taxonID	type	description	language	source
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	type_taxon	Type species: Scarabaeus ebenus De Geer, 1774: 317 by original designation.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	description	Description. (n = 555). Length 24 – 30 mm. Humeral width 12.5 – 15.1 mm. Head: Clypeus trapezoidal (Fig. 1 A). Apical clypeal teeth transverse and widely separated (3 tooth diameters). Ocular canthus rounded; without ventral or dorsal setae. Frons with 2 transverse, high tumescences. Mentum subrectangular; strongly narrowed at apex (Fig. 2 G – H). Mentum and ligula separate by a suture. Ligula wide; covered with long, dense setae; ligular lobes not fused, with a narrow emargination. Maxillar apical palpomere 1.3 times longer than the second. Maxilla with galea rectangular (2.5 times longer than wide); with 6 teeth (Fig. 4 A). Mandible tridentate (2 apical and 1 lateral triangular large tooth); apical teeth equal in shape (Fig. 5 A). Labrum rectangular (2.5 times wider than long); apex straight to slightly bilobed; with long, dense setae. Antenna with 10 antennomeres, club short (1.2 times longer than antennomeres 2 – 7). Pronotum: Apical marginal bead complete. Apex with a rounded tubercle and an oval to triangular fovea. Elytra: Inner surface of apex with irregular small tubercles. Wings: RA with dense, large peg-like setae on medial ventral region; with peg-like setae on dorsal surface (Fig. 7 A, C). Edge of RA 3 with dense setae. Abdomen: Apex of tergite IV with irregular granules like scales, not forming lines. Sternite VIII emarginate in males; entire in female. Propygidium without stridulatory area. Venter: Prosternum very wide, convex. Propleura with short, sparse setae on anterior and posterior surface, medial surface glabrous. Apex of prosternal process flat, with a deep sulcus, without dorsal setae. Metepisternum with complete carina; inner surface smooth and glabrous; outer surface homogeneously rugose, glabrous. Metasternum with large, deep, confluent punctures, without setae. Legs: Male protarsus enlarged (tarsomeres 2 – 4 2 times wider than long, tarsomere 5 1.5 longer than wide); tarsomere 4 with a lateral flat, striate projection (Fig. 10 A); inner claw bifid (Fig. 10 B). Protibia tridentate, basal tooth distant from others (Fig. 11 A). Apical margin of mesotibia and metatibia crenulate (Fig. 12 A, G). Female genitalia: Subcoxite 1.5 times wider than long. Coxite subquadrate (as long as wide); surface flat (Fig. 13 A). Subcoxite wider than and as long as coxite. Male genitalia: Phallobase 1.6 times longer than parameres. Parameres without dorsal teeth, with ventral tooth (Fig. 17 A). Internal sac without copulatory lamella nor lamellar spiny belt; accessory lamella long with an apical brush of setae (Fig. 21 A).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	diagnosis	Diagnosis. Euligyrus can be differentiated from other Pentodontini by the following: frons concave at middle; clypeus trapezoidal, clypeal teeth widely separated (Fig. 1 A); frontoclypeus with tumescences (Fig. 1 A); mentum subrectangular; galea with 6 teeth (Fig. 4 A); pronotum with apical marginal bead complete; pronotal tubercle rounded; fovea deep, oval to triangular; metasternum glabrous; protibia tridentate; protarsus of male enlarged, inner claw wide and bifid (Fig. 10 B); protarsomere 4 with a lateral flat, striate projection (Fig. 10 A); internal sac without copulatory lamella nor lamellar spiny belt, with an apical brush of setae (Fig. 21 A); surface of coxite flat (Fig. 13 A); and stridulatory apparatus consisting of an area with irregular granules like-scales on apex of sternite IV and apical inner surface of elytra with irregular small tubercles.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	discussion	Composition. Two species: Euligyrus ebenus and E. similis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	distribution	Geographic distribution. It is apparently a South American genus that dispersed to Central America and some islands of the West Indies (Fig. 31).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4FFF90AFA6186CFA23BDB8.taxon	discussion	Phylogenetic relationships. According to the phylogenetic analysis of Tomarus (sensu lato), Euligyrus is not closely related to any of the analyzed genera but shares some characters with species of Cyclocephala and Oxyligyrus (i. e., the shape of the protarsus and inner claw in males, and the shape of the prosternal process). Although its relation to other Dynastinae groups is uncertain, it is probably closer to Cyclocephalini, as suggested by Casey (1915). Taxonomic remarks. Casey (1915) described Euligyrus, which included only E. ebenus, as a subgenus of Ligyrodes due to its shared characters, such as the anterior tarsus modified in the male, the apex of the prosternal process without setae, and the transverse shape of the abdominal spiracles of segments I – III. Although the subgenus Euligyrus was not used since Cartwright (1959) synonymized the genus Ligyrus with Ligyrodes, Endrödi (1969) revalidated this subgenus but as part of the genus Ligyrus and included E. similis. Morón & Grossi (2015) placed these two species in “ Group A ” of the genus Tomarus, but the species of Euligyrus are not closely related to Tomarus and are clearly a different genus with affinities to the tribe Cyclocephalini.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4EFF93AFA61B7DFAB3BA30.taxon	description	(Figs. 1 A, 2 G, 9 A, 13 A, 14 A, 15 A, 17 A, 21 A, 24 A; 31)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4EFF93AFA61B7DFAB3BA30.taxon	description	Description. Habitus as in Fig. 24 A. Length 29.6 – 30.0 mm; humeral width 12.5 – 15.2 mm. Color brown to black. Head: Frons concave between eyes and before frontal tumescences; surface deeply rugose, nearly smooth on vertex and with sparse, deep punctures. Frontoclypeal region with 2 tumescences separated by 1.2 times a tumescence width. Clypeal surface transversely rugose. Clypeus trapezoidal, base 1.8 times as wide as apex (Fig. 1 A). Clypeal teeth transverse to widely triangular, widely separated (about 3 times a diameter tooth) (Fig. 1 A). Mandible with 2 apical and 1 lateral acute large tooth, the apical teeth equal in size and shape. Interocular distance equal to 3.5 times an eye width. Antennal club short. Pronotum: Surface with minute, sparse punctures. Apical tubercle rounded. Fovea oval, narrow (about 1 / 3 narrower than interocular distance) (Fig. 24 A). Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with transverse small tubercles without forming parallel lines. Abdomen: Apex of tergite IV with irregular tubercles like scales. Pygidial surface with deep, evenly sparse punctures, becoming nearly smooth on basal 3 rd. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, basal tooth distant from others (Fig. 11 A). Protarsus enlarged; inner claw bifid (Fig. 10 A – B). Metatibia not narrowed before apex, sides nearly parallel (Fig. 12 G). Apex of metatibia crenulate, with 7 – 8 spinules. Male genitalia: Spiculum gastrale with oval apex (Fig. 14 A). Parameres strongly contracted at apical 3 rd, apices projecting outwards; ventral margin with a small, medial tooth on each side (Figs. 15 A, 17 A).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4EFF93AFA61B7DFAB3BA30.taxon	diagnosis	Diagnosis. Euligyrus ebenus and E. similis are very similar species difficult to differentiate without examination of the shape of parameres. In some populations, like in Colombia, the pronotal fovea is oval and shallower in E. ebenus and triangular and deeper in E. similis; but in other countries like Brazil, the fovea of E. similis is less triangular and so very similar to E. ebenus. The parameres between both species are remarkably different being wider at base, strongly contracted at the apical 3 rd in E. ebenus (Fig. 15 A) while in E. similis they are slender and without strong lateral constrictions (Fig. 15 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4EFF93AFA61B7DFAB3BA30.taxon	distribution	Distribution. Brazil, Bolivia, Peru, Trinidad, Guyana, Suriname (Endrödi 1985), Venezuela (Escalona & Joly 2006), Colombia (López-García et al. 2015), Panama, Costa Rica (Ratcliffe 2003), Nicaragua, Honduras (Ratcliffe & Cave 2006), Belize, Guatemala, Mexico (Ratcliffe et al. 2013), Dominica, Guadeloupe, Martinique, and St. Lucia (Ratcliffe & Cave 2015). Locality records (Fig. 31). 398 examined specimens from CEIOC, CERPE, CEUN, CNIN, FSCA, ICN, LEUC, MPUJ, MN / UFRJ, MZUSP, NHRS, UNSM, UPN, USNM, ZMK. Some data from CMNC, Endrödi (1969) (Bolivia), Ratcliffe (2003), Escalona & Joly (2006), Ratcliffe et al. (2013), and Ratcliffe & Cave (2015). BELIZE (21). No data (7). Cayo (1): East of Roaring Creek. Stann Creek (3): Sitte Point. Toledo (7): 13 mi. NW Punta Gorda, Tranquility Lodge; Blue Creek Village; Punta Gorda; San Miguel Columbia, River Forest. BOLIVIA (1). Cochabamba (1): Chapare. BRAZIL (112). No data (6). Amazonas (12): Barcelos (160 km E); Manaus; Lago Januaca; Río Badajos. Ceará (1): Fortaleza. Espíritu Santo (3): Parque Sooretama, Linhares. Maranhao (4): Bom Jardim, Reserva Biologica Gurupi. Mato Grosso (10): Bodoquena; Diamantino, Alto Rio Arinos; no data. Mina Gerais (4): São Gonçalo do Rio Preto, Parque Estadual do Rio Preto. Pará (54): Belem; Breves; Mangabeira, Mocajuba; Obidos; Oriximina; Rio Xingú, Ponte Novo; Santarém; no data. Paraiba (1): No data. Pernambuco (10): Escada; Paudalho, Campo de Instruçao Marechal Newton Cavalcanti; Recife; Universidade Federal Rural de Pernambuco; Tamandaré. Rio Grande do Norte (2): Natal; Parnamirim. Rio de Janeiro (1): Itatiaia. Rio Grande do Sul (1): Rio Grande. Roraima (1): Caracarai, Parna Viruá. Sergipe (1): Aracaju. COLOMBIA (38). Amazonas (1): Leticia. Antioquia (4): Medellín; Puerto Berrío. Cundinamarca (3): Agua de Dios; Anapoima; Vianí. Meta (3): Remolinos. Nariño (1): Pasto. Valle del Cauca (25): Buenaventura. Vichada (1): Gaviotas. COSTA RICA (13). Heredia (1): Estación Magsasay (Parque Nacional Braulio Carrillo). Limón (10): Amubri; Cerro Cocori; Cerro Tortuguero; Hamburg Farm; Río Sardinas. Puntarenas (2): Estación Sirenas; Rancho Quemado. DOMINICA (4). St. Andrew (1): No data. St. John (1): Cabrits National Park. St. Joseph (2): Clarke Hall. FRENCH GUIANA (15). Cayenne (14): Entomotech Lodge; Risquetout (8 km W); Roura (38 km SE). St Laurent du Maroni (1): St. Jean. GUADALOUPE (6). No data (1). Base-Terre (3): Domaine de Duclos [Petit Bourg]; Piton de Sainte-Rose; Vernou. Grande-Terre (1): Grands Fonds. Marie-Galante (1): St. Louis. GUATEMALA (21). Alta Verapaz (1): Cahabón. Baja Verapaz (1): San Rafael Chilasco. Izabal (15): Biotopo Manati; Cayuga; El Estor; Morales, Finca Jocolo; Morales, Finca Firmeza; La Esmeralda; Río Dulce, Cayo Quemado; Santo Tomás de Castilla. Petén (3): El Ceibal; Sayaxche. Suchitepéquez (1): Finca Parraxe. GUYANA (10): Blairmont. Mazaruni-Potaro (8): Takutu Mountains. HONDURAS (36). Atlántida (33): La Ceiba; Parque Nacional Pico Bonito Rio Zacate; Reserva Cuero & Salado Sendero Olingo. Gracias a Dios (1): Reserva de la Biosfera Rio Plátano. Yoro (2): Parque Nacional Pico Bonito, El Portillo; Suyapa, Victoria. MARTINIQUE (13). La Trinite (2): Gros-Morne; Morne Bellevue. Le Marin (5): Anse Mitan; Le Vauclin; Monroe des Peres; Riviere-Salee; Saint Espirit. Fort-de-France (3): Croix-Rivail; Ravine Vilaine. Saint-Pierre (2): Champflore; Fonds-Saint-Denis. MEXICO (31). Chiapas (8): Bonampak; Ocosingo, Montes Azules. Oaxaca (1): Tuxtepec. Tabasco (2): Villahermosa (6 km W Teapa). Veracruz (19): Catemaco; Coatzacoalcos; El Plan; Las Choapas; Los Tuxtlas; Nautla (10 mi. N); Presidio; Sontecomapa. Doubtful record (1): Distrito Federal, Ciudad Universitaria. NICARAGUA (1). Zelaya (1): Campus Uracan Buffields. PANAMA (10). Colón (4): Madden Dam; Pipeline Road (km 2 – 4). Colón (1): Santa Rita Ridge. Panamá (3): Barro Colorado; El Llano-Carti Road km 8; Margarita. San Blas (2): Nusagandi. PERU (2). Loreto (2): Explorama Inn (25 mi NE Iquitos); Iquitos. ST. LUCIA (32). No data (31). Micoud (1): Escap Community. SURINAME (8). No data (2). Paramaribo (6): Paramaribo. TRINIDAD (34). No data (2). Arima (1): Simla, W. Beebe Tropical Rest. Couva Tabaquite-Talparo (1): Caparo. Point Fortin (7): Point Fortin. Puerto España (4): Puerto España. Río Claro-Mayaro (6): Mayaro Beach. Tunapuna Piarco (13): Centeno; Monroe Blue (1 mi. W); Mount St. Benedict Abby; St. Augustine. VENEZUELA (15). Amazonas (2): Puerto Ayacucho; San Juan de Manapiare. Apuré (1): Fundo Morichalote circa Rio Quitaparo; Santa Rosa, Río Cinaruco, Sakaní. Aragua (1): Tiara. Bolívar (3): Ciudad Bolivar; Reserva Forestal de Imataca, Camp; Río Grande, El Palmar; Río Guaniamo. Guárico (2): Nicolasito. Parque Nacional Aguaro-Guariquito. Miranda (3): Estación Río Negro cerca Capaya. Táchira (1): San Cristóbal. NO DATA (1). Natural history. The species has been reported as a pest of Colocasia esculenta (L.) known commonly as “ papachina ” or “ malanga ” (Piedrahíta et al. 2007). Some specimens were collected in roots of Musaceae.