Trait evolution, community assembly, and the phylogenetic structure of ecological communities

Taxa co-occurring in communities often represent a nonrandom sample, in phenotypic or phylogenetic terms, of the regional species pool. While heuristic arguments have identified processes that create community phylogenetic patterns, further progress hinges on a more comprehensive understanding of the interactions between underlying ecological and evolutionary processes. We created a simulation framework to model trait evolution, assemble communities (via competition, habitat filtering, or neutral assembly), and test the phylogenetic pattern of the resulting communities. We found that phylogenetic community structure is greatest when traits are highly conserved and when multiple traits influence species membership in communities. Habitat filtering produces stronger phylogenetic structure when taxa with derived (as opposed to ancestral) traits are favored in the community. Nearest-relative tests have greater power to detect patterns due to competition, while total community relatedness tests perform better with habitat filtering. The size of the local community relative to the regional pool strongly influences statistical power; in general, power increases with larger pool sizes for communities created by filtering but decreases for communities created by competition. Our results deepen our understanding of processes that contribute to phylogenetic community structure and provide guidance for the design and interpretation of empirical research.     

Neutral processes and species sorting in benthic microalgal community assembly: effects of tidal resuspension

Benthic microalgae (BMA) provide vital food resources for heterotrophs and stabilize sediments with their extracellular secretions. A central goal in ecology is to understand how processes such as species interactions and dispersal, contribute to observed patterns of species abundance and distribution. Our objectives were to assess the effects of sediment resuspension on microalgal community structure. We tested whether taxa‐abundance distributions could be predicted using neutral community models (NCMs) and also specific hypotheses about passive migration: (i) As migration decreases in sediment patches, BMA α‐diversity will decrease, and (ii) As migration decreases, BMA community dissimilarity (β‐diversity) will increase. Co‐occurrence indices (checkerboard score and variance ratio) were also computed to test for deterministic factors, such as competition and niche differentiation, in shaping communities. Two intertidal sites (mudflat and sand bar) differing in resuspension regime were sampled throughout the tidal cycle. Fluorometry and denaturing gradient gel electrophoresis were utilized to investigate diatom community structure. Observed taxa‐abundances fit those predicted from NCMs reasonably well (R² of 0.68–0.93), although comparisons of observed local communities to artificial randomly assembled communities rejected the null hypothesis that diatom communities were assembled solely by stochastic processes. No co‐occurrence tests indicated a significant role for competitive exclusion or niche partitioning in microalgal community assembly. In general, predictions about relationships between migration and species diversity were supported for local community dynamics. BMA at low tide (lowest migration) exhibited reduced α‐diversity as compared to periods of immersion at both mudflat and sand bar sites. β‐diversity was higher during low tide emersion on the mudflat, but did not differ temporally at the sand bar site. In between‐site metacommunity comparisons, low‐ and high‐resuspension sites exhibited distinct community compositions while the low‐energy mudflats contained higher microalgal biomass and greater α‐diversity. To our knowledge this is the first study to test the relevance of neutral processes in structuring marine microalgal communities. Our results demonstrate a prominent role for stochastic factors in structuring local BMA community assembly, although unidentified nonrandom processes also appear to play some role. High passive migration, in particular, appears to help maintain species diversity and structure communities in both sand and muddy habitats.     

Biogeographical patterns of abundant and rare bacterial biospheres in paddy soils across East Asia

Soil bacterial communities play fundamental roles in ecosystem functioning and often display a skewed distribution of abundant and rare taxa. So far, relatively little is known about the biogeographical patterns and mechanisms structuring the assembly of abundant and rare biospheres of soil bacterial communities. Here, we studied the geographical distribution of different bacterial sub-communities by examining the relative influence of environmental selection and dispersal limitation on taxa distributions in paddy soils across East Asia. Our results indicated that the geographical patterns of four different bacterial sub-communities consistently displayed significant distance–decay relationships (DDRs). In addition, we found niche breadth and dispersal rates to significantly explain differences in community assembly of abundant and rare taxa, directly affecting the strength of DDRs. While conditionally rare and abundant taxa displayed the strongest DDR due to higher environmental filtering and dispersal limitation, moderate taxa sub-communities had the weakest DDR due to greater environmental tolerance and dispersal rate. Random forest models indicated that soil pH (9.13%–49.78%) and average annual air temperature (16.59%–46.49%) were the most important predictors of the variation in the bacterial community. This study advances our understanding of the intrinsic links between fundamental ecological processes and microbial biogeographical patterns in paddy soils.     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Arbuscular mycorrhizal fungal communities in plant roots are not random assemblages

