SYSTEMATIC REVIEW article
Front. Ecol. Evol.
Sec. Evolutionary Ecology of Social Behavior
Abundant empirical evidence of multilevel selection revealed by a bibliometric review
Provisionally accepted
- 1Santo Tomás University, Santiago, Chile
- 2Vrije Universiteit Amsterdam, Amsterdam, Netherlands
- 3Binghamton University, Binghamton, United States
- 4University of California Santa Barbara, Santa Barbara, United States
- 5Nova Southeastern University, Fort Lauderdale, United States
- 6Indiana University, Bloomington, United States
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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.
Keywords: artificial selection, Breeding, contextual analysis, Cultural Evolution, indirect genetic effects, multicellularity, Units of selection, Group selection
Received: 23 Nov 2025; Accepted: 05 Jan 2026.
Copyright: © 2026 Marín, Clark, Philson, Eldakar and Wade. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
* Correspondence: César Marín
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