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<document id="29F71CD27973CA21C42DEF78FD2D618E" 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="AF8BC118B8C0561788F1062974FE8977" 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) niger Illiger 1798" docType="treatment" 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">
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<mods:title id="D664EC5BD14D5B4A6026246C1CBA7E77">Towards a revision of the Palaearctic species of Aphodius Hellwig, 1798, subgenus Liothorax Motschulsky, 1860 (Coleoptera, Scarabaeidae, Aphodiinae)</mods:title>
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(
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<emphasis id="F4C6B0B4F20044A3E16F763419AD0092" italics="true">niger</emphasis>
Illiger, 1798
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<figureCitation id="220E7885C20228A07F80BA3110FB0937" 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 a</figureCitation>
, 
<figureCitation id="2C5A8F22DF1E3EE87F8F9D79EA6D1A31" 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 e</figureCitation>
, 
<figureCitation id="46BB9A5CB2F93CED57130CE98AC78EC1" 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 a – c</figureCitation>
, 
<figureCitation id="A8C027A70F1FA78FC104E49D74D82CAF" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Aphodius niger group, pronotal bases 1 a A. niger, Sweden, Södermanland, Tullgarn b A. cf. niger, CZ, České Budějovice c A. muscorum, HU, Hortobágyi d A. felix sp. nov., paratype." figureDoi="10.3897/zookeys.1207.117225.figure15" httpUri="https://binary.pensoft.net/fig/1097476">15 a, b</figureCitation>
, 
<figureCitation id="35A1AB950621997C67EE0CFA782AFBF7" 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 l, m</figureCitation>
, 
<figureCitation id="1DB8D17A6B941208B8E8C84D53CF26D7" 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 i</figureCitation>
, 
<figureCitation id="E8F97FFC1EAAA0A6FF5956D766819515" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Elytral apices a A. (L.) niger, SV, Södermanland, Tullgarn b A. (Nialus) varians, FR, left elytron c A. (Calamosternus) granarius, CY, right elytron d A. (C.) hyxos, CY, left elytron e A. (Agoliinus) lapponum, GB, Cumbria, both elytra f A. (Agrilinus) constans, IT, Sardinia, left elytron. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure19" httpUri="https://binary.pensoft.net/fig/1097484">19 a</figureCitation>
, 
<figureCitation id="E89A1C2A68A5A981FCA3B0B359A2EEB3" 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 a – c</figureCitation>
, 
<figureCitation id="7D0AF7DA82361512ADC2A9B0E58DA45C" 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 a – f</figureCitation>
, 
<figureCitation id="0CD05893EEF6F4CD0E160C81725A303B" 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">24 a</figureCitation>
, 
<figureCitation id="C9D3F6584339539A9D44F0C5472E945D" 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 a</figureCitation>
, 
<figureCitation id="6A6AC5EB2508669DEBEE8D8A901AE1EE" 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">
<quantity id="E4755080BC079D5F42C7074B65F0111B" metricMagnitude="4" metricUnit="kg" metricValue="2.6" unit="t" value="26.0">26 t</quantity>
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, 
<figureCitation id="097A555D11CA87F369754439F1908927" captionStart="Figure 31" captionStartId="F31" captionText="Figure 31. Mitotic chromosomes, arranged as karyotypes a, b A. niger, ♂, SV, Tullgarn a plain b the same nucleus C-banded (1 replicate of autosome 5 missing) c A. niger ♀, SV, Tullgarn, C-banded d, e A. niger, ♂♂, GB, Hampshire, New Forest, C-banded e with 3 B-chromosomes f, g A. muscorum, ♂, HU, Hortobágyi f plain g the same nucleus C-banded h A. muscorum, ♂, HU, Hortobágyi, C-banded i, j A. muscorum, ♀, HU, Kisújszállás i plain j C-banded k, l A. muscorum, ♀, HU, Kisújszállás k plain l the same nucleus C-banded. Scale bar: 5 μm" figureDoi="10.3897/zookeys.1207.117225.figure31" httpUri="https://binary.pensoft.net/fig/1097515">31 a, b, d, e</figureCitation>
, 
<figureCitation id="C9131A7AAEE25CB50DDD1AFD0CE263DA" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Meiotic chromosomes, metaphase I from testis a – d A. niger, GB, Hampshire, New Forest, White Moor a, c plain b, d the same nuclei C-banded e – g A. muscorum, HU, Hortobágyi e plain f, g the same nucleus f plain g C-banded h A. bameuli paratype, FR, Corsica, by Lac de Melo, plain i, j A. krelli IT, Sardinia, paratype i plain j the same nucleus C-banded k, l A. wilsonae, SP, Areños k plain l the same nucleus C-banded. The sex bivalent is indicated by an arrow. Scale bar: 5 μm." figureDoi="10.3897/zookeys.1207.117225.figure34" httpUri="https://binary.pensoft.net/fig/1097526">34 a – d</figureCitation>
</paragraph>
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<taxonomicName id="4E0E2DB354D3ADFF2AFC4FBC59EE3AB9" ID-CoL="FHT7" authority="Illiger, 1798: 24" authorityName="Illiger" authorityPageNumber="24" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="A1D89CFDBDEB8E637E998CB0B26ED929" italics="true">Aphodius niger</emphasis>
Illiger, 1798: 24
</taxonomicName>
. Conserved as valid name: International Commission for Zoological Nomenclature Opinion 2009 (2005: 45).
