2nd Review of the paper 

“Easy, fast and reproducible Stochastic Cellular Automata with chouca”
by Alexandre Génin, Guillaume Dupont, Daniel Valencia, Mauro Zucconi, M. Isidora Ávila-Thieme, Sergio Navarrete, and Evie A. Wieters

Available at
https://www.biorxiv.org/content/10.1101/2023.11.08.566206v5.full.pdf

The authors present a software which makes the application of a particular type of Cellular Automata (CA) easier. It departs from the R (statistics software) environment and allows a subsequent C++ compilation (backend). Thus, larger grids can be run faster compared to a model that uses an interpreted language like R. The rule specification of models using the described software is limited to an operation of 2d rectangular grids. Cell state transition rules can use probability functions depending on the cells own state, the state of a 4-cell or 8-cell neighbourhood, and depending on the overall grid state. The current implementation would exclude descriptions of particle movement processes (like a random version of Langton’s ant). Thus, there remains a considerable development potential to expand the applicability of the approach to a wider range of ecologically relevant interaction types that can be captured in other modelling environments. 

The paper is well written and publishable in the present form. It is clear and understandable in particular to those, who are not programmers but who could be interested in an application of the software to model ecological interactions. The revision the authors considered the comments of the previous review in a reasonable way. The following statements explain the reviewer’s view and are not necessarily requirements for further update.

Some details of the paper to be looked at:

The abstract does not fully inform about the limitations of the approach. Readers may be disappointed to learn in the text, that (random) movement processes cannot be run in the current implementation. This excludes a wide range of stochastic ecological CA applications. It would be reasonable to clarify this already in the abstract.

The keywords could be more informative by adding “stochastic” to cellular automata and also mention the name of the software.

The limitation of the software to rectangular grids has the problem, that for many processes, discretisation artefacts prevail, which cannot be avoided by using a higher grid resolution. For an analysis of the relevance of results compared to empirical observations, it would be reasonable to confirm findings with simulations using an alternative grid topology, e.g. triangular, hexagonal, or even a Voronoi tessellation of a simulated area. This could be a further development perspective.

The paper states in line 101 ff: “Probabilities of transition are assumed to depend only on (i) the proportion of neighbors in each state, captured by the vector 𝒒 = (𝑞1, . . . , 𝑞𝑛), (ii) the proportion of cells in a given state in the whole landscape, 𝒑 = (𝑝1, . . . , 𝑝𝑛), and (iii) a set of constant model parameters 𝜽.  chouca has been primarily designed for modeling the dynamics of sessile organisms over space, which do not move and reproduce through the dispersal of propagules.”
Comment: this is the scope of the _gap-model_ approach, elaborated e.g. by Shugart in the 1980ies to study ecological succession dynamics. The term is widely known in ecology as a model type. It might be useful to add that term (see e.g. Shugart, H. H., & West, D. C. 1981. Long-term dynamics of forest ecosystems: Computer simulation models, which allow for numerous seedlings and the long lives of large trees, predict how forests will respond to different management techniques. American Scientist, 69(6), 647-652.)

Line 135 ff: The paper explains an example application description to illustrate the supported interaction types. It uses a transition between 3 different cell states. The example is presented very clear and well understandable. 

In the part “graphical explorations” (line 227 ff), the authors show another 3-state model (host – parasitized- empty). This is a special case of an excitable media pattern where the according states are usually named excitable, excited, and refractory. The underlying interaction process occurs in self-organisation dynamics in various contexts not only in ecology but also in physiology, physics, and chemistry. To inform readers that this interaction type can be run with the software, it would be useful to include the term “Excitable Media” in the text.

The paper states in line 269 ff: “Because SCA are defined on grids, a natural application is to compare their output to empirical raster data, such as remote-sensing images, to infer local-scale ecological interactions from landscape-wide spatial patterns”
Comment: such a comparison can be biased because of CA discretisation artefacts. On a rectangular grid with a 4-cell neighbourhood processes in vertical or horizontal direction require less iteration steps than those in diagonal direction towards the same distance. This can influence resulting simulation pattern and thus the comparison with empirical data (depending on rule-specification, the effect might be less pronounced with an 8-cell neighbourhood). Practical applications require a consideration of discretisation effects. It could be useful to point to this requirement when discussion comparisons with empirical pattern.

The paper states in line 315 ff: “Applying such an approach to empirical data would require further testing of model assumptions, for example, to investigate whether facilitation occurs on the recruitment of new plants instead of on the mortality of adult plants – this can be done by simply changing the model definition above.” 
Comment: the authors use the described model to explain, how the search for a parameter set can be done that approximates an observed pattern best. They show how a pattern comparison with systematically varied parameter identifies the size of 2 parameter that were used to generate the target pattern. The demonstration is suitable and understandable, however, from an ecological perspective, the model is questionable due to simplification but illustrates the pattern evaluation procedure well. 

