Searching for common patterns in parasite ecology: species and host contributions to beta-diversity in helminths of South African ungulates and fleas of South American rodents.

We searched for common patterns in parasite ecology by investigating species and host contributions to the beta-diversity of infracommunities (=assemblages of parasites harboured by a host individual) in helminths of three species of South African ungulates and fleas of 11 species of South American rodents, assuming that a comparison of patterns in distinctly different parasites and hosts would allow us to judge the generality or, at least, commonness of these patterns. We used data on species' composition and numbers of parasites and asked whether (i) parasite species' attributes (life cycle, transmission mode, and host specificity in helminths; possession of sclerotized combs, microhabitat preference, and host specificity in fleas) or their population structure (mean abundance and/or prevalence) and (ii) host characteristics (sex and age) affect parasite and host species' contributions to parasite beta-diversity (SCBD and HCBD, respectively). We found that parasite species' morphological and ecological attributes were mostly not associated with their SCBD. In contrast, parasite SCBD, in both ungulates and rodents, significantly increased with either parasite mean abundance or prevalence or both. The effect of host characteristics on HCBD was detected in a few hosts only. In general, parasite infracommunities' beta-diversity appeared to be driven by variation in parasite species rather than the uniqueness of the assemblages harboured by individual hosts. We conclude that some ecological patterns (such as the relationships between SCBD and parasite abundance/prevalence) appear to be common and do not differ between different host-parasite associations in different geographic regions, whereas other patterns (the relationships between SCBD and parasite species' attributes) are contingent and depend on parasite and host identities.

Nestedness and beta diversity of gastrointestinal helminth communities in common warthogs, Phacochoerus africanus (Suidae), at 2 localities in South Africa.

Few studies have investigated the ecological interactions between wild species of Suidae and their parasites, leaving our knowledge concerning this host­parasite system fragmented. In the present study, we applied network studies to analyse community nestedness in helminth assemblages of common warthogs, Phacochoerus africanus (Gmelin) (Suidae). Helminth data were compiled from 95 warthogs, including young and adult males and females, from 2 different conservation areas in Mpumalanga and Limpopo Provinces, South Africa, collected monthly over a period of 1 year each. The aim was to study the effect of host sex, age and season of sampling on the structure of helminth infracommunities harboured by the warthogs and to search for non-random structural patterns in the warthog­helminth interaction networks. Furthermore, we investigated the influence of a warthog's age, sex and season of sampling on beta diversity and dark diversity of their helminth infracommunities. Lastly, we asked whether the effects of host sex, age and sampling season on helminth communities differed between the 2 localities. We found that helminth communities of warthogs were nested and host­parasite interactions were influenced by all 3 factors as well as combinations thereof. However, the resulting patterns differed at the 2 localities, indicating that local environmental processes are important drivers of community structure.

Impact of host sex and age on the diversity of endoparasites and structure of individual-based host-parasite networks in nyalas (Tragelaphus angasii Angas) from three game reserves in KwaZulu-Natal province, South Africa.

In recent years, numerous studies have examined the effect of host sex and age on the structure of parasite communities in several host taxa under various environmental conditions and in different geographic regions. However, the influence of such factors on the structure of host-parasite networks has received less attention, and remarkably few studies have been carried out on large terrestrial mammals. In this study, we investigated the effects of host age and sex on the parasite infra- and component communities of nyalas (Tragelaphus angasii) and on the structure of individual-based nyala-endoparasite networks. We also aimed to evaluate to what extent these effects vary spatially and if they are mediated by conservation management. Based on a data set of internal macroparasites of 74 nyalas from three game reserves in KwaZulu-Natal province, we found that host age strongly influenced parasite community structure as well as the structure of parasite-nyala networks, whereas host sex played a minor role. However, the effects of both host sex and age were mediated by environmental conditions and thus led to different patterns at the three localities. Our findings highlight that host-parasite communities from different localities should not be pooled when conducting host-parasite network and community studies as this may bias results and mask patterns that are typical for a given locality.

Some gastrointestinal nematodes and ixodid ticks shared by several wildlife species in the Kruger National Park, South Africa.

