plazi/treatments-xml
2
0
Fork 0
Code Issues Pull requests Releases Activity
treatments-xml/data/F1/52/8B/F1528B080739546A933AEF6EB42F1903.xml

479 lines
74 KiB
XML
Raw Blame History

<document ID-CLB-Dataset="54713" ID-DOI="http://dx.doi.org/10.3897/evolsyst.7.104597" ID-GBIF-Dataset="bdbda4ca-7ebb-41c9-afa3-2066664a1552" ID-Pensoft-Pub="2535-0730-2-211" ID-Pensoft-UUID="2450BCB137055BB6B2E6185BCDA42840" ID-ZooBank="921F39DA40BB47968E362194D1E9A138" ModsDocID="2535-0730-7-2-211" checkinTime="1687815010481" checkinUser="pensoft" docAuthor="Yamamoto, Shuhei" docDate="2023" docId="F1528B080739546A933AEF6EB42F1903" docLanguage="en" docName="EvolutSyst 7(2): 211-235" docOrigin="Evolutionary Systematics 7 (2)" docPubDate="2023-06-26" docSource="http://dx.doi.org/10.3897/evolsyst.7.104597" docTitle="Cretognathus Yamamoto 2023, gen. nov." docType="treatment" docUuid="0EF18D3E-DC80-4E51-A14E-982C18DA3E51" docUuidSource="ZooBank" docVersion="5" id="2450BCB137055BB6B2E6185BCDA42840" lastPageNumber="211" masterDocId="2450BCB137055BB6B2E6185BCDA42840" masterDocTitle="The smallest stag beetles (Coleoptera, Lucanidae): hidden paleodiversity in mid-Cretaceous Kachin amber from northern Myanmar" masterLastPageNumber="235" masterPageNumber="211" pageNumber="211" updateTime="1732689453509" updateUser="ExternalLinkService">
<mods:mods id="3DEC9F2EF66E660A9B3D8493FB69A9B3" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="2836F7D9072E23766564AB4DD8B776F6">
<mods:title id="5F265C660F3357BCCB8735CB94D79B64">The smallest stag beetles (Coleoptera, Lucanidae): hidden paleodiversity in mid-Cretaceous Kachin amber from northern Myanmar</mods:title>
</mods:titleInfo>
<mods:name id="8525743201DBFE520B6CE676CBD3F261" type="personal">
<mods:role id="C0070213F4E4E3CDF6132F9A32D54496">
<mods:roleTerm id="CECC3AD6CEE0B00921F4107E1FCC88F8">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="F3B203C4BF816C3FF73318F5F6F6A81A">Yamamoto, Shuhei</mods:namePart>
<mods:nameIdentifier id="622D6C223EB5FE4B82E0E375C2084D2B" type="ORCID">https://orcid.org/0000-0002-4162-8457</mods:nameIdentifier>
<mods:affiliation id="956960D926EB38A3C79F86123FD09A37">The Hokkaido University Museum, Hokkaido University, Kita 10 - jo, Nishi 8 - chome, Kita-ku, Sapporo 060 - 0810, Japan</mods:affiliation>
<mods:nameIdentifier id="115D8DC0FE01CCC3F35172E4EE83E7A1" type="email">s.yamamoto.64@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource id="959F9C3AA7AEA4157A9700AEEE1E10E1">text</mods:typeOfResource>
<mods:relatedItem id="B67F830A8B63E1899C4B5CDB1B8832FD" type="host">
<mods:titleInfo id="261867A6AF03146C384BF81D70552B4F">
<mods:title id="B453A56F66E4F33E3629974386619786">Evolutionary Systematics</mods:title>
</mods:titleInfo>
<mods:part id="3220EB7702FC4B80F1A2121B19A791FB">
<mods:date id="F15005E25028B1CA43EF726F85F6E7BD">2023</mods:date>
<mods:detail id="8FD4D3F2DBF1E39CCA50956889811002" type="pubDate">
<mods:number id="1A5B204AA84BBABDFB67A9FA0860BEC9">2023-06-26</mods:number>
</mods:detail>
<mods:detail id="577F8D15641608157640E71C287B86E8" type="volume">
<mods:number id="3A0D89F34C78AA59B7199CDFE07AA1D2">7</mods:number>
</mods:detail>
<mods:detail id="0E0D55F3924ABAFD075353C7FFBA9DC5" type="issue">
<mods:number id="F19364047E31BA340F1005DBC856653C">2</mods:number>
</mods:detail>
<mods:extent id="8F901487951C3901E8739D596FC7CCCC" unit="page">
<mods:start id="E0189B01AE489E91F57CCD123DB296C4">211</mods:start>
<mods:end id="E6DC131742FC4C6E02FB06C18BDBA640">235</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location id="05CE3C453F949347741CCBF69D5EED85">
<mods:url id="7F31101044AAA04B4B70D65C1E9F7DCA">http://dx.doi.org/10.3897/evolsyst.7.104597</mods:url>
</mods:location>
<mods:classification id="2F836A879654C95FE78A9C76229400DE">journal article</mods:classification>
<mods:identifier id="5D014970EED564AB329088229D5DCF2F" type="DOI">http://dx.doi.org/10.3897/evolsyst.7.104597</mods:identifier>
<mods:identifier id="B7CBBA620943A64AD54ECC6EAF46C73D" type="Pensoft-Pub">2535-0730-2-211</mods:identifier>
<mods:identifier id="4AC36CAEFFE7D513B7B5289B8EAB056F" type="ZooBank">921F39DA40BB47968E362194D1E9A138</mods:identifier>
<mods:identifier id="2AC5FFE1896A85C2B9CF0E59825E9DD0" type="Pensoft-UUID">2450BCB137055BB6B2E6185BCDA42840</mods:identifier>
</mods:mods>
<treatment id="F1528B080739546A933AEF6EB42F1903" ID-GBIF-Taxon="210478290" LSID="urn:lsid:zoobank.org:act:0EF18D3E-DC80-4E51-A14E-982C18DA3E51" httpUri="http://treatment.plazi.org/id/F1528B080739546A933AEF6EB42F1903" lastPageNumber="211" pageId="0" pageNumber="211" scope_family="Lucanidae" scope_order="Coleoptera" scope_subFamily="Aesalinae" scope_superFamily="Scarabaeoidea" scope_tribe="Ceratognathini">
<subSubSection id="D59566B865CD6B274FC6229BCC3F2104" pageId="0" pageNumber="211" type="nomenclature">
<paragraph id="072E041C79A046E442D7663476E3A66A" pageId="0" pageNumber="211">
Genus
<taxonomicName id="DA6DAC1E89435F91D74E07963748E990" LSID="https://zoobank.org/0EF18D3E-DC80-4E51-A14E-982C18DA3E51" authority="Yamamoto, 2023" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus" status="gen. nov.">Cretognathus</taxonomicName>
<taxonomicNameLabel id="EFC359C7D05C54C6C64785515CB5DAD3" pageId="0" pageNumber="211">gen. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="C00E8C05139E779D592BF54A283DFE2A" pageId="0" pageNumber="211" type="description">
<paragraph id="3876FE99C3D4DEB414AA0704A997E561" pageId="0" pageNumber="211">
<figureCitation id="6A1334F01DE8FF3E5018D78E800E70AD" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. General view of amber pieces with lucanid inclusions. A, B. Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205, showing dorsal (A) or lateral (B) views of the holotype; C. Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206; D. Ceratognathini gen. et sp. indet. 1, SEHU- 0000121207; E. Ceratognathini gen. et sp. indet. 2, SEHU- 0000121208. Scale bars: 3.0 mm (A, B); 1.0 cm (C); 5.0 mm (D, E)." httpUri="https://binary.pensoft.net/fig/868859" pageId="0" pageNumber="211">Figs 1C</figureCitation>
<figureCitation id="FC787F6367350FFDE3319F1EC819497B" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. General habitus of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Dorsal view; B. Ventral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868864" pageId="0" pageNumber="211">, 6</figureCitation>
<figureCitation id="8DFBBC42F2E528C79057D8CCE4DAD23A" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Head with right antenna, dorsal view; B. Head with right antenna, ventral view; C. Enlargement of A; D. Right antenna, ventral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (A, B); 0.2 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211">, 7</figureCitation>
<figureCitation id="940E51CCAC374E5B09EA9A984F8AEF59" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">, 8</figureCitation>
