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 Subgenus Anagrylius Casey, 1915 revised status Anagrylius Casey 1915: 204 (as subgenus). Type species: Heteronychus tumulosus Burmeister, 1847 by monotypy. Description. ( n = 225). Length 13.2–18.8 mm . Humeral width 6.2–9.0 mm. Head: Clypeus trapezoidal ( Fig. 1D ). Apical clypeal teeth transverse to conical, widely separated. Ocular canthus rounded; without ventral or dorsal setae. Frons with 2 tumescences ( Fig. 1D ). Mentum subrectangular ( Fig. 2A ), slightly narrowed at apex; apical margin with long, dense setae. Mentum and ligula separated by a suture ( Fig. 2A ). Ligula wide; covered with long, dense setae; ligular lobes not fused, with a narrow emargination. Maxillar apical palpomere 1.5–1.7 times longer than the second. Maxilla with galea subrectangular (2 times longer than wide), with 6 teeth ( Fig. 3A ). Mandible tridentate (2 apical acute teeth and 1 lateral acute to rounded lobe); tooth 1 narrower than tooth 2. Labrum rectangular (2.5 times wider than long); apex straight to slightly bilobed, with long, dense setae. Antenna with 10 antennomeres, club long (1.5 times longer than antennomeres 2–7). Pronotum: Apical marginal bead complete.Apex with an inconspicuous, rounded tubercle and a small narrow, concavity. Elytra: Inner surface of apex with transverse small tubercles forming 160–166 parallel lines. Wings: RA with dense, large peg-like setae on medial ventral region ( Fig. 7B, D ); with peg-like setae on dorsal surface. Edge of RA 3 with dense setae. Abdomen: Apex of tergite IV with short, transverse, parallel carinae. Sternite VIII emarginate in male; entire in female. Propygidium without stridulatory area. Venter: Prosternum convex. Propleura with long, sparse setae on anterior and posterior surfaces, setae on medial surface shorter and sparser. Apex of prosternal process spherical to flat, with dense setae, not covering all the surface. Metepisternum with incomplete carina; inner surface punctate, with long setae, outer surface homogeneously, glabrous. Metasternum with large, deep, confluent punctures; with long setae, denser on anterior corners. Legs: Male protarsi simple; inner claw simple. Protibia tridentate, teeth equidistant ( Fig. 11C ). Apical margin of mesotibia and metatibia crenulate ( Fig. 12C, I ). Female genitalia: Subcoxite 2 times wider than long. Coxite subquadrate (as long as wide); surface strongly concave. Subcoxite wider and shorter than coxite ( Fig. 13C ). Male genitalia: Phallobase 1.6 times longer than parameres. Parameres without dorsal teeth, with or without a small apical ventral tooth. Internal sac with copulatory lamellae; lamellar spiny belt long; a large accessory lamella with flat base and a thick acute projection ( Fig. 21C ). Diagnosis. Clypeus trapezoidal, apex with 2 conical to transverse small teeth ( Fig. 1D ). Antennal club long. Mentum subrectangular, separated from the ligula by a suture ( Fig. 2A ). Ligula wide; covered with long, dense setae; ligular lobes not fused, with a narrow emargination. Galea with 6 teeth ( Fig. 3A ). Pronotum with apical marginal bead complete and slightly elevated at middle to shape an inconspicuous tubercle, fovea reduced to a narrow, small concavity. Male protarsi simple. Protibia tridentate, teeth equidistant. Apical margin of mesotibia and metatibia crenulate ( Fig. 12C, I ). Internal sac of aedeagus with copulatory lamella, with a long lamellar spiny belt ( Fig. 21C ); accessory lamella with flat base and a thick acute projection (T-shaped) ( Fig. 21C ). Surface of coxite strongly concave ( Fig. 13C ). Stridulatory apparatus consisting of a diagonal line of transverse, short carinae on apex of sternite IV and apical inner surface of elytra with transverse, small tubercle forming parallel lines. Composition. Ligyrus ( Anagrylius ) cuniculus and L. ( Anagrylius ) moroni . Geographic distribution. The genus is found in the Caribbean from the Atlantic Coast of Brazil , along the northern coast of South America, the West Indies, Belize , Mexico , and southern Florida in the United States . Taxonomic remarks. Casey (1915) described Anagrylius as a subgenus of Ligyrus to include L. tumulosus , which was different in the following respects: bidentate maxilla, clypeal teeth more separate, and head larger. The author did not dissect completely the mouth parts as the maxilla has not two but six teeth. Prell (1937) described Ligyrus ( Anagrylius ) cicatricosus in this subgenus, but Anagrylius was not considered valid by Endrödi (1969) or other recent authors. Key to the species of the subgenus Anagrylius 1 Apical angles of clypeus rounded; clypeal teeth triangular to transverse. First and second apical teeth of mandible equal in size and not joined each other. Parameres with parallel sides, without ventral teeth ( Figs. 15D , 17D )................................................................................... Ligyrus ( Anagrylius ) cuniculus ( Fabricius, 1801 ) - Apical angles of clypeus acute; clypeal teeth conical ( Fig. 1D ). First apical tooth of mandible small and joined to the second. Parameres with parallel sides and a small apical ventral tooth each side ( Figs. 15E , 17E )............................................................................... Ligyrus ( Anagrylius ) moroni ( López-García & Deloya, 2019 )