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<document id="2F61295D999EFC5BA99A96642D23EE70" ID-DOI="10.3897/zookeys.1207.117225" ID-ZooBank="94F18819-5AF5-4100-AB35-AA3C3976EE80" ID-publisher-id="117225" URI-arpha="255F6420-5705-5F50-9167-509E6C76FA1C" URI-zoobank="http://zoobank.org/94F18819-5AF5-4100-AB35-AA3C3976EE80" article-type="research-article" checkinTime="1721676390257" checkinUser="pensoft" docAuthor="Angus, Robert B., Maté, Jason F., Angus, Elizabeth M. &amp; Král, David" docDate="2024" docId="C85001FB61315D4FBAF4B413BEB88D37" docLanguage="en" docName="ZooKeys 1207: 205-299" docOrigin="ZooKeys 1207" docSource="https://zookeys.pensoft.net/article/117225/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.generic" docStyleName="PensoftTaxPub.0000.journal_article.generic" docTitle="Aphodius (Liothorax) bellumgerens Angus, Maté, Angus &amp; Král, 2024, sp. nov." docType="treatment" docUuid="892D4998-D52B-4B6B-BF0D-ADA21A086F2F" docUuidSource="ZooBank" docVersion="1" lastPageNumber="299" masterDocId="255F642057055F509167509E6C76FA1C" masterDocTitle="Towards a revision of the Palaearctic species of Aphodius Hellwig, 1798, subgenus Liothorax Motschulsky, 1860 (Coleoptera, Scarabaeidae, Aphodiinae)" masterLastPageNumber="299" masterPageNumber="205" pageNumber="205" updateTime="1721676524171" updateUser="pensoft">
<mods:mods id="3F302F3106B273976CEB893A4DB4C20A" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="447FEFCD2B415F3E278EE778C97F213D">
<mods:title id="F4A2A02A334836E609A2AF78E91EB3E0">Towards a revision of the Palaearctic species of Aphodius Hellwig, 1798, subgenus Liothorax Motschulsky, 1860 (Coleoptera, Scarabaeidae, Aphodiinae)</mods:title>
</mods:titleInfo>
<mods:name id="6ADD9E8F51B2EEF5FF6B234B8DF90C75" type="personal">
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<mods:namePart id="6F04022DA7C3B89C349776A29232F058">Angus, Robert B.</mods:namePart>
<mods:nameIdentifier id="6F7E2D204628165B6B6800C2C20A542F" type="ORCID">0000-0002-3860-5617</mods:nameIdentifier>
<mods:affiliation id="A5DCCACA9C5E6D73A92E93D59E17A6EB">Department of Life Sciences (Insects), The Natural History Museum, Cromwell Road, London SW 7 5 BD, UK</mods:affiliation>
</mods:name>
<mods:name id="59E4A63255FB7475C59CA039B7A6C06E" type="personal">
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<mods:namePart id="0CA40B3DB84B2CC36916965CADBF77B4">Maté, Jason F.</mods:namePart>
<mods:nameIdentifier id="6A492F32CD2A9548F039950B903E5CD8" type="ORCID">0000-0002-2505-7113</mods:nameIdentifier>
<mods:affiliation id="297E040ADF3E1C7F21233873771DE61C">c / Henares 16, Velilla de San Antonio, Madrid, 28891, Spain</mods:affiliation>
</mods:name>
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<mods:namePart id="25AD3B789AC53801C0C45F3CF413C05F">Angus, Elizabeth M.</mods:namePart>
<mods:affiliation id="01AA87FD7606BDA754A8E65D0AF0D259">Biomedical Imaging Unit, Level B South Block, Mail point 12, General Hospital, Southampton SO 16 6 YD, UK</mods:affiliation>
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<mods:namePart id="B2DE85FF16FA1C09DE8F744210D24D97">Král, David</mods:namePart>
<mods:nameIdentifier id="80A98E96C6BA9D20C4883DCCF75D172A" type="ORCID">0000-0002-7930-1927</mods:nameIdentifier>
<mods:affiliation id="725AFCBF1EBE8EDA37873D8D32A42B5D">Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ- 128 00, Praha 2, Czech Republic</mods:affiliation>
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<mods:date id="A76BE0509A10E9B67B834DD8EE259902">2024</mods:date>
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<subSubSection id="84D244F69DF0DC53E9FF53571DBB15F6" type="nomenclature">
<paragraph id="39A6D143C3DD16D164EA2441F0FE701C">
<taxonomicName id="8EE300AE0D0BD973E6EFD375C5F10971" authorityName="Angus &amp; Maté &amp; Angus &amp; Král" authorityYear="2024" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="bellumgerens" status="sp. nov." subGenus="Liothorax">
<emphasis id="BAEC7452CCB7C9502443BFAEB38E248A" italics="true">Aphodius</emphasis>
(
<emphasis id="09427E5CCB2719AD2BF681BAE03120C2" italics="true">Liothorax</emphasis>
)
<emphasis id="0A4D5608D00F606B0843C56ACEA5C2F8" italics="true">bellumgerens</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F9BCE3AE76ACDBDC71D150CB7F60E582" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="432C79F235C6A4E17ABC227586F484B2" type="description">
<paragraph id="A8C8052DC7EEDD928E56729365352CA7">
<figureCitation id="BEAAA872B3001CD405892E66515F61AB" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. a – g whole beetles a A. niger, SV, Tullgarn b A. muscorum, holotype HU, Vörös tó c A. felix sp. nov., holotype, IT, Campo Felice d A. bameuli sp. nov., holotype, Corsica, by Lac de Melo e A. krelli sp. nov., holotype, IT, Sardinia f A. bellumgerens sp. nov., holotype, IT, Sicily, Piano Battaglia g A. wilsonae, holotype, SP, Provincia de Burgos, Balneario de Corconte. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure4" httpUri="https://binary.pensoft.net/fig/1097462">Figs 4 f</figureCitation>
,
<figureCitation id="A7A14D3B0552400EEFFD72CC3DEEC590" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. a – l A. niger group, heads and pronota a, b A. isikdagensis, paratypes, TR Ḉamildere a ♂ b ♀ b ’ detail of right lateral part of frons and gena, different orientation b ” detail of right posterior of pronotum, different orientation c, d A. alberti sp. nov., TR, Rize, Ovitdagi c holotype d ♀ paratype e A. niger, GB, Hampshire, New Forest f A. muscorum, HU, Hortobágyi g A. bameuli sp. nov., holotype, FR, Corsica, by Lac de Melo h A. krelli sp. nov., holotype, IT, Sardinia i A. felix sp. nov., holotype, IT, Campo Felice j A. bellumgerens sp. nov., holotype, IT, Sicily, Piano Battaglia k, l A. wilsonae, SP k holotype, Provincia de Burgos, Balneario de Corconte l paratype, Provincia de Madrid, Manzanares el Real. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure6" httpUri="https://binary.pensoft.net/fig/1097465">6 j</figureCitation>
