PONE-D-24-16876Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogenyPLOS ONE

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Reviewer #1: In this manuscript, the authors re-evaluate the status of a large-bodied predatory dinosaur skeleton described in 1931 and destroyed in 1944 during WWII. The authors also introduce a novel phylogenetic analysis of the predatory dinosaurs which adds - among others - new elements on the relationships among the large-bodied group Allosauroidea (i.e., the lineage including the specimen destroyed in 1944).

The phylogenetic analysis and the implications of that analysis are well-argumented and would surely improve our knowledge on the evolution of these dinosaurs, and I do not request any significant edit of that section of the manuscript (see just a few notes and comments in the pdf attached to this revision). Yet, I have several concerns about the taxonomic interpretation of the particular specimen lost in 1944, which is claimed by the authors to represent a new genus and species ("Alllissaurus markgrafi") and has been removed from the well-known taxon Carcharodontosaurus saharicus.

Since my concerns are both epistemological and methodological and stem from the logical and chronological sequence of the events involved, I first provide, below, a brief chronology of them (see the manuscript for the details of each step).

1) Depéret and Savornin (1925-27) erected Megalosaurus saharicus, based on two non-associated teeth from the "mid-Cretaceous" of Algeria.

2) Stromer (1931) erected Carcharodontosaurus, based on a partial skeleton from the "mid-Cretaceous" of Egypt. Stromer considered Carcharodontosaurus and "Megalosaurus saharicus" the same species and diagnosed Carcharodontosaurus saharicus from the Egyptian specimen.

3) The Egyptian specimen was destroyed during WWII.

4) Sereno et al. (1996) described a partial skull from the "mid-Cretaceous" of Morocco and referred it to Carcharodontosaurus saharicus, based on Stromer (1931).

5) Brusatte and Sereno (2007) explicitly designated the Moroccan specimen as neotype of C. saharicus. This decision was based on the following arguments: 5a) the two teeth described by Depéret and Savornin (1925-27) likely pertain to different individuals; 5b) the diagnostic tooth characters originally cited are no longer diagnostic at the specific or generic level; and 5c) both Egyptian and Algerian specimens cannot be used as type material; 5d) the Moroccan specimen "is identical to both the original holotypic teeth and Stromer’s Egyptian material".

In this new manuscript, the authors introduce two recently rediscovered original photos of the Egyptian specimen prior to its destruction, and list a series of features which could differentiate that material from the Moroccan specimen. Despite the specimen being destroyed, the authors consider the published material sufficient for erecting a new genus and species, distinct from Carcharodontosaurus saharicus. Note that these statements challenge the statements 5c) and 5d) by Brusatte and Sereno (2007).

The thesis discussed in the manuscript is thus the following:

6) The Moroccan specimen does not belong to the same genus and/or species of the Egyptian specimen.

7) Even if the Egyptian specimen is lost, its description and the new photographs justify it as holotype for the formal erection of a genus and species .

Both 6) and 7) are problematic, and I encourage the authors to reconsider them carefully:

Statement 7) is explicitly challenged by the International Code of Zoological Nomenclature (ICZN, 1999), Recommendation 73B: "An author should designate as holotype a specimen actually studied by him or her, not a specimen known to the author only from descriptions or illustrations in the literature". The Egyptian material, actually lost in 1944, should not be used to erect any new taxon.

The ICZN recommendation must be followed for a series of epistemological reasons. The first is the non-reproducibility of the observations reported in support of the new species: the irreversible destruction of the specimen prevents any objective evaluation of the diagnostic features introduced. Accordingly, any emendation of the diagnosis cannot be falsified. The second is the non-testable nature of a series of ambiguous interpretations of the published photos used by the authors to suggest some features not explicitly reported by Stromer (1931).

For example, the authors wrote:

"An expansion of the narial fossa to the nasal in the form of a supranarial fossa, as it is present

in Acrocanthosaurus, Concavenator, Carcharodontosaurus and Meraxes [11,18,43,44] is not

clearly visible in the photo of SNSB-BSPG 1922 X 46, but Stromer [1] described the area of

the anterior end of the nasal lateral to the median premaxillary process as slightly concave.

