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Robustostrongylus aferensis gen. nov. et sp. nov. (Nematoda: Trichostrongyloidea) in Kob (Kobus kob) and Hartebeest (Alcelaphus buselaphus jacksoni) (Artiodactyla) from Sub-saharan Africa, with Further Ruminations on the Ostertagiinae
Abstract
Abomasal nematodes (Ostertagiinae: Trichostrongyloidea), representing a previously unrecognized genus and species, were discovered in kob (Kobus kob) and kongoni (hartebeest) (Alcelaphus buselaphus jacksoni) from Uganda during surveys of ungulate parasites in the 1960s. Robustostrongylus aferensis gen. nov. et sp. nov. is characterized by a ventriculus-like, bilobed valve at the junction of the esophagus and intestine, a synlophe with unusually robust ridges, cervical papillae and excretory pore situated posterior to the mid-length of the esophagus, a unique body form and large diameter in males and females, a relatively anterior position for the vulva, and strongly convoluted and spiraled ovarian tracks in females. Bursal structure is 2-1-2, with subequal Rays 2/3, strongly reduced and robust Rays 8, and relatively narrow Rays 9/10 contained within a reduced, laterally inflated dorsal lobe. Spicules are filamentous and tripartite; the gubernaculum is cryptic, alate, and heart-shaped in the anterior. Robustostrongylus aferensis, with narrow filamentous spicules that trifurcate distally near 80%, paired arcuate "0" papillae that terminate in bulbous expansions, and a reduced dorsal lobe and ray most closely resembles species of Longistrongylus. A suite of unique characters, consistent in males and females, however, unequivocally distinguishes specimens of R. aferensis from all ostertagiines with either a 2-1-2 or 2-2-1 bursal pattern. Among 15 genera of the Ostertagiinae in the global fauna, 5 are entirely limited in distribution to Africa, including Africanastrongylus, Hamulonema, Longistrongylus, Pseudomarshallagia, and Robustostrongylus gen. nov.; species among 5 additional genera, including Cervicaprastrongylus, Hyostrongylus, Marshallagia, Ostertagia, and Teladorsagia, also occur in Africa, but they are represented as mosaics, with diversity centered in Eurasia or the Holarctic.
... Based on the literature, 5 of the 15 genera within the subfamily Ostertagiinae (i.e. Teladorsagia, Ostertagia, Orloffia, Spiculopteragia, Marshallagia) should be considered as polymorphic (Hoberg et al., 2009). ...
This paper focuses on the species diversity among the Trichostrongylidae Leiper, 1912 (Nematoda: Strongylida), and complexity of the family systematics. Polymorphism (subfamilies: Ostertagiinae, Cooperiinae and Haemonchinae), the presence of cryptic species (genus: Teladorsagia) and hybridization (genera: Cooperia, Haemonchus and Ostertagia) are presented and discussed, considering both morphological and molecular evidence. Some of these phenomena are common, nevertheless not sufficiently understood, which indicates the need for expanding the current state of knowledge thereof. Within the Trichostrongylidae, species distinction supported merely by morphological features is difficult, and requires confirmation by means of molecular methods. The parasitic nematode taxonomy is complicated mainly by the genus Teladorsagia, but complexity may also be expected among other Ostertagiinae (e.g. in the genera Ostertagia and Marshallagia). The data presented here show that the members of the Trichostrongylidae can significantly complicate unambiguous species identification. Hence, it is essential to consider the phenomena mentioned, to gather valid and comparable data on the biodiversity of this family.
... In the Arctic, ungulates are important sources of food and income as well as a focus of traditional activities for indigenous peoples (AMAP, 2002; Nancarrow and Chan, 2010). Additionally, they provide an important habitat for various helminth, protozoan and arthropod parasites (Hoberg et al., 2008a; Kutz et al., 2009b). Parasites can cause significant clinical and subclinical disease in wildlife and consequently influence the dynamics and trajectory of wildlife populations (Hudson and Dobson, 1997; Hudson and Greenman, 1998; Irvine et al., 2000). ...
Parasites play an important role in the structure and function of arctic ecosystems, systems that are currently experiencing an unprecedented rate of change due to various anthropogenic perturbations, including climate change. Ungulates such as muskoxen, caribou, moose and Dall's sheep are also important components of northern ecosystems and are a source of food and income, as well as a focus for maintenance of cultural traditions, for northerners. Parasites of ungulates can influence host health, population dynamics and the quality, quantity and safety of meat and other products of animal origin consumed by people. In this article, we provide a contemporary view of the diversity of nematode, cestode, trematode, protozoan and arthropod parasites of ungulates in arctic and subarctic North America and Greenland. We explore the intricate associations among host and parasite assemblages and identify key issues and gaps in knowledge that emerge in a regime of accelerating environmental transition.
... Specimens of L. meyeri were not available for sectioning; transverse sections suitable for illustration were not available from all species. Nematodes were identified based on morphological criteria previously defined for the genus and respective species (Le Roux, 1931;Gibbons, 1977Gibbons, , 1981Boomker and Durette-Desset, 1997;Hoberg et al., 2008, Hoberg, Abrams, andPilitt, 2009). Nomenclature for ungulate hosts is consistent with Wilson and Reeder (1993). ...