<figureCitation id="D50E490884C54548A894A24F4392BDF3" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Right protibia and protarsus with arrow showing row of small teeth between large spines along outer edge of protibia, dorsal view; B. Right protibia with arrow showing row of small teeth between large spines along outer edge of protibia, ventral view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Left metatibia and metatarsus, dorsolateral view; E. Left mesotarsus, ventrolateral view; F. Left mesotarsus, dorsolateral view; G. Left metatarsus, dorsolateral view; H. Left metatarsus, ventrolateral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, elytron epg, epipleural gutter along outer margin of elytron; mds, mid-dorsal spines of meso- and metatibia; mdt 1 - 2, mid-dorsal teeth 1 - 2 of protibia; mst 1 - 5, mesotarsomeres 1 - 5; msti, mesotibia; mtt 1 - 5, metatarsomeres 1 - 5; mtti, metatibia; pt 1 - 5, protarsomeres 1 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.3 mm (A, C, D); 0.2 mm (B, E - H)." httpUri="https://binary.pensoft.net/fig/868867" pageId="0" pageNumber="211">, 9</figureCitation>
<figureCitation id="2B02705B803F91CBB227410E62CA3615" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Prosternum with arrow showing subcontiguous procoxae, ventral view; B. Meso- and metathorax, ventral view; C. Abdomen, ventral view; D. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view. Abbreviations: msf, mesofemur; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ps, prosternum; v 1 - 5, ventrites 1 - 5. Scale bars: 0.3 mm (A); 0.2 mm (B); 0.5 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868868" pageId="0" pageNumber="211">, 10</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="381921933EE5FDDB37F47C96B808FD63" pageId="0" pageNumber="211" type="type species">
<paragraph id="625078E9868957118714429391F05B51" pageId="0" pageNumber="211">Type species.</paragraph>
<paragraph id="E7FD31D5C01AEF34258D25514A286D6E" pageId="0" pageNumber="211">
<taxonomicName id="1DFD5F31A5701860F67FAFC17225E82E" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="A32C6DA25DF22B8E1F52DC78AF454F76" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
sp. nov., here designated.
</paragraph>
</subSubSection>
<subSubSection id="B9514442FA63E9604716AFD5AAF3762C" pageId="0" pageNumber="211" type="differential diagnosis">
<paragraph id="39F8A32E2BF18CF54C4E4615D68BA670" pageId="0" pageNumber="211">Differential diagnosis.</paragraph>
<paragraph id="1F6E6E273282D8CF44E394627E6DED7B" pageId="0" pageNumber="211">
<taxonomicName id="F4C3584A26789E5D0FA314387B4309A4" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="4CB0B6DB6AA4CF8CD17B9E26ED9CE300" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. is easily separated from all known extant ceratognathin genera from the Australian Region, except the extant
<taxonomicName id="9602A9A03094F81D0C8B7B0D32112421" baseAuthorityName="Holloway" baseAuthorityYear="1962" class="Insecta" family="Lucanidae" genus="Holloceratognathus" kingdom="Animalia" lsidName="Holloceratognathus passaliformis" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="passaliformis">
<emphasis id="7EF5FEBC5A17DCE278392D5779099E4C" italics="true" pageId="0" pageNumber="211">Holloceratognathus passaliformis</emphasis>
</taxonomicName>
(Holloway), by the complete lack of modified scales on the dorsal surface (Figs
<figureCitation id="7A7738244047BFBB40017F138D09AD16" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. General habitus of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Dorsal view; B. Ventral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868864" pageId="0" pageNumber="211">6A</figureCitation>
,
<figureCitation id="5F4B1F45E5F7A20B97765CB54A84D602" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8</figureCitation>
). It can be distinguished from
<taxonomicName id="E5E1D85D1683AC01CFF060CEF7357E1A" lsidName="H. passaliformis" pageId="0" pageNumber="211" rank="species" species="passaliformis">
<emphasis id="D9A890C01F52B65B65AB5DE9A85BFDF2" italics="true" pageId="0" pageNumber="211">H. passaliformis</emphasis>
</taxonomicName>
by the absence of several derived features, such as widely explanate pronotal and elytral margins and distinctly paddle-shaped tibiae (
<bibRefCitation id="67362D13698C01BBF15926A288C37FE9" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">Holloway 2007</bibRefCitation>
).
<taxonomicName id="384428BBCFD93523502AB4B0DA846E4A" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="6026DB5A38776F808232845FAC162538" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. is somewhat similar to the recently described ceratognathin genus
<taxonomicName id="8C889EE39328A13F94AB174A7A9835FC" authorityName="Li &amp; Cai" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Oncelytris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oncelytris" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="87A53A30FC5D94B331D4B00E5F51F7DB" italics="true" pageId="0" pageNumber="211">Oncelytris</emphasis>
</taxonomicName>
Li &amp; Cai, 2023 (in Li Y-D et al. 2023) from Kachin amber in the absence of dosal scales (Li Y-D et al. 2023). However, the distinctly tuberculate elytra of
<taxonomicName id="87EB96A7BE6003A4860DBF03D903FC56" authorityName="Li &amp; Cai" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Oncelytris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oncelytris" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="881ED56DC9020FAF38B246E8B0B5BC7C" italics="true" pageId="0" pageNumber="211">Oncelytris</emphasis>
</taxonomicName>
are unusual in the family (Li Y-D et al. 2023), and
<taxonomicName id="6C64F22F520B1363AEA56A071781A4CD" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="A5BD808148C373FEE1A7C22C15480040" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. apparently lacks such distinct tubercles on the dorsum of elytra.
</paragraph>
<caption id="E34F2F217D53CB8EA6F25B0915E6D8C1" httpUri="https://binary.pensoft.net/fig/868864" pageId="0" pageNumber="211" start="Figure 6" startId="F6">
<paragraph id="64CF95DDC1FC7161E66051D12266B0EF" pageId="0" pageNumber="211">
<emphasis id="1B7415E882885E1E89C4F4B377D2E03A" bold="true" pageId="0" pageNumber="211">Figure 6.</emphasis>
General habitus of
<taxonomicName id="CAAC54E82E9CCF843F1BBE0B76944467" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="619C40090D405ECE804F34353078DCDC" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
gen. et sp. nov., holotype, SEHU-0000121206.
<emphasis id="DE4F1D155EB16CB6660C323A85CF61AF" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Dorsal view;
<emphasis id="CC75F5C7477805979B2F3A2D40F16D8D" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Ventral view. Scale bars: 1.0 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="64368637F194A576CD2F557BF7577073" pageId="0" pageNumber="211" type="diagnosis">
<paragraph id="F268363A794392F5B3F1321DFE1BDE0D" pageId="0" pageNumber="211">Diagnosis.</paragraph>
<paragraph id="B0CE6F50C07CABA920913029C1B1BD8F" pageId="0" pageNumber="211">
Body very small, moderately elongate oval, rather flattened; body length well below 3.0 mm. Dorsal surface lacks neither modified scales nor setae (Figs
<figureCitation id="8F87A0290F19C5FA6CDBC110ADD8E615" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. General habitus of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Dorsal view; B. Ventral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868864" pageId="0" pageNumber="211">6A</figureCitation>
,
<figureCitation id="D10A48FE9BE255B85E1C0EAA1E23357D" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8</figureCitation>
). Head with broadly rounded anterior margin (Fig.