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	description	(Figs. 2 H, 3 D, 4 A, 5 A, 7 A, C, 10 A – B, 11 A, 12 A, 12 G, 15 B, 17 B, 24 B; 31)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	materials_examined	Male holotype (HNHM) “ Venezuela / Maracay / ges P. Vogl // Holotypus / Ligyrus / similis / Endr. ”. Male paratype (HNHM) “ Venezuela / Maracay / ges P. Vogl // Jan. - Febr. / 1935 // Allotypus / Ligyrus / similis / Endr. ”. Male paratype (HNHM) “ Venezuela / Maracay / ges P. Vogl // Jan. - Febr. / 1935 // Paratypus / Ligyrus / similis / Endr. ” Type locality: Maracay, Venezuela.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	description	Description. Habitus as in Fig. 24 B. Length 25.0 – 27.5 mm; humeral width 12.0 – 12.5 mm. Color brown to black. Head: Frons concave between eyes and before frontal tubercles; surface deeply rugose, nearly smooth on vertex and only with sparse, deep punctures. Frontoclypeal region with 2 tumescences separated by 1.2 times a tumescence width. Clypeal surface transversely rugose. Clypeus trapezoidal, base 1.8 times as wide as apex (Fig. 1 A). Clypeal teeth transverse to widely triangular, widely separated (4 times a tooth diameter). Mandible with 2 apical and 1 lateral large, acute tooth, the apical teeth equal in size and shape. Interocular distance equal to 3.5 times an eye width. Antennal club short. Pronotum: Surface with minute, sparse punctures. Apical tubercle rounded. Fovea triangular to oval, narrow (about 1 / 3 narrower than interocular distance) (Fig. 24 B). Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with small tubercles not forming parallel lines. Abdomen: Apex of tergite IV with irregular tubercles-like scales. Pygidial surface with deep, evenly sparse punctures, becoming nearly smooth on basal 3 rd. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, basal tooth distant from others (Fig. 11 A). Protarsus enlarged; inner claw bifid (Fig. 10 A – B). Metatibia not narrowed before apex, sides nearly parallel (Fig. 12 G). Apex of metatibia crenulate, with 6 – 7 spinules. Male genitalia: Spiculum gastrale with oval apex. Parameres slender; with apex acute; apices slightly expanded outwards; ventral margin with a medial, small tooth each side (Figs. 15 B, 17 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	diagnosis	Diagnosis. See the Diagnosis of Euligyrus ebenus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	distribution	Distribution. Venezuela, Ecuador, Brazil, Bolivia, Peru, Trinidad, Guyana, Suriname (Endrödi 1985), French Guiana (Escalona & Joly 2006), Colombia (Neita-Moreno 2011), Panama (Ratcliffe 2003), and Costa Rica (LópezGarcía & Deloya 2018). Locality records (Fig. 31). 157 examined specimens from ANDES, CEIOC, CERPE, FSCA, HNHM, IAvH, ICN, IEXA, LEUC, MEFLG, MEKRB, MN / UFRJ, MZUSP, UNAB, UNSM, USNM. Some data from CMNC and Escalona & Joly (2006). BOLIVIA (7). Beni (3): Guayaramerin; Rurrenabaque. Santa Cruz (4): Buena Vista; Fauna & Flora Hotel, 3.7 km SS Buena Vista. BRAZIL (75). Amapá (8): Amapá; Porto Santana. Amazonas (28). Coarí; Benjamin Constant, Río Itecoal; Ipixuna, Río Gregorio; Lago Grande; Itacoatiara, Costa do Siripá; Manaus; Río Preto; Terezinha, Jiha de Careiro. Goiás (4): Campinas. Cristalina. Marahnão (4): Carolina Pov; Campo Grande; Fortaleza dos Nogueira, Fazenda Santa Maria; Mirador Parque Estadual, Povoado Pindaiba. Mato Grosso (2): Utiariti, Rio Papagaio; Mina Gerais (1): Unaí, Faz; Bolivia. Para (28): Cachimbo; Corcovado, Breves; Maicuru; Oriximiná; Río Javi-Caracurú. COLOMBIA (13). Amazonas (4): Leticia. Antioquia (1): Apartadó. Caldas (1): Palestina, Vereda Santágueda, Granja Montelindo, Valle de Santágueda. Casanare (1): Parque Monterrey. Chocó (2): Riosucio, Cacarica; Utria. Meta (4): Cabuyaro, Vereda Yarico; Puerto Gaitán. Río Duda, Parque Nacional Natural Tinigua, Centro de Investigaciones Ecológicas La Macarena; San Miguel, Finca Yarmato. COSTA RICA (2): Puntarenas (2): Coto. ECUADOR (6). Manabi (1): Pedernales. Sucumbios (5): Limoncocha; Estación Biologica Coca La Selva (175 km SE). GUYANA (1): No data. PANAMA (8). Colón (2): Madden Dam. Los Santos (1): Corozal. Panamá (5): Barro Colorado; Camino del oleoducto. PERU (11). No data (6). Huánuco (3): Aucayacu; Tingo María. Loreto (1): Sarayacu. San Martin (1): Tarapoto. SURINAME (4). Para (4): Zanderij. VENEZUELA (147). No data (2). Amazonas (70): Cerro de la Neblina; La Esmeralda; Ocamo; Parque Nacional Duida Marahuaka, Culebra; Parque Nacional Parima Tapirapeco, Parima; San Carlos de Río Negro; San Fernando de Atabapo; San Simón del Cocuy; Santa Lucía; Surumoni. Aragua (3): Maracay. Barinas (2): Barrancas; Río Caparo. Bolívar (13): Anacoco; Caicara, San Juan de Manapiare; Kanarakuni; Los Pijiguaos; Salto Las Babas; Río Caura, Salto Para; Reserva Forestal Imataca; Río Grande, El Palmar. Carabobo (24): Central Tacarigua; Samán Mocho; Tacarigua; Urama; Valencia. Delta Amacuro (1): Coporito. Falcón (3): Miranda; Sanare; Yacaral. Monagas (2): Carapito. Portuguesa (5): San Nicolás. Táchira (5): Complejo Hidroeléctrico Leonardo Ruíz Pineda, Las Cuevas; Cordero; La Grita; Michelena. Yaracuy (1): La Hoya. Zulia (15): Colón; Perijá; Tres Bocas. Natural history. Euligyrus similis was reported as an important pest of cultivated banana, and occasionally attacking sugar cane and palms (Bactris gacipaes (Kunth )) (Arecaceae) in some localities of Amazonas, Brazil (Pamplona et al. 1994). Some specimens were collected in oil palm plantations. Genus Proculigyrus López-García & Deloya new genus	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	type_taxon	Type species. Ligyrus cicatricosus Prell, 1937, here designated	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	description	Description. (n = 33). Length 16.5 – 25.2 mm. Humeral width 8.2 – 13.3 mm. Head: Clypeus trapezoidal. Apical clypeal teeth transverse, small, widely separated. Ocular canthus acute; without ventral or dorsal setae. Frons with 2 tumescences. Mentum subquadrate (as long as wide); slightly narrowed at apex; apical margin with long, dense setae. Maxillary apical palpomere 1.5 times longer than the second. Galea of maxilla rectangular (2.5 times longer than wide); with 3 dorsal and 3 ventral teeth. Mandible tridentate (2 apical acute teeth and a lateral lobe acute and directed upwards). Labrum as long as wide; apex evenly rounded with long, dense setae. Antenna with 10 antennomeres, club short (as long as antennomeres 2 – 7). Pronotum: Apical marginal bead incomplete on middle 1 / 5. Apex without tubercle, with a small swelling and shallow, narrow fovea. Elytra: Inner surface of apex with transverse tubercles forming approximately 85 parallel lines (Fig. 8 D). Wings: RA with dense, large, peg-like setae on medial ventral region; without peg-like setae on dorsal surface. Abdomen: Tergite IV with transverse, inconspicuous striae on each side (Fig. 8 A). Sternite VIII emarginate in both sexes; emargination narrow in female, wider in male. Propygidium without stridulatory area. Venter: Prosternum acute. Propleura with a few, sparse setae on anterior and posterior surfaces, medial surface glabrous. Apex of prosternal process spherical, with dense dorsal setae. Metespisternum with complete carina; inner surface smooth, outer surface evenly rugose; without setae. Metasternum with small, deep, confluent punctures and with long setae, denser on anterior corners. Legs: Male protarsus simple; inner claw simple. Protibia tridentate, basal tooth distant from the others (Fig. 11 B). Apical margin of mesotibia and metatibia dentate (Fig. 12 B, H). Female genitalia: Subcoxite 1.8 times wider than long, outer sides narrowed towards apex (basal margin 1.5 times wider than apical margin). Coxite subrounded (as long as wide); surface strongly concave. Subcoxite as wide as coxite; shorter than coxite (Fig. 13 B). Male genitalia: Parameres approximately 1 / 2 shorter than phallobase. Parameres with a ventral, truncate tooth on each side; without dorsal teeth. Internal sac without copulatory lamellae or lamellar spiny belt and with 12 accessory lamellae with a flat, oval base (comma-like) (Fig. 21 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	diagnosis	Diagnosis. Clypeus trapezoidal, apex with 2 transverse, small teeth (Fig. 1 B). Mentum subquadrate (Fig. 2 F). Galea with 6 teeth (3 dorsal, 3 ventral) (Fig. 4 B). Pronotum with apical marginal bead incomplete on middle 1 / 5 (Fig. 1 B). Apex without tubercle, with a small swelling and shallow, depression. Surface of pronotum, elytra, and pygidium with large, dense punctures (Fig. 24 C). Metepisternum glabrous. Male protarsus simple. Protibia tridentate, basal tooth distant from others (Fig. 11 B). Apical margin of mesotibia and metatibia dentate (Fig. 12 C, I). Sternite VIII of female narrowly emarginate. Coxite surface strongly concave (Fig. 13 B). Internal sac of aedeagus without copulatory lamella; accessory lamellae with a flat, oval base (comma-like lamellae) (Fig. 21 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	distribution	Distribution. It is distributed from the Mexican tropics (south of Trans-Mexican volcanic belt) to the Chocó biogeographic region in South America. Natural history. The genus is found in cloud forests and tropical humid forests. Most individuals have been collected with light traps between at the beginning of the rainy season (April – May), but it is in general a little abundant group and poorly represented in collections. The larval stages have not been described.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4DFF95AFA61FF4FCC7BA4C.taxon	etymology	Etymology. The name Proculigyrus comes from the Latin word “ procul ” that means far and refers to the phylogenetic position of this genus with respect to Ligyrus, in which the type species was originally described. Proculigyrus is masculine in gender for the purposes of nomenclature. Taxonomic remarks. Prell (1937) described P. cicatricosus in the subgenus Anagrylius Casey, 1915 of the genus Ligyrus, probably due to the trapezoidal shape of the clypeus and shape of the pronotal fovea and mandible. Endrödi (1969) did not recognized this group and placed this species in the subgenus Ligyrus by having mandibles with three teeth. However, several other differential characters from mouthparts and internal sac were found, suggesting this species is a different genus and not related to Ligyrus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4BFF94AFA61F28FB4BBA30.taxon	description	(Figs. 1 B, 2 F, 4 B, 5 B, 8 A, D, 9 B, 11 B, 12 B, 12 H, 13 B, 14 B, 15 C, 17 C, 21 B, 24 C; 40)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4BFF94AFA61F28FB4BBA30.taxon	materials_examined	Male lectotype designated by Endrödi (1969) (ZMHB) “ Mexico // Ligyrus / (Anagrylius) / cicatricosus Prell / ♂ - Type // Lectotypus / Ligyrus / cicatricosus / Prell / Endrody ”. Female paralectotype (ZMHB) “ Mexiko / Coll / Dr. ITZINGER // Ligyrus / cicatricosus Prell / ♀ - Type // Paratypus / Ligyrus / cicatricosus / Prell ”. Type locality: Mexico.