The synlophe, or system of longitudinal cuticular ridges characteristic of some trichostrongyloid nematodes, is examined in detail for 6 of 8 species in Longistrongylus (Ostertagiinae) that occur in ungulates across sub-Saharan Africa. Among the species of Longistrongylus examined, 5 are characterized by a tapering pattern laterally in the cervical zone (anterior to the esophageal-intestinal junction), which is largely consistent among multiple male and female specimens; in contrast, for Longistrongylus meyeri the lateral pattern is parallel. The synlophe is bilaterally symmetrical, with ridges extending from the base of the cephalic expansion to near the caudal extremity in males and females. Ridges are acutely pointed, with perpendicular orientation and absence of gradient as viewed in transverse section. Species-specific patterns in conjunction with the numbers of ridges may serve to augment an array of diagnostic characters for species of Longistrongylus and contribute to increasingly accurate identification of female specimens. Among 5 of 6 species examined in the current study, the numbers of ridges in males was equal to or exceeded that observed in females, a pattern seen only in Africanastrongylus among the 15 genera of the Ostertagiinae. The differential numbers of ridges in males and females may represent another character among the suite of attributes that in part diagnose the genus Longistrongylus.
The independence of Pseudomarshallagia and its placement among the medium stomach worms of ungulates,
Ostertagiinae, is confirmed based on comparative morphological studies of the synlophe and genital attributes among male and female
specimens. An emended description of Pseudomarshallagia elongata is presented based on a series of specimens in sheep from northern
Ethiopia. Pseudomarshallagia elongata is retained among the 15 genera of the Ostertagiinae based on presence of a prominent
esophageal-intestinal valve, paired ‘‘0’’ papillae, a modified accessory bursal membrane containing the paired ‘‘7’’ papillae, and
configuration of the copulatory bursa. The structure of the synlophe in males and females is also typical and within the range of
variation demonstrated for Type II and Type A cervical patterns among other ostertagiines. We emphasize the importance of
continued survey and inventory of parasite faunas to establish the limits of diversity on local, regional, and global scales. Consequently
we strongly encourage routine deposition of voucher specimens from all survey activities designed to explore faunal diversity. Although
this is particularly important in regions that remain poorly known, such as Africa, the principle certainly applies more broadly.
Marshallagia lichtenfelsi sp. n. is a dimorphic ostertagiine nematode occurring in the abomasum of mountain goats, Oreamnos americanus, from the Western Cordillera of North America. Major and minor morphotype males and females are characterized and distinguished relative to the morphologically similar Marshallagia marshalli / Marshallagia occidentalis from North America and Marshallagia dentispicularis, along with other congeners, from the Palearctic region. The configuration of the convoluted and irregular synlophe in the cervical region of males and females of M. lichtenfelsi is apparently unique, contrasting with a continuous and parallel system of ridges among those species of Marshallagia, including M. marshalli/M. occidentalis, which have been evaluated. Specimens of M. lichtenfelsi are further defined by the rectangular form of the accessory bursal membrane (width > length) in the major morphotype and by the trapezoidal Sjöberg's organ in the minor morphotype, in addition to specific attributes of the spicules and spicule tips. We regard 12 species, including the proposed new taxon, to be valid. Primary diagnostic characters are reviewed for Marshallagia and a framework is presented for standardization of future descriptions incorporating the synlophe in males and females and the structure of the spicules and genital cone in major and minor morphotype males. The center of diversity for species of Marshallagia is the mountain-steppe region of central Eurasia where 11 species (including the Holarctic M. marshalli) are recognized in association with Caprini, Rupicaprini, and Antelopinae; only 2 species occur in the Nearctic. In this assemblage, M. lichtenfelsi is endemic to North America and limited in host distribution to mountain goats. An intricate history for refugial isolation and population fragmentation demonstrated for mountain goats and wild sheep indicates the potential for considerable cryptic diversity for Marshallagia and other nematodes. Shifting patterns of contact and sympatry among assemblages of ungulates during the Pleistocene are consistent with geographic and host colonization as a process involved in diversification of these parasites.
Collections to explore helminth diversity among free-ranging ungulates in the North American Arctic revealed the occurrence of a third male, or " davtiani ," morphotype for Teladorsagia boreoarcticus . Designated as T. boreoarcticus forma (f.) minor B, the males occurred with T. boreoarcticus f. major and T. borearcticus f. minor A in endemic populations of muskoxen (Ovibos moschatus wardi) and barrenground caribou (Rangifer tarandus groenlandicus) on Victoria Island, Nunavut, Canada, and in muskoxen and Peary caribou ( Rangifer tarandus pearyi ) on Banks Island, Northwest Territories, Canada. These specimens differ from conspecific morphotypes in the structure of the genital cone and Sjöberg's organ. Relative to T. boreoarcticus f. minor A, specimens of T. boreoarcticus f. minor B are consistently smaller, and mean dimensions for the bursa and spicules do not overlap. The robust spicules are similar in form, particularly in the relative length of the dorsal and ventral processes, but mean total length is substantially less in specimens of T. boreoarcticus f. minor B. Differences that distinguish the minor morphotypes of T. boreoarcticus parallel those demonstrated for the T. trifurcata and T. davtiani morphotypes in association with T. circumcincta sensu stricto. New host and geographic records include the 3 morphotypes of T. boreoarcticus in muskoxen and Peary caribou from Banks Island and in barrenground caribou from Victoria Island. Recognition of the ubiquitous nature of cryptic species emphasizes the need to effectively develop and use our collections-based resources and museum archives to build a robust understanding of the biosphere. Field inventory should include provisions for integrative approaches that preserve specimens suitable for comparative morphology, multi-faceted molecular investigations, and population genetics.