</treatmentCitation>
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="SECID0ERHAM" type="type material">
<paragraph id="8C4673081A4C6A8AB8474F49BC3A6C37">
<heading id="D6E7AD72E274EC1D2B9EB55C6AB5447B" reason="title">Type material examined.</heading>
</paragraph>
<paragraph id="1E5B9A35DC23479DB7912B4474581B6E">
<materialsCitation id="213FF3365280D9359CCF7FB51AB06DED" collectionCode="ZMB" specimenCount="1" typeStatus="lectotype">
<emphasis id="F25A69A0193055CCA4240FA10D7661B9" bold="true" italics="true">
<typeStatus id="86F416316612102089DA959023133D1A" type="lectotype">Lectotype</typeStatus>
</emphasis>
, unsexed, “ Suec ”, designated by 
<bibRefCitation id="4F0A8120B7743E579F9F5CDDFF910416" author="Krell F-T &amp; Mann DJ &amp; Angus RB &amp; Maté JF" etAl="et al." firstAuthor="Krell" issue="2" journalOrPublisher="Bulletin of Zoological Nomenclature" pagination="127-131" refId="B27" refString="Krell F-T, Mann DJ, Angus RB, Maté JF (2003) Case 3254 Aphodius niger Illiger, 1798 (Insecta, Coleoptera): Proposed conservation of the specific name. Bulletin of Zoological Nomenclature 60 (2): 127–131." title="Case 3254 Aphodius niger Illiger, 1798 (Insecta, Coleoptera): Proposed conservation of the specific name." volume="60" year="2003">Krell et al. (2003)</bibRefCitation>
(
<abbrev id="ABBRID0EAIAM" xlink_title="Humboldt Museum">
<collectionCode id="90A5AF045D57F53D91A81D1DD0614C25" country="Germany" httpUri="http://grbio.org/cool/8syf-d21i" name="Museum für Naturkunde Berlin (Zoological Collections)" type="Museum">ZMB</collectionCode>
</abbrev>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EEIAM" type="material">
<paragraph id="CD968201A0D2E7F94E5678073DA49473">
<heading id="FA08EB652F0FA6CDFCF3A0B74CEEB7F4" reason="title">Additional material examined.</heading>
</paragraph>
<paragraph id="B38CFE09D22C877C52A268F3633310C7">
SV: 
<collectingRegion id="6AAC84F658A8134A8B793D4AEF9A9B4B" country="Sweden" name="Sodermanland">Sodermanland</collectingRegion>
, Holo, Tulgarn, Nasuden. 
<named-content content-type="dwc:verbatimCoordinates" id="NCID0ENIAM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[17.620000,58.950000]}">
<geoCoordinate id="A82BA3EC82A78FF1510C7EF261C10056" degrees="58.95" direction="north" orientation="latitude" precision="555" value="58.95">58.95 ° N</geoCoordinate>
, 
<geoCoordinate id="97FF730A282DC0C3786A8647096D3872" degrees="17.62" direction="east" orientation="longitude" precision="555" value="17.62">17.62 ° E</geoCoordinate>
</named-content>
. Trampled bare organic mud. 
<date id="CE5A738E070F805D9685EBFBAAC3F315" value="2011-05-15">15. v. 2011</date>
. Hans-Erik &amp; Livia Wanntorp. 
