treatments-xml/data/21/73/75/21737574B2615B2FA919858AE9561D34.xml

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<mods:title id="3CB4D93E62061376AEA3C258B92C15D5">Towards a revision of the Palaearctic species of Aphodius Hellwig, 1798, subgenus Liothorax Motschulsky, 1860 (Coleoptera, Scarabaeidae, Aphodiinae)</mods:title>
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<mods:namePart id="F7D51DB720C5661E4DE52EB6BEBF985E">Maté, Jason F.</mods:namePart>
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<mods:affiliation id="F58D58292F5F82276D9D93690C0D5C9D">Biomedical Imaging Unit, Level B South Block, Mail point 12, General Hospital, Southampton SO 16 6 YD, UK</mods:affiliation>
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(
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<emphasis id="D9E37088BD828B6F35D392E48F5FDDF7" italics="true">rusakovi</emphasis>
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<figureCitation id="1193F73A663D7E07866CFDE8617E90A1" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. a – k whole beetles a A. plagiatus, CZ, Moravia b A. plagiatus, MG, Mongolia c A. jakutorum, holotype, RU, East Siberia d L. rodrigoi sp. nov., holotype SP, Aranjuez e A. rutilipennis (A. ressli, holotype) TR, Hatay, Iskenderun f A. rutilipennis, large ♀, CY, Akrotiri g A. rutilipennis (A. cypricola, holotype) CY h A. chellala sp. nov., holotype, AG, Chellala i A. rusakovi, ♂ paratype, RU, Orenburg j A. discoides, neotype (A. bytinskisalzi, holotype) IS, Kuneitra k A. discoides, black form from TR, Karakurt. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure2" httpUri="https://binary.pensoft.net/fig/1097460">Figs 2 i</figureCitation>
, 
<figureCitation id="1261AC824F2FFC02C54056360436A0F1" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. a – p A. plagiatus group, heads and pronota a – c A. plagiatus plagiatus a GB, Norfolk b CZ, Moravia, Dobre Pole c RU, St Petersburg d A. p. sinoplagiatus ssp. nov., paratype, CH, Qinghai, Gangca e – g A. discoides e neotype IS Kuneitra f TR, Kyzil Dag g Turkey Muş h A. discoides, black ♂, TR, Karakurt i – k A. rutilipennis i A. ressli, holotype, TR, Hatay j CY, Akrotiri k A. cypricola, holotype, CY l A. chellala sp. nov., holotype, AG, Chellala m A. rusakovi, paratype ♂, RU, Orenburg n A. ballerioi sp. nov., paratype ♂, TR, Rize o, p L. kraatzi o RU, Astrakhan p SR. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure5" httpUri="https://binary.pensoft.net/fig/1097463">
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<figureCitation id="88D723A88A1BBCBFF2C4D9D52360CE9F" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Aphodius plagiatus group, pronotal bases 2 a A. discoides, TR b A. kraatzi, SK c L. rusakovi, paratype, RU, Volgograd oblast’." figureDoi="10.3897/zookeys.1207.117225.figure14" httpUri="https://binary.pensoft.net/fig/1097474">14 c</figureCitation>
, 
<figureCitation id="535836C2F2DBD1989FF1BB1033E8920F" 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 f</figureCitation>
, 
<figureCitation id="161AC2BA52679385B3B0E72B4884F003" captionStart="Figure 20" captionStartId="F20" captionText="Figure 20. Metaventrites, A. plagiatus group a A. plagiatus plagiatus, GB, Studland, ♂ b A. p. sinoplagiatus ssp. nov., paratype, CH, Gansu, ♀ c, d A. rutilipennis, CY c ♂ d ♀ e, f A. chellala sp. nov., paratypes, AG e ♂ f ♀ g, h A. kraatzi, RU, Orenburg, ♂ i A. kraatzi, SK, ♀ j A. rodrigoi sp. nov., holotype, ♂, SP, Provinicia de Madrid, Aranjuez k, l A. discoides k neotype ♂, IS, Kuneitra l TR, ♀ m, n A. rusakovi m paratype, ♂, RU, Orenburg n ♀ RU, Volgograd oblast’. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure20" httpUri="https://binary.pensoft.net/fig/1097486">