Africanastrongylus giganticus n. sp. is described based on large ostertagiine nematodes occurring in the abomasum of African buffalo, Syncerus caffer, from Uganda; this represents the second species recognized in the genus. Specimens of A. giganticus are characterized by large size (15-19 mm in total length), a strongly tapering synlophe in the cervical region, and a great number of ridges at all levels of the body (maximum 72 attained in the third quarter); numbers of ridges exceed that reported among any known genera and species of the Ostertagiinae. We refer A. giganticus to this genus based on a strongly tapering lateral synlophe, relatively large numbers of ridges at all levels of the body, miniscule cervical papillae, poorly demarcated divisions of the ovejector, absence of vulval cuticular inflations, and the presence of slightly protruding lips at the vulva. It is distinguished from its congener, Africanastrongylus buceros, in total length, maximum number of ridges (68-72 vs. 53, respectively), structure and disposition of the synlophe, presence of strongly spiraled ovarian tracks, and eggs that are distributed in 3 or more rows in the uterus. A superficial resemblance to Longistrongylus meyeri, the only other large ostertagiine in the African fauna, is evident; these species, however, are distinct based on the synlophe and other characters. Recognition of a second species of Africanastrongylus represented by nematodes of large size suggests that prior reports of L. meyeri in Syncerus caffer may be attributable to A. giganticus.
Les auteurs signalent, pour la seconde fois en Ethiopie, la présence, dans la caillette des moutons, de Pseudomarshallagia elongata Roetti, 1941. Le parasite est redécrit et le sous-genre Pseudomarshallagia est érigé en genre avec 2 espèces: Pseudomarshallagia thalae et Pseudomarshallagia elongata
Aim: To integrate ecological fitting, the oscillation hypothesis and the taxon pulse hypothesis into a coherent null model for the evolution of complex host–parasite associations.
Location: Global.
Methods: This paper reviews and synthesizes literature that focuses on phylogenetic analyses and reciprocal mapping of a model system of hosts and their parasites to determine patterns of host–parasite associations and geographical distributions through time.
Results: Host-switching and geographical dispersal of parasites are common phenomena, occurring on many temporal and spatial scales. Diversification involving both co-evolution and colonization explains complex host–parasite associations. Across the expanse of Earth history, the major radiations in host– parasite assemblages have been preceded by ecological disruption, ecological breakdown and host-switching in a context that can be defined by the concept of ecological fitting. This cyclical process sets the stage for co-diversification during periods of relative stability, punctuated by host-switching during episodes of regional to global environmental disruption and climatological change.
Main conclusions: Most observed host–parasite associations can be explained by an historical interaction between ecological fitting, oscillation (episodes of increasing host range alternating with isolation on particular hosts) and taxon pulses (cyclical episodes of expansion and isolation in geographical range). Major episodes of environmental change appear to be the main drivers for both the persistence and diversification of host–parasite systems, creating opportunities for host-switching during periods of geographical expansion and allowing for co-evolution and co-speciation during periods of geographical isolation.
A parasitological perspective in biodiversity survey and inventory provides powerful insights into the history, structure, and maintenance of the biosphere. Parasitology contributes a powerful conceptual paradigm or landscape that links ecology, systematics, evolution, biogeography, behavior, and an array of biological phenomena from the molecular to the organismal level across the continuum of microparasites to macroparasites and their vertebrate and invertebrate hosts. Effective survey and inventory can be strategically focused or can take a synoptic approach, such as that represented by the All Taxa Biodiversity Inventory. We argue that parasitology should be an integral component of any programs for biodiversity assessment on local, regional, or global scales. Taxonomists, who constitute the global taxasphere, hold the key to the development of effective surveys and inventories and eventual linkage to significant environmental and socioeconomic issues. The tax-asphere is like a triage team. The "battlefield" is the biosphere, and the "war" is human activities that degrade the biosphere. Sadly, at the point in time that we reali/,e we have documented only a tiny portion of the world's diversity, and want to document more, we find that one of the most rare and declining groups of biologists is the taxasphere. This taxonomic impediment, or critical lack of global taxonomic expertise recognized by Sys-tematics Agenda 2000 and DIVERSITAS, prevents initiation and completion of biodiversity research programs at a critical juncture, where substantial components of global diversity are threatened. The Convention for Biological Diversity mandates that we document the biosphere more fully, and as a consequence, it is necessary to revitalize the taxasphere. One foundation for development of laxonomic expertise and knowledge is the Global Taxonomy Initiative and its 3 structural components: (1) systematic inventory, (2) predictive classifications, and (3) systematic knowledge bases. We argue that inclusion of parasites is critical to the success of the Global Taxonomy Initiative. Predictive databases that integrate ecological and phylogenetic knowledge from the study of parasites are synergistic, adding substantially greater ecological, historical, and biogeographic context for the study of the biosphere than that derived from data on free-living organisms alone. A Biodiversity Perspective knowledge. They can (1) focus on local, region
Aim To integrate ecological fitting, the oscillation hypothesis and the taxon pulse hypothesis into a coherent null model for the evolution of complex host–parasite associations.
Location Global.
Methods This paper reviews and synthesizes literature that focuses on phylogenetic analyses and reciprocal mapping of a model system of hosts and their parasites to determine patterns of host–parasite associations and geographical distributions through time.
Results Host-switching and geographical dispersal of parasites are common phenomena, occurring on many temporal and spatial scales. Diversification involving both co-evolution and colonization explains complex host–parasite associations. Across the expanse of Earth history, the major radiations in host–parasite assemblages have been preceded by ecological disruption, ecological breakdown and host-switching in a context that can be defined by the concept of ecological fitting. This cyclical process sets the stage for co-diversification during periods of relative stability, punctuated by host-switching during episodes of regional to global environmental disruption and climatological change.
Main conclusions Most observed host–parasite associations can be explained by an historical interaction between ecological fitting, oscillation (episodes of increasing host range alternating with isolation on particular hosts) and taxon pulses (cyclical episodes of expansion and isolation in geographical range). Major episodes of environmental change appear to be the main drivers for both the persistence and diversification of host–parasite systems, creating opportunities for host-switching during periods of geographical expansion and allowing for co-evolution and co-speciation during periods of geographical isolation.
