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 Tomarus maternus ( Prell, 1937 ) ( Figs. 4G , 5H , 6E , 7G , 8I, M , 13I , 16K , 20B , 22C , 28B ; 46 ) Ligyrus maternus Prell 1937: 89 . Original combination. Male lectotype designated by Endrödi (1969: 70) (ZMHB) “Gf. Madre de Dios / III. Peru 500 m . // Tomarus / maternus Prell / ♂ -Type // Lectotypus / Ligyrus / maternus / Prell / Endrody”. Female paralectotype (ZMHB) “Gf. Madre de Dios / III. Peru 500 m . // Tomarus / maternus Prell / ♀ -Type // Paratypus / Ligyrus / maternus / Prell” Type locality: Madre de Dios , Peru . Male paralectotype “ Umbria / Guines Fluss / Columbien // Paratypus / maternus / Prell // Tomarus gyas / Erichson, 1848 / Det. López-García M.M. 2016” corresponds to a male of Tomarus gyas (aedeagus dissected) . Description. Habitus as in Fig. 28B . Length 24.0–25.0 mm; humeral width 12.0– 12.5 mm . Color dark reddish brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 transverse tubercles separated by about 3.0–3.5 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth triangular, separated by 1.0–1.5 tooth diameters. Mandible with 2 apical teeth and a lateral, slightly rounded tooth ( Fig. 5H ). Mentum abruptly constricted at apical 3rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.8 times an eye width. Pronotum: Surface smooth on disc, with minute punctures on lateral margins. Apical tubercle small, rounded, not visible in lateral view. Fovea shallow, striate and about 1/2 as wide as interocular distance. Scutellum: Surface with deep, small punctures forming 2 lines parallel to margins. Elytra: First interval punctate, punctures smaller than on other intervals, sutural stria complete. Inner surface of apex with rounded, small tubercles not forming parallel lines ( Fig. 8M ). Abdomen: Apex of tergite IV with a triangular wide area with small and large, irregular tubercles not forming defined lines ( Fig. 8I ). Pygidial surface rugose on basal 3rd and on lateral angles in male, nearly smooth in female; disc with minute punctures; apex regularly rounded. Venter: Apex of prosternal process flat, oval to rounded. Metasternum with minute setae. Legs: Protibia tridentate, rarely with an additional small, basal denticle. Metatibia with sides nearly parallel. Apex of metatibia crenulate, with 14–17 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 2 pairs of dorso-lateral teeth; the main teeth short, wide, acute; the secondary teeth rounded ( Figs. 16K , 20B ). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 8 spine-like accessory lamellae, without granules at base ( Fig. 22C ). Diagnosis. Tomarus maternus can be recognized by the nearly impunctate pronotum; small pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1/3 as wide as interocular distance); scutellum with deep, small punctures forming 2 lines parallel to margins; protibia tridentate; and apex of metatibia with 14–17 spinules; parameres with 2 pairs of dorsal lateral teeth, the main teeth short, wide, acute, and the secondary teeth rounded ( Figs. 16K , 20B ); spine-like accessory lamellae of internal sac without granules at base ( Fig. 22C ). Distribution. Bolivia , Peru , Ecuador , Colombia , Venezuela , Panama , and Costa Rica . Locality records ( Fig. 46 ). 82 examined specimens from FSCA, MERKB, UNSM, USAAC, USNM, and ZMHB. Some records from CMNC and Endrödi (1969) , Ratcliffe (2003) , and Escalona & Joly (2006) . BOLIVIA (10). Beni (2): Rurrenabaque. Cochabamba (4): Chaparé. La Paz (2): Coroico, Guanay; Río Bopi. Santa Cruz (2): Buena Vista. COLOMBIA (14). Chocó (1): Riosucio. Cundinamarca (2): La Esperanza; Quetame. Meta (8): Villavicencio. Valle del Cauca (3): Buenaventura; Calima; Dagua. COSTA RICA (12). Limón (12): Amubri; Hamburg Farm; Hitoy Cerere. ECUADOR (27). Guayas (1): Guayaquil. Napo (12): Lago Agrio; Jatun Sacha; Misahualli; Puerto Agrio; Puerto Napo; Río Arajuno. Zamora-Chinchipe (3): Yantzaza. Imprecise data (3). Ecuador. PANAMA (1). Darien (1): Cana. PERU (27). Ayacucho (2): La Mar, Santa Rosa. Cuzco (6): Pilcopata. Huánuco (13): Tingo María. Madre de Dios (4): Shintuya. San Martín (1): Huagalla. Imprecise data (1): Peru. VENEZUELA (2). Portuguesa (1): San Nicolás. Táchira (1): San Cristobal. Natural history. Tomarus maternus is mainly found below 450 m in tropical forests and near mangroves, and adults are attracted to lights.