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<figureCitation id="2BD1FF7CC013DEED03C6A8D9C44D5350" 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 h, v</figureCitation>
, 
<figureCitation id="C6322BF941514CB3F8137B40418829B7" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Aphodius plagiatus group aedeagi a – c A. plagiatus plagiatus, GB, Studland, Dorset, aedeagus with extruded endophallus a, b SEM images a whole aedeagus b endophallus in more detail c stacked photograph of endophallus d – g A. discoides, neotype (Aphodius bytinskisalzi, holotype, IS, Kuneitra) d, e, e ’ SEM images f, g stacked photographs h – n cleared preparations mounted on slides h A. p. plagiatus, GB, Norfolk i A. p. sinoplagiatus? “ China, Pekin ”, Fry collection i ’ teeth on neck of endophallus in the region marked with an arrow in i j, k A. plagiatus sinoplagiatus ssp. nov., paratypes j CH, Gangca k CH, Tsaidam l Aphodius jakutorum Balthasar, lectotype, RU, East Siberia m A. rodrigoi sp. nov., holotype, SP, Aranjuez, dissected aedeagus m ’ endophallus m ” endophallic teeth in area indicated by arrow in m ’ at higher magnification n A. discoides, TR, Karacadag n ’ teeth on neck of endophallus, in area indicated by arrow in n at higher magnification o A. discoides, black specimen from TR, Karakurt, o ’ teeth on neck of endophallus, in area indicated by arrow in o at higher magnification p, p ’ A. rusakovi p paratype, RU, Orenburg p ’ RU Volgograd obl. Fastov, paramere, lateral q – v SEM images and stacked photographs q, r A. rutilipennis, CY, with endophallus inflated q SEM image q ’ (bottom right of plate) dorsal surface of endophallus in area indicated by arrow in q at higher magnification, to show the hair-like setae r stacked photograph of the same specimen s, s ’ A. chellala sp. nov., paratype with endophallus extruded s stacked photograph s ’, s ”, s ” ’ SEMs of the same specimen s ”, s ” ’ at higher magnifications t, u L. chellala sp. nov., holotype with the endophallus extruded t stacked photograph u SEM image v – z photographs of cleared specimens mounted in DMHF on slides v, w A. rutilipennis v CY w TR w ’ teeth on neck of endophallus, in area indicated by arrow in w at higher magnification x A. chellala sp. nov., paratype, AG, Taguin (La Smala) x ’ teeth on neck of endophallus, in area indicated by arrow in x at higher magnification y, z A. kraatzi y KZ, Syr Darya z AF, Kabul z ’ teeth on neck of endophallus, in area indicated by arrow in z at higher magnification Scale bar 1: 1 mm (a, d, f – t, v – z); 0.4 mm (n ’, o ’, s ”, w ’, x ’, z ’); 0.17 mm (e ’, m ”, q’ s ” ’). Scale bar 2: 1 mm (b, c, e, g and s ”)." figureDoi="10.3897/zookeys.1207.117225.figure22" httpUri="https://binary.pensoft.net/fig/1097490">
22 p, p ’, 
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<taxonomicName id="3BFBE2C39A38CF1B9C714E4C877A807F" ID-CoL="FGDV" authority="Gusakov, 2004: 6" authorityName="Gusakov" authorityPageNumber="6" authorityYear="2004" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="rusakovi" subGenus="Liothorax">
<emphasis id="6F2F9141C423322A11CC458ED723E27E" italics="true">Aphodius</emphasis>
(
<emphasis id="49DD720BC28E5F8DD46CF4B8E3ECE609" italics="true">Liothorax</emphasis>
) 
<emphasis id="14B89E83A985AA47DF37DB94F8B8CFC5" italics="true">rusakovi</emphasis>
<comment id="2CFF2E27368963F4E3D9887662C6DE4F">
<bibRefCitation id="D005895BA1507387AFBFCCC6212E2CA3" author="Gusakov AA" refId="B19" refString="Gusakov AA (2004) Novye vidy plastinchtousykh zhukov palearcticheskoj fauny. [New species of scarab beetles from the Palaearctic region]. Izdatel'skij dom Ejdos, Kaluga, 1 - 38." year="2004">Gusakov, 2004: 6</bibRefCitation>
.
