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  • The initial submission of this article was received on August 15th, 2018 and was peer-reviewed by 2 reviewers and the Academic Editor.
  • The Academic Editor made their initial decision on August 31st, 2018.
  • The first revision was submitted on October 3rd, 2018 and was reviewed by the Academic Editor.
  • A further revision was submitted on October 9th, 2018 and was reviewed by the Academic Editor.
  • The article was Accepted by the Academic Editor on October 16th, 2018.

Version 0.3 (accepted)

· Oct 16, 2018 · Academic Editor


Thank you addressing my final comments and adding the missing reference. Looking forward to seeing you manuscript published.

# PeerJ Staff Note - this decision was reviewed and approved by Andrew Farke, a PeerJ Section Editor covering this Section #

There was one final requirement raised by the Section Editors - specifically, they feel that the reference to the future deposition of the specimen should be removed (to avoid confusion, and in case the deposition doesn't happen as planned). Specifically, they request that the following text be removed from the manuscript: "and will be deposited in the Three Gorges Entomological Museum, Chongqing, China after 2027" (present at lines 13-14, pg 5, Materials and Methods). You can work with our Production group to remove this text.

Version 0.2

· Oct 5, 2018 · Academic Editor

Minor Revisions

Thank you for adressing our comments and suggestions. particularly adding information on apomorphic features linking previous finds to parasitic beetles. Your manuscript is as good as accepted. I just found some minor issues which i would you to address for publication:

Line 49: it would help to clarify or mention an example of what you consider a relatively small insect - what is the smallest insect that has been documented as a host

Link 65: thank you for integrating this new reference (Nagler and Haug 2015), but please also add it to the references

Line 262: add space between "body," and a

Line 273: please also take this opportunity to thank the reviewers.

Version 0.1 (original submission)

· Aug 31, 2018 · Academic Editor

Minor Revisions

This is a nice description of the first unambiguous Cretaceous strepsipteran immature. I would love to see this published, but there are still some minor, but crucial points I would like to see addressed before publication.

The main points are:

Direct versus indirect evidence of parasitism: you present an important case of direct evidence for parasitic immatures. The general importance of direct evidence rather than indirect evidence for inferring parasitic life stages has been discussed before (e.g., Nagler & Haug 2015). I feel it would benefit the reader by highlighting that presence of parasitic larval stages is often inferred based on the present of adult representatives (not just in Strepsiptera; see Nagler and Haug, 2015) and not by direct finds of parasitic larval stages making this an exceptional find.

Bearing on interpretation of previously reported putative strepsipteran immatures: I agree that these are the first unambiguous crown-group strepsipteran immatures. However, I miss a discussion on what a stem-group strepsipteran immature could have looked like. I concur that this is hard to do, but at least some informed suggestions can be made. You can potentially speak of stasis in this lineage – although ideally one would have multiple samples / taxa demonstrating this to be since their inception. How can you rule out that different strategies or intermediate morphologies (with that seen in their sister-lineage) not existed in early representative or now-extinct lineages of Strepsiptera ? Particularly, since this group has an ancient origin inferred to be traced back to the Late Carboniferous or Early Permian... Morphological intermediate lineages could be extinct today. Based on the discovery of a single immature, you can only make limit inference on other larval stages present in this interval. The rejection of previous interpretation and re-assignment should be based on characters and is most likely discussed in other articles. I am sure these are trivial to you, but it is currently not obvious to non-expert readers and should be at least briefly discussed. Does it share more characters with parasitic beetles than with Strepsiptera? How much do we know about larvae across both sister lineages. Even so can you rule out that this state represented in beetles could be ancestral for Strepsiptera? I therefore feel you should rephrase these aspects and express yourself more carefully relating to some of these aspects. Some suggestions are made in the annotated pdf.

Please address the comments of the reviewers and those listed in the annotated pdf, in addition to these points.

Suggested reference:
Nagler C, and Haug JT. 2015. From Fossil Parasitoids to Vectors: Insects as Parasites and Hosts. Advances in Parasitology 90:137-200.

