What Is the Real Unit of Selection? | Lisa Lloyd
Why It Matters
This typology sharpens conceptual tools for evolutionary theory, helping researchers and philosophers avoid category errors when attributing adaptation to genes, organisms, groups, or species, and guiding empirical and theoretical work on how complex adaptations arise. It matters for interpreting evolutionary data and for debates over which level should be the focus of selection-based explanations.
Summary
Philosopher Lisa Lloyd outlines a four-part framework for the long-running debate over the ‘‘unit of selection’’ in evolutionary biology: the reproducer (replicator subset that transmits traits), the interactor (the phenotype or target that interacts with the environment), the manifestor (the accumulated, engineering-style adaptations that build up over time), and the ultimate beneficiary (the long-term entity that gains from selection). She distinguishes simple selection-product changes—shifts in allele frequencies like industrial melanism—from true engineering adaptations that require cumulative, mechanistic construction. Lloyd traces how these distinctions clarify disputes over gene-, individual-, group- and species-level selection and builds on work by Dawkins, Hull, Griesemer, Gould and Williams. The framework reframes debates by separating processes (interaction and reproduction) from outcomes (manifested adaptations and beneficiaries).
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