<specimenCount id="DCA5CA9F90C331EEF4444B3A656E0EEB" count="4" type="male">4 ♂♂</specimenCount>
, chromosome preps 2: 
<date id="1C84CD0D9208D46AE11025732463CD6A" value="2001-05-24">24. v. 2001</date>
, 1: 
<date id="CA2A7F0B30B977CBA756B2822C6135D9" value="2011-05-25">25. v. 2011</date>
, 2: 
<date id="68E671D1DC8A0A7F50D3BC92ADADD0F0" value="2011-05-25">25. v. 2011</date>
, 1: 
<date id="120AB96A9234774F75CB8CFE50DC221E" value="2011-05-31">31. v. 2011</date>
, R. B. 
<collectingRegion id="C8077C25C0C8ED0F79FE5D11BD1C29AE" country="United Kingdom" name="Angus">Angus</collectingRegion>
; 
<specimenCount id="F674A0B5E707EBE48FF29C50C82901DF" count="7" type="female">7 ♀♀</specimenCount>
, chromosome preps 3: 
<date id="1D711F19B833BE035B40AD32BF1420BA" value="2011-05-24">24. v. 2011</date>
, 4: 
<date id="645047CBBC03A4BE60AAEBC275391ACD" value="2011-05-24">24. v. 2011</date>
, 9: 
<date id="DB7FC9FE5279CD8C1B882666F10A3BFC" value="2011-05-25">25. v. 2011</date>
, 2: 
<date id="5825ECE91922AEED9D13C57D9D6BC532" value="2011-05-31">31. v. 2011</date>
, 3: 
<date id="A3BC13BC80D18D22AFE1FC0F43B87B6F" value="2011-05-31">31. v. 2011</date>
, 4: 
<date id="DAE909C83A5FE054CFDA260F673CB9AB" value="2015-05-31">31. v. 2015</date>
, 5: 
<date id="18225223DEDAC16456D62954BA5B97D3" value="2011-05-31">31. v. 2011</date>
, R. B. 
<collectingRegion id="74A5E96FCE665FD3B89D99197EE86E69" country="United Kingdom" name="Angus">Angus</collectingRegion>
(
<named-content id="F1027E7393285F463647335A9F371325" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="720CAAE6CC05BDD9C77B6CBCE41BE37F" 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="65CF96CCF242F0049B62A22464ACAAED" count="1" type="male">1 ♂</specimenCount>
same data, chromosome prep. 2: 
<date id="2B886B3D64EC4ED1FE08A51DDC04A686" value="2011-05-24">24. v. 2011</date>
(
<named-content id="D7B692B9238CE9DEAEC514457F92693A" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-prague-0" xlink_title="National Museum Prague">
<collectionCode id="FF6CD3379F3998C39D0A154425BAAFF9" country="Czech Republic" httpUri="http://grbio.org/cool/fx6k-enrh" name="National Museum (Prague)" type="">NMP</collectionCode>
</named-content>
).
</paragraph>
<paragraph id="0480F8677DA123747B36C820C5F107D6">
GB: 
<collectingRegion id="9722F7FA8B4DF797CF310130DAA1F6E2" country="United Kingdom" name="Hampshire">Hampshire</collectingRegion>
, New Forest. Long Slade Bottom. 
<named-content content-type="dwc:verbatimCoordinates" id="NCID0E6IAM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-1.619000,50.801000]}">
<geoCoordinate id="05ED48EE86E904138A6C80CE7B17B0B5" degrees="50.801" direction="north" orientation="latitude" precision="55" value="50.801">50.801 ° N</geoCoordinate>
, 
<geoCoordinate id="C48770F04278479FDA466131D720666A" degrees="1.619" direction="west" orientation="longitude" precision="55" value="-1.619">1.619 ° W</geoCoordinate>
</named-content>
. Temporary pool after heavy rain. 
<date id="34C8D5623525208937D041CF5426B472" value="2002-05-30">30. v. 2002</date>
, R. B. 
<collectingRegion id="422121C30CE6E235949BFABE4FB80271" country="United Kingdom" name="Angus">Angus</collectingRegion>
. 
<specimenCount id="9D88CC7E7685A02D1B3E52D8AEBC5A5A" count="4" type="male">4 ♂♂</specimenCount>
, 
<specimenCount id="6175475EC479F0AFBA9AA4AA69F961A3" count="4" type="female">4 ♀♀</specimenCount>
, 5 unsexed. Chromosome preps 1–3: 31. v., 2: 1. vi, 4: 
<date id="1D74AA6DC4180348377C97144541A388" value="2011-06-01">1. vi. 2011</date>
, R. B. 
