Comparative tests of parasite species richness in primates

Some hosts harbor diverse parasite communities, whereas others are relatively parasite free. Many factors have been proposed to account for patterns of parasite species richness, but few studies have investigated competing hypotheses among multiple parasite communities in the same host clade. We used a comparative data set of 941 host-parasite combinations, representing 101 anthropoid primate species and 231 parasite taxa, to test the relative importance of four sets of variables that have been proposed as determinants of parasite community diversity in primates: host body mass and life history, social contact and population density, diet, and habitat diversity. We defined parasites broadly to include not only parasitic helminths and arthropods but also viruses, bacteria, fungi, and protozoa, and we controlled for effects of uneven sampling effort on per-host measures of parasite diversity. In nonphylogenetic tests, body mass was correlated with total parasite diversity and the diversity of helminths and viruses. When phylogeny was taken into account, however, body mass became nonsignificant. Host population density, a key determinant of parasite spread in many epidemiological models, was associated consistently with total parasite species richness and the diversity of helminths, protozoa, and viruses tested separately. Geographic range size and day range length explained significant variation in the diversity of viruses.     

Helminth species diversity of mammals: parasite species richness is a host species attribute

Studies investigating parasite diversity have shown substantial geographical variation in parasite species richness. Most of these studies have, however, adopted a local scale approach, which may have masked more general patterns. Recent studies have shown that ectoparasite species richness in mammals seems highly repeatable among populations of the same mammal host species at a regional scale. In light of these new studies we have reinvestigated the case of parasitic helminths by using a large data set of parasites from mammal populations in 3 continents. We collected homogeneous data and demonstrated that helminth species richness is highly repeatable in mammals at a regional scale. Our results highlight the strong influence of host identity in parasite species richness and call for future research linking helminth species found in a given host to its ecology, immune defences and potential energetic trade-offs.     

Habitat availability does not explain the species richness patterns of European lentic and lotic freshwater animals

Aim In Europe, the relationships between species richness and latitude differ for lentic (standing water) and lotic (running water) species. Freshwater animals are highly dependent on suitable habitat, and thus the distribution of available habitat should strongly influence large‐scale patterns of species richness. We tested whether habitat availability can account for the differences in species richness patterns between European lentic and lotic freshwater animals. Location Europe. Methods We compiled occurrence data of 1959 lentic and 2445 lotic species as well as data on the amount of lentic and lotic habitats across 25 pre‐defined biogeographical regions of European freshwaters. We used the range of elevation of each region as a proxy for habitat diversity. We investigated the relationships between species richness, habitat availability and habitat diversity with univariate and multiple regression analyses. Results Species richness increased with habitat availability for lentic species but not for lotic species. Species richness increased with elevational range for lotic species but decreased for lentic species. For both groups, neither habitat availability nor diversity could account for previously reported latitudinal patterns in species richness. For lotic species, richness declined with latitude, whereas there was no relationship between habitat availability and latitude. For lentic species, richness showed a hump‐shaped relationship with latitude, whereas available habitat increased with latitude. Main conclusions Habitat availability and diversity are poor predictors of species richness of the European freshwater fauna across large scales. Our results indicate that the distributions of European freshwater animals are probably not in equilibrium and may still be influenced by history, namely the recurrent European glaciations and possible differences in post‐glacial recolonization. The distributions of lentic species appear to be closer to equilibrium than those of lotic species. This lends further support to the hypothesis that lentic species have a higher propensity for dispersal than lotic species.     

Plant species richness of nature reserves: the interplay of area, climate and habitat in a central European landscape

Aim To detect regional patterns of plant species richness in temperate nature reserves and determine the unbiased effects of environmental variables by mutual correlation with operating factors. Location The Czech Republic. Methods Plant species richness in 302 nature reserves was studied by using 14 explanatory variables reflecting the reserve area, altitude, climate, habitat diversity and prevailing vegetation type. Backward elimination of explanatory variables was used to analyse the data, taking into account their interactive nature, until the model contained only significant terms. Results A minimal adequate model with reserve area, mean altitude, prevailing vegetation type and habitat diversity (expressed as the number of major habitat types in the reserve) accounted for 53.9% of the variance in species number. After removing the area effect, habitat diversity explained 15.6% of variance, while prevailing vegetation type explained 29.6%. After removing the effect of both area and vegetation type, the resulting model explained 10.3% of the variance, indicating that species richness further increased with habitat diversity, and most obviously towards warm districts. After removing the effects of area, habitat diversity and climatic district, the model still explained 9.4% of the variance, and showed that species richness (i) significantly decreased with increasing mean altitude and annual precipitation, and with decreasing January temperature in the region of the mountain flora, and (ii) increased with altitudinal range in regions of temperate and thermophilous flora. Main conclusions We described, in quantitative terms, the effects of the main factors that might be considered to be determining plant species richness in temperate nature reserves, and evaluated their relative importance. The direct habitat effect on species richness was roughly equal to the direct area effect, but the total direct and indirect effects of area slightly exceeded that of habitat. It was shown that the overall effect of composite variables such as altitude or climatic district can be separated into particular climatic variables, which influence the richness of flora in a context‐specific manner. The statistical explanation of richness variation at the level of families yielded similar results to that for species, indicating that the system of nature conservation provides similar degrees of protection at different taxonomic levels.     

