Intestinal helminths of spiny mice (Acomys cahirinus dimidiatus) from St Katherine's Protectorate in the Sinai, Egypt

Spiny mice, Acomys cahirinus dimidiatus, inhabiting the wadis close to St Katherine in the mountains of the Sinai peninsula, were trapped and their helminth parasites were studied. Sixty one mice provided faeces for analysis and 27 were killed and autopsied. Six species of helminths were recorded (the spirurid nematodes, Protospirura muricola (74.1%) and Mastophorus muris (11.1%), the oxyuroid nematodes, Dentostomella kuntzi (59.3%), Aspiculuris africana (3.7%), and Syphacia minuta (3.7%) and the hymenolepidid cestode Rodentolepis negevi (18.5%)). The spirurids were the dominant species present, accounting for up to 0.87% of total host body weight. Analysis of worm weights and lengths suggested that transmission had been taking place in the months preceding our study. No sex difference in the prevalence or abundance of spirurids was detected. Significant differences were identified in the abundance of total nematode burdens and the mean helminth species richness between the three wadis which provided multiple captures of mice. There was also a marked effect of host age on both parameters. A highly significant positive correlation between spirurid egg counts and total worm biomass indicated that non-invasive techniques based on egg counts could be used to quantify worm burdens and when this technique was applied to a larger sample size (n = 61), a significant difference between sites but no host sex or age effects were detected for spirurid faecal egg counts. The data suggest that there are differences between helminth component communities infecting spiny mice in different neighbouring wadis, a hypothesis which will be explored further through our continuing studies in the Sinai.     

A comparison of the intestinal helminth communities of Equidae in Southern Africa

The intestinal helminth communities of 8 horses, 12 donkeys, 21 Hartmann's mountain zebras, and 44 Burchell's zebras were compared using the original data from 6 studies in South Africa and Namibia. Necropsy and worm recovery techniques were comparable between the studies. Sixty helminth species (58 nematode, 1 cestode, and 1 trematode species) were recorded. There were significant differences in the helminth community structures of the 4 Equus species. The helminth communities of the 2 closely related zebra subspecies were most similar, and they jointly shared 7 helminth species with donkeys and only 1 with horses. Geographic variation and host-mixing contributed to the helminth species composition. Multiple confamilial species infections were the norm in the donkeys and zebra subspecies, and no single-species infection was recorded for the Strongylidae. Congeneric species were commonly recorded in 3 genera (Cyathostomum, Cylicocyclus, and Cylicostephanus). The shape of the occupancy frequency distributions for the donkeys and zebra subspecies was multimodal, with no clear satellite or core modes. Despite the presence of environmental variability and comparatively low parasite-host specificity, the phylogenetic signal within Equus helminth communities remains strong.     

Population biology of multispecies helminth infection: competition and coexistence

The role that interspecific interactions play in shaping parasite communities is uncertain. To date, models of competition between helminth species have assumed that interaction occurs through parasite-induced host death. To our knowledge, there has been no theoretical exploration of other forms of competition. We examine models in which competition acts at the point of establishment within the host, and at the time of egg production by the adult worm. The models used are stochastic and we allow hosts to vary in their rate of exposure to infective larvae. We derive the Lotka-Volterra model of competition when exposure is homogenous and thus demonstrate that two helminth species cannot coexist on a single limiting resource. We show that coexistence of species is promoted by heterogeneity in host exposure provided that the rates of exposure to the two species are not perfectly correlated, and, if they are positively correlated, provided that the degree of heterogeneity in host exposure is similar for the two competing helminth species. These results are robust to the mechanism of competition.     

