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Noah Fierer - One of the best experts on this subject based on the ideXlab platform.
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Co-habiting amphibian species harbor unique skin Bacterial Communities in wild populations
The ISME Journal, 2012Co-Authors: Valerie J Mckenzie, Noah Fierer, Rob Knight, Robert M Bowers, Christian L LauberAbstract:Although all plant and animal species harbor microbial symbionts, we know surprisingly little about the specificity of microbial Communities to their hosts. Few studies have compared the microbiomes of different species of animals, and fewer still have examined animals in the wild. We sampled four pond habitats in Colorado, USA, where multiple amphibian species were present. In total, 32 amphibian individuals were sampled from three different species including northern leopard frogs ( Lithobates pipiens ), western chorus frogs ( Pseudacris triseriata ) and tiger salamanders ( Ambystoma tigrinum ). We compared the diversity and composition of the Bacterial Communities on the skin of the collected individuals via barcoded pyrosequencing of the 16S rRNA gene. Dominant Bacterial phyla included Acidobacteria , Actinobacteria , Bacteriodetes , Cyanobacteria , Firmicutes and Proteobacteria . In total, we found members of 18 Bacterial phyla, comparable to the taxonomic diversity typically found on human skin. Levels of Bacterial diversity varied strongly across species: L. pipiens had the highest diversity; A. tigrinum the lowest. Host species was a highly significant predictor of Bacterial community similarity, and co-habitation within the same pond was not significant, highlighting that the skin-associated Bacterial Communities do not simply reflect those Bacterial Communities found in their surrounding environments. Innate species differences thus appear to regulate the structure of skin Bacterial Communities on amphibians. In light of recent discoveries that some bacteria on amphibian skin have antifungal activity, our finding suggests that host-specific bacteria may have a role in the species-specific resistance to fungal pathogens.
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Spatial variability in airborne Bacterial Communities across land-use types and their relationship to the Bacterial Communities of potential source environments
The ISME Journal, 2011Co-Authors: Robert M Bowers, Rob Knight, Shawna Mcletchie, Noah FiererAbstract:Although bacteria are ubiquitous in the near-surface atmosphere and they can have important effects on human health, airborne bacteria have received relatively little attention and their spatial dynamics remain poorly understood. Owing to differences in meteorological conditions and the potential sources of airborne bacteria, we would expect the atmosphere over different land-use types to harbor distinct Bacterial Communities. To test this hypothesis, we sampled the near-surface atmosphere above three distinct land-use types (agricultural fields, suburban areas and forests) across northern Colorado, USA, sampling five sites per land-use type. Microbial abundances were stable across land-use types, with ∼10^5–10^6 Bacterial cells per m^3 of air, but the concentrations of biological ice nuclei, determined using a droplet freezing assay, were on average two and eight times higher in samples from agricultural areas than in the other two land-use types. Likewise, the composition of the airborne Bacterial Communities, assessed via bar-coded pyrosequencing, was significantly related to land-use type and these differences were likely driven by shifts in the sources of bacteria to the atmosphere across the land-uses, not local meteorological conditions. A meta-analysis of previously published data shows that atmospheric Bacterial Communities differ from those in potential source environments (leaf surfaces and soils), and we demonstrate that we may be able to use this information to determine the relative inputs of bacteria from these source environments to the atmosphere. This work furthers our understanding of Bacterial diversity in the atmosphere, the terrestrial controls on this diversity and potential approaches for source tracking of airborne bacteria.
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Bacterial Communities associated with the lichen symbiosis
Applied and Environmental Microbiology, 2011Co-Authors: Scott T Bates, Rob Knight, Garrett W G Cropsey, Gregory J Caporaso, Noah FiererAbstract:Lichens are commonly described as a mutualistic symbiosis between fungi and “algae” (Chlorophyta or Cyanobacteria); however, they also have internal Bacterial Communities. Recent research suggests that lichen-associated microbes are an integral component of lichen thalli and that the classical view of this symbiotic relationship should be expanded to include bacteria. However, we still have a limited understanding of the phylogenetic structure of these Communities and their variability across lichen species. To address these knowledge gaps, we used bar-coded pyrosequencing to survey the Bacterial Communities associated with lichens. Bacterial sequences obtained from four lichen species at multiple locations on rock outcrops suggested that each lichen species harbored a distinct community and that all Communities were dominated by Alphaproteobacteria. Across all samples, we recovered numerous Bacterial phylotypes that were closely related to sequences isolated from lichens in prior investigations, including those from a lichen-associated Rhizobiales lineage (LAR1; putative N2 fixers). LAR1-related phylotypes were relatively abundant and were found in all four lichen species, and many sequences closely related to other known N2 fixers (e.g., Azospirillum, Bradyrhizobium, and Frankia) were recovered. Our findings confirm the presence of highly structured Bacterial Communities within lichens and provide additional evidence that these bacteria may serve distinct functional roles within lichen symbioses.