Parasite surveys were conducted for 1­2 years in the Kruger National Park (KNP), South Africa on blue wildebeest, impalas, greater kudus, common warthogs and scrub hares. The host associations of some of the gastrointestinal nematode species infecting ≥60% of at least one of the five host species, were determined. These were Agriostomum gorgonis, Cooperia acutispiculum, Cooperia connochaeti, Cooperia hungi, Cooperia neitzi, Cooperioides hamiltoni, Gaigeria pachyscelis, Haemonchus bedfordi, Haemonchus krugeri, Haemonchus vegliai, Impalaia tuberculata, Longistrongylus sabie, Strongyloides papillosus, Trichostrongylus deflexus and Trichostrongylus thomasi. Although the prevalence of Trichostrongylus falculatus did not exceed 50% in any host species, it was present in all five hosts. Nematodes in the KNP range from those exhibiting strict host associations to generalists. Nematode-host associations may be determined by host feeding patterns and habitat use. Eight ixodid tick species were commonly collected from the same animals and in 2­3 year long surveys from plains zebras and helmeted guinea fowls: Amblyomma hebraeum, Amblyomma marmoreum, Hyalomma truncatum, Rhipicephalus appendiculatus, Rhipicephalus decoloratus, Rhipicephalus evertsi evertsi, Rhipicephalus simus and Rhipicephalus zambeziensis. Host specificity was less pronounced in ixodid tick species than in nematodes and the immature stages of five tick species infested all host species examined.

Nematodes found in Nile crocodiles in the Kruger National Park, South Africa, with redescriptions of Multicaecum agile (Wedl, 1861) (Heterocheilidae) and Camallanus kaapstaadi Southwell & Kirshner, 1937 (Camallanidae).

Sixteen Nile crocodiles were collected in the Kruger National Park, South Africa and vicinity during 2010 and 2011. A total of 11 nematode species representing six families were recovered. Heterocheilids were the dominant group, comprising five species, with Dujardinascaris madagascariensis (Chabaud & Caballero, 1966) being the most prevalent (75%), followed by Ingwenascaris sprenti Junker & Mutafchiev, 2017 (68.8%), which was also the second most numerous nematode. While less prevalent (31.3%), Typhlophoros kwenae Junker & Mutafchiev, 2017 was the most abundant species. Micropleura huchzermeyeri Junker & Mutafchiev, 2017 (Micropleuridae) was collected from five crocodiles and Crocodylocapillaria sp. (Capillariidae) occurred in a single host. Three nematodes, Camallanus kaapstaadi Southwell & Kirshner, 1937, Spirocamallanus sp. (both Camallanidae) and Ascarophis sp. (Cystidicolidae), are considered accidental infections, likely ingested with the hosts' prey. Our findings of D. dujardini (Travassos, 1920), D. madagascariensis and Multicaecum agile (Wedl, 1861) in South Africa constitute new geographical records. Crocodylocapillaria sp. represents a new host and geographical record, while T. kwenae, I. sprenti and M. huchzermeyeri have been described as new species during the course of this survey. Multicaecum agile is here redescribed based on light and scanning electron microscopy. Previously undescribed morphological characters of C. kaapstaadi, typically a parasite of Xenopus spp. (Amphibia: Pipidae), but here found in two Nile crocodiles, are also presented.

Gastrointestinal helminths from the common warthog, Phacochoerus africanus (Gmelin) (Suidae), in KwaZulu-Natal Province, South Africa, with comments on helminths of Suidae and Tayassuidae worldwide.

Thirty warthogs, Phacochoerus africanus, were collected in the Pongola Game Reserve, South Africa and examined for helminths. Gastrointestinal helminth assemblages comprised Gastrodiscus aegyptiacus, the cestode genus Moniezia and seven species of nematodes. A single warthog harboured a metacestode of Taenia hydatigena in the mesenteries. No helminths were found in the heart, lungs or liver of the warthogs. Probstmayria vivipara and Murshidia spp. were the most prevalent as well as abundant helminth species, followed by Physocephalus sexalatus. The incidence of Moniezia did not differ between hosts of different sex or age. Numbers of Murshidia spp. were not affected by host sex, but were higher in adults than in juveniles. Conversely, burdens of Trichostrongylus thomasi were not affected by host age, but were higher in males than in females. While not highly significant, helminth assemblages in male warthogs were more species rich than in females. Helminth communities in the three genera of wild sub-Saharan suids are largely unique, but Ph. africanus and Hylochoerus meinertzhageni share more worm species with each other than with Potamochoerus larvatus, possibly because the former two are more closely related. Overlap between helminth communities of African wild suids and those of other suids and Tayassuidae worldwide is limited.

