Paper · 2025 · ISME Communications

Assembly and co-occurrence networks of nitrogen-fixing bacteria associated with epiphyllous liverworts in fragmented tropical forests

Adriel M Sierra; Dennis Alejandro Escolástico-Ortiz; Charles E Zartman; et al.

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citations
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Open access · cc-by

Abstract

Abstract Understanding the spatial dynamics of plant-associated microbial communities is increasingly urgent in the context of habitat loss and the biodiversity crisis. However, the influence of reduced habitat size and connectivity on the assembly mechanisms underlying microbial associations is fundamental to advancing microbial ecology and conservation. In the Brazilian Amazon, we investigated nitrogen-fixing (diazotrophic) bacterial communities associated with two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida, across 11 forest sites within the Biological Dynamics of Forest Fragments Project landscape. Using amplicon sequencing targeting the nitrogenase gene (nifH), we characterized diazotroph community diversity, inferred assembly mechanisms through null models, and analyzed co-occurrence network structure. Host-specific associations were evident: C. surinamensis predominantly hosted Hassallia, while R. flaccida was primarily associated with Fischerella. Despite habitat fragmentation, diazotrophic richness and composition remained similar across habitats of different sizes, consistent with strong homogenizing dispersal. Network analyses revealed that smaller fragments harbored more modular communities with fewer module hubs, pronounced shifts in key species relative abundance, and reduced network robustness. Our findings underscore the influence of habitat size on the stability of liverwort-associated diazotrophs, with smaller fragments exhibiting lower key species specificity and disruption of microbe-microbe interactions. Our results emphasize the importance of conserving large, connected forest habitats to maintain the functional integrity of phyllosphere N-fixing microbiota.

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The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 10

Observationobservedoriginal resultmeasured

Claim · model-written summary

Cololejeunea surinamensis predominantly hosted Hassallia diazotrophs.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Interactor
HassalliaHassalliae1symbiont
Outcome
predominantly hostede1

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e1

“C. surinamensis predominantly hosted Hassallia”

✓ located verbatim in the source · abstract, characters 887–933 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Radula flaccida was primarily associated with Fischerella diazotrophs.

Taxon
Radula flaccidaRadula flaccidae0
Interactor
FischerellaFischerellae2symbiont
Outcome
primarily associatede2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Fischerella
Fischerella · InteractorYoannis Domínguez · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e2

“R. flaccida was primarily associated with Fischerella”

✓ located verbatim in the source · abstract, characters 941–994 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Diazotrophic richness and composition remained similar across habitats of different sizes despite habitat fragmentation.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
habitat fragmentatione3
Outcome
diazotrophic richness and compositione3
Direction
remained similare3
Setting
across habitats of different sizese3

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e3

“Despite habitat fragmentation, diazotrophic richness and composition remained similar across habitats of different sizes”

✓ located verbatim in the source · abstract, characters 996–1116 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments harboured more modular liverwort-associated diazotroph networks.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse4
Outcome
modular communitiese4
Direction
moree4

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e4

“smaller fragments harbored more modular communities”

✓ located verbatim in the source · abstract, characters 1196–1247 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments had fewer module hubs in the diazotroph networks.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse5
Outcome
module hubse5
Direction
fewere5

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e5

“smaller fragments harbored more modular communities with fewer module hubs”

✓ located verbatim in the source · abstract, characters 1196–1270 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments showed reduced network robustness.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse6
Outcome
network robustnesse6
Direction
reducede6

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e6

“smaller fragments harbored more modular communities with fewer module hubs, pronounced shifts in key species relative abundance, and reduced network robustness”

✓ located verbatim in the source · abstract, characters 1196–1355 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments showed pronounced shifts in key species relative abundance.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse6
Outcome
key species relative abundancee6
Direction
pronounced shiftse6

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e6

“smaller fragments harbored more modular communities with fewer module hubs, pronounced shifts in key species relative abundance, and reduced network robustness”

✓ located verbatim in the source · abstract, characters 1196–1355 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments had lower key species specificity.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse7
Outcome
key species specificitye7
Direction
lowere7

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e7

“smaller fragments exhibiting lower key species specificity and disruption of microbe-microbe interactions”

✓ located verbatim in the source · abstract, characters 1470–1575 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Smaller fragments showed disruption of microbe-microbe interactions.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Treatment
smaller fragmentse7
Outcome
microbe-microbe interactionse7
Direction
disruptione7

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e7

“smaller fragments exhibiting lower key species specificity and disruption of microbe-microbe interactions”

✓ located verbatim in the source · abstract, characters 1470–1575 · show in text ↑
Mechanismauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Similar diazotroph richness and composition across habitat sizes is consistent with strong homogenizing dispersal.

Taxon
Cololejeunea surinamensisCololejeunea surinamensise0
Outcome
diazotrophic richness and compositione3
Qualifier
strong homogenizing dispersale7

Verbatim

e0

“two epiphyllous liverworts, Cololejeunea surinamensis and Radula flaccida”

✓ located verbatim in the source · abstract, characters 468–541 · show in text ↑
e3

“diazotrophic richness and composition remained similar across habitats of different sizes”

✓ located verbatim in the source · abstract, characters 1027–1116 · show in text ↑
e7

“consistent with strong homogenizing dispersal”

✓ located verbatim in the source · abstract, characters 1118–1163 · show in text ↑