Deconstructing latitudinal species richness patterns in the ocean: does larval development hold the clue?

The consistent decrease in species richness with latitude shows several exceptions among marine organisms. We hypothesize that contrasting latitudinal diversity gradients can be explained by differences in critical life-history attributes, such as mode of larval development (MLD). We deconstructed latitudinal species richness patterns of marine benthic invertebrates according to MLD to elucidate differences in patterns of species richness and to reveal underlying processes. The patterns of species richness were remarkably similar across taxa within MLD but differed between MLD. Species richness decreased polewards in planktotrophic species and increased in direct developers. Temperature explained most of the variation in species richness. Low temperature at high latitudes may generally favour direct developing species, but, together with low chlorophyll- a concentration, limit the distribution of planktotrophic species. The contrasting influence of temperature on different MLDs might be explained by its effect on the length of planktonic life and on brooding costs.     

An ecological perspective on bacterial biodiversity

Bacteria may be one of the most abundant and species-rich groups of organisms, and they mediate many critical ecosystem processes. Despite the ecological importance of bacteria, past practical and theoretical constraints have limited our ability to document patterns of bacterial diversity and to understand the processes that determine these patterns. However, recent advances in molecular techniques that allow more thorough detection of bacteria in nature have made it possible to examine such patterns and processes. Here, we review recent studies of the distribution of free-living bacterial diversity and compare our current understanding with what is known about patterns in plant and animal diversity. From these recent studies a preliminary picture is emerging: bacterial diversity may exhibit regular patterns, and in some cases these patterns may be qualitatively similar to those observed for plants and animals.     

Two oceans, two taxa and one mode of development: latitudinal diversity patterns of South American crabs and test for possible causal processes

The latitudinal gradient of species diversity is a widely recognized but poorly understood phenomenon. In marine systems, differences in dispersal abilities among species may pose an additional problem in identifying the processes that affect diversity. We compared latitudinal diversity gradients along two parallel continental coasts, the east and west coasts of South America, of two groups of Crustacea (Brachyura and Anomura), which exclusively exhibit planktonic development. We also evaluated the species‐area and the energy‐input hypotheses. Diversity decreased with increasing latitude for both groups in both oceans. Results suggest that the spatial structure of sea surface temperature (SST) explains diversity of both groups at large, but not small (< 5°), scales. Range size and latitude were not correlated. We hypothesize that SST differentially affects taxa with contrasting modes of development, influencing patterns of diversity. We suggest that developmental modes of marine organisms should be considered in future diversity analyses.     

Patterns of diversity in microscopic animals: are they comparable to those in protists or in larger animals?

Aim General patterns of biodiversity, such as latitudinal gradients and species‐area relationships, are found consistently in a wide range of organisms, but recent results for protist diversity suggest that organisms shorter than 2 mm do not display such patterns. We tested this prediction in bdelloid rotifers, pluricellular metazoans smaller than 2 mm, but with size and ecology comparable to protists. Location A single valley in northern Italy was surveyed in detail and compared to all available faunistic data on bdelloids worldwide. Methods We analysed 171 local assemblages of bdelloid rotifers living in 5 systems of dry mosses and submerged mosses in running water and in lakes. We compared patterns of alpha, beta, and gamma diversity, and nestedness of metacommunities, with those known from protists and larger organisms. Results Bdelloid rotifers showed low local species richness (alpha diversity), with strong habitat selection, as observed in larger organisms. The number of species differed among systems, with a higher number of species in dry than in aquatic mosses. There was no hierarchical structure or exclusion of species in the metacommunity pattern within each system. Local diversity for the entire valley was surprisingly high compared with worldwide bdelloid diversity, similar to observed patterns in protists. Main Conclusions Bdelloid rotifers have some of the peculiarities of protist biodiversity, although at slightly different spatial scales, thus confirming the idea of a major change in biodiversity patterns among organisms shorter than 2 mm. However, bdelloids show stronger habitat selection than protists. We suggest two possible explanations for the observed patterns: (1) dispersal is very rare, and not all bdelloid clones are arriving everywhere; and (2) dispersal is effective in displacing propagules, but environmental heterogeneity is very high and prevents many species from colonizing a given patch of moss.     

