A1 Refereed original research article in a scientific journal
Normative assembly rule reveals fairness in microbial communities; 
Authors: Kuosmanen, Teemu; Rantanen, Juhani; Kičiatovas, Dovydas; Pausio, Sanna; Friman, Ville-Petri; Hiltunen, Teppo; Mustonen, Ville
Editors: Koskella Britt
Publisher: Public Library of Science (PLoS)
Publication year: 2026
Journal: PLoS Biology
Article number: e3003872
Volume: 24
Issue: 6
eISSN: 1545-7885
DOI: https://doi.org/10.1371/journal.pbio.3003872
Publication's open availability at the time of reporting: Open Access
Publication channel's open availability : Open Access publication channel
Web address : https://doi.org/10.1371/journal.pbio.3003872
Self-archived copy’s web address: https://research.utu.fi/converis/portal/detail/Publication/527081776
Self-archived copy's licence: CC BY
Self-archived copy's version: Publisher`s PDF
Understanding and predicting how communities assemble is a paramount challenge in ecology. Here we address these questions normatively by comparing the observed species abundance distribution to a game-theoretically fair distribution based on each species’ Shapley value. By analyzing in total 56 distinct community outcomes, we assess how fairly biomass is distributed in microbial communities displaying both competitive and cooperative interactions in different growth conditions. We find examples of fair communities that closely follow their Shapley value across all environments as well as counterexamples where the true abundances deviate from the species’ objective contribution to community biomass. Next, we develop a fair assembly rule based on the recursive definition of Shapley value and show that also unfair community compositions are consistent with the principles of fair assembly after the lower-level competitive outcomes are known. Our results give unique empirical insights into the distributive function of ecological dynamics and lay down the theoretical foundations of what might become a normative community assembly theory.
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Funding information in the publication:
This work was funded in part by the Research Council of Finland (Multidisciplinary Center of Excellence in Antimicrobial Resistance Research, grants 346126 and 364232 to TH; grants 346128 and 364234 to VM). https://www.aka.fi/en/ The funders did not play any role in the study design, data collection and analysis, decision to publish, or preparation of the manuscript.