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 ) villosus ( Burmeister, 1847 ) ( Figs. 2D , 4E , 5E , 6B , 11G , 12 F, L , 14D , 15T , 18L , 26I ; 35 ) Podalgus villosus Burmeister, 1847: 120 . Original combination. Male lectotype designated by Endrödi (1969: 58) (MLUH) “villosus / Demophon Dej / Chili Patag. // Lectotypus / Ligyrus / ( Podalgus / villosus Burm. / Endrody” Type locality: Chile . Oryctes nitidicollis Solier, 1851: 79 . Synonym. Type locality: Santa Rosa , Santiago , and Coquimbo ( Chile ) . Type specimens not examined. The original description, including some illustrations, and the distribution coincides with L. villosus . Description. Habitus as in Fig. 26I . Length 14.8–18.6 mm ; humeral width 7.8–10.1 mm . Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex, only with sparse, small punctures. Frontoclypeal region with a carina, widely interrupted at middle. Ocular canthus acute, with 6 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus short, base 3 times wider than apex. Clypeal teeth triangular, separated by a tooth diameter. Mandible with 2 apical and 1 lateral, rounded tooth, tooth 1 longer and narrower than tooth 2 ( Fig. 5E ). Maxilla subrectangular; galea with 4 teeth ( Fig. 4E ). Apex of labrum rounded ( Fig. 6B ). Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface without or with minute punctures. Apex without tubercle (with a small swelling) or fovea. Elytra: First interval punctate, punctures similar to 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 11 transverse, parallel, long carinae. Pygidial 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 ( Fig. 11G ). Protarsus of male simple, inner claw simple. Metatibia triangular. Apex of metatibia entire to slightly crenulate, with 17–20 spinules ( Fig. 12L ). 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. Male genitalia: Spiculum gastrale thin, apex truncate ( Fig. 14D ). Parameres with apical 4th narrowed, without minute spines; latero-ventral teeth short, triangular ( Figs. 15T , 18L ). Diagnosis. Ligyrus villosus can be recognized by a short clypeus, frontal carinae widely interrupted at middle; pronotum without tubercle or fovea (only with a small swelling) ( Fig. 26I ); pronotal punctation absent or minute; metatibia strongly triangular ( Fig. 12L ); coxite surface flat; parameres with apical 4th narrowed, without minute spines; lateral teeth of parameres short, triangular ( Figs. 15T , 18L ). Geographic distribution. Ligyrus villosus is found in Argentina , Peru , and Chile ( Neita-Moreno & Ratcliffe 2017 ). Locality records ( Fig. 35 ). 190 specimens examined from EMAC, IEXA, FSCA, MEKRB, MLUH, MN/UFRJ, MZUSP. Records from Argentina are taken from Neita-Moreno & Ratcliffe (2017) . ARGENTINA (17). Neuquén (17): Colón Cura ; Quilquihue ; Rinconada ; Zapala. CHILE (179). Aconcagua (2): Los Molles ( 10 km S Pichidangui ). Araucania (2): Temuco ; Villarica. Atacama (1): Vallenar ( 15 km N). Biobío (4): Concepción ; Hualgui ; Salto de Laja. Coquimbo (10): La Serena ; Los Vilos ; Tongoy ; Ñague ; Vicuña. Los Ríos (1): Rincón de Piedra ( 20 km SE Valdivia ). Maule (37): Curanipe ; El Coigo ; El Pantanillo ( 17 km SE; Constitución) ; Los Niches ; Los Quenes (6 Km S ) ; Pr. Forel Carrizalillo ; Rio Teno ( 40 km E Curico ) ; Talca. Ñuble (29): Portezuelo ; Recinto ; Rio Pinto ( E of Chilian ). O’Higgins (2): Olivar Alto. Santiago (20): Cerro San Cristóbal ; Cordillera Reserva Rio Clarillos ; El Toyo ; Los Condes ; Maipú ; Pilay ; Punta Yeso. Valparaiso (42): Angol ; Limache ; Mantagua ; Rio Blanco ; Quillota ; San Antonio. No data (26). PERU (3). Arequipa (2): Socabaya, Lara. Lima (1): Lima. Natural history. Gutiérrez (1945) observed that L. villosus was nocturnal, sometimes highly abundant. Males in Chile flew searching for females and copulation occurred under the soil, in the same hole from where the females emerged. This species occasionally attacks the roots of sunflower, alfalfa, and maize, but not becoming a significant pest ( Gutiérrez 1945 ). It is a common species in blueberry crops, where larvae feed on roots causing weakening or even death of young plants ( Cisternas 2013 ).