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<paragraph id="C68D0924CA74DF4A2EB002DFBE8EEC5B">
<heading id="A6D67A50C6D25E5BB2787F0E9C78EA48" reason="title">Type material examined.</heading>
</paragraph>
<paragraph id="2A8CE819E79688CC943661D3B6843477">
<specimenCount id="EA5DB39CBB2641B4A59461E5FF899195" count="2" type="generic" typeStatus="paratype">2 paratypes</specimenCount>
from the 
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oblast’, 
<collectingCountry id="599DC08D1547DDF816947AAF7FDCCB85" name="Russia">Russia</collectingCountry>
(
<named-content id="E4FB26A94043F8D46CB42D86168F9C13" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/natural-history-museum-london" xlink_title="The Natural History Museum, London">
<collectionCode id="FFA25036636F3C63C48B0127C47A5B39" 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>
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).
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<paragraph id="7D475BA7BB8323CCE37441804215D47A">
<heading id="58DF04B7BF598803B4971452C507D5D4" reason="title">Additional material examined.</heading>
</paragraph>
<paragraph id="D1E96CC23E59B389B3444C1FE8298DF6">
RU: 
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, 
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, 
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<geoCoordinate id="AF04EF649D17D70AEA0A73001875A585" degrees="54.056" direction="north" orientation="latitude" precision="55" value="54.056">54.056 ° N</geoCoordinate>
, 
<geoCoordinate id="8071E3DC453CBC785C573AC393B5F34A" degrees="48.519" direction="east" orientation="longitude" precision="55" value="48.519">48.519 ° E</geoCoordinate>
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(
<named-content id="BFB2EF81AA0FB56DC88E38F8B6CE5AD3" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/russian-academy-sciences-zoological-institute-zoological-museum" xlink_title="Zoological Institute, Russian Academy of Sciences St Petersburg">
<collectionCode id="AD0B69580852A96BC9E49CD5672ECBD1" country="Russia" httpUri="http://grbio.org/cool/z2y8-2jfr" name="Russian Academy of Sciences, Zoological Institute, Zoological Museum" type="Museum">ZIN</collectionCode>
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, 
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oblast’, 
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<geoCoordinate id="9DD4AE6CDFD61AD93FBC3F4E8377929E" degrees="48.7" direction="north" orientation="latitude" precision="5555" value="48.7">48.7 ° N</geoCoordinate>
, 
<geoCoordinate id="908876044E52D778473C3F11057AD3AD" degrees="44.5" direction="east" orientation="longitude" precision="5555" value="44.5">44.5 ° E</geoCoordinate>
</named-content>
, Fastov. leg. Grebennikov. (
<named-content id="98A103EB47901C83DD0E29A1497280A3" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/russian-academy-sciences-zoological-institute-zoological-museum" xlink_title="Zoological Institute, Russian Academy of Sciences St Petersburg">
<collectionCode id="0A1BAAE126EFD7374A9C30F67252F4D9" country="Russia" httpUri="http://grbio.org/cool/z2y8-2jfr" name="Russian Academy of Sciences, Zoological Institute, Zoological Museum" type="Museum">ZIN</collectionCode>
</named-content>
).
