Same place, different time: Leopards (Panthera pardus kotiya) employ temporal partitioning as a co-existence mechanism in a human-dominated, unprotected agricultural landscape in Sri Lanka

Andrew M. Kittle; Anjali C. Watson

doi:10.1371/journal.pone.0335296

Peer Review History

Original SubmissionOctober 7, 2025
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There is no requirement to cite these works unless the editor has indicated otherwise. 6. 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 If you have cited papers that have been retracted, please include the rationale for doing so in the manuscript text, or remove these references and replace them with relevant current references.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 If you need to cite a retracted article, indicate the article’s retracted status in the References list and also include a citation and full reference for the retraction notice.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 1. Is the manuscript technically sound, and do the data support the conclusions? Reviewer #1: Yes Reviewer #2: Partly ******** 2. Has the statistical analysis been performed appropriately and rigorously? -->?> Reviewer #1: Yes Reviewer #2: Yes ****** 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: This study provides insights into leopard density, site use, and activity patterns within the human-dominated Central Highlands of Sri Lanka, where tea estates dominate the region. The study fills a critical knowledge gap in the ecology of carnivores and their coexistence outside of protected areas and is highly relevant to the field of carnivore research. Overall, the study is technically sound, and the data adequately support the study’s interpretations and conclusions. However, as the protected-area datasets were collected during different time periods than the tea estate dataset, the author should also highlight the potential impact of seasonal or temporal variations on the activity patterns, as it might partially affect the differences observed. Reviewer #2: Title and Abstract The title would benefit from capitalising major words for consistency with scientific style and including the scientific name at first mention. The abstract contains several long sentences that reduce readability; tightening these and presenting the key findings more concisely would improve accessibility. The scientific name should also appear upon first mention in the abstract. Introduction The discussion of global protected areas (lines 37–48) feels broader than necessary for a study centred on Sri Lanka. This section could be more concise and more directly linked to the local context. The section on human–wildlife conflict (lines 49–66) is relevant but currently repetitive; streamlining it would improve flow. The term “top carnivore” reads awkwardly and could be replaced with “apex predator” for clarity and consistency with later usage. Lines 67–78 could also be condensed. The material in lines 79–93 provides the clearest initial framing of the study and may be more effective if moved earlier. “UNESCO” should be written in full on first use (line 84). Finally, the use of first person (“we”) in lines 94–103 should be avoided to maintain an objective tone. Materials and Methods The Methods are generally strong, detailed, and scientifically rigorous, though several areas would benefit from clarification. The study area description (lines 106–131) contains extensive climatic detail, but the relevance of specific rainfall patterns should be explained more clearly in relation to leopard ecology or study design. The explanation of estate structure and settlement patterns (lines 128–149) is useful but would be strengthened by explicitly linking these features to expected leopard behaviour. The camera trap design (lines 150–173) is well justified, though the rotation of cameras across three rounds (lines 138–144) requires clearer explanation to ensure the population closure assumption is met. The decision to use only left flank images (lines 159–166) is reasonable, but the potential impact of excluding individuals should be acknowledged. The SECR modelling approach (lines 167–196) is robust, though the effort weighting method (line 175) is unconventional and would benefit from justification or citation. The iterative buffer selection process (lines 174–196) is thorough but could be summarised more concisely, with full details placed in supplementary material. The site utilisation analysis (lines 197–243) is clearly structured, though the use of Google Earth for settlement distance (lines 200–205) should be justified, and the treatment of plantation forests as equivalent to natural forest (lines 214–220) may require ecological explanation. The handling of multicollinearity (lines 221–243) is appropriate, though the rationale for selecting quadratic terms (lines 221–223) could be strengthened. The activity time analysis (lines 244–287) is well explained, but the Z test comparisons (lines 267–287) may require mention of multiple comparison considerations. Overall, the Methods are sound and replicable, and clarifying a few analytical decisions would further strengthen the study. Results The Results are clearly organised and generally well presented. The density results (lines 288–304) are straightforward, with the