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4BFF94AFA61F28FB4BBA30.taxon	description	Description. Habitus as in Fig. 24 C. Length 16.5 – 25.2 mm. Humeral width 8.2 – 13.3 mm. Color black, legs lighter. Head: Frons slightly concave between eyes and behind frontoclypeal tubercles; surface with deep, large, confluent punctures, nearly smooth on vertex and only with sparser punctures. Clypeus trapezoidal (Fig. 1 B). Apical clypeal teeth transverse, small, separated by 5 tooth diameters. Ocular canthus acute. Frons with 2 tumescences. Interocular distance 3.7 times the transverse ocular diameter. Antennal club short. Pronotum: Surface with punctures evenly sparse, deep, large, weakly umbilicate. Apex without defined tubercle, with an inconspicuous swelling and small fovea. Elytra: Inner surface of apex with transverse tubercles forming about 85 parallel lines (Fig. 8 D). Abdomen: Tergite IV with 6 transverse, inconspicuous striae (Fig. 8 A). Sternite VIII of female with narrow emargination. Pygidial surface completely punctate (Fig. 9 B); punctures subequal in size to those of elytral apex, ocellate, minutely setigerous. Surface evenly convex in male, slightly convex in female. Venter: Apex of prosternal process spherical, with dense setae. Metespisternum without setae. Metasternum with small, deep, confluent punctures; with long setae, denser on anterior corners. Legs: Male protarsus simple; inner claw simple. Protibia tridentate, basal tooth distant from the others (Fig. 11 B). Apical margin of mesotibia and metatibia dentate (Fig. 12 B, H). Apex of metatibia with 4 – 7 spinules. Female genitalia: Subcoxite 1.8 times wider than long, outer sides narrowed towards apex (basal margin 1.5 times wider than apical margin). Coxite subrounded (as long as wide); surface strongly concave. Subcoxite as wide as coxite; shorter than coxite (Fig. 13 B). Male genitalia: Spiculum gastrale T-shaped, thickened (Fig. 14 B). Parameres with a ventral, truncate tooth on each side; without dorsal teeth (Figs. 15 C, 17 C). Internal sac with 12 accessory lamellae like spines and with a flat, oval base (Fig. 21 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4BFF94AFA61F28FB4BBA30.taxon	diagnosis	Diagnosis. It is the only known species in the genus and can be differentiated from other Pentodontini by the combination of the following characters: clypeus trapezoidal, clypeal teeth small, frontoclypeus with tumescences (Fig. 1 B); apex of mesotibia and metatibia dentate (Fig. 12 B, H); mentum subquadrate (Fig. 2 F); protarsus of male simple; pronotum and pygidium with deep, large punctures; and parameres with wide, acute medial teeth and an apical long projection directed outwards on each side (Figs. 15 C, 17 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4BFF94AFA61F28FB4BBA30.taxon	distribution	Distribution. Mexico, Guatemala (Ratcliffe et al. 2013), El Salvador, Honduras (Ratcliffe & Cave 2006), Costa Rica (Ratcliffe 2003), and Colombia (López-García et al. 2015). Locality records (Fig. 40). 33 examined specimens from FSCA, ICN, IEXA, UNSM, USNM, ZMHB. Some records from Ratcliffe (2003), Ratcliffe & Cave (2006), and Ratcliffe et al. (2013). COLOMBIA (1). Valle del Cauca (1): Buenaventura, Río San Juan. COSTA RICA (1). Puntarenas (1): Rancho Quemado. EL SALVADOR (23). Ahuachapan (3): Caserío San Miguelito; Parque Nacional El Imposible. La Libertad (16): San Andrés; Santa Tecla. Santa Ana (3): Hacienda Montecristo, Cerro Miramundo, Metapán. San Salvador (1): San Salvador. GUATEMALA (31). Alta Verapaz (1): Finca Chulac, Mayanaquel Mining Company. Baja Verapaz (7): Biotopo de Quetzal; Carretera a Chilasco; Purulha. Chiquimula (1): Las Presas, arriba de Concepción Las Minas. El Progreso (2): El Rancho; Finca El Riscosa (5 km N El Pacayal). Escuintla (3): Peñón de Siquinalá; Santa Lucía Cotzumalguapa. Huehuetenango (1): Barillas, Malpaís. Izabal (5): Cerro Negro Norte (1 km W Honduras); Morales, Finca Firmeza. Suchitepequez (1): Finca El Vesubio (12 km N Patulul). Zacapa (10): Finca Santa Cruz, Marble Quarry road (NE Teculután); La Union (5 - 6 km S); road to San Lorenzo Marmol Mine (NW Tuculután); Teculután. HONDURAS (28). Atlántida (2): La Ceiba. Comayagua (3): El Taladro; Siguatepeque. Cortés (6): Cofradia (9.3 km NW); Lago de Yojoa; Parque Nacional Cusuco. El Paraíso (11): Capire (8.3 km SE); El Paraíso. Francisco Morazán (1): Cerro Uyuca. Olancho (2): Montaña del Malacate. Yoro (3): Parque Nacional Pico Pijol. MEXICO (11). Chiapas (5): Cañón El Chorreadero (5 mi. E Chiapa de Corzo); El Aguacero (16 km W Ocozocautla); Suchiapa (5 km SE). Oaxaca (2): Cuicatlán (15 mi. N); Salina Cruz (5 km N). Tabasco (1): Villa Hermosa. Veracruz (3): Puente Nacional; Veracruz. Natural history. Specimens have been found mainly in cloud forests up to 2,300 m but also in tropical humid forests near sea level. Most individuals have been collected with light traps between April and May.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	materials_examined	Type species: Scarabaeus gibbosus DeGeer, 1774 by subsequent designation (Casey 1915: 178).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	materials_examined	Type species: Scarabaeus relictus Say, 1825: 194 by original designation.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	materials_examined	Type species: Podalgus villosus Burmeister, 1847: 120 here designated.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	materials_examined	Type species: Ligyrus ruginasus LeConte, 1856: 20 by original designation.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	description	Description. (n = 5,691). Length 11.3 – 21.8 mm. Humeral width 5.9 – 11.5 mm. Head: Clypeus triangular, subtriangular, or trapezoidal (Fig. 1 C – G). Apex of clypeus with 1 or 2 triangular to transverse teeth, not or slightly separated. Ocular canthus rounded to acute, with or without ventral setae, without dorsal setae. Frons with tumescences or high carina, entire or interrupted at middle. Mentum triangular to subrectangular (Fig. 2 A – E); surface flat to strongly convex. Mentum and ligula separate or fused dorsally. Ligula reduced or well developed, covered with short, sparse or long, dense setae; ligular lobes fused or not. Maxillary apical palpomere 2.0 – 2.5 times longer than the second. Maxilla with galea rectangular (3.0 – 3.5 times longer than wide); with 3 – 6 well-developed teeth, sometimes with 1 to 2 rudimentary teeth. Mandible with 2 apical teeth and a lateral widely rounded lobe; tooth 1 with apex acute, longer and narrower than tooth 2. Labrum subquadrate (as long as wide) to rectangular; apex rounded to slightly bilobed; with long, dense setae. Antenna with 10 antennomeres, club long to very long (1.5 – 2.0 times longer than antennomeres 2 – 7). Pronotum: Apical marginal bead complete. Apex with or without tubercle; without fovea or with deep, wide fovea. Elytra: Inner surface of apex with transverse, small tubercles forming 50 – 166 parallel lines. Wings: RA with dense, large peg-like setae on medial ventral region; with peg-like setae on dorsal surface (Fig. 7 B – E). Edge of RA 3 with dense to sparse setae. Abdomen: Apex of tergite IV with a diagonal line of 9 – 14 short to large, transverse, parallel carinae. Sternite VIII emarginate in male; entire in female. Pygidium without rugosity. Propygidium without stridulatory area. Venter: Prosternum narrow, acute to slightly rounded. Propleura with long, dense setae on all the surface, sparser on medial area. Apex of prosternal process spherical to flat; with or without dorsal long setae. Metepisternum with incomplete, very thin carina; inner surface with deep, large punctures, with long setae; outer surface homogeneously rugose, with long, dense setae. Metasternum with small, deep, confluent punctures; with long, dense setae. Legs: Male protarsus simple or enlarged; inner claw simple or enlarged (Fig. 10 C – E). Protibia tridentate, sometimes with denticles (Fig. 11 E); basal tooth slightly distant from others. Metafemur with well-defined anterior row of punctures. Apical margin of mesotibia and metatibia entire to crenulate. Female genitalia: Subcoxite 1.5 – 2.0 times wider than long. Coxite subquadrate (as long as wide) to subrectangular (wider than long); surface flat to strongly concave. Subcoxite wider and subequal to longer than coxite. Male genitalia: Phallobase 1.2 – 1.3 times longer than parameres. Parameres without dorsal teeth (rarely with a dorsal long teeth), with ventral tooth. Internal sac with copulatory lamella and short to long lamellar spiny belt; accessory lamella simple (entire structure with shapes variable among the species).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	diagnosis	Diagnosis. Ligyrus can be differentiated from other Pentodontini by the following: frontoclypeus with carina or tumescences; antennal club long to very long (1.5 – 2.0 times as long as antennomeres 2 – 7); mentum triangular to subrectangular (Fig. 2 A – E); ligula narrow to wide; ligular lobes fused or not, with a narrow emargination; galea with 3 – 6 teeth (Fig. 3 A – C); pronotum with apical marginal bead complete; pronotal tubercle small to large; fovea deep, oval to triangular or absent; metepisternum with long setae; protibia tridentate, with or without additional denticles, basal tooth distant from others (Fig. 11 C – G); protarsus of male simple or enlarged (Fig. 10 C); internal sac with copulatory lamella and lamellar spiny belt, accessory lamellae simple (Fig. 21 C – E); surface of coxite flat to strongly concave (Fig. 13 C – G); and stridulatory apparatus consisting of a diagonal strip of short, transverse, parallel carinae on apex of sternite IV (Fig. 8 B – C) and apical inner surface of elytra with transverse small tubercles forming parallel lines (Fig. 8 E – F).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	synonymic_list	Composition. Eighteen species. Tomarus adoceteus, T. pullus, and P. cicatricosus considered by Morón & Grossi (2015) in the genus Ligyrus are excluded and placed in other genera, while L. ruginasus and L. peninsularis (previously in Oxygrylius) are included in Ligyrus, as well as three species described recently in the genus Tomarus: L. spinipenis, L. moroni, and L. paranaensis. It is difficult to assess whether the fossil species Ligyrus compositus Wickham, 1911 and L. effetus Wickham, 1914 from Florissant, Colorado (Late Eocene) belong to Ligyrus, but any diagnostic character of the genus coincides with the descriptions or illustration of the fossils presented by Wickham (1911, 1914).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	distribution	Geographic distribution. Species of Ligyrus are found from southeastern Canada to northern Argentina, including the West Indies, in deserts and dry to humid forests (mainly pine / oak and cloud forests). No species are found in Amazonia.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD4AFF96AFA61FF4FA22B807.taxon	discussion	Phylogenetic relationships. Ligyrus (sensu Morón & Grossi 2015) was found as a heterogeneous group of three subgenera Ligyrus (sensu stricto), Anagrylius, and Ligyrodes, which coincided in part with the Casey (1915) subgenera. Ligyrus (sensu stricto) is paraphyletic with respect to Oxygrylius as its species are closely related, especially those from North America. Taxonomic remarks. Burmeister (1847) described the genus Ligyrus to group some species of Heteronychus and Podalgus with a kind of stridulatory apparatus on the inner surface of the elytra, which is not found in other Dynastinae. Casey (1915) separated the group in three genera (Ligyrus, Ligyrodes, and Oxygrylius) and several subgenera as shown in Table 1. Then, Endrödi (1969) presented a different classification using part of the Casey subgenera for Ligyrus and considering Oxygrylius as a different genus. The name Ligyrus was maintained as a valid name for more than 150 years until Ratcliffe (2003) revalidated the name Tomarus by having priority over Ligyrus, and his proposal was widely accepted. Morón & Grossi (2015) separated the genus into Ligyrus and Tomarus, grouping the species by the number of mandibular teeth, and presence of a pronotal fovea. Ligyrus (sensu Morón & Grossi 2015) coincides only in part with the classification presented in this work. Oxygrylius has been treated as a separate genus for most authors except Hardy (1964), who considered it a synonym of Bothynus (name used erroneously for species of Ligyrus). Hardy (1964) found a specimen of L. gibbosus with triangular apex as in Oxygrylius, and so he thought it could be caused by a mutation in a single character. That is consistent with the results of the cladistic analysis, as there are many other characters supporting the grouping of these two species with Ligyrus, particularly with Ligyrus (sensu stricto) (i. e., presence of frontoclypeal carina, mentum triangular, galea rectangular, coxite strongly concave, structure of the stridulatory apparatus, shape of accessory lamella, etc.).