</paragraph>
<paragraph id="D0C443892BBB8128C480748EEE0C76B1">
KZ or UZ 
<specimenCount id="314B84E8C706A6BD7A918209FF882886" count="1" type="female">1 ♀</specimenCount>
“ Syr-Darja. / B. v. Bodemeyer. ” (
<named-content id="F028F0277033614851C5EFE15F8B3747" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-prague-0" xlink_title="National Museum Prague">
<collectionCode id="7CC30681ED602863D264A1AE9EDA20A4" country="Czech Republic" httpUri="http://grbio.org/cool/fx6k-enrh" name="National Museum (Prague)" type="">NMP</collectionCode>
</named-content>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0E35HK" type="Differential diagnosis">
<paragraph id="1EDEF8169C3FAC90841E328357AA3AB6">
<heading id="F5A10CB70DA91619AD27A6571350A263" reason="title">Differential diagnosis.</heading>
</paragraph>
<paragraph id="707345FE00D77ACB0ABA68753181A4C0">
A conspicuously narrow parallel-sided species, comparable with 
<taxonomicName id="E97D62F61FE77F85ACEC36CB4D95D514" authorityName="Harold" authorityYear="1868" class="Insecta" family="Scarabaeidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" phylum="Arthropoda" rank="species" species="kraatzi">
<emphasis id="DE2EDB2D0E7F5AA030E0C7DA4F2BC759" italics="true">A. kraatzi</emphasis>
</taxonomicName>
but larger, length 
<quantity id="30C33B3FF2B5B6753E6AD9B8C78A328B" metricMagnitude="-3" metricUnit="m" metricValue="5.5" metricValueMax="6.5" metricValueMin="4.5" unit="mm" value="5.5" valueMax="6.5" valueMin="4.5">4.5–6.5 mm</quantity>
and base of pronotum completely bordered by a fine impressed line, this sometimes weaker medially (Fig. 
<figureCitation id="DC6C994C0E9538A4929561248170654A" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Aphodius plagiatus group, pronotal bases 2 a A. discoides, TR b A. kraatzi, SK c L. rusakovi, paratype, RU, Volgograd oblast’." figureDoi="10.3897/zookeys.1207.117225.figure14" httpUri="https://binary.pensoft.net/fig/1097474">14 c</figureCitation>
). Protibial spur of male strongly incurved apically (Fig. 
<figureCitation id="05B217CBF28F6362D976F330933DE6F5" 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 v</figureCitation>
). 
<specimenCount id="A5BD59A5D9ECE04AA37ABF81A2766590" count="1" type="female">♀</specimenCount>
with spermatheca larger (Fig. 
<figureCitation id="A4A7474F10150B2C1B70FEA0E36685D7" 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="611561B709AFAEF9E0289FE561DC355B" metricMagnitude="1" metricUnit="m" metricValue="2.6" unit="m" value="26.0">26 m</quantity>
, n
</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EZ6HK" type="redescription">
<paragraph id="FA9496A02D6837A3DF418826E837395C">
<heading id="A6AE2A58AF32242A1B0154D232271619" reason="title">Redescription.</heading>
</paragraph>
<paragraph id="751A7F311DCA72D50DE913E0E3E8CE49">
General appearance (Fig. 
<figureCitation id="8462DB43FADC37D1834F6819227EF0CD" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. a – k whole beetles a A. plagiatus, CZ, Moravia b A. plagiatus, MG, Mongolia c A. jakutorum, holotype, RU, East Siberia d L. rodrigoi sp. nov., holotype SP, Aranjuez e A. rutilipennis (A. ressli, holotype) TR, Hatay, Iskenderun f A. rutilipennis, large ♀, CY, Akrotiri g A. rutilipennis (A. cypricola, holotype) CY h A. chellala sp. nov., holotype, AG, Chellala i A. rusakovi, ♂ paratype, RU, Orenburg j A. discoides, neotype (A. bytinskisalzi, holotype) IS, Kuneitra k A. discoides, black form from TR, Karakurt. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure2" httpUri="https://binary.pensoft.net/fig/1097460">2 i</figureCitation>
). Glossy black, length 
<quantity id="1486BAAD1461EB669022518EEA2BC2ED" metricMagnitude="-3" metricUnit="m" metricValue="5.1" metricValueMax="5.5" metricValueMin="4.7" unit="mm" value="5.1" valueMax="5.5" valueMin="4.7">4.7–5.5 mm</quantity>
, width 
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. Head fairly closely punctate, sometimes slightly rugose on the frons and anterolateral parts of the clypeus, and occasionally rugose over all except the central part of the clypeal bulge. Double punctation scarcely apparent except when the punctation is sparser on the clypeal bulge. Clypeus emarginated anteriorly, edges of this emargination bluntly rounded (head and pronotum: Fig. 
