Introductions, spread and colonization of new localities by fish helminth and crustacean parasites in the British Isles: a perspective and appraisal

Most parasites are disseminated by movements of infected hosts. The increasing extent and rapidity of anthropochore fish movements are causing increased concern related to awareness of their potential and known capacity for disseminating parasites. This paper puts these data in perspective by examining examples of actual and potential translocations of fish helminthes and crustaceans by anthropochore movements offish into and throughout the British Isles, and by distinguishing the processes of dissemination and invasiveness from those of colonization and establishment. An investigation of the British fish and helminth parasite fauna suggests that: (1) the range of many species is not well known, many are local in distribution and appearances beyond the range may reflect detection and patchiness, not translocation; (2) taxonomic problems in many groups hinder detection and determination of range; (3) most parasites possess the attributes of good colonizers so the natural expansion and contraction of ranges are often not noticed or recognized as such and the importance of parasite introductions by natural movements of fish or avian hosts is generally underestimated; (4) invasions are far commoner than colonizations, since conditions for establishment may be very restricted and transmission windows very narrow in time and space; (5) successful colonizations and translocations tend to be documented and attract attention whereas invasions resulting in failed colonizations are seldom observed and more seldom documented, thus biasing our perception. Given the extensive history of fish introductions to, and translocations within, the British Isles it is surprising how few fish helminths and crustaceans have invaded the country successfully (16 species: 11·4%) and how many still show restricted distributions. The majority (68·7%) of introduced helminths are associated with fish having ornamental varieties. Barriers of colonization are more effective than those to invasion and it is clear that most translocations and invasions fail. It is right to be concerned about the dangers, but it is also important to put anthropochore factors in perspective.     

Patterns in the composition and richness of helminth communities in brown trout, Salmo trutta, in a group of reservoirs

Helminth community composition and richness were studied in brown trout, Salmo trutta , in 10 reservoirs of broadly similar age and characteristics situated close to each other in a well-defined region of south-west England. Communities were compared using cluster and ordination analyses, and possible correlations between helminth richness and a number of environmental variables were investigated. The hypothesis that the helminth communities should show high degrees of similarity and that large differences between reservoirs and a high degree of clustering would be unrecognizable was refuted. Levels of community similarity were low and comparable to those determined for helminths in salmonids in natural lakes. Trout in some reservoirs exhibited very distinctive helminth faunas and clustering of reservoirs was apparent. No single factor, including reservoir size and presence of piscivorous birds, had a predominant influence on community richness or composition but rather a multiplicity of local factors was believed to influence these parameters. The results indicate that local factors promoting distinctiveness have a greater influence on the composition and richness of fish helminth communities in lakes and reservoirs than do regional factors promoting similarity.     

长江中下游三个湖泊中鳜消化道寄生蠕虫群落的组成与多样性

鱼类在食物网中的营养位置对其消化道寄生蠕虫的群落结构有重要作用。本研究调查了梁子湖、洞庭湖、鄱阳湖中鳜(Siniperca chuatsi)消化道寄生蠕虫群落的组成和多样性。在3个湖泊中共发现11种寄生蠕虫, 优势种均为范尼道佛吸虫(Dollfustrema vaneyi), 频率分布中感染有12种寄生虫的样本占65%, 单个样本中最大物种数为6。3个湖泊中平均物种丰富度为1.532.13, Brillouin多样性指数为0.120.33, 其中鄱阳湖中的多样性最高。梁子湖和洞庭湖之间的Jaccard相似度和百分比相似指数最高。通过比较食物网中不同营养位置鱼类的消化道寄生蠕虫群落结构, 发现鳜的消化道寄生虫群落的物种丰富度与多样性水平都高于植食性和杂食性鱼类。研究还讨论了宿主食物组成对消化道寄生蠕虫群落结构的影响。       

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.     