,
<figureCitation id="6F9AC1E870B30A360DDBC72C3F4FD5AD" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. a – i epipharynxes a – c A. niger a SV, Tullgarn b, c GB, Hampshire, New Forest d A. niger?, CZ, Hradec Králové e A. muscorum, HU, Hortobagyi f, g A. bellumgerens sp. nov., IT Piano Battaglia, paratypes h, i A. wilsonae paratypes, SP h El Vellón i Manzanares el Real. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure9" httpUri="https://binary.pensoft.net/fig/1097468">9 f, g</figureCitation>
,
<figureCitation id="C70C0CB392C8B4AE417390EE0BB603A1" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Aphodius niger group, pronotal bases 2 a A. bameuli sp. nov., paratype, FR, Corsica b A. krelli sp. nov., paratype, IT, Sardinia c, d A. wilsonae, paratypes, SP, Manzanares el Real e A. isikdagensis, paratype, TR, Ḉamildere." figureDoi="10.3897/zookeys.1207.117225.figure16" httpUri="https://binary.pensoft.net/fig/1097478">16 d</figureCitation>
,
<figureCitation id="1B9FB4662EF7FB09058596A6F4DFC516" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Elytral sculpture of interstices 2 or 3 a – z stacked photographs a A. plagiatus plagiatus, CZ, Moravia b, c A. p. sinoplagiatus ssp. nov., paratypes, CH b Gangca, Qinghai c Gansu d A. rutilipennis (A. ressli, holotype), TR, Hatay e A. chellala sp. nov., holotype f A. rusakovi, paratype, RU, Orenburg g, h A. discoides g neotype, IS, Kuneitra h TR, Kizildag i A. discoides, black specimen, TR, Karakurt j A. kraatzi, SK k A. isikdagensis, paratype, TR, Ḉamildere l, m A. niger, Sweden, Södermanland, Tullgarn n – q A. muscorum, HU n holotype, Vörös-tó o Hortobágyi p, q Kisjúszállás r, s A. niger? r CZ, Hradec Králové s CZ, České Budějovice t A. muscorum? SK, Bol u A. bellumgerens sp. nov., paratype, IT, Sicily v A. felix sp. nov., paratype, IT w A. wilsonae, paratype, SP x A. bameuli sp. nov., paratype, FR, Corsica y A. krelli sp. nov., paratype, IT, Sardinia z A. alberti sp. nov., paratype, TR, Rize, Ovitdag. a’ – c ’ A. plagiatus, at higher magnification a ’, c ’ stacked photographs a ”, c ” SEM images a ’, a ” A. p. sinoplagiatus ssp. nov., CH, Gansu c ”, c ’ A. p. plagiatus, CZ, Moravia. Scale bars: 0.1 mm." figureDoi="10.3897/zookeys.1207.117225.figure17" httpUri="https://binary.pensoft.net/fig/1097480">17 u</figureCitation>
,
<figureCitation id="A027A9067D1DCC980A2EC863C21473F6" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Legs a – h A. plagiatus group, stacked photographs mid tarsi and apical portions of tibiae a A. plagiatus sinoplagiatus ssp. nov., paratype, CH, Gangca, Qinghai b A. rodrigoi sp. nov. holotype, SP, Aranjuez c A. rutilipennis (A. ressli, holotype), TR, Hatay d A. chellala sp. nov., holotype, AG, Chellala e A. discoides, neotype, IS, Keneitra f, g A. kraatzi f SV g RU, Karasuk, W. Siberia h A. rusakovi, paratype ♂, RU, Orenburg i – t A. niger group, mid tarsi, and tibial apices i – o SEM images p – t stacked photographs i A. niger, Sweden, Södermanland, Tullgarn j A. muscorum, HU, Hortobágyi k, l A. bameuli sp. nov., paratypes, FR, Corsica m, n A. krelli sp. nov., paratypes, IT, Sardinia o A. wilsonae, paratype SP p, q A. felix sp. nov, IT, Abruzzo, Campo Felice p paratype q holotype r A. bellumgerens sp. nov., paratype, IT, Sicily s A. isikdagensis, paratype, TR, Ḉamildere t A. alberti sp. nov., holotype, TR, Rize, Ovitdag u, v fore tibiae and tarsi of ♂♂ to show the spur. u A. bameuli sp. nov., paratype, FR, Corsica v A. rusakovi, paratype, RU, Orenburg w, x apex of hind tibia to show the short spines w A. muscorum, HU x A. wilsonae, paratype, SP. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure18" httpUri="https://binary.pensoft.net/fig/1097482">18 r</figureCitation>
,
<figureCitation id="6B14B33BBF8D533C88975BBEB18B5FA5" captionStart="Figure 21" captionStartId="F21" captionText="Figure 21. Metaventrites, A. niger group a – d A. niger a, c, d SV, Södermanland, Tullgarn b GB, Hampshire, New Forest a, b ♂ c, d ♀ e, g – j A. muscorum e holotype ♀, HU, Vörös-tó f A. felix sp. nov., paratype ♂, IT, Campo Felice g ♂, HU, Hortobágyi h, i ♀♀, HU, Kisujszallas j ♀, HU, Josvafo k A. felix sp. nov. paratype ♀, IT, Campo Felice l, m A. bellumgerens sp. nov., paratypes, IT, Sicily, Piano Battaglia l ♂ m ♀ n, o A. alberti sp. nov., paratypes, TR, Rize, Ovitdag n ♂, o ♀ p – r A. bameuli sp. nov., paratypes, FR, Corsica p ♂ q, r ♀ s, t A. krelli sp. nov., paratypes, IT, Sardinia s ♂ t ♀ u – w A. wilsonae, paratypes u, v SP, Manzanares el Real u ♂ v ♀ w ♀, SP, El Vellon x, y A. isikdagensis, paratypes, TR, Ḉamildere x ♂ y ♀. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure21" httpUri="https://binary.pensoft.net/fig/1097488">21 l, m, p – t</figureCitation>
,
<figureCitation id="E7AB88BC26D387045C2499543D6F2E9F" captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. A. niger group aedeagi, cleared preparations mounted in DMHF on slides a – g, m – o dorsoventral view h – l, p right lateral view a A. niger, SV, Södermanland, Tullgarn b A. bameuli sp. nov., paratype c A. isikdagensis, paratype, TR, Ḉamildere d A. krelli sp. nov., paratype, IT, Sardinia e – f A. alberti sp. nov., TR, Rize, Ovitdag e paratype, after squashing f the same specimen, before squashing g, h, j A. alberti sp. nov., different paratypes i A. alberti sp. nov., paratype, paramere k A. isikdagensis?, TR, Artvin l A. alberti?, Armenia m A. bellumgerens sp. nov., paratype, IT, Sicily, Piano Battaglia n A. bellumgerens, paratype IT, Sicily, Nebrodi o A. wilsonae, paratype, SP, Manzanares el Real p A. muscorum, HU, Hortobági. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure24" httpUri="https://binary.pensoft.net/fig/1097494">
