Dear Dr. Hex,

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PLOS One

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Additional Editor Comments:

Dear Authors,

I have now received feedback from two reviewers. While reviewer 2 has recommended minor revisions, they have pointed out to their comments to me that the sampling regime for this study is an issue, but the study can be considered as a positive first step. You will see that reviewer 1 has made several suggestions regarding the reporting of the sampling regime and statistics, which I think are good suggestions. I hope you can take these comments into consideration and revise your manuscript for further reviewing. I look forward to receiving your revised manuscript.

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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: Yes

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

Reviewer #1: No

Reviewer #2: Yes

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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

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Reviewer #1: I would like to thank the authors for the opportunity to review this study on the social environment and allostatic load of Phyllostomus hastatus. The integration of high-resolution 4K infrared video with cutting-edge DNA methylation age estimation represents a commendable technical effort to bridge behavioral ethology with physiological aging. The description of male "wing-locking" and the complex "allogrooming huddles" in females provide valuable new insights into the social landscape of this species.

However, the primary concern of this review is a "scaling discrepancy" between the provided data and the broad conclusions regarding chronic survival. Specifically, the study relies on an 11-day behavioral "snapshot" to support claims of lifetime allostatic load and chronic physiological wear-and-tear. Furthermore, the limited group-level replication (using only a single bachelor group) and the application of Bayesian mixed-effects models with a very low number of random effect levels (N=3) raise questions regarding the generalizability and statistical stability of the findings.

While the behavioral observations are rich and technically sound, the manuscript would benefit significantly from a more cautious framing of the causal links between short-term breeding-season aggression and long-term mortality. My detailed comments below are intended to help the authors refine these interpretations and address the statistical limitations inherent in the current study design.

Introduction

L35: The authors mention that sociality evolves partly to manage "predation dilution”. In the comparison between harem groups (in elevator shafts) and bachelors (in stairwells), is there a difference in predation risk or environmental stability? If bachelors are more exposed to predators or the elements, this could contribute to mortality independent of agonistic interactions. Please discuss if these ecological factors were considered.

L44-46: The authors state that allostatic load results from "chronic, day-to-day exposure" to social stimuli. However, the study duration is only 11 days (L116).

Please clarify how 11 days of observation can be characterized as "chronic" exposure, or revise the text to acknowledge that this study captures a acute peak in social tension (breeding season) rather than a chronic, year-round baseline.

L48-50: Since this study does not measure these physiological markers directly, please ensure the Discussion does not over-interpret "wing-locking" or "grooming" as direct proxies for these complex internal processes without citing the specific limitations of behavioral-only data.

Methodology

L119-121, L125: The study compares two harem groups against only one bachelor group. In behavioral ecology, an N=1 for a social niche makes it difficult to distinguish between a species-wide trait and the unique "culture" or personality dynamics of that specific group.

Please justify the inclusion of only one bachelor group and discuss how this limited replication affects the generalizability of the "male-as-stressful" claim.

L168-170: The authors assume all unbanded individuals in the bachelor group are male. However, the capture of a female in that exact location indicates a non-negligible presence (~7.7%) of females in supposed male-only spaces.

Please provide a sensitivity analysis or clarify the proportion of interactions involving "assumed" males. This is critical for validating the claim that the observed agonistic behaviors are strictly sex-specific.

L116- L118: The data collection period is limited to 11 days during the breeding season. The abstract, however, links these interactions to long-term "allostatic load" and "survival" (L30-31).

Please temper the discussion to acknowledge that these data represent a seasonal "snapshot." Linking 11 days of breeding-season behavior to lifetime physical wear and tear requires a more cautious interpretation.

L145-146: There is a significant difference in total footage hours: 40h for harems versus 25.5h for bachelors.

Please clarify if the observation windows (AM vs. PM) were perfectly balanced across groups. A lower sampling effort for bachelors increases the risk of "burst bias," where a single morning of high activity could disproportionately skew the results.

L166-167: Identifying the "sexually dimorphic chest gland" via infrared video in a crowded roost is technically challenging.

Please report an inter-rater reliability score (e.g., Cohen’s Kappa) to demonstrate that multiple coders reached the same conclusions regarding the sex and status of unbanded individuals.

Statistical analysis

L180, L185: The authors utilize "group ID" as a random effect (L180, L185). In Bayesian mixed-effects modeling, estimating the variance of a random effect from only three levels is mathematically unstable.

Please justify the use of a random effect with only three levels (N=3). Alternatively, consider treating "Group ID" as a fixed effect to provide more stable estimates (McElreath, 2018).

