The DOI matches its Crossref record; 8 of the 8 references it cites were found in a registry or an index.
Capturing the superorganism: a formal theory of group adaptation
Abstract
Adaptation is conventionally regarded as occurring at the level of the individual organism. However, in recent years there has been a revival of interest in the possibility for group adaptations and superorganisms. Here, we provide the first formal theory of group adaptation. In particular: (1) we clarify the distinction between group selection and group adaptation, framing the former in terms of gene frequency change and the latter in terms of optimization; (2) we capture the superorganism in the form of a ‘group as maximizing agent’ analogy that links an optimization program to a model of a group‐structured population; (3) we demonstrate that between‐group selection can lead to group adaptation, but only in rather special circumstances; (4) we provide formal support for the view that between‐group selection is the best definition for ‘group selection’; and (5) we reveal that mechanisms of conflict resolution such as policing cannot be regarded as group adaptations.
Overview
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Citations
Cites
The genetical theory of natural selection
Fisher, R. A. (1930) · doi:10.5962/bhl.title.27468 · Cited by 17,617
The genetical evolution of social behaviour. I
Hamilton, W. D. (1964) · Journal of Theoretical Biology · doi:10.1016/0022-5193(64)90038-4 · Cited by 15,725
Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection.
WEST, S. A., GRIFFIN, A. S., & GARDNER, A. (2007) · Journal of Evolutionary Biology · doi:10.1111/j.1420-9101.2006.01258.x · Cited by 1,889
Self-citationWilson, D. S. (1975) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.72.1.143 · Cited by 1,259
The major evolutionary transitions
Szathmáry, E., & Smith, J. M. (1995) · Nature · doi:10.1038/374227a0 · Cited by 1,095
Bacteria as Multicellular Organisms
Shapiro, J. A. (1988) · Scientific American · doi:10.1038/scientificamerican0688-82 · Cited by 524 · cited 2×
“‘superorganisms’ (Wheeler, 1911; Marais, 1925; Wells et al., 1929; Emerson, 1932; Wynne-Edwards, 1962, 1993; Seeley, 1989, 1997; Wilson & Sober, 1989; Ho¨lldobler & Wilson, 1990, 2009; Shapiro & Dworkin, 1997; Shapiro, 1998; Sober & Wilson, 1998; Reeve & Ho¨lldobler, 2007; Wilson & Wilson, 2007).”
Relatedness and the fraternal major transitions
Queller, D. C. (2000) · Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rstb.2000.0727 · Cited by 301
Hamilton, W. D. (2001) · doi:10.1093/oso/9780198503361.001.0001 · Cited by 246
Cited by
Cooperation between non-kin in animal societies
Clutton-Brock, T. (2009) · Nature · doi:10.1038/nature08366 · Cited by 951
Beyond society: the evolution of organismality
Queller, D. C., & Strassmann, J. E. (2009) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2009.0095 · Cited by 440
Major evolutionary transitions in individuality
West, S. A., Fisher, R. M., Gardner, A., & Kiers, E. T. (2015) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1421402112 · Cited by 431
Self-citationApplying evolutionary biology to address global challenges
Carroll, S. P., Jørgensen, P. S., Kinnison, M. T., Bergstrom, C. T., Denison, R. F., Gluckman, P., Smith, T. B., Strauss, S. Y., & Tabashnik, B. E. (2014) · Science · doi:10.1126/science.1245993 · Cited by 314
Boomsma, J. J., & Gawne, R. (2018) · Biological Reviews · doi:10.1111/brv.12330 · Cited by 304
Metadata
- Authors
- GARDNER, A., & GRAFEN, A.
- Venue
- Journal of Evolutionary Biology
- Publisher
- Oxford University Press (OUP)
- Published
- 2009-03-13
- Volume
- 22
- Issue
- 4
- Pages
- 659-671
- ISSN
- 1010-061X (ISSN-L)
- 1420-9101
- OpenAlex
- W2121396649
- Cited by
- 362
- References
- 8
- FWCI
- 82.49
- Type
- journal article
- Language
- en
- Open access
- closed
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review
How to cite
Cite CoChat for the checks on this page, and the paper for its own findings.
- This page
- CoChat. (2026). Citation check: Capturing the superorganism: a formal theory of group adaptation. https://app.cochat.ai/papers/10.1111/j.1420-9101.2008.01681.x
- The paper
- GARDNER, A., & GRAFEN, A. (2009). Capturing the superorganism: a formal theory of group adaptation. Journal of Evolutionary Biology. https://doi.org/10.1111/j.1420-9101.2008.01681.x
Data sources
Bibliographic data: Semantic Scholar, OpenAlex, Crossref.
Contains information from the Semantic Scholar Academic Graph Datasets, made available under the ODC Attribution License (ODC-BY 1.0).
Semantic Scholar’s data is described in: Kinney, R., et al. (2023). The Semantic Scholar Open Data Platform. arXiv:2301.10140.