However, a clearly defined rim of such a fossa, as it is present in Carcharodontosaurus ([18]:

Fig 134A) is not at all indicated in Stromers figures, nor visible on the photograph."

The authors implicitly remark that the development of key features is ambiguous once the literature is compared with the photos/illustration. What should be considered the valid source? In this case, the narial fossa is described by Stromer (1931) as a slight concavity, yet such feature is not illustrated in the figures/photos.

The authors also wrote:

"The antorbital fossa extends onto the lateral surface of the nasals, which represents an

allosauroid synapomorphy [7,47], and is clearly visible in lateral view on the nasals of SNSBBSPG

1922 X 46. This is visible in the photograph of the specimen and also indicated in

Stromer’s reconstruction of the skull (Figs 2 and 4B, D)"

Contrary to the authors, I hardly differentiate an antorbital fossa of the nasal in the photo of the Egyptian specimen from the homologous part in the Moroccan specimen. The description by Stromer (1931) is ambiguous on such feature, so in this case we have to trust the interpretation of the text and figures by the authors. Note that the lack of such fossa is in agreement with the other late-diverging carcharodontosaurids. Maybe, I am over-conservative and prefer not speculate on elements of an old photo which was not explicitly created for illustrating anatomical details. The authors might be right in their interpretation of the images, but without direct observation of the specimen, there is no way for eventually falsifying their statement. If they are wrong, such error is fixed in their diagnosis of the taxon and could not be corrected by any independent observation. The drawings by Stromer do not match all details claimed from the photos, so what of the two images should we trust? Some features claimed by the authors could be non-biological in origin, due to taphonomy or damage, or even just photographic artifacts. If the authors have "over-interpreted" some features in the only available images of the now destroyed Egyptian specimen, nobody could falsify them. Thus, the diagnosis of "Alllissaurus markgrafi" is epistemologically problematic and cannot be considered a proper scientific statement.

Stated in a simpler way: if the authors are wrong in some interpretations, how could them be falsified without an available specimen? A couple of old black-and-white photos alone (one of which clearly was not meant to illustrate anatomical details but just the whole mounted material) is not sufficient evidence that all features listed by the authors as diagnostic are actually natural apomorphies.

Other features of the Egyptian specimen, even if genuine, could not be significant from a taxonomic perspective. For example, the "nasal horn" and the more extensive maxillary subcutaneous ornamentation (claimed to be absent or less developed in the Moroccan specimen) could be ontogeny-related and/or sexually-dimorphic features widespread among the carcharodontosaurids once a richer sample is available.

Statement 6) is an even more complex issue. The authors apparently seem unaware of the full implications of their main hypothesis: if we accept that the lost Egyptian specimen could be used as valid type of a taxon, and also endorse the taxonomic distinction of the Egyptian and Moroccan specimens, this has relevant nomenclatural implications:

First, if the Moroccan specimen does not belong to the same taxon as the Egyptian specimen, then the rationale for erecting a neotype of C. saharicus by Brusatte and Sereno (2007) from the former specimen is no more valid. In fact, the neotype was introduced under the assumption that all specimens mentioned belong to a single taxon. If the two specimens belong to different taxa, one cannot be used as the neotype which replaces the destruction of the other: the neotype sensu Brusatte and Sereno (2007) is thus invalidated! Furthermore, if the lost Egyptian specimen is now considered valid as holotype of "Alllissaurus", then it means that it has always been the valid type for a carcharodontosaurid taxon lacking a neotype: in that case, it has always been the most appropriate neotype of Carcharodontosaurus in spite of the Moroccan specimen, because Stromer (1931) used it to erect and diagnose that genus. Accordingly, the genus name "Alllissaurus" is automatically a junior synonym of the genus name Carcharodontosaurus.

