Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea

Round 1

Reviewer 1 Report

Overall, I find the manuscript to be an interesting and valuable addition to our knowledge about the fossil beetle families. The author put significant effort to trace the taxonomic history of the fossil taxa and summarize the current knowledge, providing also his comments and opinions. It will serve as a good reference for future research on extinct beetle groups. I recommend it for publication after the language check (the style in some cases and multiple misspellings) and some minor, technical changes (see comments below):

In Abstract:

- Jarzembowski, Wang et Zheng. 2019: dot should be replaced by a comma

- Jarzembowski, Yan, Wang et Zhang. 2013: dot should be replaced by a comma

- Tan et Ren 2009: comma is missing

- Tan et Ren. 2009: dot should be replaced by a comma


In Introduction:

- line 235: “curried out” should be replaced with “carried out”

- line 253: ZMUC is currently changed to NHMD that stand for Natural History Museum of Denmark


In SYSTEMATIC PART:

- line 312: replace “XXth” with “20th”

- line 323: dash before the year span (– 303.4–298.9 Ma) looks confusing, I suggest to remove it here and thereafter

- line 383: recent should start from small letter, not capital

- lines 386-387: change the sentence “Also, there are some problems in the systematic placement of the taxon Jurodidae Ponomarenko, 1985 [8] has some problems.” to “Also, there are some problems in the systematic placement of the taxon Jurodidae Ponomarenko, 1985 [8]”

- line 395: remove one “member” word from the sentence

- line 547: change “more ior less” to “more or less”

- line 715: change “elytal venation” to “elytral venation”

- line 753: change “impossibly to clartify” to “impossible to clarify”

- line 927: replace “époque” with “epoch”

- lines 929-930: change “the groups slightly structurally modified groups” part of the sentence

- line 936: using a word “probabilistic” could suggest there is an involvement of the theory of probability taken into account, I believe the author meant that the assignment of fossils is rather arbitrary depending on the author’s idea when diagnostic characters are not observable

- lines 965-966: “94.3 - 89.3 Ma” should be written with a dash, not a hyphen

- line 1043: change “synonynymized” to “synonymized”

- line 1049: “130.0 - 125.5 Ma” should be written with a dash, not a hyphen

- line 1386: change “pronotum ha/s” to “pronotum has”

- line 1421: change “promiment” to “prominent”

- line 1483: Rhopalomma should be in italics

- line 1552: change “Mosozoic” to “ Mesozoic”

- lines 1734-1735: in the sentence “Later it was transferred at first to Zygadenia [110] and later authomatically to the restored” remove one of the “later” word and change “authomatically” to “automatically”

- line 1738: change “synonymymized” to “synonymized”

- lines 1765- 1769: names of species and genera should be in italics

- line 2188: change “arganic” to “organic”

- line 2204: change”07” to “0.7”

- line 2258: change”06” to “0.6”

- line 2283: change “Pononarenko” to “Ponomarenko”

- lines 2479-2480: what is “modern amber”? I suppose there is some mistake here

- line 2597: Priacma should be in italics

- line 2681 and 2718: “48.6 - 40.4 Ma” should be written with a dash, not a hyphen

Author Response

All recommendations of the reviewer 1 are very useful and accepted. I greatly appreciate him for them, although the “recent”, because this adjective in some places in the text has common meaning but in others has terminological sense (Recent epoch, Recent fauna).

Author Response File: Author Response.pdf

Reviewer 2 Report

Geosciences mdpi: review of Kirejtshuk ts

This is a comprehensive account of current phylistic (v. cladistic) classification of 'primitive' (archostematan) fossil beetles focusing on genera, the cupedine part augmenting Kirejtshuk et al. (2016). It is proposed as first part of a series on fossil beetles, a hyperdiverse order of insects.

A tribal (contra family) classification by other authors is rejected which is not surprising as the genera are based on similarities and distinguished by hiatuses with some (inevitable) uncertainties. (Whilst an attractively simple methodology leaving genera in free space, similarities can be widely shared, and hiatuses be due to unique specialisation, neither of which reveal relationships: 'primitive' genera in particular need careful search for shared advanced characters to reveal the underlying natural classification, especially when they are morphologically close to ancestral beetle forms, as in this study.)  Phylogenetic  considerations (e.g. testing for monophyly and recognition of homoplasy) are, however, reserved by the author for a future paper. It is thus only possible to comment on the current working classification, comparing the narrative with previous systems (notably the phylistic monograph by Ponomarenko, 1969), and especially in the light of recent finds in Cretaceous/tertiary amber. It could be published subject to revision based on several questions/comments included below (and those of the second reviewer).

