Helminth parasites of intermingling axis deer, wild swine and domestic cattle from the island of Molokai, Hawaii

Helminth infections of axis deer (Cervus axis), wild swine (Sus scrofa) and domestic cattle (Bos taurus) were studied among intermingling herds on the Puu-O-Hoku Ranch, Molokai, Hawaii. Twenty-four species of helminths were collected from the 10 deer, 10 swine and 10 cattle. Capillaria bovis, Cooperia punctata, Haemonchus contortus and Trichostrongylus axei infected both axis deer and cattle, whereas Gongylonema pulchrum infected both axis deer and wild swine. None of the species of helminths occurred in both wild swine and cattle nor was any species found in all three hosts. Wild swine and domestic cattle supported separate and distinct helminth communities. In contrast, the helminth community of axis deer appeared to be derived from the helminth communities of cattle and wild swine and consisted only of those species capable of parasitizing either a broad range of ruminants or many mammalian taxa.     

Helminth infections of wild boars (Sus scrofa) in the Bursa province of Turkey

The present study aimed to investigate the status of helminth infections in wild boars in the Bursa province of Turkey. For this purpose, during 2007-2008, 27 wild boars were necropsied and examined for helminths. Individual samples of tongue and diaphragm from 27 necropsied wild boars and an additional 22 tongue and diaphragm samples provided by hunters were examined by trichinoscopy and artificial digestion for Trichinella spp. larvae. Twenty animals (74%) were identified as being infected with at least one helminth species. Twelve species of helminths were detected, with the following prevalence rates: Metastrongylus apri (59%), Metastrongylus salmi (52%), Metastrongylus pudendotectus (52%), Dicrocoelium dendriticum (33%), Globocephalus urosubulatus (22%), Macracanthorhynchus hirudinaceus (19%), Gongylonema pulchrum (11%), Physocephalus sexalatus (7%), Trichuris suis (7%), Ascarops strongylina (4%), Hyostrongylus rubidus (4%) and Taenia hydatigena larvae (4%). Generally, lungworms were the predominant helminths. The highest mean abundance was observed for M. pudendotectus, and the lowest was determined for T. hydatigena larvae. Significant differences in the prevalence and intensity were found for D. dendriticum with respect to host age and sex, respectively. The mean intensity of M. pudendotectus was significantly influenced by the sex and age of the wild boars. This study is the first report describing the presence of M. salmi, M. pudendotectus, D. dendriticum, G. urosubulatus, M. hirudinaceus, P. sexalatus, A. strongylina and H. rubidus in wild boars in Turkey. All analysed muscle samples were negative for Trichinella spp. larvae.     

Factors affecting the helminth community structure of adult collared peccaries in southern Texas

Four species of nematodes (Gongylonema pulchrum, Parabronema pecariae, Texicospirura turki, and Physocephalus sexalatus) and one species of cestode (Moniezia sp.) comprised the helminth fauna of adult collared peccaries (Tayassu tajacu) from the plains in southern Texas. The community structure of the helminth fauna of peccaries from this region was basically dissimilar to that from the more humid Gulf coastal prairies of southern Texas in composition (by the conspicuous absence of certain species) and relative abundance of shared species. The distributions of each of the three common species of helminths (G. pulchrum, T. turki, and P. sexalatus) were overdispersed. The effects of selected habitat variables operating across host subpopulations (delineated by condition and sex) and of the extrinsic variable of season on the dispersion patterns of the three common species of helminths were examined. The hypothesis that heterogeneity within the host population, rather than across the collective host population, is the main factor generating overdispersion in natural parasite populations was not confirmed for the three common species of helminths. Overdispersion in P. sexalatus resulted from seasonal changes across the collective host population, with the greatest abundances occurring during the cool season. Aggregated abundances of G. pulchrum resulted from variation generated across host sex subpopulations, while the dispersion patterns of T. turki appeared to be unaffected by the habitat variables examined in this study.     

