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<document ID-DOI="http://dx.doi.org/10.3897/evolsyst.7.104597" 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="2AAF24EF723E5FE590D7ECE00179912A" 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="Protonicagus mandibularis Yamamoto 2023, sp. nov." docType="treatment" docUuid="9689A395-8B60-46ED-BFC0-9E87158D2CA6" docUuidSource="ZooBank" docVersion="1" 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="1687815010481" updateUser="pensoft">
<mods:mods id="C44F9AED488527425F5A7C6EC21AB4A9" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="01F9B0ACF90FB68AC0EC51F263A1DCA1">
<mods:title id="632CDE5D0DE492CF3B2F56F75A2D9763">The smallest stag beetles (Coleoptera, Lucanidae): hidden paleodiversity in mid-Cretaceous Kachin amber from northern Myanmar</mods:title>
</mods:titleInfo>
<mods:name id="7C9E3723FB23DA9E5D04AB89DBECD77E" type="personal">
<mods:role id="986AF9DEF4CDEC06B13A2E2BACC1D140">
<mods:roleTerm id="F5F4F9957CAEDEDD4301C99E8D62843E">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="E3D2C04A20BC66C0708C9533C67F0BEE">Yamamoto, Shuhei</mods:namePart>
<mods:nameIdentifier id="1D29F047D0825BC8785F602ABE8FABE3" type="ORCID">https://orcid.org/0000-0002-4162-8457</mods:nameIdentifier>
<mods:affiliation id="EBF73AF0A2834E2C89F407FCDF70C593">The Hokkaido University Museum, Hokkaido University, Kita 10 - jo, Nishi 8 - chome, Kita-ku, Sapporo 060 - 0810, Japan</mods:affiliation>
<mods:nameIdentifier id="72283F965CBC164AC6A36883EEFF06B5" type="email">s.yamamoto.64@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource id="238CC7F2959347D88C11CC5C642C289D">text</mods:typeOfResource>
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<mods:title id="DBA95EBB4E8D4513F02441524C593073">Evolutionary Systematics</mods:title>
</mods:titleInfo>
<mods:part id="9547D1BAC08EA5D7E0CD6A206743B56F">
<mods:date id="5A8D791138FFF5317E44EE8027CE5756">2023</mods:date>
<mods:detail id="BF215BB6ECC0F798235A5FF4283CC4B7" type="pubDate">
<mods:number id="988346BEC1C6BE5D7A84A9AF1514123B">2023-06-26</mods:number>
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<mods:detail id="175998EA93A3AC152D315AF67A98D45B" type="volume">
<mods:number id="E78589179E409CE8EE9BE88B05910D81">7</mods:number>
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<mods:number id="67D2A6B6B3BD4CECBEA4B672CC15DF80">2</mods:number>
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<mods:url id="A1CCAD8539D88998455BD3DF5AC844B3">http://dx.doi.org/10.3897/evolsyst.7.104597</mods:url>
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<mods:classification id="1E71B382F706B0231FEEB6D8FC22EF44">journal article</mods:classification>
<mods:identifier id="8864DC77063EC41AD88B5D9080D83659" type="DOI">http://dx.doi.org/10.3897/evolsyst.7.104597</mods:identifier>
<mods:identifier id="9EEA988AB1133A581F42B16FBCD0E0DC" type="Pensoft-Pub">2535-0730-2-211</mods:identifier>
<mods:identifier id="9FC16B22EDAEF460A2FCC7A540075138" type="ZooBank">921F39DA40BB47968E362194D1E9A138</mods:identifier>
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<treatment id="2AAF24EF723E5FE590D7ECE00179912A" LSID="urn:lsid:zoobank.org:act:9689A395-8B60-46ED-BFC0-9E87158D2CA6" httpUri="http://treatment.plazi.org/id/2AAF24EF723E5FE590D7ECE00179912A" lastPageNumber="211" pageId="0" pageNumber="211">
<subSubSection id="170C77C8BE582BF952F3F67558B1774F" pageId="0" pageNumber="211" type="nomenclature">
<paragraph id="770B18C7DB7A952E7D5F5EB1C818CB71" pageId="0" pageNumber="211">
<taxonomicName id="0DD86A59209FF168EB7D2BA1C745645B" LSID="https://zoobank.org/9689A395-8B60-46ED-BFC0-9E87158D2CA6" authority="Yamamoto, 2023" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis" status="sp. nov.">Protonicagus mandibularis</taxonomicName>
<taxonomicNameLabel id="8FFDA27DA09258C5427C15C24FA41E78" pageId="0" pageNumber="211">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="0AB24CE45C754DD52F4282D332CD9113" pageId="0" pageNumber="211" type="description">
<paragraph id="DB9CE1F7DC360A1304BFD6E98CCACC8D" pageId="0" pageNumber="211">
<figureCitation id="1A689E92A8FCFAB4D7F301A0B06E5CE1" 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 1A, B</figureCitation>
<figureCitation id="2F09FE5292909B7FC920AFBCAE94A473" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">, 2</figureCitation>
<figureCitation id="1A57DF300D05D500FABDFE19946073D6" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">, 3</figureCitation>
<figureCitation id="980DCD14F4CA0ED6249395E2E2774E96" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">, 4</figureCitation>
<figureCitation id="364D5CC2349599FA791FC995640E6ECB" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">, 5</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="3CC0E1B5806CF23B9A7ED37F737F1360" pageId="0" pageNumber="211" type="materials_examined">
<paragraph id="CD1AD3B0ACF4E293BD2E5DFC87850919" pageId="0" pageNumber="211">Material examined.</paragraph>
<paragraph id="5C017D02C850B564D4BC35370085CEFD" pageId="0" pageNumber="211">
<emphasis id="CD78E1D2C013E30056AF8D43F77A5443" bold="true" italics="true" pageId="0" pageNumber="211">
<typeStatus id="B9851C958F185B819ED9D99263282967">Holotype</typeStatus>
</emphasis>
(sex undertemined), a complete adult preserved in a somewhat cuboid yellowish amber with spherical upper surface, approximately
<quantity id="B264E2B0B8B640ACEA5A5F4325F7C913" metricMagnitude="-3" metricUnit="m" metricValue="7.2" unit="mm" value="7.2">7.2 mm</quantity>
<normalizedToken id="F93FA7CE4D20ADD8111B1F8A1294D6E4" originalValue="×">x</normalizedToken>
<quantity id="329E095B14CA29AB875E46FB3BFF561E" metricMagnitude="-3" metricUnit="m" metricValue="5.4" unit="mm" value="5.4">5.4 mm</quantity>
<normalizedToken id="0D97B01012B73E56BF02DDDEF817FF77" originalValue="×">x</normalizedToken>
<quantity id="110BE44561850D23C4A48711CB32CC04" metricMagnitude="-3" metricUnit="m" metricValue="3.1" unit="mm" value="3.1">3.1 mm</quantity>
in size (Fig.