Comprehensive and Comparative Analysis of the Helminth Fauna of the Dominant Colonial Seabird Species on the Murman Coast

This work reports the results of the long-term study of the helminth fauna of the dominant colonial seabird species (common guillemots, Brünnich’s guillemots, kittiwakes, great black-backed gulls, and herring gulls) in the western, central, and eastern parts of the Murman coast. The presence of 52 parasitic worm species (18 trematode species, 21 cestode species, 11 nematode species, and 2 acanthocephalan species) was demonstrated. In the western, central, and eastern regions, the respective numbers of 32, 43, and 34 helminth species were registered. Twenty species of seabird parasites were found to be ubiquitous along the Murman coastal zone. Local foci of the trematode Maritrema arenaria and cestode Laricanthus lateralis infestation along the western part of the Murmansk coast were revealed. The highest helminth species diversity (38 species) was demonstrated for the herring gull; the lowest (7 species), for the common guillemot. Four parasite species were common for the helminth fauna of all bird species. A significant number of helminth infections were mixed ones (complexes of two or three bird species with the common helminth species in different combinations). The highest similarity of the helminth fauna was revealed in the pairs of the common guillemot and the Brünnich’s guillemot and of the great black-backed gull and the herring gull. A few parasitic worm species detected only in a single bird species were registered. The particularities and the possible causes of the geographical distribution and specificity of helminthes were analyzed, and a comprehensive assessment of the parasitological situation in the Murman coastal zone was performed.     

Interspecific associations among larval helminths in fish

Various processes can generate associations between the larvae of different helminth species in their fish intermediate or paratenic host. We investigated the pairwise associations among larval helminth species in eight different fish populations, using two different coefficients of associations, in order to determine in what situations they are strongest. All helminth species included use the fish studied as either their second intermediate host or their paratenic host, and are acquired by the fish when it ingests an infected first intermediate host. The intensity of infection correlated positively with fish length for most helminth species. Pairs of species which both exhibited positive correlations with fish length tended to be more strongly associated with one another, although this tendency was not pronounced. Similarity in life cycle had a more important influence on pairwise associations. Among the 62 pairwise associations that could be computed, pairs of helminth species that shared both first intermediate hosts and definitive hosts were the most strongly associated, followed by pairs that shared only one other host, and finally by pairs that did not share other hosts. The results suggest that assemblages of larval helminth parasites in fish are not random collections of locally available species, but rather structured packets of larval parasites that travel together along common transmission routes.     

Helminth-modified pulmonary immune response protects mice from allergen-induced airway hyperresponsiveness

It has been shown that the presence of certain helminth infections in humans, including schistosomes, may reduce the propensity to develop allergies in infected populations. Using a mouse model of schistosome worm vs worm + egg infection, our objective was to dissect the mechanisms underlying the inverse relationship between helminth infections and allergies. We have demonstrated that conventional Schistosoma mansoni egg-laying male and female worm infection of mice exacerbates airway hyperresponsiveness. In contrast, mice infected with only schistosome male worms, precluding egg production, were protected from OVA-induced airway hyperresponsiveness. Worm-infected mice developed a novel modified type 2 cytokine response in the lungs, with elevated allergen-specific IL-4 and IL-13 but reduced IL-5, and increased IL-10. Although schistosome worm-only infection is a laboratory model, these data illustrate the complexity of schistosome modulation of host immunity by the worm vs egg stages of this helminth, with the potential of infections to aggravate or suppress allergic pulmonary inflammation. Thus, infection of mice with a human parasitic worm can result in reduced airway inflammation in response to a model allergen.     

Inter- and intra-morph patterns in helminth communities of sympatric whitefish morphs

Infection patterns of trophically transmitted helminth parasites were compared with feeding ecology in two sympatric whitefish Coregonus lavaretus morphs from two lake systems in northern Norway. In both lakes, the pelagic morph was an obligate zooplanktivore, while the benthic morph utilized both the benthivore and zooplanktivore trophic niches. The differences in niche utilization between the two morphs were associated with differences in trophic morphology (gill raker numbers), suggesting that they were genetically dissimilar and reproductively isolated. The benthic morph had the highest number of helminth species, probably because they exhibited a broader niche width compared to the pelagic morph. In both lakes, the species composition and intensities of helminths reflected the trophic diversification of the whitefish ecotypes with respect to different habitat choice (benthic v . pelagic) and dietary specialization (benthivore v . zooplanktivore feeding strategies within the benthic whitefish morph). Zooplanktivorous fish from both morphs acquired parasites mainly from pelagic copepods and in almost equal quantities. The benthivore feeders within the benthic morph had the highest proportion of parasites with transmission stages from benthic organisms. Host feeding behaviour seemed to be a major determinant of the helminth community structure, and helminths appeared to be useful indicators of long-term trophic specialization of whitefish ecotypes.     