<figureCitation id="C8FE247D01F0F4E4491F0D42DF612DB7" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. a – p A. plagiatus group, heads and pronota a – c A. plagiatus plagiatus a GB, Norfolk b CZ, Moravia, Dobre Pole c RU, St Petersburg d A. p. sinoplagiatus ssp. nov., paratype, CH, Qinghai, Gangca e – g A. discoides e neotype IS Kuneitra f TR, Kyzil Dag g Turkey Muş h A. discoides, black ♂, TR, Karakurt i – k A. rutilipennis i A. ressli, holotype, TR, Hatay j CY, Akrotiri k A. cypricola, holotype, CY l A. chellala sp. nov., holotype, AG, Chellala m A. rusakovi, paratype ♂, RU, Orenburg n A. ballerioi sp. nov., paratype ♂, TR, Rize o, p L. kraatzi o RU, Astrakhan p SR. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure5" httpUri="https://binary.pensoft.net/fig/1097463">
<quantity id="329FA1ACCC274D268075F13774869D0F" metricMagnitude="0" metricUnit="m" metricValue="5.0" unit="m" value="5.0">5 m</quantity>
</figureCitation>
). Frontoclypeal suture interrupted medially in 
<specimenCount id="B809864FDFE2A05102E3C2864992CEFB" count="1" type="female">♀</specimenCount>
, in 
<specimenCount id="96C89D77A9396194E36816CCA395361C" count="1" type="male">♂</specimenCount>
sometimes with a very weak tubercle, viewed with suitable lighting (Fig. 
<figureCitation id="B645B86BDE592FC9CFAB4847E9C60362" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. a – k whole beetles a A. plagiatus, CZ, Moravia b A. plagiatus, MG, Mongolia c A. jakutorum, holotype, RU, East Siberia d L. rodrigoi sp. nov., holotype SP, Aranjuez e A. rutilipennis (A. ressli, holotype) TR, Hatay, Iskenderun f A. rutilipennis, large ♀, CY, Akrotiri g A. rutilipennis (A. cypricola, holotype) CY h A. chellala sp. nov., holotype, AG, Chellala i A. rusakovi, ♂ paratype, RU, Orenburg j A. discoides, neotype (A. bytinskisalzi, holotype) IS, Kuneitra k A. discoides, black form from TR, Karakurt. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure2" httpUri="https://binary.pensoft.net/fig/1097460">2 i</figureCitation>
). Epipharynx (Fig. 
