Dear Dr. Koju,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’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.<article class="text-token-text-primary w-full focus:outline-none [--shadow-height:45px] has-data-writing-block:pointer-events-none has-data-writing-block:-mt-(--shadow-height) has-data-writing-block:pt-(--shadow-height) [&:has([data-writing-block])>*]:pointer-events-auto scroll-mt-[calc(var(--header-height)+min(200px,max(70px,20svh)))]" data-scroll-anchor="true" data-testid="conversation-turn-4" data-turn="assistant" data-turn-id="request-WEB:72d4a3d1-613a-42fc-87c4-bb51707507b5-1" dir="auto" tabindex="-1">

The reviewers identified several substantial issues that currently limit the strength of the inferences. In particular, they noted the absence of hypotheses and predictions, insufficient clarity and consistency in the description of the study design and sampling effort, and concerns regarding the robustness of the spatial and temporal analyses. In addition, aspects of the dietary analysis require greater transparency, particularly with respect to sample sizes and uncertainty. The Introduction and Discussion were also considered underdeveloped relative to the scope and importance of the topic, and would benefit from a stronger theoretical framing and clearer articulation of the study’s contribution within the broader literature.

Addressing these points will require significant revisions to multiple sections of the manuscript, including the Introduction, Methods, Results, and Discussion. The reviewers’ detailed comments are provided below and should be carefully considered in preparing a revised version.

plosone@plos.org . When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

• A letter that responds to each point raised by the academic editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'.
• 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, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

If applicable, we recommend that you deposit your laboratory protocols in protocols.io to enhance the reproducibility of your results. Protocols.io assigns your protocol its own identifier (DOI) so that it can be cited independently in the future. For instructions see: https://journals.plos.org/plosone/s/submission-guidelines#loc-laboratory-protocols . Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols .

We look forward to receiving your revised manuscript.

Kind regards,

Paulo Corti, Ph.D.

Academic Editor

PLOS One

Journal Requirements:

When submitting your revision, we need you to address these additional requirements.

1. Please ensure that your manuscript meets PLOS ONE's style requirements, including those for file naming. The PLOS ONE style templates can be found at

https://journals.plos.org/plosone/s/file?id=wjVg/PLOSOne_formatting_sample_main_body.pdf and

https://journals.plos.org/plosone/s/file?id=ba62/PLOSOne_formatting_sample_title_authors_affiliations.pdf .

2. Thank you for stating in your Funding Statement:

“This study was supported by a faculty grant from the University Grants Commission (UGC), Nepal (FRG 77/78 S&T-01 2020/21 and FRG 080/81 S&T-07), Andrew Sabin Family Foundation, Zoo New England, and the China Exploration and Research Society, Hong Kong. RCK’s effort was supported in part by the Office of Research Infrastructure Programs (ORIP) of the National Institutes of Health through grant number P51OD010425 to the Washington National Primate Research Center, USA”

Please provide an amended statement that declares *all* the funding or sources of support (whether external or internal to your organization) received during this study, as detailed online in our guide for authors at http://journals.plos.org/plosone/s/submit-now. Please also include the statement “There was no additional external funding received for this study.” in your updated Funding Statement.

Please include your amended Funding Statement within your cover letter. We will change the online submission form on your behalf.

3. Thank you for stating the following financial disclosure:

“This study was supported by a faculty grant from the University Grants Commission (UGC), Nepal (FRG 77/78 S&T-01 2020/21 and FRG 080/81 S&T-07), Andrew Sabin Family Foundation, Zoo New England, and the China Exploration and Research Society, Hong Kong. RCK’s effort was supported in part by the Office of Research Infrastructure Programs (ORIP) of the National Institutes of Health through grant number P51OD010425 to the Washington National Primate Research Center, USA”

Please state what role the funders took in the study.  If the funders had no role, please state: "The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript."

If this statement is not correct you must amend it as needed.