<figureCitation id="56606199E0845A3842243D9417461C2F" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Head with right antenna, dorsal view; B. Head with right antenna, ventral view; C. Enlargement of A; D. Right antenna, ventral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (A, B); 0.2 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211">7A</figureCitation>
); vertex simple, without a pair of protuberances (Fig.
<figureCitation id="BD99F90718A6DE13558EE172AB81ED93" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Head with right antenna, dorsal view; B. Head with right antenna, ventral view; C. Enlargement of A; D. Right antenna, ventral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (A, B); 0.2 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211">7A</figureCitation>
). Antenna non-geniculate, with three-segmented, rather strong club (Fig.
<figureCitation id="0099264852B579EF929CA50B16D1E225" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Head with right antenna, dorsal view; B. Head with right antenna, ventral view; C. Enlargement of A; D. Right antenna, ventral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (A, B); 0.2 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211">7A, B, D</figureCitation>
). Mandibles small, inconspicuous, bearly visible dorsally, with sharply pointed apices (Fig.
<figureCitation id="16C3408407145CD4E038BB68D1C69D70" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Head with right antenna, dorsal view; B. Head with right antenna, ventral view; C. Enlargement of A; D. Right antenna, ventral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (A, B); 0.2 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211">7A-C</figureCitation>
). Pronotum densely strongly punctate (Fig.
<figureCitation id="382DDC8C40BE560038F16A5D14205755" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8A</figureCitation>
); lateral pronotal margins smooth, with very narrow gutter along whole margins (Fig.
<figureCitation id="C5453CBB00983D3FCA03E22E82977D72" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8A, D</figureCitation>
). Prosternum biconcaved; prosternal process strongly narrowed and laminate between procoxae, resulting in subcontiguous procoxae (Fig.
<figureCitation id="822BB76CAAF3E0333AB8A1260D6ABE8D" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Prosternum with arrow showing subcontiguous procoxae, ventral view; B. Meso- and metathorax, ventral view; C. Abdomen, ventral view; D. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view. Abbreviations: msf, mesofemur; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ps, prosternum; v 1 - 5, ventrites 1 - 5. Scale bars: 0.3 mm (A); 0.2 mm (B); 0.5 mm (C, D)." httpUri="https://binary.pensoft.net/fig/868868" pageId="0" pageNumber="211">10A</figureCitation>
). Elytra smooth without distinct tubercles and evident rows of punctures (Fig.
<figureCitation id="C92917B1FD094C1344BBD56B57A2F745" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8B, C, E</figureCitation>
); lateral margin with narrow epipleural gutter. Outer protibial margin with one large, curved, and slender apical tooth (
<emphasis id="93F322E93E6AE4D2862F8A5459236489" italics="true" pageId="0" pageNumber="211">at</emphasis>
), one large, curved, and slender mid-dorsal tooth 2 (
<emphasis id="BFB23D6D20A63BAEFE95F2074FE9D583" italics="true" pageId="0" pageNumber="211">mdt2</emphasis>
), one smaller mid-dorsal tooth 1 (
<emphasis id="BBFF9F7A05AA009AF7D5C73C7F1E53FD" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
), and about 16 subequal smaller teeth (Fig.
<figureCitation id="E941AAF333A9BB0635079652ACE8DF6D" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Right protibia and protarsus with arrow showing row of small teeth between large spines along outer edge of protibia, dorsal view; B. Right protibia with arrow showing row of small teeth between large spines along outer edge of protibia, ventral view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Left metatibia and metatarsus, dorsolateral view; E. Left mesotarsus, ventrolateral view; F. Left mesotarsus, dorsolateral view; G. Left metatarsus, dorsolateral view; H. Left metatarsus, ventrolateral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, elytron epg, epipleural gutter along outer margin of elytron; mds, mid-dorsal spines of meso- and metatibia; mdt 1 - 2, mid-dorsal teeth 1 - 2 of protibia; mst 1 - 5, mesotarsomeres 1 - 5; msti, mesotibia; mtt 1 - 5, metatarsomeres 1 - 5; mtti, metatibia; pt 1 - 5, protarsomeres 1 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.3 mm (A, C, D); 0.2 mm (B, E - H)." httpUri="https://binary.pensoft.net/fig/868867" pageId="0" pageNumber="211">9A, B</figureCitation>
).
</paragraph>
<caption id="B83E2BEC748E7C745CD8EC29B0AA0DB3" httpUri="https://binary.pensoft.net/fig/868865" pageId="0" pageNumber="211" start="Figure 7" startId="F7">
<paragraph id="472CB9C4D3712FF820B8B820CA056434" pageId="0" pageNumber="211">
<emphasis id="0B85937B38A8F4A7868BCB29723A6F45" bold="true" pageId="0" pageNumber="211">Figure 7.</emphasis>
Details of
<taxonomicName id="819BB27E16734E965682C0DB831CBCDA" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="320A9C0AB5EBC95A387CBE0D3DB7C54F" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
gen. et sp. nov., holotype, SEHU-0000121206.
<emphasis id="A62592CBF5D195B52B48D648D53EB5BE" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Head with right antenna, dorsal view;
<emphasis id="973F7AC9CB03FDCA8BAFAAC2EBBFFAD4" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Head with right antenna, ventral view;
<emphasis id="E2B5053342108346D0818FE0954880ED" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Enlargement of
<emphasis id="F78A276DB5F49CD25537B3F50E1D038B" bold="true" pageId="0" pageNumber="211">A</emphasis>
;
<emphasis id="DE90AAF207702990863B72D946DFEA75" bold="true" pageId="0" pageNumber="211">D.</emphasis>
Right antenna, ventral view. Abbreviations: a1-10, antennomeres 1-10; ey, compound eye; lp3, labial palpomere 3; md, mandible; mp3-4, maxillary palpomeres 3-4; pn, pronotum; pom, preocular margin of head; prt, protarsus. Scale bars: 0.3 mm (
<emphasis id="B34579B63AF26DC1EA9CD8D8A1A91507" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="588644217AD11A79CBBB8677F6720A8E" bold="true" pageId="0" pageNumber="211">B</emphasis>
); 0.2 mm (
<emphasis id="9D5CA602DA708FD18030E50741721AC4" bold="true" pageId="0" pageNumber="211">C</emphasis>
,
<emphasis id="9A528EE684EF5A12F798EF2450529E92" bold="true" pageId="0" pageNumber="211">D</emphasis>
).
</paragraph>
</caption>
<caption id="93A2EEC04DC4F8090DF11C3089E85A6C" httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211" start="Figure 8" startId="F8">
<paragraph id="EBF8F9BED889A3D484F94B96BD5796A3" pageId="0" pageNumber="211">
<emphasis id="B36F61E8F011386EB4E8ADABB40AE7BF" bold="true" pageId="0" pageNumber="211">Figure 8.</emphasis>
Details of
<taxonomicName id="37B5DFCD29CDF1A065D47E8FEF712DD8" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="0F2DBF889B891DE7CCE5288934C2A80E" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
gen. et sp. nov., holotype, SEHU-0000121206.
<emphasis id="7B5588F8C18ED81AADC2B5CB90F85637" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Pronotum and elytral base, dorsal view;
<emphasis id="C2DCAF497CC5012414B16320F7261F94" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Left elytron, dorsolateral view;
<emphasis id="0638C5F0BB8828AE09DA9A543C5A1D64" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Punctation on elytra, pronotum, and scutellar shield;
<emphasis id="CBF26E63CBF930BADF50FCEE64592099" bold="true" pageId="0" pageNumber="211">D.</emphasis>
Left lateral margin of pronotum, dorsal view;
<emphasis id="781D2463956817F327B71B7EE872152D" bold="true" pageId="0" pageNumber="211">E.</emphasis>
Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (
<emphasis id="26C15B1FE4B6F5D40998F817F191203A" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="03456E8D132E8D8863D06E3F11DFC8DA" bold="true" pageId="0" pageNumber="211">B</emphasis>
,
<emphasis id="55084255085D7ADB0C556452C45FCAAE" bold="true" pageId="0" pageNumber="211">E</emphasis>
); 0.3 mm (
<emphasis id="8DAA86CB659C4DCF8B057D6D6B8E5657" bold="true" pageId="0" pageNumber="211">D</emphasis>
).