top model clearly identified and the buffer justified, though the large difference between the observed and model estimated sex ratios (lines 293–299) should be addressed in the Discussion. The comparison with other Sri Lankan studies (lines 301–316) is useful, but the text could more directly state that overlapping confidence intervals indicate no significant difference in density. The site use findings (lines 317–324) are well linked to the hypotheses, showing that overall leopard use was highest at intermediate human and dog abundance, males preferred intermediate distances from settlements, and females selected areas with higher prey availability. These patterns are well explained, though the relatively low R² values should be acknowledged. Discussion The Discussion is thoughtful and well connected to the results, though several sections could be tightened for clarity and impact. The opening discussion of density (lines 352–367) effectively compares findings with protected areas, but would benefit from explicitly noting that overlapping confidence intervals do not necessarily imply ecological equivalence. The comparison with recent high density estimates (lines 368–392) is important, though the critique of methodology could be more concise, and the suggestion of overcrowding in protected areas may require stronger evidence. The explanation of the female biased sex ratio (lines 392–416) is well supported but could be summarised more succinctly. The site utilisation discussion (lines 417–439) interprets the quadratic relationships with human and dog presence clearly, though the section could more directly connect these findings to coexistence mechanisms rather than focusing heavily on community perceptions of dog predation. The distinction between male avoidance of settlements and female prey driven space use (lines 417–436) is well articulated but could be made more concise. The activity time interpretation (lines 440–462) convincingly demonstrates temporal partitioning, though the explanation of reduced diurnal activity could be streamlined, and the comparison with Nepal may be more effective if shortened. The final section on snares (lines 462–end) is strong and policy relevant, and would benefit from a clearer link back to the central argument that behavioural adaptations alone cannot offset human caused mortality. Overall, the Discussion is robust and insightful, and with some tightening it will read as a confident, cohesive, and persuasive interpretation of your findings. ****** 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. 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Revision 1
16 Jul 2026 Author Response We have responded to all of the Reviewers comments below. Our responses are all preceded with the word "RESPONSE". Reviewers' comments: Comments to the Author Reviewer #1: This study provides insights into leopard density, site use, and activity patterns within the human-dominated Central Highlands of Sri Lanka, where tea estates dominate the region. The study fills a critical knowledge gap in the ecology of carnivores and their coexistence outside of protected areas and is highly relevant to the field of carnivore research. Overall, the study is technically sound, and the data adequately support the study’s interpretations and conclusions. RESPONSE: Thank you for your positive comments and interest in the manuscript. However, as the protected-area datasets were collected during different time periods than the tea estate dataset, the author should also highlight the potential impact of seasonal or temporal variations on the activity patterns, as it might partially affect the differences observed. RESPONSE: Thank you for this comment as seasonal patterns may be important for leopard activity times. However, all three of the datasets do in fact overlap in terms of time of year with the tea estate study from August through November 2016, the Wilpattu NP study from July through October 2015 and the Gal Oya NP study including data from > 2 years (November 2017 – June 2020). Nevertheless, we have explicitly addressed the imperfect overlap within the text in the discussion section under Leopard activity times: “It is important to note that there was not perfect overlap regarding the time of year monitored for all 3 sites; that GONP was monitored for considerably longer than both WNP and the tea estate landscape (20 months vs 4 months); and that the sites are all located within different climatic zones. As such, there may be additional factors which partially underlie these observations. However, the strong similarities in activity patterns detected between WNP and GONP, despite having varied monitoring periods and being in different climatic zones, supports the conclusion that leopards in the tea estate landscapes are indeed acting differently and employing temporal partitioning to avoid people.” Reviewer #2: Title and Abstract: The title would benefit from capitalising major words for consistency with scientific style and including the scientific name at first mention. RESPONSE: We actually did this, and then looked at the style guide for PLoS ONE and realized it calls for “sentence style” with only the first word capitalized. So we have left it as it was, except for adding the scientific name. The abstract contains several long sentences