PCSY-D-25-00047

Mosquito Dispersal in Context

PLOS Complex Systems

Dear Dr. Smith,

Thank you for submitting your manuscript to PLOS Complex Systems. After careful consideration, we feel that it has merit but does not fully meet PLOS Complex Systems's publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Please submit your revised manuscript within 60 days Oct 07 2025 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at complexsystems@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pcsy/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

Please include the following items when submitting your revised manuscript:

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If you would like to make changes to your financial disclosure, competing interests statement, or data availability statement, please make these updates within the submission form at the time of resubmission. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

We look forward to receiving your revised manuscript.

Kind regards,

C. Brandon Ogbunugafor

Academic Editor

PLOS Complex Systems

C. Brandon Ogbunugafor

Academic Editor

PLOS Complex Systems

Hocine Cherifi

Editor-in-Chief

PLOS Complex Systems

Journal Requirements:

If the reviewer comments include a recommendation to cite specific previously published works, please review and evaluate these publications to determine whether they are relevant and should be cited. There is no requirement to cite these works unless the editor has indicated otherwise.

Additional Editor Comments (if provided):

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Reviewers' Comments:

Reviewer's Responses to Questions

-->Comments to the Author

1. Does this manuscript meet PLOS Complex Systems’s publication criteria? Is the manuscript technically sound, and do the data support the conclusions? The manuscript must describe methodologically and ethically rigorous research with conclusions that are appropriately drawn based on the data presented.-->

Reviewer #1: Yes

Reviewer #2: Partly

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-->2. Has the statistical analysis been performed appropriately and rigorously?-->

Reviewer #1: N/A

Reviewer #2: N/A

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-->3. Have the authors made all data underlying the findings in their manuscript fully available (please refer to the Data Availability Statement at the start of the manuscript PDF file)?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception. The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.-->

Reviewer #1: Yes

Reviewer #2: Yes

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-->4. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS Complex Systems does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.-->

Reviewer #1: Yes

Reviewer #2: Yes

**********

-->5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)-->

Reviewer #1: 1 Overall Impression

The manuscript titled “Mosquito Dispersal in Context” by S´anchez et al. presents a compelling and timely contribution

to the field of vector ecology and spatial epidemiology. The authors propose a novel agent-based microsimulation

framework that models mosquito movement as a sequence of transitions between biologically relevant behavioral

states, including sugar-feeding, blood-feeding, resting, and oviposition. This behaviorally-informed approach stands

in contrast to traditional dispersal models, which often reduce mosquito movement to random walks or isotropic

diffusion processes.

The simulation framework is implemented in a new R package, ramp.micro, which facilitates the exploration of

how landscape configuration, resource availability, and behavioral heterogeneity shape emergent dispersal patterns.

By incorporating empirically motivated state-transition rules and realistic spatial resource distributions, the model

enables researchers to study how mosquito populations interact with fragmented environments and encounter natural

or artificial barriers.

Key findings include the observation that behavioral heterogeneity can result in complex, emergent spatial patterns

that are not well predicted by simpler movement models. For instance, the authors show that mosquitoes aggregate

into behavioral “communities” around resource-rich areas, and that the permeability of landscape barriers is highly

context-dependent, varying with the internal state of the mosquito and spatial distribution of key resources.

The manuscript is well-written, and the modeling rationale is biologically grounded. The figures and simulation

outputs are informative and illustrate key concepts effectively. The approach is extensible and has potential applications

not only for fundamental ecological questions but also for operational planning in vector control.

In sum, the work is conceptually strong, methodologically innovative, and clearly fills a gap in the current modeling

literature by integrating mosquito behavior with spatial dynamics in a flexible and transparent way.

Major point

Clarity of mathematical formulation: In general, the derivation of the equations is not clearly presented. The

authors should either provide appropriate citations or include a more detailed explanation of the relationships between

variables and the derivation process itself. This would enhance clarity and support readers in understanding the

core concepts underlying the simulation. This issue is apparent in both the main text and the Methods section. If

constrained by word limits, detailed derivations could be provided in an appendix to improve readability without

interrupting the main narrative.

Minor points

• Notation and definitions: The state transition diagram contains numerous notations, some of which (e.g.,

σf ) are not clearly introduced or defined at the point of use. To improve clarity, the authors should consider

including a table that summarizes the primary notations and symbols used throughout the manuscript.

• Definitions: Key concepts such as ”dispersal kernels” (e.g., Line 137) lack sufficient explanation (may refer

it to method section). In the section titled “Resource Landscapes and Dispersal Kernels,” formal definitions

of both ”resource landscape” and ”dispersal kernel” may need to be provided in the context of this model, or

appropriate citations may need to be added if these terms are drawn from existing literature.

Reviewer #2: The authors propose an agent-based model of mosquito dispersal and present results suggesting that population-level movement patterns are primarily driven by intrinsic behavioral dynamics, rather than by simple spatial proximity or random movement alone. The motivation is compelling and the model is interesting; however I have several comments before the paper can be accepted for publication:

[-] The organization and compartmentalization of the information in the manuscript is not optimal giving the impression that there are repeated sections in Results and Methods.