The authors might reply that Carcharodontosaurus is now anchored to the Moroccan specimen, but such nomenclatural act was historically based and conditioned to a taxon sample the authors here have just demonstrated being wrong. They wrote: "In the interest of preserving the widely used genus Carcharodontosaurus and family Carcharodontosauridae, we concur with the designation of SGM-Din 1 as the neotype of Carcharodontosaurus saharicus sensu Brusatte and Sereno". Yet, such statement is contradicted by the main argument of their manuscript, because the authors have just (implicitly) demonstrated that the rationale for designating SGM-Din 1 [the Moroccan specimen] as valid neotype for Carcharodontosaurus is no more valid, because the Moroccan and Egyptian specimens do not belong to the same genus.

The best way to preserve both Carcharodontosaurus and Carcharodontosauridae should be to designate the Egyptian specimen as neotype of C. saharicus because:

1) The Egyptian specimen - assumed by the authors as a potentially valid type specimen for a taxon - has been the explicit source for the diagnosis of Carcharodontosaurus since the erection of the genus in 1931, whereas the Moroccan specimen was introduced only in 1996 and has been (arbitrarily) considered the neotype of the same taxon of the Egyptian specimen only in 2007 to replace the destroyed material.

2) All literature prior to 2007 assumed the Egyptian specimen as the main source of Carcharodontosaurus diagnosis (and, so far, it keeps being the only available source for all postcranial features of this taxon): even the referral of the Moroccan specimen to that taxon (regardless of it being right or wrong) was merely a consequence of comparing it with the Egyptian material. Removing the latter from Carcharodontosaurus is everything but an act "preserving" Carcharodontosaurus stability and consistency.

There is another reason which justifies the removal of the Moroccan specimen from C. saharicus instead of the Egyptian one: as stated above, according to the ICZN Recommendation 73B, "an author should designate as holotype a specimen actually studied by him or her, not a specimen known to the author only from descriptions or illustrations in the literature". Given that, of the two specimens, only the Moroccan one is currently housed in a public institution, only the latter could be eventually studied by a researcher, and so it is the only one of the two which is available if a second taxon is eventually split from the Carcharodontosaurus hypodigm.

A fourth reason for challenging the exclusion of the Egyptian specimen from Carcharodontosaurus is that the support of the phylogenetic results is weak. As shown in detail by the authors, when compared to the shortest scenario found by the analyses, only two additional steps are necessary for producing a Carcharodontosaurus saharicus node including both Egyptian and Moroccan specimens, and only three additional steps for reconstructing a node with all Carcharodontosaurus specimens in the monophyletic genus (both C. saharicus and C. iguidensis material). I replicated the analyses and confirm the results. Such small step differences are not statistically significant once evaluated with the Templeton test: we cannot exclude that they are merely due to lack of overlapping data and/or even a few coding artifacts (eventually biased by some miscoding due to misinterpretation of the published source). This means that the monophyly of Carcharodontosaurus keeps being a potentially valid topology. I would not consider the phylogenetic results sufficiently robust for supporting the splitting of the Egyptian specimen from Carcharodontosaurus. The radical taxonomic revision of a key historical specimen based on such weak results is thus poorly supported and should be avoided.

In conclusion:

Nomenclatural and epistemological reasons strongly discourage the erection of a new taxon from a specimen now lost and never studied directly by anyone after 1944.

The ICZN explicitly recommends to not erect a taxon like this.

If the Egyptian and Moroccan specimens belong to distinct genera, then the rationale followed in 2007 for designating the latter as "neotype" of Carcharodontosaurus is no more valid. Note that if the authors are correct in differentiating the Egyptian and Moroccan morphs, then "Carcharodontosaurus sensu Brusatte and Sereno 2007" is a chimera based on two (or more) taxa, and the only valid (non chimerical) "version" of Carcharodontosaurus is only the one prior 1996, based uniquely on the Egyptian specimen: any argument used in 2007 to define the Moroccan specimen as the neotype of the same taxon of the Egyptian specimen is thus no more valid, and cannot be used to justify the removal of the Egyptian specimen from the genus which was explicitly erected and diagnosed from its morphology.