</paragraph>
</caption>
<caption id="9827B885AEB165FD1EEFBB2033091C3A" httpUri="https://binary.pensoft.net/fig/868867" pageId="0" pageNumber="211" start="Figure 9" startId="F9">
<paragraph id="01A263CC006922BA922AFE5A021A1160" pageId="0" pageNumber="211">
<emphasis id="AB913663CEA37BB49BEA2E251907C265" bold="true" pageId="0" pageNumber="211">Figure 9.</emphasis>
Details of
<taxonomicName id="6F94C268096DD58DA7B14A98D3D72482" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="87A85CD1E1CBF4F8987C4D0776A93DE3" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
gen. et sp. nov., holotype, SEHU-0000121206.
<emphasis id="0AF8B37F74B2B281804B74C64A3F953A" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Right protibia and protarsus with arrow showing row of small teeth between large spines along outer edge of protibia, dorsal view;
<emphasis id="E47820ACAC56B6C8363033EE73F0E2C5" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Right protibia with arrow showing row of small teeth between large spines along outer edge of protibia, ventral view;
<emphasis id="3FDA1DE74A479ED8A8220445913C87BF" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Left mesotibia and mesotarsus, dorsolateral view;
<emphasis id="98D122BEE5435F5AE23AAABCD1EFA843" bold="true" pageId="0" pageNumber="211">D.</emphasis>
Left metatibia and metatarsus, dorsolateral view;
<emphasis id="EC64CE16C731E91FB1D2809C7C0F827F" bold="true" pageId="0" pageNumber="211">E.</emphasis>
Left mesotarsus, ventrolateral view;
<emphasis id="E6ECDE022A023ECC547C9F91050199A8" bold="true" pageId="0" pageNumber="211">F.</emphasis>
Left mesotarsus, dorsolateral view;
<emphasis id="102DB681051AECE06B2AB7C7224BD147" bold="true" pageId="0" pageNumber="211">G.</emphasis>
Left metatarsus, dorsolateral view;
<emphasis id="31C1045E0FE9BB6BBEDF6815FC7E2BDB" bold="true" pageId="0" pageNumber="211">H.</emphasis>
Left metatarsus, ventrolateral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, elytron epg, epipleural gutter along outer margin of elytron; mds, mid-dorsal spines of meso- and metatibia; mdt1-2, mid-dorsal teeth 1-2 of protibia; mst1-5, mesotarsomeres 1-5; msti, mesotibia; mtt1-5, metatarsomeres 1-5; mtti, metatibia; pt1-5, protarsomeres 1-5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.3 mm (
<emphasis id="C851A8CFEC4B4205FC74435CA13C8505" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="2C3F618A41D9BFF0ED47297BCB7AACC6" bold="true" pageId="0" pageNumber="211">C</emphasis>
,
<emphasis id="42BB9067F14C3B7BDAE73D8F1B51BE72" bold="true" pageId="0" pageNumber="211">D</emphasis>
); 0.2 mm (
<emphasis id="F47A1CD76D8D74EFBC0B8E5EE1116B6B" bold="true" pageId="0" pageNumber="211">B</emphasis>
,
<emphasis id="5B72A8AFB624AABB81677923B17236CD" bold="true" pageId="0" pageNumber="211">E</emphasis>
-
<emphasis id="955E3927651085A45933EA1213034C8A" bold="true" pageId="0" pageNumber="211">H</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="6CEF03FEC044C51C0879E157B738B3B1" pageId="0" pageNumber="211" type="included species">
<paragraph id="97E5481FC5546D1789720E64DBC118DD" pageId="0" pageNumber="211">Included species.</paragraph>
<paragraph id="1C76F84AB40A47FBF68BE6C2A30452FB" pageId="0" pageNumber="211">
Only
<taxonomicName id="2B6D467EC6F538B9E1721E247B04DBBD" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="minutissimus">
<emphasis id="8CCC8587B2ED888669E958D36CDC11EC" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</subSubSection>
<subSubSection id="459133303E05377B6626CBFEC740FF2A" pageId="0" pageNumber="211" type="etymology">
<paragraph id="8D41FA40E957CC1BDC24A92C1774C587" pageId="0" pageNumber="211">Etymology.</paragraph>
<paragraph id="B48AFAFDB770E5484CEB0D7D1E8BADBF" pageId="0" pageNumber="211">
The generic name is composed of the prefix
<taxonomicName id="5B2B80851A7FD5CE76CC153CDDC96584" genus="Creto" lsidName="Creto" pageId="0" pageNumber="211" rank="genus">
<emphasis id="0613F496CDFA2B0E380FCFD5349E66BA" italics="true" pageId="0" pageNumber="211">Creto</emphasis>
</taxonomicName>
- from
<normalizedToken id="CF3C2D10C2AC1CD470B84C9EBBCDF691" originalValue="“Cretaceus”">&quot;Cretaceus&quot;</normalizedToken>
referring to the &quot;Cretaceous age&quot; of the extinct genus and the type genus
<taxonomicName id="B3478B63E66EBDF01ED1F12D55CA7A9D" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Lucanidae" genus="Ceratognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="32FE2A1E6DE50EAABA4B344F6E2F305A" italics="true" pageId="0" pageNumber="211">Ceratognathus</emphasis>
</taxonomicName>
Westwood, 1838 of
<taxonomicName id="3D58BD5140512CB03CB58E4E90BAE631" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
. It is masculine in gender.