<quantity id="73CAB3EEF85AA44615090AC48C854EBE" metricMagnitude="1" metricUnit="m" metricValue="2.4" unit="m" value="24.0">24 m</quantity>
, n
</figureCitation>
,
<figureCitation id="33836FAE71639DC2604285C5DAC8FB59" captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Aedeagal encasements (9 th abdominal sternite) a A. niger, GB, Hampshire, New Forest b A. muscorum, HU, Hortobagyi c A. felix sp. nov., IT, Campo Felice d A. alberti sp. nov., TU, Ovit Dağı e A. bellumgerens sp. nov., IT, Sicily, Piano Battaglia f A. bameuli sp. nov., FR, Corsica, Haute-Corse g A. wilsonae, SP, Cantabria, Corconte h A. kraatzi, RU, Chita Reg. i A. plagiatus, UK, Norfolk, Hunstanton j A. plagiatus, CN, Beijing k A. rodrigoi sp. nov., SP, Madrid, Aranjuez l A. chellala sp. nov., AG, Chellala, paratype m A. rutilipennis sp. nov., CY, Konakli, paratype n A. discoides sp. nov., paratype. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure25" httpUri="https://binary.pensoft.net/fig/1097496">25 e</figureCitation>
,
<figureCitation id="2B127843399A765C9374240239BD5AF9" captionStart="Figure 26" captionStartId="F26" captionText="Figure 26. Spermathecae a – e A. plagiatus plagiatus a CZ, Dobre Pole b GB, Norfolk c KZ, Karaganda d, e RU, Transbaikal, Ulan Ude f, g A. p. sinoplagiatus, CN f Gansu g holotype, CN, Qinghai h, i A. discoides, TR h Kizil dag, i Muʂ j A. rutilipennis CY, Limassol distr k A. rutilipennis?, GR Thesaloniki l A. chellala, paratype, AG, Chellala m, n A. rusakovi m paratype, RU, wet but not covered n KZ, Syr-Darja o – s A. kraatzi o, p SK q RU, Astrakhan r RU, Lenkoran s AF, Kabul t A. niger, GB, Hampshire, New Forest u A. muscorum HU, Hortobágyi v A felix, paratype, IT w, x A. bameuli, paratypes, FR y A. krelli, sp. nov., IT z A. bellumgerens sp. nov., paratype, IT a ’, b ’ A. alberti sp. nov., TR paratypes c ’ A. isikdagensis, paratype, TR Ḉamildere d ’, e ’ A. wilsonae, SP d ’ paratype, SP, Madrid e ’ SP, Soria. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure26" httpUri="https://binary.pensoft.net/fig/1097499">26 z</figureCitation>
,
<figureCitation id="A76B37E2FD63D2AC3781344A13F4CE4A" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Mitotic chromosomes arranged as karyotypes a – e A. felix sp. nov. IT, Abruzzo. Campo Felice a, b holotype ♂ a plain b the same nucleus C-banded c holotype ♂ C-banded, autosome 8 polymorphic, 9 subacrocentric d paratype ♂, C-banded, autosomes 8 and 9 metacentric e paratype ♀ f – i A. bellumgerens sp. nov., paratypes, IT, Sicily f, g A. bellumgerens sp. nov., ♂, paratype f plain g C-banded h, i A. bellumgerens sp. nov., ♀, paratype h plain i C-banded. Scale bar: 5 μm" figureDoi="10.3897/zookeys.1207.117225.figure32" httpUri="https://binary.pensoft.net/fig/1097519">32 f – i</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EMWBM" type="type material">
<paragraph id="DC70A84FFFA3D90FDCEBDE2FF7B9B4FB">
<heading id="496C2D6C985578C780F00D8C1E322E48" reason="title">Type material examined.</heading>
</paragraph>
<paragraph id="70CCFBA9EF93536A2C4E785A1E015A0C">
<materialsCitation id="E08A46708E92AD7AC95CD9A636396189" collectingDate="2013-05-01" collectingDateMax="2013-05-08" collectingDateMin="2013-05-01" collectionCode="NHMUK" county="Parco delle Madonie. Piano Battaglia. At" location="Length" municipality="Chromosome" specimenCount="1" specimenCount-male="1" stateProvince="Sicilia" typeStatus="holotype">
<emphasis id="7BBA742FD57259764C518BADF4BB8540" bold="true" italics="true">
<typeStatus id="9417278462E4CE370173600CBB0F21B2" type="holotype">Holotype</typeStatus>
</emphasis>
<specimenCount id="2B3DE2CC4304DC2FE0FFBAB8468AB3F7" count="1" type="male">♂</specimenCount>
:
<collectingRegion id="01DD026644A4FFE78F22F415B31C2378" country="Italy" name="Sicilia">Sicily</collectingRegion>
.
<collectingCounty id="1B8E128862A17496B63BDAF87182F969">Parco delle Madonie. Piano Battaglia. At</collectingCounty>
edges of pool.
<date id="46188ABD1EF5A5F0BFCD6666D0499012" value="2013-05-01">
<collectingDate id="571F3AE25CD2684810D771C7AD7459FE" value="2013-05-01">1. v. 2013</collectingDate>
</date>
. R. B. &amp; E. M.
<collectingRegion id="5B794D8234B6F2214A985F549003F784" country="United Kingdom" name="Angus">Angus</collectingRegion>
.
<collectingMunicipality id="8C437AE6C838957C63FB9FFF89010CA4">Chromosome</collectingMunicipality>
prep. 7:
<date id="CC06E0C0EBD4D1531B5E6DAF50E69AEC" value="2013-05-08">
<collectingDate id="924F938686BF0C85AEE2C6AC4CB786C2" value="2013-05-08">8. v. 2013</collectingDate>
</date>
(
<named-content id="E48C89D6012DF7B70192DAFD4805E938" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="52115F1FA3E23477FACAF2BA38476719" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" type="Museum">NHMUK</collectionCode>
</named-content>
).
<location id="681D15CB41CD59616C2A1F5DF41AD19B" LSID="urn:lsid:plazi:treatment:C85001FB61315D4FBAF4B413BEB88D37:681D15CB41CD59616C2A1F5DF41AD19B" county="Parco delle Madonie. Piano Battaglia. At" municipality="Chromosome" name="Length" stateProvince="Sicilia">Length</location>
<quantity id="DEFF1337DAEF1CD90C21DCBDDF6BB05F" metricMagnitude="-3" metricUnit="m" metricValue="3.9" unit="mm" value="3.9">3.9 mm</quantity>
, width
<quantity id="6EC832E8CE050CAE018E5C366B94ED4A" metricMagnitude="-3" metricUnit="m" metricValue="1.8" unit="mm" value="1.8">1.8 mm</quantity>
</materialsCitation>
.