Helminth parasites of the wild boar,Sus scrofa,in Luristan province,western Iran and their public health significance

Seven helminth species were obtained from 12 wild boars (Sus scrofa) during a survey from 2000 to 2001 in Luristan province, western Iran. These species include the cestode larvae Cysticercus tenuicollis (25%), C. cellulosae (8.3%), the nematodes Metastrongylus apri (41.6%), M. pudendotectus (16.6%), M. salmi (8.3%), Trichuris suis (8.3%) and the acanthocephalan Macracanthorhynchus hirudinaceus (41.6%). No trematodes were found. Seven wild boars (58.3%) were identified as having at least one helminth species. A single infection was detected in 16.6% of cases, but a three species infection covered the highest rank (25%). All these helminths have been reported from other areas of Iran including the north, northeast and southwest, but not in Luristan. Among seven helminths identified, at least three species are transmissible to humans. The public health significance of these helminths is discussed.     

Visceral helminth communities of sympatric mule and white-tailed deer from the Davis Mountains of Texas

Hybridizing populations of mule (Odocoileus hemionus) and white-tailed deer (O. virginianus) from the Davis Mountains of Texas were examined to determine similarities in species composition of their helminth communities and if abundances of helminth species in those communities varied across host species and seasonal factors. Only three cestode and three nematode species were recovered. There were very low abundances of species and little diversity in the helminth communities of both hosts. Common helminth species were shared by both deer, and the significant variance in abundances of three of the four most common helminth species appeared to result from differences in habitat preferences of the respective hosts. Our results indicated that analyses of helminth communities of deer from this geographical area do not provide a useful quantification technique for determining deer condition, degree of hybridization, or levels of intraspecific competition.     

The decay of similarity with geographical distance in parasite communities of vertebrate hosts

Aim The rate at which similarity in species composition decays with increasing distance was investigated among communities of parasitic helminths in different populations of the same host species. Rates of distance decay in similarity of parasite communities were compared between populations of fish and mammal hosts, which differ with respect to their vagility and potential to disperse parasite species over large distances. Location Data on helminth communities were compiled for several populations of three mammalian host species (Ondatra zibethicus, Procyon lotor and Canis latrans) and three fish host species (Perca flavescens, Catostomus commersoni and Esox lucius) from continental North America. Methods Distances between localities and similarity in the composition of helminth communities, the latter computed using the Jaccard index, were calculated for all possible pairs of host populations within each host species. Similarity values were then regressed against distance to see if they decayed at exponential rates, as reported for plant communities; the significance of the regressions was assessed using randomization tests. Results The number of hosts examined per population did not correlate with the number of helminth species found per population, and thus sampling effort is unlikely to have confounded the results. In four (two mammals and two fish) of the six host species, similarity in helminth communities decayed exponentially with distance. When the log of similarity is plotted against untransformed distance, the slopes obtained for the two fish species are lower than those obtained for the two mammalian host species. Main conclusions Similarity in the composition of parasite communities appears to decay exponentially with increasing distance in some host species, but not in all host species. The rate of decay is not necessarily associated with the vagility of the host. Although distance decay of similarity is generally occurring, it seems that other ecological processes, related either to the host or its habitat, can obscure it.     

Prevalence and distribution of pig helminths in the Dongting Lake Region (Hunan Province) of the People's Republic of China

