1. Parvatrema homoeotecnum sp.nov. from the oystercatcher, Haematopus ostralegus occidentalis Neumann at Aberystwyth is described and compared with other species of the genus.

2. The life cycle of this species is unique. The larval stages occur in the gastropod, Littorina saxatilis (Olivi) subsp. tenebrosa (Montagu) and include germinal sacs which have a structure and development similar to an adult digenean. There are no free-living stages and only one intermediate host.

3. The significance of this unique life cycle is discussed.

4. The family Gymnophallidae Morozov, 1955, is reviewed. Emended definitions are given for the family, subfamilies and genera. Keys, diagnostic features and brief notes of the species are included.

I am very grateful to Dr Gwendolen Rees, who suggested the investigation which led to the discovery of this species, for her advice and indispensable assistance throughout the work and the preparation of this paper. I am also grateful to the late Professor T. A. Stephenson for his interest and for the provision of working facilities; to Mr W. A. Ballantine, Mr A. H. Clarke, Jr., Mr C. Curtis, Miss G. P. F. Evans, Dr V. Fretter, Professor L. A. Harvey, Mr D. H. Jones and Dr J. Lewis who sent me specimens of Littorina saxatilis; to Professor R. M. Cable and Emerit. Professor G. R. La Rue for helpful suggestions; to Mr J. R. Hirst and Mr D. Hemingway Jones for photographic and technical assistance and to the Department of Scientific and Industrial Research for a grant which made the work possible.

Gastrointestinal pathogens are of three varieties, those that can, and often do, take the life of the host, those that infect transiently and rarely are life-threatening, and those (parasites) that establish a relatively prolonged residence or colonization of the host's alimentary tract. In the case of the second form, if infections are recurrent, both catabolic effects during the episode and failure to digest foods and/or absorb nutrients result. Similarly, catabolic wastage through activation of the acute phase response, and interference with the host's acquisition of nutrients by maldigestion, malabsorption, intestinal losses and competition with the parasite burden can impair growth and nutrition with helminthic infections. Growth and nutrition with respect to all of the macronutrients and virtually all of the micronutrients have been documented to be adversely affected by gastrointestinal pathogens. For its burgeoning importance as a worldwide health problem, both with the HIV virus as a direct intestinal pathogen and with the opportunistic gut infections occurring in the immunocompromised host, AIDS represents the emerging context of the impairment of nutritional status by intestinal pathogens.

Many parasites that use intermediate hosts are transmitted to the next host through predation. If the next host's fitness is strongly reduced by the parasite, it is under selection either to recognize and avoid infected intermediate hosts or to exclude that prey species from its diet when alternative prey are available. We investigated the predator-prey interaction between laboratory bred three-spined sticklebacks (Gasterosteus aculeatus), the second intermediate host of the cestode Schistocephalus solidus, from 2 parasitized and 1 unparasitized population, and different prey types: infected and uninfected copepods and size-matched Daphnia as alternative prey. Copepods with infective procercoids were more active, had a lower swimming ability and were easier to catch than uninfected controls. The sticklebacks preferred moving copepods. Therefore parasitized copepods were preferentially attacked and consumed. There was no effect of the sticklebacks' parent population being parasitized or not. The sticklebacks switched from Daphnia to (uninfected) copepods in the course of a hunting sequence; this switch occurred earlier in smaller fish. With this strategy the fish maximized their feeding rate: Daphnia were easier to catch than copepods but increasingly difficult to swallow when the stomach was filling up especially for smaller fish. However, there was no indication that sticklebacks from infected populations either consumed Daphnia rather than copepods or switched later in the hunting sequence to consuming copepods than fish from an uninfected population. Thus, sticklebacks did not avoid parasitized prey although S. solidus usually has a high prevalence and causes a strong fitness reduction in its stickleback host.

