Structure of the parasite communities of a coral reef fish assemblage (Labridae): testing ecological and phylogenetic host factors

The role of ecological and phylogenetic processes is fundamental to understanding how parasite communities are structured. However, for coral reef fishes, such information is almost nonexistent. In this study, we analyzed the structure of the parasite communities based on composition, richness, abundance, and biovolume of ecto- and endoparasites of 14 wrasse species (Labridae) from Lizard Island, Great Barrier Reef, Australia. We determine whether the structure of the parasite communities from these fishes was related to ecological characteristics (body size, abundance, swimming ability, and diet) and/or the phylogenetic relatedness of the hosts. We examined 264 fishes from which almost 37,000 individual parasites and 98 parasite categories (types and species) were recorded. Gnathiid and cestode larvae were the most prevalent and abundant parasites in most fishes. Mean richness, abundance, and biovolume of ectoparasites per fish species were positively correlated with host body size only after controlling for the host phylogeny, whereas no such correlation was found for endoparasites with any host variable. Because most ectoparasites have direct transmission, one possible explanation for this pattern is that increased space (host body size) may increase the colonization and recruitment of ectoparasites. However, endoparasites generally have indirect transmission that can be affected by many other variables, such as number of prey infected and rate of parasite transmission.     

The influence of a parasite community on the dynamics of a host population: a longitudinal study on willow ptarmigan and their parasites

Despite the fact that most host populations are infected by a community of different parasite species, the majority of empirical studies have focused on the interaction between the host and a single parasite species. Here, we explore the hypothesis that host population dynamics are affected both by single parasite species and by the whole parasite community. We monitored population density and breeding productivity of two populations of willow ptarmigan ( Lagopus lagopus ) in northern Norway for 8 and 11 years, respectively, and sampled eukaryotic endoparasites. We found that increasing abundances of the cestode Hymenolepis microps was associated with increased breeding mortality and reduced annual growth rate of the host population in both areas, and reduced host body mass and body condition in the area where such data were available. In one of the areas, the abundance of the nematode Trichostrongylus tenuis was associated with reductions in host body mass, body condition and breeding mortality and the filaroid nematode Splendidofilaria papillocerca was negatively related to host population growth rates. The parasite community was also negatively related to host fitness parameters, suggesting an additional community effect on host body mass and breeding mortality, although none of the parasites had a significant impact on their own. The prevalence of parasites with very different taxonomical origins tended to covary within years, suggesting that variability in the parasite community was not random, but governed by changes in host susceptibility or environmental conditions that affected exposure to parasites in general. Other variables including climate, plant production and rodent densities were not associated with the recorded demographic changes in the host population.     

The relationship between species richness and productivity in metazoan parasite communities

Biodiversity is not distributed homogeneously in space, and it often covaries with productivity. The shape of the relationship between diversity and productivity, however, varies from a monotonic linear increase to a hump-shaped curve with maximum diversity values corresponding to intermediate productivity. The system studied and the spatial scale of study may affect this relationship. Parasite communities are useful models to test the productivity-diversity relationship because they consist of species belonging to a restricted set of higher taxa common to all host species. Using total parasite biovolume per host individual as a surrogate for community productivity, we tested the relationship between productivity and species richness among assemblages of metazoan parasites in 131 vertebrate host species. Across all host species, we found a linear relationship between total parasite biovolume and parasite species richness, with no trace of a hump-shaped curve. This result remained after corrections for the potential confounding effect of the number of host individuals examined per host species, host body mass, and phylogenetic relationships among host species. Although weaker, the linear relationship remained when the analyses were performed within the five vertebrate groups (fish, amphibians, reptiles, mammals and birds) instead of across all host species. These findings agree with the classic isolationist-interactive continuum of parasite communities that has become widely accepted in parasite ecology. They also suggest that parasite communities are not saturated with species, and that the addition of new species will result in increased total parasite biovolume per host. If the number of parasite species exploiting a host population is not regulated by processes arising from within the parasite community, external factors such as host characteristics may be the main determinants of parasite diversity.     