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consistent effects of nitrogen fertilization on soil Bacterial Communities in contrasting systems
Ecology, 2010Co-Authors: Kelly S Ramirez, Noah Fierer, Rob Knight, Christian L Lauber, Mark A BradfordAbstract:: Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) through anthropogenic activities. Although the effects of increased N inputs on plant Communities have been reasonably well studied, few comparable studies have examined impacts on whole soil Bacterial Communities, though they play critical roles in ecosystem functioning. We sampled soils from two long-term ecological research (LTER) experimental N gradients, both of which have been amended with NH4NO3; a grassland at Cedar Creek (27 years of N additions) and an agricultural field at Kellogg Biological Station (8 years of N additions). By examining shifts in Bacterial Communities across these contrasting ecosystem types, we could test competing hypotheses about the direct and indirect factors that might drive Bacterial responses to elevated N inputs. Bacterial community structure was highly responsive to N additions. We observed predictable and consistent changes in the structure of the Bacterial Communities across both ecosystem types. Our results suggest that Bacterial Communities across these gradients are more structured by N and/or soil carbon availability than by shifts in the plant community or soil pH associated with the elevated nitrogen inputs. In contrast to the pronounced shifts in Bacterial community composition and in direct contrast to the patterns often observed in plant Communities, increases in N availability did not have consistent effects on the richness and diversity of soil Bacterial Communities.
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Forensic identification using skin Bacterial Communities.
Proceedings of the National Academy of Sciences of the United States of America, 2010Co-Authors: Noah Fierer, Cl Christian L Lauber, Nick Zhou, Elizabeth K. Costello, Daniel Mcdonald, Rob KnightAbstract:Recent work has demonstrated that the diversity of skin-associated Bacterial Communities is far higher than previously recognized, with a high degree of interindividual variability in the composition of Bacterial Communities. Given that skin Bacterial Communities are personalized, we hypothesized that we could use the residual skin bacteria left on objects for forensic identification, matching the bacteria on the object to the skin-associated bacteria of the individual who touched the object. Here we describe a series of studies de-monstrating the validity of this approach. We show that skin-associated bacteria can be readily recovered from surfaces (including single computer keys and computer mice) and that the structure of these Communities can be used to differentiate objects handled by different individuals, even if those objects have been left untouched for up to 2 weeks at room temperature. Furthermore, we demonstrate that we can use a high-throughput pyrosequencing-based ap-proach to quantitatively compare the Bacterial Communities on objects and skin to match the object to the individual with a high degree of certainty. Although additional work is needed to further establish the utility of this approach, this series of studies introduces a forensics approach that could eventually be used to independently evaluate results obtained using more traditional forensic practices.
Jonathan M. Adams - One of the best experts on this subject based on the ideXlab platform.
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nutrient resource availability mediates niche differentiation and temporal co occurrence of soil Bacterial Communities
Applied Soil Ecology, 2021Co-Authors: Qiang Lin, Jonathan M. Adams, Petr HeděnecAbstract:Abstract Nutrient resource availability is a crucial environmental factor in determining soil Bacterial community assembly. However, the effects of nutrient availability on niche differentiation and temporal co-occurrence patterns of Bacterial Communities remain elusive. Here, we retrieved sequencing datasets (105 amplicon samples of Bacterial 16S rRNA genes and 60 Bacterial metagenomic samples) from a laboratory incubation experiment and a coastal wetland study, to elucidate the effects of nutrient availability on niche differentiation and co-occurrence patterns of Bacterial Communities. Niche differentiation within Bacterial Communities, indicated by nearest taxon index and niche breadth of Bacterial Communities, did not significantly correlate with nutrient availability gradients in either environment, even though deterministic processes dominated Bacterial community assembly. Network analyses revealed that co-occurrence patterns of Bacterial OTUs over incubation time changed across nutrient availability gradients, and were likely to be mediated by environmental selection. There were more co-occurrences of metabolic genes under higher nutrient levels. This study revealed the non-significant correlation between nutrient availability gradients and niche differentiation within Bacterial Communities, not only in a natural environment but also in a laboratory modified environment where the nutrient availability gradient was the sole environmental variable. These findings boost the understanding of Bacterial community assembly along environmental nutrient availability gradients.