Transgenesis for Strongyloides and Parastrongyloides was accomplished in 2006 and is based on techniques derived for Caenorhabditis elegans over two decades earlier. Adaptation of these techniques has been possible because Strongyloides and related parasite genera carry out at least one generation of free-living development, with adult males and females residing in soil contaminated by feces from an infected host. Transgenesis in this group of parasites is accomplished by microinjecting DNA constructs into the syncytia of the distal gonads of free-living females. In Strongyloides stercoralis, plasmid-encoded transgenes are expressed in promoter-regulated fashion in the F1 generation following gene transfer but are silenced subsequently. Stable inheritance and expression of transgenes in S. stercoralis requires their integration into the genome, and stable lines have been derived from integrants created using the piggyBac transposon system. More direct investigations of gene function involving expression of mutant transgene constructs designed to alter intracellular trafficking and developmental regulation have shed light on the function of the insulin-regulated transcription factor Ss-DAF-16. Transgenesis in Strongyloides and Parastrongyloides opens the possibility of powerful new methods for genome editing and transcriptional manipulation in this group of parasites. Proof of principle for one of these, CRISPR/Cas9, is presented in this review.

A 37-spined Egyptian echinostome, Echinostoma liei sp.nov., is described in adult and larval stages. The parasite develops readily in the laboratory in chicks and ducklings, hamsters and rats. Its natural final host in or near irrigation ditches of the Nile delta involves the roof rat, Egyptian giant shrew and aquatic bird hosts. Developmental forms are described from infection of the NIH strain of Biomphalaria glabrata in the laboratory. B. alexandrina, is infected in the normal habitat in Egypt and contains both developmental stages in the heart or aorta and the hepatopancreas, and metacercariae encyst in the pericardium and kidney. E. liei sp.nov. is one of six very similar species characterized by 37 collar spines with a pattern of (3 + 2) corner spines in each lappet, six laterals on each side, and 15 dorsals in alternating rows; two pairs of dorsoventral and one small pair of ventro-lateral finfolds on the cercarial tail; and rodlike cystogenous material filling the cercarial encystation glands. In addition to distinctive intermediate-host specificity, differentiating characteristics of E. liei cercariae include presence of six sets of three flame cells each per side (total 36), seven oesophageal cells, eight penetration gland outlets on the dorsal lip of the oral sucker, and an absence of paraoesophageal gland cells as determined by intravital dyes. Significance of these and other cercarial traits is emphasized to aid in defining highly similar, but none the less distinct, sibling species.

The most efficient procedure for cryopreserving viable Babesia bovis organisms for in vitro cultivation consists of freezing extracellular parasites in a solution of 10% (w/v) polyvinylpyrrolidone (PVP) using a cooling rate of 20 °C/min. Although cultures can be established from thawed infected erythrocytes, the plating efficiency is relatively low. Freezing extracellular parasites resulted in plating efficiency up to 25%, when thawed and placed in culture. Glycerin or dimethyl sulphoxide (Me2SO) can be used successfully in the cryopreservation of B. bovis, but apparent toxic effects greatly decrease their efficiency. B. bovis parasites have been kept at − 196 °C for 60 days with no appreciable reduction in plating efficiency.

A collection of stocks of Trypanosoma brucei rhodesiense isolated in Kenya have been examined for electrophoretic variation in 20 enzymes. The results obtained have been analysed in order to determine whether these trypanosomes are diploid and undergo mating and to determine the genetic distance between T. b. rhodesiense, T. b. brucei and T. b. gambiense. The enzyme electrophoretic markers were further used in experiments involving cyclically transmitted mixtures of stocks aimed at detecting genetic exchange in the laboratory. No genetic exchange was detected. Two novel features of the enzyme electro phoretic results were found. Firstly, the stocks of T. b. rhodesiense were considerably more homogeneous than equivalent collections of stocks of T. b. brucei and secondly, all the stocks examined were heterozygous for two alleles of alkaline phosphatase and showed an excess of heterozygotes at the phosphoglucomutase locus. The degree to which these features are typical of T. b. rhodesiense has been examined in relation to previously published data. The results obtained strongly support the view that T. b. rhodesiense is a set of variants of T. b. brucei rather than a subspecies and a working hypothesis as to the relationship between T. b. brucei and T. b. rhodesiense is proposed to explain the enzyme electrophoretic data obtained.

