-->PONE-D-25-67537-->-->Identification of tooth traces from a Cretaceous (Maastrichtian) Edmontosaurus annectens bonebed in the Lance Formation, Wyoming, U.S.A.-->-->PLOS One

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Reviewer #1: Yes

Reviewer #2: Partly

Reviewer #3: Yes

Reviewer #4: Partly

Reviewer #5: Yes

Reviewer #6: Yes

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Reviewer #1: N/A

Reviewer #2: N/A

Reviewer #3: Yes

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Reviewer #1: Yes

Reviewer #2: Yes

Reviewer #3: Yes

Reviewer #4: Yes

Reviewer #5: Yes

Reviewer #6: Yes

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Reviewer #2: Yes

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Reviewer #4: Yes

Reviewer #5: Yes

Reviewer #6: Yes

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-->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: The manuscript is very well written, with clear explanations and well-illustrated. The limitations of interpretation are well considered and explained. Figure 10 and S1 Fig seem to be the same. You should move S2 Fig and S3 Fig into the main text rather than the Supplementary section.

Reviewer #2: The study presents results that appear to be original and not previously published, and the manuscript does not show evidence of direct duplication of figures or text from earlier peer-reviewed publications. The research question is incremental rather than groundbreaking, but this is acceptable under PLOS ONE’s scope.

However, the manuscript would benefit from a clearer positioning relative to closely related prior studies, particularly to better articulate what is novel about the present contribution beyond the presentation of tooth traces. The Abstract Introduction should more explicitly state how this work advances existing knowledge rather than reiterating established patterns. For example, only at the end of the mauscript it is cited the guide for the identification of tooth traces

488 (Fig 10) to incorporate many different types of bone perforations. In my opinion this is the most relevant part of the MS, since is a true contribution that enhances previous guideliness (showed in Fig S1).

So, only at the end of the discussion I realized that the MS is adding value other than just a description of tooth bites on some dinosaurs.

I think that of the MS is intended to be published in PLOS ONE, it needs to add value somehow, not only presenting a bit marks in a descriptive way.

I suggest re-shaping the MS focusing on the guide of tooth marks. As I see, the guide is the product of this MS.

Coupled with this, the figures need to be of better quality (although maybe they lost quality in the PDF, but please, check the original files since they were not provided) and clearly show the mark patterns matched with the guide.

See the commented PDF for few other comments.

All the best.

Reviewer #3: The manuscript constitutes an interesting contribution to understanding certain biological features commonly found in fossil vertebrates. I believe this manuscript does not require major changes.

My only comment regards a sentence in page 14 related to the possible recognition of Nanotyrannus as a juvenile Tyrannosaurus rex. This has been recently discussed by other authors that sufficiently support they are different taxa (see Zanno, L. E., & Napoli, J. G. (2025). Nanotyrannus and Tyrannosaurus coexisted at the close of the Cretaceous. Nature, 1-3). Perhaps this can be relevant to the identification of the tooth marks inflictor.

Reviewer #4: The submitted MS reports the results of investigation of tooth traces of Edmontosaurus bones from Lance Formation. This topic is more than welcome, because numerous articles have been published in recent years on the examination of bite marks, which in many cases can be used to draw palaeoecological conclusions. However, unfortunately, this manuscript does not contain any new relevant scientific information related to the examination of bite marks, but only uses previously published observations in the examination of their own material. I am not saying that there is no need to mention the examples cited in the manuscript in a publication, but I just wanted to emphasize that the current manuscript does not contain any significant new findings. The conclusion chapter also contains entirely general statements. If the authors disagree with this, it would be justified to formulate the new results listed in the conclusions more precisely, and it would be worthwhile to emphasize in the main text of the article what different or new results have been achieved in the field of bite mark examination. In summary, the article is descriptive in nature. It contains observations, but does not place them in a broader context. However, the results presented in the study may be used as data in other publications, so the editor will decide whether to publish the study.

• There are numerous great scientific publications dealing with bite marks of Mesozoic bones, but only a few studies are mentioned in the publication, which is often based on a rather arbitrary selection. I believe that the authors do not refer to fundamental works published on the subject (e.g. Fiorill, 1991; Carpenter, 1998; Jacobsen, 1998; Farlow and Holtz, 2002; Avilla et al., 2004; Buffetaut et al., 2004; Hone and Rauhut, 2009; Fiorelli, 2010; Longrich et al., 2010; Milàn et. al., 2010; Schwimmer, 2010; Bell, et al., 2012; Boyd et al., 2013, Botfalvai et al 2014 and references therein). I recommend taking a critical look at the publication mentioned and the references listed.