If the authors are right in the taxonomic distinction of the two specimens, then the Moroccan specimen was not Carcharodontosaurus since the beginning: it could be provisionally referred to an indeterminate carcharodontosaurid species (and mentioned by its collection number, like other OTUs used by the authors in their phylogenetic analysis), pending an explicit revision of its status and that of "C. iguidensis", based on direct first-hand analysis. Removing the Egyptian specimen from Carcharodontosaurus would arbitrarily invalidate over 90 years of literature based on the taxon erected by Stromer in 1931, a result that cannot be endorsed just in order to conserve an arbitrary neotype definition which the new manuscript itself has demonstrated being based on wrong assumptions.

The phylogenetic test showed that only a few steps differentiate the splitting of all specimens from the traditional lumping of them into Carcharodontosaurus. Thus, pending more robust and less problematic evidence, any taxonomic revision should be avoided. Terms like "cf. Carcharodontosaurus" or "?Carcharodontosaurus" are valid terms for the material from Morocco and Niger to remark the potential non-monophyletic nature of that name until a consensus is reached.

The authors should thus remark the contradiction in Brusatte and Sereno (2007) relative to their new study, should state the problematic use of the Moroccan specimen as "neotype" of what actually was a chimaera, have to moderate the interpretation of the features based only on the photos of the lost fossil, and must avoid the creation of an intrinsically problematic taxon which is explicitly challenged by the ICZN.

Note that the taxonomic section of the manuscript lacks any record of ZooBank registrations, making any nomenclatural act - regardless of their substantial validity - as formally invalid (e.g., if the manuscript is accepted in the current version, "Alllissaurus markgrafi" is a nomen nudum).

Reviewer #2: This paper seeks to correct a taxonomic conundrum hiding in plain sight but which the community was essentially unaware: the fact that Stromer’s Egyptian material of Carcharodontosaurus saharicus (since destroyed) was likely not referrable to that Algerian tooth taxon at all. The authors provide previously unpublished photographic evidence of the original type, which allows a more detailed comparison to fossils discovered in the late 20th and 21st Century. This comparison reveals autapomorphies to the Egyptian dinosaur, requiring a new taxon name.

The authors have done an excellent review of the (oh so many!) questionable referral of various bones to taxa without overlapping material, when the different bones are sometimes separated by kilometers. If the information in p. 10 ¶3 could be summarized in a table (in particular, the OTU used in each “separate” and “merged” configuration), that would be appreciated.

But please note that they themselves repeat an invalid referral by taking some of Stromer’s writings at their word. Stromer’s (1931, 1934) comparisons of various African dinosaurs to “Dryptosaurus” is not to the New Jersey tyrannosauroid Dryptosaurus aquilunguis; rather, it is to the dinosaur universally accepted today as Albertosaurus sarcophagus (but which was in the late 19th and early 20th Century referred to the tooth taxon Laelaps/Dryptosaurus incrassatus. See Stromer’s footnote on the bottom of page 56 in his 1934 paper:

“Nach PARKS (1928, S. 5-7) ist Dryptosaurus LAMBE (1904, S. 20, Raf. VI, Fig. 6) warhscheinlich mit Albertosaurus sarcophagus OSBORN identisch.”

And his reference in the 1931 paper to the North American Dryptosaurus refer to Lambe’s 1904 paper of the skulls which became the type and coptype of Albertosaurus sarcophagus in Osborn 1905. Thus, in this paper of confusing theropod taxonomy, some proper clarification should be made that Stromer’s use of “Dryptosaurus” is actually Albertosaurus. (See p. 20 ¶3)

(Parenthetically, it seems bizarre to me that Depéret and Savornin included the most labiolingually compressed large theropod teeth known into the same genus as dinosaurs with the most labiolingually expanded teeth (source of the trivial nomen “incrassatus”: thickened or inflated)).

An aside: I appreciate the evocative name Alllissaurus (marauder reptile), but note in advance that this name is destined to be consistently misspelled (triple “l”s are rare), and will almost assuredly be autocorrected (either in word processors or web searches) to “Allosaurus”! This isn’t a reason not to use the name by any means, but simply looking ahead

p. 35, ¶3 The authors should make clear the definition of “Allosauroidea” they are using. I am assuming (given the authorship of this manuscript) they follow Rahut and Pol (2019), namely the most inclusive clade containing Allosaurus fragilis but not Megalosaurus bucklandii or Neornithes. Some previous node-based definitions would be potentially synonymous with Allosauria or the new Carcharodontosauriformes depending on the configuration of the allosaurs they found.