The results section should focus only on providing the main results of the paper. If the model is itself a result it should be described more in detail. For instance, the authors could provide a subsection on the result section regarding model formulation: where you describe (1) Intrinsic behavioral model: describing the compartmental model of mosquito behavioral states, (2) Landscape and Transition model: Here you can provide a clear model of a discrete spatial domain, composed of distinct resource point sets (e.g., blood, egg-laying, sugar sites), and that mosquito dispersal is modeled using a Markov chain framework and describe in detail the matrices Phi…. etc...

In contrast, the subsections “Movement to Complete a Task” and “Population Structure” among others read more like methodological tools used to analyze or interpret the model’s behavior, rather than primary results. These sections would be more appropriately placed in the Methods section, as they describe techniques for quantifying and interpreting dispersal and spatial organization.

[-] Please use more standard mathematical notation. The use of {b}, {q}, and {s} to denote sets of spatial resource locations is confusing, as {b}, conventionally implies a singleton set containing the element $b$. Since these are meant to represent finite sets of points in $\mathbb{R}^2$, the authors should instead adopt standard set notation, such as $\mathcal{B}$, $\mathcal{Q}$, and $\mathcal{S}$, and provide explicit definitions. For example, \mathcal{B} := \{ \mathbf{b}_1, \mathbf{b}_2, \ldots, \mathbf{b}_M \} \subset \mathbb{R}^2, where each $\mathbf{b}_i \in \mathbb{R}^2$ denotes the coordinates of a blood-feeding site. In addition, matrices Phi are missing dimensions. In general I recommend the authors to be more rigorous with the mathematical notation through the paper to enhance clarity.

[-] The results section needs to be significantly improved. For instance, Figure 2 illustrates the emergence of spatial heterogeneity in mosquito population densities under both the BQ and BQS models, despite a uniformly random distribution of resources. However, the figure appears to be based on a single simulation instance. It would strengthen the findings considerably to assess the robustness of these spatial patterns by reporting variability across multiple simulations with different random resource placements. For instance, one could plot the average mosquito density at each site rank (e.g., sites sorted by mean density across simulations) with the x-axis representing site rank and the y-axis representing mean population density. Variability across simulations could then be shown using shaded regions (95% confidence intervals/standard deviation bands).

[-] If one of the main results is the emergence of spatial networks of interactions and aggregations, this should be explained and analyzed in greater detail within the results section. In particular, the manuscript should clarify whether these emergent structures are driven primarily by the behavioral (compartmental) model, rather than by the underlying Markovian transition probabilities. This raises a central question: to what extent are the findings dependent on the specific values chosen for the dispersal transition matrices or the compartamental model? If the behavioral rules dominate over the movement probabilities, is this simply an artifact of the parameter choices? How general is this result? The manuscript would benefit from a discussion of how the transition probabilities were selected or justified, whether there is empirical data that supports or can be used to calibrate these probabilities, and whether the conclusions hold under different parameterizations (i.e., a sensitivity analysis). At present, it is difficult to assess the robustness or generality of the findings, as there is no comprehensive analysis of how behavioral structure and dispersal probabilities jointly influence the observed spatial aggregation patterns.

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Reviewer #1: No

Reviewer #2: No

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Attachments

Attachment

Submitted filename: Review Mosquito Dispersal.pdf

PCSY-D-25-00047R1

Mosquito Dispersal in Context

PLOS Complex Systems

Dear Dr. Smith,

Thank you for submitting your manuscript to PLOS Complex Systems. After careful consideration, we feel that it has merit but does not fully meet PLOS Complex Systems's publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Please submit your revised manuscript by Mar 06 2026 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at complexsystems@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pcsy/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

Please include the following items when submitting your revised manuscript:

* A letter that responds to each point raised by the editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'. This file does not need to include responses to any formatting updates and technical items listed in the 'Journal Requirements' section below.

* A marked-up copy of your manuscript that highlights changes made to the original version. You should upload this as a separate file labeled 'Revised Manuscript with Track Changes'.

* An unmarked version of your revised paper without tracked changes. You should upload this as a separate file labeled 'Manuscript'.

If you would like to make changes to your financial disclosure, competing interests statement, or data availability statement, please make these updates within the submission form at the time of resubmission. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

We look forward to receiving your revised manuscript.

Kind regards,

C. Brandon Ogbunugafor

Academic Editor

PLOS Complex Systems

C. Brandon Ogbunugafor

Academic Editor

PLOS Complex Systems

Hocine Cherifi

Editor-in-Chief

PLOS Complex Systems

Journal Requirements:

If the reviewer comments include a recommendation to cite specific previously published works, please review and evaluate these publications to determine whether they are relevant and should be cited. There is no requirement to cite these works unless the editor has indicated otherwise.