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	type_taxon	Type species: Heteronychus tumulosus Burmeister, 1847 by monotypy.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	description	Description. (n = 225). Length 13.2 – 18.8 mm. Humeral width 6.2 – 9.0 mm. Head: Clypeus trapezoidal (Fig. 1 D). Apical clypeal teeth transverse to conical, widely separated. Ocular canthus rounded; without ventral or dorsal setae. Frons with 2 tumescences (Fig. 1 D). Mentum subrectangular (Fig. 2 A), slightly narrowed at apex; apical margin with long, dense setae. Mentum and ligula separated by a suture (Fig. 2 A). 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. 3 A). 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. 7 B, 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. 11 C). Apical margin of mesotibia and metatibia crenulate (Fig. 12 C, 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. 13 C). 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. 21 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	diagnosis	Diagnosis. Clypeus trapezoidal, apex with 2 conical to transverse small teeth (Fig. 1 D). Antennal club long. Mentum subrectangular, separated from the ligula by a suture (Fig. 2 A). Ligula wide; covered with long, dense setae; ligular lobes not fused, with a narrow emargination. Galea with 6 teeth (Fig. 3 A). 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. 12 C, I). Internal sac of aedeagus with copulatory lamella, with a long lamellar spiny belt (Fig. 21 C); accessory lamella with flat base and a thick acute projection (T-shaped) (Fig. 21 C). Surface of coxite strongly concave (Fig. 13 C). 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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	synonymic_list	Composition. Ligyrus (Anagrylius) cuniculus and L. (Anagrylius) moroni.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	distribution	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD48FF99AFA61DEFFA23B8A1.taxon	discussion	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD47FF9AAFA61C45FE10BC98.taxon	description	(Figs. 7 B, D, 9 C, 15 D, 17 D, 25 A; 32)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD47FF9AAFA61C45FE10BC98.taxon	materials_examined	Male lectotype designated by Cartwright (1959: 518) (ZMK) “ cunicu / lus. // Lectotypus / Geotrupes / cuniculus / Fabr. / Design. 1957 / B. G. Landin // Ligyrus / cuniculus (F.) / Det. F. Chalumeau / 81 ”. Type locality: America (South Carolina). Scarabaeus antillarum Palisot de Beauvois, 1811: 104. Synonym. Type specimens were lost in a shipwreck off the coast of Nova Scotia in 1798. Heteronychus tumulosus Burmeister, 1847: 101. Synonym. Male lectotype designated by Endrödi (1969) (MLUH) “ tumulosus / Pal. Beauv. / humilis Bug. / Is. Dominq. Imr. // Lectotypus / Ligyrus / (Heteronychus) / tumulosus Burm. / Endrody // Ligyrus / cuniculus (F.) / Det. F. Chalumeau 180 ”. Type locality: Dominique.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD47FF9AAFA61C45FE10BC98.taxon	description	Description. Habitus as in Fig. 25 A. Length 13.0 – 18.8 mm; humeral width 6.2 – 9.0 mm. Color dark reddish brown. Head: Frons coarsely rugopunctate, area between eyes nearly smooth. Frontoclypeal region with 2 tumescences (Fig. 2 D). Clypeus trapezoidal, base 1.5 times as wide as apex; apical angles rounded (Fig. 2 D). Clypeal teeth widely triangular, widely separated (1 tooth diameter). Mandible with 2 apical and 1 lateral, acute, large tooth, the first apical tooth small and joined to the second. Interocular distance equal to 3 times an eye width. Antennal club long, 1.5 times longer than antennomeres 2 – 7. Pronotum: Surface with minute, sparse punctures. Apical tubercle inconspicuous, rounded. Fovea small, a shallow concavity behind tubercle (Fig. 25 A). Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with transverse, small tubercles forming 166 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 14 short, transverse, parallel carinae, carinae slightly wider at middle. Pygidium with large, evenly distributed punctures (Fig. 9 C). Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate without an additional small basal tooth. Protarsus simple; inner claw simple. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia slightly crenulate, with 15 – 19 spinules. Male genitalia: Spiculum gastrale with apex widened. Parameres with sides nearly parallel, slightly contracted at the apical 4 th, apices projecting outwards, ventral margin without teeth (Figs. 15 D, 17 D).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD47FF9AAFA61C45FE10BC98.taxon	diagnosis	Diagnosis. Ligyrus cuniculus and L. moroni are similar in general appearance, shape of the mandible, tumescences on head, and small pronotal fovea and tubercle. However, L. cuniculus has the clypeal apical angles rounded (acute in L. moroni), the clypeal teeth triangular to transverse (conical in L. moroni), and the parameres without an apical, ventral tooth (Figs. 15 D, 17 D) (present in L. moroni) (Figs. 15 E, 17 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD47FF9AAFA61C45FE10BC98.taxon	distribution	Distribution. Brazil, French Guiana, Suriname, Guyana, Venezuela, Colombia, Curaçao, Trinidad and Tobago, Grenada, St. Vincent and the Grenadines, Barbados, St. Lucia, Martinique, Guadeloupe, Monserrat, Antigua, Barbuda, St. Kitts and Nevis, Saba, St. Barthelemy, St. Maarten, Anguilla, British Virgin Islands, United States Virgin Islands, Puerto Rico, Dominican Republic, Haiti, Jamaica, Cayman Islands, Cuba, Bahamas, Belize, Mexico, and the United States (Endrödi 1985; Escalona & Joly 2006; Ratcliffe & Cave 2015, 2017; Hielkema & Hielkema 2019). Locality records (Fig. 32). 200 examined specimens from CEIOC, CERPE, FSCA, ICN, IEXA, MZUSP, UNSM, and USNM. Some records from Endrödi (1969), Escalona & Joly (2006), Ratcliffe & Cave (2015, 2017), Albuquerque et al. (2016), and Hielkema & Hielkema (2019). Only counties are listed for the United States of America. ANGUILLA (5). Altamer Hotel, Shoal Bay West; Cap Juluca, Maundays Bay; Flat Cap Point; near Captain’s Beach; West End. ANTIGUA (8). Saint George (1): Antigua International Airport. Saint John (1): St. John’s. Saint Mary (1): Christian Valley Agricultural Station. Saint Paul (3): English Harbour Town; Marmora Bay; Swetes. Saint Peter (1): Big Duers. Saint Philip (1): Devil’s Bridge. BAHAMAS (19). Abaco (2): Elbow Cay; Hopetown. Andros (7): San Andros Airport; Andros Town; Fresh Creek, Androsia Forfar Field Station; Stafford Creek. Berry Islands (1): Frazier Hog Cay. North Bimini (2): Big Game Marina. South Bimini (2): No data. Eleuthera (3): Current; Governor’s Harbour; Rainbow Bay. New Providence (2): Nassau; Orange Hill. BARBADOS (9). Christ Church (2): Dover; Silver Sands. Saint James (3): Bellairs Research Institute; Holetown. Saint Lucy (1): Hope. Saint Joseph (1): Walker’s Bridge. Saint Michael (2): Cave Hill; Bridgetown; Marine Hotel. BARBUDA (1). Codrington. BELIZE (1). No data. BRAZIL (118). Alagoas (42): Delmiro. Amapá (1): Serra do Navio. Bahía (3): Dias D’Avila; San Salvador. Ceará (5): Fortaleza; Timbauba. Espírito Santo (1): No data. Maranhão (1): São Luis. Pará (4): Belem; Maicurú; Utinga. Paraíba (10): Coremas; Juazeirinho Soledade; Sta. Luzia. Pernambuco (8): Abreu e Lima, Campo de Instrucao Marechal Newton Cavalcanti; Garanhuns; Igarassu, Refúgio Charles Darwin; Olinda; Paudalho, Campo de Instruçao Marechal Newton Cavalcanti. Piauí (1): Teresina. Rio Grande do Norte (39): Mossoró; Natal; Nisia Floresta; Parnamirim; Ceará-Mirim. Rio de Janeiro (1): Buzios. Sergipe (2): Aracaju. BRITISH VIRGIN ISLANDS (5). Guana Island; Virgin Gorda, Fischer’s Core; Tortola. CAYMAN ISLANDS (5). Grand Cayman Island (3): Boatswain Point; Queen Elizabeth II Botanic Park; Mastic Trail, S trailhead. Little Cayman Island (2): North Coast Road (0.1 km W jct. Olivine Kirk Drive); Sandy Point. COLOMBIA (2). Atlántico (1): Repelón, Reserva Bijibana. Bolívar (1): Cartagena. CUBA (18). Artemisa (1): Artemisa. Camagüey (2): Camagüey; Esmeralda. Cienfuegos (2): Cienfuegos; Pepito Tey. La Habana (1): La Habana. Holguín (1): Frank Pais; Tanamo. Isla de la Juventud (2): Nueva Gerona, Sierra de Casas. Matanzas (1): Parque Nacional Ciénaga de Zapata. Mayabeque (1): Santa Cruz del Norte. Pinar del Río (2): Guane; Pinar del Río. Sancti Spíritus (1): Trinidad. Santa Clara (1): Carralillo. Santiago de Cuba (1): Guama; Siboney. CURAÇAO (2). Piscadera. DOMINICA (2). St. George (2): Roseau; St. Aroment. St. John (2): Portsmouth; Cabrits National Park. St. Joseph (3): Clarke Hall; Mouth of Layou River; Grande Savane. St. Mark (1): Scotts Head Village. St. Patrick (1): Fond Saint Jean. St. Paul (5): Canefield; Mahaut (1 mi. N); Pont Casse (2.3 mi. E); Springfield Estate. DOMINICAN REPUBLIC (54). Azua (2): Padre Las Casas (8 km NE). Baoruco (1): Neiba (5.8 km SW). Barahona (2): Barahona; Filipinas, Lorimar Mine. Dejabón (2): Dejabón; Loma de Cabrera. Distrito Nacional (2): El Cachón de la Rubia; Santo Domingo. Espaillat (1): Moca. Hato Mayor (1): Parque Nacional Los Haitises. Independencia (2): 3 km up road La Descubierta-Los Pinos. La Altagracia (7): Bayahibe; Isla Saona; Higuey; Parque del Este, Caseta Guaraguao; Parque Nacional del Est, Boca de Yuma; Nisibon. La Romana (3): Cacata; Casa de Campo; La Romana. La Vega (4): Jarabacoa; La Vega; Rio Camu (19 km NE Jarabacoa). Monseñor Nouel (1): Bonao. Monte Cristi (4): Botoncillo (5 km NNE); Monte Cristi (10 km S); Villa Elisa (9 km N). Pedernales (2): Isla Beata; Pedernales. Peravia (1): Bani. Puerto Plata (3): Cabarete; Puerto Plata. Salcedo (1): Salcedo. Samaná (2): Samaná; Sánchez. San Cristóbal (1): San Cristóbal. San José de Ocoa (1): San José de Ocoa. San Juan (3): Guanito; Rio San Juan; San Juan de la Maguana. San Pedro de Macoris (1): San Pedro de Macoris. Santiago Rodríguez (2): Sabaneta; Santiago. Santo Domingo (1): Boca Chica. Valverde (2): Los Hidalgos; Mao (Colonia Amina). FRENCH GUIANA (4). No data (3). Cayenne (1): Cayenne. GRENADA (7). St. Andrew (2): Mirabeau; Pearls Airport. St. David (1): La Sagesse Bay. St. George (3): Lance aux Epines, Coral Cove; St. George’s Botanical Garden. St. Patrick (1): Levera. GRENADINES (7). Bequia (1): Bequia. Canouan (2): Crystal Sands. Carriacou (2): Belvedere; Hillsborough. Isle a Quatre (1): No data. Mayreau (2): Saltwhistle Bay; S end of island. Union Island (3): Clifton; Chatham Bay; Richmond Bay. GUADALOUPE (11). Base-Terre (4): Parc National de la Guadeloupe; Petit Bourg; Pigeon, Chalet Sou-le-Vent; Vernou. Grande-Terre (3): Point-a-Pitre; Pointe des Chateaux; Saint-Francois. Îles des Saintes (1): Terre-de-Haut. La Désirade (1): No data. Marie-Galante (2): Marie-Galante. GUYANA (1). No data (1). HAITI (4). Artibonite (2). Des Ronville, Gonaives (5 km ENE); Saint-Michel de L’Atalaye. Ouest (1): Port-au-Prince. Sud (1): Les Cayes. JAMAICA (15). St. Andrew (4): Constant Spring; Fresh River; Kingston; Mona. St. Ann (3): Discovery Bay Marine Laboratory; Ocho Ríos; Runaway Bay. St. James (1): Montebajo Bay. Trelawny (3): Ducans; Falmouth (3 mi. S). Westmoreland (1): Negril, Crystal Waters Beach. MARTINIQUE (17). Fort-de-France (4): Bel Air; Croix-Rivial; Fort-de-France. La Trinite (3): La Trinite; Presqu’Île de la Caravelle. Le Marin (6): Anse du Ceron; La Fouquette; Morne Amerique du Sud; Sarcelle. Saint Pierre (4): Fonds-Saint-Denis; Morne a Lianes; Saint Pierre. MEXICO (2). Tabasco (2): Centla. MONSERRAT (11). Saint Anthony (1): Plymouth. Saint George (1): Riley’s Estate. Saint Peter (9): Blake’s Estate, Bottomless Ghaut; Brades; Old Towne, Palm Court; Rendezvous Bay; Woodlands, Riverside House. NETHERLANDS ANTILLES (1): Saba Island (6): The Bottom; Windwardside. PUERTO RICO (90). Aguadilla (2): Aguadilla; San Antonio. Aibonito (1): Aibonito. Añasco (1): Añasco. Arecibo (1): Arecibo. Arroyo (1): Arroyo. Barceloneta (2): Barceloneta. Bayamón (1): Bayamón. Cabo Rojo (3): Boquerón; Cabo Rojo (Barrio Arenas); Los Morrillos State Forest. Canovanas (1): Canovanas. Coamo (2): Baños de Coamo; Coamo. Comerio (1): Comerio. Corozal (1): Corozal. Dorado (1): Dorado. Fajardo (7): Fajardo; Santa Rita. Guanica (6): Bosque Estatal de Guanica; Ensenada; Guanica. Guayama (1): Guayama. Guaynabo (1): Guaynabo. Hormigueros (1): Hormigueros. Isabela (1): Estación Experimental Agrícola Isabela; Isabela. Yayuba (1): Yayuba. Juana Diaz (2): Boca Chica; Juana Diaz. Lajas (5): La Parguera; Lajas; Refugio Nacional de Vida Silvestre de Laguna Cartagena. Las Piedras (1): Pueblito del Río. Loaiza (1): Puerto Maldonado. Luquito (1): Luquito. Manati (1): Manati. Mayagüez (8): Isla de Mona, Sardinera; Manat; Mayagüez; Santurce. Ponce (13): Bosque Estabal Toro Negro; Mercedita; Ponce. Quebradillas (1): Quebradillas. Rio Grande (2): Caribbean National Forest, El Verde; Mameyes. Salinas (3): Central Aguirre; Guyama; Salinas. San Germán (1): San German. San Juan (5): Hato Rey; Munos Rivera Park; Río Piedras; San Juan. Tao Baja (1): Tao Baja. Utuado (1): Utuado. Vega Baja (2): Reserva Natural Tortuguero; Vega Baja. Vieques (1): Puerto Road. Yabucoa (1): Yabucoa. Yauco (1): Yauco. SAINT BARTHÉLEMY (5). Saint Jean; no data. ST. KITTS AND NEVIS (21). No data (1). Nevis (5). Butlers Village; Cotton Ground Village; Hurricane Cove. St. Kitts (15). Baseterre; Bottom Mattingley Heights; Cunningham Village; Flamenco Disco; Lodge Estate; Molineux Estate; Monkey Hill; Mount Misery; Needsmut; Stapleton Estate; Wingfield Estate. ST. LUCIA (19). No data (1). Anse La Raye (2). Anse La Raye; Roseau. Castries (3). Castries; Vigie, Quarter of Castries. Micoud (3). Escap Community. Praslin (8). Mons Repos, Fox Grove Inn; Praslin. Soufrière (1). Soufrière (12 mi NW). Vieux Fort (1). Vieux Fort (1 mi NE). ST. MARTIN (3). Anse St. Marcel; Mullet Bay. ST. VINCENT (9). Charlotte Parish (3): 0.2 km W Old Sandy Bay. St. Andrew (2): Emeral Valley Hotel (E Layou); Pembroke. St. George (3): Cane Hall; Kingstown; Stubbs. St. Patrick (1): Wallilabou Bay. SURINAME: Babunsanti. TRINIDAD & TOBAGO (3). Tobago (1): John Parish, Charlotteville (3 km W). Trinidad (2): Caparo; Puerto España. UNITED STATES OF AMERICA (13). Florida (11): Broward; Dade; Highlands; Lee; Key West; Martin; Monroe (including Dry Tortugas); Palm Beach; St. Lucie. South Carolina (2): Edisto Island. UNITED STATES VIRGIN ISLANDS (25). Buck Island (1): Reef National Monument. St. Croix (20): Airport; Bonne Esperance; Canaan; Christiansted; Estate Cotton Garden & Long Point; Estate Jerusalem & Figtree Hill; Estate Sprat Hall; Estate Upper Love; Golden Grove; Hams Bluff; Kingshill; Sion Farm; Two Williams. St. John (4): Estate Carolina (NE Coral Bay); Estate Concordia; Lameshur Bay. St. Thomas (2): East St. Peter. VENEZUELA (19). Anzoátegui (2): El Hatillo; Este de Boca de Uchire. Apuré (1): Fundo Quitaparo, La Florida, Río Quitaparo. Aragua (1): Turiamo. Departamento Federal (1): Isla Las Aves. Falcón (5): Boca de Aroa; Chichiriviche; San Juan. Maracaibo (1): Isla Dorada. Miranda (4): Laguna de Tacangua; Las Mercedes, Paparo; Río Chico. Monagas (1): Caripe. Nueva Esparta (1): Las Marites. Zulia (2): Maracaibo. Natural history. Ligyrus cuniculus is considered an invasive species that affects turfgrasses in Florida, where is found in high abundance (Buss 2006). According to Chalumeau & Gruner (1977), it is a common species in Martinique and Guadeloupe, where adults devour the sugarcane cuttings, and larvae are mainly saproxylophagous, feeding on decomposing sugarcane, and sometimes on dung. In Brazil, it is low in abundance and no damage to plants has been reported.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD44FF9AAFA6181DFA81BB38.taxon	description	(Figs. 1 D, 2 A, 3 A, 11 C, 12 C, 13 C, 15 E, 17 E, 21 C, 25 B; 32)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD44FF9AAFA6181DFA81BB38.taxon	materials_examined	Male holotype (MZUSP) “ Ubatuba / São Paulo / 11. iii. 1967 / Moses / 5938 // Tomarus moroni / López-García & Deloya / HOLOTYPE ”. Female paratype (MZUSP) “ Ubatuba / São Paulo / 15. ii. 1968 / Moses / 7208 // Tomarus moroni / López-García & Deloya / PARATYPE ”. Male paratype (MZUSP) “ São Sebastião / SP / praia Baraqueçaba / 2. iv. 1963 / P. Araujo col. // Tomarus moroni / López-García & Deloya / PARATYPE ”. Female paratype (MZUSP) “ Atafona / Est. Rio Janeiro / Brasil / iii. 1964 / N. Menezes // Tomarus moroni / López-García & Deloya / PARATYPE ”. Female paratype (MZUSP) “ COLEÇÃO J. LANE // S. CATARINA / Florianopolis / R. G. Nossa / xii. 1957 / J. Lane col. // Ligyrus sp. 2 / det. Endrödi 1968 // Tomarus moroni / López-García & Deloya / PARATYPE ”. Female paratype (CEIOC) “ Coroa Grande / E. do Rio / Brasil / J-F. T. Freitas / 2 - 57 // CEIOC 32162 // Tomarus moroni / López-García & Deloya / PARATYPE ”. Male paratype (CEIOC) “ Ubatuba / S. Paulo / H. Ebert / 17 - ix- 62 // COLEÇAO FIOCRUZ // CEIOC 31535 // Tomarus moroni / López-García & Deloya / PARATYPE ”. Male paratype (CEIOC) “ Frade / R. J. / ii. 74 / A. Jesus // CEIOC 31546 // Tomarus moroni / López-García & Deloya / PARATYPE ”. Male and female paratypes (USNM) “ Mongagua / S. Paulo / XI. 1957 / 2901 // BRASIL / Halik 1966 / Collection // Tomarus moroni / López-García & Deloya / PARATYPE ”. Two male paratypes (USNM) “ Mongagua / S. Paulo / XI. 1957 / 2900 // Ligyrus / gibbosus / DeGeer // Halik / det. 1957 // BRASIL / Halik 1966 / Collection // Tomarus moroni / López-García & Deloya / PARATYPE ”. Male paratype (UNSM) “ from A. A. / Barbiellini / Sao Paulo // S //? fossor // Tomarus moroni / López-García & Deloya / PARATYPE ”. Female paratype (UNSM) “ from A. A. / Barbiellini / Sao Paulo // Tomarus moroni / López-García & Deloya / PARATYPE ” Type locality: Ubatuba, São Paulo, Brazil.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD44FF9AAFA6181DFA81BB38.taxon	description	Description. Habitus as in Fig. 25 B. Length 13.2 – 16.2 mm; humeral width 6.2 – 8.7 mm. Color dark reddish brown. Head: Frons coarsely rugopunctate, area between eyes nearly smooth. Frontoclypeal region with 2 tumescences (Fig. 2 D). Clypeus trapezoidal, base 1.5 times as wide as apex; apical angles acute (Fig. 2 D). Clypeal teeth conical, widely separated (1.5 times a tooth diameter) (Fig. 2 D). Mandible with 2 apical and 1 lateral acute large tooth, the first apical tooth small and joined to the second. Interocular distance equal to 3 times an eye width. Antennal club long, 1.5 times longer than antennomeres 2 – 7. Pronotum: Surface with minute, sparse punctures. Apical tubercle inconspicuous, rounded. Fovea small, as a shallow concavity behind tubercle (Fig. 2 D). Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with transverse, small tubercles forming 160 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 14 short, transverse, parallel carinae; carinae slightly wider at middle. Pygidium with large, dense, evenly distributed punctures. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate without an additional small basal tooth. Protarsus simple; inner claw simple. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia slightly crenulate, with 10 – 14 spinules. Male genitalia: Spiculum gastrale with apex straight. Parameres with sides nearly parallel, slightly contracted at the apical 4 th, apexes outwards, ventral margin with a small apical tooth (Figs. 15 E, 17 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD44FF9AAFA6181DFA81BB38.taxon	diagnosis	Diagnosis. Ligyrus moroni and L. cuniculus are similar in general appearance, shape of the mandible, tumescences on head, and small pronotal fovea and tubercle. However, L. moroni has the clypeal apical angles acute (rounded in L. cuniculus), the clypeal teeth conical (triangular to transverse in L. cuniculus), and the parameres with a small apical ventral tooth (Fig. 15 E), not present in L. cuniculus (Fig. 15 D).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD44FF9AAFA6181DFA81BB38.taxon	distribution	Distribution. Southern Atlantic coast of Brazil in the states of Rio de Janeiro, São Paulo, Santa Catarina, and probably Paraná. Locality records (Fig. 32). 25 examined specimens from CEIOC, MN / UFRJ, MZUSP, UNSM, USNM. BRAZIL (25). No data (1). Rio de Janeiro (4). Atafona; Coroa Grande; Rio de Janeiro; Frade. Santa Catarina (3). Florianápolis, P. G. Nossa; Itajai. São Paulo (17). Mongagua; São Paulo; São Sebastian; Ubatuba. Natural history. Specimens were collected from February to April and from September to December, which coincides with the warmer and rainiest months in this region of Brazil. It is found from sea level to 500 m.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	type_taxon	Type species: Scarabaeus relictus Say 1825: 194 by original designation.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	description	Description. (n = 885). Length 16.5 – 21.8 mm. Humeral width 8 – 11.5 mm. Head: Clypeus trapezoidal to subtriangular (Fig. 1 C). Apical clypeal teeth triangular and widely separated. Ocular canthus rounded, without dorsal setae, with 1 – 5 ventral setae. Frons with 2 tumescences. Mentum subrectangular (Fig. 2 E); not strongly narrowed at apex. Mentum and ligula separated by a suture. Ligula wide; covered with long, dense setae; ligular lobes not fused, with a narrow emargination. Maxillar apical palpomere 1.3 times longer than the second. Maxilla with galea rectangular (2.5 times longer than wide; Figs. 3 B, 4 C); teeth 3 and 4 diagonal (Fig. 3 E). Mandible tridentate (2 apical rounded and lateral long, acute to truncate lobe); apical teeth equal in shape. Labrum subquadrate (Fig. 6 C); apex straight; 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 rounded to triangular fovea. Elytra: Inner surface of apex with transverse small tubercles forming 70 – 77 parallel lines. Wings: RA with dense, large peg-like setae on medial ventral region; with peg-like setae on dorsal surface. Edge of RA 3 with dense setae. Abdomen: Apex of tergite IV with a diagonal line of 11 – 13 short, transverse, parallel carinae. Sternite VIII emarginate in male; entire in female. Pygidium rugose. Propygidium without stridulatory area. Venter: Prosternum very wide, convex. Propleura with long, sparse setae on anterior and posterior surfaces, medial surface glabrous. Apex of prosternal process suboval, flat, with a deep sulcus; without dorsal setae. Metepisternum with complete, thin carina; inner surface with deep, large punctures, and with long, dense setae; outer surface homogeneously rugose, with long, sparse setae. Metasternum with large, deep, confluent punctures and long, dense setae. Legs: Male protarsus modified; protarsomere 4 with a ventral, carinate, flat projection (Fig. 10 C – D); inner claw wide and truncate, with an acute, spine-like projection (Fig. 10 E). Protibia tridentate, with an additional basal denticle (Fig. 11 D – E), and sometimes a denticle between basal and second teeth, teeth equidistant. Apical margin of mesotibia and metatibia crenulate (Fig. 12 D, J). Female genitalia: Subcoxite 1.5 times wider than long (Fig. 13 D). Coxite subquadrate (as long as wide); surface flat. Subcoxite wider and longer than coxite. Male genitalia: Phallobase 1.6 times longer than parameres. Parameres without dorsal teeth, with ventral teeth. Internal sac with copulatory lamella and long lamellar spiny belt; accessory lamella U-shaped.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	diagnosis	Diagnosis. The subgenus Ligyrodes is differentiated by the following: Trapezoidal clypeus, clypeal teeth small; frontoclypeus with 2 tumescences (Fig. 1 C); mentum subrectangular (Fig. 2 E); mentum and ligula separate by a suture; ligula wide; covered with long, dense setae; ligular lobes fused, with a narrow emargination; galea with 3 dorsal and 2 ventral teeth (the basal often bifid); pronotum without fovea or tubercle; protibia tridentate, with teeth equidistant; protarsi of male modified, inner claw wide and truncate, with an acute like-spine projection (Fig. 10 E), protarsomere 4 with a ventral flat projection (Fig. 10 D); internal sac with copulatory lamella and long lamellar spiny belt; accessory lamella U-shaped (Fig. 21 D); surface of coxite flat (Fig. 13 D). Stridulatory apparatus consisting of a diagonal line of 11 – 13 short, transverse, parallel carinae on apex of sternite IV and apical inner surface of elytra with transverse small tubercles forming 70 – 77 parallel lines.