Cladograms: Note that Alpkarakush has yet to be published; please make certain of the timing of the publication of this paper vs. the Alpkarakush paper so that the authors do not trump themselves.

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Reviewer #1: No

Reviewer #2: Yes: Thomas R. Holtz, Jr.

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PONE-D-24-16876R1Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogenyPLOS ONE

Dear Dr. Kellermann,

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.

ACADEMIC EDITOR:

Dear Dr. Kellermann,

We appreciate you submitting your manuscript to PLOS ONE and thank you for giving us the opportunity to consider your work.

I have completed my evaluation of your manuscript, which has been reviewed by two highly qualified reviewers all of whom agree it is worth to be published in PLOS ONE. However, I agree with Reviewer 1 that the taxonomic part erecting a new species is problematic. Therefore, I kindly ask you to review that part and resubmit your manuscript after addressing Reviewer 1’s comments below. Please, note that your revised submission may need to be re-reviewed.

PLOS ONE values your contribution and I look forward to receiving your revised manuscript.

Yours sincerely,

Dr. Olga Spekker

Please submit your revised manuscript by Sep 16 2024 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.

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We look forward to receiving your revised manuscript.

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Olga Spekker, Ph.D.

Academic Editor

PLOS ONE

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Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

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

Reviewer #1: (No Response)

Reviewer #2: (No Response)

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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: Partly

Reviewer #2: Yes

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

Reviewer #1: N/A

Reviewer #2: Yes

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Reviewer #1: Yes

Reviewer #2: Yes

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Reviewer #1: Yes

Reviewer #2: Yes

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6. Review Comments to the Author

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Reviewer #1: I thank the authors for the detailed reply to my comments.

I am sorry for not being completely able to follow their final conclusion, but the epistemological issues I remarked in the first review have not been addressed in a satisfactory way, and thus I cannot endorse the erection of a new genus/species based on the material destroyed in 1944.

The authors are right in listing four possible solutions to the controversy. They (the authors) have been very rigorous in following the ICZN rules, and I applaude their detailed argument. Yet, a recommendation (in this case, to not erect a new taxon) is not followed/ignored because of the Code itself, but on the basis of the Precautionary Principle, which is logically and epistemologically superior to the code itself. Accordingly, the only conclusion I endorse is the #3 among those listed by the authors (quoted below):

"We leave the Moroccon skull as neotype for Carcharodontosaurus saharicus, simply state that we think that the Egyptian material represents a different taxon, but do not propose a new binomen for it.

Just because the Egyptian material cannot be referred to the same taxon of the Moroccan material (a conclusion I can endorse) does not automatically mean that the former could be used to erect a new taxon. The authors have not addressed the main question of my review: what if the species they erect was not a real taxon, and based on erroneous interpretation of Stromer's publications? What if Stromer's words used to define such novel taxon are (even in part) erroneous? How could such taxon be falsified/revised without a testable specimen to work with? Any new species is a particular taxonomic hypothesis anchored to a testable type specimen. This is the reason for introducing type specimens housed in valid institutions: to avoid the proliferation of untestable taxonomic hypotheses. Without a testable specimen, any detail of such hypothesis is in an epistemological limbo, because we cannot replicate Stromer's observations used by the authors to erect that taxon. If two distinct authors disagree on some of the statements by Stromer used here to define the new species, how could such disagreement be solved? The objectivity principle dictates replicabile statements based on accessible specimens: in absence of such specimen, and pending the discovery of a second specimen supporting Stromer's description, no new species should be erected.

I invite the authors to follow the most conservative and most robust conclusion among those they listed, the less speculative and the one with potentially the less negative impacts: to discuss the possible differences between the Egyptian and Moroccan carcharodontosaurs according to the published material, but to avoid the formal erection of a new taxon.

The authors seem unaware that the formal erection of a new species based on lost material cannot be falsified pending additional material, being such taxon based only on Stromer's words: they based the diagnosis on Stromer's authority and not on replicable observations. The authors stated that Stromer's publications are a valid source of information: in principle, I do not question it. What I remark is that without an objective evidence (a fossil specimen) we cannot properly test the statements in those publications using an independent source.