<emphasis id="81685B0D9AB6FEF7EAF71D1D6DBFE926" bold="true" italics="true">
<typeStatus id="8049777A5218059105E34EC974AE81BB" type="paratype">Paratypes</typeStatus>
</emphasis>
:
<specimenCount id="C0487373FD2E37AD5212B02450C4A1A7" count="7" type="male">7 ♂♂</specimenCount>
,
<specimenCount id="C5A0D6CC060F460A92CAAB290FDBBB83" count="8" type="female">8 ♀♀</specimenCount>
, same data as
<typeStatus id="BD7716BB59BB840C23D7E65195730FE4" type="holotype">holotype</typeStatus>
. Chromosome preparations 1–8,
<date id="2A208068EB06E98F227E24D0810A5965" value="2012-05-07">7. v. 2012</date>
, 1–6 and 8,
<date id="64124BF3169D1FA890467BE3BA8BE4F8" value="2012-05-08">8. v. 2012</date>
(
<named-content id="9A171C189A0769DF28EA237D5E50F087" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="623238A2C08E8A4C713C63BD7A38A16A" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" type="Museum">NHMUK</collectionCode>
</named-content>
).
<specimenCount id="8FFBDDAB3FA2B56F3F5128A3FDA78277" count="7" type="male">7 ♂♂</specimenCount>
,
<specimenCount id="23D5C79232C389099B04C516041E1541" count="5" type="female">5 ♀♀</specimenCount>
, same data as
<typeStatus id="E9411F74509E9A4D9AB031FC3CB64623" type="holotype">holotype</typeStatus>
but
<date id="859C907D934A4B0F23117F7E7A4B2544" value="2018-04-23">23. iv. 2018</date>
(
<named-content id="B5A3B5FF71199EE8A199D4B0A0F484C1" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="D2EB0392DA2D314E1650D7B03C114650" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" type="Museum">NHMUK</collectionCode>
</named-content>
),
<specimenCount id="64730093F714D3AD5FDEBDE6DCB5C581" count="1" type="male">1 ♂</specimenCount>
,
<specimenCount id="E26D1307F6897E6680783AD8B98BA7FE" count="1" type="female">1 ♀</specimenCount>
, same data (
<named-content id="A7C9CE4EB99ED6147D01D6FA1464990F" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-prague-0" xlink_title="National Museum Prague">
<collectionCode id="04BE62B58EA8269DE46C8B385587D0AA" country="Czech Republic" httpUri="http://grbio.org/cool/fx6k-enrh" name="National Museum (Prague)" type="">NMP</collectionCode>
</named-content>
);
<specimenCount id="46F9D22015FF0A9F4AAC6732A7314D0B" count="12" type="male">12 ♂♂</specimenCount>
,
<specimenCount id="4226C80FCAEC47830E4EA1B7C73FDA4E" count="6" type="female">6 ♀♀</specimenCount>
,
<collectingRegion id="55447F5A86BF2163A178A6081FBCF8D9" country="Italy" name="Sicilia">Sicily</collectingRegion>
, Nebrodi, Monte Soro,
<quantity id="AA6A7F48C99EB80FC058A5413654370D" metricMagnitude="3" metricUnit="m" metricValue="1.6" unit="m" value="1600.0">1600 m</quantity>
a. s. l.
<date id="F62144BF21CC3C2BD027C9D9B79FAE37" value="2018-04-25">25. iv. 2018</date>
. R. B. &amp; E. M.
<collectingRegion id="23AC9705924D58A93F72DA2EEF8A3999" country="United Kingdom" name="Angus">Angus</collectingRegion>
. Chromosome preparations 1–4,
<date id="D56C5C7400272848209AB743728DDCDB" value="2018-04-27">27. iv. 2018</date>
, 1–5,
<date id="67FCA8772D74BFBDA7F769DE69BA7EC3" value="2018-05-01">1. v. 2018</date>
, 1–5,
<date id="D83B1E38A07EAA0DF59C959B66F1FA3B" value="2018-05-02">2. v. 2018</date>
, 1–4,
<date id="C3DE9F45D1451D2F9531C5D7A5084403" value="2015-05-10">10. v. 2015</date>
(
<named-content id="3A921F931BE84A81B7BDC0A378D0EFEB" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="1C0A202894EFF285B54BAAE912BB5970" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" type="Museum">NHMUK</collectionCode>
</named-content>
).
<specimenCount id="9A035F10A4BEB4F044A2AC6D1C7A4D43" count="1" type="male">1 ♂</specimenCount>
,
<collectingRegion id="89E894C2044F7C90D94C5B78B6DC8D70" country="Italy" name="Sicilia">Sicily</collectingRegion>
, Nebrodi, leg F. Krell (
<named-content id="A00E9976C42246191B49B27646CD8650" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/denver-museum-nature-science" xlink_title="Denver Museum of Nature and Science">
<collectionCode id="9AB1F7287C27AF08B791B8DCA32447D0" country="USA" httpUri="http://grbio.org/cool/mmzs-jqix" name="Denver Museum of Nature and Science" type="Museum">DMNS</collectionCode>
</named-content>
). 3, data as
<typeStatus id="8240067360602526F20EC4F7499B0519" type="holotype">holotype</typeStatus>
(sequenced) (
<abbrev id="ABBRID0ESXBM" xlink_title="Jason Maté Collection">
<collectionCode id="70AEFB2CD8332FA9BEFE3D87E3235627">JFMC</collectionCode>
</abbrev>
); 1
<collectingRegion id="6DA1F8F89871ECE122EB99E0B50EE236" country="Italy" name="Sicilia">SICILY</collectingRegion>
, Nebrodi, Monte Soro,
<quantity id="34FF4F014225F6EEAE7571C71D909CB9" metricMagnitude="3" metricUnit="m" metricValue="1.6" unit="m" value="1600.0">1600 m</quantity>
a. s. l.
<date id="40228545EE4F14C4FCD6179A5D2BA33A" value="2018-04-25">25. iv. 2018</date>
. R. B. &amp; E. M.
<collectingRegion id="D44F3734170A84A490CB1515EC4F2B49" country="United Kingdom" name="Angus">Angus</collectingRegion>
. Chromosome preparation 4,
<date id="99044DDB4EE992A1915A9F8EBEB2710B" value="2018-04-27">27. iv. 2018</date>
(Sequenced) (
<abbrev id="ABBRID0EWXBM" xlink_title="Jason Maté Collection">
<collectionCode id="EF67E4C7C50555A536890FB0E1CD827E">JFMC</collectionCode>
</abbrev>
).