that reduce readability; tightening these and presenting the key findings more concisely would improve accessibility. RESPONSE: Thank-you for this suggestion. We have attempted to improve readability in several ways. We have broken up the first sentence into 2 parts (Ln 14 -16). In Ln 16 we add a comma to more effectively break up the sentence. In Ln 26 we change “human-dominated” to “shared” to improve flow. In Ln 29 we break the sentence into two parts to improve flow. In Ln 31 we remove a redundant comment “and their importance” to make the sentence more concise. The scientific name should also appear upon first mention in the abstract. RESPONSE: Ok, this has been done. Introduction: The discussion of global protected areas (lines 37–48) feels broader than necessary for a study centred on Sri Lanka. This section could be more concise and more directly linked to the local context. RESPONSE: Thank you for this suggestion. We have substantially reduced this paragraph (174 to 151 words) by removing specific references to individual taxa and instead speaking more broadly about the importance of unprotected areas for wildlife. We have also removed some unnecessary wording (“It is widely accepted that…”). With regards to linking explicitly to the local context, we don’t feel that this is the right place to do it as what we intended here was to make the argument that broadly, how wildlife utilize unprotected regions need to be better understood in general, given increasing human impacts and climate change-induced movements. The link to the local context comes later. We hope this rationale is satisfactory. The section on human–wildlife conflict (lines 49–66) is relevant but currently repetitive; streamlining it would improve flow. RESPONSE: Thank you for this suggestion. We have reduced this section as well (from 262 words to 225) by drastically compressing the opening sentence and removing redundant and unnecessary wording (e.g. replace “these large predators” with “they”; remove “it is increasingly accepted that…”; remove “in terms of”, “furthermore”; remove duplicate statements about explicitly sharing space” etc). The term “top carnivore” reads awkwardly and could be replaced with “apex predator” for clarity and consistency with later usage. RESPONSE: Thank you for this suggestion. It makes much more sense to use “apex predator” and we have done this. Lines 67–78 could also be condensed. RESPONSE: Thank you for this suggestion. We have likewise reduced this section from 171 words to 153, by removing redundancies and being more concise. The material in lines 79–93 provides the clearest initial framing of the study and may be more effective if moved earlier. RESPONSE: While we see the reviewer’s point in terms of its clarity for the purposes of this study, we have attempted to frame the research by narrowing from the broad scale to the specific, and feel that this section, which deals with the specific study area under investigation should come at the end of the Introduction. We hope that this is acceptable. “UNESCO” should be written in full on first use (line 84). RESPONSE: Ok, this has been done. Finally, the use of first person (“we”) in lines 94–103 should be avoided to maintain an objective tone. RESPONSE: Ok, this has been done. Materials and Methods: The Methods are generally strong, detailed, and scientifically rigorous, though several areas would benefit from clarification. RESPONSE: Thank you for this positive assessment of the Methods section. We have attempted to improve the clarity of the section as suggested. The study area description (lines 106–131) contains extensive climatic detail, but the relevance of specific rainfall patterns should be explained more clearly in relation to leopard ecology or study design. RESPONSE: We thank the reviewer for this pertinent point. On reflection, we realize that the detailed climatic information is not highly relevant to this study – or to leopard ecology in general given their ability to thrive across a wide range of climates. Our intention had been to describe the study site to situate the reader, but realize that we went overboard on detail. As such, we have condensed this section (from 189 words to 161) to provide an overview, hopefully without burdening the reader with too much unnecessary detail. The explanation of estate structure and settlement patterns (lines 128–149) is useful but would be strengthened by explicitly linking these features to expected leopard behaviour. RESPONSE: We thank the reviewer for this insightful comment also, which again forced us to reflect on our description. The key element we are trying to get across is that this landscape has an unusually broad human footprint with people spread widely across the landscape. The impact of this is that wildlife cannot effectively separate spatially from humans at a broad scale as it can in areas where human populations are high but clustered. We have therefore added to the sentence claiming that “the human footprint is large and extensive” to clarify this: “As such, the human footprint in this landscape is large and extensive, making broad-scale spatial separation between humans and wildlife impossible.” The camera trap design (lines 150–173) is well justified, though the rotation of cameras across three rounds (lines 138–144) requires