The Palaeozoic taxa, e.g. tshekardocoleids, are a wholly stem group(s) not preserved in amber, but Cretaceous inclusions include representatives of 'living fossil' groups: ommatines/ommatids and cupedines/cupedids with implications (/) also for recent fauna. In the former living family group, the recent genera Tetraphalerus (South America) and Omma (Australia) have been traditionally recognised in the fossil record as extending back to the Mesozoic (even though the type genus has no tertiary record). The author's placement of fossil Tetraphalerus in a new fossil genus (Allophalerus gen. nov.) is not unreasonable due to morphological disparity compared with recent Brazilian Tetraphalerus;  but why is a fossil genus (Monticupes stat. nov.) also resurrected for the same genus (Tetraphalerus)?

 By contrast, Omma is retained (Omma janetae sp. nov.) and a new species of its fossil tanian sister (Cionocoleus pulcher sp. nov.) is described; this begs the question, is Polyakius gen. nov. a crypto-genus  of the former? —

And could the two rock-fossil species of Cionocoleus from the same deposit (Yixian Formation) be just preservational variants as suggested previously? —

 Furthermore,  why does C. pulcher appear to have (translucent) elytral cells , the absence of which is an original (e.g. ponomarenkoan) generic feature of Cionocoleus?

The relatively large extinct genus Brochocoleus [tanian sister of Omma + Cionocoleus] is split trichotomously (including genera Jarzembowskiops gen. nov. and Diluticupes stat. nov.) elevating two previous species groups: why therefore not propose subgenera? —

The third genus, Burmocoleus gen.nov., comprises two similar-looking species from the same amber deposit: could they be sexual variants (if not plastically deformed)?

Also in Cretaceous amber, a proposed new species , Clessidromma hirsutum sp. nov., lacks the narrow waist of the congeneric type species (a unique archostematan feature somewhat resembling  scarite ground beetles); C. hirsutum resembles the type species of Lepidomma in general shape, but differs in the elevation of the pronotum  (which Lepidomma shares with Clessidromma); Lepidomma also has a unique scaley edging (v. spines in Clessidromma)—

why is Lepidomma therefore synonymised with Clessidromma (and not even considered as a subgenus) when the intermediate similarity of the well-preserved C. hirsutum sp. nov. is clearly offset by such disparity, the latter actually acknowledged by the author? (—

In contrast, Brochocoleus is split three-ways rather than combined: is this simply because of its relative size, i.e. it is a longer-ranging genus with more included species?)

In Stegocoleus, mention that S. arkonus and S. lawrencei were based on microscopic features not analysed in other species/specimens  and shape differences were not considered taphonomically? 

Form genera (as in palaeobotany) are not recognised in palaeozoology (International Code of Zoological Nomenclature) and Zygadenia is actually now a collective for Notocupes-like species only known from organ taxa (elytra): —

why are the body fossils of this extinct group (with more taxonomic features) not referred herein to Notocupes (and not Zygadenia)?--

In amber Notocupes, N. neli is rather similar to N. ohmkuhnlei?

Why is Mallecupes prokini sp. nov. not compared with Mallecupes cleevelyi (which also has a distinctly pointed pronotum and angular temples ) rather than Mallecupes qingqingae (which does not)?—

enc.: is this a possible picture of M. prokini (from NIGPAS)?

Why are the (subtle) differences in M. qingqingae not variation? (All the above Cretaceous amber specimens come from the same Asian forest.)

Regarding , the cupedine update, worth mentioning that the tertiary genus Cupes is interpreted sensu lato? (Compared to some recent classification.) 

Will the useful (but now dated) online taxonomic catalogue of fossil beetles be updated?

Why are lns 59-94, 502-508, 908, 918, 1583-1659, 1993 underlined?

Ln 1660 etc Correct stratigraphy! (Purbecks are Berriasian but Clockhouse brickworks is Late Hauterivian, Smokejack's Early Barremian, Beare Green Late Barremian)

Lns 101-150 need completing

Figure captions. Worth mentioning that the purpose of wetting fossil specimens with ethyl alcohol  (as opposed to covering them) is to reduce surface interference/improve contrast for short periods only to facilitate observation/photography?—

Less invasive alternatives exist for amber (e.g. maple syrup) and propanol for some rock fossils.

Worth mentioning Fig. 1B is a beautiful stem beetle but is tectonically deformed?

— entmologists (like other zoologists) may be unaware of this when considering typological taxa based on fossils (which can also appear symmetrical).

Fig 23, H wrong in GPIH?