</paragraph>
<paragraph id="0463D7A18711F457D5BFDC541F013EA8">
<specimenCount id="3F48E620E049E281E5571A02484798B0" count="3" type="male">3 ♂♂</specimenCount>
,
<specimenCount id="B881718B45088A27DF7D202557E8DBBB" count="4" type="female">4 ♀♀</specimenCount>
, 37 unsexed:
<collectingCountry id="6A18A4C9B7666E66A5C94B1B87C20B1C" name="Italy">ITALY</collectingCountry>
,
<collectingRegion id="0C366A389AC48A636CEFF17EF62CA35A" country="Italy" name="Sicilia">Sicily</collectingRegion>
, Monte Soro, Tumpelufer, welkes Laub.
<quantity id="ED1623708802CC9A01D3E00C8E7770CC" metricMagnitude="3" metricUnit="m" metricValue="1.8" unit="m" value="1800.0">1800 m</quantity>
NN
<date id="392ACC3D39B3A75796159F388E776E24" value="1993-05-30">30. 5. 1993</date>
leg. U Schaffrath (
<abbrev id="ABBRID0E3XBM" xlink_title="Axel Bellmann Collection">
<collectionCode id="7A562CFDCE1821402E674FDBE508E443">ABC</collectionCode>
</abbrev>
&amp;
<abbrev id="ABBRID0EAYBM" xlink_title="Jason Maté Collection">
<collectionCode id="68B60527ED91AE86A2820564C84265F1">JFMC</collectionCode>
</abbrev>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EEYBM" type="Differential diagnosis">
<paragraph id="F15D3B1F04C777B3411CBD27E1B5F4C7">
<heading id="C08BB7A2AEA4A4EC0FAD8931C8253F97" reason="title">Differential diagnosis.</heading>
</paragraph>
<paragraph id="3545DA4D208249D6D480102D661832CB">
Most similar to
<taxonomicName id="CC064332337EC58193B8BF9F8E67246A" authorityName="Angus &amp; Maté &amp; Angus &amp; Král" authorityYear="2024" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="felix">
<emphasis id="89861B477CD3E0CD5D5A63A4475FB9FC" italics="true">A. felix</emphasis>
</taxonomicName>
, from which it is shown to be different because of its clearly larger X chromosome (Fig.
<figureCitation id="A9006F3EC4680B49ABEF5B87AA78FBEF" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Mitotic chromosomes arranged as karyotypes a – e A. felix sp. nov. IT, Abruzzo. Campo Felice a, b holotype ♂ a plain b the same nucleus C-banded c holotype ♂ C-banded, autosome 8 polymorphic, 9 subacrocentric d paratype ♂, C-banded, autosomes 8 and 9 metacentric e paratype ♀ f – i A. bellumgerens sp. nov., paratypes, IT, Sicily f, g A. bellumgerens sp. nov., ♂, paratype f plain g C-banded h, i A. bellumgerens sp. nov., ♀, paratype h plain i C-banded. Scale bar: 5 μm" figureDoi="10.3897/zookeys.1207.117225.figure32" httpUri="https://binary.pensoft.net/fig/1097519">32 f – i, a – e</figureCitation>
). The slightly smaller aedeagus, with the endophallic tooth-field clearly shorter than the parameres (Figs
<figureCitation id="92F0F55921C42C3B65F13F44CEAF0B71" captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. A. niger group, aedeagi with inflated endophalli, stacked photographs and SEM images a – e, g – w right lateral view f viewed from above a – c A. niger, SV, Södermanland, Tullgarn a stacked photograph b the same aedeagus, SEM image c detail of endophallic teeth d – f A. niger, GB, Hampshire, New Forest d stacked photograph e, f SEM images g – i A. wilsonae, SP, Areños g stacked photograph h SEM image, i detail of endophallic teeth j – l A. bameuli sp. nov., holotype j stacked photograph k SEM image l detail of endophallic teeth m – o A. krelli sp. nov., holotype m stacked photograph n SEM image o detail of endophallic teeth p – r A. bellumgerens sp. nov., paratype p stacked photograph q SEM image r detail of endophallic teeth s, t A. bellumgerens sp. nov., holotype s stacked photograph t SEM image of endophallus u A. muscorum, HU, Hortobágyi, stacked photograph v, w A. felix sp. nov., holotype, SEM images v aedeagus w detail of endophallic teeth. Black Scale bar: 0.5 mm for aedeagi and endophallus, white SEM scale on black background: 50 μm for details of endophallic teeth." figureDoi="10.3897/zookeys.1207.117225.figure23" httpUri="https://binary.pensoft.net/fig/1097492">23 p – t</figureCitation>
,
<figureCitation id="33C84B46B54808690F1E0C08D656ACEC" captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. A. niger group aedeagi, cleared preparations mounted in DMHF on slides a – g, m – o dorsoventral view h – l, p right lateral view a A. niger, SV, Södermanland, Tullgarn b A. bameuli sp. nov., paratype c A. isikdagensis, paratype, TR, Ḉamildere d A. krelli sp. nov., paratype, IT, Sardinia e – f A. alberti sp. nov., TR, Rize, Ovitdag e paratype, after squashing f the same specimen, before squashing g, h, j A. alberti sp. nov., different paratypes i A. alberti sp. nov., paratype, paramere k A. isikdagensis?, TR, Artvin l A. alberti?, Armenia m A. bellumgerens sp. nov., paratype, IT, Sicily, Piano Battaglia n A. bellumgerens, paratype IT, Sicily, Nebrodi o A. wilsonae, paratype, SP, Manzanares el Real p A. muscorum, HU, Hortobági. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure24" httpUri="https://binary.pensoft.net/fig/1097494">
<quantity id="AB73F06E9394F60659D14E2B7689A896" metricMagnitude="1" metricUnit="m" metricValue="2.4" unit="m" value="24.0">24 m</quantity>
, n
</figureCitation>
) is also distinctive.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EBZBM" type="description">
<paragraph id="844ADD4190500D7C07506242D02A9A1D">
<heading id="FDBA3D7BE3D75F4DAA4BE0EDB7C9D2BA" reason="title">Description.</heading>
</paragraph>
<paragraph id="45B9E7805DF26D0CF4E75CE68264C43B">
General appearance (Fig.