L188-189” The authors use the Region of Practical Equivalence (ROPE) to determine significance but do not define the specific boundaries used for this "null" region.

Please state the specific range used to define the ROPE so the rejection of the null hypothesis (L192) can be independently verified.

L194-195: The manuscript uses "descriptive statistics" for allogrooming because zero instances were observed in bachelors.

Please include the bachelor group in a comparative model (e.g., a Bayesian Hurdle model). This would allow for a formal statistical contrast, providing a more rigorous basis for the claim that females benefit from social buffering while males do not.

L179: The use of a zero-inflated Poisson model (L179) may not account for "overdispersion" if fights occurred in highly concentrated bursts.

Please clarify if the data was tested for overdispersion. If the variance of the aggression rate significantly exceeds the mean, a zero-inflated Negative Binomial model may be more appropriate.

Results

L250-256: Harem EV4.7 exhibited a rate of aggression more than double that of Harem CenterRight. Given this significant variation within female groups, how can the authors be certain that the higher rate in the bachelor group is a function of sex-specific niches rather than group-specific environmental factors?

L259: The text mentions Harem 'EV3.4.' Is this a typographical error intended to be Harem EV4.7 (as mentioned in L124 and L250), or was a third harem group included in the duration analysis?

L277-280: Authors describe 'allogrooming huddles' that involve grooming difficult-to-reach locations. Do you have data to suggest these huddles occur more frequently after agonistic interactions? This would provide direct evidence for the 'social buffering' hypothesis.

L265-268: Authors note that female groups have more multi-party fights (up to 5 individuals) compared to the strictly dyadic fights of males. Does this suggest that female social stress, while less physically 'intense,' is more socially 'widespread' across the group?

Discussion

L336-338: The authors provide a compelling observation regarding a spike in grooming when individuals returned covered in pollen.

Please discuss whether allogrooming in this species is primarily a functional/hygienic response to foraging residues. If the behavior is largely driven by "cleaning edible residue," it may be premature to characterize it as a "social factor contributing to differences in allostatic load" (L357) without further evidence of its psychological impact.

L354, L381: Throughout the Discussion, behavioral observations are linked to physiological wear and tear (allostatic load). While the authors acknowledge that they cannot assign causation, the concluding sentences suggest a definitive link between "day-to-day interactions" and "physiological outcomes" (L386).

Please temper the conclusion to reflect that while the correlations are consistent with the allostatic load hypothesis, this study provides no direct physiological data to confirm the mechanism in this specific population.

L356-358: The manuscript suggests that allogrooming is a key factor in buffering allostatic load and promoting survival. However, the authors also note that harem males do not participate in allogrooming (L362) yet maintain superior body condition and survival compared to bachelors (L319-320).

Please address this discrepancy. If harem males achieve high survival without allogrooming, does this imply that the absence of aggression is a more significant determinant of allostatic load than the presence of affiliative buffering?

L379: The authors cite personal observations to suggest that bachelor wounds occur year-round, justifying the 11-day "snapshot."

To strengthen this claim, please provide more detail on these observations (e.g., frequency or seasonality) or cite any available data that suggests male aggression levels do not significantly drop outside the breeding season.

Reviewer #2: Dear authors, I very much appreciate studies on social behaviour conducted in the wild. Phyllostomus hastatus is a fascinating species with respect to sociality. Although the observation period is relatively short and the sample size limited, I think your study provides valuable observations that can serve as a starting point for further research on the relationship between social environment and allostatic load (including physiological consequences and ultimately fitness). Below I provide several minor comments.

Abstract

“Valence” is only mentioned in the abstract. I recommend either removing the term or introducing it earlier in the manuscript (e.g., in the Introduction).

Methods

Line 116: “In” should be lower case.

Line 141:

Please provide the sampling rate and bit resolution of the recordings. Were recordings automatically triggered? If so, based on which threshold?

Figure S2:

Please include the spectrogram settings used (e.g., window type, FFT size, overlap). Also specify which software was used to generate the spectrograms.

Results

I have a few more general questions regarding the results:

Females appear to solicit copulations. Is this consistently the case in this species? Are there other bat species in which females actively solicit copulations? I am more familiar with species in which males either attempt forced copulations or perform elaborate courtship displays, with females making the mating decision rather than actively soliciting copulation.

Were you able to determine whether females had preferred grooming partners?

I found the fighting behaviour between males particularly interesting. The wing-lock position seems difficult for the attacked individual to counter. Are similar fighting behaviours known from other bat species?