<figureCitation id="7730FC8B477D437B1A148A0E724E6705" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. a – k epipharynxes a – f A. plagiatus plagiatus a, b GB, Hunstanton, Norfolk c CZ, Moravia, Dobré Pole d RU, Pavlovsk, St Petersburg e A. p. sinoplagiatus ssp. nov., paratype, CH, Gangca, Qinghai f A. p. sinoplagiatus?, CH, “ Pekin, coll. Fry ” g A. rodrigoi sp. nov., holotype, SP h A. discoides, TR i A. rusakovi, paratype, RU, Orenburg j A. rutilipennis, CY, Akrotiri k A. chellala, sp. nov., paratype, AG, Chellala. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure7" httpUri="https://binary.pensoft.net/fig/1097466">7 i</figureCitation>
) with anterior margin of clithra almost straight either side of the median tylus and the mesoepitorma scarcely darkened. Prophobae rather strong, grouped towards the margins of the mesoepitorma. Chaetopedia three or four each side. Pronotum highly arched transversely, weakly so longitudinally. Entire lateral margins visible from above (Figs 
<figureCitation id="AB54FD05E608ED674281E1F5F416E462" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. a – k whole beetles a A. plagiatus, CZ, Moravia b A. plagiatus, MG, Mongolia c A. jakutorum, holotype, RU, East Siberia d L. rodrigoi sp. nov., holotype SP, Aranjuez e A. rutilipennis (A. ressli, holotype) TR, Hatay, Iskenderun f A. rutilipennis, large ♀, CY, Akrotiri g A. rutilipennis (A. cypricola, holotype) CY h A. chellala sp. nov., holotype, AG, Chellala i A. rusakovi, ♂ paratype, RU, Orenburg j A. discoides, neotype (A. bytinskisalzi, holotype) IS, Kuneitra k A. discoides, black form from TR, Karakurt. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure2" httpUri="https://binary.pensoft.net/fig/1097460">2 i</figureCitation>
, 
<figureCitation id="69691973E81D60F1516A6D091C9C95E3" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. a – p A. plagiatus group, heads and pronota a – c A. plagiatus plagiatus a GB, Norfolk b CZ, Moravia, Dobre Pole c RU, St Petersburg d A. p. sinoplagiatus ssp. nov., paratype, CH, Qinghai, Gangca e – g A. discoides e neotype IS Kuneitra f TR, Kyzil Dag g Turkey Muş h A. discoides, black ♂, TR, Karakurt i – k A. rutilipennis i A. ressli, holotype, TR, Hatay j CY, Akrotiri k A. cypricola, holotype, CY l A. chellala sp. nov., holotype, AG, Chellala m A. rusakovi, paratype ♂, RU, Orenburg n A. ballerioi sp. nov., paratype ♂, TR, Rize o, p L. kraatzi o RU, Astrakhan p SR. Scale bar: 1 mm." figureDoi="10.3897/zookeys.1207.117225.figure5" httpUri="https://binary.pensoft.net/fig/1097463">
<quantity id="DA2D2B7A1A201990E9D92E0D8A569498" metricMagnitude="0" metricUnit="m" metricValue="5.0" unit="m" value="5.0">5 m</quantity>
</figureCitation>
). Surface with double punctation, this often close though sometimes sparser on the disc. In some specimens the punctures are deeply impressed, so the pronotal surface does not appear smooth. Basal margin entirely bordered, the border rather heavy (Fig. 
<figureCitation id="29084C5F72D9C33FD0E6DE78E472E322" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Aphodius plagiatus group, pronotal bases 2 a A. discoides, TR b A. kraatzi, SK c L. rusakovi, paratype, RU, Volgograd oblast’." figureDoi="10.3897/zookeys.1207.117225.figure14" httpUri="https://binary.pensoft.net/fig/1097474">14 c</figureCitation>
). Elytra noticeably elongate, parallel-sided. Striae narrow but rather deep, strial punctures separated by ca 4 × their diameters. Interstices 6–8 × the width of the striae, with distinct sparse fine punctation and fine reticulation (Fig. 
<figureCitation id="BF6A3105F53A90246D48407A484CB0B4" 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 f</figureCitation>
). Apex bluntly rounded. Metaventrite (Fig. 