The following report deals with a considerable collection of Nematodes made by Prof. A. S. Pearse, of Duke University, Durham, N.C., U.S.A., in various parts of Nigeria, and very kindly sent by him to the writer for study and determination.

Nuclear-encoded large-subunit ribosomal DNA sequences were used to infer a phylogenetic hypothesis for 17 taxa (16 nominal species) of the genera Contracaecum and Phocascaris. Phylogenetic trees based on these data have been used to assess the validity of the taxonomic distinction between these genera, which was based on the presence or absence of certain structural features, rather than on explicit hypotheses of evolutionary history. Phylogenetic hypotheses based on parsimony, likelihood, and neighbor-joining analyses of these sequence data strongly support the hypothesis that species of Phocascaris are nested within the clade of Contracaecum species hosted by phocid seals, and are more closely related to species of the Contracaecum osculatum complex than to other Contracaecum species. Alternative tree topologies representing Phocascaris as not nested within the C. osculatum complex were significantly worse interpretations of these sequence data. Phylogenetic analysis also provides strong support for the monophyly of all taxa (Contracaecum and Phocascaris) from phocid seals, which is consistent with Berland's (1964) proposal that such species form a natural group; however, his proposal to recognize all species in phocid seals as Phocascaris, with all species from birds as Contracaecum would result in a paraphyletic Contracaecum, according to the molecular phylogenetic hypothesis.

The use of naturally infected hosts in studies attempting to identify parasite-induced changes in host biology is problematical because it does not eliminate the possibility that infection may be a consequence, rather than a cause, of host trait variation. In addition, uncontrolled concomitant infections may confound results. In this study we experimentally infected individual laboratory-bred female three-spined sticklebacks Gasterosteus aculeatus L. with the pseudophyllidean cestode Schistocephalus solidus [Müller], and compared the morphology and growth patterns of infected females with sham-exposed controls over a 16-week period. Fish were fed a ration of 8% body weight per day. Non-invasive image analysis techniques allowed the growth of individual plerocercoids to be tracked in vivo throughout the course of infection, and patterns of host and parasite growth were determined. Females that developed infections diverged morphometrically from unexposed control females and exposed-uninfected females at 6 weeks post-infection, with the width of the body at the pectoral fins giving the earliest indication of infection success. When including the plerocercoid, infected females gained weight more quickly than controls, but when plerocercoid weight was removed this trend was reversed. There was no effect of infection on the increase in fish length. Plerocercoids grew at different rates in individual hosts, and exhibited measurable sustained weight increases of up to 10% per day. Final estimates of plerocercoid weight from morphometric analysis prior to autopsy were accurate to within +/−17% of actual plerocercoid weight. At autopsy, infected female sticklebacks had significantly lower perivisceral fat reserves but had developed significantly larger ovaries than controls. The results are discussed in relation to previous studies examining natural infections, and the value of utilizing experimental infections to examine ecological aspects of host–parasite interactions is discussed.