The authors are invited to address my concern: how could we falsify Stromer's words in the case they (the words) were wrong? Only a second specimen could falsify those words. But if the species is not a real taxon, such second specimen does not exist, and would never be found. In particular, in the case of non-existence of such taxon, no additional specimen actually exists, and we would never be able to say anything about the validity/revision of that taxon. In that case, the species would merely be a non-falsifiable hypothesis included in our taxonomies and phylogenies. This must be avoided.

The authors listed a series of fossil taxa based on lost material in support of their preferred option (e.g., Maraapunisaurus). Yet, this is not a solid argument. First, I note that all of them are based on very fragmentary material (so, according to the conclusion of the manuscript, all of them would be considered nomina dubia even if based on accessible material). Second, all of the listed taxa are very problematic and barely mentioned in literature, thus of limited impact and significance. The exclusion of such taxa from the realm of the valid taxa would not impact in a significant way the progress of the palaeontological knowledge.

I encourage the authors to ask themselves: is the erection of this new taxon really necessary? Do we need another problematic African theropod to the list of the controversial dinosaur taxa? The authors provided examples of already named taxa based on available material from North Africa (so, taxa which at least epistemologically are more robust than the new species they wish to introduce) which are of controversial status: is the inclusion of another problematic species really necessary? I conclude that the taxonomy and inclusiveness of the North African theropods is sufficiently controversial and does not need another problematic species. Such new taxon would likely ignite a long debate on its validity, a debate unfortunately impossible to resolve if the species is not a real taxon and no material is available. Is another (potentially endless) debate on the validity of some dinosaur really welcome?

In summary, if the authors are right, only a second specimen would support their hypothesis: unfortunately, such second specimen does not exist, and pending its discovery, we cannot accept the institution of a taxon based uniquely on two photographs and the untestable acceptance of Stromer's words.

Note that from a palaeontological perspective, I have not a priori objections to the hypothesis that the Egyptian carcharodontosaur could be distinct from the Moroccan one. What I question is the epistemological basis for such hypothesis. In conclusion, the main elements of the manuscript in its current form, excluding the erection of a new species, are sufficient for a solid and significant paper: removing any mention to a new taxon would not impact the relevance and importance of the manuscript, and would likely be considered more positively than a manuscript introducing such a controversial and problematic species actually based on a destroyed fossil and just a few photos.

Reviewer #2: This version seems to have addressed all the issues that I and the other reviewer had previously suggested.

I did come across two typographical errors:

p. 25, ¶3 “Mapusaurus” is misspelled in the middle of the paragraph.

p. 26, ¶1 “carcharodontosaurids” is misspelled on the first line of the page.

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Reviewer #1: No

Reviewer #2: Yes: Thomas R. Holtz, Jr.

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PONE-D-24-16876R2Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogenyPLOS ONE

Dear Dr. Kellermann,

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 Oct 18 2024 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.

Please include the following items when submitting your revised manuscript:

• A rebuttal 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,

Olga Spekker, Ph.D.

Academic Editor

PLOS ONE

Journal Requirements:

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.

Additional Editor Comments:

Dear Dr Kellermann,

We appreciate you submitting your manuscript to PLOS ONE and thank you for giving us the opportunity to consider your work.

I have completed my evaluation of your manuscript, which has been reviewed by three highly qualified reviewers during the different stages of the review process – all of the reviewers agree it is worth to be published in PLOS ONE. However, as I already highlighted in the last review round, I agree with one of the original reviewers that the taxonomic part erecting a new species is problematic. Therefore, I invited another reviewer in this review round, who also agrees with us that “introducing a new taxon in this circumstances is highly problematic and not warranted”, Based on the above, I kindly ask you to review that part and resubmit your manuscript after addressing the reviewer’s comments below. Unfortunately, without that, your manuscript cannot be accepted. Please, note that your revised submission may need to be re-reviewed.

PLOS ONE values your contribution and I look forward to receiving your revised manuscript.

