# Are the better cooperators dormant or quiescent?

## TL;DR

This paper investigates how different resting or dormancy times in species influence the evolution and stability of cooperation, revealing that cooperation persists only under specific timing conditions between cooperators and defectors.

## Contribution

It introduces a mathematical model linking resting stage durations to cooperation stability, showing that timing differences can promote or hinder cooperation.

## Key findings

- Cooperation cannot persist if cooperators and defectors have identical resting times.
- Distinct timing differences can stabilize cooperation as an evolutionarily stable strategy.
- The model applies to microbial, invertebrate, and plant populations with resting stages.

## Abstract

Despite the wealth of empirical and theoretical studies, the origin and maintenance of cooperation is still an evolutionary riddle. In this context, ecological life-history traits which affect the efficiency of selection may play a role, though these are often ignored. We consider here species such as bacteria, fungi, invertebrates and plants which exhibit resting stages in the form of a quiescent state or a seedbank. When quiescent, individuals are inactive and reproduce upon activation, while under seed bank parents produce offspring remaining dormant for different amount of time. We assume weak frequency-dependent selection modeled using game-theory and the prisoners dilemma (cooperation/defect) as payoff matrix. The cooperators and defectors are allowed to evolve different quiescence or dormancy times. By means of singular perturbation theory we reduce the model to a one-dimensional equation resembling the well known replicator equation, where the gain functions are scaled with lumped parameters reflecting the time scale of the resting state of the cooperators and defectors. If both time scales are identical cooperation cannot persist in a homogeneous population. If, however, the time scale of the cooperator is distinctively different from that of the defector, cooperation may become a locally asymptotically stable strategy. Interestingly enough, in the seedbank case the cooperator needs to be faster than the defector, while in the quiescent case the cooperator has to be slower. We use adaptive dynamics to identify situations where cooperation may evolve and form a convergent stable ESS. We conclude by highlighting the relevance fo these results for many non-model species and the maintenance of cooperation in microbial, invertebrate or plant populations.