<figureCitation id="F028AE1C9BE9D6B40BC8176ABCDFB6CD" 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">1A, B</figureCitation>
); specimen accession number SEHU-0000121205, housed in HUM. The
<typeStatus id="107477FC43F8B18C1491BEE8518BE7FE">holotype</typeStatus>
specimen is well preserved, but noticeable pigmentation inside the amber makes detailed observation and photography difficult. In addition, the ventral surface of the lucanid is covered with a thin whitish layer, which makes detailed observation even more difficult (Fig.
<figureCitation id="EA6CF05F4BDBDE86AAA1CF69438C5959" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2B</figureCitation>
).
</paragraph>
<caption id="A8657F3B1FFB9AFB2B945BCA4E31BE87" httpUri="https://binary.pensoft.net/fig/868859" pageId="0" pageNumber="211" start="Figure 1" startId="F1">
<paragraph id="FDA20F7A5AFEA0B5A9697EF3BFDC90A9" pageId="0" pageNumber="211">
<emphasis id="083167E87F185D350C8420C6872E1237" bold="true" pageId="0" pageNumber="211">Figure 1.</emphasis>
General view of amber pieces with lucanid inclusions.
<emphasis id="3D1DD5BECAE486AFFC159524751728A6" bold="true" pageId="0" pageNumber="211">A, B.</emphasis>
<taxonomicName id="463FCE16B0FD69F80B822FE594F68FBB" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="C3ADA8B1AF08E1911CFC548BC8E412D6" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov.,
<typeStatus id="6BA2813D70101676942D01D24E5CF671">holotype</typeStatus>
, SEHU-0000121205, showing dorsal (
<emphasis id="A917973297D3E12FA2921BAFE6A9DE4E" bold="true" pageId="0" pageNumber="211">A</emphasis>
) or lateral (
<emphasis id="7F21704AE0257BCD0FEC22A04CE5B34D" bold="true" pageId="0" pageNumber="211">B</emphasis>
) views of the
<typeStatus id="F3E4442CC4AE281C353BAECB8449985D">holotype</typeStatus>
;
<emphasis id="08A93F70C3DBB1375DAA01BAF9C2E436" bold="true" pageId="0" pageNumber="211">C.</emphasis>
<taxonomicName id="1268BE0CC3B0C6E4F4616AF2608FD981" authorityName="Yamamoto" authorityYear="2023" family="Lucanidae" genus="Cretognathus" kingdom="Animalia" lsidName="Cretognathus minutissimus" order="Coleoptera" pageId="0" pageNumber="211" rank="species" species="minutissimus">
<emphasis id="6C7B49FAFC8B2A042E7232C89F7E2293" italics="true" pageId="0" pageNumber="211">Cretognathus minutissimus</emphasis>
</taxonomicName>
gen. et sp. nov.,
<typeStatus id="B4B546903BDB8E10F1CAD9C30A94657C">holotype</typeStatus>
, SEHU-0000121206;
<emphasis id="824C960379DB3F4D137CCFB1E3EFA3A3" bold="true" pageId="0" pageNumber="211">D.</emphasis>
<taxonomicName id="AEA7D927DD60B95D4A3C6FFDB5234101" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
gen. et sp. indet. 1, SEHU-0000121207;
<emphasis id="191F0953030A0C1AF80857E97DF9443D" bold="true" pageId="0" pageNumber="211">E.</emphasis>
<taxonomicName id="460E7BFE6813018A85F994F20A527777" authorityName="Sharp" authorityYear="1899" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Ceratognathini">Ceratognathini</taxonomicName>
gen. et sp. indet. 2, SEHU-0000121208. Scale bars: 3.0 mm (
<emphasis id="BE55E7011B73E7567993E8FB984CF9EF" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="D2DCBFB187E04FFD51D56EB7B6A97F4B" bold="true" pageId="0" pageNumber="211">B</emphasis>
); 1.0 cm (
<emphasis id="5D42133FE6787B8351256F5AA97BDC5F" bold="true" pageId="0" pageNumber="211">C</emphasis>
); 5.0 mm (
<emphasis id="342DB0442D0D2495E7F1A57542B72C3F" bold="true" pageId="0" pageNumber="211">D, E</emphasis>
).
</paragraph>
</caption>
<caption id="F5A2349AD7A1538CCF7CF315225D7979" httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211" start="Figure 2" startId="F2">
<paragraph id="8E06FBA4B1A17C8098CF8C13CE481E38" pageId="0" pageNumber="211">
<emphasis id="398796BC8B0104834811664880022772" bold="true" pageId="0" pageNumber="211">Figure 2.</emphasis>
General habitus of
<taxonomicName id="665A51EC41ADF375F836C41F875EEEB1" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="A8FDB2389C9B3DC51AAFBA1CCDA3EADD" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov.,
<typeStatus id="2086BB9459953A99FB6CFA159C3B7C15">holotype</typeStatus>
, SEHU-0000121205.
<emphasis id="2DAC4E657FA21ECEE76474520DD8F4FC" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Dorsal view;
<emphasis id="34EFA156627A4BC4378A66617F283EE9" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Ventral view;
<emphasis id="F9F6218F6807D8B1569C20F3218AF45C" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Lateral view. Scale bars: 1.0 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="808F670A95D09A89BFA0E28D0F19C751" pageId="0" pageNumber="211" type="differential diagnosis">
<paragraph id="F0695D109C7EB011418E00A47AFF8545" pageId="0" pageNumber="211">Differential diagnosis.</paragraph>
<paragraph id="EB5D1C16AD3D84BFB711725419F6FF88" pageId="0" pageNumber="211">
<taxonomicName id="C8F333772202C6EFE582F2B66B52E30E" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="D33A8B878B51D56EEBB0089998CD44B2" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov. is most similar to
<taxonomicName id="150FB4E8EC33B6D0574B83DF4E7D0FCF" authorityName="Cai, Yin, Liu &amp; Huang" authorityYear="2017" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus tani" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="tani">
<emphasis id="8FD592FA1E544B5990F8F26DDE6E35DE" italics="true" pageId="0" pageNumber="211">Protonicagus tani</emphasis>
</taxonomicName>
Cai, Yin, Liu &amp; Huang, 2017, also from Kachin amber, based on external morphological features in having a similar body size (3.62
<emphasis id="83C43AB843A2CFA2ED0AB6E8B8F98105" italics="true" pageId="0" pageNumber="211">versus</emphasis>
3.71 mm in
<taxonomicName id="C8CC614D16F8C6C7BC9987D62312A589" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="5B0E8CC0E863935B681305F5B6789647" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
), preocular margin (
<emphasis id="E76ADCBBE5503363F3B9EFF6E1ADACD4" italics="true" pageId="0" pageNumber="211">sensu</emphasis>
<bibRefCitation id="DB72F68543DD44A8FBC04701E73CD66D" 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>
: fig. 3) of the eyes strongly enlarged and produced laterally, pronotal lateral margin rounded and strongly crenulate with a row of setae, similar structures and arrangements of outer teeth along the protibiae with strong spines in basal half, legs with short pretarsal claw, and each posterior margin of abdominal ventrites 2-4 armed with a row of villiform teeth. However, the new species is easily distinguished from the latter by its general habitus (narrowly elongate
<emphasis id="3A2BE3CE120B45ECC8BF49F86DF764CE" italics="true" pageId="0" pageNumber="211">versus</emphasis>
rounded and oval in
<taxonomicName id="E219E93C21F38FB1C530FCD57AB69770" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="BD44C2750C70104094337F2C6028C1C0" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
), much larger non-acute apices of mandibles (acute in
<taxonomicName id="54C1A891CC056C688DD325673D014CB4" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="B258624F08E805792FF6F358902F8EB6" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
), dorsal surface densely covered with thin and long ground setae (rather than short scales as in
<taxonomicName id="31EABF44E782B75AEDB7D74F926D1FCC" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="12AB9F3E831E900DE2FFDD45E9AD18E0" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
;
<bibRefCitation id="B872AD875E04934C94D003DA72C59BD6" 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>
: fig. 2B-D), much wider antennomeres 2-7, and obviously longer narrower metatarsi (see further details in
<bibRefCitation id="DD2F695E1F281B600CFEE8E2A9EFEF73" 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>
).