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	synonymic_list	Composition. Three extant species: Ligyrus (Ligyrodes) relictus, L. (Ligyrodes) sallaei, and L. (Ligyrodes) peruvianus. Euligyrus ebenus, which was originally included in Ligyrodes by Casey (1915), is excluded from the subgenus Ligyrodes because although this species also has the protarsi enlarged in male, the shape of the tarsomeres and claws are very different, and there are many characters that support the inclusion of E. ebenus in the genus Euligyrus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	distribution	Geographic distribution. One species in North America (United States of America and Canada), one in North and Central America, and only one of its species in South America (restricted to a small area of the Pacific Coast of Ecuador and Peru).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD43FF9CAFA619C1FA22BD98.taxon	discussion	Phylogenetic relationships. The cladistic analysis of Tomarus (sensu lato) showed that this group is more closely related to Ligyrus than Euligyrus, contrary to the idea of Casey (1915). Taxonomic remarks. Casey (1915) described the genus Ligyrodes to group three species of Ligyrus and seven new species (all currently synonyms of Ligyrus (Ligyrodes) relictus and Ligyrus (Ligyrodes) sallaei) with the male protarsus modified, apex of prosternal process without setae, and the abdominal spiracles of the segments I – III transverse. The genus Ligyrodes was synonymized by Cartwright (1959), and Endrödi (1969) recognized it as a subgenus of Ligyrus but excluded E. ebenus, which he grouped in the subgenus Euligyrus along with E. similis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD42FF9CAFA61B1AFB8BBB30.taxon	description	(Figs. 1 C, 15 F, 25 C; 33)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD42FF9CAFA61B1AFB8BBB30.taxon	materials_examined	Male holotype (ZMHB) “ Peru, Zorritos / I-IV. 1926 / Rageuzut / Petersen J. / Schubart G. // Holotypus / Ligyrus / peruvianus / Endr. ”. Female allotype (ZMHB) “ Peru, Zorritos / I-IV. 1926 / Rageuzut / Petersen J. / Schubart G. // Allotypus / Ligyrus / peruvianus / Endr. ”. Male paratype (ZMHB) “ Peru, Zorritos / I-IV. 1926 / Rageuzut / Petersen J. / Schubart G. // Paratypus / Ligyrus / peruvianus / Endr. ” Male and female paratypes (HNHM) “ Peru, Zorritos / I-IV. 1926 / Rageuzut / Petersen J. / Schubart G. // Paratypus / Ligyrus / peruvianus / Endr. ” One paratype from Guayaquil, Ecuador at Endrödi’s collection (Endrödi 1970) not examined. Type locality: Zorritos, Tumbes, Peru.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD42FF9CAFA61B1AFB8BBB30.taxon	description	Description. Habitus as in Fig. 25 C. Length 15.6 – 19.0 mm; humeral width 8.4 – 10.2 mm. Color reddish brown. Head: Frons flat between eyes and before frontal tubercles; surface deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex only with sparse, small punctures. Frontoclypeal region with 2 tumescences separated by 1 / 2 a tumescence width (Fig. 1 C). Ocular canthus wide; with 4 – 5 ventral setae. Clypeal surface rugopunctate, striae denser than those on frons. Clypeus subtriangular, base 2.5 times as wide as apex (Fig. 1 C). Clypeal teeth transverse to widely triangular, separated by 1.5 times a tooth diameter. Mandible with 2 apical and 1 lateral truncate tooth, the tooth 1 shorter and narrower than tooth 2. Interocular distance equal to 4 times an eye width. Antennal club long. Pronotum: Surface with small, sparse punctures. Apex without tubercle or fovea. Elytra: First interval punctate, punctures similar in the other intervals. Inner surface of apex with small, transverse tubercles forming about 70 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 12 transverse, parallel, short carinae. Pygidial surface with large evenly sparse punctures, with small scarce rugosity on anterior corners. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, with an additional basal denticle; teeth equidistant. Protarsus of male with inner claw wide, apex with a spine-like projection (Fig. 10 E). Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate, with 13 – 17 spinules. Male genitalia: Spiculum gastrale wide, apex truncate. Parameres slender; strongly contracted at middle; ventral margin with an apical small tooth each side (Fig. 15 F).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD42FF9CAFA61B1AFB8BBB30.taxon	diagnosis	Diagnosis. Ligyrus peruvianus can be differentiated from the other species in the genus by the protibia with basal denticle and without a denticle between the basal and second tooth (as in L. relictus Fig. 11 E); pygidium with large evenly distributed punctures and with scarce rugosity on anterior corners; the clypeus subtriangular (Fig. 1 C); and the ventral tooth of the parameres at apex (at middle in the other species) (Fig. 15 F).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD42FF9CAFA61B1AFB8BBB30.taxon	distribution	Distribution. Ecuador and Peru (Endrödi 1970). Locality records (Fig. 33). 21 examined specimens from MERKB, MNHM, USNM, ZMUB. One record from Endrödi (1970). ECUADOR (15). Guayas (1): Guayaquil. Santa Helena (14): Chanduy. PERU (7). Piura (2): Tambogrande, Progreso Bajo. Tumbes (5): Zorritos. Natural history. Specimens were collected between 70 – 100 m, in dry coastal regions of Peru and Ecuador. All the specimens with date information were found at the end of rainy season (April).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD41FF9EAFA619C1FF5EBA14.taxon	description	(Figs. 2 E, 4 C, 5 C, 6 C, 8 C, F, 11 D, 14 C, 15 G, 17 F, 25 D; 33)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD41FF9EAFA619C1FF5EBA14.taxon	description	Description. Habitus as in Fig. 25 D. Length 16.7 – 22.2 mm; humeral width 8.0 – 11.48 mm. Color dark reddish brown to black. Head: Frons slightly concave between eyes and before frontal tubercles; surface deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex with only sparse, small punctures. Frontoclypeal region with 2 tumescences separated by 1 / 2 a tumescence width. Ocular canthus rounded and wide, with 3 – 4 ventral setae. Clypeal surface rugopunctate, striae denser than those on frons. Clypeus subtriangular to trapezoidal, base 2.0 – 2.5 times as wide as apex. Clypeal teeth transverse to widely triangular, separated by 1.5 times a tooth diameter. Mandible with 2 apical and 1 lateral, truncate tooth, tooth 1 shorter and narrower than tooth 2 (Fig. 5 C). Interocular distance 3 times an eye width. Antennal club long. Pronotum: Surface with small, sparse punctures. Apex without tubercle or fovea (Fig. 25 D). Elytra: First interval punctate, punctures similar to the other intervals. Inner surface of apex with small transverse tubercles forming about 70 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 12 transverse parallel short carinae. Pygidial surface with large, evenly sparse punctures. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, with an additional basal denticle; teeth equidistant (Fig. 11 D). Protarsus of male with inner claw wide, apex with a spine-like projection (Fig. 10 E). Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate, with 6 – 9 spinules. Male genitalia: Spiculum gastrale wide, apex truncate (Fig. 14 C). Parameres short; strongly contracted at middle; ventral margin with an apical acute, tooth long on each side (Figs. 15 G, 17 F).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD41FF9EAFA619C1FF5EBA14.taxon	diagnosis	Diagnosis. Ligyrus relictus is differentiated from the other species in the genus by the protibia with basal denticle and without a denticle between the basal and second tooth (Fig. 11 D) (with a denticle in L. sallaei); pygidium with dense rugosity; clypeus trapezoidal; and parameres short, with the ventral tooth acute and placed at middle (Figs. 15 G, 17 F). Taxonomic remarks. The species was originally described in the genus Scarabaeus by Say (1825). Although it has stridulatory area on the inner face of the elytra as the other species included by Burmeister (1847) in Ligyrus, for unknown reasons he placed it in Heteronychus and later LeConte (1856) combined it in Ligyrus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD41FF9EAFA619C1FF5EBA14.taxon	distribution	Distribution. Central and eastern United States to southeastern Canada (Cartwright 1959; Endrödi 1969; Ratcliffe & Cave 2017). Locality records (Fig. 33). 432 examined specimens from CEIOC, FSCA, IEXA, MZUSP, UNSM, USNM. Localities from Canada and United States of America counties are listed from Ratcliffe & Cave (2017). CANADA (518). Manitoba (79): Aweme; Souris; Winnipeg. New Brunswick (32): Fredericton; Kouchibouguac National Park; Saint Andrews, Huntsman Marine Science Centre; Saint John. New Scotia (139): Antigonish; Cape Breton; Halifax; Iverness; Kentville; Pictou; Victoria. Ontario (142): Arnprior; Aylmer; Britannia Heights; Canfield (4 km S); Carp; Chaffey’s Lock; Chalk River; Constance Bay; Cumberland Village; Dunks Bay; Dunnville; Dunrobin; Goderich, Maitland River and Highway 21; Grand Bend; Kerr Lake; Latta; London; Mallorytown; Marmora; Merivale, Midland; Muskoka; Normandale; Ottawa; Richmond (6 mi. W); Rondeau Provincial Park; Rostrevor; Sauble Falls, road along Bruce County forest; Severn Bridge; Sharbot Lake; Wolf Lake, Nipissing. Prince Edward Island (7): Charlottetown; Cornwall; Meadowbank. Quebec (119): Aylmer; Aylmer, Queen’s Park; Beamish Hill; Bécancour; Choisy; Eardley; Farrelton; Fort Coulonge; Hull; Île Perrot; Lac Cloutier; Lac Heney, Chermin Sharp; Lanoraie; Lévis; Limbour; Lucerne; Montreal; Norway Bay; Parc Paul-Sauve; Parc Paul-Sauvé, Deux-Montagnes; Perkins; Pont-Rouge; Rigaud; Riviera, Touraine; Sainte-Foy; Terrebonne; Thurso; Touraine; Vaudreuil; Ville Laval, Duvernay. UNITED STATES OF AMERICA (2,400). Alabama (1): Mobile. Arkansas (23): Hempstead, Washington. Colorado (28): Boulder; Denver; Jefferson; Larimer; Logan; Weld; Yuma. Connecticut (19): Fairfield; Litchfield; Middlesex; New Haven; New London; Tolland. Delaware (10): New Castle. District of Columbia (25): Washington. Illinois (168): Champaign; Cook; Douglas; Jasper; Kankakee; Lake; Macon; Marshall; Mason; McLean; Peoria; Pike; Rock Island; Saline; Vermilion. Indiana (60): Brown; Elkhart; Kosciusko; La Porte; LaGrange; Lake; Porter; Steuben; Sillivan; Tippecanoe; Vigo. Iowa (91): Appanoose; Boone; Bremer; Buchanan; Carroll; Cerro Gordo; Cherokee; Davis; Decatur; Deleware; Des Moines; Dickinson; Fremont; Henry; Jefferson; Johnson; Keokuk; Linn; Louisa; Lucas; Marion; Monona; Monroe; Muscatine; Page; Plymouth; Polk; Pottawattamie; Scott; Shelby; Story; Wapello; Winneshiek. Kansas (182): Atchison; Barber; Clark; Crawford; Dickinson; Doniphan; Douglas; Harvey; Kingman; Kiowa; Labette; Neosho; Pottawatomie; Reno; Riley; Sedgwick; Shawnee; Sherman; Stafford; Sumner; Wabaunsee. Louisiana (1): Grant. Maine (27): Aroostook; Cumberland; Hancock; Kennebec; Lincoln; Penobscot; Somerset. Maryland (31): Anne Arundel; Calvert; Cecil; Dorchester; Prince George’s; Queen Anne’s; St. Mary’s; Talbot. Massachusetts (73): Barnstable; Dukes; Middlesex; Nantucket; Norfolk; Suffolk. Michigan (191): Arenac; Barry; Bay; Berrien; Branch; Calhoun; Cheboygan; Huron; Ingham; Iosco; Kalamazoo; Kent; Lake; Lenawee; Livingston; Macomb; Midland; Monroe; Oakland; Oceana; Ottawa; Shiawassee; St. Joseph; Van Buren; Washtenaw; Wayne. Minnesota (13): Clay; Hennepin; Olmsted; Otter Trail; Ramsey. Missouri (182): Adair; Atchison; Bates; Boone; Buchanan; Caldwell; Callaway; Cedar; Cooper; Howard; Jackson; Pike; McDonald; Monroe; Osage; Randolph; St. Louis; Vernon. Montana (54): Carter; Custer; Powder River; Richland; Roosevelt; Rosebud; Treasure; Yellowstone. Nebraska (275): Antelope; Blaine; Boone; Box Butte; Boyd; Brown; Buffalo; Cass; Cherry; Custer; Dakota; Dixon; Douglas; Dundy; Franklin; Gage; Garfield; Grant; Greeley; Hall; Hamilton; Harlan; Holt; Jefferson; Keith; Kimball; Lancaster; Lincoln; McPherson; Merrick; Nemaha; Otoe; Red Willow; Sarpy; Saunders; Scotts Bluff; Thomas; Valley; Wayne; Wheeler. New Hampshire (17): Rockingham; Strafford. New Jersey (42): Atlantic; Bergen; Cape May; Gloucester; Middlesex; Monmouth; Morris; Ocean; Salem; Union. New York (63): Clinton; Essex; Nassau; Onandaga; Orange; Queens; Richmond; St. Lawrence; Suffolk; Westchester. North Dakota (372): Adams; Barnes; Burleigh; Cass; Grand Forks; Morton; Ransom; Richland; Slope; Stutsman; Wells; Williams. Ohio (13): Athens; Auglaize; Champaign; Defiance; Erie; Fairfield; Stark. Oklahoma (62): Caddo; Canadian; Cleveland; Kay; Latimer; Lincoln; Logan; McClain; Osage; Ottawa; Payne; Seminole; Texas; Tulsa; Wagoner; Woods. Pennsylvania (79): Allegheny; Armstrong; Beaver; Centre; Clearfield; Crawford; Dauphin; Fayette; Forest; Lycoming; Monroe; Montgomery; Northampton; Philadephia; Venango; Westmoreland; York. Rhode Island (11): Kent; Providence; Washington. South Dakota (144): Bennett; Bon Homme; Brookings; Brown; Brule; Buffalo; Clay; Cluster; Gregory; Haakon; Hughes; Hyde; Jones; Kingsbury; Marshall; Oglala; Pennington; Tripp; Yankton; Texas (14): Bexar; Cameron; Hardin; Hemphill; Hidalgo; Kleberg; McLennan; Travis; Wheeler. Vermont (44): Addison; Chittenden; Grand Isle; Windham. Virginia (15): Arlington; Caroline; Chesapeake; Chesterfield; King George; Northampton. West Virginia (5): Mason; Preston. Wisconsin (58): Clark; Columbia; Dane; Grant; Jackson; Jefferson; Kenosha; Marinette; Milwaukee; Monroe; Polk; Racine; Richland; Sauk; Walworth; Waukesha; Winnebago; Wood. Wyoming (5): Campbell; Platte. Natural history. The larvae can be found under piles of straw and decaying hay or horse or cattle dung (Hayes 1925; Ritcher 1966). The eggs are laid between April and May, and adults emerge in August, so winter is passed as an adult under soil; it is a beneficial species that contributes to the decomposition of organic material (Hayes 1929).