Nestedness and beta diversity of gastrointestinal helminth communities in common warthogs, Phacochoerus africanus (Suidae), at 2 localities in South Africa.

Few studies have investigated the ecological interactions between wild species of Suidae and their parasites, leaving our knowledge concerning this host­parasite system fragmented. In the present study, we applied network studies to analyse community nestedness in helminth assemblages of common warthogs, Phacochoerus africanus (Gmelin) (Suidae). Helminth data were compiled from 95 warthogs, including young and adult males and females, from 2 different conservation areas in Mpumalanga and Limpopo Provinces, South Africa, collected monthly over a period of 1 year each. The aim was to study the effect of host sex, age and season of sampling on the structure of helminth infracommunities harboured by the warthogs and to search for non-random structural patterns in the warthog­helminth interaction networks. Furthermore, we investigated the influence of a warthog's age, sex and season of sampling on beta diversity and dark diversity of their helminth infracommunities. Lastly, we asked whether the effects of host sex, age and sampling season on helminth communities differed between the 2 localities. We found that helminth communities of warthogs were nested and host­parasite interactions were influenced by all 3 factors as well as combinations thereof. However, the resulting patterns differed at the 2 localities, indicating that local environmental processes are important drivers of community structure.

Dark host specificity in two ectoparasite taxa: repeatability, parasite traits, and environmental effects.

We applied the concept of dark diversity (species that may potentially inhabit a locality but are absent) to the host spectrum of a parasite and defined it as dark host specificity (DHS). We studied the trait-associated and geographic patterns of dark host specificity in fleas and gamasid mites parasitic on small mammals, asking the following questions: (a) Is dark host specificity repeatable across populations of the same species? (b) Is it associated with morphological and/or ecological species traits? (c) What are the factors associated with geographical variation in the DHS among populations of the same species? The DHS was repeatable within species with a large proportion of variance among samples, accounted for by differences between species. The average DHS of fleas, but not mites, was affected by parasite traits, with the DHS being higher in fleas with larger geographic ranges, higher characteristic abundance levels, and summer reproduction peaks. In the majority of ectoparasites, the regional DHS decreased with an increase in either structural or phylogenetic host specificity. The associations between the DHS and the environmental or host-associated characteristics of a region were revealed in a few species (eight of 22 fleas and three of 12 mites). The DHS decreased with (a) an increase in air temperature in two fleas, (b) a decrease in precipitation in two fleas, and (c) an increase in regional host species richness (in three fleas and three mites). Overall, our results suggest that dark host specificity in arthropod ectoparasites is a species-specific character associated, to a large extent, with the breadth of their host-related niches, while the influences of parasite traits and local environmental conditions are minor.

Specialisation and diversity of multiple trophic groups are promoted by different forest features.

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.

Historical biome distribution and recent human disturbance shape the diversity of arbuscular mycorrhizal fungi.

The availability of global microbial diversity data, collected using standardized metabarcoding techniques, makes microorganisms promising models for investigating the role of regional and local factors in driving biodiversity. Here we modelled the global diversity of symbiotic arbuscular mycorrhizal (AM) fungi using currently available data on AM fungal molecular diversity (small subunit (SSU) ribosomal RNA (rRNA) gene sequences) in field samples. To differentiate between regional and local effects, we estimated species pools (sets of potentially suitable taxa) for each site, which are expected to reflect regional processes. We then calculated community completeness, an index showing the fraction of the species pool present, which is expected to reflect local processes. We found significant spatial variation, globally in species pool size, as well as in local and dark diversity (absent members of the species pool). Species pool size was larger close to areas containing tropical grasslands during the last glacial maximum, which are possible centres of diversification. Community completeness was greater in regions of high wilderness (remoteness from human disturbance). Local diversity was correlated with wilderness and current connectivity to mountain grasslands. Applying the species pool concept to symbiotic fungi facilitated a better understanding of how biodiversity can be jointly shaped by large-scale historical processes and recent human disturbance.

Species pools, community completeness and invasion: disentangling diversity effects on the establishment of native and alien species.

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity-invasion relationships by separating environmental and biotic effects, especially if species' life-history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.

Dark diversity of flea assemblages of small mammalian hosts: effects of environment, host traits and host phylogeny.

An assemblage of species in a locality comprises two components, namely (i) species that are present (realised diversity) and (ii) species from the regional pool that may potentially inhabit this locality due to suitable ecological conditions, but that are absent (dark diversity). We investigated factors affecting the dark diversity of component communities of fleas parasitic on small mammals in the northern Palearctic at two scales. First, we considered the dark diversity of flea assemblages of the same host (for 13 host species) across regions and tested for the effects of environmental factors and the number of available host species on the dark diversity of within-region flea assemblages. Second, we considered the dark diversity of fleas across host species within a region (for 20 regions) and asked whether within-host dark diversity is associated with host phylogeny and/or traits. We found that the dark diversity of flea assemblages harboured by small mammals varied substantially (i) within the same host species across space (in 12 of 13 host species) and (ii) between host species within a region (in eight of 20 regions). The size of the dark diversity of flea assemblages of the same host across regions was generally affected by environmental factors (mainly by the amount of green vegetation), whereas the size of the dark diversity of flea assemblages of a host species within a region was affected by host traits (mainly by the degree of host sociality and the structure of its shelter and, to a lesser degree, by its geographic range size) but was not associated with host phylogenetic affinities. We conclude that application of the dark diversity concept to parasite communities across space or hosts allows a better understanding of the factors affecting the species richness and composition of these communities.

