Journal Pre-proof On the use of “life history theory” in evolutionary psychology Stephen C. Stearns, António M.M. Rodrigues PII: S1090-5138(20)30026-X DOI: https://doi.org/10.1016/j.evolhumbehav.2020.02.001 Reference: ENS 6308 To appear in: Evolution and Human Behavior Received date: 7 July 2019 Revised date: 3 February 2020 Accepted date: 12 February 2020 Please cite this article as: S.C. Stearns and A.M.M. Rodrigues, On the use of “life history theory” in evolutionary psychology, Evolution and Human Behavior(2020), https://doi.org/10.1016/j.evolhumbehav.2020.02.001 This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2020 Published by Elsevier. Journal Pre-proof On the use of “life history theory” in evolutionary psychology Stephen C. Stearns1 and António M.M. Rodrigues2 1Department of Ecology and Evolutionary Biology Yale University Box 208106 New Haven, CT 06520-8106 USA 2Department of Zoology University of Cambridge Downing Street Cambridge CB2 3EJ UK Email of corresponding author: [email protected] Declaration of interest: We have no financial or personal relationships with people or organizations that could bias this work. Journal Pre-proof Submission declaration: This work has not been published previously and is not under consideration elsewhere. On the use of “life history theory” in evolutionary psychology Journal Pre-proof Declaration of interest: We have no financial or personal relationships with people or organizations that could bias this work. Submission declaration: This work has not been published previously and is not under consideration elsewhere. Journal Pre-proof Journal Pre-proof Abstract We critically review the use of the term “life history theory” in recent publications on evolutionary psychology, focusing on how the idea of a fast-slow continuum is deployed in that literature. We raise four issues: First, concerning plasticity, should we expect the effects of plasticity on the developmental response of a trait to mirror the effects of selection on the mean of that trait? We conclude that we should not. Do only plastic responses to harsh or unpredictable environments accelerate maturation, or are there plausible alternatives, such as nutrition? In many situations better nutrition is a plausible alternative. Second, how should we conceive of the harshness of an environment? It has several important dimensions. It could mean an increase in the mean mortality rate, a decrease in the mean growth rate or fertility rate, or increases in the variances of any of those rates. Our judgement of harshness will also be affected by the distribution of such effects across patches in space and through generations in time. The combination and distribution of effects make important differences to predictions.Journal Pre-proof Third, where did the fast-slow idea come from, and how much does it explain? It was initially detected in comparisons across higher taxonomic levels, whose relevance to variation among individuals is unclear and where it fails to explain much of the variation. Journal Pre-proof Fourth, what sorts of processes could generate the fast-slow pattern? Here we expand on insights mentioned earlier in passing to make clear how spatial population structure and class effects generate alternative predictions. We conclude with some thoughts on the nature of theories and research strategies and on how one might respond to empirical puzzles. Keywords: life history theory; fast-slow; patterns; causes; research strategies Journal Pre-proof Journal Pre-proof 1. Introduction Because the term “life history theory” is increasingly used by some in evolutionary psychology to support explanations of developmental responses of human behavior, cognition, and physiology along a fast-slow continuum associated with psychosocial stress (Nettle and Frankenhuis 2019), this is a good time to take a step back to ask, what exactly does life history theory predict, and is the way in which it is deployed in evolutionary psychology appropriate? Specifically, we address these questions: 1. Should one expect the effects of plasticity to mirror the effects of selection on the mean of a trait? What effects can accelerate maturation? 2. How should we represent the harshness of an environment? Several combinations of the means and variances of age-specific vital rates are candidates. 3. Where did the idea of a fast-slow pattern come from, how much variation does it explain, and should we expect it to explain variation among individuals within populations? 4. What sorts of processes could each generate the fast-slow pattern? Spatial population structureJournal and class effects Pre-proof can both generate alternative predictions. These considerations reveal a general issue that arises when one borrows ideas from other fields and has surfaced in many, including our own. While borrowing can be powerful, it has pitfalls. Its power comes from a new point of view that suggests new questions – a precious commodity in science. The usual reaction is to analogize: this pattern in my field (e.g. inter-individual correlations among Journal Pre-proof traits related to behaviour, cognition, physiology, and psychosocial stress) is like that pattern in the other field (e.g. among-species correlations among life history traits), therefore what they found there should apply here. But analogies mislead if the underlying causes differ. Insights from analogies must be validated by showing that the predicted effects also result from tracing causes in the new context. That requires building and testing formal models that demonstrate that the borrowed predictions also arise naturally from the causal relationships of the new field. In formal models, careful attention to assumptions is important, for predictions are only meaningful if the assumptions hold. For example, in the population under study, can one demonstrate that individuals in which age at first birth is earlier are also those in which both psychosocial stress and risk of adult mortality are higher, or are they simply the ones who are better nourished? Why do people doing research on life history evolution find formal models useful? First, by setting down explicit assumptions and analyzing their consequences, it was, very early on, possible to distinguish between two major causal claims: r&K selection on the one hand, age-specific selection on the other. The clarifying effect of that experienceJournal shaped the field Pre-proof for decades to follow. Second, the interactions with various intrinsic tradeoffs of age-specific changes in fertility and mortality produced a complex conceptual structure that could only be handled with confidence by building mathematical models capable of keeping track of all those interactions. Third, many of the predictions depend on quantitative effects, for the costs and benefits of tradeoffs change both with trait values and with environmental gradients. Qualitative, verbal models cannot handle such effects, which can lead to major changes in predictions. Journal Pre-proof The quantitative nature of relationships is thus central. Drawing qualitative conclusions from a body of theory in which quantitative effects are important can be dangerous when predictions change qualitatively for quantitative reasons – as when the balance of benefits and costs in a trade-off shifts from positive to negative. While many of the tests of quantitative life history theory in biology have been done with qualitative comparisons, as have applications of life history theory in evolutionary psychology, there are some in which it has been possible to predict evolutionary trajectories quantitatively. For example, when Ackermann and colleagues (Ackermann et al. 2007) made quantitative predictions of changes in age at first birth and interbirth interval in bacteria undergoing experimental evolution, they found changes in age at first birth close to those predicted, but changes in interbirth interval much smaller than those predicted. This result suggested that interbirth interval is constrained in ways not previously expected and productively focused attention on why the mechanisms that determine reproductive rate are not easily changed by strong selection. Here the useful feature of the quantitative prediction was its capacity to fail in an interesting way. Those using life historyJournal theory to support Pre-proof claims in evolutionary psychology therefore need answers to these questions: What is assumed? How were the models built? When assumptions change among models, do predictions change? Is what is thought to be a general prediction in fact context-dependent? To be clear, we define the senses in which we use terms in a Glossary. [Glossary near here] Journal Pre-proof We next unpack the fast-slow continuum as it is understood in life history theory through answers to four expanded sets of questions: How do the reaction norms for age and size at maturity as functions of environmental quality evolve? Is the evolved mean population response to a deterioration in environmental