Reviewer 1 ·

Basic reporting

The study is thorough and generally the conclusions are appropriate. There are numerous minor edits needed in sentence structure and grammar. For example on line 62 and 63 the last word “yet” should be removed. This is just an example of many minor changes needed to improve the writing of the paper.

Experimental design

no comment

Validity of the findings

• There is little doubt that this specimen constitutes a strepsipteran primary larvae and therefore the interpretation of this specimen is important to science.
• It is valuable that the authors discuss and correct the use of larval terms for the first instar larvae of Strepsiptera. The terms planidium and triungulin are obviously not appropriate for Strepsiptera larva but have been repeatedly used incorrectly.
• In lines 155-156, there is no need to note that wings or wing buds are missing. No evidence of these would be expected in a strepsipteran larvae or any larvae that might be mistaken for this order, such as Meloidae, Rhipiphoridae, …
• The term hatching in line 244 in not appropriate. The larvae emerge from the female after hatching internally from an egg. The term in this sentence should be changed to emerging.
• In more than one place in the manuscript it is suggested that there must be a large number of primary larvae produced by each female but this appears to be speculation and if included should be identified as such. Based on knowledge of current strepsipterans, this could be true but the authors should not rely so heavily on this interpretation as fact. The reality is that we know nothing of the natural history of these larvae or their numbers from this geological age. Unless the unlikely event of the discovery of a gravid female of this age it may always be speculation as to the relative number of offspring. Likewise, lines 243-246 are stated as fact but is speculation for the species covered in this study.

Additional comments

This is a valuable paper that appears to only need minor changes. My suggestions are offered to try to improve an already generally good paper.


Basic reporting

The manuscript fully meets the standards and is within the scope of the journal. It provides sufficient backround and introduction to place the work into the broader field of knowledge.The text is is written in professional, unambiguous language, so an international audience can clearly understand. The sources are adequately and appropriately cited. Very important part of manuscript are the figures, which are well described, in a high quality and sufficient resolution (as much as possible). The text is organized into coherent subsections.
The results are definitely ‘self-contained’, while results are relevant to the hypothesis.

Experimental design

It is clearly stated how research fills and identified knowledge gap. The investigation was conducted to very high technical standard, the methods are comprehensive and described sufficiently to be reproducible.

Validity of the findings

Conclusion are limited to supporting results and are appropriately stated. Speculations are identified as such.

Additional comments

According to my opinion, there are a few notes that need more details or to be slightly modified, usually to ensure that an audience can clearly understand the text. I cannot find any serious weakness.

Line 19: Mesozoic origin – it could be „at least“ the Mesozoic origin
Line 21: „one of the most“ or the most?
Line 42: Bahiaxenidae probably also leave their host – is it supported by references?
Line 50: It is not clear, whether labrum is completly missing, or it is „largely reduced“ (written in Lines 163-166) / or fused with other mouthparts – „lacking separate labrum“ (Lines 204, 205), so I would prefer "not present as separate element".
Line 56: „enable the minute larvae to jump very efficiently“ – not true for the family Stylopidae, so I would kindly suggest "for the most" of families"
Line 84: I am not familiar with an abbreviation „s.b.“
Line 85-87: Is the „planidium“ always „legless“? Aren’t also first instar larvae of Meloidae (with legs) reffered as „planidium“? This term is derived from the Greek word, which should mean „wanderer“, so I assume the term is not such a problem in case of Strepsiptera.
Lines 87-94: Although I agree with neutral term „primary larva“ for first instars of Strepsiptera, term „triungulinid“ (= triungulin-like) also would not be such a problem for case of Strepsiptera, because it is not the same as „triungulin“ (without -id).
Line 101: I would suggest to add the final years again – "endoparasitism is extended back by ca. 50 milion years to minimum age of ca. 100 mya"
Line 106: Better to use mya consistantly than „Ma“
Line 163: (st in Figs. 1A, B, 3A)
Line 187: Sternites IV-VI with two setae, not Sternites III-IV
Line 197: Mengenilla chobauti
Line 251: Typo (missing spacebar) „body, a …“
Line 357: Typo (missing spacebar) „plate, mx…“

After correcting minor ambiguities, I am very pleased to suggest the manuscript to be accepted for the journal.

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