Discussion

Line 319:

Could you briefly clarify how body condition was defined?

Line 377:

I would suggest rephrasing this statement — the observation period most likely does not fully capture seasonal variation in social behaviour.

**********

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Reviewer #1: No

Reviewer #2: No

**********

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Dear Dr. Hex,

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.

Please submit your revised manuscript by Aug 28 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'.
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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.

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As the corresponding author, your ORCID iD is verified in the submission system and will appear in the published article. PLOS supports the use of ORCID, and we encourage all coauthors to register for an ORCID iD and use it as well. Please encourage your coauthors to verify their ORCID iD within the submission system before final acceptance, as unverified ORCID iDs will not appear in the published article. Only  the individual author can complete the verification step; PLOS staff cannot  verify ORCID iDs on behalf of authors.

We look forward to receiving your revised manuscript.

Kind regards,

Anindita Bhadra, PhD

Academic Editor

PLOS One

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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.

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Additional Editor Comments (if provided):

I am happy to see the revised version of the manuscript. Please respond to Reviewer 1, and submit your revised manuscript.

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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: (No Response)

**********

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

Reviewer #1: Yes

Reviewer #2: (No Response)

**********

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

The PLOS Data policy

Reviewer #1: Yes

Reviewer #2: (No Response)

**********

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

Reviewer #1: Yes

Reviewer #2: (No Response)

**********

Reviewer #1: I thank the authors for their careful and constructive revision. The revised manuscript is substantially improved. In particular, the addition of a second observation period and additional harem and bachelor-group observations directly addresses my earlier concern about limited group-level replication. The revised dataset now provides a more convincing basis for comparing affiliative and agonistic interactions across sex-biased social environments. I also appreciate the authors’ revision of the aggression-rate model to account for overdispersion, their addition of a model for allogrooming occurrence, and their more cautious framing of the behavioural nature of the study.

Overall, I find the manuscript much stronger and closer to publication. The behavioural observations remain valuable, especially the detailed descriptions of male “wing-locking” and female allogrooming. However, I recommend minor revision before acceptance. My remaining concerns are not about the central behavioural pattern, but about ensuring that the interpretation remains appropriately cautious and that several inconsistencies introduced or retained during revision are corrected.

The authors have clarified that the study does not directly test allostatic load, social buffering, or survival mechanisms. This is a substantial improvement. However, the Abstract and final Discussion still give considerable prominence to the possibility that the observed behavioural differences contribute to allostatic load and survival differences. In the Abstract, lines 34–39, the sentence linking the behavioural differences to allostatic load and survival should be tempered slightly further. I recommend phrasing this as a hypothesis generated by the present behavioural data, or as a pattern consistent with possible future tests, rather than implying that the present study demonstrates such links. Similarly, in the Discussion, lines 448–455, the authors appropriately acknowledge that causal links cannot be drawn, but the final sentence still implies that the present study characterizes how day-to-day interactions culminate into physiological outcomes. Please revise this to state that the behavioural findings generate testable hypotheses about possible physiological consequences.

In the Methods, lines 139–141, the manuscript states that harem groups roost in the elevator shaft and bachelor groups roost in the adjacent stairwell. In the response letter, the authors further explain that they have no evidence that one location is more exposed to predators or environmental conditions than the other. This clarification should be added briefly to the manuscript, either in the Methods or Discussion, so that readers can evaluate whether roost-location differences may confound the harem–bachelor comparison.

The expanded dataset is a major improvement. However, in the Methods, lines 136–150, one bachelor roost location, B-ST3.4, was sampled in both 2025 and 2026 and treated as independent because bachelor-group membership is presumed to be labile. This may be reasonable, but it should remain explicit wherever the manuscript refers to “three bachelor groups.” The conclusions should be framed as evidence from sex-biased social environments at this study site rather than as broad species-wide generalization.

The shift from “sex-specific” to “sex-biased” social environments is appropriate. The capture data also support the claim that bachelor groups are strongly male-biased. However, in the Methods, lines 185–195, some sex assignments still rely on assumptions from roost membership and visual criteria. Please state clearly that coding was performed primarily by a single observer and that formal inter-rater reliability for sex/status assignment could not be calculated. If possible, the representative image showing the chest-gland/genitalia criteria should be included in the Supplementary Information.

In the Methods, lines 159–168, the increased total footage reduces the earlier concern about burst bias. However, because AM/PM sampling was not perfectly balanced, I recommend including the AM/PM distribution table from the response letter in the Methods or Supplement. This would make the sampling structure transparent to readers and would help show that the higher aggression rate in bachelor groups is unlikely to be an artefact of a single active time period.