Factors related to diversity of decomposer fungi in tropical forests

Recent studies suggest that host-preferences are common among certain groups of tropical fungal decomposers but rare in others, and sometimes occur where we least expect them. Host preferences among microfungi and ascomycetes that decompose leaf litter are common but usually involve differences in relative frequencies more than presence/absence, so their diversity may be loosely correlated with species richness of host trees. Strong host-specificity appears to be rare among wood decomposer fungi, whereas characteristics of their substrata and habitat are very important for this group. Anthropogenic disturbance predisposed a tropical forest to subsequent hurricane damage, and the resulting direct and indirect effects on host diversity and habitat heterogeneity were reflected in the decomposer fungal community more than sixty years after the original disturbance. While species richness of dictyostelid slime molds and functional diversity of their bacterial prey increased with disturbance, the more diverse microfungi and ascomycetes were apparently negatively affected by disturbance.     

Determinants of tapeworm species richness in elasmobranch fishes: untangling environmental and phylogenetic influences

Parasite species richness is a fundamental characteristic of host species and varies substantially among host communities. Hypotheses aiming to explain observed patterns of richness are numerous, and none is universal. In this study, we use tapeworm parasites of elasmobranch fishes to examine the phylogenetic and environmental influences on the variation in species richness for this specific system. Tapeworms are the most diverse group of helminths to infect elasmobranchs. Elasmobranchs are cosmopolitan in distribution and their tapeworm parasites are remarkably host specific; therefore, making this an ideal system in which to examine global patterns in species diversity. Here, we 1) quantify the tapeworm richness in elasmobranch fishes, 2) identify the host features correlated with tapeworm richness, and 3) determine whether tapeworm richness follows a latitudinal gradient. The individual and combined effects of host size, factors associated with water temperatures (influenced by latitude and depth), host habitat, and type of elasmobranch (shark or batoid) on measures of species diversity were assessed using general linear models. These analyses included tapeworm host records for 317 different elasmobranch species (124 species were included in our analyses) and were conducted with and without taking into account phylogenetic relationships between host species. Since sharks and batoids differ substantially in body form, analyses were repeated for each host subset. On average, batoids harboured significantly more tapeworm species than shark hosts. Tapeworm richness in sharks was influenced by median depth, whereas no predictor variable included in our models could adequately account for interspecific variation in tapeworm richness in batoid hosts. The taxonomic diversity of tapeworm assemblages of sharks and batoids was influenced by median depth and median latitude, respectively. When the influence of host phylogeny is accounted for, larger hosts harbour a greater tapeworm richness, whereas hosts exploiting wider latitudinal ranges harbour more taxonomically distinct tapeworm assemblages. Species richness and taxonomic diversity of tapeworm assemblages in elasmobranch fishes are influenced by different evolutionary pressures, including host phylogenetic relationships, space constraints and geographical area. Our results suggest that ca 3600 tapeworm species have yet to be described from elasmobranch fishes.     

AVIAN BROOD PARASITISM AND ECTOPARASITE RICHNESS–SCALE‐DEPENDENT DIVERSITY INTERACTIONS IN A THREE‐LEVEL HOST–PARASITE SYSTEM

Brood parasitic birds, their foster species and their ectoparasites form a complex coevolving system composed of three hierarchical levels. However, effects of hosts’ brood parasitic life‐style on the evolution of their louse (Phthiraptera: Amblycera, Ischnocera) lineages have never been tested. We present two phylogenetic analyses of ectoparasite richness of brood parasitic clades. Our hypothesis was that brood parasitic life‐style affects louse richness negatively across all avian clades due to the lack of vertical transmission routes. Then, narrowing our scope to brood parasitic cuckoos, we explored macroevolutionary factors responsible for the variability of their louse richness. Our results show that taxonomic richness of lice is lower on brood parasitic clades than on their nonparasitic sister clades. However, we found a positive covariation between the richness of cuckoos’ Ischnoceran lice and the number of their foster species, possibly due to the complex and dynamic subpopulation structure of cuckoo species that utilize several host species. We documented diversity interactions across a three‐level host parasite system and we found evidence that brood parasitism has opposing effects on louse richness at two slightly differing macroevolutionary scales, namely the species richness and the genera richness.     