Yours sincerely,

Dr Olga Spekker

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Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

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

Reviewer #3: (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 #3: Partly

**********

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

Reviewer #3: Yes

**********

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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 #3: Yes

**********

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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 #3: Yes

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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 #3: I congratulate the authors on their detective work and agree with them in many regards (e.g., authors need to be careful when comparing seemingly coeval assemblages from different parts of North Africa). The descriptions and observations are good for the most part and overall the MS is well-written.

Having said that, I concur with previous reviewers that it is highly problematic to publish a new genus and species name for a set of fossils that were destroyed many decades ago, based on drawings and photographs (I should add that I examined one of the photographs first hand). Renaming old - and most importantly lost - material could set a very problematic precedent indeed. It would be an entirely different situation if relevant new material had been found in the Bahariya Fm, but that is not the case here. The two other reviewers have already made some valid points, which I think have not been fully addressed. I am adding a few more here.

(1) The difficulty of reconciling Stromer's observations with the (sometimes difficult to interpret) photographs and drawings (which do not always match the photographs) in absence of an actual, physical set of fossils is evident in many passages, e,g,.:

"An expansion of the narial fossa to the nasal in the form of a supranarial fossa, as it is present in Acrocanthosaurus, Concavenator, Carcharodontosaurus and Meraxes [11,18,43,44] is not clearly visible in the photo of SNSB-BSPG 1922 X 46, but Stromer [1] described the area of the anterior end of the nasal lateral to the median premaxillary process as slightly concave. However, a clearly defined rim of such a fossa, as it is present in Carcharodontosaurus ([18]:Fig 134A) is not at all indicated in Stromers figures, nor visible on the photograph."

What are we to make of ambiguous aspects of the description, which cannot be resolved? As interesting as the new photograph is, the case made here to erect a new taxon is on very shaky ground, as many questions remain and the photograph, while of huge interest historically, does not offer a huge amount of new insights anatomically. This is not meant to downplay the scientific information the authors have carefully extracted from the image (well done), but simply reflects the limitations of an old photograph.

(2) Even if all of these issues are set aside, I don't think a highly convincing case can be made at this stage that this is a different genus (as opposed to, for example, a different species, as has been done with Carcharodontosaurus iguidensis, and even that would need carefully balanced arguments, see below). These decisions are obviously notoriously difficult to comment on when dealing with extinct taxa, as they are often somewhat arbitrary and based on different approaches to naming taxa (splitters vs lumpers etc). Either way, it would have been good and very important to include detailed comparisons to other theropods, known from large sample sizes (e.g. Allosaurus, Coelophysis), as these often show a rather remarkable degree of variation (reflecting everything from ontogeny to individual variation etc), especially in their skulls.

Several features highlighted by the authors may not be as unique as claimed here, including but not limited to:

- The horn-like rugosity may reflect sexual dimorphism.

- Speaking of rugosity, isolated specimens from the Kem Kem (some not accessioned in public collections) show a wide range of different degrees of rugosity (which is at odds with the author's comment that the Egyptian specimen shows ridges, furrows and rugosity that are far more pronounced than in the Moroccan Carcharodontosaurus). To be fair, as much of this material is not publicly available, one cannot blame the authors, but it might be of interest to them.

- Comparisons between the braincases are interesting but should come with lots of caveats: the Berlin specimen (endocast) is not very detailed (i.e. it is not necessarily the most faithful and undistorted endocast) and meaningful comparisons are consequently difficult.

- Note that teeth matching the shape of Stromer's specimen have been found in Morocco (the authors only mention one such tooth, figure 8E, but others have been collected), so in this regard, the Egyptian specimen is not necessarily unique. Note also that carcharodontosaurid teeth found in the Kem Kem show quite a bit of variation, likely reflecting different positions in the jaws.

Some other aspects may well represent meaningful differences, but for reasons outlined above (and below) I maintain that introducing a new taxon in these circumstances is highly problematic and not warranted.