Visceral helminth communities of an insular population of feral swine

Nine species of helminths, all nematodes, were recovered from the viscera of 48 feral swine (Sus scrofa) from Cumberland Island, Georgia. Both the overdispersed frequency distributions and the abundances of the four common species of helminths (Stephanurus dentatus, Metastrongylus apri, M. pudendotectus and Gongylonema pulchrum) did not vary significantly across the main and interactive effects of host sex and/or seasons. Whether or not the present low population densities of feral swine on Cumberland Island has influenced the pattern of fluctuations in abundances of helminth species across seasons as often observed in helminth communities from other hosts was not resolved. The apparent recent decline in prevalences and abundances, and the loss of certain species from the helminth communities of feral swine on the island may be explained partially by the decreasing transmission potentials of direct life cycle species caused by a recent marked reduction of numbers of individuals in the host population. Conversely, the apparent increased prevalence and abundance of three species of helminths (S. dentatus, M. apri and M. pudendotectus) may be related to their common utilization of earthworms as paratenic or intermediate hosts. Gongylonema pulchrum was the only helminth in which abundances seemed to remain unchanged. This was the only species that was not strictly host specific to feral swine. We found no evidence that helminth infections were responsible for morbidity or mortality in this feral swine population.     

Helminth parasites of the wild rabbit Oryctolagus cuniculus near Malham Tarn, Yorkshire, UK

Between 1992 and 1996, 95 rabbits from the immediate locality of Malham Tarn, North Yorkshire, UK were examined for the presence of helminth parasites. All the examinations took place in late September or October. Three species of nematodes, Graphidium strigosum, Passalurus ambiguus and Trichostrongylus retortaeformis and two species of cestodes, Taenia pisiformis and Cittotaenia pectinata were identified. There were no associations between helminth species richness and year of sampling, host weight or sex. A logistic model was fitted to the prevalence data from these helminths as was an over-dispersed Poisson model to the worm burden data. Graphidium strigosum was the most frequently identified species with an average prevalence of 78%. The mean prevalence and intensity of Graphidium infection were significantly effected by sampling year. The lower than normal rainfall recorded at the Tarn during the years 1995 and 1996 may have be one reason for this pattern. The worm burden of G. strigosum was significantly positively associated with rabbit body weight. The intensity of infection with P. ambiguus was significantly higher in female rabbits. There was a significant non-linear relationship between P. ambiguus worm burden and rabbit weight (P = 0.002) with worm burdens being highest in the 1000 g to 1499 g weight cohort. Trichostrongylus retortaeformis was only identified in 1994 and male rabbits harboured significantly higher worm burdens than females (48 vs. 7, P = 0.022). Over the five years, the average Taenia pisiformis prevalence was 31% and there was a significant positive association between worm burden and rabbit weight (P = 0.001). Cittotaenia pectinata had a prevalence of 37% over the whole study period with no interactions between prevalence or intensity and body weight, year of sampling or rabbit sex. All five helminths showed an overdispersed distribution with k values less than 1.     

Inequalities in size and intensity-dependent growth in a mermithid nematode parasitic in beach hoppers

Inequality in body sizes is a common feature in populations of helminth parasites, with potential consequences for egg production and population genetics. Inequalities in body lengths and the effects of intraspecific competition on worm length were studied in a species of mermithid nematode parasitic in the crustacean Talorchestia quoyana (Amphipoda: Talitridae). The majority of the 753 worms recovered were relatively small, and an analysis using a Lorenz curve and Gini coefficient suggested that there were no marked inequalities in body lengths among the worms. Total worm length in the 356 infected amphipods (i.e. the sum of the lengths of all the worms in a host) increased steadily as a function of the number of worms per amphipod, whereas the length of the longest worm per amphipod peaked in amphipods harbouring intermediate numbers of worms. This last result was not significantly accounted for by the observed increase in host size with increasing intensity of infection, but resulted from a correlation between worm length and host size. As the number of worms per amphipod increased, the relative sizes of the second-, third-, and fourth-longest worms per host increased markedly. This means that relative inequalities in sizes become less pronounced, i.e. subordinate worms get closer in size to the longest worm, as the number of worms per host increases. The main consequence of this phenomenon is that worm sizes in the mermithid population are more homogeneous than they would be if intraspecific competition had stronger effects on worm growth.     