The prevalence of helminths in pigs was investigated in five rural communities situated on the embankment of Dongting Lake in Zhiyang County, Hunan Province, People's Republic of China, in an area known to be endemic for Schistosoma japonicum. The helminth prevalences identified on the basis of faecal egg count analysis were: Oesophagostomum spp. (86.7%), Ascaris suum (36.7%), Metastrongylus spp. (25.8%), Strongyloides spp. (25.8%), Trichuris suis (15.8%), Globocephalus spp. (6.7%), Gnathostoma spp. (4.2%), Schistosoma japonicum (5.0%) and Fasciola spp. (1.3%). Post mortem examinations of a small number of pigs depositing eggs of different helminth species revealed the presence of Oesophagostomum dentatum, O. quadrispinulatum, A. suum, Metastrongylus apri, M. pudendotectus, T. suis, G. hispidum and Ascarops dentata. Prevalences of all helminths, with the exception of Oesophagostomum spp., were higher in young pigs (< 8 months old) compared with adult pigs. Prevalences of trematodes were very low, especially for S. japonicum which had decreased dramatically compared with previous reports from this area of P.R. China, whereas prevalences of nematodes were generally in agreement with those reported from other Yangtze River Provinces. Results from helminth prevalence studies in pigs, conducted in other provinces of P.R. China between 1987 and 1997, are presented and discussed. It was concluded that a government helminth control programme, implemented in 1995 to control S. japonicum infection in pigs in Hunan Province, may have resulted in a greatly reduced prevalence of S. japonicum in pigs in this region.     

Effects of season and physical condition on the gastrointestinal helminth community of white-tailed deer from the Texas Edwards Plateau

Eighty-six adult female white-tailed deer, Odocoileus virginianus (Zimmermann), collected over a 12-mo period in the Texas Edwards Plateau, harbored six species of nematodes (Haemonchus contortus, Gongylonema pulchrum, Oesophagostomum venulosum, Ostertagia ostertagi, Cooperia sp., and Apteragia odocoilei), and two cestodes (Moniezia sp. and Taenia hydatigena). The patterns of distribution of the three common species of gastrointestinal helminths (H. contortus, O. venulosum, and G. pulchrum) were overdispersed. When analyzed for the main and interactive effects of the extrinsic and intrinsic variables of season and physical condition, respectively, aggregated abundances in H. contortus and O. venulosum appeared to result from the main effect of seasonal changes operating over the collective populations of these two species rather than from the intrinsic factor of physical condition operating within selected subpopulations. Abomasal parasite counts do not appear to be a useful index for monitoring herd condition of white-tailed deer from this geographic region.     

Helminth communities of three sympatric species of shorebirds (Charadrii) from four summer seasons at Bristol Bay, Alaska

Helminth communities in sympatric black turnstones (Arenaria melanocephala), ruddy turnstones (Arenaria interpres), and dunlin (Calidris alpina) were examined over 4 summers in Bristol Bay, Alaska. The compound community, made up of component communities of all 3 species of hosts for 4 summer seasons (n=164), consisted of 43 helminth species, with cestodes, especially Anomotaenia clavigera, accounting for 47% of the helminth species and 95% of the abundance. The black turnstone had significantly higher species richness and abundance than either the ruddy turnstone or dunlin. The congeneric black and ruddy turnstone component communities were the most similar, and the dunlin's was the least similar. New helminth species continued to be acquired in all 3 host species during years 2 to 4. There was no significant difference for abundance among sample years for each of the 3 species of host. The 3 component communities all included a predictable suite of helminths with 1 dominant species and 4 to 5 associates, a large number of less-predictable species, and a greater prevalence and abundance of cestode species. Consistencies over time included high diversity, low evenness, low species richness (<5), and continued recruitment of small numbers of helminth species with low prevalence and abundance. There was minimal circulation of helminth species between the dunlin and the 2 turnstone species, indicating a considerable degree of specialization, particularly among species of cestodes.     