Seeing through sedimented waters: environmental DNA reduces the phantom diversity of sharks and rays in turbid marine habitats.

BACKGROUND: Sharks and rays are some of the most threatened marine taxa due to the high levels of bycatch and significant demand for meat and fin-related products in many Asian communities. At least 25% of shark and ray species are considered to be threatened with extinction. In particular, the density of reef sharks in the Pacific has declined to 3-10% of pre-human levels. Elasmobranchs are thought to be sparse in highly urbanised and turbid environments. Low visibility coupled with the highly elusive behaviour of sharks and rays pose a challenge to diversity estimation and biomonitoring efforts as sightings are limited to chance encounters or from carcasses ensnared in nets. Here we utilised an eDNA metabarcoding approach to enhance the precision of elasmobranch diversity estimates in urbanised marine environments. RESULTS: We applied eDNA metabarcoding on seawater samples to detect elasmobranch species in the hyper-urbanised waters off Singapore. Two genes-vertebrate 12S and elasmobranch COI-were targeted and amplicons subjected to Illumina high-throughput sequencing. With a total of 84 water samples collected from nine localities, we found 47 shark and ray molecular operational taxonomic units, of which 16 had species-level identities. When data were compared against historical collections and contemporary sightings, eDNA detected 14 locally known species as well as two potential new records. CONCLUSIONS: Local elasmobranch richness uncovered by eDNA is greater than the seven species sighted over the last two decades, thereby reducing phantom diversity. Our findings demonstrate that eDNA metabarcoding is effective in detecting shark and ray species despite the challenges posed by the physical environment, granting a more consistent approach to monitor these highly elusive and threatened species.

Throwing light on dark diversity of vascular plants in China: predicting the distribution of dark and threatened species under global climate change.

BACKGROUND: As global climate change accelerates, ecologists and conservationists are increasingly investigating changes in biodiversity and predicting species distribution based on species observed at sites, but rarely consider those plant species that could potentially inhabit but are absent from these areas (i.e., the dark diversity and its distribution). Here, we estimated the dark diversity of vascular plants in China and picked up threatened dark species from the result, and applied maximum entropy (MaxEnt) model to project current and future distributions of those dark species in their potential regions (those regions that have these dark species). METHODS: We used the Beals probability index to estimate dark diversity in China based on available species distribution information and explored which environmental variables had significant impacts on dark diversity by incorporating bioclimatic data into the random forest (RF) model. We collected occurrence data of threatened dark species (Eucommia ulmoides, Liriodendron chinense, Phoebe bournei, Fagus longipetiolata, Amentotaxus argotaenia, and Cathaya argyrophylla) and related bioclimatic information that can be used to predict their distributions. In addition, we used MaxEnt modeling to project their distributions in suitable areas under future (2050 and 2070) climate change scenarios. RESULTS: We found that every study region's dark diversity was lower than its observed species richness. In these areas, their numbers of dark species are ranging from 0 to 215, with a generally increasing trend from western regions to the east. RF results showed that temperature variables had a more significant effect on dark diversity than those associated with precipitation. The results of MaxEnt modeling showed that most threatened dark species were climatically suitable in their potential regions from current to 2070. DISCUSSIONS: The results of this study provide the first ever dark diversity patterns concentrated in China, even though it was estimated at the provincial scale. A combination of dark diversity and MaxEnt modeling is an effective way to shed light on the species that make up the dark diversity, such as projecting the distribution of specific dark species under global climate change. Besides, the combination of dark diversity and species distribution models (SDMs) may also be of value for ex situ conservation, ecological restoration, and species invasion prevention in the future.

Changes in potential mammal diversity in national parks and their implications for conservation.

Observed species richness (OSR) is a widely used and well-studied biodiversity metric. However, non-observed species in favorable ecosystems are also relevant. Two metrics that include observed and potential species were recently defined: potential biodiversity (hereafter potential species richness-PSR) and geometric mean of favorabilities (GMF). We used these metrics to evaluate the national park network of mainland Spain at two time periods (2002 and 2015), using terrestrial mammals on a UTM 100-km2 grid. PSR and GMF are based on the favorability function, a species distribution model that assesses how favorable an area is for the presence of a species, over and above its prevalence in the study area. For each park and for the whole network, we calculated the mean and sum of OSR, PSR, and GMF in each time period, as well as changes between periods. OSR and PSR were higher inside than outside the park network in both time periods. Thus, although the network covers a very small proportion of the country, it performs well for the representation of mammal species and their favorable areas. However, mean PSR was lower in 2015 than in 2002 inside the national park network, whereas the opposite was the case outside the network. Mountainous Parks generally not only concentrated highly favorable areas for mammals, but they also showed less favorable areas in 2015 compared to 2002, although the reduction was moderate to low. This is a result to consider for future analyses because if the tendency increases, it may have consequences for the conservation of mammals and for the adequacy of the national park network.