<figureCitation id="683303A2244539AD8ACAB00696F97466" captionStart="Figure 20" captionStartId="F20" captionText="Figure 20. Metaventrites, A. plagiatus group a A. plagiatus plagiatus, GB, Studland, ♂ b A. p. sinoplagiatus ssp. nov., paratype, CH, Gansu, ♀ c, d A. rutilipennis, CY c ♂ d ♀ e, f A. chellala sp. nov., paratypes, AG e ♂ f ♀ g, h A. kraatzi, RU, Orenburg, ♂ i A. kraatzi, SK, ♀ j A. rodrigoi sp. nov., holotype, ♂, SP, Provinicia de Madrid, Aranjuez k, l A. discoides k neotype ♂, IS, Kuneitra l TR, ♀ m, n A. rusakovi m paratype, ♂, RU, Orenburg n ♀ RU, Volgograd oblast’. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.1207.117225.figure20" httpUri="https://binary.pensoft.net/fig/1097486">
<quantity id="C4060086E179FBAA3BB0C4E67E5EBF90" metricMagnitude="1" metricUnit="m" metricValue="2.0" unit="m" value="20.0">20 m</quantity>
, n
</figureCitation>
) with median diamond- shaped plate fairly strongly punctate and no obvious sexual dimorphism. Legs with longer spur of mid tibiae slightly longer than the basal segment (Fig. 
<figureCitation id="C38DC50AFF4B6AA4A61326236D3D2D86" 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 h</figureCitation>
), and in males the spur of the anterior tibiae is distinctly hooked (Fig. 
<figureCitation id="A65AAE19066602C189CC7CB9A67F379A" 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 v</figureCitation>
).
</paragraph>
<paragraph id="012164FF37B7A118BF803037643F5E7F">
Aedeagus (Fig. 
<figureCitation id="961EA71E795467B562D8E1C679A7CC33" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Aphodius plagiatus group aedeagi a – c A. plagiatus plagiatus, GB, Studland, Dorset, aedeagus with extruded endophallus a, b SEM images a whole aedeagus b endophallus in more detail c stacked photograph of endophallus d – g A. discoides, neotype (Aphodius bytinskisalzi, holotype, IS, Kuneitra) d, e, e ’ SEM images f, g stacked photographs h – n cleared preparations mounted on slides h A. p. plagiatus, GB, Norfolk i A. p. sinoplagiatus? “ China, Pekin ”, Fry collection i ’ teeth on neck of endophallus in the region marked with an arrow in i j, k A. plagiatus sinoplagiatus ssp. nov., paratypes j CH, Gangca k CH, Tsaidam l Aphodius jakutorum Balthasar, lectotype, RU, East Siberia m A. rodrigoi sp. nov., holotype, SP, Aranjuez, dissected aedeagus m ’ endophallus m ” endophallic teeth in area indicated by arrow in m ’ at higher magnification n A. discoides, TR, Karacadag n ’ teeth on neck of endophallus, in area indicated by arrow in n at higher magnification o A. discoides, black specimen from TR, Karakurt, o ’ teeth on neck of endophallus, in area indicated by arrow in o at higher magnification p, p ’ A. rusakovi p paratype, RU, Orenburg p ’ RU Volgograd obl. Fastov, paramere, lateral q – v SEM images and stacked photographs q, r A. rutilipennis, CY, with endophallus inflated q SEM image q ’ (bottom right of plate) dorsal surface of endophallus in area indicated by arrow in q at higher magnification, to show the hair-like setae r stacked photograph of the same specimen s, s ’ A. chellala sp. nov., paratype with endophallus extruded s stacked photograph s ’, s ”, s ” ’ SEMs of the same specimen s ”, s ” ’ at higher magnifications t, u L. chellala sp. nov., holotype with the endophallus extruded t stacked photograph u SEM image v – z photographs of cleared specimens mounted in DMHF on slides v, w A. rutilipennis v CY w TR w ’ teeth on neck of endophallus, in area indicated by arrow in w at higher magnification x A. chellala sp. nov., paratype, AG, Taguin (La Smala) x ’ teeth on neck of endophallus, in area indicated by arrow in x at higher magnification y, z A. kraatzi y KZ, Syr Darya z AF, Kabul z ’ teeth on neck of endophallus, in area indicated by arrow in z at higher magnification Scale bar 1: 1 mm (a, d, f – t, v – z); 0.4 mm (n ’, o ’, s ”, w ’, x ’, z ’); 0.17 mm (e ’, m ”, q’ s ” ’). Scale bar 2: 1 mm (b, c, e, g and s ”)." figureDoi="10.3897/zookeys.1207.117225.figure22" httpUri="https://binary.pensoft.net/fig/1097490">22 p, p</figureCitation>
’) length ca 
<quantity id="1CA0F6241E4FE40981ED3596C08D7E92" metricMagnitude="-3" metricUnit="m" metricValue="1.1" unit="mm" value="1.1">1.1 mm</quantity>
, parameres relatively short and blunt-ended, somewhat downturned apically. Endophallus with scales but no hairs or bristles.