</paragraph>
<caption id="F8578013CDEF29285C657CA362F67DE0" httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211" start="Figure 3" startId="F3">
<paragraph id="4317CB37F0C9399061DBEAB3A8385F8B" pageId="0" pageNumber="211">
<emphasis id="9EC2D470459C51DFA41A1AA9840FAA45" bold="true" pageId="0" pageNumber="211">Figure 3.</emphasis>
Details of
<taxonomicName id="F643022CBEA85B6256B461BB5C5B8FDF" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="B473D0B8FDF948B94627CA19873459ED" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov., holotype, SEHU-0000121205.
<emphasis id="1A0E503E11C27262616272728B437600" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Head with left antenna, dorsal view;
<emphasis id="83C6013550769377CF2C516156846FB6" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Head, dorsolateral view;
<emphasis id="3C1D8CFF56F2E420D6328973EB6BC78B" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Pronotum, left lateral half, dorsal view;
<emphasis id="9265FD89CAA9F5E09D47339F4D4632F8" bold="true" pageId="0" pageNumber="211">D.</emphasis>
Pronotum, dorsolateral view;
<emphasis id="800324407695806DE0E08BE3F61E90F0" bold="true" pageId="0" pageNumber="211">E.</emphasis>
Elytral base with shoulder, dorsolateral view. Abbreviations: a1-10, antennomeres 1-10; ey, compound eye; lb, labrum; lp3, labial palpomere 3; md, mandible; mp3-4, maxillary palpomeres 3-4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (
<emphasis id="A71A4338F72557AFFA904F004814C4A8" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="CA14A2B0F927A0AB0FB2030E3AF51278" bold="true" pageId="0" pageNumber="211">B</emphasis>
); 0.5 mm (
<emphasis id="0CCA5AAA18B29B2FD128E548E12C9879" bold="true" pageId="0" pageNumber="211">C</emphasis>
,
<emphasis id="F43207B44B547760DE275ABDE1667178" bold="true" pageId="0" pageNumber="211">D</emphasis>
,
<emphasis id="05CC392426F9B7754C7D0CD38C996E48" bold="true" pageId="0" pageNumber="211">E</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="1C3C264645608F5410CD4DC5CA4016B1" pageId="0" pageNumber="211" type="diagnosis">
<paragraph id="02C29F647A158CC901A3FC82AEFCD064" pageId="0" pageNumber="211">Diagnosis.</paragraph>
<paragraph id="AD863E8A8F2A948BF862E29933FA9FC2" pageId="0" pageNumber="211">
Body small (ca. 3.6 mm), moderately elongate, subparallel sided (Fig.
<figureCitation id="70E99372C34CE963237372202FB2F366" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2A, B</figureCitation>
). Dorsal surface covered with thin and long setae, lacking modified scales (Fig.
<figureCitation id="EE4B651451D48405F530DCBE62F750A9" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3</figureCitation>
). Antenna with spherical to transverse antennomeres 2-7 and three-segmented small club (Fig.
<figureCitation id="08F62A9EE3EF7B823B6FB839276384A6" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
). Mandibles shorter than head, mostly visible dorsally, rather strongly curved and flattened, with rounded apices and nearly truncate apico-inner margins (Fig.
<figureCitation id="1A2C8CEBE8159F64D3B5C8430E9B3264" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A, B</figureCitation>
). Pronotum strongly and very densely punctate (Fig.
<figureCitation id="C77318B97353ADD2206710628BF1343C" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3C, D</figureCitation>
); lateral pronotal margins weakly crenulate, each with a row of setae (Fig.
<figureCitation id="52AF49317E27C158DB89042494594B99" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3C, D</figureCitation>
,
<emphasis id="F79EB97E04F9AB2423E38700FACF079B" italics="true" pageId="0" pageNumber="211">rsp</emphasis>
). Outer protibial edge with one large apical tooth (
<emphasis id="EEC29B8CD262EFE2A98D70A2318FA05A" italics="true" pageId="0" pageNumber="211">at</emphasis>
), one large and one smaller mid-dorsal teeth (
<emphasis id="F9D389307FD8067EAF53757202BAC1B6" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
-
<emphasis id="F242643F877049732E5AA693C559E5FB" italics="true" pageId="0" pageNumber="211">2</emphasis>
), and less than ten generally subequal smaller tooth-like spines (Fig.
<figureCitation id="AE70416E17C4C480CA0237BF8DD4EDC8" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4A, B</figureCitation>
). Metatarsi long and slender, but shorter than metatibiae (Fig.
<figureCitation id="B68A48F3B932A9D469F4DCA2DB3F17FD" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="AE480F3ABA04C31840CADAAE4D2859A2" pageId="0" pageNumber="211" type="description">
<paragraph id="0CFB273BC53CA3E6BDC8DF83FFC35B07" pageId="0" pageNumber="211">Description.</paragraph>
<paragraph id="B9929730FE01BC8E6893C75A468D0C94" pageId="0" pageNumber="211">
<emphasis id="649B67C477FDAD3CEDE2C2209FDEDAFF" bold="true" italics="true" pageId="0" pageNumber="211">Body</emphasis>
(Fig.