Richness and diversity of helminth communities in the Japanese grenadier anchovy, Coilia nasus, during its anadromous migration in the Yangtze River, China

To determine the relationship between the species richness, diversity of helminth communities, and migration distance during upward migration from coast to freshwater, helminth communities in the anadromous fish Coilia nasus were investigated along the coast of the East China Sea, the Yangtze Estuary, and 3 localities on the Yangtze River. Six helminth species were found in 224 C. nasus . Changes in salinity usually reduced the survival time of parasites, and thus the number of helminth species and their abundance. Except for the 2 dominant helminths, the acanthocephalan Acanthosentis cheni and the nematode Contracaecum sp., mean abundance of other 4 species of helminths was rather low (<1.0) during the upward migration in the Yangtze River. Mean abundance of the 2 dominant helminths peaked in the Yangtze Estuary and showed no obvious decrease among the 3 localities on the Yangtze River. Mean species richness, Brillouin's index, and Shannon index were also highest in the estuary (1.93 ± 0.88, 0.28 ± 0.25, and 0.37 ± 0.34, respectively) and did not exhibit marked decline at the 3 localities on the Yangtze River. A significant negative correlation was not seen between the similarity and the geographical distance (R = -0.5104, P = 0.1317). The strong salinity tolerance of intestinal helminths, relatively brief stay in the Yangtze River, and large amount of feeding on small fish and shrimp when commencing spawning migration perhaps were responsible for the results.     

Biogeography of helminth parasites of freshwater fishes in Mexico: the search for patterns and processes

Aim To uncover and describe patterns of biogeography of helminth parasites in freshwater fishes of Mexico, and to understand processes that determine them. Three predictions about host‐specificity, faunal exchange in transitional areas, and the biogeographical ‘core’ fauna, are evaluated, all of which follow from a fundamental hypothesis: that parasites show characteristic associations with particular host clades. The parasite fauna of the southern Mexican cichlids and of the fishes of the Mesa Central are examined as case studies that reflect Neotropical and Nearctic historical influences. Location The region covered in this study includes most of Mexico, with emphasis on six biogeographical areas: the Yucatán Peninsula (area 1), the Grijalva‐Usumacinta drainage (area 2), the Papaloapan and Pánuco drainages (area 3), the Balsas drainage (area 4), the Lerma‐Santiago drainage (area 5), and the Bravo drainage (area 6). Methods A parasite data base containing all the records of helminth parasites of freshwater fishes of Mexico was filtered to extract records of adult helminth parasites in freshwater fishes from the six biogeographical areas designated in this study. Jaccard's similarity coefficients and cluster analyses (using upgma ) were used to analyse the extent of faunal similarity between the designated biogeographical areas and between host (fish) families. Taxonomic composition of parasite assemblages in different host groups was also qualitatively compared from summary data. These data were used to test the three main predictions. Results To date, 184 species of helminths (120 as adults) have been recorded from 127 freshwater fishes in Mexico (almost 33% of the total fish diversity of Mexico). Of these parasite species, 69 are digenetic flukes, 51 are nematodes, 33 are monogeneans, 25 are tapeworms, and only six are acanthocephalans. The data and analyses from the six biogeographical areas corroborate the predictions that: (1) the adult parasite fauna is largely circumscribed by higher levels of monophyletic host taxa (families, orders, etc.), and that this pattern is independent of areas; (2) areas within a certain biogeographical region, and consequently with similar fish composition (e.g. areas 1, 2 and 3) have more similar parasite faunas compared to areas with less similar fish faunal composition; and (3) ‘core’ parasite faunas persist to some extent in transitional areas with limited host‐sharing. Main conclusions Helminth biodiversity in Mexican freshwater fishes is determined by the historical and contemporary biogeography of their hosts. Host lineage specificity, mainly at the level of the host family, appears to be an important factor in the distribution of the parasites. Most fish families (Characidae, Cichlidae, Pimelodidae, Ictaluridae, Catsotomidae, Goodeidae, Atherinidae) possess their own characteristic ‘core’ helminth fauna, with limited host‐sharing in transitional areas (e.g. areas 3 and 4). A re‐evaluation of the helminth fauna of Mexican cichlids questions the hypothesis that cichlids lost parasites during the colonization of Mexico from South America. The evidence supports the idea that they acquired new parasites by host switching, possibly from marine or brackish‐water percomorphs. In contrast, the parasite fauna of the Mesa Central remains enigmatic and reflects the region's history of endemicity with historical marine and Nearctic connections.     

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.     

