Environmental Filtering by pH and Salinity Jointly Drives Prokaryotic Community Assembly in Coastal Wetland Sediments

Understanding the microbial community assembly is an essential topic in microbial ecology. Coastal wetlands are an important blue carbon sink, where microbes play a key role in biogeochemical cycling of nutrients and energy transformation. However, the drivers controlling the distribution patterns and assembly of bacterial and archaeal communities in coastal wetland are unclear. Here we examined the diversity, co-occurrence network, assembly processes and environmental drivers of bacterial and archaeal communities from inshore to offshore sediments by the sequencing of 16S rRNA gene amplicons. The value of α- and β-diversity of bacterial and archaeal communities generally did not change significantly (P > 0.05) between offshore sites, but changed significantly (P < 0.05) among inshore sites. Sediment pH and salinity showed significant effects on the diversity and keystone taxa of bacterial and archaeal communities. The bacterial and archaeal co-occurrence networks were inextricably linked with pH and salinity to formed the large network nodes, suggesting that they were the key factors to drive the prokaryotic community. We also identified that heterogeneous and homogeneous selection drove the bacterial and archaeal community assembly, while the two selections became weaker from offshore sites to inshore sites, suggesting that deterministic processes were more important in offshore sites. Overall, these results suggested that the environmental filtering of pH and salinity jointly governed the assembly of prokaryotic community in offshore sediments. This study advances our understanding of microbial community assembly in coastal wetland ecosystems.

Coexistence patterns and diversity in a trait-based metacommunity on an environmental gradient

The dynamics of trait-based metacommunities have attracted much attention, but not much is known about how dispersal and spatial environmental variability mutually interact with each other to drive coexistence patterns and diversity. Here, we present a spatially explicit model of competition for two essential resources in a metacommunity on a one-dimensional environmental gradient. We find that both the strength of dispersal and the range of spatial environmental variability affect coexistence patterns, spatial structure, trait distribution, and local and regional diversity. Without dispersal, species are sorted according to their optimal growth conditions on the gradient. With the onset of dispersal, source-sink effects are initiated, which increases the effects of environmental filtering and interspecific competition and generates trait lumping, so that only a few species from an environment-defined trait range can survive. Interestingly, for very large dispersal rates, species distributions become spatially homogeneous, but nevertheless two species at the extreme ends of the trade-off curve can coexist for large environmental variability. Local species richness follows a classic hump-shaped dependence on dispersal rate, while local and regional diversity exhibit a pronounced peak for intermediate values of the environmental variability. Our findings provide important insights into the factors that shape the structure of trait-based metacommunities.

The Joint Contributions of Environmental Filtering and Spatial Processes to Macroinvertebrate Metacommunity Dynamics in the Alpine Stream Environment of Baima Snow Mountain, Southwest China

While macroinvertebrates are extensively investigated in many river ecosystems, meta-community ecology perspectives in alpine streams are very limited. We assessed the role of ecological factors and temporal dynamics in the macroinvertebrate meta-community assembly of an alpine stream situated in a dry-hot valley of Baima Snow Mountain, China. We found that spatial structuring and environmental filtering jointly drive the structure of macroinvertebrate meta-community, with relative contributions to the variance in community composition changing over time. RDA ordination and variation partitioning indicate that environmental variables are the most important predictors of community organization in most scenarios, whereas spatial determinants also play a significant role. Moreover, the explanatory power, identity, and the relative significance of ecological factors change over time. Particularly, in the years 2018 and 2019, stronger environmental filtering was found shaping community assembly, suggesting that deterministic mechanisms predominated in driving community dynamics. However, spatial factors had a stronger predictive power on meta-community structures in 2017, implying conspicuous dispersal mechanisms which may be owing to increased connectivity amongst sites. Thereby, we inferred that the alpine stream macroinvertebrate metacommunity composition can be regulated by the interaction of both spatial processes and environmental filtering, with relative contributions varying over time. Based on these findings, we suggest that community ecology studies in aquatic systems should be designed beyond single snapshot investigations.