clearer explanation to ensure the population closure assumption is met. RESPONSE: Thank-you for the positive feedback. Regarding the rotation of cameras, we have added a sentence explicitly detailing the duration of the entire study (all 3 rounds combined) and justifying (with references which link it to a study which provides a biological underpinning) this duration as fitting within a closed population. In addition, we have included specific mention of running the ‘closure’ test in ‘secr’. The decision to use only left flank images (lines 159–166) is reasonable, but the potential impact of excluding individuals should be acknowledged. RESPONSE: Ok, thank you. We have added a couple of explanatory sentences as follows: “Employing only a single flank has the potential to result in individuals being excluded which can impact density estimates. However, the fact that the three images excluded were lacking in clarify sufficient to unambiguously identify or sex the individuals, alleviates this concern.” The SECR modelling approach (lines 167–196) is robust, though the effort weighting method (line 175) is unconventional and would benefit from justification or citation. RESPONSE: Ok, we have added a citation for an earlier publication which utilizes this same weighting method (Kittle AM, Watson AC, Samaranayake PKL. Edge effects and distribution of prey forage resources influence how an apex predator utilizes Sri Lanka’s largest protected area. J. Zool. 2021; 314(1): 31 – 42. doi: 10.1111/jzo.12870) and added it to the existing citation (Efford MG, Borchers DL, Mowat G. Varying effort in capture-recapture studies. Methods Ecol. Evol. 2013; 4(7): 629-636. doi: 10.1111/2041-210X.12049) for the theoretical underpinning. The iterative buffer selection process (lines 174–196) is thorough but could be summarised more concisely, with full details placed in supplementary material. RESPONSE: Ok, thank you for this suggestion. We have summarized this section in the main text by alluding only to the general process employed and directing readers to Table S2 for details. We have then moved the more detailed description to the caption for Table S2 (which shows the iterative buffer size estimation process). This has streamlined this section considerably. The site utilisation analysis (lines 197–243) is clearly structured, though the use of Google Earth for settlement distance (lines 200–205) should be justified RESPONSE: Thank you for the comment. We attempted to justify this method in the original ms by stating that “available GIS layers did not have sufficiently fine grain to identify all communities on the landscape”. However, we have more clearly justified this in the revision by stating: “Distance to human settlement was determined using the “distance measure” tool on Google Earth because many estate communities are small clusters of dwellings which are not represented on available GIS layers, but are visible on Google Earth maps.” , and the treatment of plantation forests as equivalent to natural forest (lines 214–220) may require ecological explanation. RESPONSE: Thank you for this comment. As above, we attempted to explain this in the original manuscript as follows: “forests were considered all treed landscapes inclusive of plantation forests as leopards are generalists [58] and these habitats are used by leopards, especially in landscape mosaics [59]” with the citations supporting the justification. However, we realize that this was insufficient and poorly worded. We have now tried to further clarify this approach by stating: “forests were considered all treed landscapes inclusive of plantation forests, which here are typically Eucalyptus-dominated stands originally planted to provide fuelwood for the tea factories. Natural and plantation stands were treated as the same since leopards are adept at utilizing a wide range of habitats [58] including plantation patches, which are especially important in landscape mosaics [59] despite their inferior quality from a biodiversity perspective” The handling of multicollinearity (lines 221–243) is appropriate, though the rationale for selecting quadratic terms (lines 221–223) could be strengthened. RESPONSE: Thank you for this comment. We agree that this wording was a little weak. We have added two references here to strengthen the support for this approach (Balme GA, Hunter LTB, Slotow R. Feeding habitat selection by hunting leopards Panthera pardus in a woodland savanna: prey catchability versus abundance. Anim. Behav. 2007; 74(3): 589-598. doi: 10.1016/j.anbehav.2006.12.014 and Hopcraft JGC, Anderson TM, Pérez-Vila S, Mayemba E, Olff, H. Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers. J. Anim. Ecol. 2012; 81: 201-213. doi: 10.1111/j.1365-2656.2011.01885.x). We also added an explanatory sentence: “If continuous variables are acting quadratically, model coefficients for the quadratic term should be negative, which indicates that although a leopard might select for a particular feature (e.g. elevation) or resource (e.g. dog abundance), it avoids areas where the feature is extreme or resource overabundant, resulting in a non-linear relationship [63].” The activity time analysis (lines 244–287) is well explained, but the Z test comparisons (lines 267–287) may require mention of multiple comparison considerations. RESPONSE: Thank you for this comment. We have added some explanatory detail here to make clear that there was more than one comparison undertaken, so that the description of the method now reads: “To compare leopard activity patterns between the protected (WNP and GONP) and unprotected (tea estate landscape) areas, we compared the independent proportions of leopard observations for each of the time periods between the unprotected site and each protected site in turn.” Overall, the Methods are sound and replicable, and clarifying a few analytical decisions would further strengthen the study. RESPONSE: Thank you for this positive and supportive comment. We have tried to address the reviewers concerns and improved clarification of the analytical decisions that were pointed out. We agree that this improves the interpretability of the manuscript. Results: The Results are clearly organised and generally well presented. RESPONSE: We appreciate that the reviewer felt this was the case. The density results (lines 288–304) are straightforward, with the top model clearly identified and the buffer justified, though the large difference between the observed and model estimated sex ratios (lines 293–299) should be addressed in the Discussion. RESPONSE: Thank you for bringing up this important point. We have addressed the large observed difference between the observed and model estimates sex ratios more directly than we had previously in the Discussion section about adult sex ratios as follows: “This model allowed for variation between sexes to influence the spatial parameter related to home range size (σ), but not the probability of capture at home range centre (g0). The much larger σ seen for males compared to females (x̅ = 2409 vs 737) indicates substantial home range size variation resulting in an even more female-biased ASR than was apparent from raw observations, as the latter would have likely included observations of males far from home range centres.” The comparison with other Sri Lankan studies (lines 301–316) is useful, but the text could more directly state that overlapping confidence intervals indicate no significant difference in density. RESPONSE: Although we felt that we had been quite clear about this in the original ms, we have altered the language slightly to make the implication even more clear. It now reads: “The observed density was lower than what has been found in Sri Lanka’s best protected National Parks, but in almost every case 95% CIs overlap, indicating that this difference is not significant (Table 3)”. We hope that this small change is sufficient to clarify the issue. The site use findings (lines 317–324) are well linked to the hypotheses, showing that overall leopard use was highest at Attachments Attachment Submitted filename: PONE-D-25-54311.R1 - Response to Reviewers for Resubmission.docx https://doi.org/10.1371/journal.pone.0335296.r002
21 Jul 2026 Decision Letter - James Edward Brereton, Editor Same place, different time: Leopards (Panthera pardus kotiya) employ temporal partitioning as a co-existence mechanism in a human-dominated, unprotected agricultural landscape in Sri Lanka. PONE-D-25-54311R1 Dear Dr. Kittle, 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, James Edward Brereton, MSc Academic Editor PLOS One Additional Editor Comments (optional): Reviewers' comments: https://doi.org/10.1371/journal.pone.0335296.r003
Formally Accepted
Acceptance Letter - James Edward Brereton, Editor PONE-D-25-54311R1 PLOS One Dear Dr. Kittle, I'm pleased to inform you that your manuscript has been deemed suitable for publication in PLOS One. Congratulations! Your manuscript is now being handed over to our production team. At this stage, our production department will prepare your paper for publication. This includes ensuring the following: * All references, tables, and figures are properly cited * All relevant supporting information is included in the manuscript submission, * There are no issues that prevent the paper from being properly typeset You will receive further instructions from the production team, including instructions on how to review your proof when it is ready. Please keep in mind that we are working through a large volume of accepted articles, so please give us a few days to review your paper and let you know the next and final steps. Lastly, if your institution or institutions have a press office, please let them know about your upcoming paper now to help maximize its impact. If they'll be preparing press materials, please inform our press team within the next 48 hours. 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. You will receive an invoice from PLOS for your publication fee after your manuscript has reached the completed accept phase. If you receive an email requesting payment before acceptance or for any other service, this may be a phishing scheme. Learn how to identify phishing emails and protect your accounts at https://explore.plos.org/phishing. If we can help with anything else, please email us at customercare@plos.org. Thank you for submitting your work to PLOS One and supporting open access. Kind regards, PLOS ONE Editorial Office Staff on behalf of Mr. James Edward Brereton Academic Editor PLOS One https://doi.org/10.1371/journal.pone.0335296.r004

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