In the Methods, lines 217–223, and Results, lines 338–344, the added Bernoulli model is an acceptable improvement for a descriptive study. However, the manuscript should make clear that this model estimates the probability of observing allogrooming in a 10-minute interval. It does not directly test social buffering. Any wording linking allogrooming to buffering or allostatic load should remain explicitly hypothetical.

In the Discussion, lines 416–436, the response to the harem-male discrepancy is only partly satisfactory. If harem males have better condition/survival than bachelor males but were not observed to participate in allogrooming, this suggests that reduced exposure to intense bachelor aggression may be an important alternative or complementary mechanism to allogrooming-based buffering. I recommend that the Discussion state this more clearly. Allogrooming should be framed as a potential female-specific social benefit, not as a general explanation for all harem-associated survival advantages.

In the Discussion, lines 437–448, the added capture-record information on wounds across months strengthens the argument that bachelor males may experience aggression beyond the immediate observation window. However, wound records are indirect evidence and do not quantify year-round aggression rates. The current cautious framing is mostly appropriate, but please avoid implying that equivalent aggression rates persist outside the mating season.

Several statistical and terminology issues should be corrected.

In the Methods, lines 197–200, the text still refers to “three focal groups,” although the revised design includes six group-level observations. Please correct this.

In the Methods, lines 217–219, please correct “brsm” to “brms.”

In the Supporting Information, lines 663–665 and 677–679, the statement that Bulk and Tail ESS are “similar to effect sizes” is incorrect. ESS refers to effective sample size, not effect size. Please correct this in the supplementary table captions.

In the Methods, lines 210–215, and Supporting Information, lines 645–649 and 656–660, the ROPE boundaries are now reported, which resolves the earlier concern. However, the main text should clarify whether the ROPE is defined on a standardized scale, consistent with the Supplementary figure captions.

Several small textual, figure, and numerical inconsistencies should also be corrected before publication.

In the Abstract, lines 25–27, the species name appears misspelled as “Phylostomus hastattus.” Please correct to Phyllostomus hastatus.

In the Introduction, lines 44–46, please correct “individualsfrom.”

In the Methods, lines 153–154, please correct the sentence break in “Mammalogists. and were approved…”

In the Methods, lines 163–166, please correct the typographical artefact “ly?the harem male.” In the Results, lines 302–307, the text refers to “H-EV4.4,” but the Methods and Table 2 list “H-EV4.2.” Please correct for consistency.

In the Figure 2 caption, lines 348–352, the caption refers to duration of allogrooming bouts in “the two harem groups,” but the revised study includes three harem groups.

In the Supporting Information, lines 631–635, the caption still refers to “three focal social groups,” although the revised manuscript discusses six group-level observations.

Please also check numerical consistency between the Results and Table 1. In the Results, lines 225–230, the authors report 394 bachelor-group agonistic interactions, but Table 1 appears to list 386 bachelor male–bachelor male interactions. Similarly, the Results report 197 allogrooming instances, whereas Table 1 appears to list 196 female–female allogrooming interactions. If these differences are due to unknown individuals, excluded classifications, or rounding/counting decisions, this should be explained.

For the supplementary allogrooming probability figure, the y-axis should be labelled more clearly as “Predicted probability of observing allogrooming in a 10-min interval.” The axis should ideally be bounded from 0 to 1, and group labels should be standardized to match the manuscript.

Reviewer #2: (No Response)

**********

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Reviewer #1: No

Reviewer #2: No

**********

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Attachments

Attachment

Submitted filename: Review report.docx

Affiliative and agonistic interactions distinguish sex-biased social groups of greater spear-nosed bats, Phyllostomus hastatus

PONE-D-25-60173R2

Dear Dr. Hex,

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.

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Kind regards,

Anindita Bhadra, PhD

Academic Editor

PLOS One

Additional Editor Comments (optional):

I am happy to see that you have revised the manuscript with inputs from the reviewers. I am happy to accept the manuscript in it's current form.

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

Reviewer #1: All comments have been addressed

**********

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

Reviewer #1: Yes

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

Reviewer #1: Yes

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

The PLOS Data policy

Reviewer #1: Yes

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

Reviewer #1: Yes

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Reviewer #1: The authors have satisfactorily addressed all the comments and concerns raised in the previous review. The revised manuscript is clear, and I have no further substantive comments. I therefore recommend acceptance of the manuscript in its present form.

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Reviewer #1: No

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