Modular marvels: The ecology, evolvability, and energetics of bryozoan polymorphism

<p>Modularity is a fundamental concept in biology. Most taxa within the colonial invertebrate phylum Bryozoa have achieved division of labor through the development of specialized modules (polymorphs), and this group is perhaps the most outstanding exemplar of the phenomenon. This thesis addresses several gaps in the literature concerning the morphology, ecology, energetics, and evolvability of bryozoan polymorphism. It has been over 40 years since the last review of bryozoan polymorphism, and here I provide a comprehensive update that describes the diversity, morphology, and function of bryozoan polymorphs and the significance of modularity to their evolutionary success. While the degree of module compartmentalization is important for the evolution of polymorphism in bryozoans, this does not appear to be the case for other colonial invertebrates. To facilitate data collection, I developed a classification system for polymorphism in cheilostome bryozoans. While classification systems exist for bryozoan colony form, the system presented here is the first developed for polymorphism. This system is fully illustrated and non-hierarchical, enabling swift classification and statistical comparisons at many levels of detail. Understanding community assembly is a key goal in community ecology, but previous work on bryozoan communities has focused on colony form rather than polymorphism. Environmental filtering influences community assembly by excluding ill-adapted species, resulting in communities with similar functional traits. An RLQ (a four-way ordination) analysis incorporating spatial data was run on a dataset of 642 species of cheilostomes from 779 New Zealand sites, to investigate environmental filtering of colony form and zooid polymorphism. This revealed environmental filtering of colony form: encrusting-cemented taxa were predominant in shallow environments with hard substrata (200 m). Furthermore, erect taxa found in shallow environments with high current speeds were typically jointed. Surprisingly, polymorphism also followed environmental gradients. External ovicells (brood chambers) were more common in deeper, low oxygen water than immersed and internal ovicells. This may reflect the oxygen needs of the embryo or increased predation intensity in shallow environments. Bryozoans with costae (rib-like spines) tended to be found in deeper water as well, while bryozoans with calcified frontal shields were found in shallow environments with a higher concentration of CaCO₃. Avicularia (defensive grasping structures) were not related to environmental conditions, and changes in pivot bar structure with depth likely represent a phylogenetic signal. Factors influencing community assembly were somewhat partitioned by levels of organization, since colony form responds to environmental conditions, while the effects of evolutionary history, predation, and environmental conditions were not well-separated for zooid-level morphology. Finally, rootlets may have been a key innovation that allowed cementing taxa to escape hard substrata, potentially contributing to the cheilostome radiation. Despite the diversity of life on earth, many morphologies have not been achieved. Morphology can be limited by a variety of constraints (developmental, historical, biomechanical) and comparing the distribution of realized forms in a theoretical form-space (i.e. “morphospace”) can highlight which constraints are at play and potential functions. If traits cluster around biomechanical optima, then morphology may be shaped by strong selective pressures. In contrast, a well-explored (filled) morphospace suggests weak constraints and high morphological evolvability. Here, constraints on morphospace exploration were examined for 125 cheilostome bryozoan species from New Zealand. The mandible morphospaces for avicularia (beak-like polymorphs) were visualized using Coordinate-Point Extended Eigenshape analysis. Mechanical advantage, moment of inertia, drag, peak force, and rotational work required to close the mandible were calculated for theoretical (n=47) and real mandibles (n=224) to identify biomechanical optima. The volume and surface of area of the parcel of water passed through by the closing mandible (referred to as the “domain”) was also calculated. The theoretical morphospace of avicularia is well-explored, suggesting they are highly evolvable and have relaxed developmental constraints. However, there may be constraints within lineages. A well-developed fulcrum (complete pivot bar) may be an evolutionary pre/corequisite to evolving mandibles with extreme moments of inertia such as setose and highly spathulate forms. The most common mandible shape, triangular, represents a trade-off between maximizing domain size, minimizing energetic cost (force and construction material), and minimizing the potential for breakage. This suggests that they are well suited for catching epibionts, representing the first empirical evidence for avicularian function. Tendon length and mechanical advantage are limited by tendon width, which itself is constrained by the base width of the mandible. This explains the low mechanical advantage of setose mandibles and suggests that they are unable to grasp epibionts. The calories required to close the mandible of an avicularium (estimated from rotational work) are quite small (1.24 x 10⁻¹⁶ to 8.82 x 10⁻¹¹ cal). Overall, this thesis highlights the complexity of bryozoan polymorphism and suggests cheilostome avicularia could provide a unique evolutionary system to study due to their apparent lack of strong developmental constraints. Future studies into the ecology of polymorphism should focus on the degree of investment (polymorph abundance within a colony) rather than presence or absence.</p>