Genetic structure of a foundation species: scaling community phenotypes from the individual to the region

Understanding the local and regional patterns of species distributions has been a major goal of ecological and evolutionary research. The notion that these patterns can be understood through simple quantitative rules is attractive, but while numerous scaling laws exist (e.g., metabolic, fractals), we are aware of no studies that have placed individual traits and community structure together within a genetics based scaling framework. We document the potential for a genetic basis to the scaling of ecological communities, largely based upon our long-term studies of poplars (Populus spp.). The genetic structure and diversity of these foundation species affects riparian ecosystems and determines a much larger community of dependent organisms. Three examples illustrate these ideas. First, there is a strong genetic basis to phytochemistry and tree architecture (both above- and belowground), which can affect diverse organisms and ecosystem processes. Second, empirical studies in the wild show that the local patterns of genetics based community structure scale up to western North America. At multiple spatial scales the arthropod community phenotype is related to the genetic distance among plants that these arthropods depend upon for survival. Third, we suggest that the familiar species-area curve, in which species richness is a function of area, is also a function of genetic diversity. We find that arthropod species richness is closely correlated with the genetic marker diversity and trait variance suggesting a genetic component to these curves. Finally, we discuss how genetic variation can interact with environmental variation to affect community attributes across geographic scales along with conservation implications.     

Dispersal limitations and historical factors determine the biogeography of specialized terrestrial protists

Recent studies show that soil eukaryotic diversity is immense and dominated by micro‐organisms. However, it is unclear to what extent the processes that shape the distribution of diversity in plants and animals also apply to micro‐organisms. Major diversification events in multicellular organisms have often been attributed to long‐term climatic and geological processes, but the impact of such processes on protist diversity has received much less attention as their distribution has often been believed to be largely cosmopolitan. Here, we quantified phylogeographical patterns in Hyalosphenia papilio, a large testate amoeba restricted to Holarctic Sphagnum‐dominated peatlands, to test if the current distribution of its genetic diversity can be explained by historical factors or by the current distribution of suitable habitats. Phylogenetic diversity was higher in Western North America, corresponding to the inferred geographical origin of the H. papilio complex, and was lower in Eurasia despite extensive suitable habitats. These results suggest that patterns of phylogenetic diversity and distribution can be explained by the history of Holarctic Sphagnum peatland range expansions and contractions in response to Quaternary glaciations that promoted cladogenetic range evolution, rather than the contemporary distribution of suitable habitats. Species distributions were positively correlated with climatic niche breadth, suggesting that climatic tolerance is key to dispersal ability in H. papilio. This implies that, at least for large and specialized terrestrial micro‐organisms, propagule dispersal is slow enough that historical processes may contribute to their diversification and phylogeographical patterns and may partly explain their very high overall diversity.     

Comparing classical community models: theoretical consequences for patterns of diversity

Abstract: Mechanisms proposed to explain the maintenance of species diversity within ecological communities of sessile organisms include niche differentiation mediated by competitive trade-offs, frequency dependence resulting from species-specific pests, recruitment limitation due to local dispersal, and a speciation-extinction dynamic equilibrium mediated by stochasticity (drift). While each of these processes, and more, have been shown to act in particular communities, much remains to be learned about their relative importance in shaping community-level patterns. We used a spatially-explicit, individual-based model to assess the effects of each of these processes on species richness, relative abundance, and spatial patterns such as the species-area curve. Our model communities had an order-of-magnitude more individuals than any previous such study, and we also developed a finite-size scaling analysis to infer the large-scale properties of these systems in order to establish the generality of our conclusions across system sizes. As expected, each mechanism can promote diversity. We found some qualitative differences in community patterns across communities in which different combinations of these mechanisms operate. Species-area curves follow a power law with short-range dispersal and a logarithmic law with global dispersal. Relative-abundance distributions are more even for systems with competitive differences and trade-offs than for those in which all species are competitively equivalent, and they are most even when frequency dependence (even if weak) is present. Overall, however, communities in which different processes operated showed surprisingly similar patterns, which suggests that the form of community-level patterns cannot in general be used to distinguish among mechanisms maintaining diversity there. Nevertheless, parameterization of models such as these from field data on the strengths of the different mechanisms could yield insight into their relative roles in diversity maintenance in any given community.     