References not checked versus text but check ln 2963? and 3332 gap

Author Response

Some recommendations of the reviewer 2 are directed to improve the text and accepted with many thanks and others have proposed as a discussions and need some comments.
In particular:
- “ but why is a fossil genus (Monticupes stat. nov.) also resurrected for the same genus (Tetraphalerus)?” – first of all because of different elytral venation !!! this and other diagnostic characters are added in the comparison of Allophalerus.
-“By contrast, Omma is retained (Omma janetae sp. nov.) and a new species of its fossil tanian sister (Cionocoleus pulcher sp. nov.) is described; this begs the question, is Polyakius gen. nov. a crypto-genus of the former?” — each of the mentioned genera are provided by a diagnosis including the main characters necessary to identify every genus – the reviewer is right that new species of Cionocoleus from Burmese amber is certainly quite distinct from other congeners known from compression fossils in stone – in particular the new species has very different head which is markedly shorter than that in other congeners (with known head) and with very larger eyes – therefore a new subgenus was proposed for the new species vfrom Burmese amber.
- “The relatively large extinct genus Brochocoleus [tanian sister of Omma + Cionocoleus] is split trichotomously (including genera Jarzembowskiops gen. nov. and Diluticupes stat. nov.) elevating two previous species groups: why therefore not propose subgenera?” — all mentioned taxa have rather clear diagnoses and well separated from other generic ones – therefore the rank of them should be comparable.
- “The third genus, Burmocoleus gen.nov., comprises two similar-looking species from the same amber deposit: could they be sexual variants (if not plastically deformed)?” – yes, a probability of sexual dimorphism can be supposed – however the sexual dimorphism in the elytral base is unknown and therefore it is reasonable that this and other characters should be preliminarily better to interpret as specific rather than intraspecific.
- “Also in Cretaceous amber, a proposed new species , Clessidromma hirsutum sp. nov., lacks the narrow waist of the congeneric type species (a unique archostematan feature somewhat resembling scarite ground beetles); C. hirsutum resembles the type species of Lepidomma in general shape, but differs in the elevation of the pronotum (which Lepidomma shares with Clessidromma); Lepidomma also has a unique scaley edging (v. spines in Clessidromma)— why is Lepidomma therefore synonymised with Clessidromma (and not even considered as a subgenus) when the intermediate similarity of the well-preserved C. hirsutum sp. nov. is clearly offset by such disparity, the latter actually acknowledged by the author?” — at the moment all three species have comparable differences and therefore these three species can be treated as three species of one genus or three same supraspecific taxa can be proposed for each of them – probably further species of this group will clarify the situation and may be it will be reasonable to change their interpretation.
- “In contrast, Brochocoleus is split three-ways rather than combined: is this simply because of its relative size, i.e. it is a longer-ranging genus with more included species? In Stegocoleus, mention that S. arkonus and S. lawrencei were based on microscopic features not analysed in other species/specimens and shape differences were not considered taphonomically?” – yes further revision of this group is very advisable.
- “Form genera (as in palaeobotany) are not recognised in palaeozoology (International Code of Zoological Nomenclature) and Zygadenia is actually now a collective for Notocupes-like species only known from organ taxa (elytra)” — this is rather delicate problem and this group can be split after a detailed analysis.
- “why are the body fossils of this extinct group (with more taxonomic features) not referred herein to Notocupes (and not Zygadenia)?”—perhaps I do not understand the question – nevertheless I agree that this group need a serious revision. “In amber Notocupes, N. neli is rather similar to N. ohmkuhnlei?” – I suppose that at the moment all Echinocups would be better to temporally consider as separate species till a detailed revision.
- “Why is Mallecupes prokini sp. nov. not compared with Mallecupes cleevelyi (which also has a distinctly pointed pronotum and angular temples ) rather than Mallecupes qingqingae (which does not)?” — the diagnoses of both species are included in the paper – if a further specimens will give additional information their taxonomic position can be revised. - “enc.: is this a possible picture of M. prokini (from NIGPAS)?” – this is the specimen from the Hamburg collection.
- “Why are the (subtle) differences in M. qingqingae not variation? (All the above Cretaceous amber specimens come from the same Asian forest.)” – such viewpoint should be proved. Mallecupes is very similar and apparently closely related to the Сenozoic Cupes – both can be treated also as only subgenera of the same genus. However the adequate choice between these alternatives can be done after study of more specimens.
- “Regarding , the cupedine update, worth mentioning that the tertiary genus Cupes is interpreted sensu lato? (Compared to some recent classification.)” – horizontal classification can be different from that elaborated with usage of the data from other geological time. - “Will the useful (but now dated) online taxonomic catalogue of fossil beetles be updated?” – of course - “Why are lns 59-94, 502-508, 908, 918, 1583-1659, 1993 underlined?” – this is a result of convertation of fonts in several soft-ware and all underlinings are corrected.
- Ln 1660 etc Correct stratigraphy! (Purbecks are Berriasian (in MS this age is mentioned) but Clockhouse brickworks is Late Hauterivian (in MS Valanginian/Hauterivian is mentioned FOSSILWORS: 136.4 - 130.0 Ma – 130 is the boundary according to ISC of 2018), Smokejack's Early Barremian (in MS this age is mentioned), Beare Green Late Barremian (in MS is mentioned FOSSILWORS: Late/Upper Barremian (130.0 - 125.5 Ma) it is corrected on ???? Barremian without LOWER) - “Lns 101-150 need completing” – it will be done in the proof
- “Worth mentioning Fig. 1B is a beautiful stem beetle but is tectonically deformed?” - certainly
— “entmologists (like other zoologists) may be unaware of this when considering typological taxa based on fossils (which can also appear symmetrical).” – this is natural process

Author Response File: Author Response.pdf