Exponential decay in community similarity as a function of distance is a ubiquitous phenomenon in biogeography. Thus, for parasite communities, pairwise similarity decreases with increasing geographical distance between host populations. This biogeographical rule should also apply along other dimensions characterizing the separation between communities. Since host-switching and phylogenetic affinities among host species affect the evolution of parasite faunas across host phylogenetic space the same way as dispersal and environmental gradients affect the assembly of local communities in geographical space, an exponential decay in similarity of parasite faunas with increasing host phylogenetic distance should be observed. This prediction is tested using data on metazoan parasites of 45 species of Canadian freshwater fishes belonging to 5 families. Across all host species, pairwise similarity in the composition of parasite faunas decayed exponentially, though not strongly, with increasing phylogenetic distance between hosts (measured as the number of substitutions per site along DNA sequences). A meta-analysis of correlations computed for separate fish families indicates only a very weak overall relationship. Data distribution indicates that phylogenetically close host species tend to share many of their parasites, while phylogenetically distant hosts have roughly equal chances of harbouring very similar or very dissimilar parasite faunas. The same pattern was seen when monogenean and trematode parasites were analysed separately, whereas no significant decay was observed for cestodes or nematodes, suggesting different patterns of host-switching and parasite colonization among these taxa. The results show that similarity in species composition decreases, though weakly, with increasing distance in the same manner in phylogenetic space as it does in geographical space.

Eight taxa have recently been proposed as being encompassed by the genus Trichinella on the basis of allozyme and biological data. In this paper we show that an analogous 8 taxon structure for this genus results from the random amplified polymorphic DNAs (RAPDs). Five 10-mer or 20-mer primers were used under different polymerase chain reaction (PCR) conditions to produce multiband RAPD fingerprints from muscle larvae of 40 isolates of Trichinella spp. The resulting RAPD data were analysed following the numerical taxonomic approach, and the resulting classification was compared to that derived from allozyme data. The agreement found between allozymes and RAPDs, while supporting the polyspecific structure of the genus Trichinella, confirms the potential of RAPDs as a tool for the detection of cryptic species. The selected primers were tested on individual muscle larvae in an attempt to standardize a RAPD assay for the routine identification of the 8 taxa of Trichinella. Only 1 of the 5 primers yielded reproducible fingerprints from the single larvae. Using this primer, the 5 species and the 3 other taxa of the genus Trichinella can be identified in a single assay without the need for massive in vivo parasite production.

The survival of Heligmosomoides polygyrus was monitored during primary infections in female C57Bl10, NIH and BALB/c mice at low and high intensities of infection. Survivorship curves were fitted for each data set and analysed. C57Bl10 mice, given either low or high intensities of infection, harboured parasites for 28–37 weeks, heavier infections surviving marginally but significantly longer. Essentially the survivorship curves of H. polygyrus in C57Bl10 mice could be accounted for by senility, the increased probability of worms with a longer life-span occurring at high infection intensities and, possibly, by a contribution from host-protective immune mechanisms in the terminal stages of infection. The pattern of survivorship was different in NIH and BALB/c mice. NIH mice showed weak but significant density-dependent suppression of parasite loss and infections in this strain did not exceed 27·5 weeks in duration. Primary infections in BALB/c mice were briefer still and showed marked dependence on parasite density. Thus low-level infections lasted 10–15 weeks whereas heavier infections survived for 21–34 weeks. The data suggested that both strains developed host-protective responses to adult H. polygyrus and that parasite survival was curtailed earlier than would be expected if senility alone was involved. The hybrid strains (C57Bl10 × NIH)F1 and (B10G × NIH)F1 both expelled H. polygyrus in a dose-dependent manner, worm loss commencing within 10 weeks of infection. In some experiments worm loss was clearly evident by weeks 4 and 6. These hybrid strains showed gene complementation in that adult worms were cleared considerably earlier than in parental strains.