We investigated whether arbuscular mycorrhizal fungal (AMF) communities in plant roots are random subsets of the local taxon pool or whether they reflect the action of certain community assembly rules. We studied AMF small subunit rRNA gene sequence groups in the roots of plant individuals belonging to 11 temperate forest understorey species. Empirical data were compared with null models assuming random association. Distinct fungal species pools were present in young and old successional forest. In both forest types, the richness of plant-AMF associations was lower than expected by chance, indicating a degree of partner selectivity. AMF communities were generally not characteristic of individual plant species, but those associated with ecological groups of plant species - habitat generalists and forest specialists - were nonrandom subsets of the available pool of fungal taxa and differed significantly from each other. Moreover, these AMF communities were the least distinctive in spring, but developed later in the season. Comparison with a global database showed that generalist plants tend to associate with generalist AMF. Thus, the habitat range of the host and a possible interaction with season played a role in the assembly of AMF communities in individual plant root systems.     

Convergence of fish communities from the littoral zone of reservoirs

1. Understanding factors that regulate the assembly of communities is a main focus of ecology. Human‐engineered habitats, such as reservoirs, may provide insight into these assembly processes because they represent novel habitats that are subjected to colonization by fishes from the surrounding river basin or transported by humans. By contrasting community similarity within and among reservoirs from different drainage basins to nearby stream communities, we can test the relative constraints of reservoir habitats and regional species pools in determining species composition of reservoirs. 2. We used a large spatial database that included intensive collections from 143 stream and 28 reservoir sites within three major river basins in the Great Plains, U.S.A., to compare patterns of species diversity and community structure between streams and reservoirs and to characterize variation in fish community structure within and among major drainage basins. We expected reservoir fish faunas to reflect the regional species pool, but would be more homogeneous that stream communities because similar species are stocked and thrive in reservoirs (e.g. planktivores and piscivores), and they lack obligate stream organisms that are not shared among regional species pools. 3. We found that fish communities from reservoirs were a subset of fishes collected from streams and dominant taxa had ecological traits that would be favoured in lentic environments. Although there were regional differences in reservoir fish communities, species richness, patterns of rank abundance and community structure in reservoir communities were more homogonous across three major drainage basins than for stream communities. 4. The general pattern of convergence of reservoir fish community structure suggests their assembly is constrained by local factors such as habitat and biotic interactions, and facilitated by the introduction of species among basins. Because there is a reciprocal transfer of biota between reservoirs and streams, understanding factors structuring both habitats is necessary to evaluate the long‐term dynamics of impounded river networks.     

Rhizosphere shapes the associations between protistan predators and bacteria within microbiomes through the deterministic selection on bacterial communities

The assembly of bacterial communities in the rhizosphere is well-documented and plays a crucial role in supporting plant performance. However, we have limited knowledge of how plant rhizosphere determines the assembly of protistan predators and whether the potential associations between protistan predators and bacterial communities shift due to rhizosphere selection. To address this, we examined bacterial and protistan taxa from 443 agricultural soil samples including bulk and rhizosphere soils. Our results presented distinct patterns of bacteria and protistan predators in rhizosphere microbiome assembly. Community assembly of protistan predators was determined by a stochastic process in the rhizosphere and the diversity of protistan predators was reduced in the rhizosphere compared to bulk soils, these may be attributed to the indirect impacts from the altered bacterial communities that showed deterministic process assembly in the rhizosphere. Interestingly, we observed that the plant rhizosphere facilitates more close interrelationships between protistan predators and bacterial communities, which might promote a healthy rhizosphere microbial community for plant growth. Overall, our findings indicate that the potential predator–prey relationships within the microbiome, mediated by plant rhizosphere, might contribute to plant performance in agricultural ecosystems.     

Effects of pulse versus steady recruitment on sessile marine communities

Variation in patterns of propagule establishment (recruitment) has important effects on population dynamics and the structure of some communities. Most experimental studies have varied recruitment by changing the nature of a single event early in community development, but recruitment can also vary from steady rates of arrival to highly episodic 'pulse' events, causing differences in the temporal spacing of individuals recruiting into patches. We examined whether two different temporal patterns of recruitment of sessile invertebrates affected temperate marine communities in southeastern Australia in two experiments that were run at different times at the same site and that manipulated several different species. Target species entered communities as either a single pulse of recruits within a 2-week period or steady input of the same total number of recruits over a longer time period (5-6 weeks). The pattern of recruitment had variable effects on community structure. The colonial ascidian Botryllus schlosseri did not have a strong influence on community structure whether it recruited in a single pulse or steadily. The cover of B. schlosseri was higher when recruitment occurred as a single pulse. In a second experiment, botryllid ascidians caused changes in the composition of communities when they recruited steadily compared to when they did not recruit or didemnids recruited, but caused no differences in communities when they recruited in a shorter pulse. In contrast, recruitment frequency of didemnid ascidians had little effect, though their presence/absence caused community differences. Though we found that different temporal recruitment patterns can alter community composition, the life history and ecology of particular taxa as well as differences in environmental background processes are likely to influence the strength of these effects.     