<figureCitation id="4B56E50032E6BAC51059E65472D6226F" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. a – g whole beetles a A. niger, SV, Tullgarn b A. muscorum, holotype HU, Vörös tó c A. felix sp. nov., holotype, IT, Campo Felice d A. bameuli sp. nov., holotype, Corsica, by Lac de Melo e A. krelli sp. nov., holotype, IT, Sardinia f A. bellumgerens sp. nov., holotype, IT, Sicily, Piano Battaglia g A. wilsonae, holotype, SP, Provincia de Burgos, Balneario de Corconte. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure4" httpUri="https://binary.pensoft.net/fig/1097462">4 f</figureCitation>
). Length:
<quantity id="FF1FAF0C76F124A554B3E32775C9EA14" metricMagnitude="-3" metricUnit="m" metricValue="4.0" metricValueMax="4.4" metricValueMin="3.6" unit="mm" value="4.0" valueMax="4.4" valueMin="3.6">3.6–4.4 mm</quantity>
(
<specimenCount id="B47BEB33A3D65B0BD5E3881A0652E9AA" count="1" type="male">♂</specimenCount>
), 4.0–
<quantity id="D8D5BEB12728E3874D30D9730BECA157" metricMagnitude="-3" metricUnit="m" metricValue="4.8" unit="mm" value="4.8">4.8 mm</quantity>
(
<specimenCount id="7F4086D6A037B61A998AD291ACF503E6" count="1" type="female">♀</specimenCount>
), width:
<quantity id="898B344F191282D0B8362D9C479A94B1" metricMagnitude="-3" metricUnit="m" metricValue="1.7999999999999998" metricValueMax="1.95" metricValueMin="1.65" unit="mm" value="1.7999999999999998" valueMax="1.95" valueMin="1.65">1.65–1.95 mm</quantity>
(
<specimenCount id="178BC854EB7F4B15E4EC285592D45837" count="1" type="male">♂</specimenCount>
), 1.7–2.0 mm (
<specimenCount id="CDEC9303BC04992532B5EB90B7A48799" count="1" type="female">♀</specimenCount>
). Glossy black, apical segments plus claws of tarsi dark brown.
</paragraph>
<paragraph id="57D2BD27DEA0B62E11CDBAE2E8C182FA">
Head with no trace of reticulation but with sparse double punctures, the punctures separated by ca 2 × the diameter of the larger punctures. Frontoclypeal suture indistinct but present, without tubercles but slightly raised at ends of median section, before turning to run to the anterior margins of the genae. Anterior margin of clypeus emarginated medially, rounded at sides of the emargination. Antennae and palpi black. Epipharynx (Fig.
<figureCitation id="0F8AEE15B70981F2A42F2294B0DA6D32" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. a – i epipharynxes a – c A. niger a SV, Tullgarn b, c GB, Hampshire, New Forest d A. niger?, CZ, Hradec Králové e A. muscorum, HU, Hortobagyi f, g A. bellumgerens sp. nov., IT Piano Battaglia, paratypes h, i A. wilsonae paratypes, SP h El Vellón i Manzanares el Real. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure9" httpUri="https://binary.pensoft.net/fig/1097468">9 f, g</figureCitation>
) with anterior margin of clithra emarginate each side of projecting median tylus, with a few fine acropariae. Chaetopedia 4 or 5 each side, stout. Chaetopariae forming a single line each side, closely set, stout. Apophobae forming a narrow band of fine setae outside the chaetopariae, prophobae in a narrow field each side of the mesoepitorma. Head and pronotum (Fig.
<figureCitation id="BA9FF2121FBCD1E30AE7768A0049C2BC" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. a – l A. niger group, heads and pronota a, b A. isikdagensis, paratypes, TR Ḉamildere a ♂ b ♀ b ’ detail of right lateral part of frons and gena, different orientation b ” detail of right posterior of pronotum, different orientation c, d A. alberti sp. nov., TR, Rize, Ovitdagi c holotype d ♀ paratype e A. niger, GB, Hampshire, New Forest f A. muscorum, HU, Hortobágyi g A. bameuli sp. nov., holotype, FR, Corsica, by Lac de Melo h A. krelli sp. nov., holotype, IT, Sardinia i A. felix sp. nov., holotype, IT, Campo Felice j A. bellumgerens sp. nov., holotype, IT, Sicily, Piano Battaglia k, l A. wilsonae, SP k holotype, Provincia de Burgos, Balneario de Corconte l paratype, Provincia de Madrid, Manzanares el Real. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure6" httpUri="https://binary.pensoft.net/fig/1097465">6 j</figureCitation>
): pronotum: surface either smooth and glossy as head or slightly dull, with double punctation, size of the large punctures stronger than on head, but variable, these punctures separated by ca 2 × their diameter. Small punctures tiny dots. Mid-line impunctate or with very fine punctures. Pronotum more or less hemicylindrical, highly arched transversely, in mid-line weakly arched longitudinally. Sides of pronotum bulging outwards in basal 1 / 3, so that the basal 1 / 3 of the lateral margins is not visible from above. Lateral margins completely bordered, this border extending along lateral 1 / 3 of basal margin as a very fine impressed line. Median 1 / 3 of basal margin without an impressed border, as in the
<taxonomicName id="7F45E86FF7175FB8A08366D5172918C7" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="wilsonae">
<emphasis id="B7BD64FE2ADED0CF0CFB754462DC24EA" italics="true">A. wilsonae</emphasis>
</taxonomicName>
shown in Fig.
<figureCitation id="719548E24193C9F265C14E7C8AAD30DE" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Aphodius niger group, pronotal bases 2 a A. bameuli sp. nov., paratype, FR, Corsica b A. krelli sp. nov., paratype, IT, Sardinia c, d A. wilsonae, paratypes, SP, Manzanares el Real e A. isikdagensis, paratype, TR, Ḉamildere." figureDoi="10.3897/zookeys.1207.117225.figure16" httpUri="https://binary.pensoft.net/fig/1097478">16 d</figureCitation>
. Scutellum pentagonal, sides parallel in basal 1 / 2, then converging to blunt apical point. Elytra: sides either almost parallel over basal 2 / 3 or more distinctly rounded to widest point 2 / 3 of the way from the base, bluntly rounded to apex. Interstices flat, ca 6 × the width of striae, glossy black, sometimes with very fine reticulation, especially over basal 1 / 4 (Fig.
<figureCitation id="ECCB4DA2FC5E4D747B9CAD7B898AB7FB" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Elytral sculpture of interstices 2 or 3 a – z stacked photographs a A. plagiatus plagiatus, CZ, Moravia b, c A. p. sinoplagiatus ssp. nov., paratypes, CH b Gangca, Qinghai c Gansu d A. rutilipennis (A. ressli, holotype), TR, Hatay e A. chellala sp. nov., holotype f A. rusakovi, paratype, RU, Orenburg g, h A. discoides g neotype, IS, Kuneitra h TR, Kizildag i A. discoides, black specimen, TR, Karakurt j A. kraatzi, SK k A. isikdagensis, paratype, TR, Ḉamildere l, m A. niger, Sweden, Södermanland, Tullgarn n – q A. muscorum, HU n holotype, Vörös-tó o Hortobágyi p, q Kisjúszállás r, s A. niger? r CZ, Hradec Králové s CZ, České Budějovice t A. muscorum? SK, Bol u A. bellumgerens sp. nov., paratype, IT, Sicily v A. felix sp. nov., paratype, IT w A. wilsonae, paratype, SP x A. bameuli sp. nov., paratype, FR, Corsica y A. krelli sp. nov., paratype, IT, Sardinia z A. alberti sp. nov., paratype, TR, Rize, Ovitdag. a’ – c ’ A. plagiatus, at higher magnification a ’, c ’ stacked photographs a ”, c ” SEM images a ’, a ” A. p. sinoplagiatus ssp. nov., CH, Gansu c ”, c ’ A. p. plagiatus, CZ, Moravia. Scale bars: 0.1 mm." figureDoi="10.3897/zookeys.1207.117225.figure17" httpUri="https://binary.pensoft.net/fig/1097480">17 u</figureCitation>
). Sutural interstices weakly raised to suture. Striae narrow, shallow, the strial punctures scarcely impinging on their margins. Median diamond-shaped plate of metaventrite with sparse fairly fine punctures, sometimes flattened in males or with a median depression in females (Fig.
