Environmental filtering governs consistent vertical zonation in sedimentary microbial communities across disconnected mountain lakes.

Subsurface microorganisms make up the majority of Earth's microbial biomass, but ecological processes governing surface communities may not explain community patterns at depth because of burial. Depth constrains dispersal and energy availability, and when combined with geographic isolation across landscapes, may influence community assembly. We sequenced the 16S rRNA gene of bacteria and archaea from 48 sediment cores across 36 lakes in four disconnected mountain ranges in Wyoming, USA and used null models to infer assembly processes across depth, spatial isolation, and varying environments. Although we expected strong dispersal limitations across these isolated settings, community composition was primarily shaped by environmental selection. Communities consistently shifted from domination by organisms that degrade organic matter at the surface to methanogenic, low-energy adapted taxa in deeper zones. Stochastic processes-like dispersal limitation-contributed to differences among lakes, but because these effects weakened with depth, selection processes ultimately governed subsurface microbial biogeography.

Diversity-stability relationships across organism groups and ecosystem types become decoupled across spatial scales.

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity-stability relationship [DSR]). At broader spatial extents, regional-scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long-term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ-diversity on metacommunity variability, but ß-diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ-diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial ß-diversity to reduce spatial synchrony.

Scaling up and down: movement ecology for microorganisms.

Movement is critical for the fitness of organisms, both large and small. It dictates how individuals acquire resources, evade predators, exchange genetic material, and respond to stressful environments. Movement also influences ecological and evolutionary dynamics at higher organizational levels, such as populations and communities. However, the links between individual motility and the processes that generate and maintain microbial diversity are poorly understood. Movement ecology is a framework linking the physiological and behavioral properties of individuals to movement patterns across scales of space, time, and biological organization. By synthesizing insights from cell biology, ecology, and evolution, we expand theory from movement ecology to predict the causes and consequences of microbial movements.

Seed banks alter metacommunity diversity: The interactive effects of competition, dispersal and dormancy.

Dispersal and dormancy are two common strategies allowing for species persistence and the maintenance of biodiversity in variable environments. However, theory and empirical tests of spatial diversity patterns tend to examine either mechanism in isolation. Here, we developed a stochastic, spatially explicit metacommunity model incorporating seed banks with varying germination and survival rates. We found that dormancy and dispersal had interactive, nonlinear effects on the maintenance and distribution of metacommunity diversity. Seed banks promoted local diversity when seed survival was high and maintained regional diversity through interactions with dispersal. The benefits of seed banks for regional diversity were largest when dispersal was high or intermediate, depending on whether local competition was equal or stabilising. Our study shows that classic predictions for how dispersal affects metacommunity diversity can be strongly influenced by dormancy. Together, these results emphasise the need to consider both temporal and spatial processes when predicting multi-scale patterns of diversity.

Stabilising role of seed banks and the maintenance of bacterial diversity.

Coexisting species often exhibit negative frequency dependence due to mechanisms that promote population growth and persistence when rare. These stabilising mechanisms can maintain diversity through interspecific niche differences, but also through life-history strategies like dormancy that buffer populations in fluctuating environments. However, there are few tests demonstrating how seed banks contribute to long-term community dynamics and the maintenance of diversity. Using a multi-year, high-frequency time series of bacterial community data from a north temperate lake, we documented patterns consistent with stabilising coexistence. Bacterial taxa exhibited differential responses to seasonal environmental conditions, while seed bank dynamics helped maintain diversity over less-favourable winter periods. Strong negative frequency dependence in rare, but metabolically active, taxa suggested a role for biotic interactions in promoting coexistence. Together, our results provide field-based evidence that niche differences and seed banks contribute to recurring community dynamics and the long-term maintenance of diversity in nature.

Microbial community assembly in a multi-layer dendritic metacommunity.

A major goal of metacommunity ecology is to infer the local- and regional-scale processes that underlie community assembly. In dendritic ecological networks, branching patterns and directional flow can alter the balance between local and regional factors during assembly. Vertical habitat structure may further affect community assembly in dendritic metacommunities. In this study, we analyzed the bacterial metacommunity of a fifth-order mountain stream network to assess differences in community assembly (1) between planktonic and benthic habitats, (2) across spatial scales, and (3) between headwater and downstream regions of the network. Using taxonomic and phylogenetic null modeling, we found habitat-specific spatial patterns of community assembly across the dendritic network. Compositional differences between planktonic and benthic communities were maintained by variable selection, but we also found evidence of local dispersal limitation between the two habitats. Planktonic community assembly was scale dependent, transitioning from homogeneous selection at local scales to variable selection at regional scales, while benthic community assembly was less scale dependent. Variable selection structured headwaters in both habitat types, but downstream communities were primarily structured by homogeneous selection, especially in sediments. Taken together, our results show that vertical habitat structure contributes to the scale-dependent processes of community assembly across the dendritic metacommunity.

Metabolic insight into bacterial community assembly across ecosystem boundaries.

The movement of organisms across habitat boundaries has important consequences for populations, communities, and ecosystems. However, because most species are not well adapted to all habitat types, dispersal into suboptimal habitats could induce physiological changes associated with persistence strategies that influence community assembly. For example, high rates of cross-boundary dispersal are thought to maintain sink populations of terrestrial bacteria in aquatic habitats, but these bacteria may also persist by lowering their metabolic activity, introducing metabolic heterogeneity that buffers the population against species sorting. To differentiate between these assembly processes, we analyzed bacterial composition along a hydrological flow path from terrestrial soils through an aquatic reservoir by sequencing the active and total (active + inactive) portions of the community. When metabolic heterogeneity was ignored, our data were consistent with views that cross-boundary dispersal is important for structuring aquatic bacterial communities. In contrast, we found evidence for strong species sorting in the active portion of the aquatic community, suggesting that dispersal may have a weaker effect than persistence strategies on aquatic community assembly. By accounting for metabolic heterogeneity in complex communities, our findings clarify the roles of local- and regional-scale assembly processes in terrestrial-aquatic meta-ecosystems.

Dormancy in Metacommunities.

Although metacommunity ecology has improved our understanding of how dispersal affects community structure and dynamics across spatial scales, it has yet to adequately account for dormancy. Dormancy is a reversible state of reduced metabolic activity that enables temporal dispersal within the metacommunity. Dormancy is also a metacommunity-level process because it can covary with spatial dispersal and affect diversity across spatial scales. We develop a framework to integrate dispersal and dormancy, focusing on the covariation they exhibit, to predict how dormancy modifies the importance of species interactions, dispersal, and historical contingencies in metacommunities. We used empirical and modeling approaches to demonstrate the utility of this framework. We examined case studies of microcrustaceans in ephemeral ponds, where dormancy underlies metacommunity dynamics, and identified constraints on the dispersal and dormancy strategies of bromeliad-dwelling invertebrates. Using simulations, we showed that dormancy can alter classic metacommunity patterns of diversity in ways that depend on dispersal-dormancy covariation and spatiotemporal environmental variability. We propose that dormancy may also facilitate evolution-mediated priority effects if locally adapted seed banks prevent colonization by more dispersal-limited species. Last, we present testable predictions for the implications of dormancy in metacommunities, some of which may fundamentally alter our understanding of metacommunity ecology.