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD40FFA3AFA61F91FA87BEC8.taxon	description	(Figs. 3 B, E, 10 C – E, 11 E, 12 D, J, 13 D, 15 H, 17 G, 21 D, 25 E; 33)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD40FFA3AFA61F91FA87BEC8.taxon	materials_examined	Female lectotype designated by Cartwright (1959) (BMNH) “ LECTO- / TYPE // TYPE // Sp. figured. // Almolonga, / Mexico. / Hoege. // Ligyrus / sallaei / Bates / B. C. A., Col., II (2) / Ligyrus / sallaei. // Lectotype / B. sallaei Bates / Cartwright 1959 ”. Male paralectotype (BMNH) “ PARA- / LECTO- / TYPE // TYPE // mexico city / H. H. S. // Ligyrus / sallaei / Bates ♂ // B. C. A., Col., II (2) / Ligyrus / sallaei. ” Female paralectotype (BMNH) “ PARA- / LECTO- / TYPE // Cubulco, / Vera Paz. / Champion. // B. C. A., Col., II (2) / Ligyrus / sallaei. ” Two female paralectotypes (BMNH) “ PARA- / LECTO- / TYPE // Mexico / City / Hoege. // Ligyrus / sallaei ♀ // B. C. A., Col., II (2) / Ligyrus / sallaei. ” Male paralectotype (MNHN) “ Mexico // Ligyrus / sallaei / Bates ♂ // Ex-Musaeo / H. W. Bates / 1892 // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2015 // PARALECTOTYPE // MNHN / EC 6740 ”. Female paralectotype (MNHN) “ Mexico // Ligyrus / sallaei / Bates ♀ // Ex-Musaeo / H. W. Bates / 1892 // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2015 // PARALECTOTYPE // MNHN / EC 6741 ”. Male paralectotype (MNHN) “ Mexico. / Hoege. // Ligyrus / sallaei / Bates ♂ // Ex-Musaeo / H. W. Bates / 1892 // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2014 // PARALECTOTYPE // MNHN / EC 6742 ”. Male paralectotype (MNHN) “ Mexico / City. / Hoege. // Ligyrus / sallaei / Bates // H. W. Bates / Biol. Cent. Amer. // Museum Paris / ex Coll. / R. Oberthur // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2014 // PARALECTOTYPE // MNHN / EC 6743 ”. Female paralectotype (MNHN) “ Oaxaca, / Mexico. / Hoege. // Ligyrus / sallaei / Bates // H. W. Bates / Biol. Cent. Amer. // Museum Paris / ex Coll. / R. Oberthur // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2015 // PARALECTOTYPE // MNHN / EC 6744 ”. Male paralectotype (MNHN) “ Jalapa // Ligyrus / sallaei / Bates // Ex-Musaeo / H. W. Bates / 1892 // Museum Paris / ex Coll. / R. Oberthur // PARALECTOTYPE / Ligyrus / sallaei Bates, 1888 // PARALECTOTYPE // Tomarus sallaei / (Bates, 1888) / det F. Dupuis 2014 // MNHN / EC 6745 ”. Another 11 paralectotypes at MNHN not examined. Type locality: Almolonga, Mexico. Ligyrodes propinquus Casey, 1915: 183. Synonym. Male lectotype (USNM) “ San Angel / D. F. Mex / Wickham // CASEY / bequest / 1925 // TYPE USNM / 48606 // propinquus / Csy ” here designated. Female paralectotype (USNM) “ San Angel / D. F. Mex / Wickhem // CASEY / bequest / 1925 // PARATYPE USNM / 48606 // propinquus / Csy ”. Type locality: San Angel, Mexico. Ligyrodes aztecus Casey, 1915: 183. Synonym. Male lectotype (USNM) “ Cuernavaca / Mor. Mex. / Wickhem // CASEY / bequest / 1925 // TYPE USNM / 48605 // aztecus / Csy ” here designated. Type locality: Cuernavaca, Morelos, Mexico.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD40FFA3AFA61F91FA87BEC8.taxon	description	Description. Habitus as in Fig. 25 E. Length 18.5 – 21.2 mm; humeral width 10.4 – 11.5 mm. Color dark reddish brown to black. Head: Frons slightly concave between eyes and before frontal tubercles; surface deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex with only sparse, small punctures. Frontoclypeal region with 2 tumescences separated by 1 / 2 a tumescence width. Ocular canthus rounded and wide; with 4 – 5 ventral setae. Clypeal surface rugopunctate, striae denser than those on frons. Clypeus trapezoidal, base 2 times wider than apex. Clypeal teeth transverse to widely triangular, separated by 1.5 times a tooth diameter. Mandible with 2 apical and 1 lateral truncate tooth, tooth 1 shorter and narrower than tooth 2. Interocular distance equal to 3 times an eye width. Antennal club long. Pronotum: Surface with small, sparse punctures. Apex without tubercle or fovea. Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with small transverse tubercles forming about 77 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 13 transverse, parallel, short carinae. Pygidial surface with rugosity on base and anterior corners or with dense, large punctures (female). Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, with a basal denticle and another denticle between basal and second tooth; teeth equidistant (Fig. 11 E). Protarsus of male with inner claw wide, apex with a spine-like projection (Fig. 10 C – E). Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate, with 6 – 7 spinules. Male genitalia: Spiculum gastrale wide, apex truncate. Parameres long, slightly contracted at middle, ventral margin with an apical rounded tooth each side (Figs. 15 H, 17 G).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD40FFA3AFA61F91FA87BEC8.taxon	diagnosis	Diagnosis. Ligyrus sallaei can be differentiated from the other species in the genus by the protibia with a basal denticle and another denticle between the basal and second tooth (Fig. 11 E); pygidium with dense to sparse rugosity on base and anterior angles; clypeus trapezoidal; and parameres long, with the ventral tooth rounded and placed at the middle (Figs. 15 H, 17 G).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD40FFA3AFA61F91FA87BEC8.taxon	distribution	Distribution. Southern United States, Mexico, Guatemala, Nicaragua, El Salvador, Honduras, and Costa Rica. Locality records (Fig. 33). 496 examined specimens from BMNH, CEIOC, CERPE, CIIDIR, CNIN, CZUG, EMAC, FSCA, IEXA, ITVO, UNSM, USNM. Some records from Ratcliffe (2003), Ratcliffe & Cave (2006, 2017), and Ratcliffe et al. (2013). Only counties are listed for the United States of America. COSTA RICA (16). Alajuela (1): Palmareña. Guanacaste (9): Agua Buena; Finca Jenny; Parque Nacional Guanacaste (Estaciones Lomas Barbudal, Murciélago, Palo Verde); La Pacífica; Nacaome; Playa Naranjo; Tierras Morenas. Puntarenas (5): Reserva Carara; Estación La Casona; Estación Sirena; Monteverde. San José (1): Escazú. EL SALVADOR (72). Ahuachapan (2): El Imposible. Cabañas (16): Cinquera. Chalatenango (1): La Montaña. La Libertad (2): San Andrés. Morazán (44): Cerro Cacahuatique. San Salvador (1): San Salvador. Santa Ana (6): Parque Los Andes; Parque Montecristo. GUATEMALA (97). Alta Verapaz (3): Coban; Senahu. Tactic. Baja Verapaz (7): Chilasco; Cubulco; Niño Perdido; Purulha. Chimaltenango (2): Chimaltenango; Tecpan. Chiquimula (6): Chanmagua; Cerro Montecristo; Concepción Las Minas; Esquipulas. El Progreso (2): El Rancho. Finca El Riscosa. Escuintla (5): Finca Eminencia; Finca Santa Cecilia; La Gomera; Palin; Sipacate. Guatemala (12): Amatitlán; Fraijanes; Guatemala; Mixco; Puerta Tarada; San José Pinula; Santa Catarina. Huehuetenango (16): Chiantla; Jacaltenango; La Democracia; La Laguna; Nenton. Izabal (2): Finca Firmeza; Morales. Jalapa (3): Jalapa; Mataquescuintla; Miramundo. Petén (1): Boca del Monte. Quetzaltenango (1): Colomba Costa Cuca. Sacatepéquez (4): Antigua Guatemala; Cerro Alux; San Lucas Sacatepéquez. San Marcos (1): San Rafael Pie de Cuesta. Santa Rosa (6): Barberena; Laguna El Pino; Los Esclavos; Mal País; Pueblo Nuevo; Chiquimulilla. Sololá (3): Panajachel; San José Chacaya; Santa María Visitación. Suchitepéquez (10): Finca El Vesubio; Los Tarrales; Santa Bárbara. Zacapa (11): La Unión; Río Hondo; San José; San Lorenzo; Teculután; Usumatlán. HONDURAS (81). Atlántida (20): Curla; La Ceiba; Parque Nacional Pico Bonito. Comayagua (5): La Soledad; Siguatepeque. Copán (2): La Entrada; San Agustín. Cortés (2): Lago de Yojoa; Parque Nacional Cusuco. El Paraiso (8): Barrio Las Flores; Capire; Cifuentes; Morocelí. Francisco Morazán (10): El Zamorano. La Paz (2): Jesús de Otoro; Reserva Guajiquiro. Lempira (3): Erandique. Ocotepeque (7): El Portillo, Nueva Ocotepeque; Reserva Guisayote. Olancho (12): Montaña del Melacate; Parque Nacional La Muralla. Yoro (10): Linda Vista; Parque Nacional Pico Pijol; San Rafael; Suyapa Victoria. MEXICO (866). Aguascalientes (6): Aguascalientes; Calvillo; Pabellón de Arteaga. Chiapas (82): Aguacero; Bonampak; Chicoasén; Cinco Cerros; Cintalapa; Comitán; Cuxtepeques; El Chorreadero; El Jardín; Francisco Villa; Ixtapa; Laguna Bélgica Lagunas de Montebello; Las Rosas; Los Ángeles; Mirador El Caminero; Palenque; Posas de Berriozabal; Rizo de Oro; Simojovel; Teopisca; Tuxtla Gutiérrez; Villaflores. Chihuahua (1): Puente Los Hoyos. Coahuila (4): La Babia; Rancho Golondrina; Río Sabinas; Saltillo. Colima (5): Laguna El Jabalí. Distrito Federal (10): Chapultepec; Guadalupe; Los Dinamos; Xochimilco. Durango (1): Canelas. Guanajuato (4): Celaya; Guanajuato; Irapuato. Guerrero (38): Atoyac; Acapulco; Ahuacuotzingo; Almolonga; Chilapa de Álvarez; Chilpancingo; Iguala; Juan R. Escudero; Las Granadas; Malinaltepec; Mochitlán; Ocotito; Olinalá; Papanoa; Puerto de Gallo; Quelchultenango; Taxco de Alarcón; Teloloapan; Tixtla; Villa Hermosa. Hidalgo (30): Actopan; Cañón Metztitlán (Venados) Chapulhuacan; Guerrero Mill; Ixmiquilpan; Las Milpullas; Mezquititlán; Molango; Otongo; Tepeji del Río; Tianguiztango; Zacualtipán. Jalisco (67): Acatic; Ajijic; Amatitán; Ameca; Arroyo Colorado; Autlan; Cañada Río Mismaloya; Casimiro Castillo; Chapala; Cuitzmala; El Tamarindo; Guadalajara; Las Barrancas; Magdalena (14 km W); Mascota; Melaque; Mixtlan; Ocotlán; Ocuililla; Paso Los Mazos; Poncitlán; San Antonio; Sierra de Manantlán; Talpa de Allende; Tecolotlán; Tepatitlán; Tequila; Tizapán; Zapiltitic; Zapopan; México (58): Amecameca. Atizapan; Avandaro; Chalma, Calmita; Chapingo; Coacalco; Coatepec Harinas; Culhuacán; Ixtapan de Oro; Malinanco; Malinantenango; Nanchititla; Naucalpan; Ocoyoacac; Parque El Salto de Chihuahua; San José Deguedo; San José Villa de Allende; Sierra de Nanchititla; Temascaltepec; Texcoco; Tonatico; Valle de Bravo; Villa Corono; Villa de Allende. Michoacán (47): Acambaro; Agua Blanca; Apatzingan; Aquila; Cerro El Águila; Coalcoman; Cojumatlan; Cotija; El Sabino; Jerecuaro; Jiquilpan; Jungapeo de Juárez; Juquilpan; Lago Chapala; Laureles; Morelia; San Lorenzo; Senguio; Tacambaro; Tuxpan; Tzararacua; Uruapan; Villa Victoria. Morelos (65): Cañón de Lobos; Chamilpa; Cuautla; Cuernavaca; Jantetelco; Jojutla; Juitepec; Ignacio Bastida (2 km S); Oaxtepec; Ocuituco; Tepoztlán; Tlaquiltenango; Tlayacapan; Valle de Vásquez; Vista Hermosa; Xochicalco; Xochitepec; Yautepec; Zacatepec; Zacualpan. Nayarit (57): Acaponeta (21 mi. S); Agua Milpa; Chapalilla; Compostela; El Pichón; El Venado; El Zopilote; Huajicori (6.2 mi. N); Jesús María; Las Adjuntas; Los Sabinos; Pochotitán; Río Huaynamota; Río Santiago; San Blas; Santa María del Oro; Tepetiltic; Tepic; Volcán Ceboruco. Nuevo León (7): Linares; Los