Soil Microbial Substrate Properties and Microbial Community Responses under Irrigated Organic and Reduced-Tillage Crop and Forage Production Systems

Changes in soil microbiotic properties such as microbial biomass and community structure in response to alternative management systems are driven by microbial substrate quality and substrate utilization. We evaluated irrigated crop and forage production in two separate four-year experiments for differences in microbial substrate quality, microbial biomass and community structure, and microbial substrate utilization under conventional, organic, and reduced-tillage management systems. The six different management systems were imposed on fields previously under long-term, intensively tilled maize production. Soils under crop and forage production responded to conversion from monocropping to crop rotation, as well as to the three different management systems, but in different ways. Under crop production, four years of organic management resulted in the highest soil organic C (SOC) and microbial biomass concentrations, while under forage production, reduced-tillage management most effectively increased SOC and microbial biomass. There were significant increases in relative abundance of bacteria, fungi, and protozoa, with two- to 36-fold increases in biomarker phospholipid fatty acids (PLFAs). Under crop production, dissolved organic C (DOC) content was higher under organic management than under reduced-tillage and conventional management. Perennial legume crops and organic soil amendments in the organic crop rotation system apparently favored greater soil microbial substrate availability, as well as more microbial biomass compared with other management systems that had fewer legume crops in rotation and synthetic fertilizer applications. Among the forage production management systems with equivalent crop rotations, reduced-tillage management had higher microbial substrate availability and greater microbial biomass than other management systems. Combined crop rotation, tillage management, soil amendments, and legume crops in rotations considerably influenced soil microbiotic properties. More research will expand our understanding of combined effects of these alternatives on feedbacks between soil microbiotic properties and SOC accrual.     

A study of the internal parasites of small rodents from woodland areas in Surrey

The occurrence of helminth parasites and leptospires from the Long-tailed field mouse, Apodemus sylvaticus , the Bank vole, Clethrionomys giareolus and the Short-tailed vole, Microtus agrestis has been studied from four woodland areas in Surrey. The composition of the helminth fauna of these rodents, particularly A. sylvaticus and C. giareolus , differs from that of previous studies carried out in the British Isles in consisting largely of cestodes and fewer species of digeneans and nematodes. On the other hand the degree of infection of the rodents with nematodes as in the case of A. sylvaticus infected with the trichostrongylid Nematospiroides dubius and theoxyurid Syphacia stroma and of C. giareolus with thecap-illarid Capillaria muris sylvatici is greater than the cestode and digenean infections and is probably linked with the monoxenous nature of the nematode life cycles.
Differences in the number, incidence and degree of infection of mice and voles with species of helminths is undoubtedly linked with differences in the feeding habits of the respective hosts but the present study has also confirmed that infection levels in both leptospires and helminths are influenced by the age/sex relationships of the host together with season and habitat. The relationship between the distribution of rodents infected with leptospires and some helminths suggests similarities in the mode of their transmission.     

Does organic farming benefit farmland birds in winter?

The generally higher biodiversity on organic farms may be influenced by management features such as no synthetic pesticide and fertilizer inputs and/or by differences in uncropped habitat at the site and landscape scale. We analysed bird and habitat data collected on 48 paired organic and conventional farms over two winters to determine the extent to which broad-scale habitat differences between systems could explain overall differences in farmland bird abundance. Density was significantly higher on organic farms for six out of 16 species, and none on conventional. Total abundance of all species combined was higher on organic farms in both years. Analyses using an information-theoretic approach suggested that both habitat extent and farm type were important predictors only for starling and greenfinch. Organic farming as currently practised may not provide significant benefits to those bird species that are limited by winter food resources, in particular, several declining granivores.     

