre as espécies Haemonchus contortus e Haemonchus placei não tem sido considerada. Vários relatos de H. contortus, particularmente em bovinos, podem se tratar na verdade da infecção dos animais por H. placei. A identificação correta das espécies é, portanto, fundamental. Além das medidas dos espículos dos exemplares machos, outros detalhes morfológicos, tais como a sínlofe, devem ser avaliados com o objetivo de auxiliar na diferenciação das espécies. Mensurações das larvas infectantes, obtidas em coproculturas, podem também indicar a espécie de Haemonchus presente. Esse procedimento pode ser útil especialmente em estudos que não envolvem a necropsia de animais, como é o caso de testes destinados a avaliar a resistência anti-helmíntica em rebanhos.

Haemonchus contortus infection in sheep: parasite fecundity correlates with worm size and host lymphocyte counts.

Two experiments were conducted to elucidate the timing and nature of the sheep immune response to Haemonchus contortus (Barber's pole worm). The first experiment examined the establishment of H. contortus populations and the immune response by comparing a bolus infection of third-stage larvae in naïve sheep with a group previously primed by a trickle infection. The second experiment used staggered doses of ivermectin-resistant larvae to compare the development of adult worms during different durations of trickle infection with ivermectin-sensitive larvae. Infections successfully generated pathological signs of haemonchosis such as anaemia. Image analysis software was used to measure the area and perimeter of worms collected at post-mortem, and the number of eggs present in individual adult females (fecundity) was significantly correlated with worm size. A significant inverse correlation was found between blood lymphocyte counts and worm fecundity. The absence of correlation between worm fecundity and other leukocyte and erythrocyte counts highlighted the specificity of the lymphocyte response. This is the first report of a link between haematology profiles and worm fecundity in haemonchosis. The correlation observed between adult worm size and egg content leads to the hypothesis that egg production in H. contortus is limited by immune regulation of worm size and presumably growth. Mean worm size and fecundity declined as sheep received more prolonged trickle infections before necropsy, confirming previous reports that immune responses to adult worms are enhanced by ongoing larval challenge. Immunohistochemical results showed trends consistent with a Th2 (humoral) immune response which has been implicated in reducing nematode burdens in several species.

Genetic and phenotypic differences between isolates of Haemonchus contortus in Australia.

We believe this study is the first to consider the genetic and phenotypic divergence between isolates of Haemonchus contortus in Australia. Microsatellite markers have been used to investigate genetic divergence, whilst phenotypic divergence has been considered through individual worm morphology, isolate life history traits and the effect of isolates upon the host. The results are discussed in the context of the likely introduction of H. contortus to Australia, its recent isolation, and the characteristics of sheep and goat farming which might act to either isolate or distribute parasites. We conclude that there is significant observable genetic divergence between isolates of H. contortus in Australia. The divergence may have been under-estimated in this study due to a variety of factors. Phenotypic divergence is also observed, and potentially has significant implications for both economic losses due to haemonchosis on individual properties and for decisions regarding the regulation of stock movements in Australia.

Morphological characterization of Haemonchus contortus in goats (Capra hircus) and sheep (Ovis aries) in Penang, Malaysia.

The large stomach worm, Haemonchus contortus is an important pathogen of goats (Capra hircus) and sheep (Ovis aries). This paper describes characteristics of surface cuticular ridges (synlophe) of H. contortus adults from the two hosts. There were more ridges in H. contortus from goats compared to that from sheep. Total body length, vulvar morphology, spicule length and cervical papillae had been considered as markers of physical adaptation and were studied and described.

Phylogeny for species of Haemonchus (Nematoda: Trichostrongyloidea): considerations of their evolutionary history and global biogeography among Camelidae and Pecora (Artiodactyla).

Phylogenetic analysis of 25 morphological characters among the 12 species of Haemonchus resulted in 1 most parsimonious tree (60 steps; consistency index = 0.67, retention index = 0.80). Monophyly for Haemonchus was diagnosed by 3 unequivocal synapomorphies, including the asymmetric origin of the dorsal ray, relative size of the ventral rays, and the presence of a barb on each spicule tip. Species of Haemonchus have complex histories with respect to host and geographic associations: (1) origins in Africa with basal diversification in antelopes (H. krugeri, H. lawrencei, H. dinniki, H. horaki), (2) independent events of colonization for those species in Caprini and Bovinae (H. contortus, H. placei, H. bedfordi, H. similis), (3) colonization and development of core host associations within Camelidae (H. longistipes) and among Antilopinae, Tragelaphini, and Giraffidae (H. mitchelli, H. okapiae, H. vegliai), and (4) geographically widespread species that are represented only by those that have been translocated with domestic stock. The North American fauna is characterized by 3 introduced and exotic species, H. placei, H. contortus, H. similis, which emphasizes the importance of continued documentation of faunal diversity in the context of predictive foundations derived from phylogenetic studies. Satellite associations for species of Haemonchus, particularly among Cervidae and Camelidae in the Neotropics and Cervidae, Antilocapridae, and possibly wild Caprinae in the Nearctic, have been a consequence of introductions and exchange of parasites at historical interfaces for managed and natural ecosystems. Such distributions are emblematic of the overriding significance of anthropogenic factors as determinants of the global distributions for pathogenic parasites in domestic and wild ruminants.

Morphological identification of nematode larvae of small ruminants and cattle simplified.

A simplified system has been developed to facilitate identification of nematode larvae of the common nematodes of cattle, sheep and goats. Firstly, in addition to the characteristics conventionally used for identification (such as the shape of the cranial extremity and numbers of intestinal cells), the lengths of the infective sheath tails of infective larvae of each genus/species are related to that of Trichostrongylus spp. instead of using measurements for differentiation. For instance, if the mean length of the sheath tail (the distance the sheath extends caudad beyond the caudal tip of the larva) of Trichostrongylus spp. is assumed to be "X", then that of Haemonchus contortus is 2-2.7"X", and that of Oesophagostomum spp. from sheep is 4-7"X", etc. Secondly, by estimating the proportion of the sheath tail of a larva comprised of a terminal thin whip-like filament, identification is aided, particularly in those L3 of species that resemble one another closely, such as Chabertia ovina and Oesophagostomum venulosum or Oesophagostomum columbianum. After some practice with the system it is usually necessary to measure only one or two sheath tails of L3 in a mixed population, whereupon the identity of most of the remaining L3 can be estimated in relation to those measured, without a need for further measurements. The keys were found to facilitate differential larval identification and are particularly useful for training.