<figureCitation id="BB9FC7410F8B987F2A712AB0F4589129" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2</figureCitation>
) elongate oval, small, 3.62 mm long (measured from apex of clypeus to apex of elytra), 1.45 mm wide, moderately covex dorsoventrally. Color uniformly dark brown; antennae and mouthparts slightly paler. Dorsum (Figs
<figureCitation id="579162DC24BC508599304B5C32FF5E02" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2A</figureCitation>
,
<figureCitation id="66146A06C50FBBACBA8D61D1BA071F66" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3</figureCitation>
) without modified scales, densely deeply punctate, setiferous punctures on pronotum and elytra large, each bearing suberect thin seta.
</paragraph>
<paragraph id="055D96BB4D4DCB5175108AD90C3E6CDD" pageId="0" pageNumber="211">
<emphasis id="F2744F59CDD95F26213C8885B7F62D7E" bold="true" italics="true" pageId="0" pageNumber="211">Head</emphasis>
(Fig.
<figureCitation id="B55079750512529BD5C09A305463F584" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A, B</figureCitation>
) small, rather strongly transverse, shorter than half pronotal length and narrower than half pronotal width, 0.44 mm long (measured from apex of clypeus to anterior margin of pronotum) and 0.57 mm wide across eyes; ventral side not well visible; vertex (Fig.
<figureCitation id="17B9AAB68D575397C421E0ABDF9CA84E" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
) densely punctate; preocular margins (Fig.
<figureCitation id="C4F35F42AF1AC4BAE8291F85B9B6EC4C" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
,
<emphasis id="8CE8F06A794C25E03A834D097BF6D13A" italics="true" pageId="0" pageNumber="211">pom</emphasis>
) conspicuous, strongly enlarged, protruding anterolaterally. Compound eye (Fig.
<figureCitation id="6E127CC40786BAC32B28B7644C47B2F2" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A, B</figureCitation>
,
<emphasis id="3B88B560D569A54BB86935C614D369E8" italics="true" pageId="0" pageNumber="211">ey</emphasis>
) relatively large, entire, not divided by ocular canthus. Antenna (Fig.
<figureCitation id="6A7917388AC0C4E7BC28DAB57213BE7C" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
,
<emphasis id="A0C9EDA63E6A4DD6B2304707F38425A2" italics="true" pageId="0" pageNumber="211">a1-10</emphasis>
) 10-segmented, non-geniculate, with three-segmented weakly lamellate club, 0.66 mm long (left antenna); antennomere 1 (scape) elongate, approximately 2.7 times longer than wide, sparsely bearing nine long and thin bristles, not covered with many setae; antennomere 2 (pedicel) very small, spherical, wider than long, with probably only a single bristle, attached to tip of scape; antennomeres 3-6 evenly weakly widened apicad; antennomere 3 small, as long as wide, narrower than preceding antennomere, weakly dilated apically; antennomere 4 moderately transverse, slightly wider than preceding antennomere; antennomere 5 rather strongly transverse, wider than preceding antennomere; antennomere 6 strongly transverse, moderately wider than preceding antennomere; antennomere 7 strongly transverse, slightly shorter than preceding antennomere; antennomeres 8-10 forming small and relatively loose club, strongly asymmetrical, each not coherent. Mandibles (Fig.
<figureCitation id="62BD89EB2C4DCC7A02A9802CED56628A" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A, B</figureCitation>
,
<emphasis id="49F1D31560D5F1FD6C0317ACC3C9A659" italics="true" pageId="0" pageNumber="211">md</emphasis>
) mostly visible dorsally, slightly asymmetrical, small, shorter than head, bent strongly inward, dorsoventrally flattened and widened, not extending beyond maxillary palps, with rounded, non-acute apex and nearly truncate apico-inner margin. Labrum (Fig.
<figureCitation id="83FAB0C2EABA19007FE585094B042C18" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
,
<emphasis id="F378DA97BA1BEA0F46FCCE6C909E1C5B" italics="true" pageId="0" pageNumber="211">lb</emphasis>
) small, transverse, with nearly truncate apex. Clypeus with produced anterior margin. Maxillary palpus (Fig.
<figureCitation id="328545737226E90907AFFFC92628B960" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A, B</figureCitation>
,
<emphasis id="E1A2BE9A18D4FDD2DC0F7B60CE6A7192" italics="true" pageId="0" pageNumber="211">mp3-4</emphasis>
) 4-segmented, moderately long, slender; palpomere 2 small; palpomere 3 spindle-shaped, widest in middle, about half length as terminal palpomere, with a long bristle near apex on outer margin; palpomere 4 very long, slender, fusiform, slightly bent inward near base, narrower than preceding palpomere. Labial palpus (Fig.
<figureCitation id="22B3447ABE783BDF3B9FE1CA41277BBF" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3B</figureCitation>
,
<emphasis id="DBA153BBED264582DDB12AFE93950D5E" italics="true" pageId="0" pageNumber="211">lp3</emphasis>
) probably three-segmented, long; palpomere 3 very long, slender, fusiform, slightly curved.
</paragraph>
<paragraph id="609CAEBAEE32A90D617E8CE7B5E710BE" pageId="0" pageNumber="211">
<emphasis id="723C1CA6BA4B7F1CF945F79360837DFF" bold="true" italics="true" pageId="0" pageNumber="211">Pronotum</emphasis>
(Figs
<figureCitation id="7AF9D2263A3D950331569E2517EE9498" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2A</figureCitation>
,
<figureCitation id="EC6BD9432C76BFC75C8A8D8AD2F1E51C" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3C, D</figureCitation>
) transverse, widest in middle, 1.39 mm wide and 1.06 mm long (along midline), dorsally convex, with weak basolateral constriction; lateral margin (Fig.
<figureCitation id="E8CA546C048852F669845BD008A9C0C1" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3C, D</figureCitation>
) broadly rounded, modetately crenulate along whole margin, equipped with a row of ventrolaterally-directed long setae; surface simple and even; anterior corners slightly produced with obtuse angles, whereas those of posterior ones right-angled. Procoxal cavity externally closed behind. Prosternum short, seemingly strongly transverse; prosternal process between procoxae very narrow, resulting in subcontiguous procoxae (Fig.
<figureCitation id="D55CF5FACB90EAF12DC416E72A68E47F" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5A, C</figureCitation>
, arrow).
</paragraph>
<paragraph id="6B3CBA8DC93D865A1D47189729C5B8AC" pageId="0" pageNumber="211">
<emphasis id="EABF12FBF95B938224C077F1CA805A1B" bold="true" italics="true" pageId="0" pageNumber="211">Elytra</emphasis>
(Figs
<figureCitation id="07730A71685DBD0B1CE30142BC1D6D1A" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2A, C</figureCitation>
,
<figureCitation id="CC16B954B374645E58420D013430F6A7" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3E</figureCitation>
) complete, narrowly elongate, nearly subparallel-sided in anterior three-quarters, with left elytron 2.24 mm long (measured from pronotal posterior end to apex of elytron) and 0.70 mm wide; lateral margins serrulate, each with a row of setae; surface smooth without tubercles, but densely covered with deep dense setiferous punctures uniformly, seemingly not forming longitudinal rows (Fig.