Resistance to rust fungi in Lolium perenne depends on within-species variation and performance of the host species in grasslands of different plant diversity

The hypothesis that plant species diversity and genetic variation of the host species decrease the severity of plant diseases is supported by studies of agricultural systems, but experimental evidence from more complex systems is scarce. In an experiment with grassland communities of varying species richness (1, 2, 4, 8, 16, and 60 species) and functional group richness (1, 2, 3, and 4 functional groups), we used different cultivars of Lolium perenne (perennial ryegrass) to study effects of biodiversity and cultivar identity on the occurrence and severity of foliar fungal diseases caused by Puccinia coronata (crown rust) and P. graminis (stem rust). Cultivar monocultures of perennial ryegrass revealed strong differences in pathogen susceptibility among these cultivars. Disease intensity caused by both rust fungi decreased significantly with growing species richness of species mixtures. The response to the diversity gradient was related to the decreased density and size of the host individuals with increasing species richness. The occurrence of other grass species known to be possible hosts of the pathogens in the experimental mixtures did not promote disease intensity in L. perenne, indicating that there was a high host specificity of pathogen strains. Differences in pathogen susceptibility among perennial ryegrass cultivars persisted independent of diversity treatment, host density and host individual size, but resulted in a cultivar-specific pattern of changes in pathogen infestation across the species-richness gradient. Our study provided evidence that within-species variation in pathogen susceptibility and competitive interactions of the host species with the environment, as caused by species diversity treatments, are key determinants of the occurrence and severity of fungal diseases. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

Context dependent effects of ectomycorrhizal species richness on tree seedling productivity

While there has been much recent interest about the relationships between plant diversity and plant productivity, much remains unknown about how the diversity of mycorrhizal fungi affects plant productivity. We investigated the effects of ectomycorrhizal fungal community composition and diversity on the productivity and growth characteristics of seedlings of two tree species ( Pinus sylvetris and Betula pendula ) as well as their interactions with each other. This involved setting up a mycorrhizal fungal diversity gradient from one to eight species using a design previously demonstrated to be able to separate diversity effects from compositional effects. We found that the eight mycorrhizal fungal species differed in their effects on seedling productivity and that the nature of effects was determined by the fertility of the substrate. Fungal species richness effects were also important in affecting seedling productivity over and above what could be explained by "sampling effect" but only in some situations. For B. pendula in a low fertility substrate there were clear positive causative effects between fungal species richness and productivity with the eight species treatment having over double the productivity of any of the eight monoculture treatments; no diversity effects were, however, detected in a high fertility substrate. For P. sylvestris in a high fertility substrate there were significant negative effects of fungal diversity on productivity while in a low fertility substrate no effects were apparent. The possible mechanistic bases for these results are discussed. The growth of P. sylvestris relative to that of B. pendula when grown in combination was unaffected by mycorrhizal treatments. Our results provide clear evidence that effects of mycorrhizal fungal diversity on productivity are context dependent and may be positive, negative or neutral depending on the situation considered.     

Cascade effects of crop species richness on the diversity of pest insects and their natural enemies

Understanding how plant species richness influences the diversity of herbivorous and predatory/parasitic arthropods is central to community ecology.We explore the effects of crop species richness on the diversity of pest insects and their natural enemies.Using data from a four-year experiment with five levels of crop species richness,we found that crop species richness significantly affected the pest species richness,but there were no significant effects on richness of the pests’natural enemies.In contrast,the species richness of pest insects significantly affected their natural enemies.These findings suggest a cascade effect where trophic interactions are strong between adjacent trophic levels,while the interactions between connected but nonadjacent trophic levels are weakened by the intermediate trophic level.High crop species richness resulted in a more stable arthropod community compared with communities in monoculture crops.Our results highlight the complicated cross-trophic interactions and the crucial role of crop diversity in the food webs of agro-ecosystems.     