• Surangular fossil HRS05499: The authors analyse the perforations/lesions visible on this fossil in a separate section. In the end, they conclude that these are not bite marks, but are normal foramina found in a triceratopsini surangular. The figure used as evidence (Figure 3) is of such poor resolution that I could not make out anything on it. It is possible that this is a mistake of the journal and only the generated PDF file contains such a poor quality figure. In any case, based on the available material, I cannot judge whether the authors are right on this issue. Comparing the marks on the angular bone of HRS05499 with those on other triceratopsine angular bones would help clarify this issue. Furthermore, it is necessary to list other anatomical features that link this bone element to triceratopsine dinosaurs. Are there other bones from the site that originate from ceratopsian dinosaurs?

• The authors discuss at length that bite marks can often be confused with anatomically present foramina. I don't really understand why this needs to be emphasized. A detailed examination of the bite marks begins with determining the taxonomic classification of the bone being examined. Bones that cannot be classified taxonomically cannot be examined for bite marks, as in such cases we do not know what foramina the original bone contained. In my opinion, this is a conclusion that everyone knows even without the article.

Figures:

• I realize that this is not the authors' fault, but the PDF figures sent by the journal are of very poor resolution. Therefore, in many cases, I was unable to verify the evidence that the authors wrote about in the text. So, at the moment, I can only hope that the authors have created images with much better resolution, where everything is much clearer. The following figures were the most problematic in this regard: Figure 2B-C; Figure 3; Figure 4A;

• Figure 1: For me, this is not a very informative figure. I know that it is not a very important figure in this study, but I still felt I should point out that it does not provide any information about the site. A sedimentary section, a photograph of the bone-bearing beds, an excavation picture, etc. would improve this figure and enhance the appearance of the article.

Reviewer #5: Overall I like this manuscript and it's attempt to more clearly define and confine what is a tooth trace on a fossil, as these have important ecological and behavioural implications. The main issue I have, and it's not a major one, is the writing style and organization of this manuscript could be improved to increase readability (especially for a more general audience). I suggest another pass through to tighten and moving around results sections would make it flow better.

But overall a god manuscript and I look forward to seeing it published without much alteration of the core of the text.

Reviewer #6: This manuscript by Siviero and colleagues provides a rigorous and appropriately cautious approach to identifying the organisms behind tooth traces in a dinosaur bonebed from the Late Cretaceous. The research is well documented, with sufficient information to follow and reproduce the analysis in other settings. Similarly, I feel that the assignment of tooth marks to Tyrannosaurus is well supported with the evidence presented. A number of minor points are outlined below, but I wish to call attention to three in particular:

1) Determining the etiology of marks on bone has a significant body of research in paleoanthropology and archaeology, particularly in distinguishing human-created tool marks from other biotic and abiotic causes. I would suggest incorporating some of this research here, because it is directly relevant. This includes wear marks from trampling or soil compaction (e.g., Behrensmeyer et al., 1986; Dominguez-Rodrigo et a., 2009; Fiorillo, 1989; Olsen and Shipman, 1988; Rozada et al., 2018).

2) Similarly, a few additional paleontological papers on tooth marks related to crocodilian and archosaur feeding, etc., could be cited. This includes Drumheller et a., 2023; Njau and Gilber, 2016; Erickson and Olson, 1996; and citations therein.

3) The ultimate identification of HRS05499 is ambiguous. Can you definitively say it is ceratopsid based on other aspects of morphology? Or is it a weird Edmontosaurus that has a ceratopsid-like foramen pattern? Ceratopsid and hadrosaurid surangulars should be fairly distinct in overall shape (not just the pattern of foramina), so I think this should be addressed in more detail. It is not clear from the text as currently written.

Minor Comments:

- The recent Siviero et al. 2025 reference may now be added, because it includes some of the same sample.

- lines 91-92: Are the inferences around Knethichnus and Linichnus related to the same citation as earlier in the paragram? I would add the citation here again, if approriate, because it is fairly well separated in the text.

- line 180: Can the teeth be referred to Borealosuchus definitively? Tooth morphology is pretty flexible in crocs, so I personally prefer to be a bit more cautious - e.g., "Eusuchia indet., likely from Borealosuchus sternbergii". They're likely not Brachychampsa, of course!