Predicting species establishment using absent species and functional neighborhoods.

Species establishment within a community depends on their interactions with the local environment and resident community. Such environmental and biotic filtering is frequently inferred from functional trait and phylogenetic patterns within communities; these patterns may also predict which additional species can establish. However, differentiating between environmental and biotic filtering can be challenging, which may complicate establishment predictions. Creating a habitat-specific species pool by identifying which absent species within the region can establish in the focal habitat allows us to isolate biotic filtering by modeling dissimilarity between the observed and biotically excluded species able to pass environmental filters. Similarly, modeling the dissimilarity between the habitat-specific species pool and the environmentally excluded species within the region can isolate local environmental filters. Combined, these models identify potentially successful phenotypes and why certain phenotypes were unsuccessful. Here, we present a framework that uses the functional dissimilarity among these groups in logistic models to predict establishment of additional species. This approach can use multivariate trait distances and phylogenetic information, but is most powerful when using individual traits and their interactions. It also requires an appropriate distance-based dissimilarity measure, yet the two most commonly used indices, nearest neighbor (one species) and mean pairwise (all species) distances, may inaccurately predict establishment. By iteratively increasing the number of species used to measure dissimilarity, a functional neighborhood can be chosen that maximizes the detection of underlying trait patterns. We tested this framework using two seed addition experiments in calcareous grasslands. Although the functional neighborhood size that best fits the community's trait structure depended on the type of filtering considered, selecting these functional neighborhood sizes allowed our framework to predict up to 50% of the variation in actual establishment from seed. These results indicate that the proposed framework may be a powerful tool for studying and predicting species establishment.

Large-scale dark diversity estimates: new perspectives with combined methods.

Large-scale biodiversity studies can be more informative if observed diversity in a study site is accompanied by dark diversity, the set of absent although ecologically suitable species. Dark diversity methodology is still being developed and a comparison of different approaches is needed. We used plant data at two different scales (European and seven large regions) and compared dark diversity estimates from two mathematical methods: species co-occurrence (SCO) and species distribution modeling (SDM). We used plant distribution data from the Atlas Florae Europaeae (50 × 50 km grid cells) and seven different European regions (10 × 10 km grid cells). Dark diversity was estimated by SCO and SDM for both datasets. We examined the relationship between the dark diversity sizes (type II regression) and the overlap in species composition (overlap coefficient). We tested the overlap probability according to the hypergeometric distribution. We combined the estimates of the two methods to determine consensus dark diversity and composite dark diversity. We tested whether dark diversity and completeness of site diversity (log ratio of observed and dark diversity) are related to various natural and anthropogenic factors differently than simple observed diversity. Both methods provided similar dark diversity sizes and distribution patterns; dark diversity is greater in southern Europe. The regression line, however, deviated from a 1:1 relationship. The species composition overlap of two methods was about 75%, which is much greater than expected by chance. Both consensus and composite dark diversity estimates showed similar distribution patterns. Both dark diversity and completeness measures exhibit relationships to natural and anthropogenic factors different than those exhibited by observed richness. In summary, dark diversity revealed new biodiversity patterns which were not evident when only observed diversity was examined. A new perspective in dark diversity studies can incorporate a combination of methods.

Measuring size and composition of species pools: a comparison of dark diversity estimates.

Ecological theory and biodiversity conservation have traditionally relied on the number of species recorded at a site, but it is agreed that site richness represents only a portion of the species that can inhabit particular ecological conditions, that is, the habitat-specific species pool. Knowledge of the species pool at different sites enables meaningful comparisons of biodiversity and provides insights into processes of biodiversity formation. Empirical studies, however, are limited due to conceptual and methodological difficulties in determining both the size and composition of the absent part of species pools, the so-called dark diversity. We used >50,000 vegetation plots from 18 types of habitats throughout the Czech Republic, most of which served as a training dataset and 1083 as a subset of test sites. These data were used to compare predicted results from three quantitative methods with those of previously published expert estimates based on species habitat preferences: (1) species co-occurrence based on Beals' smoothing approach; (2) species ecological requirements, with envelopes around community mean Ellenberg values; and (3) species distribution models, using species environmental niches modeled by Biomod software. Dark diversity estimates were compared at both plot and habitat levels, and each method was applied in different configurations. While there were some differences in the results obtained by different methods, particularly at the plot level, there was a clear convergence, especially at the habitat level. The better convergence at the habitat level reflects less variation in local environmental conditions, whereas variation at the plot level is an effect of each particular method. The co-occurrence agreed closest the expert estimate, followed by the method based on species ecological requirements. We conclude that several analytical methods can estimate species pools of given habitats. However, the strengths and weaknesses of different methods need attention, especially when dark diversity is estimated at the plot level.