</paragraph>
</subSubSection>
<subSubSection id="SECID0ESBAM" type="remarks">
<paragraph id="60C81B5729902C52EDFBBA9220C6725D">
<heading id="19ADDF39C08444C6855FFA32B39F61C7" reason="title">Remarks.</heading>
</paragraph>
<paragraph id="4B20F95518D297C576769908998D53D3">
Distributed mainly in southern 
<collectingCountry id="98CB042226EFC702D46B508712C7BFB9" name="Russia">Russia</collectingCountry>
, 
<collectingCountry id="EC485BF06482FC3A10B8B5D758E3CD6F" name="Ukraine">Ukraine</collectingCountry>
, and 
<collectingCountry id="14FE5E6BDF4F1E5093D63D123CAE3354" name="Kazakhstan">Kazakhstan</collectingCountry>
(Fig. 
<figureCitation id="36B35940EC1AA454170EF40185DDE9FE" 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 b, c</figureCitation>
). 
<bibRefCitation id="38DCBDBB0ECD5E56984A544E19C6630A" author="Gusakov AA" firstAuthor="Gusakov" refId="B19" refString="Gusakov AA (2004) Novye vidy plastinchtousykh zhukov palearcticheskoj fauny. [New species of scarab beetles from the Palaearctic region]. Izdatel’skij dom Ejdos, Kaluga, 1–38." year="2004">Gusakov (2004)</bibRefCitation>
lists the 
<typeStatus id="64548D007E0930FABDFE2A45919D1C68" type="holotype">holotype</typeStatus>
and 
<specimenCount id="A73EB2F838D65BFED7ADA34480F4E851" count="14" type="generic" typeStatus="paratype">14 paratypes</specimenCount>
as having been collected in a dried-out rivulet at Ilek, 
<quantity id="72455446C9D6F934C5AA9EC1A3E404A7" metricMagnitude="5" metricUnit="m" metricValue="1.2" unit="km" value="120.0">120 km</quantity>
SSW of Orenburg on the southern Urals, with two further 
<typeStatus id="F22B18B79F3373D1048C8DEA03E9B1BB" type="paratype">paratypes</typeStatus>
from the Rostov and 
<collectingRegion id="7D3C9217492229605412455EFFC89EF4" country="Ukraine" name="Kharkiv">Kharkov</collectingRegion>
districts of 
<collectingCountry id="972AE048B76F9A8884B6FDFB9663DEAE" name="Ukraine">Ukraine</collectingCountry>
and four more from the Temirsky region and Uralsk oblast’ in 
<collectingCountry id="9EB703242FC9CF4B043F5DF2A43D2C70" name="Kazakhstan">Kazakhstan</collectingCountry>
The Ulyanovsk specimen (
<named-content id="8C5DA249CABE12F4AB0BEA97D93F792D" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/russian-academy-sciences-zoological-institute-zoological-museum" xlink_title="Zoological Institute, Russian Academy of Sciences St Petersburg">
<collectionCode id="745E977EDC14FDD1D81D7123BAD6E922" country="Russia" httpUri="http://grbio.org/cool/z2y8-2jfr" name="Russian Academy of Sciences, Zoological Institute, Zoological Museum" type="Museum">ZIN</collectionCode>
</named-content>
) is labelled as having been taken in a damp saline area (solonchak).
</paragraph>
</subSubSection>
</treatment>
</document>