Population dynamics across selected habitat variables of the helminth community in coyotes, Canis latrans, from south Texas

The effects of selected intrinsic variables operating on host subpopulations and of extrinsic variables across the collective host population on the distributions of 10 common helminth species from coyotes in south Texas were compared. The intrinsic variables of host sex and presence and severity of sarcoptic mange had little effect on the distributions of most helminth species. The combined influences of (1) seasonal changes across the collective host population and (2) host subpopulations delineated by age were responsible for the overdispersed distributions of Oncicola canis, Physaloptera rara, and Protospirura numidica. Overdispersion in Ancylostoma caninum, Alaria marcianae, and Spirocerca lupi populations resulted almost exclusively from the heterogeneity of factors contributing to their rates of establishment, survival, and reproduction as generated across host age subpopulations. Aggregated abundances of A. caninum and A. marcianae occurred in young hosts, but were cumulative in older animals infected with S. lupi. The hypothesis that heterogeneity within the host population, rather than across the collective host population, is the main factor generating overdispersion in natural populations was confirmed for only 3 of 10 helminth species (A. caninum, A. marcianae, and S. lupi). The effects of extrinsic factors operating across the collective host population appeared to be equal to, or in some instances of greater importance than, these forces operating over host subpopulations in determining the dispersion patterns of some helminth species (O. canis, P. rara, and P. numidica). The distributions of Taenia pisiformis, Mesocestoides lineatus, Oslerus osleri, and Toxascaris leonina were not affected, or only minimally influenced, by these intrinsic or extrinsic variables. The effects of these habitat variables on dispersion patterns are highly correlated with the life cycle and mode of transmission of the respective helminth species.     

Structure in parasite component communities in wild rodents: predictability, stability, associations and interactions .... or pure randomness?

Experimental data establish that interactions exist between species of intestinal helminths during concurrent infections in rodents, the strongest effects being mediated through the host's immune responses. Detecting immune-mediated relationships in wild rodent populations has been fraught with problems and published data do not support a major role for interactions in structuring helminth communities. Helminths in wild rodents show predictable patterns of seasonal, host age-dependent and spatial variation in species richness and in abundance of core species. When these are controlled for, patterns of co-infection compatible with synergistic interactions can be demonstrated. At least one of these, the positive relationship between Heligmosomoides polygyrus and species richness of other helminths has been demonstrated in three totally independent data-sets. Collectively, they explain only a small percentage of the variance/deviance in abundance data and at this level are unlikely to play a major role in structuring helminth communities, although they may be important in the more heavily infected wood mice. Current worm burdens underestimate the possibility that earlier interactions through the immune system have taken place, and therefore interactions may have a greater role to play than is immediately evident from current worm burdens. Longitudinal studies are proposed to resolve this issue.     

Relationship between structure and stability of a Halipegus occidualis component population in green frogs: a test of selective treatment

The aggregated nature of helminth parasite populations has led to the suggestion that selectively treating heavily infected hosts can efficiently reduce parasite abundance and morbidity within a host population. Moreover, it has been postulated that a selective treatment protocol might have long-term effects on parasite transmission by disrupting the stability attributed to aggregated parasite population distributions by theoretical models. Long-term investigation has demonstrated year-to-year consistency in the population dynamics of Halipegus occidualis in green frogs from Charlie's Pond, North Carolina. In 1996, removal of all but 1 worm from each frog with > or =15 worms reduced the estimated component worm population by 45%, thereby decreasing mean intensity and aggregation (variance-to-mean ratio) of H. occidualis in the frogs by 85% and 63%, respectively. The following year, mean intensity, aggregation, and host colonization trends returned to pretreatment levels, indicating no effect of worm removal and demonstrating the stability of this host-parasite system. Although this result might be attributable to inefficient treatment or the presence of infection reservoirs, it is suggested that parasite population stability in this system might be governed by prevalence rather than intensity of adult worms. Therefore, repeated selective treatment might effectively modify intensity-dependent morbidity in similar host-parasite systems but should not affect further parasite transmission.     