Helminth communities of native and introduced fishes in Lake Pátzcuaro, Michoacán, México

In Lake Pátzcuaro in the Mesa Central of México, a total of 19 species of helminths was found in 598 fishes and comprised five digeneans, two monogeneans, four cestodes, one acanthocephalan and seven nematodes, of which ten species were represented by larval or immature states. The richest and most diverse helminth communities were found in the native carnivorous goodeid Alloophorus robustus. In general, the helminth communities in the different fish species were not particularly species rich and the parasite assemblages were numerically dominated by larvae of the bird trematode, Posthodiplostomum minimum. Patterns of helminth community richness and diversity were similar to those previously observed in north-temperate freshwater fishes. Most enteric helminths occurred with low abundance and only a small proportion of the gut helminth communities was numerically dominated by any one species. Helminths dominating their enteric communities showed some level of host specificity. Helminth communities in carnivorous fish species were generally richer than those in herbivores and detritivores, with the exception of the predominantly herbivorous Goodea atripinnis. The helminth fauna of introduced fishes, Cyprinus carpio, Micropterus salmoides and Oreochromis niloticus , consisted of either few or no host-specific adult helminth(s) translocated from their original geographical areas and by larval stages of helminths of piscivorous birds. Based on the geological history of the area and the biogeography of the endemic fish fauna, it is hypothesized that host-switching and relationships with the nearctic fauna have been fundamental in determining the helminth fauna of the endemic fish hosts.     

The structure of helminth communities

The existence of both interactive and isolationist communities of helminths, each of which may be produced by any one of several processes, necessitates a pluralistic view of helminth community structure. A scheme involving three hierarchical scales of communities of helminths is proposed. Interactions between species occur in infracommunities, and should be looked for there. Interactions are often clearer in smaller groups of species, either among core species or within guilds. Infracommunities are samples of helminth communities at two larger scales: the helminth community in the host population provides core species, largely specialists, and that in the host community provides the generalists and the satellite species. The richness of each of the two large-scale communities is affected by various ecological, historical and evolutionary factors. The different concepts which have been applied to parasite communities are based on these factors.     

Host sharing and host manipulation by larval helminths in shore crabs: cooperation or conflict?

Larval helminths of different species that share the same intermediate host and are transmitted by predation to the same definitive host may cooperate in their attempts to manipulate the behaviour of the intermediate host, while at the same time having conflicts of interests over the use of host resources. A few studies have indicated that intermediate hosts harbouring larval helminths have altered concentrations of neurotransmitters in their nervous system, and thus measuring levels of neurotransmitters in host brains could serve to assess the respective and combined effect of different helminth species on host behaviour. Here, we investigate potential cooperation and conflict among three helminths in two species of crab intermediate hosts. The acanthocephalan Profilicollis spp., the trematode Maritrema sp. and an acuariid nematode, all use Macrophthalmus hirtipes (Ocypodidae) as intermediate host, whereas Profilicollis and Maritrema also use Hemigrapsus crenulatus (Grapsidae). All three helminths mature inside gulls or other shore birds. There was a significant decrease in the mean volume of Profilicollis cystacanths as the intensity of infection by this parasite increased in H. crenulatus, the only host in which this was investigated; however, there was no measurable effect of other helminth species on the size of acanthocephalans, suggesting no interspecific conflict over resource use within crabs. There was, in contrast, evidence of a positive interspecific association between the two most common helminth species: numbers of Profilicollis and Maritrema were positively correlated among crabs, independently of crab size, in M. hirtipes but not H. crenulatus. More importantly, we found that the total number of larval helminths per crab correlated significantly, and negatively, with concentrations of serotonin in crab brains, again only in M. hirtipes; numbers of each parasite species separately did not covary in either crab species with serotonin or dopamine, the other neurotransmitter investigated in this study. The relationship with serotonin appears due mainly to numbers of Profilicollis and Maritrema and not to nematodes. This is the first demonstration of a potentially synergistic manipulation of host behaviour by different helminth species, one that appears host-specific; our results also point toward the neurobiological mechanism underlying this phenomenon.     

