Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision Author López-García, Margarita M. 0000-0003-2796-2931 margaralopezg@gmail.com Author Deloya, Cuauhtémoc 0000-0002-4774-140X cuauhtemoc.deloya@inecol.mx text Zootaxa 2022 2022-11-22 5211 1 1 119 http://dx.doi.org/10.11646/zootaxa.5211.1.1 journal article 202083 10.11646/zootaxa.5211.1.1 bb97b9dd-2acc-4244-8f30-72bef07b9ed9 1175-5326 7345007 5754769C-B747-4714-BDD9-7D5509D48BEB Ligyrus ( Ligyrus ) bidentulus Fairmaire, 1892 ( Figs. 1G , 13F , 15J , 18B , 25G ; 35 ) Ligyrus bidentulus Fairmaire, 1892: 244 . Male neotype designated by López-García & Deloya (2019: 128) (USNM) “ La Rioja / Tinogasta / v.965 Kohler // ARGENTINA / 1968 Colln. / J. Daguerre // Ligyrus bidentulus / Fairmaire, 1892 / NEOTYPE ”. Type locality: La Rioja , Argentina. Description. Habitus as in Fig. 25G . Length 12.1–19.3 mm ; humeral width 5.8–10.7 mm . Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and with only sparse, small punctures. Frontoclypeal region with a carina, narrowly interrupted at middle ( Fig. 1G ). Ocular canthus acute; with 8 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex ( Fig. 1G ). Clypeal teeth triangular, slightly separated ( Fig. 1G ). Mandible with 2 apical and 1 lateral, rounded tooth, tooth 1 longer and narrower than tooth 2. Maxilla subrectangular; galea with 3 dorsal and 2 ventral teeth. Apex of labrum straight to rounded. Interocular distance equal to 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex with small tubercle and shallow fovea. Elytra: First interval punctate, punctures similar to the other intervals. Inner surface of apex with small, transverse tubercles forming about 115 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 10 transverse, parallel, short carinae. Pygidium: Surface with small to large punctures, denser on base and anterior corners. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate; basal tooth slightly distant from others. Protarsus of male simple, inner claw simple. Metatibia triangular. Apex of metatibia entire, with 25–29 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subquadrate (as long as wide); surface flat. Subcoxite wider and subequal in length to coxite ( Fig. 13F ). Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with apical 4th narrowed, without minute spines; lateral ventral teeth short, triangular ( Figs. 15J , 18B ). Diagnosis. Ligyrus bidentulus can be recognized by the very long antennal club (2 times longer than antennomeres 2–7); clypeus long, strongly contracted towards apex ( Fig. 1G ); clypeal teeth triangular, long, and not separated; frontal carinae narrowly interrupted at middle ( Fig. 1G ); pronotum with small tubercle and narrow fovea ( Fig. 25G ); parameres with apical 4th narrowed, without minute spines; lateral teeth of parameres short, triangular ( Figs. 15J , 18B ). Distribution. Argentina ( Endrödi 1985 ) and Uruguay ( Neita-Moreno & Ratcliffe 2017 ). Locality records ( Fig. 35 ). 58 examined specimens from MERKB, MNHM, MNHN, USNM, ZMUB. Some records from Neita-Moreno & Ratcliffe (2017) . ARGENTINA (101). Buenos Aires (9): Bahía Blanca ; Buenos Aires ; Carmen de Patagones ; Necochea ; Olavarria ; San Blas ; Villa Gesell. Catamarca (3): Ancafri ; Sinquil. Chaco (1): Resistencia. Chubut (6): Dolavon ; Estación de Esquel ; Golfo de San Jorge ; Península Valdés. Entre Ríos (3): Gualeguaychu. Jujuy (1): Posta de Lozano. La Pampa (1): Chacharramendi. La Rioja (9): Jaguel ; La Unión ; Mascasin ; Patquía ; Tinogasta ; Villa Unión. Mendoza (19): Bardas Blancas ; Cerro Guadal ; Mendoza ; El Nihuil ( 60 km S) ; Las Heras ; La Valle. Misiones (1): Posadas. Neuquén (14): Añelo ; Bajada del Marucho ; Bajada del Agrio ; Collón-Curá ; Covunco ; Huncal ; La Negra ; La Pintada ; Picún Leufú ; Piedra del Águila ; Pilmatué ; Planicie Banderita ; Río Salado ; Zalapa-Catuto. Río Negro (15): Allen ; Catriel ; Cipolletti ; Conesa ; El Bolsón ; Fray Luís Beltrán ; General Gómez ; General Roca ; Paso Flores ; Piedra del Águila ; Río Negro ; San Antonio ; Valcheta ; Villa Regina. Salta (14): Cafayate (desierto) ; Coronel Olleros ; Las Lajitas ; Salta. San Luís (1): Paso Patria. Tucumán (1): Tafi de Valle. No data (3). URUGUAY (1). Canelones (1): Banda Oriental . Natural history. Adults are attracted to lights at night. Most records are from arid and desert areas.