Current knowledge on B chromosomes in natural populations of helminth parasites: a review

Helminths, traditionally classified into three phyla Platyhelminthes, Nemathelminthes and Acanthocephala, are a phylogenetically broadly diversified group of invertebrates, characterised by a parasitic life style. Current estimates of the helminth species diversity are at least 23-40,000 platyhelminthes, 10-26,000 nematodes and 1,200 acanthocephalans. Recent information on helminth karyotypes is fragmentary, and basic karyological data are known from approximately 1.1% of known species. Supernumerary chromosomes have been reported in selected populations of only 11 digenean flukes (Platyhelminthes), 1 thorny-headed worm (Acanthocephala) and 4 roundworms (Nematoda), which represent 3.6, 7.7 and 1.3% of the total number of species cytogenetically analysed to date within respective helminth groups. B chromosome presence was not generally associated with heteromorphic sex chromosomes as they occurred both in hermaphroditic flukes and dioecious helminths, and in species having male or female heterogametic sex chromosomes (ZW of schistosomes, XO of acanthocephalans and XY of nematodes). Numbers of B chromosomes varied from 1 to 10. Most often, Bs represented one or two of the smallest elements of the complement but they could be much bigger in some digenean flukes. B chromosomes showed a diverse morphology, including telocentric to metacentric structure. There is no detailed banding or ultrastructural study of Bs in the majority of helminth carriers. Assumptions on the possible relation between the occurrence of Bs in endoparasitic helminths and extreme environments are discussed.     

Basophils amplify type 2 immune responses, but do not serve a protective role, during chronic infection of mice with the filarial nematode Litomosoides sigmodontis

Chronic helminth infections induce a type 2 immune response characterized by eosinophilia, high levels of IgE, and increased T cell production of type 2 cytokines. Because basophils have been shown to be substantial contributors of IL-4 in helminth infections, and because basophils are capable of inducing Th2 differentiation of CD4(+) T cells and IgE isotype switching in B cells, we hypothesized that basophils function to amplify type 2 immune responses in chronic helminth infection. To test this, we evaluated basophil function using the Litomosoides sigmodontis filaria model of chronic helminth infection in BALB/c mice. Time-course studies showed that eosinophilia, parasite Ag-specific CD4(+) T cell production of IL-4 and IL-5 and basophil activation and IL-4 production in response to parasite Ag all peak late (6-8 wk) in the course of L. sigmodontis infection, after parasite-specific IgE has become detectable. Mixed-gender and single-sex worm implantation experiments demonstrated that the relatively late peak of these responses was not dependent on the appearance of circulating microfilariae, but may be due to initial low levels of parasite Ag load and/or habitation of the developing worms in the pleural space. Depletion of basophils throughout the course of L. sigmodontis infection caused significant decreases in total and parasite-specific IgE, eosinophilia, and parasite Ag-driven CD4(+) T cell proliferation and IL-4 production, but did not alter total worm numbers. These results demonstrate that basophils amplify type 2 immune responses, but do not serve a protective role, in chronic infection of mice with the filarial nematode L. sigmodontis.     

Nestedness in colonization-dominated systems: helminth infracommunities of Rana vaillanti Brocchi (Anura: Ranidae) in Los Tuxtlas, Veracruz, Mexico

Colonization probabilities of parasite species often are determined by the habitat preference and vagility of host individuals. Although extinction-based interpretations have been investigated for nested subset patterns of parasite infracommunities, the low relative frequency of nestedness in colonization-dominated systems makes the determination and interpretation of nested infracommunities of broad ecological importance. In these systems, ontogenetic shifts in habitat preference or diet of the host have the potential to produce nested subset patterns of parasite infracommunities. Helminth infracommunity structure was investigated for 76 Rana vaillanti individuals collected from Laguna Escondida, Los Tuxtlas, Veracruz, Mexico, in 1998. Pooled helminth infracommunities were significantly nested, as were penetrating and ingested helminth infracommunities when considered separately. Richness, diversity, and evenness of the helminth infracommunities were not correlated with host size, and did not differ between host sexes, suggesting that the structure of infracommunities simply is a product of the interaction between host individuals and their landscape mediated by individual differences in vagility. It is hypothesized that individual differences in recruitment can produce nested subset infracommunity patterns when the habitats or habitat preferences of hosts are themselves nested.