Helminth communities in avocets: importance of the compound community

This paper reports patterns of similarity and overlap in species presence and patterns of linear distribution of intestinal helminths in 22 avocets from 4 populations. Avocets collected from ephemeral bodies of water in Alberta and Manitoba had communities composed largely of species that are avocet specialists plus some that are host generalists. The composition of helminth communities in these hosts was similar to that reported in earlier surveys of avocet helminths. There was little evidence for competition between helminth species in these communities. In contrast, avocets collected from permanent bodies of water in Alberta had communities composed largely of species that are specialists in various duck species, particularly lesser scaup. These helminths were superimposed on the normal community, fitting into linear gaps along the intestine but also overlapping the distributions of avocet specialists. These lesser scaup specialists exhibit interactive patterns amongst themselves and, to some extent, with avocet specialists. Helminth communities in avocets from ephemeral bodies of water have vacant niches and are largely isolationist in nature. Those in avocets from permanent bodies of water are saturated and are more interactive in nature.     

Species associations among larval helminths in an amphipod intermediate host

Larval helminths that share the same intermediate host may or may not also share the same definitive hosts. If one or more of these helminth species can manipulate the phenotype of the intermediate host, there can be great advantages or severe costs for other helminths resulting from co-occurring with a manipulator, depending on whether they have the same definitive host or not. Among 2372 specimens of the amphipod Echinogammarus stammeri collected from the river Brenta, northern Italy, there was a positive association between two acanthocephalan species with the same fish definitive hosts, the relatively common Pomphorhynchus laevis and the much less prevalent Acanthocephalus clavula.The number of cystacanths of P. laevis per infected amphipod, which ranged from one to five, did not influence the likelihood that the amphipod would also host A. clavula. A third acanthocephalan species, Polymorphus minutus,which matures in birds, showed no association with either of the two other species. These results show that associations among helminth species in intermediate hosts are not random, and are instead the product of selection favouring certain pathways of transmission.     

The helminth community of the wood mouse, Apodemus sylvaticus, in the Sierra Espuna, Murcia, Spain

The helminth community of the wood mouse, Apodemus sylvaticus, in the Sierra Espuna was characterized after a complete analysis of its helminth community component and infracommunity structure relative to host age, sex and year of capture. The helminth community comprised 13 species: one trematode, four cestodes and eight nematodes. The cestode Pseudocatenotaenia matovi and the nematode Syphacia frederici were the most prevalent and abundant helminth species, respectively. Sixty four percent of mice analysed presented helminths with a direct cycle and 42% presented helminths with an indirect cycle. The helminth community presents a low diversity with infracommunities usually made up of only one or two helminth species. Host age and year of capture seem to play a major role in determining species richness and helminth diversity, but not in determining the abundance of helminths. Host sex does not seem to affect the infection rate nor the diversity. Further studies on more samples of wood mice and other small mammal species in this regional park are needed to explore any possible interactions between helminth communities in the host populations.     

The predictability of helminth community structure in space: a comparison of fish populations from adjacent lakes

Patterns in helminth community structure can suggest that various processes are acting to shape parasite communities into organised, non-random assemblages of species. It is not clear, however, whether a pattern observed in one host population at one time would be observed again at another time, or at the same time in a different but comparable host population. Here, we test the repeatability of parasite community structure in space, and to a lesser extent time, with data on helminth parasites of two fish species, perch Perca fluviatilis and roach Rutilus rutilus, collected in different seasons from four adjacent lakes in Central Finland. Since populations of the same fish species harbour the same parasite species and were sampled in the same way, we would expect similar patterns in the structure of their helminth parasites if the same structuring processes are acting in all lakes. We found that no pairwise association between the most common helminth species were observed consistently between seasons within lakes, or among lakes during the same season. Similarly, nested subset patterns of species assembly were observed in some samples, but not consistently between seasons or among lakes. The lack of repeatability in space and between seasons shown by these analyses indicates that although helminth community structure often departs from randomness, it does not do so in a consistent and predictable manner. There may be some general, large-scale processes acting to structure helminth communities, but local or seasonal influences can often either mask their action, or play more important roles themselves.     