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Distinctive Bacterial Communities in the Rhizoplane of Four Tropical Tree Species
Microbial Ecology, 2012Co-Authors: Yoon Myung Oh, Larisa Lee-cruz, Rusea Go, N. Ainuddin, Raha Abdul Rahim, Noraini Shukor, Jonathan M. AdamsAbstract:It is known that the microbial community of the rhizosphere is not only influenced by factors such as root exudates, phenology, and nutrient uptake but also by the plant species. However, studies of Bacterial Communities associated with tropical rainforest tree root surfaces, or rhizoplane, are lacking. Here, we analyzed the Bacterial community of root surfaces of four species of native trees, Agathis borneensis , Dipterocarpus kerrii , Dyera costulata , and Gnetum gnemon , and nearby bulk soils, in a rainforest arboretum in Malaysia, using 454 pyrosequencing of the 16S rRNA gene. The rhizoplane Bacterial Communities for each of the four tree species sampled clustered separately from one another on an ordination, suggesting that these assemblages are linked to chemical and biological characteristics of the host or possibly to the mycorrhizal fungi present. Bacterial Communities of the rhizoplane had various similarities to surrounding bulk soils. Acidobacteria, Alphaproteobacteria, and Betaproteobacteria were dominant in rhizoplane Communities and in bulk soils from the same depth (0–10 cm). In contrast, the relative abundance of certain Bacterial lineages on the rhizoplane was different from that in bulk soils: Bacteroidetes and Betaproteobacteria, which are known as copiotrophs, were much more abundant in the rhizoplane in comparison to bulk soil. At the genus level, Burkholderia , Acidobacterium , Dyella , and Edaphobacter were more abundant in the rhizoplane. Burkholderia , which are known as both pathogens and mutualists of plants, were especially abundant on the rhizoplane of all tree species sampled. The Burkholderia species present included known mutualists of tropical crops and also known N fixers. The host-specific character of tropical tree rhizoplane Bacterial Communities may have implications for understanding nutrient cycling, recruitment, and structuring of tree species diversity in tropical forests. Such understanding may prove to be useful in both tropical forestry and conservation.
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Distinctive Phyllosphere Bacterial Communities in Tropical Trees
Microbial Ecology, 2012Co-Authors: Dharmesh Singh, Rusea Go, Raha Abdul Rahim, Jongsik Chun, Jonathan M. AdamsAbstract:Recent work has suggested that in temperate and subtropical trees, leaf surface Bacterial Communities are distinctive to each individual tree species and dominated by Alpha- and Gammaproteobacteria. In order to understand how general this pattern is, we studied the phyllosphere Bacterial community on leaves of six species of tropical trees at a rainforest arboretum in Malaysia. This represents the first detailed study of ‘true’ tropical lowland tree phyllosphere Communities. Leaf surface DNA was extracted and pyrosequenced targeting the V1–V3 region of 16S rRNA gene. As was previously found in temperate and subtropical trees, each tree species had a distinctive Bacterial community on its leaves, clustering separately from other tree species in an ordination analysis. Bacterial Communities in the phyllosphere were unique to plant leaves in that very few operational taxonomic units (0.5%) co-occurred in the surrounding soil environment. A novel and distinctive aspect of tropical phyllosphere Communities is that Acidobacteria were one of the most abundant phyla across all samples (on average, 17%), a pattern not previously recognized. Sequences belonging to Acidobacteria were classified into subgroups 1–6 among known 24 subdivisions, and subgroup 1 (84%) was the most abundant group, followed by subgroup 3 (15%). The high abundance of Acidobacteria on leaves of tropical trees indicates that there is a strong relationship between host plants and Acidobacteria in tropical rain forest, which needs to be investigated further. The similarity of phyllosphere Bacterial Communities amongst the tree species sampled shows a significant tendency to follow host plant phylogeny, with more similar Communities on more closely related hosts.