The natural history of fig-pollinating wasps and their associated species-specific nematodes allows the measurement of many parameters which are relevant to testing hypotheses concerning host-parasite ecology and evolution. Within fig wasps species, it is possible to estimate lifetime reproductive success of foundress wasps as a function of presence or absence of nematode parasitism (virulence). Across species, there is a wide range of host population structures which, in turn, results in a range of opportunities for either horizontal or vertical nematode transmission. Therefore, estimates of virulence can be related to opportunities for transmission across a group of closely related hosts and their parasites. Further, the dynamics of the nematode infections over ecological and short-term evolutionary timescales can be monitored, giving added insight into the interpretation of the virulence estimates. Moreover, several scales of longer term evolutionary relationships are either known directly from fossil evidence or can be inferred from molecular data, providing deeper temporal context for the observed patterns. This combination of attributes permits detailed testing of hypotheses concerning the factors that potentially influence the evolution of virulence in host-parasite systems, and further, population and simulation models of the system that incorporate the parameter estimates can clarify the interpretation of how those factors act. There is little evidence suggesting that intermediate and long-term evolutionary relationships explain current levels of virulence. That is, it appears that virulence can change rapidly relative to speciation events, and that the nematodes do not tend to become ‘benign over time’. Instead, it appears that host population structure can influence the evolution of parasite virulence by affecting the relative opportunities for horizontal to vertical transmission, which, in turn, influences the relative costs and benefits of virulence to the nematodes. At one level, increased opportunities for horizontal transmission decouple the reproductive interests of the individual nematodes from those of the individual hosts that they are directly parasitizing, thereby reducing the cost of virulence to individual nematodes. At another level, increased opportunities of horizontal transmission also increases the relative frequency of hosts infected by multiple strains of nematodes. This promotes the evolution of more virulent forms by increasing the relative importance of within-host competition among nematode strains, thereby favouring strains that ‘eat more host sooner’. An interesting property of the fig-nematode systems is that the proportion of infected hosts does not change dramatically through time. This finding implies that there can be considerable negative effects on survival of infected hosts in addition to the previously documented reductions in fecundity of infected foundresses, because the latter are insufficient to account for the observed stability of wasp infection rates.

To facilitate the completion of their life-cycle, many helminth parasites have evolved the ability to manipulate the behaviour of their intermediate host in order to make it more likely to be eaten by the parasite's definitive host. Here, we determined whether the cestode Eubothrium salvelini modifies the behaviour of its intermediate host, the copepod Cyclops vernalis, and makes it more susceptible to predation by brook trout, Salvelinus fontinalis, the parasite's final host. Following the experimental infection of copepods, the spontaneous activity of infected and control subjects was quantified weekly. In addition, we regularly quantified predation by individual brook trout fry on known numbers of infected and control copepods. At approximately the time when the cestode larvae became infective to fish (2–3 weeks following infection), the infected copepods started to swim more actively than uninfected controls. Also at that time, infected individuals became more likely to be captured by fish than uninfected ones. Copepod size and intensity of infection had no significant effect on their behaviour or their risk of being eaten by fish. Thus cestode- induced changes in copepod swimming activity can lead to infected copepods becoming highly vulnerable to fish predators, and may have resulted from selection on the parasite to increase its transmission success

1. Histological data are given about the location and frequency of the different stages of the life history of D. folliculorum in the pilo-sebaceous apparatus of man. These data are examined statistically and the relative durations of each stage and their movements in the follicle are deduced from them.

2. A method of in vitro culture is described and data are given about the longevity of the various stages of the life history.

3. Experiments on the behaviour of the different motile stages of the mite are described. It is concluded that the deutonymph is the distributive stage, and that distribution occurs over the skin surface.

4. The life history is reconstructed by a synthesis of the evidence presented. The life-cycle lasts approximately 14½ days, the life span of each stage of it being: ovum 60 hr., larva 36 hr., protonymph 72 hr., deutonymph 60 hr., adult 120 hr. Interval between copulation and oviposition 12 hr.

This work was largely carried out in the Department of Pathology of the Institute of Dermatology, London, and I wish to thank the Director of Pathology, Dr J. O. Oliver, for making available to me the facilities of his department, and for his advice and interest during the progress of the work. The photographs were prepared by Mr R. H. Lunnan of the Photographic Department of the Institute of Dermatology. I am indebted to the Dean of the Institute of Dermatology for permission to publish the photographs.

I am grateful to Mr T. E. Hughes of the Department of Zoology, Birkbeck College, University of London, for his advice at all times, and to Miss Hilda Davies of the Department of Statistics, University of Sheffield, for help with statistical methods and to Mr W. Moseley, who prepared the text figures.