Pentastome assemblages of the Nile crocodile, Crocodylus niloticus Laurenti (Reptilia: Crocodylidae), in the Kruger National Park, South Africa.

Thirty-two specimens of the Nile crocodile, Crocodylus niloticus Laurenti (Reptilia: Crocodylidae), from the Kruger National Park, South Africa, and its vicinity were examined for pentastomid parasites during 1995 to 1999 and 2010 to 2011. Pentastomid parasites occurred throughout the year and were widespread in the study area with an overall prevalence of 97% and an overall mean abundance of 23.4 (0-81). Pentastome assemblages comprised six species in three sebekid genera: Alofia nilotici Riley et Huchzermeyer, 1995, A. simpsoni Riley, 1994, Leiperia cincinnalis Sambon, 1922, Sebekia cesarisi Giglioli in Sambon, 1922, S. minor (Wedl, 1861) and S. okavangoensis Riley et Huchzermeyer, 1995. The possible influence of host age, gender and geographic location (river system) on pentastome prevalence, abundance and species richness was investigated. Generally, neither host age, gender nor locality did affect infracommunities, likely because all hosts examined were adult or subadult and displayed comparable foraging behaviour, resulting in similar exposure pathways to fish intermediate hosts. Additionally, the longevity of pentastomids would contribute to accumulative infections as hosts mature. Structuring of pentastome assemblages was observed in as far as S. minor was the dominant species based on overall prevalence and abundance, followed by the equally common species S. cesarisi and L. cincinnalis. With an overall prevalence ranging from 34% to 41% and relatively low abundances, A. nilotici, A. simpsoni and S. okavangoensis form the rarer component of pentastome communities.

Ufudia, a replacement name for Pelonia Junker & Boomker, 2002 (Pentastomida: Sebekidae) from South African terrapins.

In order to accommodate a new species of pentastomid parasite, Pelonia africana Junker & Boomker, 2002 (Sebekidae), Junker & Boomker (2002) described the new genus Pelonia. This genus name is, however, preoccupied by Pelonia Grube, 1859 (Acari). Grube (1859) did not designate a type species, but Oudemans (1900: 141) recorded Pelonia as a possible synonym of Cepheus Koch, 1835 and listed P. foliosa as its type species, allegedly designated by Grube. Subías et al. (2012), accepted the synonymy of Pelonia with Cepheus Koch, 1835 (Acari: Compactozetidae). Norton & Ermilov (2014) provided a brief summary of the nomenclatural history of the mite genus Pelonia and proposed P. foliosa as being the nymph of Cepheus cepheiformis (Nicolet, 1855).

Redescription of Setaria graberi Shoho in Troncy, Graber & Thal, 1976 (Nematoda: Filarioidea) based on specimens from Redunca arundinum (Bovidae) in South Africa.

The filarial onchocercid Setaria graberi Shoho in Troncy, Graber & Thal, 1976 is redescribed from the abdominal cavity of Southern reedbuck, Redunca arundinum (Boddaert), in South Africa, including illustrations and scanning electron micrographs of important morphological features. Morphometric data for this species are provided for the first time. Setaria graberi is characterised by the possession of bifid deirids, and females having a distinctly bulbous tail. The slightly raised peribuccal crown forms a dumbbell-shaped unit with the cephalic elevations in apical view; the dorsal and ventral elevations, spaced 73-115 µm apart in females and 71-93 µm in males, carry two well-separated tips each. In dorsoventral view, the cephalic elevations appear more or less rectangular with a slightly notched apex and are narrow in comparison to the width of the anterior end. They are triangular in lateral view. Four cephalic and four external labial papillae are arranged in a laterally elongated rectangle each. The species is distinguished from other Setaria Viborg, 1795 species that possess bifid deirids or occur in members of the same host genus. The presence of S. graberi in R. arundinum in South Africa constitutes a new host and geographic record.

Nematodes of the small intestine of African buffaloes, Syncerus caffer, in the Kruger National Park, South Africa.

The abundance and distribution of parasitic helminths in populations of African buffaloes, Syncerus caffer, have not been well documented. A total of 28 buffaloes of different ages and sexeswere sampled in the Kruger National Park, South Africa, for nematodes of the small intestine. Three nematode species were identified, namely Cooperia fuelleborni, Cooperia hungi and Trichostrongylus deflexus, with C. hungi being a new country record for African buffalo in South Africa. The overall prevalence was 71%and the average number of worms was 2346 (range: 0-15 980). This is a small burden for such a large mammal. Sex, age and body condition of the buffaloes had no significant effect on worm occurrence.