Body size,trophic level,and the use of fish as transmission routes by parasites

Within food webs, trophically transmitted helminth parasites use predator–prey links for their own transfer from intermediate prey hosts, in which they occur as larval or juvenile stages, to predatory definitive hosts, in which they reach maturity. In large taxa that can be used as intermediate and/or definitive hosts, such as fish, a host species’ position within a trophic network should determine whether its parasite fauna consists mostly of adult or larval helminths, since vulnerability to predation determines an animal’s role in predator–prey links. Using a large database on the helminth parasites of 303 fish species, we tested whether the proportion of parasite species in a host that occur as larval or juvenile stages is best explained by their trophic level or by their body size. Independent of fish phylogeny or habitat, only fish body length emerged as a significant predictor of the proportion of parasites in a host that occur as larval stages from our multivariate analyses. On average, the proportion of larval helminth taxa in fish shorter than 20 cm was twice as high as that for fish over 100 cm in length. This is consistent with the prediction that small fishes, being more vulnerable to predation, make better hosts for larval parasites. However, trophic level and body length are strongly correlated among fish species, and they may have separate though confounded effects on the parasite fauna exploiting a given species. Helminths show varying levels of host specificity toward their intermediate host when the latter is the downstream host involved in trophic transmission toward an upstream definitive host. Given this broad physiological compatibility of many helminths with fish hosts, our results indicate that fish body length, as a proxy for vulnerability to predators, is a better predictor of their use by helminth larvae than their trophic level based on diet content.     

Helminth communities of the lesser sandeel Ammodytes tobianus L. off the west coast of Ireland

The helminth parasites present in 412 lesser sandeels (Ammodytes tobianus) taken from June 1996 to May 1997 from the Aran Islands on the west coast of Ireland were examined. Ten helminth parasite species were recorded, and more than 92% of the sandeels were infected with at least 1 helminth species. Seven of the species were digeneans, including Brachyphallus crenatus, Hemiurus communis, Derogenes varicus, Lecithaster gibbosus, Opechona bacillaris, Cryptocotyle lingua, and Galactosomum lacteum; 2 nematodes, including Hysterothylacium sp. and Contracaecum sp.; and 1 cestode, Scolex pleuronectis. Three of the 7 digenean species were either larvae or immature. Only 2 species, the digeneans G. lacteum and H. communis, had prevalences greater than 50%. The dominant species was G. lacteum, accounting for 67% of all parasites present. The relationship between spawned groups, host length, and season versus the abundance, prevalence, species richness, and the total number of parasites in the infracommunities was investigated. No difference was found between the parasite communities of the 2 spawning races of the host population. Mean abundance and prevalence of the different parasite species showed seasonal variation. Numbers of parasite species and numbers of parasites increased with fish length. The role of A. tobianus as an intermediate host for helminths was assessed; it was determined that most were infectious to birds or mammals, with the majority of the parasite species being autogenic (infectious to fish). The mean number of parasites per fish was nearly a quarter of the value recorded for A. tobianus in the North Sea, where a much higher intensity of infection was recorded.     

Fish immune responses to experimental and natural infection with helminth parasites

Selected features of the responses by fish to helminth parasites are discussed and comparison is made where appropriate with mammals. These include: (i) Factors influencing host specificity and consideration of the mechanisms that underpin the restriction of some parasites in their host spectrum, (ii) How fish leucocytes kill helminth larvae, with emphasis on the role of released oxygen (ROS) and nitrogen (RNS) free radicals from macrophages, (iii) Immune evasion strategies used by fish helminths, including invasion of immunologically privileged sites, encystment, adsorption of host proteins on the parasite surface, and high surface membrane turnover, (iv) Potential immunogens for vaccination and use for immunodiagnosis of infection, and (v) Natural and induced protection against helminths, with emphasis on the potential for future vaccination strategies.     