Taxonomic and functional dissimilarities of soil bacterial communities are more related to environmental dissimilarity than geographic distance

The processes governing soil bacteria biogeography are still not fully understood. It remains unknown how the importance of environmental filtering and dispersal differs between bacterial taxonomic and functional biogeography, and whether their importance is scale-dependent. We sampled soils at 195 plots across the Tibet plateau, with distances among plots ranging from 20 m to 1 550 km. Taxonomic composition of bacterial community was characterized by 16S amplicon sequencing, and functional community composition by qPCR targeting 9 functional groups involved in N dynamics. Twelve climatic and soil characteristics were also measured. Both taxonomic and functional dissimilarities were more related to environmental dissimilarity than geographic distance. Taxonomic dissimilarity was mostly explained by soil pH and organic matter, while functional dissimilarity was mostly linked to moisture, temperature and N, P and C availabilities. The roles of environmental filtering and dispersal were, however, scale-dependent and varied between taxonomic and functional dissimilarities, with distance affecting taxonomic dissimilarity over short distances (<~300 km) and functional dissimilarity over long distances (>~600 km). The importance of different environmental predictors varied across scales more for functional than taxonomic dissimilarity. Our results demonstrate how biodiversity dimension (taxonomic versus functional) and spatial scale strongly influence the conclusions derived from bacterial biogeography studies.

Modular marvels: The ecology, evolvability, and energetics of bryozoan polymorphism

<p>Modularity is a fundamental concept in biology. Most taxa within the colonial invertebrate phylum Bryozoa have achieved division of labor through the development of specialized modules (polymorphs), and this group is perhaps the most outstanding exemplar of the phenomenon. This thesis addresses several gaps in the literature concerning the morphology, ecology, energetics, and evolvability of bryozoan polymorphism. It has been over 40 years since the last review of bryozoan polymorphism, and here I provide a comprehensive update that describes the diversity, morphology, and function of bryozoan polymorphs and the significance of modularity to their evolutionary success. While the degree of module compartmentalization is important for the evolution of polymorphism in bryozoans, this does not appear to be the case for other colonial invertebrates. To facilitate data collection, I developed a classification system for polymorphism in cheilostome bryozoans. While classification systems exist for bryozoan colony form, the system presented here is the first developed for polymorphism. This system is fully illustrated and non-hierarchical, enabling swift classification and statistical comparisons at many levels of detail. Understanding community assembly is a key goal in community ecology, but previous work on bryozoan communities has focused on colony form rather than polymorphism. Environmental filtering influences community assembly by excluding ill-adapted species, resulting in communities with similar functional traits. An RLQ (a four-way ordination) analysis incorporating spatial data was run on a dataset of 642 species of cheilostomes from 779 New Zealand sites, to investigate environmental filtering of colony form and zooid polymorphism. This revealed environmental filtering of colony form: encrusting-cemented taxa were predominant in shallow environments with hard substrata (200 m). Furthermore, erect taxa found in shallow environments with high current speeds were typically jointed. Surprisingly, polymorphism also followed environmental gradients. External ovicells (brood chambers) were more common in deeper, low oxygen water than immersed and internal ovicells. This may reflect the oxygen needs of the embryo or increased predation intensity in shallow environments. Bryozoans with costae (rib-like spines) tended to be found in deeper water as well, while bryozoans with calcified frontal shields were found in shallow environments with a higher concentration of CaCO₃. Avicularia (defensive grasping structures) were not related to environmental conditions, and changes in pivot bar structure with depth likely represent a phylogenetic signal. Factors influencing community assembly were somewhat partitioned by levels of organization, since colony form responds to environmental conditions, while the effects of evolutionary history, predation, and environmental conditions were not well-separated for zooid-level morphology. Finally, rootlets may have been a key innovation that allowed cementing taxa to escape hard substrata, potentially contributing to the cheilostome radiation. Despite the diversity of life on earth, many morphologies have not been achieved. Morphology can be limited by a variety of constraints (developmental, historical, biomechanical) and comparing the distribution of realized forms in a theoretical form-space (i.e. “morphospace”) can highlight which constraints are at play and potential functions. If traits cluster around biomechanical optima, then morphology may be shaped by strong selective pressures. In contrast, a well-explored (filled) morphospace suggests weak constraints and high morphological evolvability. Here, constraints on morphospace exploration were examined for 125 cheilostome bryozoan species from New Zealand. The mandible morphospaces for avicularia (beak-like polymorphs) were visualized using Coordinate-Point Extended Eigenshape analysis. Mechanical advantage, moment of inertia, drag, peak force, and rotational work required to close the mandible were calculated for theoretical (n=47) and real mandibles (n=224) to identify biomechanical optima. The volume and surface of area of the parcel of water passed through by the closing mandible (referred to as the “domain”) was also calculated. The theoretical morphospace of avicularia is well-explored, suggesting they are highly evolvable and have relaxed developmental constraints. However, there may be constraints within lineages. A well-developed fulcrum (complete pivot bar) may be an evolutionary pre/corequisite to evolving mandibles with extreme moments of inertia such as setose and highly spathulate forms. The most common mandible shape, triangular, represents a trade-off between maximizing domain size, minimizing energetic cost (force and construction material), and minimizing the potential for breakage. This suggests that they are well suited for catching epibionts, representing the first empirical evidence for avicularian function. Tendon length and mechanical advantage are limited by tendon width, which itself is constrained by the base width of the mandible. This explains the low mechanical advantage of setose mandibles and suggests that they are unable to grasp epibionts. The calories required to close the mandible of an avicularium (estimated from rotational work) are quite small (1.24 x 10⁻¹⁶ to 8.82 x 10⁻¹¹ cal). Overall, this thesis highlights the complexity of bryozoan polymorphism and suggests cheilostome avicularia could provide a unique evolutionary system to study due to their apparent lack of strong developmental constraints. Future studies into the ecology of polymorphism should focus on the degree of investment (polymorph abundance within a colony) rather than presence or absence.</p>