Community structure of helminth parasites of the tuna, Euthynnus affinis, from the Visakhapatnam coast, Bay of Bengal

An analysis is made of the community structure of the helminth parasites of the tuna Euthynnus affinis collected off the coast of Visakhapatnam, Bay of Bengal. The helminth fauna comprised 23 species, consisting of two monogeneans, 18 digeneans, one larval cestode, one nematode and one acanthocephalan. Didymozoids, represented by 14 species, were the dominant members of the parasite spectrum. The parasite assemblage is characterized by species richness as well as a high diversity. Most of the parasites found, the didymozoids in particular, were host specialists, exhibiting a high degree of host specificity. The fauna comprised three core species, four satellite species and many secondary species, with no potential for interaction among them but the degree of predictability is high in that each infracommunity comprised two or three core species superimposed by a few secondary species. Many host factors such as the varied diet, high vagility, long life span and endothermy appear to have contributed to the development of the species rich and diverse parasite communities in the tuna. There was evidence for a decrease in parasite density and an increase in diversity with increase in host size, indicating that host size has a profound influence on the component community structure.     

Host population density and body mass as determinants of species richness in parasite communities: comparative analyses of directly transmitted nematodes of mammals

Epidemiological theory predicts positive correlations between host population density or body mass and species richness among parasite communities. Here I test these predictions by a comparative study of communities of directly transmitted mammalian parasites, gastrointestinal strongylid nematodes. I use data from 45 species of mammals, representing examination of 17 200 individual hosts. The variable studied was the average number of gastrointestinal strongylid nematode species per host population, and three different methods were used to obtain estimates of parasite species richness that are unbiased by number of host individuals examined. Analyses were done using the phylogenetically independent contrast method. Host population density and parasite species richness were strongly positively correlated when the effects of host body weight had been controlled for. Controlling for other variables did not change this, and the relationship was found regardless of method used to correct for uneven sampling effort among host species. A positive relationship between parasite species richness and host body weight was also found, but the effect of host densities had to be controlled for to see this. These relationships between host traits and species richness of directly transmitted parasites are stronger than patterns found using data on indirectly transmitted mammalian parasites, and suggests that links between host traits and parasite species richness are stronger than previously suggested. The results are consistent with parasite species richness being positively linked to pathogen transmission rates and reductions in transmission rates possibly increasing extinction probabilities in parasite populations. The results also suggest that parasites may exert a cost of increases in rate of population energy usage, and thus show that pathogens may be important in generating independence between body mass and rate of population energy usage among host species.     

Fungal infections of the phytoplankton: seasonality, minimal host density, and specificity in a mesotrophic lake

Phytoplankton infections by fungal parasites in the upper, mixed layer of a mesotrophic northern temperate lake were analysed according to the following parameters: host and parasite species, host population density and prevalence of infection, resting spore formation by the parasite, and the lowest host density at which parasites appeared. The phytoplankton taxa recorded included the Cyanobacteria, Dinomastigota, Chrysophyceae, Bacillariophyceae, Chlorophyceae, Cryptophyceae and Haptophyceae, but infection was never found in the last two classes. The parasites belonged almost exclusively to the monocentric Chytridiomycetes. Fungal epidemics occurred at all times of the year. Parasites appeared at population densities as low as about 1 cell ml⁻¹ in some host species, with infection prevalence sometimes exceeding 80%. The proportion of the total phytoplankton biovolume infected by fungi was usually much <1%, but occasionally reached 10%. Parasitism proved to be highly species-specific, with one parasite species usually infecting only one host species. In the case of Zygorhizidium planktonicum , which infected both Asterionella formosa and Synedra acus , there is evidence that two species-specific formae speciales , each infecting only one of these two host species, are present in the lake.     

Communities of metazoan parasites in open water fishes of Cold Lake, Alberta

Communities of metazoan parasites in ten species of fishes from Cold Lake, Alberta are described and compared. Relative abundances in the overall community of parasites in the lake were estimated using data on abundance in each host species, plus estimates of the relative abundances of the species of hosts. Parasites of the numerically dominant salmonid fishes dominated the overall community, with over half of the individual parasites being Metechinorhynchus salmonis . Exchange of parasites between host species was greatest between related and/or abundant host species. Parasite communities in cisco and whitefish, Coregonus spp., were relatively rich in species and diverse (Simpson's index) compared with communities in those species in other lakes in North America; communities in lake trout, Salvelinus spp., and the non-salmonid fishes were poor in species and low in diversity compared with communities in other lakes. Parasite species overlaps (Jaccard index) between related host species in Cold Lake were greater than overlaps within host species between lakes. The same pattern is seen in data from some other lakes. These features support the hypothesis of Wisiewski (1958); that the parasite community within an ecosystem is characterized by parasites of the numerically dominant hosts.     