Dissecting NDVI–species richness relationships in Hawaiian dry forests

Aim A growing body of research has used the normalized difference vegetation index (NDVI) as a proxy for productivity to predict species richness. Yet the mechanisms that produce the relationship between NDVI and species richness remain unclear because of correlated biotic and abiotic factors that influence NDVI. In this study we investigated different biotic and abiotic effects that potentially drive plant species richness–productivity relationships. Location Hawaiian Islands, USA. Methods We quantified woody plant species richness, structure (density, basal area and canopy height), and species composition along a precipitation gradient of 14 Hawaiian dry forest plots. We then used structural equation models combined with 10 years of satellite data to disentangle the effects of precipitation, structure and NDVI‐estimated productivity on species richness. Results Underlying the simple correlation between NDVI and species richness was the indirect effect of precipitation and direct effect of forest structure. The best‐fit model showed there was no direct effect of NDVI on species richness. Main conclusions Our results demonstrate that complex relationships drive simple correlations between species richness and productivity. Considering the mechanisms and underlying factors driving NDVI–species richness relationships could improve predictions of species diversity as satellite measures of productivity have an increasingly important role in habitat mapping, species distribution modelling and predictions for global change.     

Predictability of plant and fungal species richness of old‐growth boreal forest islands

Abstract. The fragmentation and deterioration of old‐growth forest habitat by modern forestry have become a major threat to species diversity in Fennoscandia. In order to develop a conservation strategy for the remaining diversity it is essential to identify the existing diversity and to develop appropriate conservation and monitoring programs. For these purposes indicators of conservation value for administrative prioritization are required. This study examines the predictability of plant and fungal species richness on two spatial scales on 46 isolated old‐growth forest islands (0.17 ‐ 12 ha) in a forest‐wetland mosaic. We explore (1) to what extent area, isolation and stand structure variables can explain the variation in species richness and (2) if richness patterns of individual species groups correlate. Isolation showed no relation to species richness. Area explained 50 ‐ 70% of the variation in total species richness and was positively related to the density of crustose lichens and Red‐list species in island interiors. Stand structure variables explained 28 ‐ 66% of the residual variation in total species richness after controlling for island size, and 15 ‐ 73% of the variation in density of species in island interiors. The highest predictability of species richness was found among substrate‐specific fungi and Red‐list species. Different stand structure variables were found to explain richness in the different species groups, and only among a few species groups species richness correlated. Thus, species richness of one single species group is unlikely to be a good indicator for total biodiversity. The results show that measurements of stand size and stand structure variables may be a strong complementary tool, and sometimes a substitute to extensive species inventories when one aims to estimate and monitor plant and fungal species diversity in old‐growth Picea abies forests.     

Human land use promotes the abundance and diversity of exotic species on Caribbean islands

Human land use causes major changes in species abundance and composition, yet native and exotic species can exhibit different responses to land use change. Native populations generally decline in human‐impacted habitats while exotic species often benefit. In this study, we assessed the effects of human land use on exotic and native reptile diversity, including functional diversity, which relates to the range of habitat use strategies in biotic communities. We surveyed 114 reptile communities from localities that varied in habitat structure and human impact level on two Caribbean islands, and calculated species richness, overall abundance, and evenness for every plot. Functional diversity indices were calculated using published trait data, which enabled us to detect signs of trait filtering associated with impacted habitats. Our results show that environmental variation among sampling plots was explained by two Principal Component Analysis (PCA) ordination axes related to habitat structure (i.e., forest or nonforest) and human impact level (i.e., addition of man‐made constructions such as roads and buildings). Several diversity indices were significantly correlated with the two PCA axes, but exotic and native species showed opposing responses. Native species reached the highest abundance in forests, while exotic species were absent in this habitat. Human impact was associated with an increase in exotic abundance and species richness, while native species showed no significant associations. Functional diversity was highest in nonforested environments on both islands, and further increased on St. Martin with the establishment of functionally unique exotic species in nonforested habitat. Habitat structure, rather than human impact, proved to be an important agent for environmental filtering of traits, causing divergent functional trait values across forested and nonforested environments. Our results illustrate the importance of considering various elements of land use when studying its impact on species diversity and the establishment and spread of exotic species.     