Environmental filtering affects soil fungal community composition more than dispersal limitation at regional scales

The relative importance of dispersal limitation versus environmental filtering for community assembly has received much attention for macroorganisms. These processes have only recently been examined in microbial communities. Instead, microbial dispersal has mostly been measured as community composition change over space (i.e., distance decay). Here we directly examined fungal composition in airborne wind currents and soil fungal communities across a 40 000 km² regional landscape to determine if dispersal limitation or abiotic factors were structuring soil fungal communities. Over this landscape, neither airborne nor soil fungal communities exhibited compositional differences due to geographic distance. Airborne fungal communities shifted temporally while soil fungal communities were correlated with abiotic parameters. These patterns suggest that environmental filtering may have the largest influence on fungal regional community assembly in soils, especially for aerially dispersed fungal taxa. Furthermore, we found evidence that dispersal of fungal spores differs between fungal taxa and can be both a stochastic and deterministic process. The spatial range of soil fungal taxa was correlated with their average regional abundance across all sites, which may imply stochastic dispersal mechanisms. Nevertheless, spore volume was also negatively correlated with spatial range for some species. Smaller volume spores may be adapted to long-range dispersal, or establishment, suggesting that deterministic fungal traits may also influence fungal distributions. Fungal life-history traits may influence their distributions as well. Hypogeous fungal taxa exhibited high local abundance, but small spatial ranges, while epigeous fungal taxa had lower local abundance, but larger spatial ranges. This study is the first, to our knowledge, to directly sample air dispersal and soil fungal communities simultaneously across a regional landscape. We provide some of the first evidence that soil fungal communities are mostly assembled through environmental filtering and experience little dispersal limitation.     

Co-occurrence of ectoparasites of marine fishes: a null model analysis

We used null model analysis to test for nonrandomness in the structure of metazoan ectoparasite communities of 45 species of marine fish. Host species consistently supported fewer parasite species combinations than expected by chance, even in analyses that incorporated empty sites. However, for most analyses, the null hypothesis was not rejected, and co-occurrence patterns could not be distinguished from those that might arise by random colonization and extinction. We compared our results to analyses of presence–absence matrices for vertebrate taxa, and found support for the hypothesis that there is an ecological continuum of community organization. Presence–absence matrices for small-bodied taxa with low vagility and/or small populations (marine ectoparasites, herps) were mostly random, whereas presence–absence matrices for large-bodied taxa with high vagility and/or large populations (birds, mammals) were highly structured. Metazoan ectoparasites of marine fishes fall near the low end of this continuum, with little evidence for nonrandom species co-occurrence patterns.     

Nearly a decade‐long repeatable seasonal diversity patterns of bacterioplankton communities in the eutrophic Lake Donghu (Wuhan,China)

Uncovering which environmental factors govern community diversity patterns and how ecological processes drive community turnover are key questions related to understand the community assembly. However, the ecological mechanisms regulating long‐term variations of bacterioplankton communities in lake ecosystems remain poorly understood. Here we present nearly a decade‐long study of bacterioplankton communities from the eutrophic Lake Donghu (Wuhan, China) using 16S rRNA gene amplicon sequencing with MiSeq platform. We found strong repeatable seasonal diversity patterns in terms of both common (detected in more than 50% samples) and dominant (relative abundance >1%) bacterial taxa turnover. Moreover, community composition tracked the seasonal temperature gradient, indicating that temperature is a key environmental factor controlling observed diversity patterns. Total phosphorus also contributed significantly to the seasonal shifts in bacterioplankton composition. However, any spatial pattern of bacterioplankton communities across the main lake areas within season was overwhelmed by their temporal variabilities. Phylogenetic analysis further indicated that 75%–82% of community turnover was governed by homogeneous selection due to consistent environmental conditions within seasons, suggesting that the microbial communities in Lake Donghu are mainly controlled by niche‐based processes. Therefore, dominant niches available within seasons might be occupied by similar combinations of bacterial taxa with modest dispersal rates throughout different lake areas.     

