The Overview, Design concepts and Details (ODD) protocol for describing Individual-and Agent-Based Models (ABMs) is now widely accepted and used to document such models in journal articles. As a standardized document for providing a consistent, logical and readable account of the structure and dynamics of ABMs, some research groups also find it useful as a workflow for model design. Even so, there are still limitations to ODD that obstruct its more widespread adoption. Such limitations are discussed and addressed in this paper: the limited availability of guidance on how to use ODD; the length of ODD documents; limitations of ODD for highly complex models; lack of su icient details of many ODDs to enable reimplementation without access to the model code; and the lack of provision for sections in the document structure covering model design rationale, the model's underlying narrative, and the means by which the model's fitness for purpose is evaluated. We document the steps we have taken to provide better guidance on: structuring complex ODDs and an ODD summary for inclusion in a journal article (with full details in supplementary material; Table ); using ODD to JASSS, ( ) , http://jasss.soc.surrey.ac.uk/ / / .html Doi: . /jasss.point readers to relevant sections of the model code; update the document structure to include sections on model rationale and evaluation. We also further advocate the need for standard descriptions of simulation experiments and argue that ODD can in principle be used for any type of simulation model. Thereby ODD would provide a lingua franca for simulation modelling.

Movement behavior is an essential element of fundamental ecological processes such as competition and predation. Although intraspecific trait variation (ITV) in movement behaviors is pervasive, its consequences for ecological community dynamics are still not fully understood. Using a newly developed individual-based model, we analyzed how given and constant ITVs in foraging movement affect differences in foraging efficiencies between species competing for common resources under various resource distributions. Further, we analyzed how the effect of ITV on emerging differences in competitive abilities ultimately affects species coexistence. The model is generic but mimics observed patterns of among-individual covariation between personality, movement and space use in ground-dwelling rodents. Interacting species differed in their mean behavioral types along a slow-fast continuum, integrating consistent individual variation in average behavioral expression and responsiveness (i.e. behavioral reaction norms). We found that ITV reduced interspecific differences in competitive abilities by 5-35% and thereby promoted coexistence via an equalizing mechanism. The emergent relationships between behavioral types and foraging efficiency are characteristic for specific environmental contexts of resource distribution and population density. As these relationships are asymmetric, species that were either 'too fast' or 'too slow' benefited differently from ITV. Thus, ITV in movement behavior has consequences for species coexistence but to predict its effect in a given system requires intimate knowledge on how variation in movement traits relates to fitness components along an environmental gradient.

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The pace-of-life-syndrome (POLS) hypothesis posits that suites of traits are correlated along a slow-fast continuum due to life-history trade-offs. Despite widespread adoption, environmental conditions driving the emergence of POLS remain unclear. A recently proposed conceptual framework of POLS suggests that a slow-fast continuum should align to fluctuations in densitydependent selection. We tested three key predictions made by this framework with an ecoevolutionary agent-based population model. Selection acted on responsiveness (behavioral trait) to inter-patch resource differences and the reproductive investment threshold (life-history trait).Across environments with density fluctuations of different magnitudes, we observed the emergence of a common axis of trait covariation between and within populations, i.e. the evolution of a POLS. Slow-type (fast-type) populations with high (low) responsiveness and low (high) reproductive investment threshold were selected at high (low) population densities and less (more) intense and frequent density fluctuations. In support of the predictions, fast-type populations contained a higher degree of variation in traits and were associated with higher intrinsic reproductive rate (r0) and higher sensitivity to intraspecific competition (γ), pointing to a universal trade-off. While our findings support that POLS aligns with density-dependent selection, we discuss possible mechanisms which may lead to alternative evolutionary pathways.