- line 323: You might add the newest Nanotyrannus papers here that more definitively settle the issue

- Erickson and Olson, 1996, discusses other bite marks referable to Tyrannosaurus rex; this should be cited.

Behrensmeyer, A.K., Gordon, K.D., and Yanagi, G.T. 1986. Trampling as a cause of bone surface damage and pseudo-cutmarks. Nature, 319:768-771.

https://doi.org/10.1038/319768a0

Dominguez-Rodrigo, M., S. de Juana, A.B. Galan, and M. Rodriguez, 2009. A new protocol to differentiate trampling marks from butchery cut marks. Journal of Archaeological Science, 1-12.

Drumheller, S. K., D’Amore, D. C., Njau, J. K. 2023. Taphonomic approaches to bite-mark analyses in the fossil record and applications to archosaurian paleobiology. In IUP Crocodylian Paleobiology.

Erickson, G. M., & Olson, K. H. (1996). Bite marks attributable to Tyrannosaurus rex: preliminary description and implications. Journal of Vertebrate Paleontology, 16(1), 175-178.

Fiorillo, A.R., 1989. An experimental study of trampling: implications for the fossil record. Bone Modification, 2, 61-71.

Njau, J. and Gilbert, H. 2016. Standardizing terms for crocodile-induced bite marks on bone surfaces in light of the frequent bone modification equifinality found to result from crocodile feeding behavior, stone tool modification, and trampling. FOROST (Forensic Osteology) Occasional Publications, 3:1-13.

Olsen, S.L. and P. Shipman, 1988. Surface Modification on Bone: Trampling versus Butchery. Journal of Archaeological Science, 15, 535-553.

Rozada, L., Allain, R., & Tournepiche, J. F. (2018). Trampling experiments on bones in fine and soft sediments. Quaternaire. Revue de l'Association française pour l'étude du Quaternaire, 29(1), 39-44.

Siviero, B. C., Rega, E., Nick, K. E., & Chadwick, A. V. (2025). Analysis of pseudopathologies in Edmontosaurus annectens bones: taphonomic implications from biogenetic and diagenetic bone alterations from a Cretaceous bonebed in the Lance Formation, Wyoming. Journal of Vertebrate Paleontology, e2600392.

--Andrew A. Farke

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Reviewer #1: No

Reviewer #2: No

Reviewer #3: Yes:  Rodolfo A. Coria

Reviewer #4: No

Reviewer #5: Yes:  T. Alexander Dececchi

Reviewer #6: Yes:  Andrew A. Farke

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Attachments

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Submitted filename: PONE-D-25-67537-B. Siviero.pdf

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Submitted filename: PONE-D-25-67537_reviewer.pdf

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Submitted filename: PONE-D-25-67537TAD.pdf

Identification of tooth traces from a Cretaceous (Maastrichtian) Edmontosaurus annectens bonebed in the Lance Formation, Wyoming, U.S.A.

PONE-D-25-67537R1

Dear Dr. C. T. Siviero,

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,

Furqan A. Shah

Academic Editor

PLOS One

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Reviewer #1: All comments have been addressed

Reviewer #3: All comments have been addressed

Reviewer #6: (No Response)

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

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented. -->

Reviewer #1: Yes

Reviewer #3: Yes

Reviewer #6: Yes

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

Reviewer #1: N/A

Reviewer #3: Yes

Reviewer #6: N/A

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

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). 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 #3: Yes

Reviewer #6: Yes

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

PLOS ONE 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 #3: Yes

Reviewer #6: (No Response)

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-->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 #3: I believe the authors have fullfilled all comments and reviews. I have no further comments hoping to see this ms out soon.

Reviewer #6: The authors have adequately addressed my comments from the previous version of this manuscript. I have only a handful of minor edits to suggest - at this point, I do not need to see the manuscript again.

Comments:

- Add the Griffin et al. paper also, relative to Nanotyrannus as a valid taxon (10.1126/science.adx8706)

- line 118, line 158 - replace "monospecific" with "monodominant" as to the bonebed type, to ensure consistency throughout

- check italics for all genus/species names (esp. in references - e.g., reference 38 and others)

--Andrew A. Farke

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Reviewer #1: Yes:  Raúl Esperante

Reviewer #3: Yes:  Rodolfo A. Coria

Reviewer #6: Yes:  Andrew A. Farke

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