Comparing the richness of metazoan ectoparasite communities of marine fishes: controlling for host phylogeny

Parasite communities are the product of acquisitions and losses of parasite species during the evolutionary history of their host. When comparing the parasite communities of different host species to assess the role of ecological variables as determinants of parasite species richness, a correction must be made for the possible phylogenetic inheritance of parasites from ancestral hosts independent of host ecology. We performed a comparative analysis of the metazoan ectoparasite communities on the heads and gills of 111 species of marine fish. The influences of host body size, host schooling behaviour and water temperature were tested after controlling for both sampling and phylogenetic effects. Overall, water temperature correlated positively with both parasite species richness and abundance, whereas fish size only correlated with parasite abundance. The correlation across all fish species between water temperature and parasite species richness was dependent on an outlier point. The results, however, generally held when fish from different biogeographical areas (Pacific and Atlantic) were analysed separately. In all analyses, parasite species richness always correlated strongly with parasite abundance. There was no evidence that schooling fish taxa harboured richer or more abundant ectoparasite communities than their non-schooling sister taxa, possibly because of the small number of contrasts available for that test. Overall, whereas both water temperature and host size affect the number of parasite individuals that can be harboured by a fish, only temperature appears important as a determinant of ectoparasite community richness. Received: 30 May 1996 / Accepted: 23 October 1996     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

Patterns in metazoan parasite communities of some oyster species

Metazoan parasite communities of Crassostrea gigas and Ostrea edulis from Great Britain, Crassostrea virginica from Mexico, and Saccostrea commercialis from Australia are described and summarized in terms of species composition, species richness, total number of individuals and dominance. Metazoan parasite communities in all host species were composed of turbellarians and the metacercarial stage of digeneans, with the exception of S. commercialis where only metacercariae were found. Arthropods, including one copepod and one mite species, were present only in British oyster species. All metazoan parasite communities of oysters had few species and low density of individuals. Richest communities were found in C. virginica at both component and infracommunity level. The least diverse component community occurred in S. commercialis. Infracommunities in O. edulis and S. commercialis never exceeded one species per host. The host response against parasites is suggested as the principal factor responsible for depauperate parasite communities of oysters. Environmental factors characteristic of tropical latitudes are likely to have enhanced both the number of species and the densities of parasites per host in the infracommunities of C. virginica.     

An ecological law and its macroecological consequences as revealed by studies of relationships between host densities and parasite prevalence

Epidemiological models predict a positive relationship between host population density and abundance of macroparasites. Here I lest these by a comparative study. I used data on communities of four groups of parasites inhabiting the gastrointestinal tract of mammals, nematodes of the orders Oxyurida. Ascarida. Enoplida and Spirurida. respectively. The data came from 44 mammalian species and represent examination of 16886 individual hosts. I studied average prevalence of all nematodes within an order in a host species, a measure of community level abundance, and considered the potential confounding effects of host body weight, fecundity, age at maturity and diet. Host population density was positively correlated with parasite prevalence within the order Oxyurida, where all species have direct life cycles. Considering the effects of other variables did not change this. This supports the assumption that parasite transmission rate generally is a positive function of host population density_: It also strengthens the hypothesis that host densities generally act as important determinants of species richness among directly transmitted parasites and suggests that negative influence of such parasites on host population growth rate increase with increasing host population density among host species. Within the other three nematode orders, where a substantial number of the species have indirect life cycles, no relationships between prevalence and host population density were seen, Again. considering the effects of other variables did not affect this conclusion. This suggests that host population density is a poor predictor of species richness of indirectly transmitted parasites and that effects of such parasites on host population dynamics do not scale with host densities among species of hosts.     

