Recently, microbiologists have established the existence of biogeographic patterns among a wide range of microorganisms. The focus of the field is now shifting to identifying the mechanisms that shape these patterns. Here, we propose that four processes - selection, drift, dispersal and mutation - create and maintain microbial biogeographic patterns on inseparable ecological and evolutionary scales. We consider how the interplay of these processes affects one biogeographic pattern, the distance-decay relationship, and review evidence from the published literature for the processes driving this pattern in microorganisms. Given the limitations of inferring processes from biogeographic patterns, we suggest that studies should focus on directly testing the underlying processes.

Soils are immensely diverse microbial habitats with thousands of co-existing bacterial, archaeal, and fungal species. Across broad spatial scales, factors such as pH and soil moisture appear to determine the diversity and structure of soil bacterial communities. Within any one site however, bacterial taxon diversity is high and factors maintaining this diversity are poorly resolved. Candidate factors include organic substrate availability and chemical recalcitrance, and given that they appear to structure bacterial communities at the phylum level, we examine whether these factors might structure bacterial communities at finer levels of taxonomic resolution. Analyzing 16S rRNA gene composition of nucleotide analog-labeled DNA by PhyloChip microarrays, we compare relative growth rates on organic substrates of increasing chemical recalcitrance of >2,200 bacterial taxa across 43 divisions/phyla. Taxa that increase in relative abundance with labile organic substrates (i.e., glycine, sucrose) are numerous (>500), phylogenetically clustered, and occur predominantly in two phyla (Proteobacteria and Actinobacteria) including orders Actinomycetales, Enterobacteriales, Burkholderiales, Rhodocyclales, Alteromonadales, and Pseudomonadales. Taxa increasing in relative abundance with more chemically recalcitrant substrates (i.e., cellulose, lignin, or tannin–protein) are fewer (168) but more phylogenetically dispersed, occurring across eight phyla and including Clostridiales, Sphingomonadalaes, Desulfovibrionales. Just over 6% of detected taxa, including many Burkholderiales increase in relative abundance with both labile and chemically recalcitrant substrates. Estimates of median rRNA copy number per genome of responding taxa demonstrate that these patterns are broadly consistent with bacterial growth strategies. Taken together, these data suggest that changes in availability of intrinsically labile substrates may result in predictable shifts in soil bacterial composition.

Nitrogen (N) availability is increasing in many ecosystems due to anthropogenic disturbance. We used a nucleotide analog technique and sequencing of ribosomal RNA genes to test whether N fertilization altered active fungal communities in two boreal ecosystems. In decaying litter from a recently burned spruce forest, Shannon diversity decreased significantly with N fertilization, and taxonomic richness declined from 44 to 33 operational taxonomic units (OTUs). In soils from a mature spruce forest, richness also declined with N fertilization, from 67 to 52 OTUs. Fungal community structure in litter differed significantly with N fertilization, primarily because fungi of the order Ceratobasidiales increased in abundance. We observed similar changes in fungal diversity and community structure with starch addition to litter, suggesting that N fertilization may affect fungal communities by altering plant carbon inputs. These changes could have important consequences for ecosystem processes such as decomposition and nutrient mineralization. r

Soil microbes are among the most abundant and diverse organisms on Earth. Although microbial decomposers, particularly fungi, are important mediators of global carbon and nutrient cycling, the functional roles of specific taxa within natural environments remain unclear. We used a nucleotide-analog technique in soils from the Harvard Forest to characterize the fungal taxa that responded to the addition of five different carbon substrates-glycine, sucrose, cellulose, lignin, and tannin-protein. We show that fungal community structure and richness shift in response to different carbon sources, and we demonstrate that particular fungal taxa target different organic compounds within soil microcosms. Specifically, we identified eleven taxa that exhibited changes in relative abundances across substrate treatments. These results imply that niche partitioning through specialized resource use may be an important mechanism by which soil microbial diversity is maintained in ecosystems. Consequently, high microbial diversity may be necessary to sustain ecosystem processes and stability under global change.

Despite the important role of phages in marine systems, little is understood about how their diversity is distributed in space. Biogeographic patterns of marine phages may be difficult to detect due to their vast genetic diversity, which may not be accurately represented by conserved marker genes. To investigate the spatial biogeographic structure of marine phages, we isolated over 400 cyanophages on Synechococcus host strain WH7803 at three coastal locations in the United States (Rhode Island, Washington, and southern California). Approximately 90% of the cyanophage isolates were myoviruses, while the other 10% were podoviruses. The diversity of isolates was further characterized in two ways: (i) taxonomically, using conserved marker genes and (ii) phenotypically, by testing isolates for their ability to infect a suite of hosts, or their “host range.” Because host range is a highly variable trait even among closely related isolates, we hypothesized that host range phenotypes of cyanophage isolates would vary more strongly among locations than would taxonomic composition. Instead, we found evidence for strong biogeographic variation both in taxonomic composition and host range phenotypes, with little taxonomic overlap among the three coastal regions. For both taxonomic composition and host range phenotypes, cyanophage communities from California and Rhode Island were the most dissimilar, while Washington communities exhibited similarity to each of the other two locations. These results suggest that selection imposed by spatial variation in host dynamics influence the biogeographic distribution of cyanophages.