Community ecology of the metazoan parasites of namorado sandperches, Pseudopercis numida Miranda-Ribeiro, 1903 and P. semifasciata Cuvier, 1829 (Perciformes: Pinguipedidae), from the coastal zone of the State of Rio de Janeiro, Brazil

One hundred and twenty-eight specimens of namorado sandperches, 62 P. numida and 66 P. semifasciata, collected between October 2002 and June 2003 off the Cabo Frio, Rio de Janeiro, Brazil (Lat 23 degrees S and Long 42 degrees W), were examined to study their metazoan parasites. Parasite communities of these fish were composed basically of endoparasites, mainly digenean and cestodes species, with low prevalence and abundance but having high parasite species richness values (at the component community level). Among these values, that found for P. numida is the highest so far recorded for marine fishes from the Neotropical Region. Thirty-nine species of metazoan parasites were collected: 36 from P. numida and 28 from P. semifasciata. Twenty-five parasite species were common to both species of namorado sandperches. Pseudopercis numida and P. semifasciata are new host records for all parasite species collected, with the exception of Microcotyle pseudopercis. Choanodera sp., Leurodera decora, Neolebouria georgenascimentoi, and Proctoeces sp. which were recorded for the first time in the South American Atlantic Ocean. Gnathia sp. from P. numida and Scolex pleuronectis from P. semifasciata were the species having the greatest dominance frequency. Parasite abundance in P. numida and P. semifasciata were positively correlated with the host total length. Only in P. numida was parasite species richness correlated positively with the host total length. No significant differences between endoparasite infracommunities of P. numida and P. semifasciata were detected. The ectoparasites of P. numida had higher values for parasite abundance, parasite richness, Brillouin index, evenness index, and Berger-Parker index than those of the ectoparasites of P. semifasciata. Comparisons among all ecto- and endoparasites showed the ectoparasites of P. numida as the most heterogeneous group. Low similarity values were observed among the three types of parasite infracommunities of the two hosts.     

Linking ecology with parasite diversity in Neotropical fishes

A comparative analysis was performed to seek large-scale patterns in the relationships between a set of fish species traits (body size, type of environment, trophic level, schooling behaviour, depth range, mean habitat temperature, geographical range, ability to enter brackish waters and capability of migration) and the diversity of their metazoan parasite assemblages among 651 Neotropical fish species. Two measurements of parasite diversity are used: the species richness and the taxonomic distinctness of a fish's parasite assemblage, including all metazoan parasites, ectoparasites only, or endoparasites only. The results showed that, on this scale, the average taxonomic distinctness of parasite assemblages was clearly more sensitive to the influence of host traits than parasite species richness. Differences in the taxonomic diversification of the parasite assemblages of different fish species were mainly related to the fish's environment (higher values in benthic–demersal species), trophic level (positive correlation with increasing level), temperature (positive correlation with temperature in marine ectoparasites, negative in endoparasites; positive for all groups of parasites in freshwater fishes) and oceanic distribution (higher values in fish species from the Pacific Ocean than those of the Atlantic). The results suggest that, among Neotropical fish species, only certain key host traits have influenced the processes causing the taxonomic diversification of parasite assemblages.     

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     

Body size vs abundance among parasite species: positive relationships?

Across species, abundance usually correlates negatively with body size. This intuitive pattern may result from size-dependent resource requirements in habitats where only finite amounts of resources are available. Among parasite species, it is possible that some resource limitations arc less severe than for free-living animals although this may depend on the type of parasites. The interspecific relationship between body size and abundance (measured as prevalence and intensity of infection) among parasites was tested in two groups of parasites. Among helminth endoparasites of fish, parasite body size correlated positively with prevalence and negatively with intensity of infection. Among copepod ectoparasites of fish, body size correlated positively with both prevalence and intensity. These trends were observed after controlling for the confounding influences of phylogeny and sampling effort. These contrasting patterns may result from the more intense link between body size and intensity-dependent regulation in endoparasites than in ectoparasites. The results of this comparative analysis suggest that parasite body size could be an important factor determining aspects of parasite abundance and distribution, including aggregation.     