Invasion and initial replication of ultraviolet irradiated waterborne infective stages of Myxobolus cerebralis results in immunity to whirling disease in rainbow trout

Myxobolus cerebralis is a microscopic metazoan parasite (Phylum Myxozoa: Myxosporea) associated with salmonid whirling disease. There are currently no vaccines to minimise the serious negative economical and ecological impacts of whirling disease among populations of salmonid fish worldwide. UV irradiation has been shown to effectively inactivate the waterborne infective stages or triactinomyxons of M. cerbralis in experimental and hatchery settings but the mechanisms by which the parasite is compromised are unknown. Treatments of triactinomyxons with UV irradiation at doses from 10 to 80 mJ/cm(2) either prevented (20-80 mJ/cm(2)) or significantly inhibited (10 mJ/cm(2)) completion of the parasite life cycle in experimentally exposed juvenile rainbow trout (Oncorhynchus mykiss). However, even the highest doses of UV irradiation examined (80 mJ/cm(2)) did not prevent key steps in the initiation of parasite infection, including attachment and penetration of the epidermis of juvenile rainbow trout as demonstrated by scanning electron and light microscopy. Furthermore, replication of UV-treated parasites within the first 24h following invasion of the caudal fin was suggested by the detection of concentrations of parasite DNA by quantitative PCR comparable to that among fish exposed to an equal concentration of untreated triactinomyxons. Subsequent development of parasites treated with an 80 mJ/cm(2) dose of UV irradiation however, was impaired as demonstrated by the decline and then lack of detection of parasite DNA; a trend beginning at 10 days and continuing thereafter until the end of the study at 46 days post parasite exposure. Treatments of triactinomyxons with a lower dose of UV irradiation (20 mJ/cm(2)) resulted in a more prolonged survival with parasite DNA detected, although at very low concentrations, in fish up to 49 days post parasite exposure. The successful invasion but only short-term survival of parasites treated with UV in rainbow trout resulted in a protective response to challenges with fully infective triactinomyxons. Prior treatments of juvenile rainbow trout with UV-treated triactinomyxons (10 and 20 mJ/cm(2)) resulted in a reduced prevalence of infection and significantly lower concentrations of cranial myxospores (two direct measures of the severity of whirling disease) compared with trout receiving no prior treatments when assessed 5 months post parasite exposure to fully infective triactinomyxons.     

Helminths of stocked rainbow trout (Salmo gairdneri) with special reference to Clinostomum complanatum

Rainbow trout (Salmo gairdneri) stocked into a small prairie dug-out were examined for helminths at harvest time. Five species of helminths were found (Diplostomum sp., Diplostomulum scheuringi, Clinostomum complanatum, Crepidostomum farionis and Pomphorhynchus bulbocolli); C. farionis and P. bulbocolli mature in fish while the remaining species utilize fish-eating birds as definitive hosts. Examination of stomach contents indicated that all invertebrate hosts required for the life-cycles of these helminths were present in the dug-out. The most conspicuous and prevalent helminth was C. complanatum as fish were "wormy" and unmarketable due to the presence of high numbers of metacercariae. Metacercariae of C. complanatum were recovered from most organs of rainbow trout. The low survival of stocked fish suggests that C. complanatum may cause some host mortality, but the condition of infected and uninfected fish was similar. It appears that a community of ichthyoparasites can become established in a population of stocked fish in a single growing season in north temperate regions.     

The effect of local environmental variables on the helminth parasite communities of the pointedbelly frog Leptodactylus podicipinus from ponds in the Pantanal wetlands

Understanding the patterns of species distribution and abundance has been at the core of ecology. In general, these patterns are determined by species dispersion as well as by abiotic and biotic environmental conditions. Similarly, host-parasite relations and the structure of parasite assemblages are also shaped by environmental conditions and landscape composition. Herein, we assessed the influence of environmental variables and parasite species dispersion on the structure of helminth parasites communities in the frog Leptodactylus podicipinus. We sampled 10 ponds and recorded area, depth, altitude, pH, dissolved oxygen, salinity, temperature, and extent of soil, water, and vegetation cover as well as the distances between the ponds. We collected 121 frogs and found 9 helminth taxa; 2 of them were core species (prevalence higher than 50%), which contributed to the relatively high similarity observed among the ponds. Most of the helminths showed some variation in the frequencies of occurrence among communities from different ponds. The change in species composition among ponds was explained by the environmental variables but not by the distance between the ponds. Moreover, the results indicated that local processes (variation in environmental conditions) were more important than the regional processes (species distribution) in determining the structure of parasite communities. The variation in helminth communities among ponds in response to moderate differences in pond environmental characteristics points to the potential of helminth species as indicators of environmental conditions.     

Has the introduction of brown trout altered disease patterns in native New Zealand fish?