<collectingRegion id="0D3A0E90293BEE4E139AE0D9AEA2C75F" country="United Kingdom" name="Angus">Angus</collectingRegion>
(
<named-content id="9A260F78F8C6DF6C8DBB548F9AE5DB18" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="84EB037B3D6A830B56D32227D47D8CC3" 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="F6DBA67248F607EDD6185B892A5C0630" count="1" type="male">1 ♂</specimenCount>
, same data. (
<named-content id="70905218842D272EF445C3646736F933" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-prague-0" xlink_title="National Museum Prague">
<collectionCode id="24283CA7EC558CEC10FDB364CB797B44" country="Czech Republic" httpUri="http://grbio.org/cool/fx6k-enrh" name="National Museum (Prague)" type="">NMP</collectionCode>
</named-content>
); New Forest, White Moor Pond. 
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EPJAM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-1.607000,50.820000]}">
<geoCoordinate id="D1536232211E47E39AF5D00043CF403F" degrees="50.820" direction="north" orientation="latitude" precision="55" value="50.82">50.820 ° N</geoCoordinate>
, 
<geoCoordinate id="46EF09FFAD31D4B034BE2EFF6417FE45" degrees="1.607" direction="west" orientation="longitude" precision="55" value="-1.607">1.607 ° W</geoCoordinate>
</named-content>
. 
<date id="D51B178E25D5629CEAF2DECC4930632E" value="2015-10-02">2. x. 2015</date>
, 
<date id="F85ABD2EB0C4F9AF92EEBF25F39CD16C" value="2016-05-02">2. v. 2016</date>
, R. B. 
<collectingRegion id="ACD7740E8E5E160333A0BDF58B81B524" country="United Kingdom" name="Angus">Angus</collectingRegion>
. 
<specimenCount id="66EBCDEBE6CFFC2C443F0A8D853EC9C9" count="8" type="male">8 ♂♂</specimenCount>
, 
<specimenCount id="1B2AC82ACBB3A2A29299AE28E0AC8300" count="7" type="female">7 ♀♀</specimenCount>
, 3 unsexed. Chromosome preps: 3: 
<date id="1C4E761A17CD83D946F1EA2BB21E8F79" value="2015-10">x. 2015</date>
, 5: 
<date id="071DDF65553EE1DF7D7533750DE306C8" value="2016-05">v. 2016</date>
. New Forest, Balmer Lawn. 
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EXJAM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-1.567000,50.828000]}">
<geoCoordinate id="09ADC8DD03A4525E9C44FFA9A3136CE0" degrees="50.828" direction="north" orientation="latitude" precision="55" value="50.828">50.828 ° N</geoCoordinate>
, 
<geoCoordinate id="69387DA92148F51C32A3BED1244037E1" degrees="1.567" direction="west" orientation="longitude" precision="55" value="-1.567">1.567 ° W</geoCoordinate>
</named-content>
. At edges of pond and ditch. 
<date id="87C7638D1D97DA6F8313F5AEBC121C34" value="2002-05">V. 2002</date>
. R. B. 
<collectingRegion id="C38026BF2024ECD7F3C311BFA90E1AB4" country="United Kingdom" name="Angus">Angus</collectingRegion>
(
<named-content id="830FD7AA73B99736DB0840EEA92D9454" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="8998A8F82D1E77E021AB639001803771" 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>
). Inkpen Common, Berkshire 
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EDKAM" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-1.453000,51.375000]}">
<geoCoordinate id="565497BBB6C216B26C0E9324EB3AD6CC" degrees="51.375" direction="north" orientation="latitude" precision="55" value="51.375">51.375 ° N</geoCoordinate>
, 
<geoCoordinate id="AF1479C0B9BEB21C26DF36F350798869" degrees="1.453" direction="west" orientation="longitude" precision="55" value="-1.453">1.453 ° W</geoCoordinate>
</named-content>
(fide 
<bibRefCitation id="826798CA12790F010725F3613F08D1A3" author="Mann DJ &amp; Garvey L" firstAuthor="Mann" issue="3" journalOrPublisher="Coleopterist" pagination="138-142" refId="B31" refString="Mann DJ, Garvey L (2014) Aphodius (Liothorax) niger (Illiger) (Scarabaeidae: Aphodiinae) inside and outside the New Forest. Coleopterist 23 (3): 138–142." title="Aphodius (Liothorax) niger (Illiger) (Scarabaeidae: Aphodiinae) inside and outside the New Forest." volume="23" year="2014">Mann and Garvey (2014)</bibRefCitation>
, not examined) 1 unsexed (
<named-content id="1F513F612843479FFF0E482AB88A2C1D" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/oxford-university-museum-natural-history" xlink_title="Oxford University Museum">
<collectionCode id="6DB108FD22AF434568397A2ACC852CCE" country="United Kingdom" httpUri="http://grbio.org/cool/g6yf-wft0" name="Oxford University Museum of Natural History" type="">OUM</collectionCode>
</named-content>
).