Rob Knight - One of the best experts on this subject based on the ideXlab platform.
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Co-habiting amphibian species harbor unique skin Bacterial Communities in wild populations
The ISME Journal, 2012Co-Authors: Valerie J Mckenzie, Noah Fierer, Rob Knight, Robert M Bowers, Christian L LauberAbstract:Although all plant and animal species harbor microbial symbionts, we know surprisingly little about the specificity of microbial Communities to their hosts. Few studies have compared the microbiomes of different species of animals, and fewer still have examined animals in the wild. We sampled four pond habitats in Colorado, USA, where multiple amphibian species were present. In total, 32 amphibian individuals were sampled from three different species including northern leopard frogs ( Lithobates pipiens ), western chorus frogs ( Pseudacris triseriata ) and tiger salamanders ( Ambystoma tigrinum ). We compared the diversity and composition of the Bacterial Communities on the skin of the collected individuals via barcoded pyrosequencing of the 16S rRNA gene. Dominant Bacterial phyla included Acidobacteria , Actinobacteria , Bacteriodetes , Cyanobacteria , Firmicutes and Proteobacteria . In total, we found members of 18 Bacterial phyla, comparable to the taxonomic diversity typically found on human skin. Levels of Bacterial diversity varied strongly across species: L. pipiens had the highest diversity; A. tigrinum the lowest. Host species was a highly significant predictor of Bacterial community similarity, and co-habitation within the same pond was not significant, highlighting that the skin-associated Bacterial Communities do not simply reflect those Bacterial Communities found in their surrounding environments. Innate species differences thus appear to regulate the structure of skin Bacterial Communities on amphibians. In light of recent discoveries that some bacteria on amphibian skin have antifungal activity, our finding suggests that host-specific bacteria may have a role in the species-specific resistance to fungal pathogens.
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Spatial variability in airborne Bacterial Communities across land-use types and their relationship to the Bacterial Communities of potential source environments
The ISME Journal, 2011Co-Authors: Robert M Bowers, Rob Knight, Shawna Mcletchie, Noah FiererAbstract:Although bacteria are ubiquitous in the near-surface atmosphere and they can have important effects on human health, airborne bacteria have received relatively little attention and their spatial dynamics remain poorly understood. Owing to differences in meteorological conditions and the potential sources of airborne bacteria, we would expect the atmosphere over different land-use types to harbor distinct Bacterial Communities. To test this hypothesis, we sampled the near-surface atmosphere above three distinct land-use types (agricultural fields, suburban areas and forests) across northern Colorado, USA, sampling five sites per land-use type. Microbial abundances were stable across land-use types, with ∼10^5–10^6 Bacterial cells per m^3 of air, but the concentrations of biological ice nuclei, determined using a droplet freezing assay, were on average two and eight times higher in samples from agricultural areas than in the other two land-use types. Likewise, the composition of the airborne Bacterial Communities, assessed via bar-coded pyrosequencing, was significantly related to land-use type and these differences were likely driven by shifts in the sources of bacteria to the atmosphere across the land-uses, not local meteorological conditions. A meta-analysis of previously published data shows that atmospheric Bacterial Communities differ from those in potential source environments (leaf surfaces and soils), and we demonstrate that we may be able to use this information to determine the relative inputs of bacteria from these source environments to the atmosphere. This work furthers our understanding of Bacterial diversity in the atmosphere, the terrestrial controls on this diversity and potential approaches for source tracking of airborne bacteria.