Life history strategy influences parasite responses to habitat fragmentation

Anthropogenic habitat use is a major threat to biodiversity and is known to increase the abundance of generalist host species such as rodents, which are regarded as potential disease carriers. Parasites have an intimate relationship with their host and the surrounding environment and it is expected that habitat fragmentation will affect parasite infestation levels. We investigated the effect of habitat fragmentation on the ecto- and endoparasitic burdens of a broad niche small mammal, Rhabdomys pumilio, in the Western Cape Province, South Africa. Our aim was to look at the effects of fragmentation on different parasite species with diverse life history characteristics and to determine whether general patterns can be found. Sampling took place within pristine lowland (Fynbos/Renosterveld) areas and at fragmented sites surrounded and isolated by agricultural activities. All arthropod ectoparasites and available gastrointestinal endoparasites were identified. We used conditional autoregressive models to investigate the effects of habitat fragmentation on parasite species richness and abundance of all recovered parasites. Host density and body size were larger in the fragments. Combined ecto- as well as combined endoparasite taxa showed higher parasite species richness in fragmented sites. Parasite abundance was generally higher in the case of R. pumilio individuals in fragmented habitats but it appears that parasites that are more permanently associated with the host’s body and those that are host-specific show the opposite trend. Parasite life history is an important factor that needs to be considered when predicting the effects of habitat fragmentation on parasite and pathogen transmission.     

Infracommunity structure of parasites of Hemigymnus melapterus (Pisces: Labridae) from Lizard Island, Australia: the importance of habitat and parasite body size

This study describes the community of all metazoan parasites from 14 individuals of thicklip wrasse, Hemigymnus melapterus, from Lizard Island, Australia. All fish were parasitized, and 4,649 parasite individuals were found. Twenty-six parasite species were identified although only 6 species were abundant and prevalent: gnathiid isopods, the copepod Hatschekia hemigymni, the digenean Callohelmis pichelinae, and 3 morphotypes of tetraphyllidean cestode larvae. We analyzed whether the body size and microhabitat of the parasites and size of the host affected understanding of the structure of the parasite community. We related the abundance, biovolume, and density of parasites with the host body size and analyzed the abundances and volumetric densities of some parasite species within microhabitats. Although the 2 most abundant species comprised 75% of all parasite individuals, 4 species, each in similar proportion, comprised 85% of the total biovolume. Although larger host individuals had higher richness, abundance, and biovolume of parasites than smaller individuals, overall parasite volumetric density actually decreased with the host body size. Moreover, parasites exhibited abundances and densities significantly different among microhabitats; some parasite species depended on the area available, whereas others selected a specific microhabitat. Parasite and habitat size exhibited interesting relationships that should be considered more frequently. Considerations of these parameters improve understanding of parasite community structure and how the parasites use their habitats.     

Parasites of coral reef fish larvae: its role in the pelagic larval stage

The pelagic larval stage is a critical component of the life cycle of most coral reef fishes, but the adaptive significance of this stage remains controversial. One hypothesis is that migrating through the pelagic environment reduces the risk a larval fish has of being parasitised. Most organisms interact with parasites, often with significant, detrimental consequences for the hosts. However, little is known about the parasites that larval fish have upon settlement, and the factors that affect the levels of parasitism. At settlement, coral reef fishes vary greatly in size and age (pelagic larval duration), which may influence the degree of parasitism. We identified and quantified the parasites of pre-settlement larvae from 44 species of coral reef fishes from the Great Barrier Reef and explored their relationship with host size and age at settlement, and phylogeny. Overall, less than 50% of the larval fishes were infected with parasites, and over 99% of these were endoparasites. A Bayesian phylogenetic regression was used to analyse host-parasite (presence and intensity) associations. The analysis showed parasite presence was not significantly related to fish size, and parasite intensity was not significantly related to fish age. A phylogenetic signal was detected for both parasite presence and intensity, indicating that, overall, closely related fish species were likely to have more similar susceptibility to parasites and similar levels of parasitism when compared to more distantly related species. The low prevalence of infection with any parasite type and the striking rarity of ectoparasites is consistent with the ‘parasite avoidance hypothesis’, which proposes that the pelagic phase of coral reef fishes results in reduced levels of parasitism.

     

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.     

Determinants of parasite species richness on small taxonomical and geographical scales: Lamellodiscus monogeneans of northwestern Mediterranean sparid fish

Determinants of parasite species richness have been investigated in a host-parasite system comprising fish of the family Sparidae and their monogenean gill ectoparasites of the genus Lamellodiscus. This study was carried out on a small geographical scale in the northwestern Mediterranean Sea. Host phylogenetic relationships were taken into account by phylogenetic eigenvector regression which required the reconstruction of a phylogenetic tree for the sparid fish species using mtDNA sequences. Several ecological variables potentially acting on Lamellodiscus species richness were considered. Host body size and host migratory behaviour appeared to be the main determinants of parasite species richness in this system. It is concluded that structuring of monogenean communities is controlled more by ecological than evolutionary factors.     