</paragraph>
</subSubSection>
<subSubSection id="91FEBAB3C6C8B82FE72D3FFD21DC2180" pageId="0" pageNumber="211" type="systematic placement and comparison">
<paragraph id="7BD7CC62F4A55B40891C50AD41CB12E1" pageId="0" pageNumber="211">Systematic placement and comparison.</paragraph>
<paragraph id="B8873B900D9E92D837E72E13366A03F0" pageId="0" pageNumber="211">
<taxonomicName id="9F4F9FCAAD320C3E2B939AD279728C6A" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="0AFF1E209E1202D9C1FAA88859144C39" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. can be unambiguously assigned to the scarabaeoid family
<taxonomicName id="0E472484DE0A62654F19683FD8BC1022" authorityName="Latreille" authorityYear="1804" class="Insecta" family="Lucanidae" kingdom="Animalia" lsidName="" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="family">Lucanidae</taxonomicName>
based on the moderately elongate and rather dorsoventrally flattened body shape, 10-segmented antennae with a relatively long antennomere 1 and three-segmented, non-coherent and lamellate apical club, 5-5-5 tarsal formula, protibial structures with characteristic outer teeth, strongly developed bisetose arolium between the pretarsal claws, and five visible abdominal ventrites (e.g.,
<bibRefCitation id="817D7B37660BADDB5105E66E919BDD66" author="Ratcliffe, BC" editor="Arnett, R" journalOrPublisher="II). Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton" pageId="0" pageNumber="211" pagination="6 - 9" refId="B67" refString="Ratcliffe, BC, 2002. Lucanidae Latreille 1804. In: Arnett, R, Thomas, MC, Skelley, PE, Frank, JH, Eds., American Beetles (Vol. II). Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton: 6 - 9" title="Lucanidae Latreille 1804." volumeTitle="American Beetles (Vol." year="2002">Ratcliffe 2002</bibRefCitation>
;
<bibRefCitation id="91059A3E96E86DC292BEA112167B449C" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">Holloway 2007</bibRefCitation>
;
<bibRefCitation id="F4A4977F885B459F10390FC346967F90" DOI="https://doi.org/10.1016/j.cretres.2017.06.008" author="Cai, C" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="211" pagination="109 - 112" refId="B9" refString="Cai, C, Yin, Z, Liu, Y, Huang, D, 2017. Protonicagus tani gen. et sp. nov., the first stag beetles from Upper Cretaceous Burmese amber (Coleoptera: Lucanidae: Aesalinae: Nicagini). Cretaceous Research 78: 109 - 112, DOI: https://doi.org/10.1016/j.cretres.2017.06.008" title="Protonicagus tani gen. et sp. nov., the first stag beetles from Upper Cretaceous Burmese amber (Coleoptera: Lucanidae: Aesalinae: Nicagini)." url="https://doi.org/10.1016/j.cretres.2017.06.008" volume="78" year="2017">Cai et al. 2017</bibRefCitation>
;
<bibRefCitation id="7835A40980A5CF86A82C5223CC892810" author="Reid, C" editor="Ślipinski, A" journalOrPublisher="2). Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, Victoria" pageId="0" pageNumber="211" pagination="391 - 416" refId="B68" refString="Reid, C, 2019. Lucanidae Latreille, 1804. In: Ślipinski, A, Lawrence, JF, Eds., Australian Beetles (Vol. 2). Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, Victoria: 391 - 416" title="Lucanidae Latreille, 1804." volumeTitle="Australian Beetles (Vol." year="2019">Reid 2019</bibRefCitation>
). Furthermore, the new genus can be safely attributed to the plesiomorphic extant subfamily
<taxonomicName id="E8A9C89FE8E137EB00EB982A1EFF10FB" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Aesalinae">Aesalinae</taxonomicName>
based on its non-geniculate antennae with a three-segmented club, complete eyes (not divided by ocular canthus), small mandibles, non-striate elytra, and strongly narrowed prosternal process with subcontiguous procoxae (e.g.,
<bibRefCitation id="2C3F73B1946DB4C100DD743E3169481D" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">Holloway 2007</bibRefCitation>
;
<bibRefCitation id="F0664AC27343DD78D73C93003A9A1D8C" author="Reid, C" editor="Ślipinski, A" journalOrPublisher="2). Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, Victoria" pageId="0" pageNumber="211" pagination="391 - 416" refId="B68" refString="Reid, C, 2019. Lucanidae Latreille, 1804. In: Ślipinski, A, Lawrence, JF, Eds., Australian Beetles (Vol. 2). Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, Victoria: 391 - 416" title="Lucanidae Latreille, 1804." volumeTitle="Australian Beetles (Vol." year="2019">Reid 2019</bibRefCitation>
). Among the three tribes of
<taxonomicName id="3080038D9E7E8756F3E3BD67BA002CA5" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Aesalinae">Aesalinae</taxonomicName>
,
<taxonomicName id="C47EFAC8E1899EF8D3D606E0ECF7503F" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="F281E36C05B4FB51C1A117EBDDCDA1BC" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. is separated from
<taxonomicName id="93C68973276444B7C8D2F08720FD0975" authorityName="Leconte" authorityYear="1861" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Nicagini">Nicagini</taxonomicName>
by a combination of the following character states (Paulsen 2005): complete absence of evident ground scales (setae) on the dorsum, much longer setae of the arolium, and particularly the features of the outer protibial edge (i.e., apical tooth (
<emphasis id="5BAA2C3E8D56B6625C9684EC64158622" italics="true" pageId="0" pageNumber="211">at</emphasis>
) much narrower and pointed, mid-dorsal tooth (
<emphasis id="BBE4873BBE4487EFC9872773D4BC58D0" italics="true" pageId="0" pageNumber="211">mdt2</emphasis>
) much longer and slenderer, located anteriorly, mid-dorsal tooth (
<emphasis id="ABB1FC2C5A86A84C70A1EBA9F4294765" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
) narrower with acute apex, and the presence of subequal smaller teeth between apical tooth and mid-dorsal tooth 2) (see Fig.
<figureCitation id="C4AD5D53ADB1F07F87D39138E04D5B43" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Details of extant species of Aesalinae. A, D, G, K. Nicagus japonicus (tribe Nicagini), male, showing head with left antenna in dorsal view (A), prosternal process (intercoxal process of procoxae) with subcontiguous procoxae in ventral view (D), right protibia in dorsal view (G), or right metatarsus in ventral view (K) based on two specimens; B, E, H, J, L. Ceratognathus cf. niger (tribe Ceratognathini), female, showing head with left antenna in dorsal view (B), prosternal process (intercoxal process of procoxae) with subcontiguous procoxae in ventral view (E), right protibia in dorsal view (H), pronotum and left elytron with scales (J), or left metatarsus in ventral view (L) based on same specimen; C, F, I. Aesalus asiaticus asiaticus (tribe Aesalini), female, showing left antenna in dorsal view (C), prosternal process (intercoxal process of procoxae) with separated procoxae in ventral view (F), or right protibia in dorsal view (I) based on two specimens. Arrows show either smooth (G) or serrate outer edge (H) between mid-dorsal tooth 1 and mid-dorsal tooth 2 of each protibia. Abbreviations: a 1, a 2, a 8 - 10, antennomeres 1, 2, 8 - 10; as, apical spur; at, apical tooth of protibia; cl, claw; ey, compound eye; md, mandible; mdt 1 - 2, mid-dorsal teeth 1 - 2 of protibia; mtt 5, metatarsomere 5; psp, prosternal process between procoxae; ra, rod of arolium; scl, scales on head, pronotum, and elytra; sa, setae on rod of arolium." httpUri="https://binary.pensoft.net/fig/868873" pageId="0" pageNumber="211">15</figureCitation>
for the comparison). Furthermore, the new genus is easily distinguished from
<taxonomicName id="097B295D6EA942364BB58E3D3A6C1ABD" authorityName="MacLeay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Aesalini">Aesalini</taxonomicName>
in having a strongly narrowed prosternal process and subcontiguous procoxae (see
<bibRefCitation id="1AD66BEB32D1F002A96E5A423B631895" DOI="https://doi.org/10.11646/zootaxa.2069.1.2" author="Huang, H" journalOrPublisher="Zootaxa" pageId="0" pageNumber="211" pagination="18 - 42" refId="B28" refString="Huang, H, Bi, W-X, Li, L-Z, 2009. Discovery of a second species of Aesalini from continental China, with description of the new species and its third instar larva (Coleoptera: Scarabaeoidea: Lucanidae). Zootaxa 2069 (1): 18 - 42, DOI: https://doi.org/10.11646/zootaxa.2069.1.2" title="Discovery of a second species of Aesalini from continental China, with description of the new species and its third instar larva (Coleoptera: Scarabaeoidea: Lucanidae)." url="https://doi.org/10.11646/zootaxa.2069.1.2" volume="2069" year="2009">Huang et al. 2009</bibRefCitation>
). Overall, the general morphology of
<taxonomicName id="B7519FFAA76AF2AA25A32E07524F945E" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="CD130F8231D46CDB81562387731D843B" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. matches the remaining tribe
<taxonomicName id="CA59C1F920566E997C86F4B2D8F68B3E" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
, particularly based on the structures of the protibial edges. In fact, the new genus has the typical pattern of protibial spines. Consequently, it is best placed within
<taxonomicName id="2ED3C016802AF44B589036C19CD75E55" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
.