</paragraph>
<paragraph id="0FF665D631719665AFBF6A28BF38294B">
RU: 
<collectingRegion id="BC1FC490C4F9EB7D4F400A3AA8DF44BA" country="Russia" name="Yaroslavl'">Yaroslavl</collectingRegion>
oblast’, Berditsino, 
<geoCoordinate id="E7C6CB9DBB559DF25D9B0DCCE7BCBEDF" degrees="57.454" direction="north" orientation="latitude" precision="55" value="57.454">57.454 ° N</geoCoordinate>
, 
<geoCoordinate id="47E64EA5914C3FBA12DDFEA0A1E1291C" degrees="40.1108" direction="east" orientation="longitude" precision="5" value="40.1108">40.1108 ° E</geoCoordinate>
. leg. Yakovlev. 
<specimenCount id="19EB0375C08180E1FF16CCC1EAF9AE3F" count="1" type="male">1 ♂</specimenCount>
(
<named-content id="EF6B5EFCBA9B7E79922FFE6247B340C0" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/naturhistorisches-museum-wien-0" xlink_title="Naturhistorisches Museum, Wien">
<collectionCode id="463947FF1F1A91DF12BB50378B7CD71A" country="Austria" httpUri="http://grbio.org/cool/c7yt-hh77" name="Naturhistorisches Museum, Wien" type="">NMW</collectionCode>
</named-content>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EWKAM" type="Differential diagnosis">
<paragraph id="8B3C07DD2FDA8129F59FE1C0C97BAE5C">
<heading id="19A4F054291FD95BA278F7F378F8177B" reason="title">Differential diagnosis.</heading>
</paragraph>
<paragraph id="FD8492977465EA3A018C5C7CE4C26E00">
<taxonomicName id="48527262B01AEDC7E5BE2BC2331ECFA1" authorityName="Illiger" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="0AB315111688D1711A50AC0D50FEE83A" italics="true">Aphodius niger</emphasis>
</taxonomicName>
is immediately distinguished from all other 
<taxonomicName id="89DF6286ACDC1AE75605D18E082B8025">
<emphasis id="8E4E13319C691D3B81C6CA5BC930ED69" italics="true">Liothorax</emphasis>
species
</taxonomicName>
(and all 
<taxonomicName id="6DF04FC7B7C3EE966C5BE83D20875F79" authorityName="Leach" authorityYear="1815" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="subFamily" subFamily="Aphodiinae">Aphodiinae</taxonomicName>
whose chromosomes are known) by its long, almost entirely euchromatic, Y chromosome (Fig. 
<figureCitation id="79FF85C641D389E21B965B24AFF3BA9B" captionStart="Figure 31" captionStartId="F31" captionText="Figure 31. Mitotic chromosomes, arranged as karyotypes a, b A. niger, ♂, SV, Tullgarn a plain b the same nucleus C-banded (1 replicate of autosome 5 missing) c A. niger ♀, SV, Tullgarn, C-banded d, e A. niger, ♂♂, GB, Hampshire, New Forest, C-banded e with 3 B-chromosomes f, g A. muscorum, ♂, HU, Hortobágyi f plain g the same nucleus C-banded h A. muscorum, ♂, HU, Hortobágyi, C-banded i, j A. muscorum, ♀, HU, Kisújszállás i plain j C-banded k, l A. muscorum, ♀, HU, Kisújszállás k plain l the same nucleus C-banded. Scale bar: 5 μm" figureDoi="10.3897/zookeys.1207.117225.figure31" httpUri="https://binary.pensoft.net/fig/1097515">31 a, b, d, e</figureCitation>
). The basal segment of the mesotarsus is always clearly longer than the longer mesotibial spur (Fig. 
<figureCitation id="104556791C34007426DFD7188DE4AA93" 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 i</figureCitation>
). Elytra glossy black, though interstices have fine reticulation. Central part of metaventrite without reticulation (Fig. 