Microhabitat distributions and species interactions of ectoparasites on the gills of cichlid fish in Lake Victoria,Tanzania

Heterogeneous exposure to parasites may contribute to host species differentiation. Hosts often harbour multiple parasite species which may interact and thus modify each other’s effects on host fitness. Antagonistic or synergistic interactions between parasites may be detectable as niche segregation within hosts. Consequently, the within-host distribution of different parasite taxa may constitute an important axis of infection variation among host populations and species. We investigated the microhabitat distributions and species interactions of gill parasites (four genera) infecting 14 sympatric cichlid species in Lake Victoria, Tanzania. We found that the two most abundant ectoparasite genera (the monogenean Cichlidogyrus spp. and the copepod Lamproglena monodi) were non-randomly distributed across the host gills and their spatial distribution differed between host species. This may indicate microhabitat selection by the parasites and cryptic differences in the host–parasite interaction among host species. Relationships among ectoparasite genera were synergistic: the abundances of Cichlidogyrus spp. and the copepods L. monodi and Ergasilus lamellifer tended to be positively correlated. In contrast, relationships among morphospecies of Cichlidogyrus were antagonistic: the abundances of morphospecies were negatively correlated. Together with niche overlap, this suggests competition among morphospecies of Cichlidogyrus. We also assessed the reproductive activity of the copepod species (the proportion of individuals carrying egg clutches), as it may be affected by the presence of other parasites and provide another indicator of the species specificity of the host–parasite relationship. Copepod reproductive activity did not differ between host species and was not associated with the presence or abundance of other parasites, suggesting that these are generalist parasites, thriving in all cichlid species examined from Lake Victoria.     

Monogenean body size,but not reproduction,increases with infracommunity density

Body size reveals a plethora of life-history, ecological, and evolutionary information about a species. It plays a critical role in success or failure during competitive, reproductive, or predator–prey interactions. Typically, there is a negative relationship between body size and population density in natural populations and communities. I analysed this relationship within and among multiple populations of two prominent monogenean parasites (>90% prevalence) on Lepomis macrochirus in three lakes in New Jersey (USA), using multiple regression models. To elucidate the causes and benefits of this relationship, I also measured host body condition via a regression index, and reproductive output of the parasite community by measuring parasite eggs shed from the host. The relationship between body size and density of infrapopulations (parasites of a single species on a single host) was positive, and the strength of this relationship for both species depended on which lake they occupied, indicating the potential for Allee effects. This relationship persists at the infracommunity level, where there was a similar positive relationship between a community weighted mean body size and density. However, this relationship did not result in greater reproductive success as measured by infracommunity egg production per individual per 24 h or egg size. The cause of this relationship also remains elusive; it was not explained by host condition or age. The results suggest that there is either no reproductive advantage to this increase in body size or the advantage conferred was not related to these measured fitness components. These findings indicate that researchers should be cautious using body size as a proxy for fitness or reproduction, while also raising further questions about the nature of the relationship between parasites on a host and that between those parasites and the host.     

Bottom–up regulation of parasite population densities in freshwater ecosystems

Theory predicts the bottom–up coupling of resource and consumer densities, and epidemiological models make the same prediction for host–parasite interactions. Empirical evidence that spatial variation in local host density drives parasite population density remains scarce, however. We test the coupling of consumer (parasite) and resource (host) populations using data from 310 populations of metazoan parasites infecting invertebrates and fish in New Zealand lakes, spanning a range of transmission modes. Both parasite density (no. parasites per m²) and intensity of infection (no. parasites per infected hosts) were quantified for each parasite population, and related to host density, spatial variability in host density and transmission mode (egg ingestion, contact transmission or trophic transmission). The results show that dense and temporally stable host populations are exploited by denser and more stable parasite populations. For parasites with multi‐host cycles, density of the ‘source’ host did not matter: only density of the current host affected parasite density at a given life stage. For contact‐transmitted parasites, intensity of infection decreased with increasing host density. Our results support the strong bottom–up coupling of consumer and resource densities, but also suggest that intraspecific competition among parasites may be weaker when hosts are abundant: high host density promotes greater parasite population density, but also reduces the number of conspecific parasites per individual host.     