<figureCitation id="9FFFF78914320AE010E2908A1943437F" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Head with left antenna, dorsal view; B. Head, dorsolateral view; C. Pronotum, left lateral half, dorsal view; D. Pronotum, dorsolateral view; E. Elytral base with shoulder, dorsolateral view. Abbreviations: a 1 - 10, antennomeres 1 - 10; ey, compound eye; lb, labrum; lp 3, labial palpomere 3; md, mandible; mp 3 - 4, maxillary palpomeres 3 - 4; pn, pronotum; pom, preocular margin of head; rsp, row of setae along outer margin of pronotum. Scale bars: 0.3 mm (A, B); 0.5 mm (C, D, E)." httpUri="https://binary.pensoft.net/fig/868861" pageId="0" pageNumber="211">3A</figureCitation>
). Mesocoxal cavities narrowly separated. Metaventrite (Fig.
<figureCitation id="20A3A8A8F43C3746C4D24CC68799E7AC" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5</figureCitation>
,
<emphasis id="4F76CD08C4DB24682C0BDDDAF97668B4" italics="true" pageId="0" pageNumber="211">mtv</emphasis>
) transverse, much longer than mesoventrite; surface generally densely punctured, with short but prominent discrimen (median longitudinal sulcus; see Fig.
<figureCitation id="321AEB9278DF50102079DFB3489EB2BD" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5</figureCitation>
, arrow).
</paragraph>
<paragraph id="9B58F0904AB60F40FED1E4D47D073329" pageId="0" pageNumber="211">
<emphasis id="0D0EB2D0473F4344018DFB47A75CCFAD" bold="true" italics="true" pageId="0" pageNumber="211">Legs</emphasis>
(Figs
<figureCitation id="F6AF55203E700EB458A97345BF3E8E03" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2</figureCitation>
-
<figureCitation id="52F7A3027D8D440F40E3034C7F560C11" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5</figureCitation>
) relatively short. Protibia (Fig.
<figureCitation id="F593FE2D3AA6D5148ECB8EB4CFE24125" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2A, B</figureCitation>
) robust, flattened, weakly curved, gradually widened apically, with one large apical spur; external margin armed with three strong teeth in apical half, i.e., one large, broadly pointed apical tooth (
<emphasis id="B71156AFBDB411384578425A549D8852" italics="true" pageId="0" pageNumber="211">at</emphasis>
), one large, broadly pointed mid-dorsal tooth 2 (
<emphasis id="690A23A1554CECE996265247B05C2127" italics="true" pageId="0" pageNumber="211">mdt2</emphasis>
), and one smaller but still prominent mid-dorsal tooth 1 (
<emphasis id="C134C9610DCA73289C33F056F91A9A3F" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
) (
<emphasis id="57F4B57A923ABFBF26C68548DDDE2D24" italics="true" pageId="0" pageNumber="211">sensu</emphasis>
<bibRefCitation id="80E71A9A0FD618B40BC9F587E16731A5" 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>
: fig. 1), with 9 short, acute, and generally subequal smaller tooth-like spines (8 subcontiguous ones between base and
<emphasis id="60183E898B1136C79FF0F5E7C6C69C2D" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
, 1 between
<emphasis id="82C8FB1233702E29D7F5D8290F40AFC6" italics="true" pageId="0" pageNumber="211">mdt1</emphasis>
and
<emphasis id="6F066A2BE9BBB1B3474AC20F3312A9CE" italics="true" pageId="0" pageNumber="211">mdt2</emphasis>
, and 0 between
<emphasis id="118AB4EF5A3B9C3DA8DEF54D1110328A" italics="true" pageId="0" pageNumber="211">mdt2</emphasis>
and
<emphasis id="AD74A42EC313C4F7DEC4A99CEB2ECB26" italics="true" pageId="0" pageNumber="211">at</emphasis>
, see arrows on Fig.
<figureCitation id="F5FCE09FBCF324D25DC7FDB7285546BE" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4A, B, E</figureCitation>
for the latter condition). Procoxa widely transverse. Mesotibia with three small mid-dorsal spines at middle (Fig.
<figureCitation id="4627C7F722A53C59CA49AF16FE359991" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4H</figureCitation>
,
<emphasis id="7F0F5B7A82307A9323FA9BA517E8EE6B" italics="true" pageId="0" pageNumber="211">mds</emphasis>
), with paired apical spurs (Fig.
<figureCitation id="F15913CC58A21ECE3E308BD14889FFEB" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4C, G, H</figureCitation>
,
<emphasis id="DADF5506B7BBB05C6B0366985B2F283F" italics="true" pageId="0" pageNumber="211">as</emphasis>
). Metacoxae (Fig.
<figureCitation id="6DABD561DAEDB0B52F71BC901BD67761" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5</figureCitation>
,
<emphasis id="09C987FD9F979D7E8F238A02C731650B" italics="true" pageId="0" pageNumber="211">mtc</emphasis>
) transverse, subcontiguous. Metatibia (Fig.
<figureCitation id="4F938490CA4EB163FEA22B8D297560DA" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4B</figureCitation>
) slender, only slightly longer than metatarsus, with three small mid-dorsal spines located at slightly beyond middle apically (Fig.
<figureCitation id="F475F40081A313D35310C3F4D68A9857" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4D, F</figureCitation>
,
<emphasis id="D05241456CC950FD16B5BD03103AC7F1" italics="true" pageId="0" pageNumber="211">mds</emphasis>
) and paired prominent apical spurs (Fig.
<figureCitation id="6DF9ACFE72E804A681F5A0BA2149D105" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4D, F</figureCitation>
,
<emphasis id="856FE9A42E209BAFF87C431F2E991C8A" italics="true" pageId="0" pageNumber="211">as</emphasis>
). All tarsi 5-segmented, each slender and rather long; tarsomere 1 longer than tarsomere 2, tarsomeres 2-4 subequal in length, tarsomere 5 longest, as long as tarsomeres 2-4 combined (Fig.
<figureCitation id="2AEA954FC381F17052B7186CC1AD870B" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4A, C-H</figureCitation>
). Pretarsal claw (Fig.
<figureCitation id="29B63D5DF76D57316BEB9E729A40AFEE" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4A, C-H</figureCitation>
,
<emphasis id="F682178A0434456C6579F373A6C80B41" italics="true" pageId="0" pageNumber="211">cl</emphasis>
) short, simple, moderately curved, shorter than tarsomeres 3 and 4 combined. Arolium (Fig.