Patterns of interactions of a large fish-parasite network in a tropical floodplain

1.?Describing and explaining the structure of species interaction networks is of paramount importance for community ecology. Yet much has to be learned about the mechanisms responsible for major patterns, such as nestedness and modularity in different kinds of systems, of which large and diverse networks are a still underrepresented and scarcely studied fraction. 2.?We assembled information on fishes and their parasites living in a large floodplain of key ecological importance for freshwater ecosystems in the Paraná River basin in South America. The resulting fish-parasite network containing 72 and 324 species of fishes and parasites, respectively, was analysed to investigate the patterns of nestedness and modularity as related to fish and parasite features. 3.?Nestedness was found in the entire network and among endoparasites, multiple-host life cycle parasites and native hosts, but not in networks of ectoparasites, single-host life cycle parasites and non-native fishes. All networks were significantly modular. Taxonomy was the major host's attribute influencing both nestedness and modularity: more closely related host species tended to be associated with more nested parasite compositions and had greater chance of belonging to the same network module. Nevertheless, host abundance had a positive relationship with nestedness when only native host species pairs of the same network module were considered for analysis. 4.?These results highlight the importance of evolutionary history of hosts in linking patterns of nestedness and formation of modules in the network. They also show that functional attributes of parasites (i.e. parasitism mode and life cycle) and origin of host populations (i.e. natives versus non-natives) are crucial to define the relative contribution of these two network properties and their dependence on other ecological factors (e.g. host abundance), with potential implications for community dynamics and stability.     

Community ecology of metazoan parasites of the anchovy Anchoa tricolor (Osteichthyes: Engraulidae) from the coastal zone of the State of Rio de Janeiro, Brazil.

Between October 2001 and March 2002, 103 specimens of A. tricolor from Angra dos Reis (23 degrees 01' S, 44 degrees 19' W), in the coastal zone of the State of Rio de Janeiro, Brazil, were analyzed in order to study their metazoan parasite infracommunities. Ten species of metazoan parasites were collected: 4 digeneans, 1 cestode, 1 acantocephalan, 2 nematodes, 1 copepod, and 1 hirudinean; 77.7% of the fishes were parasitized by one or more metazoan, with a mean of 3.5 +/- 6.2 parasite/fish. Digenean was the most dominant with 4 species that accounted for 53.2% of the total parasites collected; Ergasilus sp. was the most abundant species. Abundance and prevalence of Parahemiurus merus (Linton, 1910) were positively correlated with the total length of host. Relationships between total body length of fish and both total parasite abundance and mean parasite species richness were observed. Mean parasite diversity of species was correlated to host's total length, with significant differences found between male and female fishes. Two pairs of larval species showed significant positive association and covariation. The metazoan parasite infracommunities of A. tricolor presented dominance of larval endoparasites; correlation of parasite abundance, diversity, and species richness with host total length; and low number of parasite interspecific relationships. The parasite community of A. tricolor showed some similarities with the parasite community of another South American Atlantic engraulid.     

Evolutionary associations between host traits and parasite load: insights from Lake Tanganyika cichlids

Parasite diversity and abundance (parasite load) vary greatly among host species. However, the influence of host traits on variation in parasitism remains poorly understood. Comparative studies of parasite load have largely examined measures of parasite species richness and are predominantly based on records obtained from published data. Consequently, little is known about the relationships between host traits and other aspects of parasite load, such as parasite abundance, prevalence and aggregation. Meanwhile, understanding of parasite species richness may be clouded by limitations associated with data collation from multiple independent sources. We conducted a field study of Lake Tanganyika cichlid fishes and their helminth parasites. Using a Bayesian phylogenetic comparative framework, we tested evolutionary associations between five key host traits (body size, gut length, diet breadth, habitat complexity and number of sympatric hosts) predicted to influence parasitism, together with multiple measures of parasite load. We find that the number of host species that a particular host may encounter due to its habitat preferences emerges as a factor of general importance for parasite diversity, abundance and prevalence, but not parasite aggregation. In contrast, body size and gut size are positively related to aspects of parasite load within, but not between species. The influence of host phylogeny varies considerably among measures of parasite load, with the greatest influence exerted on parasite diversity. These results reveal that both host morphology and biotic interactions are key determinants of host–parasite associations and that consideration of multiple aspects of parasite load is required to fully understand patterns in parasitism.     