Ecosystem engineers maintain a rare species of butterfly and increase plant diversity

We evaluated whether ecosystem engineers can accomplish two conservation goals simultaneously: (1) indirectly maintain populations of an endangered animal through habitat modification and (2) increase riparian plant diversity. We tested for effects of a prominent ecosystem engineer, the beaver Castor canadensis, on populations of St. Francis' satyr butterfly Neonympha mitchellii francisci and plant species richness and composition. We performed our test by surveying riparian vegetation communities in all stages of beaver‐influenced wetland succession. We found that beavers created wetland habitats that supported plant species not found elsewhere in riparian zones and increased plant species diversity across the landscape by creating a novel combination of patch types. Our results confirmed what others have found about engineering effects on plant diversity, but these results further demonstrated a case where ecosystem engineers indirectly maintain populations of rare animals by modifying the composition and diversity of plant communities within wetlands. Our research demonstrates how an ecosystem engineer can influence habitat availability and composition of plant communities important for an endangered insect, and maintain overall plant species diversity by increasing habitat heterogeneity.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

Wood decomposition is more strongly controlled by temperature than by tree species and decomposer diversity in highly species rich subtropical forests

While the number of studies on the role of biodiversity on ecosystem functioning is steadily increasing, a key component of biogeochemical cycling in forests, dead wood decay, has been largely neglected. It remains widely unknown whether and how dead wood decay is affected by diversity loss in forests. We studied the hierarchical effects of tree species diversity on wood decay rates in a subtropical forest landscape in southeast China via its influence on fungal OTU richness and invertebrate diversity using piecewise structural equation models. The experiment was conducted in natural forest plots that span a wide gradient of tree species diversity embedded in a heterogeneous topography. To account for interactions between macro‐invertebrates and fungi, that potentially modify the influence of tree biodiversity and climate on dead wood decay, we compared a macro‐invertebrate exclusion treatment with a control treatment that allowed access to all types of decomposers. Diversity effects of trees on wood decay rates were mostly negative and mediated by the diversity of macro‐invertebrates. However, the effects of tree species diversity or fungal OTU richness and macro‐invertebrate diversity on wood decay rates were comparatively weak. Temperature affected decay rates positively and had the strongest influence in all treatments. While the exclusion of macro‐invertebrates did not lead to a reduction of wood decay rates, our results suggest that they may however have a mediating role in the process. In the presence of invertebrates the predictability of wood decay rates was higher and we observed a tendency of a stronger temperature control. Our results suggest that there is evidence for diversity effects on wood decomposition, but the temperature control is still more important. Thus, an increase in mean annual temperature will increase carbon and nutrient turnover through wood decomposition in subtropical forest irrespective of biotic composition.     

Fungal diversity increases soil fungistasis and resistance to microbial invasion by a non resident species

Biodiversity decline is a major concern for ecosystem functioning. Recent research efforts have been mostly focused on terrestrial plants, while, despite their importance in both natural and artificial ecosystems, little is known about soil microbial communities. This work aims at investigating the effects of fungal species richness on soil invasion by non resident microbes. Synthetic fungal communities with a species diversity ranging from 1 to 8 were assembled in laboratory microcosms and used in three factorial experiments to assess the effect of diversity on soil fungistasis, microbial invasion of soil amended with plant litter and of plant rhizosphere. The capability of different microbes to colonize environments characterized by different resident microbial communities was measured. The number of microbial species in the microcosms positively affected soil fungistasis that was also induced more rapidly in presence of synthetic communities with more species. Moreover, the increase of resident fungal diversity dramatically reduced the invasibility of both soil and plant rhizosphere. We found lower variability of soil fungistasis and invasibility in microcosms with higher species richness of microbial communities. Our study pointed out the existence of negative relationships between fungal diversity and soil invasibility by non resident microbes. Therefore, the loss of microbial species may adversely affect ecosystem functionality under specific environmental conditions.     

Local and regional influences on patterns of parasite species richness of central European fishes

Local, regional and global influences on the patterns of parasite species richness of 39 freshwater fish species from Central Europe were investigated. Host local abundance and host occurrence were considered respectively as local and regional factors, while host geographical range in longitude and latitude was considered as a global factor. Influences of size, ecology and behavior of hosts were also included in a comparative analysis using the independent contrasts method. We considered host habitat, host diet, host shoaling behavior and mobility. We found a positive relationship between local occurrence of fish and global range of their distribution. We confirmed previous findings showing the importance of host behavior and ecology on the variability of parasite species richness. Second, we showed how a global pattern, such as host geographical range, may affect the variability in parasite species richness through its effects on local abundance and distribution of hosts. A negative relationship between endoparasite species richness and host longitudinal range was found. This suggests that fish with eastern distribution live in the boundary of their distribution in Central Europe far from their center of distribution, which should also be characterized by a higher diversity of parasites.