As a positive point it has to be mentioned, that the connection to R can help in various statistical evaluations. The authors state in line 348 ff that the “code used for this work is freely-available at Zenodo under a CC-BY license (Génin, 2024), along with chouca version v0.1.99 used at the time of writing”. This is a good condition to work with the software package and an achievement of the authors with regard to the relevance of their contribution.

Génin et al present a framework to implement a kind of simplistic cellular automata with probabilitstic state transitions. They use illustrative ecological examples to show the functionality. Because they apply R as a computational framework, it can be expected that it makes the access for users easier compared to genuine programming. However, the approach is syntactically rather limited and excludes important ecological interaction types, e.g. it operates on 2d rectangular grids only and excludes direct cell-to-cell relation that would allow 1:1 mutual state change of cells. Besides specific application cases, its benefit could be in the introduction to spatial modelling in a didactical context. For research applications, in most practical cases a higher flexibility and complexity in rule-specification might be required.

In the following, comments to particular statements in the text are added. Because the paper does not provide line numbers, the commented sentences are copied below.

Specific comments

The paper states
“Stochastic cellular automata (SCA) are models that describe spatial dynamics using a grid of cells that switch between discrete states over time, depending only on the current state (Markovian processes).”
Comment
This is a rather narrow specification which excludes an application for many practical requirements. If for the characterisation of a cell’s state continuous variables are excluded, many relevant processes cannot be modelled.g. the concentration of a substance, continuous variation in pressure, temperature, etc. at a particular grid position. This is in particular relevant for ecological applications.

The paper states
“de novo implementation of SCA for each specific system and application represents a major barrier for many practitioners.”
Comment
It is questionable if this is actually a barrier. Even without object-oriented programming, it is not such a challenge to develop a CA. For a rectangular CA with a von Neumann neighbourhood, two nested FOR-loops together with conventional calculation would do as a main component to set the layout for an iteration cycle.

The paper states
“we built chouca, an R package that translates intuitive SCA model definitions into compiled code,”
Comment
this may be a major advantage compared to interpreted languages. The authors present a figure (Fig. 3) which compares runtimes of different grid size and indicate a considerable efficiency.

The paper states
“ad hoc implementations found in the literature,”
Comment
here, a reference would be useful to clarify which implementations the authors have in mind 

The paper states
“Conway’s game of life,”
Comment
a reference would be nice

The paper states
“The probabilities of a cell switching from one state to another is assumed to depend on model parameters, the global state of the system (the proportion of cells in each state), and the local neighborhood of the focal cell.”
Comment
Frequently, the change of the cell’s states in CAs are considered to depend on the state of the cell itself and its neighbours. The specification here, expands that and offers to operate on two levels of neighbourhood – a local and a global one. This is possible, however, the evaluation of the entire grid state to specify a new cells state is usually not required as a CA standard. Nevertheless, under particular condition, this could be an interesting option.

The paper states
“In many cases, the explicit numerical simulation must be run, which is often done on small grids to reduce computation time.”
Comment
What do the authors understand as a “small grid”? Since computational potentials expand with technical developments, this statement is relative and tends to lose relevance. Is the grid size stated below (p.7: “a typical 128 x 128 grid (Figure 3)” a small one?

The paper states
“This opens the possibility of errors in code and often makes it difficult to reproduce model simulations.”
Comment
this is a widespread issue throughout ecological modelling. It may be a useful contribution of this approach, if for this specific segment in modelling, result reproduction may become easier.

The paper states
The R package chouca works with 2-dimensional rectangular grids of cells (a “landscape”).
Comment
In an upcoming version, the authors should expand it to 3d grids, then allowing to represent also waterbodies or processes referring to atmospheric layers. 
A further step then could be to facilitate the use of triangular or hexagonal grids. To have an alternative to rectangular grids would allow for tests of the extent of discretisation artefacts – which are quite relevant for rectangular grids with regard to ecological processes.

The paper states
“It is important to note that this excludes cellular automata in which an intermediate distance of interaction is considered (e.g. through a dispersion kernel; Muthukrishnan et al., 2016), or those in which a preferential direction exists (e.g. modeling water redistribution on a slope; Mayor et al., 2013). Other types of SCA not fitting these constraints are those in which two cells swap their respective state, e.g. when modelling the movements of a predator in a landscape (Pascual et al., 2002)” … “In practice, this functional form is flexible enough to approximate the probabilities of transition of many ecological models”
Comment
this excludes a very large extend (if not the majority) of potential ecological CA applications. If not even the change of a state’s cell with its neighbour could be represented, then the whole class of pattern generating processes like Diffusion Limited Aggregation as well as a majority of trophic interaction modelling would be excluded. 

The paper states
“Arid systems provide a good illustration of this approach: in those systems, plants often facilitate each other, which results in their aggregation into patches, and has important consequences for the resilience of those systems to changes in aridity”
Comment
even more often, in arid systems plants limit each other through competition for water access and give rise to gaps in vegetation cover. Facilitation is a process that helps in the survival of seedlings and is usually only a part of the relevant interactions. To describe plant dynamics under arid conditions might be more convincing by using “inhibition” rather than “facilitation”…