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.     

Ecological determinants of parasite acquisition by exotic fish species

Disease‐mediated threats posed by exotic species to native counterparts are not limited to introduced parasites alone, since exotic hosts frequently acquire native parasites with possible consequences for infection patterns in native hosts. Several biological and geographical factors are thought to explain both the richness of parasites in native hosts, and the invasion success of free‐living exotic species. However, the determinants of native parasite acquisition by exotic hosts remain unknown. Here, we investigated native parasite communities of exotic freshwater fish to determine which traits influence acquisition of native parasites by exotic hosts. Model selection suggested that five factors (total body length, time since introduction, phylogenetic relatedness to the native fish fauna, trophic level and native fish species richness) may be linked to native parasite acquisition by exotic fish, but 95% confidence intervals of coefficient estimates indicated these explained little of the variance in parasite richness. Based on R²‐values, weak positive relationships may exist only between the number of parasites acquired and either host size or time since introduction. Whilst our results suggest that factors influencing parasite richness in native host communities may be less important for exotic species, it seems that analyses of general ecological factors currently fail to adequately incorporate the physiological and immunological complexity of whether a given animal species will become a host for a new parasite.     

Conservatism of host specificity in parasites

As information becomes available for many groups of organisms a general pattern of phylogenetic conservatism in ecological characters or morphological traits is now widely recognized. Conversely, conservatism of external ecological attributes throughout a lineage is still a contentious theme in ecology. Moreover, the studies exploring this topic have focused on free-living organisms, and have ignored parasites. The main external ecological attribute of parasite species is certainly their host specificity, which is a key determinant of both their range size and local abundance. We address the subject of conservatism and predictability of host specificity using 2 large databases concerning, respectively, ectoparasites and endoparasites. We found a significant positive relationship between the numbers of host species infested by flea sister species. Moreover, this result was consistent whether we used sympatric or allopatric flea species, suggesting no influence of the mode of speciation on this conservatism of specificity. Additionally, our results showed that congeneric helminth species have more similar host taxonomic diversities than expected by chance, although this conservatism is due mostly to trematodes. Whilst there is evidence of conservatism, the moderate levels preclude robust prediction of host specificity for one species based on that of closely related species.     

Structure of populations and ecological nishes of ectoparasites in the parasite communities of small forest mammals

The paper reports the results of eight-year investigations on the ectoparasites of rodents and insectivores carried out in southern taiga of the Ilmen-Volkhov lowland (Novgorod Region) and Kurgolovsky reserve (Leningrad Region). Twelve species of small mammals were captured including three dominate species--bank vole Clethrionomys glareolus (2722 specimens), common shrew Sorex araneus (1658 specimens), and wood mouse Apodemus uralensis (367 specimens). Parasite community of the bank vole comprises 34 species of mites, ticks, and insects, the community of common shrew comprises 25 species, and the community of A. uralensis includes 28 species. Taxonomic diversity of the ectoparasite communities was shown to be based on the diversity of types of parasitism and ecological nishes of the host body. Permanent ectoparasites are found to be represented by 2 species of lie and 14 species of acariform mites. The group of temporary parasites includes 13 species of fleas, 10 species of gamasid mites. 3 ixodid species and 1 Trombiculidae. There is a common pool of temporary parasites of small mammals in the ecological system of taiga. Significance of different shrew and rodent species as hosts were found to be dependent on the population density in possible hosts and many other factors. Species diversity in the parasite communities of different small mammal species is dependent on the number of possible ecological nishes in the host body. Actual infill of these nishes by ectoparasites is usually lesser than potential one. Species composition of temporary parasites, their occurrence and abundance changes according to season. Interspecific competition in the temporary parasite species can decrease because of the seasonal disjunction of their population peaks. Diversification of the ecological niches of ectoparasites allow simultaneous feeding of more parasite individuals on one host, than in the case of parasitising of single species or several species with similar ecological nishes. The distribution of parasites on their hosts was also studied. The aggregative distribution has been found in ixodid larvae only, and the distribution of fleas was close to the Poisson distribution. Deviations from the aggregative distribution can be an effect of several independent factors, including limited ability of small mammals for providing numerous parasites with food. On the most part of hosts simultaneous parasitizing of no more than 1-3 individuals of each tick, mite, and flea species was registered. Excessive infestation by ectoparasites may probably be limited by effective reactions of self-purification in the mammal hosts.     