<figureCitation id="9676EA38057E90641CDAF1D194659916" captionStart="Figure 21" captionStartId="F21" captionText="Figure 21. Metaventrites, A. niger group a – d A. niger a, c, d SV, Södermanland, Tullgarn b GB, Hampshire, New Forest a, b ♂ c, d ♀ e, g – j A. muscorum e holotype ♀, HU, Vörös-tó f A. felix sp. nov., paratype ♂, IT, Campo Felice g ♂, HU, Hortobágyi h, i ♀♀, HU, Kisujszallas j ♀, HU, Josvafo k A. felix sp. nov. paratype ♀, IT, Campo Felice l, m A. bellumgerens sp. nov., paratypes, IT, Sicily, Piano Battaglia l ♂ m ♀ n, o A. alberti sp. nov., paratypes, TR, Rize, Ovitdag n ♂, o ♀ p – r A. bameuli sp. nov., paratypes, FR, Corsica p ♂ q, r ♀ s, t A. krelli sp. nov., paratypes, IT, Sardinia s ♂ t ♀ u – w A. wilsonae, paratypes u, v SP, Manzanares el Real u ♂ v ♀ w ♀, SP, El Vellon x, y A. isikdagensis, paratypes, TR, Ḉamildere x ♂ y ♀. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure21" httpUri="https://binary.pensoft.net/fig/1097488">21 l, m</figureCitation>
).
</paragraph>
<paragraph id="F1E89886013AE066F1BED6ED879D6524">
Legs: Long spur of mid tibiae clearly longer than basal segment of tarsi, exceeding it by a distance approximately equal to the width of the basal margin of the second tarsal segment (Fig.
<figureCitation id="CE6C5FB72D21882DBFDACBC9B9339C62" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Legs a – h A. plagiatus group, stacked photographs mid tarsi and apical portions of tibiae a A. plagiatus sinoplagiatus ssp. nov., paratype, CH, Gangca, Qinghai b A. rodrigoi sp. nov. holotype, SP, Aranjuez c A. rutilipennis (A. ressli, holotype), TR, Hatay d A. chellala sp. nov., holotype, AG, Chellala e A. discoides, neotype, IS, Keneitra f, g A. kraatzi f SV g RU, Karasuk, W. Siberia h A. rusakovi, paratype ♂, RU, Orenburg i – t A. niger group, mid tarsi, and tibial apices i – o SEM images p – t stacked photographs i A. niger, Sweden, Södermanland, Tullgarn j A. muscorum, HU, Hortobágyi k, l A. bameuli sp. nov., paratypes, FR, Corsica m, n A. krelli sp. nov., paratypes, IT, Sardinia o A. wilsonae, paratype SP p, q A. felix sp. nov, IT, Abruzzo, Campo Felice p paratype q holotype r A. bellumgerens sp. nov., paratype, IT, Sicily s A. isikdagensis, paratype, TR, Ḉamildere t A. alberti sp. nov., holotype, TR, Rize, Ovitdag u, v fore tibiae and tarsi of ♂♂ to show the spur. u A. bameuli sp. nov., paratype, FR, Corsica v A. rusakovi, paratype, RU, Orenburg w, x apex of hind tibia to show the short spines w A. muscorum, HU x A. wilsonae, paratype, SP. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure18" httpUri="https://binary.pensoft.net/fig/1097482">18 r</figureCitation>
). Spines on apical margin of posterior tarsi short and even on outer / ventral face, longer, sparser, and more irregular on inner / dorsal face.
</paragraph>
<paragraph id="3C54182318F93D302CC42485408D10F8">
Aedeagus (Figs
<figureCitation id="BBAFC5B0718E0E758FE8B359505BED88" captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. A. niger group, aedeagi with inflated endophalli, stacked photographs and SEM images a – e, g – w right lateral view f viewed from above a – c A. niger, SV, Södermanland, Tullgarn a stacked photograph b the same aedeagus, SEM image c detail of endophallic teeth d – f A. niger, GB, Hampshire, New Forest d stacked photograph e, f SEM images g – i A. wilsonae, SP, Areños g stacked photograph h SEM image, i detail of endophallic teeth j – l A. bameuli sp. nov., holotype j stacked photograph k SEM image l detail of endophallic teeth m – o A. krelli sp. nov., holotype m stacked photograph n SEM image o detail of endophallic teeth p – r A. bellumgerens sp. nov., paratype p stacked photograph q SEM image r detail of endophallic teeth s, t A. bellumgerens sp. nov., holotype s stacked photograph t SEM image of endophallus u A. muscorum, HU, Hortobágyi, stacked photograph v, w A. felix sp. nov., holotype, SEM images v aedeagus w detail of endophallic teeth. Black Scale bar: 0.5 mm for aedeagi and endophallus, white SEM scale on black background: 50 μm for details of endophallic teeth." figureDoi="10.3897/zookeys.1207.117225.figure23" httpUri="https://binary.pensoft.net/fig/1097492">23 p – t</figureCitation>
,
<figureCitation id="DA4D03F625049D2D8C15637F43C323E1" captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. A. niger group aedeagi, cleared preparations mounted in DMHF on slides a – g, m – o dorsoventral view h – l, p right lateral view a A. niger, SV, Södermanland, Tullgarn b A. bameuli sp. nov., paratype c A. isikdagensis, paratype, TR, Ḉamildere d A. krelli sp. nov., paratype, IT, Sardinia e – f A. alberti sp. nov., TR, Rize, Ovitdag e paratype, after squashing f the same specimen, before squashing g, h, j A. alberti sp. nov., different paratypes i A. alberti sp. nov., paratype, paramere k A. isikdagensis?, TR, Artvin l A. alberti?, Armenia m A. bellumgerens sp. nov., paratype, IT, Sicily, Piano Battaglia n A. bellumgerens, paratype IT, Sicily, Nebrodi o A. wilsonae, paratype, SP, Manzanares el Real p A. muscorum, HU, Hortobági. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure24" httpUri="https://binary.pensoft.net/fig/1097494">
<quantity id="2F175D9485E57FCD0F59E637911C9699" metricMagnitude="1" metricUnit="m" metricValue="2.4" unit="m" value="24.0">24 m</quantity>
, n
</figureCitation>
) Length ca
<quantity id="D26123A00F56CA564E8DC3BE92D8B2C6" metricMagnitude="-3" metricUnit="m" metricValue="1.0" metricValueMax="1.1" metricValueMin="0.9" unit="mm" value="1.0" valueMax="1.1" valueMin="0.9">0.9–1.1 mm</quantity>
