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 selanderi ( Cartwright, 1959 ) ( Figs. 2J , 4H , 6F , 14G , 16Q , 20H , 28H ; 45 ) Bothynus selanderi Cartwright, 1959: 537 . Original combination. Male holotype (USNM) “ MEXICO / Los Mochis / Sinaloa // at light / 22 July 1955 // RB&JM Selander // Type No / 63912 / USNM // HOLOTYPE / Bothynus / selanderi / Cartwright”. Female allotype (USNM) “ MEXICO / Los Mochis / Sinaloa // at light / 22 July 1955 // RB&JM Selander // Type No / 63912 / USNM // ALLOTYPE / Bothynus / selanderi / Cartwright”. Twelve male and 21 female paratypes (USNM) “ MEXICO / Los Mochis / Sinaloa // at light / 22 July 1955 // RB&JM Selander // Type No / 63912 / USNM // PARATYPE / Bothynus / selanderi / Cartwright”. “ MEXICO / Mazatlán / Sinaloa // 21 July 1955 // RB&JM Selander // Type No / 63912 / USNM // PARATYPE / Bothynus / selanderi / Cartwright”. Male paratype (MNHN) “Los Mochis / Sinaloa , Mex / 22-VII-1955 // R&J Selander // PARATYPE / Bothynus / selanderi / Cartwright // PARATYPE // MNHN / EC6732 ”. Male paratype (MNHN) “Los Mochis / Sinaloa , Mex / 22-VII-1955 // R&J Selander // PARATYPE / Bothynus / selanderi / Cartwright // PARATYPE // MNHN / EC6733 ”. Type locality: Los Mochis, Sinaloa , Mexico. Description. Habitus as in Fig. 28H . Length 21.0–30.0 mm; humeral width 10.1–14.5 mm . Color dark reddish brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles separated by about 4.5–5.0 tubercle diameters. Clypeus narrowed towards apex, base 2.5 times wider than apex. Clypeal teeth triangular, separated by 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 ( Fig. 4H ). Interocular distance equal to 3.5 times an eye width. Pronotum: Surface with small punctures, punctures denser and deeper on anterior and posterior angles. Apical tubercle prominent, acute, slightly recurved. Subapical fovea deep, rounded, as wide as interocular distance or slightly narrower, equal in both sexes, surface transversely rugose. Scutellum: Surface with minute, sparse punctures. Elytra: Punctures on first interval as large as those on other intervals. Inner surface of apex with rounded, large tubercles forming 17 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, rounded tubercles forming 11–12 diagonal, parallel lines. Pygidium with dense rugosity on basal 3rd. Apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with minute setae. Legs: Protibia tridentate, sometimes with a basal small denticle. Apex of metatibia crenulate, with 13–15 spinules. Male genitalia: Spiculum gastrale without basal plates ( Fig. 14G ). Phallobase 1.1 times longer than parameres. Parameres without dorsal teeth, apex wide, triangularly dilated ( Figs. 16Q , 20H ). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like, accessory lamellae, with dense granules at base. Diagnosis. Tomarus selanderi can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent ( Fig. 28H ); pronotal fovea shallow, as wide as or narrower than interocular distance; protibia tridentate, rarely with an additional basal denticle; apex of metatibia with 14–15 spinules; parameres without dorsal teeth; apex wide, triangularly dilated ( Figs. 16Q , 20H ). Distribution. Tomarus selanderi is an endemic species found along the Pacific coast of Mexico ( Cartwright 1959 ; Endrödi 1985 ; Ratcliffe et al. 2013 ). Locality records ( Fig. 45 ). 205 examined specimens from CAFESI , EMAC , FSCA , CNIN , IEXA , MNHN , UNSM , USNM . Some data from Ratcliffe et al. (2013) . MEXICO (214). Colima (2): Manzanillo. Guerrero (2): Acapulco. Jalisco (17): Careyes; Chamela; Guadalajara; Melaque; Puerto Vallarta. Michoacán (1): Jungapeo. Nayarit (85): Acaponeta; El Venado ; Ruiz; San Blas ; 15 km S San Blas ; Sayulita; Singayta; Tecuala; Yago. Oaxaca (2): Pinotepa Nacional; Puerto Escondido. Sinaloa (98): El Mármol ; Escuinapa; Los Mochis; Mazatlan; Novalato; Piedra Blanca. Sonora (6): Esperanza, Guaymas, Hermosillo . Natural history. Specimens have been collected between 30–1500 m ( Ratcliffe et al. 2013 ) in dry forests of Pacific coast of Mexico .