Additional Editor Comments (if provided):

Thank you for the submission. It appears like one of the reviewers was dissatisfied with the effort in the revision. I recognize that not all recommended changes are always necessary in a revision, but one of the reviewers felt like their suggestions were mostly dismissed. Should you choose to submit a revision, I'd strongly suggest that you address the concerns more squarely.

[Note: HTML markup is below. Please do not edit.]

Reviewers' Comments:

Reviewer's Responses to Questions

-->Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.-->

Reviewer #1: All comments have been addressed

Reviewer #2: (No Response)

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-->2. Does this manuscript meet PLOS Complex Systems's publication criteria? Is the manuscript technically sound, and do the data support the conclusions? The manuscript must describe methodologically and ethically rigorous research with conclusions that are appropriately drawn based on the data presented.-->

Reviewer #1: Yes

Reviewer #2: No

**********

-->3. Has the statistical analysis been performed appropriately and rigorously?-->

Reviewer #1: Yes

Reviewer #2: N/A

**********

-->4. Have the authors made all data underlying the findings in their manuscript fully available (please refer to the Data Availability Statement at the start of the manuscript PDF file)?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception. The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.-->

Reviewer #1: Yes

Reviewer #2: Yes

**********

-->5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS Complex Systems does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.-->

Reviewer #1: Yes

Reviewer #2: Yes

**********

-->6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)-->

Reviewer #1: (No Response)

Reviewer #2: The authors' rebuttal did not address the substance of my previous comments. While they acknowledge that the requested analyses are important, they dismiss them as ''a different paper''. I would like to emphasize that these are not optional extensions; rather, they are fundamental requirements for establishing the validity, robustness, and interpretability of a the proposed microsimulation modeling framework. Specifically, the authors state that their main finding is ''mosquitoes will leave an area that lacks a resource and that this will give structure to mosquito populations'' As written, this is not an actual supported finding. A scientific claim of this type must be backed by compelling evidence rather than a single illustrative simulation. While I agree that algorithms are an important contribution, the conclusions remain speculative; therefore, I recommend rejecting the paper.

**********

-->7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

Do you want your identity to be public for this peer review?   If you choose “no”, your identity will remain anonymous but your review may still be made public.

For information about this choice, including consent withdrawal, please see our Privacy Policy.-->

Reviewer #1: None

Reviewer #2: No

**********

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-->While revising your submission, we strongly recommend that you use PLOS’s NAAS tool (https://ngplosjournals.pagemajik.ai/artanalysis) to test your figure files. NAAS can convert your figure files to the TIFF file type and meet basic requirements (such as print size, resolution), or provide you with a report on issues that do not meet our requirements and that NAAS cannot fix.-->-->

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Reproducibility:

To enhance the reproducibility of your results, we recommend that authors of applicable studies deposit laboratory protocols in protocols.io, where a protocol can be assigned its own identifier (DOI) such that it can be cited independently in the future. Additionally, PLOS ONE offers an option to publish peer-reviewed clinical study protocols. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols

Mosquito Dispersal in Context

PCSY-D-25-00047R2

Dear Dr. Smith,

We're pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you'll receive an e-mail detailing the required amendments. When these have been addressed, you'll receive a formal acceptance letter and your manuscript will be scheduled for publication.

An invoice for payment will follow shortly after the formal acceptance. To ensure an efficient process, please log into Editorial Manager at https://www.editorialmanager.com/pcsy/ click the 'Update My Information' link at the top of the page, and double check that your user information is up-to-date. For questions related to billing, please contact billing support at https://plos.my.site.com/s/.

If your institution or institutions have a press office, please notify them about your upcoming paper to help maximize its impact. If they'll be preparing press materials, please inform our press team as soon as possible -- no later than 48 hours after receiving the formal acceptance. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information, please contact complexsystems@plos.org.

Kind regards,

C. Brandon Ogbunugafor

Academic Editor

PLOS Complex Systems

Additional Editor Comments (optional):

Reviewers' comments:

Reviewer's Responses to Questions

-->Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.-->

Reviewer #2: All comments have been addressed

--------------------

-->2. Does this manuscript meet PLOS Complex Systems's publication criteria? Is the manuscript technically sound, and do the data support the conclusions? The manuscript must describe methodologically and ethically rigorous research with conclusions that are appropriately drawn based on the data presented.-->

Reviewer #2: Yes

--------------------

-->3. Has the statistical analysis been performed appropriately and rigorously?-->

Reviewer #2: No

--------------------

-->4. Have the authors made all data underlying the findings in their manuscript fully available (please refer to the Data Availability Statement at the start of the manuscript PDF file)?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception. The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.-->

Reviewer #2: Yes

--------------------

-->5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS Complex Systems does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.-->

Reviewer #2: Yes

--------------------

-->6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)-->

Reviewer #2: The authors addressed the comments.

--------------------

-->7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

Do you want your identity to be public for this peer review?   If you choose “no”, your identity will remain anonymous but your review may still be made public.

For information about this choice, including consent withdrawal, please see our Privacy Policy.-->

Reviewer #2: No

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