, paramere length ca
<quantity id="270FDC45E9D37239CFE3FA78D4A81484" metricMagnitude="-4" metricUnit="m" metricValue="4.5" metricValueMax="4.699999999999999" metricValueMin="4.3" unit="mm" value="0.44999999999999996" valueMax="0.47" valueMin="0.43">0.43–0.47 mm</quantity>
, basal piece length ca
<quantity id="B4BA374DD0042B6B4AF1066113112632" metricMagnitude="-4" metricUnit="m" metricValue="5.5" metricValueMax="6.0" metricValueMin="5.0" unit="mm" value="0.55" valueMax="0.6" valueMin="0.5">0.5–0.6 mm</quantity>
, tooth field length ca
<quantity id="38592FB64459A8E7ACAE951FFCE5F6B9" metricMagnitude="-4" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" unit="mm" value="0.35" valueMax="0.4" valueMin="0.3">0.3–0.4 mm</quantity>
. This is one of the few species in which the length of the endophallic tooth-field is less than that of the parameres. Longest teeth on endophallus ca 47 μm long.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E31BM" type="etymology">
<paragraph id="F8F1CF82FBEABBA4B2BD41A84B27C144">
<heading id="EBE4313E3274B34B5F98C975DCDF3E4C" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="E82023A50607D9826D6433AFE9A140D4">
<taxonomicName id="A3066E71CE44DB4DEE2191F09168331D" authorityName="Angus &amp; Maté &amp; Angus &amp; Král" authorityYear="2024" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="bellumgerens">
<emphasis id="AA577DA42E66ADED4AF2A95E2EB8D8A6" italics="true">bellumgerens</emphasis>
</taxonomicName>
– Latin, waging war, adjective, named for the
<typeStatus id="DABEF0879C4E1343581748E392C5F880">type</typeStatus>
locality, Piano Battaglia (the battlefield).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EM2BM" type="remarks">
<paragraph id="415D1AD5D310FE2C39F3DB0891B540F8">
<heading id="BD3FE0DCD2C283F0D9E75C3882E810A5" reason="title">Remarks.</heading>
</paragraph>
<paragraph id="89489D87E4B45E6AB8282ADBAE6F02DE">
So far known only from
<collectingRegion id="47F044D9F84C480C07F2023641979228" country="Italy" name="Sicilia">Sicily</collectingRegion>
(Fig.
<figureCitation id="3D03B8AE96896A5A2C1B53633C566430" captionStart="Figure 29" captionStartId="F29" captionText="Figure 29. a distribution of niger - group taxa in Western and Central Europe b distribution of Liothorax species in Central and Eastern Palearctic c distribution of plagiatus - group species in Central and Eastern Palearctic. The record of isikdagensis with the question mark indicates a doubtful specimen from Artvin Çam, TR (Ziani Coll)." figureDoi="10.3897/zookeys.1207.117225.figure29" httpUri="https://binary.pensoft.net/fig/1097509">29 a</figureCitation>
). Many of the
<typeStatus id="BC7EEFF6613ACD8968ABA3E72C8A457C">types</typeStatus>
were taken by swishing the water at the edges of the pool on the Piano Battaglia, washing the beetles from the banks so that they floated on the water. This pool dries out completely in the summer. The Monte Soro specimens were floating and clinging to plants in a shallow pool, following recent rain (Fig.
<figureCitation id="919A61DB8380AEDBBFC04D72F401CC4D" captionStart="Figure 27" captionStartId="F27" captionText="Figure 27. Habitat pictures a A. niger, GB, Hampshire, New Forest b A. muscorum, HU, Hortobagyi c A. niger, SW, Tullgarn d A. krelli sp. nov., IT, Sardinia, Badde Salighes e A. bameuli sp. nov., FR, Corsica, Haute-Corse, Lac de Melo f A. wilsonae, SP, Cantabria, Corconte g A. bellumgerens sp. nov., IT, Sicily, Nebrodi." figureDoi="10.3897/zookeys.1207.117225.figure27" httpUri="https://binary.pensoft.net/fig/1097502">
<quantity id="24CC5644593ABD234CFF37C7E1431419" metricMagnitude="-2" metricUnit="kg" metricValue="2.7" unit="g" value="27.0">27 g</quantity>
</figureCitation>
). We have seen specimens from
<collectingRegion id="990EAD83AAB701486E1FFA7EF5B3278B" country="Italy" name="Calabria">Calabria</collectingRegion>
(Sta Christina, Cippo di Garibaldi) which are apparently morphologically identical but have not been included in the
<typeStatus id="A01961F843180F1922611F0A999B9F7C">type</typeStatus>
series due to the scarcity of material and lack of karyological or molecular data. RBA has recently received living material from
<collectingRegion id="95FC303B6A2082D036D4FDC407C48960" country="Italy" name="Calabria">Calabria</collectingRegion>
: Villaggio Mancusa,
<named-content content-type="dwc:verbatimCoordinates" id="NCID0E42BM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[16.640850,39.107270]}">
<geoCoordinate id="451F97B239513F6B3D2889F95A6C9DEB" degrees="39.10727" direction="north" orientation="latitude" precision="1" value="39.10727">39.10727 ° N</geoCoordinate>
,
<geoCoordinate id="F9A5684CF17BAE0FA781056183C9623B" degrees="16.64085" direction="east" orientation="longitude" precision="1" value="16.64085">16.64085 ° E</geoCoordinate>
</named-content>
, from which chromosomes indicating
<taxonomicName id="97537084ACFB9D0FBA7CE4789B09B0EF" authorityName="Angus &amp; Maté &amp; Angus &amp; Král" authorityYear="2024" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="felix">
<emphasis id="F3D63C0111516AF3A3387347B70B82D5" italics="true">A. felix</emphasis>
</taxonomicName>
, not
<taxonomicName id="C386355B956BF8A40E64F72CFCBD0053" authorityName="Angus &amp; Maté &amp; Angus &amp; Král" authorityYear="2024" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="bellumgerens">
<emphasis id="2DDB1B651E7C4401B471B4D6E715ED70" italics="true">A. bellumgerens</emphasis>
</taxonomicName>
have been obtained. These data will be published in due course.
</paragraph>
</subSubSection>
</treatment>
</document>