Parasites of South African wildlife. XIX. The prevalence of helminths in some common antelopes, warthogs and a bushpig in the Limpopo province, South Africa.

Little work has been conducted on the helminth parasites of artiodactylids in the northern and western parts of the Limpopo province, which is considerably drier than the rest of the province. The aim of this study was to determine the kinds and numbers of helminth that occur in different wildlife hosts in the area as well as whether any zoonotic helminths were present. Ten impalas (Aepyceros melampus), eight kudus (Tragelaphus strepsiceros), four blue wildebeest (Connochaetes taurinus), two black wildebeest (Connochaetes gnou), three gemsbok (Oryx gazella), one nyala (Tragelaphus angasii), one bushbuck (Tragelaphus scriptus), one waterbuck (Kobus ellipsiprymnus), six warthogs (Phacochoerus aethiopicus) and a single bushpig (Potamochoerus porcus) were sampled from various localities in the semi-arid northern and western areas of the Limpopo province. New host-parasite associations included Trichostrongylus deflexus from blue wildebeest, Agriostomum gorgonis from black wildebeest, Stilesia globipunctata from the waterbuck and Fasciola hepatica in a kudu. The mean helminth burden, including extra-gastrointestinal helminths, was 592 in impalas, 407 in kudus and blue wildebeest, 588 in black wildebeest, 184 in gemsbok, and 2150 in the waterbuck. Excluding Probstmayria vivipara, the mean helminth burden in warthogs was 2228 and the total nematode burden in the bushpig was 80. The total burdens and species richness of the helminths in this study were consistently low when compared with similar studies on the same species in areas with higher rainfall. This has practical implications when animals are translocated to areas with higher rainfall and higher prevalence of helminths.

Proctocaecum gairhei n. sp. (Digenea: Cryptogonimidae: Acanthostominae) from Gavialis gangeticus (Gmelin) in Nepal and a revised cladogram of Proctocaecum Baugh, 1957.

Proctocaecum gairhei n. sp. is described from the gharial Gavialis gangeticus (Gmelin) in Nepal. The new taxon can be distinguished from all other species of Proctocaecum Baugh, 1957 by the combination of the following morphological characters: a single row of 23 cephalic spines, lateral anal pores opening at uneven levels, a forebody accounting for 16% of the total body length (TBL), an oral to ventral sucker width ratio of 1:0.7, an oral sucker to pharynx width ratio of 1:0.6 and uterine loops that occupy 59-67% of the TBL. While sharing some morphological characteristics with Acanthostomum slusarskii Kalyankar, 1977 from Crocodylus palustris (Lesson) in India, P. gairhei n. sp. is distinguished from the latter by possessing the solid muscular gonotyl diagnostic for Proctocaecum, by the number of cephalic spines, position of the anal pores and egg size. The existing cladogram for Proctocaecum was expanded using the character information of P. gairhei n. sp. The host range of Proctocaecum is now known to include all three families of the order Crocodylia.

Genetic characterization and phylogenetic position of Echinococcus felidis (Cestoda: Taeniidae) from the African lion.

Echinococcus felidis had been described in 1937 from African lions, but was later included in Echinococcus granulosus as a subspecies or a strain. In the absence of any genetic characterization, most previous records of this taxon from a variety of large African mammals remained unconfirmed due to the lack of diagnostic criteria and the possible confusion with the sympatric E. granulosus sensu stricto, Echinococcus ortleppi and Echinococcus canadensis. In this study, we obtained taeniid eggs from lion feces in Uganda and amplified DNA from individual eggs. Mitochondrial and nuclear DNA sequences showed similarities with those of other Echinococcus spp., but high values of percentage divergence of mitochondrial genes indicated the presence of a distinct species. In a second step, we compared this material with the preserved specimens of adult E. granulosus felidis, which had been identified morphologically approximately 40 years ago in South Africa. All DNA fragments (<200 bp) that could be amplified from the adults showed 100% similarity with the Ugandan material. In the phylogenetic tree of Echinococcus which was constructed from the mitochondrial genes, E. felidis is positioned as a sister taxon of E. granulosus sensu stricto. The data obtained will facilitate the development of diagnostic tools necessary to study the epidemiology of this enigmatic parasite.