<figureCitation id="86C1042F3494E0C0319D1C4B0D970269" 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 a – d</figureCitation>
). Endophallic teeth clearly longer than high. Length of longest endophallic teeth at least 40 μm.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E6LAM" type="redescription">
<paragraph id="C9702EE6C78F4F20FE3B4F59BA9167DC">
<heading id="CB613ECB9021B0A69573A6B1C79981B7" reason="title">Redescription.</heading>
</paragraph>
<paragraph id="FAC89330199CA16CED67EB378846D1FF">
General appearance (Fig. 
<figureCitation id="639452F0AA4CDFE272A8F9A55D7C15E8" 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 a</figureCitation>
). Length 
<quantity id="AB918F647DF53B757F9BDCEE51982DF1" metricMagnitude="-3" metricUnit="m" metricValue="4.35" metricValueMax="4.5" metricValueMin="4.2" unit="mm" value="4.35" valueMax="4.5" valueMin="4.2">4.2–4.5 mm</quantity>
(
<specimenCount id="6180AB73245FF5450A5A65E975F65323" count="1" type="male">♂</specimenCount>
), 
<quantity id="86180F5E246B0BB9C411C7549D2F702E" metricMagnitude="-3" metricUnit="m" metricValue="4.55" metricValueMax="5.0" metricValueMin="4.1" unit="mm" value="4.55" valueMax="5.0" valueMin="4.1">4.1–5 mm</quantity>
(
<specimenCount id="AFCDCE607BF1AA373ACAF9A566217194" count="1" type="female">♀</specimenCount>
), width 
<quantity id="6FC2E7286462D77A8B6342D7E3C0FEA0" metricMagnitude="-3" metricUnit="m" metricValue="1.9" metricValueMax="2.0" metricValueMin="1.8" unit="mm" value="1.9" valueMax="2.0" valueMin="1.8">1.8–2 mm</quantity>
(
<specimenCount id="DF7D4BFAAE097B756F599673C7D4F9EC" count="1" type="male">♂</specimenCount>
), 
<quantity id="2C4EB9827DE60A4BB7F37D759F023CBE" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="2.2" metricValueMin="1.8" unit="mm" value="2.0" valueMax="2.2" valueMin="1.8">1.8–2.2 mm</quantity>
(
<specimenCount id="07ACC0ABE57056484470789B21FFCEC1" count="1" type="female">♀</specimenCount>
) (Swedish material), 
<quantity id="DC3F56186B6F5A75A78BC867D72A192E" metricMagnitude="-3" metricUnit="m" metricValue="4.6" metricValueMax="4.8" metricValueMin="4.4" unit="mm" value="4.6" valueMax="4.8" valueMin="4.4">4.4–4.8 mm</quantity>
(
<specimenCount id="B9BBE306EC324955DEB8FFB66C18086F" count="1" type="male">♂</specimenCount>
), 
<quantity id="5E234F6FC9C7B8CB09AC1243DD8148CB" metricMagnitude="-3" metricUnit="m" metricValue="4.75" metricValueMax="5.1" metricValueMin="4.4" unit="mm" value="4.75" valueMax="5.1" valueMin="4.4">4.4–5.1 mm</quantity>
(
<specimenCount id="BBEBC0EBC36EDB761E962F8FB9018B3A" count="1" type="female">♀</specimenCount>
), width 2.0– 
<quantity id="799BB1D7E11680D30CC36161EC74FAAC" metricMagnitude="-3" metricUnit="m" metricValue="2.2" unit="mm" value="2.2">2.2 mm</quantity>
(
<specimenCount id="AC52931AF4886EB6FE8AD5CB1B2E6BF3" count="1" type="male">♂</specimenCount>
and 
<specimenCount id="6C9A8DC45EF8F3353155F1BFB8326C75" count="1" type="female">♀</specimenCount>
) (English Material).
</paragraph>
<paragraph id="D77ABE4D3855D3A11FD88A5EF30D88BA">
Head black, without obvious brownish margin. Anterior margin of clypeus excised over middle 1 / 3, the lateral edges of this excision bluntly rounded. Frontoclypeal suture very fine, without any trace of a tubercle, straight-transverse over middle 1 / 2, angled forward to run to the clypeo-genal junction laterally. The lateral parts of the suture may be almost completely effaced. Clypeus raised in a rounded bulge medially. Surface moderately strongly punctate with double punctation. Larger punctures separated by ca their own diameter over most of the head, but sparser on clypeal bulge. Antennae and palpi blackish brown. Epipharynx (Fig. 