Please include this amended Role of Funder statement in your cover letter; we will change the online submission form on your behalf.

4. We note that your Data Availability Statement is currently as follows: [All relevant data are within the manuscript and its Supporting Information files]

Please confirm at this time whether or not your submission contains all raw data required to replicate the results of your study. Authors must share the “minimal data set” for their submission. PLOS defines the minimal data set to consist of the data required to replicate all study findings reported in the article, as well as related metadata and methods (https://journals.plos.org/plosone/s/data-availability#loc-minimal-data-set-definition).

For example, authors should submit the following data:

- The values behind the means, standard deviations and other measures reported;

- The values used to build graphs;

- The points extracted from images for analysis.

Authors do not need to submit their entire data set if only a portion of the data was used in the reported study.

If your submission does not contain these data, please either upload them as Supporting Information files or deposit them to a stable, public repository and provide us with the relevant URLs, DOIs, or accession numbers. For a list of recommended repositories, please see https://journals.plos.org/plosone/s/recommended-repositories.

If there are ethical or legal restrictions on sharing a de-identified data set, please explain them in detail (e.g., data contain potentially sensitive information, data are owned by a third-party organization, etc.) and who has imposed them (e.g., an ethics committee). Please also provide contact information for a data access committee, ethics committee, or other institutional body to which data requests may be sent. If data are owned by a third party, please indicate how others may request data access.

5. We note that Figures 1 & 2 in your submission contain [map/satellite] images which may be copyrighted. All PLOS content is published under the Creative Commons Attribution License (CC BY 4.0), which means that the manuscript, images, and Supporting Information files will be freely available online, and any third party is permitted to access, download, copy, distribute, and use these materials in any way, even commercially, with proper attribution. For these reasons, we cannot publish previously copyrighted maps or satellite images created using proprietary data, such as Google software (Google Maps, Street View, and Earth). For more information, see our copyright guidelines: http://journals.plos.org/plosone/s/licenses-and-copyright.

We require you to either (1) present written permission from the copyright holder to publish these figures specifically under the CC BY 4.0 license, or (2) remove the figures from your submission:

a. You may seek permission from the original copyright holder of Figures 1 &2to publish the content specifically under the CC BY 4.0 license.

We recommend that you contact the original copyright holder with the Content Permission Form (http://journals.plos.org/plosone/s/file?id=7c09/content-permission-form.pdf) and the following text:

“I request permission for the open-access journal PLOS ONE to publish XXX under the Creative Commons Attribution License (CCAL) CC BY 4.0 (http://creativecommons.org/licenses/by/4.0/). Please be aware that this license allows unrestricted use and distribution, even commercially, by third parties. Please reply and provide explicit written permission to publish XXX under a CC BY license and complete the attached form.”

Please upload the completed Content Permission Form or other proof of granted permissions as an "Other" file with your submission.

In the figure caption of the copyrighted figure, please include the following text: “Reprinted from [ref] under a CC BY license, with permission from [name of publisher], original copyright [original copyright year].”

b. If you are unable to obtain permission from the original copyright holder to publish these figures under the CC BY 4.0 license or if the copyright holder’s requirements are incompatible with the CC BY 4.0 license, please either i) remove the figure or ii) supply a replacement figure that complies with the CC BY 4.0 license. Please check copyright information on all replacement figures and update the figure caption with source information. If applicable, please specify in the figure caption text when a figure is similar but not identical to the original image and is therefore for illustrative purposes only.

The following resources for replacing copyrighted map figures may be helpful:

USGS National Map Viewer (public domain): http://viewer.nationalmap.gov/viewer/

The Gateway to Astronaut Photography of Earth (public domain): http://eol.jsc.nasa.gov/sseop/clickmap/

Maps at the CIA (public domain): https://www.cia.gov/library/publications/the-world-factbook/index.html and https://www.cia.gov/library/publications/cia-maps-publications/index.html

NASA Earth Observatory (public domain): http://earthobservatory.nasa.gov/

Landsat: http://landsat.visibleearth.nasa.gov/

USGS EROS (Earth Resources Observatory and Science (EROS) Center) (public domain): http:

Earth (public domain): http://www.naturalearthdata.com/

6. Please remove your figures from within your manuscript file, leaving only the individual TIFF/EPS image files, uploaded separately. These will be automatically included in the reviewers’ PDF.