(3) Geography. " Since Stromer’s referral of the Egyptian specimen to the Algerian M. saharicus is questionable and the Moroccan material was found in closer proximity to it, we accept the designation of the latter as neotype of C.

saharicus" - - - This is not convincing. If one looks at the geographical range of extant animals like Nile crocodiles, lions, hippos etc it is evident that they range across huge parts of the African continent (certainly historically). There is no particularly convincing reason to believe that theropod dinosaurs were not ranging across vast areas of Africa, so we need to be careful when arguing for taxonomic assignments based on a distance in km from one site to another. These arguments should be based on unambiguous anatomical observations etc, and not on the distance from Morocco to Algeria vs the distance from Algeria to Egypt...

(4) The authors argue for an exceptional case (avoiding the strong recommendation that scientists not designate as type material specimens the author only knows from descriptions and illustrations, but has not studied first hand) by stating that "1) The material described by Stromer [1] has played a pivotal role for our understanding of carcharodontosaurid anatomy, phylogeny and evolution". This is a problematic statement in the sense that, as this paper amply demonstrates, very little is actually known about the Egyptian material. It certainly played an important historical role, but to claim this long lost material played a pivotal role in carcharodontosaurid phylogeny and evolution is somewhat at odds with one of the key messages of this paper: that we know so little about this Egyptian material that much of what people have suggested about this material in the past - including its name and the referral of other material to it - is now in doubt according to the authors. If we contrast this with, say, Acrocanthosaurus, which has been described in great detail, the difference becomes quite clear.

In summary, I don't think it is wise or warranted to erect a new taxon in these unusual circumstances. Note also that, contrary to one claim made here in response to reviewer comments, taxa do not have to be named to be noticed in other papers. Indeed, the unnamed Kem Kem abelisaur, or the (previously unnamed) Kem Kem tapejarid were not neglected at all and found their way into numerous papers. The authors need not worry, I am sure that this paper is going to elicit a great deal of interest and it will be an essential reference for many future papers on Cretaceous North African dinosaurs - going down a slippery slope and naming a long-destroyed set of bones is not necessary to achieve this.

A few other comments:

- "S1. Select measurements of Alllissaurus markgrafi in mm." I assume this is from a previous version of the MS?

- The age of the Bahariya Fm (and the Kem Kem Group, and possibly roughly coeval strata in Algeria) is highly uncertain, and the Cenomanian age for many of these North African assemblages published by many authors in the past is based on very limited data (and a decent dose of circular reasoning).

- "Although we do agree that differences between Sauroniops and Carcharodontosaurus saharicus pointed out by Cau et al. [186–188] are intriguing, we agree with Cau et al [186] that this single frontal bone does warrant the erection of a new genus or species and instead consider Sauroniops a nomen dubium, pending the discovery of new

material." - - - - I think the authors meant to say "we disagree with Cau et al etc.

- When discussing faunal overlap, it is important to be clear about what the comparisons are based on. It is worth noting that only one sauropod genus is known from the Kem Kem Group and only two have been described from the Bahariya Fm. There may well be overlap, including at the genus level, and the discovery of isolated titanosaur remains in Morocco (which may turn out to be very similar to Paralititan, for example) certainly leaves some room for potential overlap. Outcrops in the Kem Kem are far more extensive, and A LOT more material has been found, therefore we need to be careful when it comes to how much importance we give to the presence/absence of some of the crocodyliforms etc. We are really not comparing like with like at all, so we need to be careful when deciding how much importance we give to the presence of seemingly identical or very similar taxa based on isolated remains (as the authors correctly point out), but also when we give too much importance to the absence of shared taxa (something the authors are not doing here) because, again, we are comparing one locality that has barely been explored in the last century and another one that has been explored - exploited one might say - by large numbers of commercial collectors (many Kem Kem specimens, including most of the crocodyliforms the authors refer to here as examples of taxa unique to Morocco, were commercially acquired and often illegaly exported) in addition to a handful of scientific expeditions.

- and it seems that the anteromedial corners of the fossae were overhang by a lamina, as it is the case in other carcharodontosaurids [52]. - - - This should read "as is the case".

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Reviewer #3: No

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Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny

PONE-D-24-16876R3

Dear Dr. Kellermann,

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.

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Olga Spekker, Ph.D.

Academic Editor

PLOS ONE

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