Finally, I wish to express my thanks to Professor I. Chester Jones and Dr E. T. B. Francis of the Department of Zoology, University of Sheffield, for reading the manuscript and for their help in its preparation.

Changes in the variance to mean ratio and the parameter k of the negative binomial distribution were used to study temporal changes in the degree of aggregation of the monogenean parasite, Gyrodactylus turnbulli in free-running laboratory populations of the guppy, Poecilia reticulata. The parasite undergoes recurrent epidemic cycles in the host population under conditions of regular immigration of uninfected guppies. During the early phase of the epidemic, heterogeneity among fish together with direct reproduction are thought to contribute to the increasing degree of aggregation. During the increasing phase of the epidemic cycle, parasites become increasingly aggregated in the host population, presumably because of the direct reproduction of the parasite on the surface of a single host. As the peak prevalance and abundance are approached, the parasites become less aggregated with lowest clumping occurring during the declining phase of the cycle. This is thought to be a function of density-dependent death of infected hosts, and density-dependent reduction in parasite survival and reproduction on hosts that recover from infection. This study clearly indicates that the variance to mean ratio and the parameter k of the negative binomial distribution do not quantify the same aspect of the frequency distribution. It is suggested that the variance to mean ratio is a better measure when the prevalence and−or mean burden are changing and when the tail of the distribution is of particular interest, and that k may be a preferred parameter when the zero class or the lightly infected hosts are of primary interest.

Cyclosporin A (CsA), a cyclic undecapeptide with powerful properties of immunosuppression, acts on parasitic infections in laboratory animals in various ways. The outcome of drug administration in vivo varies with timing of treatment relative to infection, route of administration, dose and number of treatments applied. CsA is clearly antiparasitic against malaria, schistosomes, adult tapeworms, metacestodes and filarial nematodes. By contrast, it acts as an immunomodulator against trypanosomes and Giardia, by exacerbating infection; in the case ofLeishmania spp. the drug acts variously. In some other infections CsA acts both as an antiparasite drug and as an immunosuppressant (Toxoplasma, avian coccidiosis and gastrointestinal nematodes).

Immunity to N. brasiliensis in rats is stimulated primarily by the adult worms. An initial infection consisting solely of adult worms terminated at the same time as an infection consisting of all the parasitic stages.

Immunity is not stage specific. Immunity stimulated by adult worms acts not only on adult stages inhibiting their reproduction but also inhibits the development to maturity of larvae in a challenge infection.

Rats infected solely with female adult worms, whether egg producing or sterile, are more resistant to reinfection than rats infected with male worms only.

Immunity stimulated by a single normal infection lasts a long time but it does not depend on the persistence of adult worms in the intestine of the rats.

This work was done during the tenure of a Commonwealth Scholarship awarded by the Association of Commonwealth Universities.

It is shown that under the conditions that prevailed in New Zealand in the late 1950s, Taenia hydatigena was hyperendemic, the life-cycle being regulated by a density-dependent constraint in the form of acquired immunity, and T. ovis was rare. The control measures that caused Echinococcus granulosus, which was endemic at the time, to decline towards extinction reduced T. hydatigena and T. ovis to endemic status only. A non-linear integrodifferential equation model, which was previously linearized to describe the life-cycle of E. granulosus in dogs and sheep in New Zealand, is used to describe the life-cycles of T. hydatigena and T. ovis. The model is then used to compare and contrast the population dynamics of these three species. The model is used to demonstrate that the endemic steady state is structurally unstable, and may be asymptotically unstable to small perturbations. It is also shown that despite the lower infection pressure experienced by the intermediate host in the endemic state, the numbers of larvae in sheep may be higher than in the hyperendemic state. Finally it is shown that the partial success of the control measures against T. hydatigena may have caused an increase in the numbers and prevalence of T. ovis larvae in sheep due to the reciprocal immunity between the two species.