Of all the morphological characters generally used in classification of the Trichostrongyloidea, the evolution of the caudal bursa seems to illustrate the general evolution of the subfamily most clearly. The Ostertagiinae are divided into two groups. The first group, arising from Graphidium, a parasite of Lagomorphs, is characterized by the isolation of ray 4 and the grouping of ray 2 with ray 3 and ray 5 with ray 6 (arrangement of type 2-1-2). This includes Marshallagia Orloff, 1933, Longistrongylus Le Roux, 1931 and Ostertagia Ransom, 1907. The second group arising from Hyostrongylus (parasites of Lagomorphs and Ungulates--usually primitive), is characterized by the grouping of ray 2 with ray 3, ray 6 with ray 5, and the isolation of ray 6 (arrangement type 2-2-1). This consists of Spiculopteragia (Orloff, 1933), Teladorsagia Andreeva and Satubaldin, 1953 and Gazellostrongylus Yeh, 1956. Evolution in both groups involves the elongation of ray 4 and, especially in the first group, a shortening of rays 8, 5 and 6, as well as the dorsal ray which becomes more and more deeply divided. The following synonymies are recognized: - Bergheia Drozdz, 1965, synonym of Hyostrongylus Hall, 1921. - Grosspiculagia (Orloff, 1933). Ostertagiella Andreeva, 1956, Camelostrongylus Orloff, 1933 and Orloffia Drozdz, 1965, synomyms of Marshallagia. - Bigalkea Monnig, 1931, Bigalkenema Ortlepp, 1963, Kobusinema Ortlepp, 1963, Ostertagia (Costarcuata) Schulz and Kadenazii, 1950, Pseudomarshallagia (Roetti, 1941), synonyms of Longistrongylus. - Gruhneria Sarwar, 1956, Capreolagia Schulz, Andreeva and Kadenazii, 1954, Skrjabinagia (Kassimov, 1942), Muflonagia Schulz, Andreeva and Kadenazii, 1954, Sjobergia Sarwar, 1956, Ostertagiana Dikov, 1963, synomyms of Ostertagia.(ABSTRACT TRUNCATED AT 250 WORDS)
A morphologically based cladistic analysis of 40 genera included within the Trichostrongyloidea (Amidostomatidae, Dromaeostrongylidae and Trichostrongylidae) is proposed. Two genera were used as outgroups, one from the Strongylina and the other from the Ancylostomatina. Seven genera do not appear in the matrix because some significant morphological characters remain unknown for these genera. Nonetheless, except for Moguranema which is excluded as incertae sedis, a likely systematic position could be assigned to them based on the morphological characters that are known.
The crisis of emerging infectious disease stems from the absence of comprehensive taxonomic inventories of the world's parasites, which includes the world's pathogens. Recent technological developments raise hopes that the global inventory of species, including potential pathogens, can be accomplished in a timely and cost-effective manner. The phylogenetics revolution initiated by systematists provides a means by which information about pathogen transmission dynamics can be placed in a predictive framework. Increasingly, that information is widely available in digital form on the Internet. Systematic biology is well positioned to play a crucial role in efforts to be proactive in the arena of emerging parasitic and infectious diseases.
Parasites are integral components of the biosphere. Host switching correlated with events of episodic climate change is ubiquitous in evolutionary and ecological time. Global climate change produces ecological perturbations, which cause geographical and phenological shifts, and alteration in the dynamics of parasite transmission, increasing the potential for host switching. The intersection of climate change with evolutionary conservative aspects of host specificity and transmission dynamics, called ecological fitting, permits emergence of parasites and diseases without evolutionary changes in their capacity for host utilization.
In the polymorphic Teladorsagia circumcincta (morphs circumcincta and trifurcata), a sheep and goat line (SGL) and a goat line (GL) have been previously described on the basis of the malate dehydrogenase allozyme polymorphism (MDH-2) and of the morphology of the dorsal ray. The GL were never found alone in 1 host, so the status of species was not given to these 2 lines. To investigate further whether there are other genetic markers that will delineate them, we collected T. circumcincta worms from goat and sheep at 8 farms in Touraine (west-central France). The worms were identified individually as being SGL or GL on the basis of MDH-2 polymorphism. This distinctiveness was corroborated by sequences of the β-tubulin isotype 1 gene, the second internal transcribed spacer (ITS2) of their rDNA, and the nicotinamide dehydrogenase (ND4) gene of their mDNA. The extent of the divergence in the 3 additional genetic markers was such that SGL and GL may be considered as 2 species. A third putative species was found in the SGL line based exclusively on the ND4 gene. These findings suggest that T. circumcincta is a species complex and that further investigation is required on a wider geographic scale.
Bovid (essentially antelope) biostratigraphy and phylogeny indicate a widespread environmental change in Africa near 2.5 Myr. Morphological and ecological analyses of the taxa lead to the inference that across sub-Saharan Africa a spreading of open grassland, probably caused by global reduction in temperature and associated changes in rainfall, occurred at around this time. -from Author
In most parts of the world helminth infestations are responsible for such a heavy wastage of sheep that their control has undoubtedly become one of the most important charges that devolve upon the flock-owner. It might be thought that in tropical regions suited to the raising of sheep the problem of helminth control would cease to be urgent; and that climatic conditions, the general aridity of land chosen in such countries for sheep-rearing, and its consequently smaller carrying capacity, would contribute to form an environment distinctly unfavourable both to the survival of the free-living stages of parasitic worms and to their chances of reaching a suitable intermediary or definitive host. Such areas are not usually well watered, and it certainly does result from the paucity of permanent streams and standing water that few species of trematodes are represented. Those species that do occur, e.g. Fasciola hepatica, F. gigantica, are restricted to particular localities and are easily subjected to control.(Received December 19 1932)
Only three of the four species assigned to the genus Cooperioides are considered valid and are redescribed. The fourth species Cooperioides kenyensis Daubney, 1933 is considered a synonym of Cooperioides hamiltoni (Mönnig, 1932) Mönnig, 1933. A key to the species of the genus is given and their hosts and geographical localities are listed.