</paragraph>
<paragraph id="ED3ED5EE0393C04FA724241242BEE595" pageId="0" pageNumber="211">
<taxonomicName id="B81A05BD3C5F32C1628535366364AEA4" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
is a small tribe with only four extant genera (Table
<tableCitation id="455EC4A296D1F762DACDB7E35331BE91" captionStart="Table 1" captionStartId="T1" captionText="Table 1. General overview of Aesalinae based on Schoolmeesters (2023), Li Y-D et al. (2023), and this study." httpUri="http://table.plazi.org/id/58609FC0C8857E9C424565BE269CBB23" pageId="0" pageNumber="211" tableUuid="58609FC0C8857E9C424565BE269CBB23">1</tableCitation>
):
<taxonomicName id="5C2CAE816678BF5A36EBA98B769F60F6" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Lucanidae" genus="Ceratognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="2C568FB9A7AD25015382FADE1E8F4789" italics="true" pageId="0" pageNumber="211">Ceratognathus</emphasis>
</taxonomicName>
Westwood, 1838;
<taxonomicName id="B334D6AE85B9EF3DC2ECCAA1B793B17C" authorityName="Paulsen &amp; Mondaca" authorityYear="2006" class="Insecta" family="Lucanidae" genus="Hilophyllus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hilophyllus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="F5E29AC7B72AAA6E60FA0891B276631D" italics="true" pageId="0" pageNumber="211">Hilophyllus</emphasis>
</taxonomicName>
Paulsen &amp; Mondaca, 2006;
<taxonomicName id="1A0B7718E71BBA488F5D3415523F1961" authorityName="Nikolajev" authorityYear="1998" class="Insecta" family="Lucanidae" genus="Holloceratognathus" kingdom="Animalia" lsidName="Holloceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="0B2E87189BB3713D9AC4DCA701218D33" italics="true" pageId="0" pageNumber="211">Holloceratognathus</emphasis>
</taxonomicName>
Nikolajev, 1998; and
<taxonomicName id="74C05D3D55A5D6A52BF307DB51383BA9" authorityName="Parry" authorityYear="1843" class="Insecta" family="Lucanidae" genus="Mitophyllus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mitophyllus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="D54F690510DE17A95B7C0D37DE2EC5E1" italics="true" pageId="0" pageNumber="211">Mitophyllus</emphasis>
</taxonomicName>
Parry, 1843 (
<bibRefCitation id="4902DB7FCC2089B8F6D9A36D37D6EC77" DOI="https://doi.org/10.1080/03014223.1998.9517578" author="Holloway, BA" journalOrPublisher="Journal of the Royal Society of New Zealand" pageId="0" pageNumber="211" pagination="641 - 656" refId="B21" refString="Holloway, BA, 1998. A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae). Journal of the Royal Society of New Zealand 28 (4): 641 - 656, DOI: https://doi.org/10.1080/03014223.1998.9517578" title="A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae)." url="https://doi.org/10.1080/03014223.1998.9517578" volume="28" year="1998">Holloway 1998</bibRefCitation>
,
<bibRefCitation id="AD483866C541B512D1CBD54099BBEBBE" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">2007</bibRefCitation>
;
<bibRefCitation id="EFD0282462BE05F0069530106DB05207" DOI="https://doi.org/10.11646/zootaxa.1191.1.1" author="Paulsen, MJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="211" pagination="1 - 19" refId="B61" refString="Paulsen, MJ, Mondaca, J, 2006. Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species. Zootaxa 1191 (1): 1 - 19, DOI: https://doi.org/10.11646/zootaxa.1191.1.1" title="Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species." url="https://doi.org/10.11646/zootaxa.1191.1.1" volume="1191" year="2006">Paulsen and Mondaca 2006</bibRefCitation>
;
<bibRefCitation id="84C668DF6E3F22E1BD0FE97222CD5215" author="Paulsen, MJ" journalOrPublisher="Insecta Mundi" pageId="0" pageNumber="211" pagination="1 - 25" refId="B59" refString="Paulsen, MJ, 2013. A new genus for the Neotropical species of Aesalus Fabricius, with descriptions of eight new species (Coleoptera: Lucanidae: Aesalinae). Insecta Mundi 0325: 1 - 25" title="A new genus for the Neotropical species of Aesalus Fabricius, with descriptions of eight new species (Coleoptera: Lucanidae: Aesalinae)." volume="0325" year="2013">Paulsen 2013</bibRefCitation>
). In addition, the extinct genus
<taxonomicName id="B6E40A7C599C233D80AC05D621025222" authorityName="Li &amp; Cai" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Oncelytris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oncelytris" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="FF5F024367956A8AD9ED63C5EE8FE308" italics="true" pageId="0" pageNumber="211">Oncelytris</emphasis>
</taxonomicName>
from Kachin amber was very recently described (Li Y-D et al. 2023). Although these extant genera share externally similar habitus and body parts, they are mainly identified by the different structures and arrangements of the smaller protibial outer teeth-like spines (
<bibRefCitation id="7C8E685E016A1282646299883089F551" DOI="https://doi.org/10.1080/03014223.1998.9517578" author="Holloway, BA" journalOrPublisher="Journal of the Royal Society of New Zealand" pageId="0" pageNumber="211" pagination="641 - 656" refId="B21" refString="Holloway, BA, 1998. A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae). Journal of the Royal Society of New Zealand 28 (4): 641 - 656, DOI: https://doi.org/10.1080/03014223.1998.9517578" title="A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae)." url="https://doi.org/10.1080/03014223.1998.9517578" volume="28" year="1998">Holloway 1998</bibRefCitation>
,
<bibRefCitation id="8D44011328B44341A8550206DB31FD1A" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">2007</bibRefCitation>
;
<bibRefCitation id="9B0473562465D93BCADDDA234329DE44" DOI="https://doi.org/10.11646/zootaxa.1191.1.1" author="Paulsen, MJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="211" pagination="1 - 19" refId="B61" refString="Paulsen, MJ, Mondaca, J, 2006. Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species. Zootaxa 1191 (1): 1 - 19, DOI: https://doi.org/10.11646/zootaxa.1191.1.1" title="Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species." url="https://doi.org/10.11646/zootaxa.1191.1.1" volume="1191" year="2006">Paulsen and Mondaca 2006</bibRefCitation>
). Unlike
<taxonomicName id="64CA56AFD71FC56AE095BABE1FFF7BD3" authorityName="Nikolajev" authorityYear="1998" class="Insecta" family="Lucanidae" genus="Holloceratognathus" kingdom="Animalia" lsidName="Holloceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="1A30B94513FBA950FA1180F48F6D5F79" italics="true" pageId="0" pageNumber="211">Holloceratognathus</emphasis>
</taxonomicName>
which has numerous variably sized small spines, the new genus has numerous small, equal spines elsewhere along the outer edges of the protibiae (cf.
