Abundant empirical evidence of multilevel selection revealed by a bibliometric review

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Abstract

Natural selection is based on the notion of differential reproduction between entities, often characterized as a struggle between individual organisms. However, natural selection can act at all levels of biological organization, thus being termed ‘multilevel selection’ (MLS). A common misconception is that MLS lacks empirical support. To address this, we conducted a bibliometric review of 2,950 Web of Science/Scopus-indexed scientific articles. Our goal was documenting the range of taxa/systems, levels, and research topics/tools where MLS has been used to understand natural selection across levels. We found 280 studies providing empirical support for MLS: 100 were performed in situ, 180 were laboratory experiments. The studies span a vast range of organisms, from viruses to humans and eusocial insects. While 90.4% of studies focused on some form of organismal group (demes, colonies, aggregates), the remaining 9.6% explored selection at other levels (communities, cells, nuclei). We classified these 280 studies into research categories such as artificial selection, breeding through group selection, indirect/social genetic effects, and contextual analysis, among others. In contextual analysis studies, the strength of selection was comparable across levels. Contrary to common notions, there is solid empirical support for the utility and importance of MLS in explaining natural selection and evolution.
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Abstract

Natural selection is based on the concept of differential reproduction between entities, often characterized as a struggle between individual organisms. However, natural selection can act at all levels of biological organization, thus being termed “multilevel selection” (MLS). A common misconception is that selection across levels of biological organization lacks empirical support. To address this, we conducted a bibliometric review of 2,950 Web of Science/Scopus-indexed scientific articles, to document the range of taxa and research topics where MLS has been used to understand natural selection across levels. The 280 studies providing empirical support for selection at more than one level spanned a vast range of organisms, from viruses to humans to eusocial insects. They included research done both in natural populations (100) and in laboratory experiments (180). While 90.4% of studies focused on selection among organismal groups (e.g., demes, colonies, aggregates), another 9.6% explored selection across other levels (genetic elements, nuclei, cells, or multispecies communities). We classified studies by topic including artificial selection, breeding through group selection, indirect genetic effects, and contextual analysis, among others. Contrary to common notions, we found solid empirical support for the utility and importance of MLS in explaining natural selection and evolution. DOI https://doi.org/10.32942/X25S84 Subjects Life Sciences

Keywords

animal and plant breeding, artificial selection, contextual analysis, Epistasis, group selection, units of selection Dates Published: 2025-07-31 16:04 Last Updated: 2026-02-20 18:20 Older Versions License CC-By Attribution-NonCommercial-NoDerivatives 4.0 International Additional Metadata Conflict of interest statement: The authors declare no conflict of interest. Data and Code Availability Statement: The database used for this Review is available at Zenodo: https://doi.org/10.5281/zenodo.16633276 Language: English

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