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The Order Pelagibacterales (SAR11) is the most abundant group of heterotrophic bacterioplankton in global oceans and comprises multiple subclades with unique spatiotemporal distributions. Subclade IIIa is the primary SAR11 group in brackish waters and shares a common ancestor with the dominant freshwater IIIb (LD12) subclade. Despite its dominance in brackish environments, subclade IIIa lacks systematic genomic or ecological studies. Here, we combine closed genomes from new IIIa isolates, new IIIa MAGS from San Francisco Bay (SFB), and 460 highly complete publicly available SAR11 genomes for the most comprehensive pangenomic study of subclade IIIa to date. Subclade IIIa represents a taxonomic family containing three genera (denoted as subgroups IIIa.1, IIIa.2, and IIIa.3) that had distinct ecological distributions related to salinity. The expansion of taxon selection within subclade IIIa also established previously noted metabolic differentiation in subclade IIIa compared to other SAR11 subclades such as glycine/serine prototrophy, mosaic glyoxylate shunt presence, and polyhydroxyalkanoate synthesis potential. Our analysis further shows metabolic flexibility among subgroups within IIIa. Additionally, we find that subclade IIIa.3 bridges the marine and freshwater clades based on its potential for compatible solute transport, iron utilization, and bicarbonate management potential. Pure culture experimentation validated differential salinity ranges in IIIa.1 and IIIa.3 and provided detailed IIIa cell size and volume data. This study is an important step forward for understanding the genomic, ecological, and physiological differentiation of subclade IIIa and the overall evolutionary history of SAR11.

 

Beach aquifers, located in the subsurface of sandy beaches, are unique ecosystems with steep chemical and physical gradients resulting from the mixing of terrestrial fresh groundwater and saline groundwater from the sea. While work has rapidly progressed to understand the physics and chemistry in this environment, much less is known about the microorganisms present despite the fact that they are responsible for vital biogeochemical processes. This paper presents a review of the current state of knowledge of microbes within beach aquifers and the mechanisms that control the beach aquifer microbiome. We review literature describing the distribution and diversity of microorganisms in the freshwater-saltwater mixing zone of beach aquifers, and identify just 12 papers. We highlight knowledge gaps, as well as future research directions: The understanding of beach aquifer microorganisms is informed primarily by 16S ribosomal RNA gene sequences. Metagenomics and metatranscriptomics have not yet been applied but are promising approaches for elucidating key metabolic and ecological roles of microbes in this environment. Additionally, variability in field sampling and analytical methods restrict comparison of data across studies and geographic locations. Further, documented evidence on the migration of microbes within the beach aquifer is limited. Taking into account the physical transport of microbes through sand by flowing groundwater may be critical for understanding the structure and dynamics of microbial communities. Quantitative measurements of rates of elemental cycling in the context of microbial diversity need further investigation, in order to understand the roles of microbes in mediating biogeochemical fluxes from the beach aquifer to the coastal ocean. Lastly, understanding the current state of beach aquifers in regulating carbon stocks is critical to foster a better understanding of the contribution of the beach aquifer microbiome to global climate models. 

Ecological factors contributing to depth-related diversification of marine Thaumarchaeota populations remain largely unresolved. To investigate the role of potential microbial associations in shaping thaumarchaeal ecotype diversification, we examined co-occurrence relationships in a community composition dataset (16S rRNA V4-V5 region) collected as part of a 2-year time series in coastal Monterey Bay. Ecotype groups previously defined based on functional gene diversity—water column A (WCA), water column B (WCB) and Nitrosopumilus-like clusters—were recovered in the thaumarchaeal 16S rRNA gene phylogeny. Networks systematically reflected depth-related patterns in the abundances of ecotype populations, suggesting thaumarchaeal ecotypes as keystone members of the microbial community below the euphotic zone. Differential environmental controls on the ecotype populations were further evident in subnetwork modules showing preferential co-occurrence of OTUs belonging to the same ecotype cluster. Correlated abundances of Thaumarchaeota and heterotrophic bacteria (e.g., Bacteroidetes, Marinimicrobia and Gammaproteobacteria) indicated potential reciprocal interactions via dissolved organic matter transformations. Notably, the networks recovered ecotype-specific associations between thaumarchaeal and Nitrospina OTUs. Even at depths where WCB-like Thaumarchaeota dominated, Nitrospina OTUs were found to preferentially co-occur with WCA-like and Nitrosopumilus-like thaumarchaeal OTUs, highlighting the need to investigate the ecological implications of the composition of nitrifier assemblages in marine waters.

 

Ammonia‐oxidizing archaea (AOA) of the phylumThaumarchaeotaare key players in nutrient cycling, yet large gaps remain in our understanding of their ecology and metabolism. Despite multiple lines of evidence pointing to a central role for copper‐containing nitrite reductase (NirK) in AOA metabolism, the thaumarchaealnirKgene is rarely studied in the environment. In this study, we examine the diversity ofnirKin the marine pelagic environment, in light of previously described ecological patterns of pelagic thaumarchaeal populations. Phylogenetic analyses show thatnirKbetter resolves diversification patterns of marineThaumarchaeota, compared to the conventionally used marker geneamoA. Specifically, we demonstrate that the three major phylogenetic clusters of marinenirKcorrespond to the three ‘ecotype’ populations of pelagicThaumarchaeota. In this context, we further examine the relative distributions of the three variant groups in metagenomes and metatranscriptomes representing two depth profiles in coastal Monterey Bay. Our results reveal thatnirKeffectively tracks the dynamics of thaumarchaeal ecotype populations, particularly finer‐scale diversification patterns within major lineages. We also find evidence for multiple copies ofnirKper genome in a fraction of thaumarchaeal cells in the water column, which must be taken into account when using it as a molecular marker.

 

Nitrification plays a key role in marine ecosystems whereThaumarchaeotaare thought to be responsible for most of the ammonia oxidation in the water column. Over a 2‐yr, near‐monthly time series at two sites in Monterey Bay we observed repeatable seasonal and depth‐based patterns ofThaumarchaeotaecotype abundance that highlighted a clear delineation between populations in shallow euphotic (< 50 m) vs. deeper mesopelagic (60–500 m) depths. Euphotic depths show greater seasonality and influence from light, while mesopelagic waters have trends based on water mass and other covarying features with depth. Three major ecotypes were recovered: aNitrosopumilus‐like (NP) group, aNitrosopelagicus‐like ecotype containing “shallow” water column A (WCA) members, and an ecotype affiliated with the “deep” water column B (WCB)Thaumarchaeota. These ecotypes show a strong depth distribution, with WCB dominant at ≥ 200 m depth and WCA most abundant in surface (5–100 m) waters. The NP ecotype was found throughout the water column with the highest abundance in summer, and was the only ecotype showing a correlation with measured nitrification rates. We also found three abundant taxa related toNitrospina—the major nitrite‐oxidizing bacteria in the ocean; these showed clear connections to each of the threeThaumarchaeotaecotypes, suggesting a specific relationship between both steps of nitrification. Our results support the importance of ecotype‐based analysis ofThaumarchaeotaand show that their abundance and distribution are controlled based on their water column position, with a distinct shift at 50 m between euphotic and mesopelagic depths.