<figureCitation id="45BA84A89AB9A766FFECE537462DDB5A" 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 a – c</figureCitation>
) with the anterior margin of the clithra strongly excised either side of the median tylus and with a few fine acropariae. Chaetopedia stout, 3–6 each side. Chaetopariae ca as stout as chaetopedia, forming a close-set line each side. Apophobae very fine, arranged in a line outside that of the chaetopariae; prophobae vine, quite long, arranged against the sides of the mesoepitorma. Pronotum (Fig. 
<figureCitation id="A7955718A7ED8A2E6138BF67D86D1115" 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 e</figureCitation>
) hemicylindrical, black, highly arched transversely, weakly so longitudinally, lateral parts of surface bulging outwards so that the lateral margins are often not visible from above in their basal 1 / 3. Surface glossy black, with double punctation, the larger punctures separated by 1–2 × their diameter, sparser medially and sometimes petering out in anterior 1 / 8 where only the fine punctures are present. Scutellum elongate, pentagonal, glossy black with a few sparse punctures. Elytra black, interstices flat, 6–8 × the width of the striae, finely reticulate and with sparse fine punctures (Fig. 
<figureCitation id="32D810C09A80118CCB649F6D4D3088F9" 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 l, m</figureCitation>
), the reticulation stronger towards the apex (Fig. 
<figureCitation id="DEB47EF21CD4A715D0C29B2DC29B5B86" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Elytral apices a A. (L.) niger, SV, Södermanland, Tullgarn b A. (Nialus) varians, FR, left elytron c A. (Calamosternus) granarius, CY, right elytron d A. (C.) hyxos, CY, left elytron e A. (Agoliinus) lapponum, GB, Cumbria, both elytra f A. (Agrilinus) constans, IT, Sardinia, left elytron. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure19" httpUri="https://binary.pensoft.net/fig/1097484">19 a</figureCitation>
). Striae glossy, without reticulation, with a single row of punctures separated by ca 2 × their diameter and excising the strial margins. Metaventrite moderately punctate, the median plate not reticulate, sometimes distinctly concave in males Fig. 
<figureCitation id="ACB77BE4B86F24FFF6974F19F9127312" 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 a</figureCitation>
but sometimes almost flat with a median impressed line, as in females (Fig. 
<figureCitation id="A2A9AEA476A2C5743B2282756E7BCCC5" 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 b, c</figureCitation>
). Legs blackish brown, mesotarsi always with basal segment longer than longer mesotibial spur (Fig. 
<figureCitation id="F3920F2A0B655A11C7DE016A1F71CA52" 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 i</figureCitation>
).
</paragraph>
<paragraph id="AAFF7B9DAA835F382B006D40EAD45A24">
Aedeagus (Figs 
<figureCitation id="C0ED862E2D8BA8F3652FF34A649AC89C" 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 a – f</figureCitation>
, 
<figureCitation id="FCCDD66E059CF1FC51A2647F7AFC93A1" 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">24 a</figureCitation>
) length ca 
<quantity id="6F12E5DA7F83CE443AFBFE110028E05D" metricMagnitude="-3" metricUnit="m" metricValue="1.2" unit="mm" value="1.2">1.2 mm</quantity>
, paramere length ca 
<quantity id="6B84CF9A26136AFDE6290B46D9672A90" metricMagnitude="-4" metricUnit="m" metricValue="4.8" unit="mm" value="0.48">0.48 mm</quantity>
, basal piece length ca 
<quantity id="4C80F36694A53D5A9F34031B30BF6179" metricMagnitude="-4" metricUnit="m" metricValue="6.8999999999999995" unit="mm" value="0.69">0.69 mm</quantity>
, tooth-field length ca 
<quantity id="F867F7775126E2E69ACC2169D93A7BF0" metricMagnitude="-4" metricUnit="m" metricValue="6.7" unit="mm" value="0.67">0.67 mm</quantity>
(measurements from the cleared preparation; Fig. 
