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 colombianus López-García & Gasca-Álvarez, 2014 ( Figs. 9G , 11I , 16D , 19D , 27D ; 44 ) Tomarus colombianus López-García & Gasca-Álvarez, 2014 in López-García et al . 2014: 579 . Original combination. Male holotype (CIUQ) “Colombia—Valle / La Bocana-B/ventura / Altura: 5 m . s.n.m / Fecha: 16/04/04 / Método: Directo / Col: Serna F. // Tomarus colombianus / López-García & / Gasca-Álvarez / HOLOTIPO // CIUQ 1505”. Female paratype (CIUQ) “ COLOMBIA : Dep. Valle del Cauca / Buenaventura / La Bocana / 15.iv.2004 / Light trap / 2 m altitud / Rivas L. Coll // Tomarus colombianus / López-García & / Gasca-Álvarez / PARATIPO // CIUQ 1504”. Female paratype (CIUQ) “Colombia- Valle Cauca / B/ventura-Bocana / Altura: 2 m . s.n.m / Fecha: Mayo/04 / Método: Directo-Bosque / Col: Mendoza L. // Tomarus colombianus / López-García & / Gasca-Álvarez / PARATIPO // CIUQ 1507”. Male parat–pe (ICN) “Colombia—Valle / La Bocana-B/ventura / Altura: 5 m . s.n.m / Fecha: 17/04/04 / Método: Directo / Col: Serna F.” Female paratype (ICN) “ Huila / Neiva / 5.iii.1974 / Bagos A. I. // Tomarus colombianus / López-García & / Gasca-Álvarez / PARATIPO. Type locality: La Bocana, Valle del Cauca , Colombia. Description. Habitus as in Fig. 27D . Length 18.0–21.0 mm; humeral width 8.5–10.5 mm . Color dark reddish brown. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles separated by about 6 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth triangular, separated by less than a tooth diameter. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth. Mentum abruptly constricted at apical 3rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.2 times an eye width. Pronotum: Surface with minute punctures; denser and deeper on anterior and lateral angles, disc impunctate. Apical tubercle small, rounded, not visible in lateral view. Subapical fovea shallow, elongate, narrow (1/3 the interocular distance); equal in both sexes, surface rugopunctate. Scutellum: Surface with deep, large punctures forming 2 lines parallel to margins. Elytra: Punctures on first interval as large as those on other intervals. Inner surface of apex with rounded, large tubercles forming 14–16 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with small, irregular tubercles forming 9–10 diagonal nearly parallel lines. Pygidium with dense rugosity on basal 3rd ( Fig. 9G ), apex rounded. Venter: Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate, with a basal, small denticle ( Fig. 11I ). Apex of metatibia crenulate, with 9–11 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with 2 dorsal teeth on each side; basal tooth acute, longer than apical ( Figs. 16D , 19D ). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like accessory lamellae, with dense granules at base. Diagnosis. Tomarus colombianus can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal surface nearly smooth ( Fig. 27D ); pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1/3 as wide as interocular distance); protibia tridentate, with an additional basal denticle ( Fig. 11I ); apex of metatibia with 9–11 spinules; parameres with 2 dorsal teeth on each side, main teeth long, widely separate from secondary teeth ( Figs. 16D , 19D ). Distribution. Colombia ( López-García et al . 2014 ). Locality records ( Fig. 44 ). Five specimens examined from ICN and CIUQ. COLOMBIA (5). Huila (1): Neiva. Valle del Cauca (4): Buenaventura. Natural history. The few known specimens were found in tropical humid forest near a mangrove swamp and Guandal forest. They were collected by hand and at light traps.