7. 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.

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

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

Reviewer #1: Partly

Reviewer #2: Partly

**********

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

Reviewer #1: No

Reviewer #2: N/A

**********

3. Have the authors made all data underlying the findings in their manuscript fully available??>

The PLOS Data policy

Reviewer #1: Yes

Reviewer #2: Yes

**********

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

Reviewer #1: Yes

Reviewer #2: Yes

**********

Reviewer #1: The manuscript addresses an important question about the coexistence of large carnivores. However, the strength of the inferences is limited by (i) the lack of explicit hypotheses/predictions, (ii) inconsistencies and omissions in the design/sampling, and (iii) spatial/temporal conclusions based on very simple methodological decisions without sensitivity analysis. I recommend major revision.

1) Introduction

Hypotheses and predictions are missing. It remains to be clarified what patterns are expected (spatial, temporal, and trophic) and what mechanisms sustain them (competition, exploitation vs. interference, anthropogenic subsidies, prey availability).

I recommend adding hypotheses and predictions

2) Methodology – Design and effort

-Inconsistencies in number of sites/cameras/duration. Twenty-six cameras in 20 locations are mentioned, but Figure 1 shows 19 locations. In addition, the operating time is reported first, followed by the number of cameras (reverse order).

I recommend: (a) standardizing figures in text/figures/tables; (b) describing whether there was >1 camera per location and under what criteria; (c) presenting an effort table with camera days per location and per year, failures, and relocations. The experimental design should be better explained.

-Long study period (2018–2025) without stratification. They report an average of ~761 camera days, which implies that cameras were not active throughout the entire period and possible temporal bias in location.

I recommend: (a) detailing the actual windows of activity for each camera; (b) indicating whether the spatial distribution changed between years; (c) stratifying analyses (or including covariates) by year/season to avoid mixing ecologically dissimilar periods that inflate overlaps.

-Camera model and recording mode. The exact model is requested (line 98). Further on (line 166), it is indicated that there were images and videos, but earlier only images are mentioned.

I recommend specifying the brand/model, sensitivity/intervals, and for each period whether photos, videos, or both were captured, and how they were integrated into the analysis.

3) Methodology – Space and time

-Spatial overlap is calculated using 100 m buffers around detections and altitude limits defined with the same data. This approach is overly simplistic and potentially circular.

I recommend: (a) replacing or supplementing with use estimators (e.g., use density/kernel)); (b) reporting sensitivity analyses to buffer width and altitude limits

-Temporal activity collapsed into 7 years. It is not appropriate to speak of “synchrony” of the three predators with multi-year averaged data.

Request: (a) report Δ with CI and n per pair; (b) stratify by year/season or add a supplementary analysis showing the temporal stability of the patterns.

4) Methodology – Diet

-Sample sizes by species not reported. To interpret trophic overlap, the number of feces analyzed per species (DNA/micro-histology) and the criteria for selecting the histological “subset” are needed.

5) Results

-Temporal synchrony (l. 260–261): Avoid statements of synchrony for the three predators without temporal stratification.

-Spatial overlap: Strong conclusions are based on buffer + altitudinal clipping.

-Diet: Include n per species analyzed and Pianka's CI (bootstrap per scat).

To support the Discussion and conclusions, the manuscript should: (i) state hypotheses/predictions and mechanisms; (ii) clarify and tabulate design/effort (with temporal stratification); (iii) replace or, at least, subject to sensitivity analysis the spatial analysis based on buffers; and (iv) strengthen the diet section with n per species. These improvements are necessary to ensure reproducibility and inferential validity.