Helminths as vectors of pathogens in vertebrate hosts: a theoretical approach

Pathogens frequently use vectors to facilitate transmission between hosts and, for vertebrate hosts, the vectors are typically ectoparasitic arthropods. However, other parasites that are intimately associated with their hosts may also be ideal candidate vectors; namely the parasitic helminths. Here, we present empirical evidence that helminth vectoring of pathogens occurs in a range of vertebrate systems by a variety of helminth taxa. Using a novel theoretical framework we explore the dynamics of helminth vectoring and determine which host-helminth-pathogen characteristics may favour the evolution of helminth vectoring. We use two theoretical models: the first is a population dynamic model amalgamated from standard macro- and microparasite models, which serves as a framework for investigation of within-host interactions between co-infecting pathogens and helminths. The second is an evolutionary model, which we use to predict the ecological conditions under which we would expect helminth vectoring to evolve. We show that, like arthropod vectors, helminth vectors increase pathogen fitness. However, unlike arthropod vectors, helminth vectoring increases the pathogenic impact on the host and may allow the evolution of high pathogen virulence. We show that concomitant infection of a host with a helminth and pathogen are not necessarily independent of one another, due to helminth vectoring of microparasites, with profound consequences for pathogen persistence and the impact of disease on the host population.     

Helminth community composition, structure, and pattern in six dove species (Columbiformes: Columbidae) of South Texas

The helminth community composition and structure of 6 species of columbids residing in south Texas are reported and compared herein. Sixty individuals of the following species, rock pigeons (Columba livia [RP]), mourning doves (Zenaida macroura [MD]), Eurasian collared-doves (Streptopelia decaocto [ECD]), white-winged doves (Zenaida asiatica [WWD]), and common ground doves (Columbina passerina [CGD]), and 48 Inca doves (Columbina inca [ID]) were collected during the summer of 2006 and examined for helminths. Twelve helminth species were found (9 nematodes and 3 cestodes), representing 486 individuals. Nematodes numerically dominated the component community in all host species. Overall, helminth prevalence was similar among host sex within all dove species. However, prevalence of Skrjabinia bonini and Hymenolepis sp. in RPs was significantly different among host age groups (P = 0.01, P = 0.0002, respectively). Likewise, prevalence of Killigrewia delafondi was higher (P = 0.0001) in adult WWDs. Based on percent similarity and Jaccard's coefficient of community indices, helminth component communities were dissimilar, and the number of shared helminth species varied among host species. Data from this study suggest that the environment surrounding preferred host habitat and foraging strategies of each host species is the driving force behind helminth component communities. This study emphasizes the importance of examining co-occurring hosts at both local and regional scales to elucidate helminth community structure and patterns.     

Helminth community structure of sympatric eastern American toad, Bufo americanus americanus, northern leopard frog, Rana pipiens, and blue-spotted salamander, Ambystoma laterale, from southeastern Wisconsin

One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.     

Intestinal helminths of capybaras, Hydrochoerus hydrochaeris, from Venezuela

Quantitative parameters of intestinal helminth species and their potential relations to host characteristics in a population of capybaras (Hydrochoerus hydrochaeris) from Venezuela are reported for the first time. The intestines of 40 capybaras were collected during the 1992-annual harvest at Hato El Cedral. Six helminth species were found: 2 cestodes (Monoecocestus macrobursatum, M. hagmanni), 2 nematodes (Viannella hydrochoeri, Protozoophaga obesa), and 2 trematodes (Hippocrepis hippocrepis, Taxorchis schistocotyle). This is the first report for M. macrobursatum in Venezuela. Helminth abundance did not differ between sexes or age classes. Although patterns of distribution for all helminth species were overdispersed, the high prevalence found for all species (over 70%) and the high abundance observed for nematodes made it difficult to assess the effect that these helminths may produce on capybaras. Nevertheless, the negative associations found between the body condition of capybaras and helminth intensity for M. macrobursatum and V. hydrochoeri, might be pointing out potential host population regulatory role for these parasites which require further research.