Parasitologists have long studied helminth infections in wildlife species and have documented the existence of many organisms from a diversity of mammalian hosts. With this accumulation of information has come improved understanding of the significance of these organisms and the diseases they produce in their mammalian hosts. Some of the most notable examples include the metastrongyloid lungworms, Trichinella spiralis, and Elaeophora schneideri, which are covered separately in this volume. It is, however, for the group of parasites referred to as gastrointestinal nematodes that we have accumulated the most data. Only recently has progress been made in determining the significance of these strongylate nematodes with respect to their potential impact on the morbidity and mortality of the ruminants that they infect.
The accumulation of information on diseases of wild animals into a single combined volume has been slow, but progress has coincided with the proliferation of data for host and parasite interactions. Numerous references including Alaskan Wildlife Diseases (Dieterich 1981), Manual of Common Wildlife Diseases in Colorado (Adrian 1981), Field Manual of Wildlife Diseases in the Southeastern United States (Davidson and Nettles 1988), Zoo and Wildlife Medicine (Fowler 1993), and the previous editions of Parasitic Diseases of Wild Mammals (Davis and Anderson 1971) have all made significant contributions to our knowledge. Beyond North America, Dunn (1969) and Govorka et al. (1988) provided excellent compilations on the helminths in wild ruminants. In the 1971 printing of Parasitic Diseases of Mammals, however, there was no general coverage of gastrointestinal nematodes, and only T. spiralis was addressed. Herein, we present the first synoptic review of the strongylate nematodes that occur in the gastrointestinal system of wild ruminants from North America.
Fourteen of the 20 species recorded from Africa are resdescribed and illustrated. Of the remaining six species, four, namely, Cooperia borgesi, C. hippotragusi, C. minor and C. reduncai are considered species inquirendae and one, C. africana, is a possible synonym of C. punctata. The validity of C. oncophora and C. surnabada is discussed. A key to 16 species is given.
Only nine species of the genus Haemonchus are considered valid, namely, H. contortus (Rudolphi, 1803) Cobb, 1898 (type), H. bedfordi Le Roux, 1929, H. dinniki Sach, Gibbons & Lweno, 1973, H. krugeri Ortlepp, 1964, H. lawrencei Sandground, 1933, H. longistipes Railliet & Henry, 1909, H. mitchelli Le Roux, 1929, H. similis Travassos, 1914 and H. vegliai Le Roux, 1929. These are redescribed.
H. bispinosus Molin, 1860, H. placei Place, 1893, H. cervinus Baylis & Daubney, 1922, H. okapiae van den Berghe, 1937 (in part), H. tartaricus Evranova, 1940, H. contortus contortus Das & Whitlock, 1960, H. contortus cayugensis Das & Whitlock, 1960, H. contortus bangalorensis Rao & Rahman, 1967, H. contortus hispanicus Martnez Gmez, 1968, H. contortus kentuckiensis Sukhapesna, 1974 and H. contortus var. uktalensis Das & Whitlock, 1960 are considered synonyms of H. contortus (Rudolphi, 1803) Cobb, 1898.The synonymy of H. lunatus Travassos, 1914, H. atectus Lebedev, 1929, H. pseudocontortus Lebedev, 1929 and H. fuhrmanni Kamensky, 1929 with H. contortus is confirmed. H. okapiae van den Berghe, 1937 (in part) is considered a synonym of H. mitchelli Le Roux, 1929 and H. bubalis Chauhan & Pande, 1968 is considered a synonym of H. similis Travassos, 1914.
H. bovis Bonelli, 1941 and H. contortus var. kashmirensis Fotedar & Bambroo, 1965 are considered species inquirendae. An illustrated key to the species of the genus is provided.
A list of 19 polymorphic species of the generaOstertagia, Orloffia, Teladorsagia, Marshallagia andSpiculopteragia is presented. The morphs of the polymorphic species in question have identical females and show similarities in some male features, e.g. the structure of the oesophagus, the synlophe and the shape of the rays of the copulatory bursa. However, they differ by characters which are recognised as generic features, e.g. the structure of spicules, gulbernaculum and genital cone. The presence or lack of minor morphs is a generic feature in the Ostertagiinae. Each polymorphic genus has a defined type of minor morph.
In North America broad-based research networks explore the interaction of vertebrates, their characteristic arrays of pathogens and emergent disease. A diversity of programmes address the impact of environmental change on animal health, zoonoses, and human health, but as yet no comprehensive framework or strategy has emerged to develop and implement policy and planning. In a regime of climate change and ecological perturbation, the need to document and understand the health, agricultural, societal and economic impact of pathogens and emerging infectious disease is urgent. An integrated and proactive planning process linking national and international resources can lead to informed predictions aboutthe impact of environmental change and can identify pathways for potential management and mitigation. An effective and comprehensive programme will have components for establishing priorities, developing primary data for faunal structure and biodiversity, a capacity for monitoring and surveillance (including scanning and targeted activities), and linkage to historical and contemporary baselines (against which to assess change) established through archival biological collections. Field and laboratory studies are also necessary to determine developmental thresholds, tolerances and tipping points for many pathogens to establish a context for recognising current constraints and future perturbation, and to explore factors that promote emergence for a variety of pathogens, vectors and pest species. Predictive modelling and risk assessment utilising a range of scenarios for climate change is a final step in this multidisciplinary process.