Thoughts on the processes that maintain local species diversity of ectomycorrhizal fungi

Ectomycorrhizal fungi exhibit high diversity even in small monoculture forests. Roughly 20 to 35 species typically occupy such sites. Explanations for this diversity can be based on resource partitioning, disturbance, competition, or interaction with other organisms. Mycorrhizal fungi compete for two general classes of resources: host-derived carbon and soil or detritus derived mineral nutrients. Both types of resources are arrayed in space (e.g., soil depth, distance from tree) and time (e.g., season, host successional series). Some species seem to be partitioned in space and time at these scales, but the question of how widespread these patterns are remains largely unanswered. Mineral resources are distributed in discrete substrates in soil, litter, and within other soil microorganisms; the biochemical diversity exhibited by fungi may translate into differences in access to these resources among species. Small-scale natural disturbances that sever roots, mix soil horizons and litter layers, or change local pH and nutrient availability, are likely to create additional habitats for ectomycorrhizal fungi. Evidence from fruiting patterns and differences in colonization strategies suggest that such disturbances may be important for establishment of some species. Competitive replacement networks among species have the theoretical potential to increase diversity. The frequency of species replacements, observed co-infections of ectomycorrhizal fungi on single host roots, and high rates of rootlet turn-over all suggest that competition is important, but whether it plays a creative role in maintaining diversity remains to be demonstrated. Other organisms could be important in the maintenance of diversity, if they effect competition among mycorrhizal fungi. Bacteria and soil invertebrates are the most likely groups for such interactions. Technological advances in root observation and PCR methods for indentification of mycorrhizae make many of these theories testable.     

Comparative taxonomic structure of the floras of two Mediterranean-climate regions: Iberia and California

Two major events were invoked to understand recent biodiversity patterns in Mediterranean floras: northern hemisphere glaciations and historical human impacts. These two events were considered in this work, where we investigated general patterns in plant species richness and rarity attributes in two different Mediterranean regions: California and Iberia. Our goal was to assess whether comparisons of this sort provided evidence of different extinctions rates, making an effort to decouple anthropogenic from ice age‐related effects in both regions. We employed a taxonomically revised database for eight Mediterranean floras containing information on species richness for 298 families and rarity attributes for 11,834 taxa. We used summary statistics (Gini coefficient) and randomly generated models to test for general patterns of the distribution of diversity within and among taxonomic groups. We then used this general pattern among Mediterranean floras to provide a context in which to evaluate our two focal areas. Results indicated that floras of California and Iberia share the closest taxonomic structure among Mediterranean regions. Differences emerged in rarity attributes and the taxonomic identities of rarity rich groups. These findings were interpreted in the light of Pleistocene changes. In addition, a closer focus on rarity attributes allowed us to pinpoint some segments of these floras where anthropogenic activities may drive variation from general patterns, specifically for rare species in ecologically sensitive habitats.     

Evolving Concepts of Bacterial Species

The same evolutionary forces that cause diversification in sexual eukaryotes are expected to cause diversification in bacteria. However, in bacteria, the wider variety of mechanisms for gene exchange (or lack thereof) increases the range of expected diversity patterns compared to those of sexual organisms. Two parallel concepts for bacterial speciation have developed, based on ecological divergence or barriers to recombination in turn. Recent evidence from DNA sequence data shows that both processes can generate independently evolving groups that are equivalent to sexual species and that represent separate arenas within which recombination (when it occurs), selection and drift occur. It remains unclear, however, how often different processes act in concert to generate simple units of diversity, or whether a more complex model of diversity is required, specifying hierarchical levels at which different cohesive processes operate. We advocate an integrative approach that evaluates the effects of multiple evolutionary forces on diversity patterns. There is also great potential for laboratory studies of bacterial evolution that test evolutionary mechanisms inferred from population genetic analyses of multi-locus and genome sequence data.     