<bibRefCitation id="27AA0FA473C787A403F2DCE0881B4928" DOI="https://doi.org/10.1080/03014223.1998.9517578" author="Holloway, BA" journalOrPublisher="Journal of the Royal Society of New Zealand" pageId="0" pageNumber="211" pagination="641 - 656" refId="B21" refString="Holloway, BA, 1998. A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae). Journal of the Royal Society of New Zealand 28 (4): 641 - 656, DOI: https://doi.org/10.1080/03014223.1998.9517578" title="A re-evaluation of the genera of New Zealand aesaline stag beetles (Coleoptera: Lucanidae)." url="https://doi.org/10.1080/03014223.1998.9517578" volume="28" year="1998">Holloway 1998</bibRefCitation>
,
<bibRefCitation id="694B7E64E99421EB92526210F8C1A2C0" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">2007</bibRefCitation>
;
<bibRefCitation id="50AF1C9A59A1F105BFB7E05F010487EF" DOI="https://doi.org/10.11646/zootaxa.1191.1.1" author="Paulsen, MJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="211" pagination="1 - 19" refId="B61" refString="Paulsen, MJ, Mondaca, J, 2006. Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species. Zootaxa 1191 (1): 1 - 19, DOI: https://doi.org/10.11646/zootaxa.1191.1.1" title="Revision of the South American Ceratognathini (Coleoptera: Lucanidae: Aesalinae) with the description of a new genus and a new species." url="https://doi.org/10.11646/zootaxa.1191.1.1" volume="1191" year="2006">Paulsen and Mondaca 2006</bibRefCitation>
). It is somewhat difficult to compare the new taxon with the three other extant ceratognathinin genera in further detail based on the structures and arrangements of the protibial teeth, but
<taxonomicName id="CF2B98FCAD1A4EEF70ACB74286222195" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="6F2FA4BC84AE89285A639D51CF3F479A" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. has relatively large teeth-like spines between the base and mid-dorsal tooth 2 (i.e., mid-dorsal tooth 1,
<emphasis id="7FBC5AD7898281AAEF8DE287A2049C9F" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
, in this paper) along with the similar condition in
<taxonomicName id="A85DE727556B2ACBD844BCE6B0935312" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Lucanidae" genus="Ceratognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="447DED1B325B4EA545F6F2A4F62D6433" italics="true" pageId="0" pageNumber="211">Ceratognathus</emphasis>
</taxonomicName>
. Nevertheless, the new genus cannot be placed in
<taxonomicName id="3CC735A9F8D8D5D19FDD34C633CD7516" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Lucanidae" genus="Ceratognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="CE6E51D42AA99078475E388A4B2156F5" italics="true" pageId="0" pageNumber="211">Ceratognathus</emphasis>
</taxonomicName>
due to the lack of a pair of protuberances or tubercles (raised shiny patches) on the dorsal side of the head (
<bibRefCitation id="10D24A9EC605C119257B9339F1B705F2" author="Holloway, BA" journalOrPublisher="Fauna of New Zealand" pageId="0" pageNumber="211" pagination="1 - 254" refId="B22" refString="Holloway, BA, 2007. Lucanidae (Insecta: Coleoptera). Fauna of New Zealand 61: 1 - 254" title="Lucanidae (Insecta: Coleoptera)." volume="61" year="2007">Holloway 2007</bibRefCitation>
). More importantly, no ground setation or modified scales are found on the dorsal surface of
<taxonomicName id="E7F605EB4E25AA2B332AEC97A291265A" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="67F076809A554E2F62C9D065B255EC9A" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. (Fig.
<figureCitation id="4FBEB4A4777022168081ED78103E2B6F" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Details of Cretognathus minutissimus gen. et sp. nov., holotype, SEHU- 0000121206. A. Pronotum and elytral base, dorsal view; B. Left elytron, dorsolateral view; C. Punctation on elytra, pronotum, and scutellar shield; D. Left lateral margin of pronotum, dorsal view; E. Left elytral margin with elytral shoulder, dorsal view. Abbreviations: epg, epipleural gutter along outer margin of elytron; pg, gutter along outer margin of pronotum; pn, pronotum; sc, scutellar shield. Scale bars: 0.5 mm (A, B, E); 0.3 mm (D)." httpUri="https://binary.pensoft.net/fig/868866" pageId="0" pageNumber="211">8</figureCitation>
). This condition has never been observed in any recent ceratognathinin (Fig.
<figureCitation id="38CC3416CE1C63074D96E31FA8956F49" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Details of extant species of Aesalinae. A, D, G, K. Nicagus japonicus (tribe Nicagini), male, showing head with left antenna in dorsal view (A), prosternal process (intercoxal process of procoxae) with subcontiguous procoxae in ventral view (D), right protibia in dorsal view (G), or right metatarsus in ventral view (K) based on two specimens; B, E, H, J, L. Ceratognathus cf. niger (tribe Ceratognathini), female, showing head with left antenna in dorsal view (B), prosternal process (intercoxal process of procoxae) with subcontiguous procoxae in ventral view (E), right protibia in dorsal view (H), pronotum and left elytron with scales (J), or left metatarsus in ventral view (L) based on same specimen; C, F, I. Aesalus asiaticus asiaticus (tribe Aesalini), female, showing left antenna in dorsal view (C), prosternal process (intercoxal process of procoxae) with separated procoxae in ventral view (F), or right protibia in dorsal view (I) based on two specimens. Arrows show either smooth (G) or serrate outer edge (H) between mid-dorsal tooth 1 and mid-dorsal tooth 2 of each protibia. Abbreviations: a 1, a 2, a 8 - 10, antennomeres 1, 2, 8 - 10; as, apical spur; at, apical tooth of protibia; cl, claw; ey, compound eye; md, mandible; mdt 1 - 2, mid-dorsal teeth 1 - 2 of protibia; mtt 5, metatarsomere 5; psp, prosternal process between procoxae; ra, rod of arolium; scl, scales on head, pronotum, and elytra; sa, setae on rod of arolium." httpUri="https://binary.pensoft.net/fig/868873" pageId="0" pageNumber="211">15</figureCitation>
), including the
<taxonomicName id="065DA87B2502816A259A3793DE2E3ED5" authorityName="Westwood" authorityYear="1838" class="Insecta" family="Lucanidae" genus="Ceratognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ceratognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="3127462CA5C643ED4DB675AD3719FACF" italics="true" pageId="0" pageNumber="211">Ceratognathus</emphasis>
</taxonomicName>
species, with the only exception
<taxonomicName id="16A0FD8857005E22C53352F68892A9B2" lsidName="H. passaliformis" pageId="0" pageNumber="211" rank="species" species="passaliformis">
<emphasis id="A2A35AE7FBEE43918089463CFFF35543" italics="true" pageId="0" pageNumber="211">H. passaliformis</emphasis>
</taxonomicName>
(see also
<bibRefCitation id="F0D6BC226B19FA88AF30BA212AC0C6D1" author="Bartolozzi, L" journalOrPublisher="Taita Publishers, Hradec Kralove" pageId="0" pageNumber="211" refId="B2" refString="Bartolozzi, L, Zilioli, M, de Keyzer, R, 2017. The Stag Beetles of Australia, New Zealand, New Caledonia and Fiji (Coleoptera: Lucanidae). Taita Publishers, Hradec Kralove" title="The Stag Beetles of Australia, New Zealand, New Caledonia and Fiji (Coleoptera: Lucanidae)." year="2017">Bartolozzi et al. 2017</bibRefCitation>
). Interestingly, however, the other Kachin amber ceratognathinin
<taxonomicName id="D5DFC5114DB655BAF4B64F26486F9B04" authorityName="Li &amp; Cai" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Oncelytris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oncelytris" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="42BFDCF4CC1E61C2ED3B20667217E16C" italics="true" pageId="0" pageNumber="211">Oncelytris</emphasis>
</taxonomicName>
also lacks such modified scales on the elytra (Li Y-D et al. 2023).
</paragraph>
<paragraph id="0E4172A1EE47E7B3E6BDB57E13AC30B6" pageId="0" pageNumber="211">
From all of the lucanid fossils so far described from Kachin amber,
<taxonomicName id="CA5AED23F547495662E5637B1C0CD9AE" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="05EE17487AB165CEFE62A1A00D14EF4B" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. can be readily separated from
<taxonomicName id="3192064E24D344666E4D6DA184A9CC3B" class="Insecta" family="Lucanidae" genus="Anisoodontus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Anisoodontus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="FBE558B18684F3B87DCE2DBBAEEC9BB0" italics="true" pageId="0" pageNumber="211">Anisoodontus</emphasis>
</taxonomicName>
Wu, Tang &amp; Peng, 2022 (in
<bibRefCitation id="F279AF32A6E6202ABF9B425FFE46540A" DOI="https://doi.org/10.1016/j.cretres.2022.105208" author="Wu, L" journalOrPublisher="Insecta Mundi" pageId="0" pageNumber="211" refId="B87" refString="Wu, L, Tang, H, Peng, L, Zhang, H, Tong, H, 2022. The first representatives of Lucaninae from mid-Cretaceous Burmese amber based on two new species of Anisoodontus gen. nov. (Coleoptera: Scarabaeoidea: Lucanidae). Cretaceous Research 135: e105208. https://doi.org/10.1016/j.cretres.2022.105208" title="The first representatives of Lucaninae from mid-Cretaceous Burmese amber based on two new species of Anisoodontus gen. nov. (Coleoptera: Scarabaeoidea: Lucanidae). Cretaceous Research 135: e 105208." url="https://doi.org/10.1016/j.cretres.2022.105208" year="2022">Wu et al. 2022</bibRefCitation>
) (
<taxonomicName id="88A12ADBAF3FB8FE5548DC63C45748EC" authorityName="Latreille" authorityYear="1804" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Lucaninae">Lucaninae</taxonomicName>
) in having a much smaller body (11.80-17.36 mm, including mandibles, in
<taxonomicName id="8ADE26B6DC512F11014CB64C73539850" class="Insecta" family="Lucanidae" genus="Anisoodontus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Anisoodontus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="8485BA69953AE3557DCA971293DA7393" italics="true" pageId="0" pageNumber="211">Anisoodontus</emphasis>
</taxonomicName>
), non-geniculate antennae, and short, inconspicuous mandibles (cf.