Beta Diversity of Antarctic and Sub-Antarctic Benthic Communities Reveals a Major Role of Stochastic Assembly Processes

Community assembly is the result of both, deterministic and stochastic processes. The former encompasses niche-based local-scale mechanisms such as environmental filtering and biotic interactions; the latter includes ecological drift, probabilistic colonisation, and random extinctions. Using standardised sampling protocols, we show that the spatial variation in species composition (beta diversity) of shallow subtidal macrobenthic communities of sub-Antarctic (Strait of Magellan and Yendegaia Fjord [Beagle Channel]) and Antarctic (Fildes Bay [King George Island, West Antarctic Peninsula]) localities reflects a high contribution of stochastic processes to community assembly. Null model analyses indicated that random sampling from species pools of different sizes drove the observed among-locality differences in incidence- and abundance-based beta diversity. We analysed a normalised stochasticity ratio (NST), which delimits between more deterministic (&lt;50%) and more stochastic (&gt;50%) assembly. NST was notably larger than 50%, with mean values of 69.5% (95% CI = 69.2–69.8%), 62.5% (62.1–62.9%), and 72.8% (72.5–73.2%) in Strait of Magellan, Yendegaia Fjord, and Fildes Bay, respectively. Accordingly, environmental factors, such as depth, seawater temperature, salinity, and underwater light penetration, accounted for a small fraction of the spatial variation in community composition across the three localities. In this region, therefore, stochastic processes could have stronger effects on community assembly than deterministic niche-based factors. As anthropogenic biotic homogenisation continues apace, our study can give useful insights into the major ecological processes in Southern Ocean’ coastal marine communities.

Diverse recruitment to a taxonomically structured global atmospheric microbiota

Atmospheric transport is critical to dispersal of microorganisms between habitats and this underpins resilience in terrestrial and marine ecosystems globally. A key unresolved question is whether microorganisms assemble to form a taxonomically distinct, geographically variable, and functionally adapted atmospheric microbiota. Here we characterised inter-continental patterns of microbial taxonomic and functional diversity in air within and above the atmospheric boundary layer and in underlying soils for 596 globally sourced samples. Bacterial and fungal assemblages in air were taxonomically structured and deviated significantly from purely stochastic assembly. Patterns differed with location and reflected underlying surface cover and environmental filtering. Source-tracking indicated a complex recruitment process involving local soils plus globally distributed inputs from drylands and the phyllosphere. Assemblages displayed stress-response and metabolic traits relevant to survival in air, and taxonomic and functional diversity were correlated with macroclimate and atmospheric variables. Our findings highlight complexity in the atmospheric microbiota that is key to understanding regional and global ecosystem connectivity.