Phorophyte size and soil profiles differentially correlate with community structure among hemiepiphytes and nomadic vines

Tropical non-self-supporting plants such as hemiepiphytes and nomadic vines are model organisms for disentangling biotic and environmental correlates which influence their occupancy patterns. We inventoried >4000 individuals from >3000 trees ranging from 1 to 200 cm diameter at breast height (DBH) in a northeastern Amazonian upland forest to address how tree (phorophyte) size, edaphic factors and recruitment strategy influence occupancy, diversity, and compositional patterns of two vascular non-self-supporting plant functional groups. Hemiepiphytes germinate on phorophytes prior to establishing soil connections, whereas nomadic vines initiate their life cycle on the forest floor and subsequently climb phorophytes for crown access, abandoning roots replaced by adventitious connections which may reach the ground. Our results show that larger phorophytes (≥30 cm DBH) supported more species for both hemiepiphytes and nomadic vines. However, nomadic vines' occupancy probabilities saturated faster at smaller stem sizes than that of hemiepiphytes indicating differential preferences for stem sizes among the two functional groups. For smaller phorophytes (<30 cm DBH), soil correlations were stronger with nomadic vines than hemiepiphytes, whereas no significant differences were detected among functional groups in relation to edaphic factors for larger (≥ 30 cm DBH) ones. Finally, a small core group of species showed disproportionately greater abundances among large phorophytes suggesting that autogenic processes differentially promote survivability. Such interactions among phorophyte size and edaphic factors may result from the contrasting ecological requirements of hemiepiphytes and nomadic vines at the recruitment stage, demonstrating the necessity for elaborate demographic-based studies to better understand these complex plant–plant interactions. Abstract in Spanish is available with online material     

Biological and ecological traits of benthic freshwater macroinvertebrates: relationships and definition of groups with similar traits

• 1 Relating species traits to habitat characteristics can provide important insights into the structure and functioning of stream communities. However, trade‐offs among species traits make it difficult to predict accurately the functional diversity of freshwater communities. Many authors have pointed to the value of working with groups of organisms as similar as possible in terms of relationships among traits and have called for definition of groups of organisms with similar suites of attributes.
• 2 We used multivariate analyses to examine separately the relationships among 11 biological traits and among 11 ecological traits of 472 benthic macroinvertebrate taxa (mainly genera). The main objective was to demonstrate (1) potential trade‐offs among traits; (2) the importance of the different traits to separate systematic units or functional groupings; and (3) uniform functional groups of taxa that should allow a more effective use of macroinvertebrate biological and ecological traits.
• 3 We defined eight groups and 15 subgroups according to a biological trait ordination which highlighted size (large to small), reproductive traits (K to r strategists), food (animal to plant material) and feeding habits (predator to scraper and/or deposit feeder) as ‘significant’ factors determining the ordination of taxa. This ordination partly preserved phylogenetic relationships among groups.
• 4 Seven ecological groups and 13 ecological subgroups included organisms with combinations of traits which should be successively more adequate in habitats from the main channel to temporary waters, and from the crenon to the potamic sections of rivers, and to systems situated outside the river floodplain. These gradients corresponded to a gradual shift from (1) rheophilic organisms that lived in the main channel of cold oligotrophic mountain streams to (2) animals that preferred eutrophic habitats of still or temporary waters in lowlands. The groups with similar ecological traits had a more diverse systematic structure than those with similar biological traits.
• 5 Monitoring and assessment tools for the management of water resources are generally more effective if they are based on a clear understanding of the mechanisms that lead to the presence or absence of species groups in the environment. We believe that groups with similar relationships among their species traits may be useful in developing tools that measure the functional diversity of communities.

     

Mechanisms underlying the bioindicator notion: spatial association between individual sexual performance and community diversity

The bioindicator notion is an appealing concept that has received more support in applied than in basic ecology, mostly due to the difficulty in deriving general ecological rules applicable to all target organisms. However, recognizing the mechanisms that determine the association between a particular species and the well-being of many other species is important for understanding the functioning of ecosystems and the relationship among different biological levels. We examined here the processes at the individual level that cause an association between species performance and biodiversity value, by analyzing attributes that can be studied in a variety of animals with sexual reproduction, namely breeding site selection and condition-dependent sexual signals. Our study model was the Capercaillie, an indicator of forest functioning and diversity, and the associated bird community, used here as a surrogate of broader forest biodiversity. At a regional scale Capercaillie occurrence was not associated with the most diverse forest patches, but at the scale of male spring territories the sexual display grounds (arenas) were located in the oldest and less disturbed forest portions, which also hosted the richest local bird communities. Social mechanisms and conspecific cueing likely concurred with habitat-driven processes in determining the long-term persistence of traditional display grounds, which were appealing to many other species because of their structural composition. Characteristics of male vocal display that honestly advertize male quality (low frequencies and rapid song rates) were significantly correlated with high diversity values, resulting in a spatial association between individual and community performances. Costly or risky activities such as reproductive or social behaviors, which more than other attributes match gradients in habitat quality, are therefore contributing to functionally connect individuals with ecosystem health.     

Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time

A key challenge in ecological research is to integrate data from different scales to evaluate the ecological and evolutionary mechanisms that influence current patterns of biological diversity. We build on recent attempts to incorporate phylogenetic information into traditional diversity analyses and on existing research on beta diversity and phylogenetic community ecology. Phylogenetic beta diversity (phylobetadiversity) measures the phylogenetic distance among communities and as such allows us to connect local processes, such as biotic interactions and environmental filtering, with more regional processes including trait evolution and speciation. When combined with traditional measures of beta diversity, environmental gradient analyses or ecological niche modelling, phylobetadiversity can provide significant and novel insights into the mechanisms underlying current patterns of biological diversity.     

The Linnean shortfall in oceanic island biogeography: a case study in the Azores

Aim Speciation processes on islands are still poorly understood. Previous studies based on the analysis of distribution data from checklists found that the flora of the Azores archipelago differs from other island floras in the exceptionally low number of radiations and the low number of single‐island endemics. The general mechanism(s) responsible for these apparently unique patterns remained unclear. One possible explanation for the distinctiveness of the Azorean endemic flora is the lack of a consistent and critical taxonomic framework for the floras of the Atlantic archipelagos. In this study, molecular variation within a range of Azorean endemic plant lineages was analysed to determine whether inadequacies in the current taxonomy of endemics might be an explanation for the unusual diversity patterns observed in the endemic flora of the Azores. Location Azores archipelago. Method Sixty‐nine populations of eight endemic species or subspecies belonging to five genetic lineages were sampled from all Azorean islands but one. Nuclear and plastid DNA regions were sequenced, and relationships among internal transcribed spacer (ITS) region ribotypes established using statistical parsimony. Results Molecular diversity patterns differ from current taxonomic groupings, with all lineages comprising previously overlooked genetic entities. Main conclusions Recognition as distinct taxa of the genetically distinct entities discovered in this study would drastically change the diversity patterns and make them more similar to those of other Atlantic archipelagos. The results serve to highlight that current knowledge of endemic diversity on oceanic islands may be far from complete, even in relatively well‐known groups such as angiosperms. This limitation is rarely considered in macroecological and evolutionary studies that make use of data from taxonomic checklists to draw inferences about oceanic island biogeographic processes.     

Clade age and species richness are decoupled across the eukaryotic tree of life

Explaining the dramatic variation in species richness across the tree of life remains a key challenge in evolutionary biology. At the largest phylogenetic scales, the extreme heterogeneity in species richness observed among different groups of organisms is almost certainly a function of many complex and interdependent factors. However, the most fundamental expectation in macroevolutionary studies is simply that species richness in extant clades should be correlated with clade age: all things being equal, older clades will have had more time for diversity to accumulate than younger clades. Here, we test the relationship between stem clade age and species richness across 1,397 major clades of multicellular eukaryotes that collectively account for more than 1.2 million described species. We find no evidence that clade age predicts species richness at this scale. We demonstrate that this decoupling of age and richness is unlikely to result from variation in net diversification rates among clades. At the largest phylogenetic scales, contemporary patterns of species richness are inconsistent with unbounded diversity increase through time. These results imply that a fundamentally different interpretative paradigm may be needed in the study of phylogenetic diversity patterns in many groups of organisms.     

Patterns of bryophyte and vascular plant richness in European subalpine springs

The diversity of spring habitats can be determined not only by local environmental conditions, but also by large-scale biogeographical effects. The effects can differ across various groups of organisms. We compared α-, β- and γ-diversity patterns of bryophytes and vascular plants of (sub)alpine springs in three contrasting mountain ranges: Alps (Switzerland), Balkans (Bulgaria), Western Carpathians (Slovakia, Poland). We used univariate and multivariate statistics to test for the effects of pH, conductivity, altitude, slope, mean annual temperature and annual precipitation on diversity patterns of both taxonomic groups and compared diversity patterns among the regions for particular pH and conductivity classes. We identified acidophyte and basiphyte, calcifuge and calcicole species using species response modelling. All regions displayed significant relationship between conductivity and α-diversity of vascular plants. Bulgaria showed the highest α-diversity of vascular plants for the middle part of the conductivity gradient. For both taxonomic groups, the β-diversity in the middle part of gradient was highest in Swiss Alps. The total species pool was lowest in Bulgaria. The percentage of basiphyte and calcicole species was highest in the Alps. In (sub)alpine springs, mineral richness was a better determinant of vascular plant α-diversity than pH, and the extent of the alpine area did not coincide with α-diversity. Observed inter-regional differences in diversity patterns could be explained by the different proportion of limestone bedrock and different biogeographic history. The differences in α-diversity between both taxonomic groups are presumably result of the different rates of adaptation processes.     