The species of the genera Longistrongylus, Kobusinema and Bigalkenema are redescribed. Kobusinema and Bigalkenema are considered synonyms of Longistrongylus and their species are transferred as new combinations. Bigalkenema neveulemairei (Gutteres, 1947 Jansen, 1958 and Longistrongylus muraschkinzevi (Shulz and Kadenatsii, 1950), Jansen, 1958 are considered species inquirendae. A key to the species of the genus Longistrongylus is given and their hosts and geographical localities are listed.
After a historical introduction to the Ostertagiinae and several revisions of this group, it is proposed that the subfamily shall contain seven genera: Ostertagia, Hyostrongylus, Longistrongylus, Marshallagia, Mazamastrongylus, Spiculopteragia and Teladorsagia. The synonyms of these genera are to be found in the literature. In this proposal the genus Hyostrongylus should belong to the Ostertagiinae though it has probably an intermediate position between the Ostertagiinae and Graphidiinae. Cervicaprastrongylus and Tunisostertagia are regarded as synonyms of Hyostrongylus and Ostertagia respectively. The position of Ostertamia and Trichostrongyella is not clear but they could have their closest affinities with Teladorsagia and Mazamastrongylus respectively. The origin of the subfamily and the relationships of the genera with regard to their preferences for host groups and to their geographical distribution are discussed.
The nematode family Trichostrongylidae is divided into six subfamilies Trichostrongylinae, Libyostrongylinae, Haemonchinae, Cooperiinae, Ostertagiinae and Graphidiinae which are briefly reviewed. The key is based mainly on specimens examined by the authors and includes one genus in the Trichostrongylinae, three genera in the Libyostrongylinae, six in the Haemonchinae, seven in the Cooperiinae, 17 in the Ostertagiinae and two in the Graphidiinae. Pseudostertagia, Pararhabdonema, Parostertagia, Obeliscoides and Travassosius are briefly described, figured and considered as genera of uncertain affinities. Nine genera, namely, Moguranema, Neocooperia, Bergheia, Mazamastrongylus, Paramecistocirrus, Ostertagiana, Ostertamia, Graphidiella and Trichostrongyella, specimens of which have not been available to the authors for examination, are listed giving details of host, habitat and geographical location, together with a summary of the diagnostic characters as described by the original authors.
Phylogenetic analysis of the subfamilies of the Trichostrongylidae based on 22 morphological transformation series produced a single cladogram with a consistency index (CI) = 74.2%. Monophyly for the family was supported by the structure of the female tail and copulatory bursa. Two major clades are recognized: the Cooperiinae clade with the basal Cooperiinae and Libyostrongylinae+Trichostrongylinae, and the Graphidiinae clade with the basal Graphidiinae and Ostertagiinae+Haemonchinae. Dendrograms presented by Durette-Desset (1985) (CI = 56.1%) and Lichtenfels (1987), based on the key to the Trichostrongylidae by Gibbons and Khalil (1982) (CI = 59.0%), were found to be relatively inefficient in describing character evolution and in supporting putative relationships among the subfamilies. Based on the current analysis, the intestine appears to have constituted the ancestral habitat for the trichostrongylids with the stomach/abomasum having been independently colonized in each clade. Assessment of host associations suggests extensive colonization but also a high degree of coevolution with Bovidae and Cervidae for Ostertagiinae+Haemonchinae. Biogeography for this assemblage is complex, but this analysis is compatible with a Palearctic or Eurasian origin for Cooperiinae, Haemonchinae, and Ostertagiinae.
Discovery of the ostertagiine nematode Teladorsagia boreoarcticus n. sp. in muskoxen, Ovibos moschatus, from the central Canadian Arctic highlights the paucity of knowledge about the genealogical and numerical diversity of nematode faunas characteristic of artiodactyls at high latitudes across the Holarctic. Teladorsagia boreoarcticus is a dimorphic cryptic species distinguished from Teladorsagia circumcincta/Teladorsagia trifurcata in domestic sheep by a 13% divergence in the ND4 region of mitochondrial DNA, constant differences in the synlophe, and significantly longer esophageal valve, spicules, gubernaculum, and bursa. Teladorsagia boreoarcticus represents an archaic component of the North American fauna and may have a Holarctic distribution in muskoxen and caribou. Recognition of T. boreoarcticus in muskoxen, in part, corroborates hypotheses for the existence of a cryptic species complex of Teladorsagia spp. among Caprinae and Cervidae at high latitudes and indicates the importance of climatological determinants during the late Tertiary and Pleistocene on diversification of the fauna. Also reinforced is the concept of the North American fauna as a mosaic of endemic and introduced species. Discovery of a previously unrecognized species of Teladorsagia has additional implications and clearly indicates that (1) our knowledge is incomplete relative to potentially pathogenic nematodes that could be exchanged among domestic and wild caprines; (2) we do not have sufficient knowledge of the fauna to understand the ecological control mechanisms (limitations) on dissemination and host range; and (3) an understanding of historical and geographical influences on the genealogical diversity and distribution of nematode faunas in domestic and wild ruminants is requisite to define the interface between agricultural and natural ecosystems across the Holarctic.
The pattern of longitudinal ridges (synlophe) on the external cuticular surface of trichostrongylid nematodes has been shown to be of value for distinguishing species and determining relationships among higher taxa. In the process of studying Mecistocirrus digitatus, the large stomach worm of bovids of Asia that has been imported and established in the Americas, we observed remarkably similar synlophe patterns to those described for 3 species of Haemonchus and to those we examined in a species of Ashworthius. In all 3 genera, the synlophe is absent from the posterior part of the body. Only in Haemonchus does the synlophe extend beyond midbody. In both M. digitatus and Ashworthius sidemi, the synlophe extends posteriorly only about 1/4 of body length. In all 3 genera, the synlophe consists of about 30 ridges in the region of the esophagus with variation among species in specific areas, including additional pairs of subventral and subdorsal ridges and different lengths of sublateral ridges. This information is useful for identifying species and determining relationships among these large stomach worm parasites of cattle, sheep, goats, and farmed and wild cervids.