<figureCitation id="E0B3C339EEE88FA3D0260FC85E05028A" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4C, D, G, H</figureCitation>
) developed, with short rod (Fig.
<figureCitation id="AEC23503D4A374F574719B041608C647" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4D</figureCitation>
,
<emphasis id="9DC165D2F0CF50B9C35100845F452F36" italics="true" pageId="0" pageNumber="211">ra</emphasis>
), bearing 2 bristles on its apex (Fig.
<figureCitation id="D0987E08EB27FFC710F03A8CDCA13374" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Right protibia and protarsus, frontal view; B. Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view; C. Left mesotibia and mesotarsus, dorsolateral view; D. Metatibiae and metatarsi, lateral view; E. Apex of right protibia, frontal view; F. Metatibiae and metatarsi, dorsal view; G. Left mesotarsus, ventrolateral view; H. Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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 2 - 5, protarsomeres 2 - 5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (A - D, F); 0.3 mm (E, G, H)." httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211">4D</figureCitation>
,
<emphasis id="978BFC8E527448439B20C6628B9A0119" italics="true" pageId="0" pageNumber="211">sa</emphasis>
).
</paragraph>
<caption id="F11E35BEC06A4EFD672B27B430028117" httpUri="https://binary.pensoft.net/fig/868862" pageId="0" pageNumber="211" start="Figure 4" startId="F4">
<paragraph id="9D0EC8C80751B30505BC5711F1DCC4E4" pageId="0" pageNumber="211">
<emphasis id="2076BBFBC008C81E37A741878B491D95" bold="true" pageId="0" pageNumber="211">Figure 4.</emphasis>
Details of
<taxonomicName id="A49F822E5E51BD7F1D6C091BE3D74E99" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="017A96FD48DA14C07611A50F9A6307E0" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov., holotype, SEHU-0000121205.
<emphasis id="3151A20304B5F8928BDE21DE1702CFAE" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Right protibia and protarsus, frontal view;
<emphasis id="2C32FDF22B6AECD5B2FDB8CFAFBAD0F6" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Left protibia with arrow showing absence of small teeth between large spines along outer edge of protibia, frontal view;
<emphasis id="25DFA96E175129EF73AE4D59DCB2F757" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Left mesotibia and mesotarsus, dorsolateral view;
<emphasis id="48B387AC5CCA48975A419E4E501100AE" bold="true" pageId="0" pageNumber="211">D.</emphasis>
Metatibiae and metatarsi, lateral view;
<emphasis id="701D004EDC6BE0930DB9B677AB08072E" bold="true" pageId="0" pageNumber="211">E.</emphasis>
Apex of right protibia, frontal view;
<emphasis id="79A7E04D16E3DE6EC6E58FE60BFAA563" bold="true" pageId="0" pageNumber="211">F.</emphasis>
Metatibiae and metatarsi, dorsal view;
<emphasis id="7E5B6D0D13F718D826FFAF4AFCE2237D" bold="true" pageId="0" pageNumber="211">G.</emphasis>
Left mesotarsus, ventrolateral view;
<emphasis id="84E0962B8F5A8E585364A2849EC4C631" bold="true" pageId="0" pageNumber="211">H.</emphasis>
Left mesotarsus, ventral view. Abbreviations: as, apical spur; at, apical tooth of protibia; cl, claw; el, 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; pt2-5, protarsomeres 2-5; pti, protibia; ra, rod of arolium; sa, setae on rod of arolium. Scale bars: 0.5 mm (
<emphasis id="191A79C0F46A6D179C33C8C8E95B64A6" bold="true" pageId="0" pageNumber="211">A</emphasis>
-
<emphasis id="5A345E361007A7F41BE56E0BD85FE016" bold="true" pageId="0" pageNumber="211">D</emphasis>
,
<emphasis id="7C3ADBEB460D6522AE360EA66A08F53C" bold="true" pageId="0" pageNumber="211">F</emphasis>
); 0.3 mm (
<emphasis id="6ED23033863A040354ED8B4564AB24AE" bold="true" pageId="0" pageNumber="211">E</emphasis>
,
<emphasis id="56A89A7F3270D108A14F30A9F39C1CAA" bold="true" pageId="0" pageNumber="211">G</emphasis>
,
<emphasis id="CB9133B4A6A4BFBA015EB9CE4DAD240C" bold="true" pageId="0" pageNumber="211">H</emphasis>
).
</paragraph>
</caption>
<caption id="974A12BD16DEB6ED60B992F3B9B0CE96" httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211" start="Figure 5" startId="F5">
<paragraph id="5996E7031900AC3CE028DE6AC1F3836B" pageId="0" pageNumber="211">
<emphasis id="4036DEFCF9C745FF1B93DA8B78813DDF" bold="true" pageId="0" pageNumber="211">Figure 5.</emphasis>
Details of
<taxonomicName id="78F4AA804749AC2F20118A63F81D6AAA" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="263287234D941A0DAEA0EE83A63EFC07" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov., holotype, SEHU-0000121205.
<emphasis id="4870397F0636FB95AF3C6737BFA83885" bold="true" pageId="0" pageNumber="211">A.</emphasis>
Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view;
<emphasis id="04B8BD47B438473B422FF554C9893099" bold="true" pageId="0" pageNumber="211">B.</emphasis>
Metathorax and abdomen, ventral view;
<emphasis id="AB70CC46FD875160A74257315753C57F" bold="true" pageId="0" pageNumber="211">C.</emphasis>
Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v1-5, ventrites 1-5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (
<emphasis id="712499B00DED366529A9A804D3DFA200" bold="true" pageId="0" pageNumber="211">A</emphasis>
,
<emphasis id="42C423B0F4F700A7E059E6923561D2BA" bold="true" pageId="0" pageNumber="211">B</emphasis>
); 0.3 mm (
<emphasis id="2F950D864EAD5C702151541EC9ADCC19" bold="true" pageId="0" pageNumber="211">C</emphasis>
).