Fine‐scale Beta‐diversity Patterns Across Multiple Arthropod Taxa Over a Neotropical Latitudinal Gradient

Documenting how diversity patterns vary at fine‐ and broad scales may help answer many questions in theoretical and applied ecology. However, studies tend to compare diversity patterns at the same scale and within the same taxonomic group, which limits the applicability and generality of the results. Here, we have investigated whether vegetation‐dwelling arthropods from different trophic ranks and with distinct life histories (i.e., ants, caterpillars, cockroaches, and spiders) have different beta‐diversity patterns at multiple scales. Specifically, we compared their beta diversity across architecturally distinct plant species (fine‐scale process) and a latitudinal gradient of sites (broad‐scale process) along 2040 km of coastal restinga vegetation in the Neotropics. Over 50 percent of the compositional changes (β‐diversity) in ants, caterpillars, and spiders and 41 percent of those in cockroaches were explained by plant identity within each site. Even groups that do not feed on plant tissues, such as omnivores and predators, were strongly affected by plant identity. Fine‐scale variation was more important than large‐scale processes for all studied groups. Performing a cross‐scale comparison of diversity patterns of groups with distinct life histories helps elucidate how processes that act at regional scales, such as dispersal, interact with local processes to assemble arthropod communities.     

Body size determines the strength of the latitudinal diversity gradient

In most groups of organisms, the species richness decreases from the tropics to the poles. The mechanisms causing this latitudinal diversity gradient are still controversial. We present data from a comprehensive weighted meta-analysis on the strength of the latitudinal gradient in relation to body size. We sampled literature data on the correlation between species richness and latitude for a variety of organisms, ranging from trees to protozoa. In addition, own data on the presence of large-scale diversity patterns for diatoms were included, both for local and regional species richness. The strength of the latitudinal gradient was positively correlated to the size of the organisms. Strongest decreases of species richness to the poles was found for large organisms like trees and vertebrates, whereas meiofauna, protozoa and diatoms showed weak or no correlations between species richness and latitude. These results imply that latitudinal gradients are shaped by non-equilibrium (regional) processes and are persistent under conditions of dispersal limitation.     

Patterns of elevational beta diversity in micro‐ and macroorganisms

Aim While ecologists have long been interested in diversity in mountain regions, elevational patterns in beta diversity are still rarely studied across different life forms ranging from micro‐ to macroorganisms. Also, it is not known whether the patterns in turnover among organism groups are affected by the degree to which the environment is modified by human activities. Location Laojun Mountain, Yunnan Province, China. Methods The beta diversity patterns of benthic microorganisms (i.e. diatoms and bacteria) and macroorganisms (i.e. macroinvertebrates) in a stony stream were simultaneously investigated between elevations of 1820 and 4050 m. Data were analysed by using a distance‐based approach and variation partitioning based on canonical redundancy analysis. Results Analyses of community dissimilarities between adjacent sampling sites showed comparable small‐scale beta diversity along the elevational gradient for the organism groups. However, bacteria clearly showed the lowest elevational turnover when analyses were conducted simultaneously for all pairwise sites. Variation partitioning indicated that species turnover was mostly related to environmental heterogeneity and spatial gradients including horizontal distance and elevation, while purely human impacts were shown to be less important. Main conclusions The elevational beta diversity at large scales was lower for bacteria than for eukaryotic microorganisms or macroorganisms, perhaps indicative of high dispersal ability and good adaptability of bacteria to harsh environmental conditions. However, the small‐scale beta diversity did not differ among the groups. Elevation was the major driver for the turnover of eukaryotic organisms, while the turnover of bacteria was correlated more with environmental variation.