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Bacterial Communities associated with the lichen symbiosis
Applied and Environmental Microbiology, 2011Co-Authors: Scott T Bates, Rob Knight, Garrett W G Cropsey, Gregory J Caporaso, Noah FiererAbstract:Lichens are commonly described as a mutualistic symbiosis between fungi and “algae” (Chlorophyta or Cyanobacteria); however, they also have internal Bacterial Communities. Recent research suggests that lichen-associated microbes are an integral component of lichen thalli and that the classical view of this symbiotic relationship should be expanded to include bacteria. However, we still have a limited understanding of the phylogenetic structure of these Communities and their variability across lichen species. To address these knowledge gaps, we used bar-coded pyrosequencing to survey the Bacterial Communities associated with lichens. Bacterial sequences obtained from four lichen species at multiple locations on rock outcrops suggested that each lichen species harbored a distinct community and that all Communities were dominated by Alphaproteobacteria. Across all samples, we recovered numerous Bacterial phylotypes that were closely related to sequences isolated from lichens in prior investigations, including those from a lichen-associated Rhizobiales lineage (LAR1; putative N2 fixers). LAR1-related phylotypes were relatively abundant and were found in all four lichen species, and many sequences closely related to other known N2 fixers (e.g., Azospirillum, Bradyrhizobium, and Frankia) were recovered. Our findings confirm the presence of highly structured Bacterial Communities within lichens and provide additional evidence that these bacteria may serve distinct functional roles within lichen symbioses.
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consistent effects of nitrogen fertilization on soil Bacterial Communities in contrasting systems
Ecology, 2010Co-Authors: Kelly S Ramirez, Noah Fierer, Rob Knight, Christian L Lauber, Mark A BradfordAbstract:: Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) through anthropogenic activities. Although the effects of increased N inputs on plant Communities have been reasonably well studied, few comparable studies have examined impacts on whole soil Bacterial Communities, though they play critical roles in ecosystem functioning. We sampled soils from two long-term ecological research (LTER) experimental N gradients, both of which have been amended with NH4NO3; a grassland at Cedar Creek (27 years of N additions) and an agricultural field at Kellogg Biological Station (8 years of N additions). By examining shifts in Bacterial Communities across these contrasting ecosystem types, we could test competing hypotheses about the direct and indirect factors that might drive Bacterial responses to elevated N inputs. Bacterial community structure was highly responsive to N additions. We observed predictable and consistent changes in the structure of the Bacterial Communities across both ecosystem types. Our results suggest that Bacterial Communities across these gradients are more structured by N and/or soil carbon availability than by shifts in the plant community or soil pH associated with the elevated nitrogen inputs. In contrast to the pronounced shifts in Bacterial community composition and in direct contrast to the patterns often observed in plant Communities, increases in N availability did not have consistent effects on the richness and diversity of soil Bacterial Communities.
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Forensic identification using skin Bacterial Communities.
Proceedings of the National Academy of Sciences of the United States of America, 2010Co-Authors: Noah Fierer, Cl Christian L Lauber, Nick Zhou, Elizabeth K. Costello, Daniel Mcdonald, Rob KnightAbstract:Recent work has demonstrated that the diversity of skin-associated Bacterial Communities is far higher than previously recognized, with a high degree of interindividual variability in the composition of Bacterial Communities. Given that skin Bacterial Communities are personalized, we hypothesized that we could use the residual skin bacteria left on objects for forensic identification, matching the bacteria on the object to the skin-associated bacteria of the individual who touched the object. Here we describe a series of studies de-monstrating the validity of this approach. We show that skin-associated bacteria can be readily recovered from surfaces (including single computer keys and computer mice) and that the structure of these Communities can be used to differentiate objects handled by different individuals, even if those objects have been left untouched for up to 2 weeks at room temperature. Furthermore, we demonstrate that we can use a high-throughput pyrosequencing-based ap-proach to quantitatively compare the Bacterial Communities on objects and skin to match the object to the individual with a high degree of certainty. Although additional work is needed to further establish the utility of this approach, this series of studies introduces a forensics approach that could eventually be used to independently evaluate results obtained using more traditional forensic practices.
Steven W Kembel - One of the best experts on this subject based on the ideXlab platform.