A 12-month survey of the gastro-intestinal helminths of antelopes, gazelles and giraffids kept at two zoos in Belgium.

Faecal egg count patterns and clinical signs associated with gastro-intestinal (GI) nematodes of 107 zoo ruminants were monitored at fortnightly intervals for 1 year. The ruminants in this study were kept under different husbandry conditions at two sites of the Royal Zoological Society of Antwerp, the Antwerp Zoo and the Animal Park Planckendael. Artiodactylids involved were Arabian oryx (Oryx leucoryx), scimitar-horned oryx (Oryx dammah), bongos (Tragelaphus euryceros isaaci), sitatungas (Tragelaphus spekii gratus), common eland (Taurotragus oryx), impala (Aepyceros melampus), slender-horned gazelles, (Gazella leptoceros), blue wildebeest (Connochaetes taurinus taurinus), Kordofan giraffes (Giraffe camelopardalis antiquorum) and okapi (Okapia johnstoni). Nematode eggs were recovered from 586 of 1606 (36.5%) individual faecal samples, using flotation techniques. Infection levels were distinctly low at Antwerp Zoo, probably due to zero grazing and daily dung removal. At Planckendael, the herds of Arabian oryx, scimitar-horned oryx and slender-horned gazelles showed markedly higher egg counts than the other herds, with more than 10% of the faecal egg counts having more than 100 eggs per gram (epg) and maximum faecal egg counts of 600, 750 and 1350 epg, respectively. Faecal egg counts increased during the mid-grazing season (July) and peaked at the end of the grazing season (October). No clinical signs, such as loss of faecal consistency, could be correlated with faecal egg counts (P > 0.05). With the exception of significantly more Nematodirus spp. eggs that were present in juvenile eland, no differences in faecal egg counts could be found between the sexes and different age groups. Abomasa and intestines of 17 animals that died during the survey were available for total worm counts. In one Arabian oryx, four slender-horned gazelles and one sitatunga low burdens ranging from 200 to 14,300 were found. Nematode species recovered were Camelostrongylus mentulatus from the abomasa and Trichostrongylus retortaeformis, Nematodirus fillicollis, Capillaria spp. and Trichuris spp. from the intestines. Our findings suggest different nematode infection levels between herds, which are mainly due to husbandry conditions but to a lesser extent to species- or individual susceptibility. Identification of ungulates that are highly infected and knowledge of the seasonal variation of their helminths can contribute greatly to a well-adjusted species-specific management and helminth control program.

A 12-month survey of gastrointestinal helminth infections of cervids kept in two zoos in Belgium.

Infections with helminths are a major health issue in captive and wild deer. In this study, fecal egg count patterns and clinical signs associated with gastrointestinal nematodes were assessed for 12 mo in nine cervid herds kept under different husbandry conditions at two sites. At site 1, an urban zoo, fecal egg counts remained low and no clinical signs of parasitic gastroenteritis were seen in the herds of fallow deer (Dama dama), Dybowski's deer (Cervus nippon dybowski), pudu (Pudu pudu), and reindeer (Rangifer tarandus tarandus). Helminth infection at this site may have been successfully prevented by daily dung removal of the small sandy-soil enclosures, and applying routine anthelmintic treatment was not justified. At site 2, a wild animal park, involved species were red deer (Cervus elaphus hippelaphus), Nelson's elk (Cervus elaphus nelsoni), Père David's deer (Elaphurus davidianus), European elk (Alces alces alces), and reindeer (Rangifer tarandus tarandus). Nematode eggs were frequently encountered in herds of red deer, Nelson's elk, and European elk, which were kept on larger, grassy enclosures that were irregularly cleaned. The trimodal pattern of fecal egg counts in herds from the wild animal park, consisting of a small spring rise in June, a peak in October, and a small rise in February, indicates that infective larvae on pastures are the main source of infection. In addition, routine anthelmintic treatment with fenbendazole in April and July limited egg shedding, but reinfection rapidly occurred. In two European elk and one reindeer, increasing fecal egg counts were associated with loss of fecal consistency and reduced appetite. Three genera and three species of nematodes were recovered at necropsy of one red deer and three Nelson's elk: Spiculopteragia spiculoptera, Trichostrongylus spp., Nematodirus filicollis, Capillaria spp., Oesophagostomum radiatum, and Trichuris spp., with total worm counts between 950 and 8,700.