1. It is well recognised that non-indigenous species (NIS) can affect native communities via the 'spillover' of introduced parasites. However, two other potentially important processes, the 'spillback' of native parasites from a competent NIS host, where the latter acts as a reservoir leading to amplified infection in native hosts, and the 'dilution' of parasitism by a NIS host acting as a sink for native parasites, have either not been tested or largely overlooked.
2. We surveyed the helminth parasite fauna of native New Zealand fish in Otago streams that varied in the abundance of introduced brown trout Salmo trutta , to look for evidence of spillback and/or dilution. Spillover is not an issue in this system, with trout introduced as parasite-free eggs.
3. Seven native parasite species were present across 12 sites; significant inverse relationships with an index of trout abundance (i.e. dilution) were documented for three species infecting the native upland bully Gobiomorphus breviceps , and one species infecting the native roundhead galaxias Galaxias anomalus .
4. An inverse relationship between bully energy status and infection intensity of one parasite species suggests that parasite dilution could have positive effects on bully populations. Our failure to detect similar relationships for the other parasites does not preclude the possibility that dilution is beneficial to native fish, since parasites may have subtle or unmeasured impacts.
5. The parasite dilution patterns reported are compelling in that they occurred across several native host and parasite species; as such they have important implications for invasion ecology, providing an interesting contrast to the largely negative impacts reported for NIS. Mechanisms potentially responsible for the patterns observed are discussed.     

The status of brown and rainbow trout, Salmo trutta and S. gairdneri as hosts of the acanthocephalan, Pomphorhynchus laevis

The status of brown and rainbow trout as hosts of Pomphorhynchus laevis was studied in the field and by means of laboratory investigations. Field data indicated that rainbow trout might belong to the group of preferred hosts of P. laevis , whereas brown trout belonged to the group in which the parasite achieved less than optimal growth and maturation. This was confirmed by laboratory infections. In rainbow trout P. laevis attained up to three times the growth rate in brown trout and maturation occurred whereas in brown trout establishment was lower, growth slower and no parasites matured. Changes in the behaviour of infected Gammarus pulex induced by the presence of P. laevis cystacanths were such as to render the shrimps more vulnerable to predation by trout and other surface and mid-water feeding fish, and selective predation upon infeged G. pulex by fish was demonstrated. nvestigations into the stimuli necessary for eversion of cystacanths of P. laevis revealed that the most important factor was a non-specific component of bile, and it was concluded that cystacanths were likely to evert in any species of fish. Recognition of the different status of brown and rainbow trout as hosts of P. laevis still fails to explain some peculiarities in the distribution of the parasite in the British Isles, where in Britain it occurs in trout in only one river but in Ireland in all rivers throughout the country. It is suggested that the Irish parasites may constitute a different strain of P. laevis , since they use a different species of intermediate host and are better able to survive in brown trout.     

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.     

Community ecology of the parasites of adult spot, Leiostomus xanthurus, and Atlantic croaker, Micropogonias undulatus (Sciaenidae) in the Cape Hatteras region

Twenty-three species of metazoan parasites were recorded from adult spot and 26 from adult croaker. Of the 33 parasitic species found, 17 occurred in both spot and croaker. No significant differences in intensity of parasites occurred between sexes of either spot or croaker. All of the parasites had over-dispersed, or clumped, distributions among hosts. Adult spot and croaker collected offshore had much greater species-richness, diversity, and total number of individual parasites than juvenile fishes collected in adjoining estuaries during another study. The juvenile spot and croaker had less time to acquire parasites and inhabited less diverse and more confined habitats in inshore estuaries, which resulted in less diverse parasite communities than offshore fishes. The number of species and diversity of parasites in adult fish was greater in croaker than spot. However, when only gastrointestinal helminths were considered, spot had greater species-richness as well as greater numbers of individual helminths, suggesting that they had a more diverse diet and that they fed on more infected intermediate hosts than croaker. In both adult spot and croaker the mean number of parasitic species was greater than those of freshwater fishes and fewer than those for birds and mammals. The total number of individual parasites was similar to that of freshwater fishes. The opportunistic diet and the migratory habits of both spot and croaker contribute to their diverse parasite faunas. Comparison of adult spot and croaker parasite faunas collected offshore indicated that their respective parasite component communities were distinct and that similar parasite infracommunity variability existed in both hosts and that their communities were not ‘random’ samples, but restricted subsets of the compound community. Although the parasite dominance hierarchy in adults of both species varied slightly between areas and seasons sampled, there appeared to be a predictable dominant species that was accompanied by subordinate, less predictable species. However, the variability in both relative intensities and presence-absence of parasites within communities resulting from their diverse diets make them less predictable than those of other vertebrates with less diverse diets such as the lesser scaup duck and more like those of other marine fishes.     