Phylogenetic signals in host–parasite associations for Neotropical bats and Nearctic desert rodents

Hosts and their parasites have strong ecological and evolutionary relationships, with hosts representing habitats and resources for parasites. In the present study, we use approaches developed to evaluate the statistical dependence of species trait values on phylogenetic relationships to determine whether host–parasite relationships (i.e. parasite infections) are contingent on host phylogeny. If host–parasite relationships are contingent on the ability of hosts to provide habitat or resources to parasites, and if host phylogeny is an effective surrogate for among‐host variation in habitat and resource quality, host–parasite relationships should evince phylogenetic signals (i.e. be contingent on host phylogeny). Because the strength of ecological relationships between parasites and their hosts may affect the likelihood of phylogenetic signals occurring in host–parasite relationships, we hypothesized that (1) host specificity would be positively correlated with the strength of phylogenetic signals and (2) the strength of phylogenetic signals will be greater for parasites that rely more on their host throughout their life cycle. Analyses were conducted for ectoparasites from tropical bats and for ectoparasites, helminths, and coccidians from desert rodents. Phylogenetic signals were evaluated for parasite presence and for parasite prevalence. The frequency of phylogenetic signal occurrence was similar for parasite presence and prevalence, with a signal detected in 24–27% of cases at the species level and in 67% and 15% of cases at the genus level for parasites of bats and rodents, respectively. No differences in signal strength or the likelihood of detecting a signal existed between groups of parasites. Phylogenetic signal strength was correlated with host specificity, suggesting that mechanisms increasing host specificity also increase the likelihood of a phylogenetic signal in host use by parasites. Differences in the transmission mode did not affect signal strength or the likelihood of detecting a signal, indicating that variation in host switching opportunities associated with the transmission mode does not affect signal strength.     

Group-living and the richness of the parasite fauna in Canadian freshwater fishes

Summary An increased transmission of ectoparasites among individual animals is considered to be an inevitable cost of living in groups, since several kinds of ectoparasites require close proximity between large numbers of hosts for successful transmission. However, we do not know whether individuals belonging to group-living species incur a greater risk of ectoparasitism than individuals of solitary species. Here, using published data from 3 families (60 species) of Canadian freshwater fishes, I test the hypothesis that group-living species are host to more species of contagious ectoparasites (copepods and monogeneans) than solitary host species. As the different fish species have been studied with varying intensity, I used the mean number of parasite species recorded per study as a standard measure of parasite numbers. Multiple regression analyses were performed separately for each family to determine the effects of group-living and of 3 other variables (host size, age, and range) on the richness of the recorded parasite fauna. Once the effects of the other variables were removed, I found no significant effect of sociality on the richness of the parasite fauna per fish species, for contagious ectoparasites and other types of parasites. Neither of the other variables had any influence on the numbers of parasite species per fish species. These results suggest that a richer ectoparasite fauna is not a cost of group-living in fishes.     

Host abundance,durability, basidiome form and phylogenetic isolation determine fungivore species richness

Species with close associations to a specific host species, such as parasites and phytophages, make immense contributions to biodiversity. Hence, factors determining the variation in species richness among hosts are a main focus of ecological research. Investigations of determining factors of fungivorous species among host species are still scarce. Based on ecological patterns of parasites and phytophages, we hypothesized that the species richness of tree‐fungus beetles of the family Ciidae (Coleoptera) would increase with increasing basidiome size, niche diversity of the growth form, durability, increasing abundance and decreasing phylogenetic isolation of the host fungus. Our generalized least‐squares model, controlled by host phylogeny, revealed that Ciidae species richness increases with host abundance, but decreases with host phylogenetic isolation. In contrast with our prediction, Ciidae species richness was higher in annual basidiomes than in perennials. Pileate basidiomes revealed higher species richness than resupinate and stipitate basidiomes, which may be interpreted as being a result of their higher host niche diversity. The importance of host abundance, measured on the landscape scale, corroborates that fungivore species richness among macrofungal hosts is determined by factors similar to those that determine parasite and phytophage species richness among their hosts. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 699–708.