Reviewer #2: Comments to authors

I read with great interest your article titled “Niche Partitioning Facilitates Coexistence of Three Apex Predators in the Lapchi Valley, Central Himalaya, Nepal.” This study examines niche partitioning and coexistence among sympatric snow leopards, leopards, and Himalayan wolves in the Lapchi Valley of Central Nepal. Using camera trapping and dietary analyses, the authors assess spatial, temporal, and trophic overlap to explore mechanisms facilitating predator coexistence in a high-altitude Himalayan system. My primary concerns are related to some of the methods and the depth of the introduction and discussion given the nature of the topic and approaches.

Please see my specific line comments below.

Abstract

In the abstract, it would be helpful to start with one or two sentences that provide some context and clearly point out the main gap before moving into the study objectives.

Introduction

The introduction currently reads thin and would benefit from further development. Lines 47–54 discuss the threats these apex predators face, but the content is split across two paragraphs and would benefit from better organization.

While some of these points were briefly mentioned, the authors should further expand on contexts such as the role of apex predators in the ecosystem, existing mechanisms of coexistence among apex predators (e.g., different types of niche partitioning), the remaining knowledge gaps, and what makes the study system unique, etc. Expanding on these areas would provide a stronger theoretical background for the study.

Methods

Please ensure consistent citation formatting.

The spatial overlap analysis adopts a presence-based, rule-constrained approach, conceptually similar to an environmental envelope method, rather than more complex approaches such as MCP, SDM, or JSDM. The authors should more explicitly justify their methodological choice and discuss its underlying assumptions, rather than adopting alternative approaches.

Please ensure that index abbreviations are used consistently across the manuscript (e.g., “B” is used twice for different indices).

Results

The spatial overlap results are derived from a relatively small number of camera trap sites (7–11 detections per species). Given this limited spatial sampling, stronger justification is needed when interpreting these estimates, particularly statements suggesting complete spatial overlap or subsumed distributions. The authors should clarify that these results reflect detection-based, potential overlap, and discuss how sampling design and buffer choices may influence the observed patterns, or alternatively consider using additional data or complementary methods to support their conclusions.

Please ensure consistent table formatting. Please also correct the grammar in lines 299–301; there is an extra closing parenthesis.

Discussion

Please ensure consistent citation formatting throughout the manuscript.

The Discussion should be expanded and developed in greater depth, particularly with respect to the conservation implications of the findings. Given that both the topic and the analytical approaches are not particularly novel, and that similar analyses of apex and meso-predator interactions have been conducted across other regions of the Himalayas, it is especially important for the authors to clearly articulate how this study system differs from previous work and what new insights it provides. In particular, the authors should better explain why understanding niche partitioning among snow leopards, leopards, and Himalayan wolves in this specific area matters. For example, do the observed dietary patterns reflect differential coexistence strategies, human-predator relationships, or local perceptions of predators by herders? What about the spatial overlap results, and how do those ecological findings link to the cultural context? What are the potential long-term implications for prey resources, habitat use, or conflict risk given the high spatial and temporal overlap observed among these species?

**********

what does this mean? ). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy

Reviewer #1: No

Reviewer #2: No

**********

[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

To ensure your figures meet our technical requirements, please review our figure guidelines: https://journals.plos.org/plosone/s/figures

You may also use PLOS’s free figure tool, NAAS, to help you prepare publication quality figures: https://journals.plos.org/plosone/s/figures#loc-tools-for-figure-preparation.

NAAS will assess whether your figures meet our technical requirements by comparing each figure against our figure specifications.

Dear Dr. Koju,

Please submit your revised manuscript by Apr 01 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 plosone@plos.org . When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

• A letter that responds to each point raised by the academic editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'.
• 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, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

https://journals.plos.org/plosone/s/submission-guidelines#loc-laboratory-protocols . Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols .

We look forward to receiving your revised manuscript.

Kind regards,

Paulo Corti, Ph.D.