</paragraph>
</caption>
<paragraph id="515855A779C666AF2DDF004C59C3A4E9" pageId="0" pageNumber="211">
<emphasis id="E4497CA13170815BF2BB596A98B803FB" bold="true" italics="true" pageId="0" pageNumber="211">Abdomen</emphasis>
(Figs
<figureCitation id="9602CA357CDFF5496566E5EE884AD280" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. General habitus of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Dorsal view; B. Ventral view; C. Lateral view. Scale bars: 1.0 mm." httpUri="https://binary.pensoft.net/fig/868860" pageId="0" pageNumber="211">2B</figureCitation>
,
<figureCitation id="24A180DEF73E2D7B9325CB40A6386481" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Details of Protonicagus mandibularis sp. nov., holotype, SEHU- 0000121205. A. Meso- and metathorax with arrow showing discrimen (median longitudinal sulcus) on metaventrite, ventral view; B. Metathorax and abdomen, ventral view; C. Pro-, meso-, and metathorax with arrow showing subcontiguous procoxae, ventral view. Abbreviations: msf, mesofemur; msti, mesotibia; msv, mesoventrite; mtc, metacoxal cavity; mtf, metafemur; mtti, metatibia; mtv, metaventrite; pf, profemur; ph, pronotal hypomeron; v 1 - 5, ventrites 1 - 5; vt, row of villiform teeth on margins of ventrites. Scale bars: 0.5 mm (A, B); 0.3 mm (C)." httpUri="https://binary.pensoft.net/fig/868863" pageId="0" pageNumber="211">5B</figureCitation>
) with 5 visible ventrites (
<emphasis id="3DDCCFD5832E7798489A78C420B707B5" italics="true" pageId="0" pageNumber="211">v1</emphasis>
-
<emphasis id="8C3FC57C8FCB3C60C77D646C47B0C58E" italics="true" pageId="0" pageNumber="211">5</emphasis>
); central part rather swelled above; each ventrite strongly transverse; ventrites 2-4, at least, each armed with a row of large villiform teeth along posterior margin; ventrite 5 with posterior margin broadly arcuate.
</paragraph>
<paragraph id="40AAD9A8911E4844FB1409655056C9D8" pageId="0" pageNumber="211">
<emphasis id="BDF0211017C01053D701E7AEB466341C" bold="true" italics="true" pageId="0" pageNumber="211">Genitalia</emphasis>
not visible.
</paragraph>
</subSubSection>
<subSubSection id="12A224142A828A58A5950B8A5E351CF3" pageId="0" pageNumber="211" type="etymology">
<paragraph id="C7DE5D15C792107DBC452EC1FFDB2013" pageId="0" pageNumber="211">Etymology.</paragraph>
<paragraph id="5298DA30EAD0A4916F7790EBA7F4760F" pageId="0" pageNumber="211">
The specific name &quot;
<emphasis id="2E0736A17D497C582F43904D565ABCEC" italics="true" pageId="0" pageNumber="211">mandibularis</emphasis>
&quot; (Latin
<emphasis id="BEF5FE0C60A2AC31F5F74562492B388E" italics="true" pageId="0" pageNumber="211">mandibulum</emphasis>
+ -
<emphasis id="1806C11E4D01BA48C9BD270A78D9474A" italics="true" pageId="0" pageNumber="211">aris</emphasis>
) refers to its unusual shape of the mandibles.
</paragraph>
</subSubSection>
<subSubSection id="D549F8C94AA61035826D58623BA168AB" pageId="0" pageNumber="211" type="locality and horizon">
<paragraph id="C4CE00E1412009EAF23A6B96E87668EA" pageId="0" pageNumber="211">Locality and horizon.</paragraph>
<paragraph id="56280238946EA201D879A86573D858CE" pageId="0" pageNumber="211">
Hukawng Valley (
<geoCoordinate id="B3098DB80B141821F3C162DA2B0AC85A" degrees="26" direction="north" minutes="20" orientation="latitude" precision="925" value="26.333334">26°20'N</geoCoordinate>
,
<geoCoordinate id="B8B8C94CFD6EF42FBC2F8B5145576EDA" degrees="96" direction="east" minutes="36" orientation="longitude" precision="925" value="96.6">96°36'E</geoCoordinate>
), Kachin State, northern Myanmar; unnamed horizon, mid-Cretaceous, Upper Albian to Lower Cenomanian.
</paragraph>
</subSubSection>
<subSubSection id="57C0F6C4E23164C07080ED08E56514B7" pageId="0" pageNumber="211" type="systematic placement and comparison">
<paragraph id="2D4583CBA9ABE415DB7973583A6DCE1C" pageId="0" pageNumber="211">Systematic placement and comparison.</paragraph>
<paragraph id="6D73B80DE34148AA9373E320942D21B3" pageId="0" pageNumber="211">
<taxonomicName id="2D7B2D395C186F0E30D8DFBDB39000EB" authorityName="Yamamoto" authorityYear="2023" class="Insecta" family="Lucanidae" genus="Protonicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protonicagus mandibularis" order="Hymenoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="mandibularis">
<emphasis id="6AFC94F21279B89A9F3C1E64375A9229" italics="true" pageId="0" pageNumber="211">Protonicagus mandibularis</emphasis>
</taxonomicName>
sp. nov. can be assigned to the scarabaeoid family
<taxonomicName id="A118A1FC19EA934CDF2581E6AA9CB2C7" 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 narrowly elongate body shape, 10-segmented antennae with three-segmented, non-coherent, lamellate apical club, anteriorly projecting and rather developed mandibles, 5-5-5 tarsal formula, protibial structures with characteristic outer teeth, developed bisetose arolium between the pretarsal claws, and five visible abdominal ventrites (e.g.,
<bibRefCitation id="39671EC837B0AE32E4496D0E9D252D7F" 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="36C4D0938CE87F37B7A000EDF5ADD822" 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="F6A8C6394BA8881BE543AA8B539CD9C2" 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="80A86981D93A9340D38911FE2DF8C22F" 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 fossil taxon is assigned to the ancestral extant subfamily
<taxonomicName id="5FECE8B4B5AAFFFB85321CA5C4B824E3" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Aesalinae">Aesalinae</taxonomicName>
based on its non-geniculate antennae, complete eyes (not divided by ocular canthus), small mandibles, non-striate elytra, and strongly narrowed prosternal process with subcontiguous procoxae. It is easily separated from the New Zealand endemic
<taxonomicName id="823B268227E13579A722A7BAB0654454" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Lampriminae">Lampriminae</taxonomicName>
,
<taxonomicName id="3CACF059CC8ACD5B0469F8EC7178236E" authorityName="White" authorityYear="1846" class="Insecta" family="Lucanidae" genus="Dendroblax" kingdom="Animalia" lsidName="Dendroblax earlii" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="species" species="earlii">
<emphasis id="50B5120E10E0AD410C38544DD58C1A55" italics="true" pageId="0" pageNumber="211">Dendroblax earlii</emphasis>
</taxonomicName>
White with a
<taxonomicName id="4431E9867C8E164BE2D633FC96A62AF4" authorityName="Leconte" authorityYear="1861" class="Insecta" family="Lucanidae" genus="Nicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nicagus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="3934CDB137545EBE08F569934475CD38" italics="true" pageId="0" pageNumber="211">Nicagus</emphasis>
</taxonomicName>
-like body and short mandibles in both sexes, by the considerably different locations and shapes of the protibial outer teeth, distinctly smaller body size, glabrous metaventrite, and much longer linear antennomere 1 (cf.