Distinct successional patterns and processes of free-living and particle-attached bacterial communities throughout a phytoplankton bloom

1. Understanding the successional patterns of microbial communities during a phytoplankton bloom is crucial for predicting the compositional and functional stability of lake ecosystems in response to the disturbance of a bloom. Previous studies on bacterial communities associated with blooms have rarely studied the dynamics of these communities. The successional patterns of bacterial communities within different micro-habitats (i.e. water column versus particles) and mechanisms that shape these communities that differ in composition and structure remain unclear.
2. We selected a eutrophic urban lake to investigate the succession of bacterial communities during a bloom. We divided the bacterial communities into free-living (FL) and particle-attached (PA) groups based on their different lifestyles. The amplicon-based 16S rRNA gene high-throughput sequencing technology was used to obtain bacterial community composition and phylogenetic structure.
3. Our study showed distinct successional patterns between FL and PA bacterial communities, and the two bacterial lifestyles showed different responses and resilience to the bloom, in terms of diversity and relative abundance of bacterial taxa. Alpha-diversity of the PA bacterial community decreased during the bloom, whereas that of the FL bacterial community increased. More taxa in the FL bacterial community showed resilience after the disturbance than in the PA bacterial community.
4. The influence of phytoplankton blooms on the assembly of the bacterial community can be viewed as niche selection that led to the decrease in the relative importance of stochastic processes in shaping both FL and PA bacterial communities. This study shows the ecological significance of the bacterial community response to bloom events in lakes. It also shows that assembly processes differ for bacterial communities that have different lifestyles in lake ecosystems disturbed by phytoplankton blooms.

     

Effects of taxon sampling and tree reconstruction methods on phylodiversity metrics

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放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

Macrosystem community change in lake phytoplankton and its implications for diversity and function

Aim

We use lake phytoplankton community data to quantify the spatio-temporal and scale-dependent impacts of eutrophication, land-use and climate change on species niches and community assembly processes while accounting for species traits and phylogenetic constraints.

Location

Finland.

Time period

1977–2017.

Major taxa

Phytoplankton.

Methods

We use hierarchical modelling of species communities (HMSC) to model metacommunity trajectories at 853 lakes over four decades of environmental change, including a hierarchical spatial structure to account for scale-dependent processes. Using a “region of common profile” approach, we evaluate compositional changes of species communities and trait profiles and investigate their temporal development.

Results

We demonstrate the emergence of novel and widespread community composition clusters in previously more compositionally homogeneous communities, with cluster-specific community trait profiles, indicating functional differences. A strong phylogenetic signal of species responses to the environment implies similar responses among closely related taxa. Community cluster-specific species prevalence indicates lower taxonomic dispersion within the current dominant clusters compared with the historically dominant cluster and an overall higher prevalence of smaller species sizes within communities. Our findings denote profound spatio-temporal structuring of species co-occurrence patterns and highlight functional differences of lake phytoplankton communities.

Main conclusions

Diverging community trajectories have led to a nationwide reshuffling of lake phytoplankton communities. At regional and national scales, lakes are not single entities but metacommunity hubs in an interconnected waterscape. The assembly mechanisms of phytoplankton communities are strongly structured by spatio-temporal dynamics, which have led to novel community types, but only a minor part of this reshuffling could be linked to temporal environmental change.     

Abundant and rare picoeukaryotic sub‐communities present contrasting patterns in the epipelagic waters of marginal seas in the northwestern Pacific Ocean

In this work, they compared patterns of abundant and rare picoeukaryotic sub‐communities in the epipelagic waters (surface and 40–75 m depth subsurface layers) of the East and South China Seas across seasons via 454 pyrosequencing of the V4 region of 18S rDNA. They also examined the relative effects of environmental filtering, dispersal limitations and seasonality on community assembly. Their results indicated that (i) in the surface layer, abundant taxa are primarily influenced by dispersal limitations and rare taxa are primarily influenced by environmental filtering, whereas (ii) in the subsurface layer, both abundant and rare sub‐communities are only weakly influenced by environmental filtering but are strongly influenced by dispersal limitations. Moreover, (iii) abundant taxa exhibit stronger temporal variability than rare taxa. They also found that abundant and rare sub‐communities display similar spatial richness patterns that are negatively correlated with latitude and chlorophyll a and positively correlated with temperature. In summary, environmental filtering and dispersal limitations have different effects on abundant and rare picoeukaryotic sub‐communities in different layers. Thus, depth appears as an essential variable that governs the structuring patterns of picoeukaryotic communities in the oceans and should be thoroughly considered to develop a more comprehensive understanding of oceanic microbial assemblages.     