<bibRefCitation id="D577A5697965DAFF19AAB3A3F1E6406C" DOI="https://doi.org/10.1016/j.cretres.2022.105208" author="Wu, L" journalOrPublisher="Insecta Mundi" pageId="0" pageNumber="211" refId="B87" refString="Wu, L, Tang, H, Peng, L, Zhang, H, Tong, H, 2022. The first representatives of Lucaninae from mid-Cretaceous Burmese amber based on two new species of Anisoodontus gen. nov. (Coleoptera: Scarabaeoidea: Lucanidae). Cretaceous Research 135: e105208. https://doi.org/10.1016/j.cretres.2022.105208" title="The first representatives of Lucaninae from mid-Cretaceous Burmese amber based on two new species of Anisoodontus gen. nov. (Coleoptera: Scarabaeoidea: Lucanidae). Cretaceous Research 135: e 105208." url="https://doi.org/10.1016/j.cretres.2022.105208" year="2022">Wu et al. 2022</bibRefCitation>
); from
<taxonomicName id="0E28920D97DAC700B110C1D790FF5B2B" authorityName="Cai, Yin, Liu &amp; Huang" authorityYear="2017" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="9177D1F448C188664B25DC54625214A0" italics="true" pageId="0" pageNumber="211">Protonicagus</emphasis>
</taxonomicName>
(
<taxonomicName id="75F9CF0D7008FAD2B8481B55E2BE1C27" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Aesalinae">Aesalinae</taxonomicName>
:
<taxonomicName id="E0D70D9E22521168EA37E93F78F3C399" authorityName="Leconte" authorityYear="1861" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Nicagini">Nicagini</taxonomicName>
) by non-crenulate pronotal lateral margins without a row of setae, larger head with non-enlarged preocular margin of the eyes, absence of any dorsal scales or ground setae, and presence of small subcontiguous teeth-like spines between the apical tooth and mid-dorsal tooth 2 (cf.
<bibRefCitation id="AC5985917C248207C3254B843C552C58" DOI="https://doi.org/10.1016/j.cretres.2017.06.008" author="Cai, C" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="211" pagination="109 - 112" refId="B9" refString="Cai, C, Yin, Z, Liu, Y, Huang, D, 2017. Protonicagus tani gen. et sp. nov., the first stag beetles from Upper Cretaceous Burmese amber (Coleoptera: Lucanidae: Aesalinae: Nicagini). Cretaceous Research 78: 109 - 112, DOI: https://doi.org/10.1016/j.cretres.2017.06.008" title="Protonicagus tani gen. et sp. nov., the first stag beetles from Upper Cretaceous Burmese amber (Coleoptera: Lucanidae: Aesalinae: Nicagini)." url="https://doi.org/10.1016/j.cretres.2017.06.008" volume="78" year="2017">Cai et al. 2017</bibRefCitation>
; this study); from
<taxonomicName id="2A07744B461DA5B01FD0E7793B1F075C" authorityName="Bai, Zhang &amp; Qiu" authorityYear="2017" class="Insecta" family="Lucanidae" genus="Electraesalopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Electraesalopsis" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="2BEDEE691C5E7E6239773AF4F5BFEE5E" italics="true" pageId="0" pageNumber="211">Electraesalopsis</emphasis>
</taxonomicName>
Bai, Zhang &amp; Qiu, 2017 (in
<bibRefCitation id="661F97FAAC5D35EAFD3781DF900851E7" DOI="https://doi.org/10.11865/zs.201717" author="Qiu, T" journalOrPublisher="Zoological Systematics" pageId="0" pageNumber="211" pagination="390 - 394" refId="B66" refString="Qiu, T, Lu, Y, Zhang, W, Wang, S, Yang, Y, Bai, M, 2017. Electraesalopsis beuteli gen. &amp; sp. nov., the first lucanid beetle from the Cretaceous Burmese amber (Coleoptera: Scarabaeoidea). Zoological Systematics 42 (3): 390 - 394, DOI: https://doi.org/10.11865/zs.201717" title="Electraesalopsis beuteli gen. &amp; sp. nov., the first lucanid beetle from the Cretaceous Burmese amber (Coleoptera: Scarabaeoidea)." url="https://doi.org/10.11865/zs.201717" volume="42" year="2017">Qiu et al. 2017</bibRefCitation>
) (subfamily
<emphasis id="5517D50F82CC6D798A28E04DF1FBF1E0" italics="true" pageId="0" pageNumber="211">incertae sedis</emphasis>
) by the presence of much shorter, unmodified mandibles, much strongly developed antennal club, broadly rounded clypeal margin, distinctly shorter metatarsi, smaller body (5.6 mm, including mandibles, in
<taxonomicName id="E862C866543738BF377A28AFC8558382" authorityName="Bai, Zhang &amp; Qiu" authorityYear="2017" class="Insecta" family="Lucanidae" genus="Electraesalopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Electraesalopsis" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="73E61BFB91F241D11239695C42A5D333" italics="true" pageId="0" pageNumber="211">Electraesalopsis</emphasis>
</taxonomicName>
), and structures of the protibiae with their strongly developed apical teeth (cf.
<bibRefCitation id="C89A6832F9B3BF817A8FDAD704BEAE70" DOI="https://doi.org/10.11865/zs.201717" author="Qiu, T" journalOrPublisher="Zoological Systematics" pageId="0" pageNumber="211" pagination="390 - 394" refId="B66" refString="Qiu, T, Lu, Y, Zhang, W, Wang, S, Yang, Y, Bai, M, 2017. Electraesalopsis beuteli gen. &amp; sp. nov., the first lucanid beetle from the Cretaceous Burmese amber (Coleoptera: Scarabaeoidea). Zoological Systematics 42 (3): 390 - 394, DOI: https://doi.org/10.11865/zs.201717" title="Electraesalopsis beuteli gen. &amp; sp. nov., the first lucanid beetle from the Cretaceous Burmese amber (Coleoptera: Scarabaeoidea)." url="https://doi.org/10.11865/zs.201717" volume="42" year="2017">Qiu et al. 2017</bibRefCitation>
); from
<taxonomicName id="954E83688A702855E60A735F0446600E" authorityName="Li &amp; Cai" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Oncelytris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Oncelytris" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="6D249F23249DD7EBF5B6A5C8DBB65813" italics="true" pageId="0" pageNumber="211">Oncelytris</emphasis>
</taxonomicName>
by the lack of a pair of not well-separated protuberances and tuberculate elytra. Consequently, the new genus can easily be distinguished from any fossil genera described from Kachin amber. Furthermore,
<taxonomicName id="4B64335281C370E549FC52D052757D66" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Cretognathus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cretognathus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="E1041DBAEF42653BBF51A2B928196E8E" italics="true" pageId="0" pageNumber="211">Cretognathus</emphasis>
</taxonomicName>
gen. nov. is separated from the two undetermined provisional ceratognathins reported below, mainly by the shape of the protibiae and antennae (the latter can be applied only for
<taxonomicName id="3EABE65CB2EDC95704C002DED9AA1DF3" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
gen. et sp. indet. 1).
</paragraph>
</subSubSection>
</treatment>
</document>
Reference in a new issue View git blame Copy permalink