<bibRefCitation id="05C22C100735CD0EDBB4EC62133C2ADF" author="Fujita, H" journalOrPublisher="Mushi-sha, Tokyo" pageId="0" pageNumber="211" refId="B13" refString="Fujita, H, 2010. The Lucanid Beetles of the World. Mushi-sha's Iconogrpahic Series of Insects 6. Mushi-sha, Tokyo" title="The Lucanid Beetles of the World. Mushi-sha's Iconogrpahic Series of Insects 6." year="2010">Fujita 2010</bibRefCitation>
;
<bibRefCitation id="A02A3C5051353A47138490E67DBCC824" 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>
). Within
<taxonomicName id="D9EFA5024B129BF4803C295A46798AD8" authorityName="Macleay" authorityYear="1819" lsidName="" pageId="0" pageNumber="211" rank="subFamily" subFamily="Aesalinae">Aesalinae</taxonomicName>
, the new species is similar externally to the extinct genus
<taxonomicName id="32991B80441C11E8368E33421A8327A5" 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="1A3EA32B3E465D0BAB2B1A4D2F207384" italics="true" pageId="0" pageNumber="211">Protonicagus</emphasis>
</taxonomicName>
, also from Kachin amber (tribe
<taxonomicName id="EB26429404932820DE733D1139FA6173" authorityName="Leconte" authorityYear="1861" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Nicagini">Nicagini</taxonomicName>
), including the similar body size, notably enlarged lateral margins of preocular margin of the eyes, rounded lateral margin of the pronotum with strong crenulation bearing a row of setae, similar structures of the protibiae, and abdominal ventrites 2-4 each armed with a row of villiform teeth (
<bibRefCitation id="F1B6C1C1FF5427FBCA6360DC6F2345F5" 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>
). However, I found several distinct features that do not match
<taxonomicName id="60ADF0D7C5E6000FAFA36B7BD06075F4" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="988BBA586BA05A78B047A1D032F4E48D" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
, the sole member of
<taxonomicName id="F64AEBFA089742B22374F1AAC92E09F0" 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="BF6EB1981D0EE93A380D53BFA9032338" italics="true" pageId="0" pageNumber="211">Protonicagus</emphasis>
</taxonomicName>
: 1) much narrower, elongate body; 2) clearly larger non-acute mandibles; 3) absence of dense, inclined, linear scales on the pronotum and elytra, but covered with thin, long setae; 4) much wider antennomeres 2-7; and 5) obviously longer and narrower metatarsi with longer tarsomeres 1 (cf.
<bibRefCitation id="5A7DBD5504D1F60263435DA6FEC1A599" 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>
). From the sole extant genus
<taxonomicName id="D0452FCA9BD49DE9234A04D3A6C976EB" authorityName="Leconte" authorityYear="1861" class="Insecta" family="Lucanidae" genus="Nicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nicagus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="67FBD38A0BCADA36F2B61082F8C724E9" italics="true" pageId="0" pageNumber="211">Nicagus</emphasis>
</taxonomicName>
in
<taxonomicName id="9713CE9E0A676F207EBF2F4D6A015F45" authorityName="Leconte" authorityYear="1861" lsidName="" pageId="0" pageNumber="211" rank="tribe" tribe="Nicagini">Nicagini</taxonomicName>
,
<taxonomicName id="7C3B708B77A5494945D006BCB20D6095" lsidName="P. mandibularis" pageId="0" pageNumber="211" rank="species" species="mandibularis">
<emphasis id="1BC9797E4A43EB9FD6D0CF168E063082" italics="true" pageId="0" pageNumber="211">P. mandibularis</emphasis>
</taxonomicName>
sp. nov. is ruled out mainly by the characters mentioned in
<bibRefCitation id="21548E4BDEC284662B8154A6950E66A1" 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>
and described above. Therefore, I prefer to tentatively include it in
<taxonomicName id="AAE986222B7FC5AD1BB4974DAC249DF2" 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="36E58D7F5C903DCAE0BFDF5AD93AB4E5" italics="true" pageId="0" pageNumber="211">Protonicagus</emphasis>
</taxonomicName>
because of the general morphological similarity.
</paragraph>
<paragraph id="782B8EE4B228E7088D4135F62B3C4CC2" pageId="0" pageNumber="211">
Given the distinct morphological differences in lucanid species due to sexual dimorphism, it is difficult to determine whether such differences are interspecific characters within the genus or just individual variation between the different sexes of the same species. I have considered these morphological differences to be interspecific rather than sexual dimorphism and it is hereby described as a new species. This conclusion is supported by the marked morphological differences. For example, the extant sole nicagin genus
<taxonomicName id="13CE3BEABC6285DD9D58EC56DFF70AF1" authorityName="Leconte" authorityYear="1861" class="Insecta" family="Lucanidae" genus="Nicagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nicagus" order="Coleoptera" pageId="0" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="A824A78EBD6B20DC6BDD85B54A9F6EBD" italics="true" pageId="0" pageNumber="211">Nicagus</emphasis>
</taxonomicName>
has generally similar body shapes in both sexes (
<bibRefCitation id="DACCD80D4208352E4FEDCEE01A038623" DOI="https://doi.org/10.11646/zootaxa.1050.5" author="Paulsen, MJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="211" pagination="45 - 60" refId="B62" refString="Paulsen, MJ, Smith, AD, 2005. A new species of stag beetle from sand dunes in west Texas, and a synopsis of the genus Nicagus (Coleoptera: Lucanidae: Aesalinae: Nicagini). Zootaxa 1050 (1): 45 - 60, DOI: https://doi.org/10.11646/zootaxa.1050.5" title="A new species of stag beetle from sand dunes in west Texas, and a synopsis of the genus Nicagus (Coleoptera: Lucanidae: Aesalinae: Nicagini)." url="https://doi.org/10.11646/zootaxa.1050.5" volume="1050" year="2005">Paulsen and Smith 2005</bibRefCitation>
), unlike the great differences seen in the fossil presented here and
<taxonomicName id="8CDBAD60976030230BB7F1D84E80A02A" lsidName="P. tani" pageId="0" pageNumber="211" rank="species" species="tani">
<emphasis id="2702A80738C42EBAF2F3E537CC7484FC" italics="true" pageId="0" pageNumber="211">P. tani</emphasis>
</taxonomicName>
. Similarly, the body surface of
<taxonomicName id="A6E581CDEA85F4F7F2C3A9AAE3AA235F" lsidName="P. mandibularis" pageId="0" pageNumber="211" rank="species" species="mandibularis">
<emphasis id="CCB9DA3E660064BD719DE56E6A4A4897" italics="true" pageId="0" pageNumber="211">P. mandibularis</emphasis>
</taxonomicName>
sp. nov. lacks short distinct scales, which should be considered an interspecific difference rather than a sexual difference.
</paragraph>
</subSubSection>
</treatment>
</document>
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