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relationships between phyllosphere Bacterial Communities and plant functional traits in a neotropical forest
Proceedings of the National Academy of Sciences of the United States of America, 2014Co-Authors: Timothy K Oconnor, Steven W Kembel, Holly K Arnold, Stephen P Hubbell, Joseph S Wright, Jessica L GreenAbstract:The phyllosphere—the aerial surfaces of plants, including leaves—is a ubiquitous global habitat that harbors diverse Bacterial Communities. Phyllosphere Bacterial Communities have the potential to influence plant biogeography and ecosystem function through their influence on the fitness and function of their hosts, but the host attributes that drive community assembly in the phyllosphere are poorly understood. In this study we used high-throughput sequencing to quantify Bacterial community structure on the leaves of 57 tree species in a neotropical forest in Panama. We tested for relationships between Bacterial Communities on tree leaves and the functional traits, taxonomy, and phylogeny of their plant hosts. Bacterial Communities on tropical tree leaves were diverse; leaves from individual trees were host to more than 400 Bacterial taxa. Bacterial Communities in the phyllosphere were dominated by a core microbiome of taxa including Actinobacteria, Alpha-, Beta-, and Gammaproteobacteria, and Sphingobacteria. Host attributes including plant taxonomic identity, phylogeny, growth and mortality rates, wood density, leaf mass per area, and leaf nitrogen and phosphorous concentrations were correlated with Bacterial community structure on leaves. The relative abundances of several Bacterial taxa were correlated with suites of host plant traits related to major axes of plant trait variation, including the leaf economics spectrum and the wood density–growth/mortality tradeoff. These correlations between phyllosphere Bacterial diversity and host growth, mortality, and function suggest that incorporating information on plant–microbe associations will improve our ability to understand plant functional biogeography and the drivers of variation in plant and ecosystem function.
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architectural design drives the biogeography of indoor Bacterial Communities
PLOS ONE, 2014Co-Authors: James F Meadow, Timothy K Oconnor, Gwynne Mhuireach, Dale Northcutt, Maxwell Moriyama, Jeff Kline, Steven W Kembel, G Z BrownAbstract:Background Architectural design has the potential to influence the microbiology of the built environment, with implications for human health and well-being, but the impact of design on the microbial biogeography of buildings remains poorly understood. In this study we combined microbiological data with information on the function, form, and organization of spaces from a classroom and office building to understand how design choices influence the biogeography of the built environment microbiome. Results Sequencing of the Bacterial 16S gene from dust samples revealed that indoor Bacterial Communities were extremely diverse, containing more than 32,750 OTUs (operational taxonomic units, 97% sequence similarity cutoff), but most Communities were dominated by Proteobacteria, Firmicutes, and Deinococci. Architectural design characteristics related to space type, building arrangement, human use and movement, and ventilation source had a large influence on the structure of Bacterial Communities. Restrooms contained Bacterial Communities that were highly distinct from all other rooms, and spaces with high human occupant diversity and a high degree of connectedness to other spaces via ventilation or human movement contained a distinct set of Bacterial taxa when compared to spaces with low occupant diversity and low connectedness. Within offices, the source of ventilation air had the greatest effect on Bacterial community structure. Conclusions Our study indicates that humans have a guiding impact on the microbial biodiversity in buildings, both indirectly through the effects of architectural design on microbial community structure, and more directly through the effects of human occupancy and use patterns on the microbes found in different spaces and space types. The impact of design decisions in structuring the indoor microbiome offers the possibility to use ecological knowledge to shape our buildings in a way that will select for an indoor microbiome that promotes our health and well-being.
Mohammad Bahram - One of the best experts on this subject based on the ideXlab platform.
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Bacterial Communities in boreal forest mushrooms are shaped both by soil parameters and host identity
Frontiers in Microbiology, 2017Co-Authors: Mari Pent, Kadri Poldmaa, Mohammad BahramAbstract:Despite recent advances in understanding the microbiome of eukaryotes, little is known about microbial Communities in fungi. Here we investigate the structure of Bacterial Communities in mushrooms, including common edible ones, with respect to biotic and abiotic factors in the boreal forest. Using a combination of culture-based and Illumina high-throughput sequencing, we characterized the Bacterial Communities in fruitbodies of fungi from eight genera spanning four orders of the class Agaricomycetes (Basidiomycota). Our results revealed that soil pH followed by fungal identity are the main determinants of the structure of Bacterial Communities in mushrooms. While almost half of fruitbody bacteria were also detected from soil, the abundance of several Bacterial taxa differed considerably between the two environments. The effect of host identity was significant at the fungal genus and order level and could to considerable extent be ascribed to the distinct Bacterial community of the chanterelle, representing Cantharellales – the earliest diverged group of mushroom-forming basidiomycetes. These data suggest that besides the substantial contribution of soil as a major taxa source and environmental determinant of Bacterial Communities in mushrooms, the structure of these Communities is to a great degree affected by the identity of the host. Thus, bacteria inhabiting fungal fruitbodies may be non-randomly selected from environment based on their symbiotic functions and/or habitat requirements.