Species abundance distributions and numerical dominance in gastrointestinal helminth communities of fish hosts

The abundances of different species in a parasite community are never similar: there is typically one or a few numerically dominant species and many species with low abundance. Here, we determine whether basic features of parasite communities are associated with strong dominance by one or a few species, among 39 component communities of gastrointestinal helminths in marine fishes from Brazil. First, we tested whether the shape of the species abundance distribution in these communities fits that predicted by several theoretical models, using a goodness-of-fit procedure. Only the canonical lognormal model could be rejected for 5 out of 39 communities; all other comparisons of observed and predicted abundance distributions showed no significant differences, although this may be due to limited statistical power. Second, we used the ratio between the abundance of the most abundant species and either the second or third most abundant species, as indices of dominance; these show, for instance, that the dominant species in a community is typically twice, but sometimes over ten times, as abundant as the next most abundant species. We found that these ratios were not influenced by either the community's species richness, the mean number of individual parasites per host, or the taxonomic identity of the dominant species. However, the abundance ratio between the first and third most abundant species in a community was significantly correlated with an independent index of species interactivity, based on the likelihood that the different parasite species in a component community co-occur in the same host individuals: the difference in abundance between the dominant and third most abundant species was greater in communities characterized by weak interactions. These findings suggest that strong interactions may lead to greater evenness in the abundance of species, and that numerical dominance is more likely to result from interspecific differences in recruitment rates.     

Drivers of parasite community structure in fishes of the continental shelf of the Western Mediterranean: the importance of host phylogeny and autecological traits

We explored the relationships between features of host species and their environment, and the diversity, composition and structure of parasite faunas and communities using a large taxonomically consistent dataset of host-parasite associations and host-prey associations, and original environmental and host trait data (diet, trophic level, population density and habitat depth vagility) for the most abundant demersal fish species off the Catalonian coast of the Western Mediterranean. Altogether 98 species/taxa belonging to seven major parasite groups were recovered in 683 fish belonging to 10 species from seven families and four orders. Our analyses revealed that (i) the parasite fauna of the region is rich and dominated by digeneans; (ii) the host parasite faunas and communities exhibited wide variations in richness, abundance and similarity due to a strong phylogenetic component; (iii) the levels of host sharing were low and involved host generalists and larval parasites; (iv) the multivariate similarity pattern of prey samples showed significant associations with hosts and host trophic guilds; (v) prey compositional similarity was not associated with the similarity of trophically transmitted parasite assemblages; and (vi) phylogeny and fish autecological traits were the best predictors of parasite community metrics in the host-parasite system studied.     

Species affinity and infracommunity ordination of helminths of Leptodactylus chaquensis (Anura: Leptodactylidae) in two contrasting environments from northeastern Argentina

One hundred seventy-two frogs (Leptodactylus chaquensis) were collected from November 2002 to November 2003, in agricultural (n = 132) and nonagricultural (n = 40) areas. Both sites are near the city of Corrientes, Argentina. The main goals of this study were as follows: (1) to determine the helminth parasite community in agricultural and nonagricultural habitats; (2) to analyze the relationships between helminth parasites and site of infection, frog body size, and gender; (3) to identify and examine covariation and association of helminth communities; and (4) to determine the mean richness and diversity of parasite communities. The helminth compound community of this amphibian species consisted of 24 species: 19 in agricultural habitats and 18 in nonagricultural habitats. The mean richness, mean diversity, and evenness of helminths were significantly different between the habitats (P < 0.05). The body size of the host was the important factor in determining parasite richness. Both habitats exhibited differences in community ordination. The helminth species in the 2 habitats exhibited the same interspecific relationships, although differences were observed in the intensity of infection.     

Come with me if you want to live: sympatric parasites follow different transmission routes through aquatic host communities

Community composition, including the relative density of each host species, plays a vital role in the transmission of parasites or disease in freshwater ecosystems. Whereas some host species can effectively transmit parasites, others can act as dead ends (non-viable transmission routes), accumulating large numbers of parasites throughout their life, thus becoming important sinks for parasite populations. Although population sinks have been identified in certain host-parasite systems, robust field estimates of the proportions of parasites that are lost to these hosts are lacking. Here, we quantified the distribution of encysted larval hairworms (phylum Nematomorpha), common parasites in lotic ecosystems, in two subalpine stream communities of New Zealand. With parasite and host population densities calculated per m², we identified which host species most likely contributed to the transmission of three sympatric hairworm morphotypes identified in both streams, and which species acted as population sinks. We also tested for seasonal patterns and peaks in the abundance of each morphotype in the two communities over the sampling season. Finally, we tested whether hosts emerging from the streams had comparable abundances of hairworm morphotypes throughout the sampling period. For each morphotype, different key sets of host species harboured more hairworms on average (abundance) than others, depending on the stream. For one morphotype in particular, two species of hosts were found to be important population sinks that inhibited over a third of these parasites from completing their life cycle. We also observed a clear peak in abundance for another hairworm morphotype during summer. Our data suggest that hosts emerging from the streams matched their aquatic counterparts with respect to hairworm abundance, indicating no infection-dependent reduction in emergence success. Our findings suggest that, depending on relative community composition, sympatric parasites follow different host transmission pathways, some of which lead to dead ends that potentially impact overall infection dynamics. In turn, this information can help us understand the spread or emergence of disease in both freshwater and terrestrial environments, since hairworms infect terrestrial arthropods to complete their life cycle.     