The most important viral disease of farmed deer and bison is malignant catarrhal fever. The other herpesviruses which have been isolated from these species are briefly described. Other viral agents that are recognised in these animals, including adenovirus, parapox, foot and mouth disease, bluetongue, epizootic haemorrhagic disease, bovine virus diarrhoea, rotavirus and coronavirus, are also discussed. Ectoparasites of importance in this group in various parts of the world include a variety of ticks, as well as lice, keds, Oestridae, mange mites and fire ants. Helminth parasites include liver flukes (Fascioloides and Fasciola), gastrointestinal nematodes of the family Trichostrongylidae, pulmonary lungworms of the genus Dictyocaulus and extra-pulmonary lungworms of the family Protostrongylidae. Chronic wasting disease is principally important in North America, where the disease occurs in wild cervids in a limited area and has been reported in farmed deer in a small number of states in the United States of America and one province in Canada. These diseases are summarised in terms of their classification, epidemiology, clinical signs, pathology, diagnosis, treatment and control.
In the polymorphic Teladorsagia circumcincta (morphs circumcincta and trifurcata), a sheep and goat line (SGL) and a goat line (GL) have been previously described on the basis of the malate dehydrogenase allozyme polymorphism (MDH-2) and of the morphology of the dorsal ray. The GL were never found alone in 1 host, so the status of species was not given to these 2 lines. To investigate further whether there are other genetic markers that will delineate them, we collected T. circumcincta worms from goat and sheep at 8 farms in Touraine (west-central France). The worms were identified individually as being SGL or GL on the basis of MDH-2 polymorphism. This distinctiveness was corroborated by sequences of the beta-tubulin isotype I gene, the second internal transcribed spacer (ITS2) of their rDNA, and the nicotinamide dehydrogenase (ND4) gene of their mDNA. The extent of the divergence in the 3 additional genetic markers was such that SGL and GL may be considered as 2 species. A third putative species was found in the SGL line based exclusively on the ND4 gene. These findings suggest that T. circumcincta is a species complex and that further investigation is required on a wider geographic scale.
A species of Ashworthius is reported for the first time in the Western Hemisphere, and A. patriciapilittae n. sp. is described on the basis of specimens in white-tailed deer Odocoileus virginianus from Costa Rica. Among 8 known species, A. patriciapilittae is morphologically similar to A. tuyenquangi in red muntjac Muntjacas muntjak from northern Vietnam. The synlophe in A. patriciapilittae is composed of 26 ridges in the cervical zone and is continuous to the caudal extremity in males and females. Males are characterized by a complex dorsal ray and narrow trifurcate spicules (351-356 microm long) lacking an "eyelet." with dissimilar ventral and dorsal processes; the gubernaculum is 45-48% of the spicule length. Females have a prominent linguiform flap at the vulva and large eggs (108-142 microm long). The presence of A. patriciapilittae in Costa Rica is examined in the context of competing hypotheses for cospeciation or contemporary host-switching in cervids: either A. patriciapilittae is a component of an endemic Central and South American fauna that has diversified through coevolution of Ashworthius and cervid hosts or it has been introduced. Among haemonchines in the Western Hemisphere, specimens of A. patriciapilittae may be confused with 3 species of Haemonchus, including H. contortus, H. placei, and H. similis, that occur in both domestic and wild ruminants. Discovery of A. patriciapilittae emphasizes the continued need for survey and inventory to define the structure and distribution of parasite faunas in wild and domestic ruminants from the Nearctic and Neotropical regions.
The present paper is a report on a collection of helminths collected from gazelles from Tanganyika Territory. It comprises one species of trematode, Paramphistomum microbothriwn ; one species of a larval cestodc Taenia hydatigcna ; eleven species of nematodes, Trichuris spiricollis; Haemonchus contortus; Trichostrongylus probolurus; Gazellofilaria tanganyikae; Cooperioides antidorca; Paracooperia serrata; P. daubneyi; Longistrongylus meyeri; Impalaia nudicollis; Gazellostrongylus lerouxi gen. et sp. nov. (Trichostrongylidae) and Protostrongylns gazellae sp. nov. The seven last mentioned species are all new records for the gazelle.
The purpose of this paper is to bring together all the information available on the helminth parasites known to occur in domesticated animals in Kenya. Several reports have been made by various workers over the past fifty years but these are scattered throughout the literature. Of the principal authors, Daubney (1926, 1933) has recorded many of the Nematodes, and Hudson (1934) published a list of the Cestodes known to occur in mammals, birds and reptiles in East Africa. Dinnik (1951, 1952, 1954, 1959, 1961a, 1961b, 1962) and Dinnik and Dinnik (1954–55, 1954, 1960) have recorded the Trematode species found in cattle, sheep and goats in East Africa.
The use of ovejector structure in the phylogeny of the Haemonchinae is hampered by differences among nematologists in the application of terminology and the recognition of homologous parts. Some workers recognize a sphincter with 2 parts, but others recognize only the proximal, rounded part of the sphincter and include the distal cylindrical part of the sphincter with the vestibule. The results of this study demonstrate that all sphincters of Haemonchinae of ruminants have 2 parts. To encourage the application of a uniform terminology to homologous parts of the ovejector, we propose the use of the terms "sphincter 1" for the rounded part and "sphincter 2" for the cylindrical part. It is hoped that clarification of the terminology for ovejectors of the Haemonchinae will provide a model useful for improving descriptions of ovejectors throughout the Trichostrongyloidea.
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.