<figureCitation id="900B5636D8F35DCF57EA81AA422B2B18" 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">24 a</figureCitation>
). Length of longest teeth on endophallus 54–65 μm. Parameres with the sclerotised strut running from the base just mediad of the outer lateral margin to the inner apical angle. Apical soft pad of parameres rounded apically, not widened.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EVNAM" type="remarks">
<paragraph id="3C0B048FEC058E83DE0194F32D8AA5AC">
<heading id="6065637E25049C31C7F588CAFDAED703" reason="title">Remarks.</heading>
</paragraph>
<paragraph id="BB19D14C64B78E9CC7F64736F76A0DB2">
Chromosomally verified material is from the New Forest, 
<collectingCountry id="48F944AC107B59D664992452DD4FAA91" name="United Kingdom">England</collectingCountry>
and Södermannland, 
<collectingCountry id="3204A0A4B8AD94DC03DD28CFDEC0510A" name="Sweden">Sweden</collectingCountry>
. Material from Berditsino, 
<collectingCountry id="F3D6116D81DB64C1AAEBE308C310B531" name="Russia">Russia</collectingCountry>
(
<named-content id="139FF33B8244C9E63F24AC816F77DEDF" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/naturhistorisches-museum-wien-0" xlink_title="Naturhistorisches Museum, Wien">
<collectionCode id="3E3E90B7534DC8F65575E1DE7DCA7759" country="Austria" httpUri="http://grbio.org/cool/c7yt-hh77" name="Naturhistorisches Museum, Wien" type="">NMW</collectionCode>
</named-content>
) is taken to belong to this species. In the New Forest 
<taxonomicName id="E6A2DA8934E4B57282EECE3C93EEDB63" authorityName="Illiger" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="81E58BA875DA299C4BCE3C5CCBA8C09B" italics="true">A. niger</emphasis>
</taxonomicName>
is typically taken at the edges of pools and ditches in April-May, but when large numbers are taken it is almost always due to inwash from the surrounding grassland after heavy rain (Fig. 
<figureCitation id="4843AAAFA2F35CF435D7A5DA0FA3D9BB" 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">27 a</figureCitation>
). In Britain, outside of the New Forest, 
<taxonomicName id="0319229C7AB21B19A248F5C4095598E7" authorityName="Illiger" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="1443E80F540700170202D7E10DB5F983" italics="true">A. niger</emphasis>
</taxonomicName>
has recently been found at Inkpen Common near Hungerford in Berkshire (
<bibRefCitation id="9D99333060349A9E2C76D1043DDF0A2A" author="Mann and Garvey" firstAuthor="Mann" issue="3" journalOrPublisher="Coleopterist" pagination="138-142" refId="B31" refString="Mann DJ, Garvey L (2014) Aphodius (Liothorax) niger (Illiger) (Scarabaeidae: Aphodiinae) inside and outside the New Forest. Coleopterist 23 (3): 138–142." title="Aphodius (Liothorax) niger (Illiger) (Scarabaeidae: Aphodiinae) inside and outside the New Forest." volume="23" year="2014">Mann and Garvey 2014</bibRefCitation>
). In the Swedish locality the beetles were burrowing in trampled organic mud (H. - E. Wanntorp, pers. comm., 
<date id="666FA303C9EA507D6EAC8AFF3BA6BE13" value="2011-04">April 2011</date>
) (Fig. 
<figureCitation id="8AAED65C8A8F11EBD0035402BA322F35" 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">27 c</figureCitation>
). Very occasionally 
<taxonomicName id="D0CBD20F5B17ED378534DA597502BF9A" authorityName="Illiger" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="E39696AD35ABBEB633BD0D167731D7CE" italics="true">A. niger</emphasis>
</taxonomicName>
is taken in dung, as by J. Bergsten in the New Forest in 1999, and sometimes in dry areas of the Swedish island of Öland (H. Lundqvist, pers. comm., 
<date id="10029805B3E4665A6C510A85BCC41FB6" value="2011-04">April 2011</date>
). These are rare, random occurrences and may sometimes result from the beetle taking shelter from drought. One further feature of the occurrence of 
<taxonomicName id="25844280BC3D774588E82C1286712541" authorityName="Illiger" authorityYear="1798" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="niger">
<emphasis id="29670FC19070E7195736D719A47C6B1F" italics="true">A. niger</emphasis>
</taxonomicName>
is worthy of comment – it shows dramatic fluctuations in abundance. Thus, in 2002 it was very abundant in its New Forest localities whereas more recently RBA has taken it only in low numbers. Similarly, H. - E. Wanntorp describes the species as being in very high numbers in the locality where he usually only took a few specimens (Fig. 
<figureCitation id="3D0E405B95D288BD176781F5C1878820" 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>
).
</paragraph>
</subSubSection>
</treatment>
</document>