Interspecific aggregation of ectoparasites on bats: importance of hosts as habitats supersedes interspecific interactions

Patterns of aggregation of species or individuals may result from combinations of interspecific interactions such as competition, facilitation, or apparent facilitation, as well as from equivalent responses to environmental factors. Host–parasite systems are ideal for the investigation of mechanisms that structure assemblages. Interspecific aggregation is documented for multiple groups that are ectoparasitic on mammals and host‐mediated apparent facilitation has been suggested to explain these aggregation patterns. To investigate the generality of this pattern and to determine likely structuring mechanisms, I analyzed species co‐occurrence, correlations of abundances, and nestedness for ectoparasite assemblages from each of 11 species of Neotropical bat. Ectoparasite assemblages on four of 11 host species exhibited significant positive co‐occurrence for the entire assemblage or for at least one pair of species in the assemblage; ectoparasites on two host species exhibited positive co‐occurrence that approached significance. There was no evidence of negative co‐occurrence. Nine species‐pairs exhibited positive abundance correlations, including seven of the eight species‐pairs that exhibited positive co‐occurrence. No species‐pair exhibited a negative correlation of abundances (i.e. density compensation). Ectoparasite assemblages from five of 11 host species exhibited nestedness, including all three assemblages that exhibited assemblage‐wide positive co‐occurrence. Multiple mechanisms associated with host characteristics may contribute to host aggregation in ectoparasite assemblages, including host body size, vagility, home range size, burrow or roost size and complexity, immunocompetence and social structure. In general, data in this study and elsewhere are not consistent with interspecific interactions among ectoparasites, including apparent facilitation, being primary structuring mechanisms of ectoparasite assemblages on mammalian hosts. Rather, host behavior and ecology are likely to affect the frequency of host–ectoparasite encounters and of conspecific host interactions that facilitate transfer of ectoparasites, thereby, molding patterns of ectoparasite co‐occurrence, abundance and species composition on mammalian hosts. Combinations of characteristics that are primarily responsible for molding ectoparasite assemblage composition likely are host‐taxon specific.     

Speciation and diversification of parasite lineages: an analysis of congeneric parasite species in vertebrates

The evolutionary diversification of living organisms is a central research theme in evolutionary ecology, and yet it remains difficult to infer the action of evolutionary processes from patterns in the distribution of rates of diversification among related taxa. Using data from helminth parasite communities in 76 species of birds and 114 species of mammals, the influence of four factors that may either be associated with or modulate rates of parasite speciation were examined in a comparative analysis. Two measures of the relative number of congeneric parasite species per host species were used as indices of parasite diversification, and related to host body mass, host density, latitude, and whether the host is aquatic or terrestrial. The occurrence of congeneric parasites was not distributed randomly with respect to these factors. Aquatic bird species tended to harbour more congeneric parasites than terrestrial birds. Large-bodied mammal species, or those living at low latitudes, harboured more congeneric parasites than small-bodied mammals, or than those from higher latitudes. Host density had no apparent association with either measures of parasite diversification. These patterns, however, reflect only the present-day distribution of parasite diversification among host taxa, and not the evolutionary processes responsible for diversification, because the apparent effects of the factors investigated disappeared once corrections were made for host phylogeny. This indicates that features other than host body size, host density, latitude, and whether the habitat is terrestrial or aquatic, have been the key driving forces in the diversification of parasitic helminth lineages. This revised version was published online in July 2006 with corrections to the Cover Date.