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.     

Correlates of helminth community in the red-legged partridge (Alectoris rufa L.) in Spain

Between 1992 and 1996, 587 wild red-legged partridges (Alectoris rufa) from 16 Spanish provinces were examined to study the variations of helminth communities in this game species across a broad geographical area. The survey revealed 13 species of helminth parasites. Dicrocoelium sp.. Rhabdometra nigropunctata, and Cheilospirura gruweli were the most common species, whereas Raillietina bolivari, Choanotaenia infundibulum, Tetrameres sp., and Capillaria anatis were the most rare. Subulura suctoria, Heterakis gallinarum, Heterakis tenuicaudata, Capillaria contorta, Trichostrongylus tenuis, and Raillietina tetragona occurred with intermediate frequencies. The abundance of C. gruweli, S. suctoria, H. tenuicaudata, T. tenuis, and R. tetragona was inversely correlated to latitude and directly correlated to yearly mean temperature, whereas the abundance of Dicrocoelium sp. was directly correlated to latitude and inversely correlated to yearly mean temperature. The abundance of R. tetragona was inversely correlated to latitude and yearly mean humidity. The number of helminths per partridge and the number of helminth species per partridge were lower in young birds than in adults. Partridge body condition was inversely correlated to abundance of C. contorta. Richer infracommunities were linked to richer component communities. At the infracommunity level, total number of helminths per partridge and number of helminth species per partridge were inversely correlated to latitude and directly correlated to yearly mean temperature. At the component community level, both species richness and diversity (Simpson's index) were inversely correlated to latitude and directly correlated to mean temperature. Across the broad geographical range of the study area, the helminth parasite communities of red-legged partridges had marked geographical variation in their structure. Our results suggest that this variation is determined by the distribution of both intermediate and definitive hosts. We discuss the implications of this variation for the hypothesis that supplementary releases of captive-bred partridges for sport hunting can affect the helminth fauna of wild red-legged partridges.     

Distribution of the flagellate Hexamita salmonis Moore, 1922 and the microsporidian Loma salmonae Putz, Hoffman and Dunbar, 1965 in brown trout, Salmo trutta L., and rainbow trout, Salmo gairdneri Richardson, in the River Itchen (U.K.) and three of its fish farms

The occurrence of Hexamita salmonis Moore, 1922 and Loma salmonae Putz, Hoffman and Dunbar, 1965 was investigated at 10 sites on the R. Itchen (five for brown trout only, three for rainbow trout only, and two for both brown trout and rainbow trout) and at three of its nine fish farms (two for rainbow trout, one for brown trout). Hexamita salmonis was recorded in brown trout from three river sites and the farm, and in rainbow trout from both farms and four river sites. Prevalence of Hexamita salmonis in farmed rainbow trout was higher than in farmed brown trout and was consistent with the former species being more susceptible to infection. H. salmonis was at significantly higher prevalence in rainbow trout from farm no. 5 than farm no. 2 for three size classes of fish. In wild brown trout and feral rainbow trout, the highest prevalences of H. salmonis were recorded at sites in the vicinity of farm no. 2. This distribution was consistent with an area of naturally high infection levels, and with infected fish unintentionally released from farm no. 2 serving as a source of infection, the infection subsequently becoming established in the river fish. Loma salmonae was recorded in wild brown trout and in rainbow trout from both farms. This appears to be the first recording of this parasite from British salmonids and also the first recording of the parasite from brown trout. The distribution of the parasite (particularly the prevalence being higher at farm no. 2 than farm no. 5) was consistent with it being introduced into the R. Itchen via rainbow trout from farm no. 2 (and probably no. 3) much of whose stock derived from imported Californian 'Shasta' rainbow trout.