Linking community assembly and structure across scales in a wild mouse parasite community

Understanding what processes drive community structure is fundamental to ecology. Many wild animals are simultaneously infected by multiple parasite species, so host–parasite communities can be valuable tools for investigating connections between community structures at multiple scales, as each host can be considered a replicate parasite community. Like free‐living communities, within‐host–parasite communities are hierarchical; ecological interactions between hosts and parasites can occur at multiple scales (e.g., host community, host population, parasite community within the host), therefore, both extrinsic and intrinsic processes can determine parasite community structure. We combine analyses of community structure and assembly at both the host population and individual scales using extensive datasets on wild wood mice (Apodemus sylvaticus) and their parasite community. An analysis of parasite community nestedness at the host population scale provided predictions about the order of infection at the individual scale, which were then tested using parasite community assembly data from individual hosts from the same populations. Nestedness analyses revealed parasite communities were significantly more structured than random. However, observed nestedness did not differ from null models in which parasite species abundance was kept constant. We did not find consistency between observed community structure at the host population scale and within‐host order of infection. Multi‐state Markov models of parasite community assembly showed that a host's likelihood of infection with one parasite did not consistently follow previous infection by a different parasite species, suggesting there is not a deterministic order of infection among the species we investigated in wild wood mice. Our results demonstrate that patterns at one scale (i.e., host population) do not reliably predict processes at another scale (i.e., individual host), and that neutral or stochastic processes may be driving the patterns of nestedness observed in these communities. We suggest that experimental approaches that manipulate parasite communities are needed to better link processes at multiple ecological scales.     

Eutrophication and climate warming alter spatial (depth) co-occurrence patterns of lake phytoplankton assemblages

The composition and dynamics of plankton communities are critically affected by human-induced environmental changes. We analysed 33 years of phytoplankton monthly data collected in Lake Zurich (Switzerland), assigning organisms (genus level) to taxonomic groups (class, family), Reynolds associations and size categories. The aim was to understand how eutrophication and climate change have influenced taxa co-occurrence patterns within and between groups over the lake water column (14 depths, 0–135 m), using null-models to test for non-random spatial (depth) assembly. We found that the whole community showed high taxa co-occurrence levels, significantly deviating over time from random assembly concurrently with lake warming and reduced nutrient loading. This pattern was driven mostly by the depth structure of metalimnetic assemblages during summer and autumn. The prevalence of non-random spatial patterns changed for different taxonomic and functional groups, with only few significant deviations from null-model expectations. Within taxonomic and functional groups (particularly Classes and size categories), the frequency of spatial overdispersion of taxa decreased over time while the frequency of clustering increased. Our data suggest that the relative importance of mechanisms determining phytoplankton metacommunity dynamics have changed along with environmental gradients shaping water column structure.

     

Biogeography at the limits of life: Do extremophilic microbial communities show biogeographical regionalization?

Aim

Biogeographical regions are the fundamental geographical units for grouping Earth's biodiversity. Biogeographical regionalization has been demonstrated for many higher taxa, such as terrestrial plants and vertebrates, but not in microbial communities. Therefore, we sought to test empirically whether microbial communities, or taxa, show patterns consistent with biogeographical regionalization.

Location

Within halite (NaCl) crystals from coastal solar salterns of western Europe, the Mediterranean and east Africa.

Time period

Modern (2006–2013).

Major taxa studied

Archaea.

Methods

Using high‐throughput Illumina amplicon sequencing, we generated the most high‐resolution characterization of halite‐associated archaeal communities to date, using samples from 17 locations. We grouped communities into biogeographical clusters based on community turnover to test whether these communities show biogeographical regionalization. To examine whether individual taxa, rather than communities, show biogeographical patterns, we also tested whether the relative abundance of individual genera may be indicative of a community's biogeographical origins using machine learning methods, specifically random forest classification.

Results

We found that the rate of community turnover was greatest over subregional spatial scales (< 500 km), whereas at regional spatial scales the turnover was independent of geographical distance. Biogeographical clusters of communities were either not statistically robust or lacked spatial coherence, inconsistent with biogeographical regionalization. However, we identified several archaeal genera that were good indicators of biogeographical origin, providing classification error rates of < 10%.

Main conclusions

Overall, our results provide little support for the concept of biogeographical regions in these extremophilic microbial communities, despite the fact that some taxa do show biogeographical patterns. We suggest that variable dispersal ability among the halite‐associated Archaea may disrupt biogeographical patterns at the community level, preventing the formation of biogeographical regions. This means that the processes that lead to the formation of biogeographical regions operate differentially on individual microbial taxa rather than on entire communities.