Academic Editor

PLOS One

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.

Please review your reference list to ensure that it is complete and correct. If you have cited papers that have been retracted, please include the rationale for doing so in the manuscript text, or remove these references and replace them with relevant current references. Any changes to the reference list should be mentioned in the rebuttal letter that accompanies your revised manuscript. If you need to cite a retracted article, indicate the article’s retracted status in the References list and also include a citation and full reference for the retraction notice.

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

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

Reviewer #1: All comments have been addressed

Reviewer #2: All comments have been addressed

**********

2. Is the manuscript technically sound, and do the data support the conclusions??>

Reviewer #1: Yes

Reviewer #2: Yes

**********

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

Reviewer #1: Yes

Reviewer #2: Yes

**********

4. Have the authors made all data underlying the findings in their manuscript fully available??>

The PLOS Data policy

Reviewer #1: (No Response)

Reviewer #2: Yes

**********

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

Reviewer #1: Yes

Reviewer #2: Yes

**********

Reviewer #1: (No Response)

Reviewer #2: I read with great interest your article titled “Niche Partitioning Facilitates Coexistence of Three Apex Predators in the Lapchi Valley, Central Himalaya, Nepal.” The authors have effectively addressed the reviewers’ comments from the previous review round. I only have a few minor comments remaining.

Lines 49-50: Classification should be specified as by the IUCN Red List.

Lines 54-61: These sentences provide important background information about the study species, but they feel out of place after the knowledge gap has already been introduced. I suggest moving them earlier in the paragraph right after introducing snow leopards and Himalayan wolves as the dominant alpine carnivores. The paragraph could then follow a clearer logical flow: (1) Snow leopards and Himalayan wolves are the dominant carnivores in this alpine system, followed by one or two sentences describing each species; (2) in recent years, leopards have increasingly expanded into snow leopard habitat, followed by one or two sentences introducing leopards; (3) this recent range expansion (currently line 47) then naturally leads into the knowledge gap.

Line 125 and multiple instances in Discussion: Please ensure consistency in citation formatting, particularly in the use of parentheses versus brackets.

Line 277: The abbreviation of MCP has already been noted earlier in line 204。

Discussion: The conservation implications section could be strengthened by being more specific, as the current recommendations feel somewhat generic. It would also benefit from more explicitly engaging with the broader consequences of ongoing leopard range expansion and climate-driven habitat shifts. For example, do the authors expect to see similar or different range or diet shifts in snow leopards and wolves as well? What might this imply for longer-term restructuring of the predator guild under continued environmental change? How might future land-use planning or protected-area management adapt to these elevational range shifts and changing predator interactions? Expanding this section to address these forward-looking and differential management considerations would enhance the applied relevance of the study, especially in the context of rapid climate change in high-elevation ecosystems.

**********

what does this mean? ). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy

Reviewer #1: No

Reviewer #2: No

**********

[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

To ensure your figures meet our technical requirements, please review our figure guidelines: https://journals.plos.org/plosone/s/figures

You may also use PLOS’s free figure tool, NAAS, to help you prepare publication quality figures: https://journals.plos.org/plosone/s/figures#loc-tools-for-figure-preparation.

NAAS will assess whether your figures meet our technical requirements by comparing each figure against our figure specifications.

Niche Partitioning Facilitates Coexistence of Three Apex Predators in the Lapchi Valley, Central Himalaya, Nepal

PONE-D-25-54236R2

Dear Dr. Koju,

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 will be generated when your article is formally accepted. Please note, if your institution has a publishing partnership with PLOS and your article meets the relevant criteria, all or part of your publication costs will be covered. Please make sure your user information is up-to-date by logging into Editorial Manager at Editorial Manager®  and clicking the ‘Update My Information' link at the top of the page. For questions related to billing, please contact billing support .

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 onepress@plos.org.

Kind regards,

Paulo Corti, Ph.D.

Academic Editor

PLOS One