We review studies that have used molecular markers to address ecological and microevolutionary processes in parasites. Our goal is to highlight areas of research that may be of particular interest in relation to the parasitic lifestyle, and to draw attention to areas that require additional study. Topics include species identification, phylogeography, host specificity and speciation, population genetic structure, modes of reproduction and transmission patterns, and searching for loci under selection.
An understanding of genetic variation in parasite populations, and how it is partitioned, is required to underpin many areas of basic and applied research. Population genetic studies on parasitic nematode populations are still in their infancy and have been dominated by the use of single locus markers. We have used a panel of five microsatellite markers to undertake a genetic study of a number of field and laboratory populations of Teladorsagia circumcincta. High levels of polymorphism were seen in all the populations examined with the majority of diversity being within rather than between populations. There was no detectable genetic differentiation between the UK populations examined although they included both laboratory passaged and field isolates derived from different geographical regions and host species. This broadly supports previous mtDNA sequence diversity studies of this parasite in the UK and USA. However, some between-population genetic differentiation was apparent when several populations from French goats and a laboratory population from New Zealand were examined. Most notably, a population from a French goat farm, which has previously been suggested to contain a cryptic species, showed very high levels of genetic differentiation from all the other populations. Analysis of multi-locus genotypes suggested the presence of two sympatric parasite populations on this farm with little or no gene flow between them. This supports the hypothesis that parasites currently defined as T. circumcincta by routine morphological criteria comprise more than a single species.
Among genera of the ostertagiine nematodes, structural attributes of Spiculopteragia caballeroi are consistent with criteria that diagnose the genus Sarwaria. Specifically, the following characters are compatible with referral to this genus: (1) species characterized by monomorphic males; (2) tapering lateral synlophe in the cervical zone; (3) minuscule, thornlike cervical papillae; (4) length of Ray "4" < Ray "5"; (5) relatively thick or robust Ray "4"; (6) a substantially reduced dorsal lobe and dorsal ray that are disposed or curved ventrally relative to Rays "8"; and (7) a broad, laterally inflated dorsal lobe. Consequently, we propose Sarwaria caballeroi n. comb., and we provide further validation for the genus Sarwaria. Additionally, we propose that the ostertagiines are represented by 12 valid genera, including those characterized by a bursal formula of 2-2-1 (Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Spiculopteragia, and Teladorsagia in addition to Sarwaria), and those in which the lateral rays describe a 2-1-2 pattern (Camelostrongylus, Longistrongylus, Marshallagia, Orloffia, Ostertagia, and Pseudomarshallagia).
Abomasal nematodes (Ostertagiine: Trichostrongyloidea) representing a previously unrecognized genus and species are reported in African buffalo (Syncerus caffer caffer) from Kenya, Uganda, and South Africa. Africanastrongylus buceros gen. nov. et sp. nov. is characterized by a symmetrical tapering synlophe in the cervical region and a maximum of 60 ridges in males and females. Bursal structure is 2-2-1, with subequal Rays 4/5, massive Rays 8, and Rays 9/10, and a massive dorsal lobe that is reduced in length, laterally and dorsally inflated, and positioned ventral to externodorsal rays. Spicules are tripartite, and the gubernaculum is broadly alate in the anterior. A proconus is present. Among ostertagiines with a 2-2-1 bursa (Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Sarwaria, Spiculopteragia, and Teladorsagia) specimens of Africanastrongylus are differentiated from respective genera based on the structure of the cervical synlophe, patterns of dorsal, externodorsal, lateral, and ventral rays, and configuration of the genital cone, gubernaculum, and spicules. Among 13 genera of the Ostertagiinae in the global fauna, 3 are entirely limited in distribution to Africa, including Africanastrongylus, Longistrongylus, and Pseudomarshallagia. Another 5 genera including Cervicaprastrongylus, Hyostrongylus, Marshallagia, Ostertagia, and Teladorsagia are represented as mosaics, with diversity centered in Eurasia or the Holarctic. Genera not represented in the African fauna include Camelostrongylus among Caprinae and some Antelopinae from Eurasia, Mazamastrongylus and Spiculopteragia in Cervidae from the Holarctic and Eurasia, respectively, Orloffia in Cervidae and Bovidae from the Holarctic, and Sarwaria among Tragulidae and Bovinae in southern Asia. The diverse nature of the ostertagiine fauna, with a disproportionate number of endemic genera relative to other regions of the northern hemisphere, may reflect the timing of episodic expansion events for artiodactyls into Africa from Eurasia during the Tertiary and Quaternary.
Hamulonema gen. nov. is proposed for Teladorsagia hamata and Ostertagia kenyensis in the ostertagiine nematode fauna found in artiodactyl hosts from Africa. Monomorphic species representing this genus are characterized by a bilaterally symmetrical and parallel synlophe in males and females, a 2-2-1 bursal formula, an accessory bursal membrane that is strongly cuticularized and reduced, a strongly reduced dorsal lobe and ray, and robust spicules with a simple, weakly pointed, ventral process, and curved, hooklike dorsal process. Species referred to Hamulonema nov. gen. are immediately distinguished from those of Camelostrongylus, Longistrongylus, Marshallagia, Orloffia, Ostertagia, and Pseudomarshallagia in which the bursal formula is 2-1-2 in males. Hamulonema nov. gen. is distinguished from those genera having a 2-2-1 bursa, including Africanastrongylus, Cervicaprastrongylus, Hyostrongylus, Mazamastrongylus, Sarwaria, Spiculopteragia, and Teladorsagia by the structure of the synlophe, bursa, genital cone, "0" and "7" papillae, dorsal lobe, and spicules in specific instances. In the global fauna, 4 of 14 ostertagiine genera are endemic to Africa. African genera may represent morphologically divergent and discrete or historically isolated lineages reflecting a pattern of geographic and host colonization as a driver for diversification since the Miocene.
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