Rayones; Monterrey (5 mi. S). Oaxaca (82): Candelaria Loxicha; Capulalpam; Cuilapam de Guerrero; El Camarón; El Polvorín; Gelatao; Ixtlán de Juárez; Juquila Mixes; Oaxaca; Ocotlán; Pueblo Nuevo; San Bartolo Coyotepec; San José El Paraiso; San Juan de los Cues; San Pablo Villa de Mitla; Santa Catarina Juquila; Santa María El Tule; Santa María Nativitas Coatlán; Santa Rosa; Santiago Xiacui; Santiago Yosondua; Santo Reyes Papalo; Sola de Vega; Taltepec; Tehuantepec; Teotitlán; Tlacolula; Toltepec; Valle Nacional; Xoxocotlán. Puebla (48): Acatzingo de Hidalgo; Cacaloapan; Coxcatlán; Cuetzalan; El Aguacate; Hueyacan; Izúcar de Matamoros; Nuevo Necaxa; Patla; Puebla; Petlalcingo (3 mi. S); Santiago Mihuatlán; Santo Domingo Huehuetlan; Tehuacán; Tehuetzingo; Tehuixtla; Tlacotepec; Villa Juárez. Querétaro (13): Querétaro; San Juan del Río; Santa Rosa. Quintana Roo (3): Cobá. San Luís Potosí (28): Ciudad Valles; Ciudad del Maíz; El Aguacate; El Pujal; El Salto; Los Cuates; Los Cerritos; Maguey de Oriente; Matehuala; Río Verde; Santa María del Río; Sierra La Trinidad; Tamasopo; Tamazunchale; Valle de los Fantasmas. Sinaloa (53): Capilla de Taxte; Copala; Culiacán; El Batel; El Fuerte (12 mi. NE); El Zapotillo; Guamuchil (11 mi. S); Loberas; Los Chirimoyos; Los Mochis; Mazatlán; Piedra Blanca; Potrerillos; Rosario; San Ignacio (2.5 mi. S); San Lorenzo; Santa Lucía; Vado Hondo; Venadillo; Villa Unión (26 mi. NE). Sonora (43): Aconchi; Álamos; Ciudad Obregón; La Palmita; Moctezuma (14 mi. N); Navajoa; Rancho Los Alisos; Rancho Nuevo; Río Cuchujaqui; Rosario de Tesopaco; Santa Ana (45 mi. S); Yecora (12 mi. W). Tabasco (3): El Triunfo; Malpasito; Ocotlán. Tamaulipas (21): Bocatoma; Ciudad Victoria; El Salto; Estación Biológica Los Cedros; Galeana; Gómez Farias; Jaumave; Las Norias (5 km E); Palmillas; Reserva de la Biosfera El Cielo; Río Bravo; Salamanca; Soto La Marina; Tula (10 km E). Veracruz (84): Actopan; Acultzingo; Boca Andrea; Catemaco; Cerro del Vigía; Coatepec; Coatzacoalcos; Córdoba; Cotaxtla; Calcahualco; Cuitláhuac; Dos Amates; Dos de Abril; Fortín de las Flores; Huatusco; Ixhuatlán del Café; La Gloria Cardel; La Mancha; Las Trancas; Los Tuxtlas; Monte Blanca; Ocotal Chico; Orizaba; Paraje Nuevo; Peñuela; Playa Escondida; Puente Nacional; Rinconada (6 mi. SE); San Andrés Tuxtla; Tecolutla; Tlalnelhuayocan; Totoapan; Totutla; Tuzampan; Uxpanapa; Villa Rica; Volcán San Martín; Xalapa; Yucatán (2): Chichén Itzá; Mérida. Zacatecas (5): Apozol; Nochistlan; Zacatecas (8 mi. E). NICARAGUA (6). Esteli (2): Cerro Tomabu. Jinotega (3): Cerro Kilambe. Madriz (1): Somuto San Lucas. UNITED STATES OF AMERICA (45). Arizona (1): Santa Cruz. Texas (44): Atascosa; Bastrop; Bexar; Brazos; Brooks; Burleson; Caldwell; Cameron; Comal; Dimmit; Frio; Gonzales; Guadalupe; Hidalgo; Houston; Live Oak; McMullen; McLennon; Medina; Nueces; San Patricio; Sequin; Starr; Tavis; Val Verde. Natural history. Their larval stages were described by Ramírez-Salinas et al. (2004) who indicated that the larvae can be found under sheep dung or coffee pulp compost. In the material examined, there are several records of adults found in dung galleries. They are predominantly found in pine / oak forests, cloud forests, and coffee plantations. Specimens have been found at 40 m elevation, but they are more abundant between 1,000 – 2,500 m.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7DFFA2AFA61BACFA23BB58.taxon	type_taxon	Type species: Scarabaeus gibbosus De Geer, 1774 by subsequent designation (Casey 1915: 178).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7DFFA2AFA61BACFA23BB58.taxon	description	Description. (n = 4,381). Length 11.3 – 18.0 mm. Humeral width 5.9 – 10.0 mm. Head: Clypeus triangular to subtriangular (Fig. 1 E – G). Apex of clypeus with 1 or 2 triangular teeth, not or slightly separated. Ocular canthus rounded to acute, with ventral setae, without dorsal setae. Frons with carina, entire or interrupted at middle. Mentum triangular (Fig. 2 B – D), surface flat to strongly convex. Mentum and ligula separate. Ligula reduced, covered with short, sparse setae; ligular lobes fused, apex straight. Maxillary apical palpomere 2.0 – 2.5 times longer than the second. Maxilla with galea rectangular (3.0 – 3.5 times longer than wide) (Figs. 3 C, 4 D – E); with 3 – 4 well-developed teeth (Fig. 3 H), sometimes with 1 – 2 rudimentary teeth (Fig. 3 G). Mandible with 2 apical teeth and a lateral widely rounded lobe (Fig. 5 D – E); tooth 1 with apex acute, longer and narrower than tooth 2. Labrum subquadrate (as long as wide); apex rounded to slightly bilobed; with long, dense setae (Fig. 6 A – B). Antenna with 10 antennomeres, club very long (2 times longer than antennomeres 2 – 7). Pronotum: Anterior marginal bead complete. Apex without tubercle or with acute tubercle; without fovea or with deep, wide fovea. Elytra: Inner surface of apex with transverse small tubercles forming 50 – 115 parallel lines (Fig. 8 E – F). Wings: RA with dense, large peg-like setae on medial ventral region (Fig. 7 E); with peg-like setae on dorsal surface. Edge of RA 3 with dense to sparse setae. Abdomen: Apex of tergite IV with a diagonal line of 9 – 11 short to large, transverse, parallel carinae (Fig. 8 B – C). Sternite VIII emarginate in male; entire in female. Pygidium without rugosity (Fig. 9 C – E). Propygidium without stridulatory area. Venter: Prosternum narrow, acute to slightly rounded. Propleura with long, dense setae on all the surface, sparser on medial area. Apex of prosternal process spherical to slightly oval, with dorsal, long setae. Metepisternum with incomplete, very thin carina; inner surface with deep, large punctures, with long setae; outer surface homogeneously rugose, with long, dense setae. Metasternum with small, deep, confluent punctures, with long, dense setae. Legs: Male protarsus simple; inner claw simple. Protibia tridentate; basal tooth slightly distant from others (Fig. 11 F – G). Apical margin of mesotibia and metatibia entire (Fig. 12 E – F). Female genitalia: Subcoxite 1.5 – 2.0 times wider than long (Fig. 13 E – G). Coxite subquadrate to subrectangular; surface flat to strongly concave. Subcoxite 2 times wider and longer than or subequal in length to coxite. Male genitalia: Phallobase 1.2 – 1.3 times longer than parameres. Parameres without dorsal teeth (rarely with a dorsal long tooth), with ventral tooth. Internal sac with copulatory lamella and short to long lamellar spiny belt; accessory lamella irregular (Fig. 21 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7DFFA2AFA61BACFA23BB58.taxon	diagnosis	Diagnosis. The subgenus Ligyrus can be differentiated by the following: Triangular to subtriangular clypeus (Fig. 1 E – G), clypeal teeth triangular to transverse; frontoclypeus with carina entire or interrupted at middle (Fig. 1 E – G); mentum triangular (Fig. 2 B – D); mentum and ligula separate; ligula reduced; covered with short, sparse setae; ligular lobes fused; galea with 3 – 4 well-developed teeth, sometimes with 1 – 2 rudimentary teeth (Fig. 3 C, G, H); antennal club very long; pronotum with or without fovea or tubercle; protibia tridentate, basal tooth slightly distant from others (Fig. 11 F – G); protarsus of male simple, inner claw simple; internal sac with copulatory lamella and short to long lamellar spiny belt; accessory lamella irregular (Fig. 21 E); stridulatory apparatus consisting of a diagonal line of 9 – 11 short to large, transverse, parallel carinae on apex of sternite IV and apical inner surface of elytra with transverse small tubercles forming 50 – 115 parallel lines.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7DFFA2AFA61BACFA23BB58.taxon	synonymic_list	Composition. Thirteen species are recognized as members of the subgenus Ligyrus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7DFFA2AFA61BACFA23BB58.taxon	distribution	Geographic distribution. This is the most widely distributed subgenus with species ranging from Argentina to Canada, including the West Indies. Species are mainly found in dry ecosystems such as deserts and dry forests.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7BFFA4AFA619C1FB95BB11.taxon	description	(Figs. 2 B, 3 H, 11 F, 12 E, K, 13 E, 15 I, 21 E, 25 F; 34)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7BFFA4AFA619C1FB95BB11.taxon	materials_examined	Type material. Male holotype (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, 360 / msnm, 24 - vii- 12, / Alumbrado público / Héctor Calvo Gatica // Ligyrus allonasutus / López-García & Deloya / HOLOTYPE ”. Female allotype (IEXA) labeled as holotype and with yellow label “ Ligyrus allonasutus / López-García & Deloya / ALLOTYPE ”. Paratypes (6 ♂, 5 ♀): One female labeled as holotype. One male and two females (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, 360 / msnm, 01 - 31 - VIII- 12, / Alumbrado público / Héctor Calvo Gatica ” Two males and one female (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, 360 / msnm, 25 - VII- 12, / Alumbrado público / Héctor Calvo Gatica ” One male (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, 360 / msnm, 08 - IX- 12, / Alumbrado público / Héctor Calvo Gatica ” One male (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, Luz / mercurial, 350 m. / 01 - 30 - VI- 12, / Héctor Calvo Gatica ” One male (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, 360 / msnm, 25 - VIII- 12, / Excremento bovino / Héctor Calvo Gatica ” One female (IEXA) “ México, Guerrero, Juan R. / Escudero, El Terrero, Luz / mercurial, 350 m. / 25 - V- 12, / Héctor Calvo Gatica ” All paratypes with yellow label “ Ligyrus allonasutus / López-García & Deloya / PARATYPE ”.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7BFFA4AFA619C1FB95BB11.taxon	description	Description. Holotype. Habitus as in Fig. 25 F. Length 16.1 mm; humeral width 8.2 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and with only sparse, small punctures. Frontoclypeal carina straight, complete. Ocular canthus acute, with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex. Clypeal teeth triangular, separated by 1 tooth diameter. Mandible with 2 apical and 1 lateral, acute tooth, tooth 1 longer and narrower than tooth 2. Maxilla rectangular, 3.5 times longer than wide; galea with 3 dorsal teeth. Apex of labrum evenly rounded. Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, dense punctures. Apex without tubercle or fovea. Elytra: First interval punctate, punctures similar to the other intervals. Inner surface of apex with small transverse tubercles forming about 70 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 11 transverse, parallel, short carinae. Pygidial surface with small punctures, denser on base and anterior corners. Surface in lateral view strongly convex. Legs: Protibia tridentate; basal tooth slightly distant from others (Fig. 11 F). Protibial surface sparsely punctate. Protarsus of male simple, inner claw simple. Metatibia strongly contracted at apex (Fig. 12 K). Apex of metatibia entire, with 13 spinules. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with sides nearly parallel, lateral ventral teeth short (Figs. 15 I, 18 A). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thickened, elongate (Fig. 21 E). Allotype. Length 16.8 mm; humeral width 8.9 mm. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subrectangular (wider than long); surface strongly concave. Subcoxite wider and longer than coxite (Fig. 13 E). Pygidium: Surface in lateral view strongly flat. Legs: Apex of metatibia with 19 spinules. Variation. Eleven paratypes. Length 14.8 – 17.4 mm; humeral width 7.82 – 8.98 mm. Legs: Apex of metatibia with 13 – 20 spinules.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7BFFA4AFA619C1FB95BB11.taxon	diagnosis	Diagnosis. Ligyrus allonasutus can be recognized by the triangular clypeus; frontal carina straight, not interrupted; mandible with outer margin acute; pronotum without tubercle or fovea, without an apical swelling; pronotal punctation deep and large; metatibia elongate with apex strongly narrowed; parameres with sides nearly parallel; lateral ventral teeth short (Figs. 15 I, 18 A); internal sac with accessory lamella thick and elongate (Fig. 21 E); coxite of female strongly concave (Fig. 13 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD7BFFA4AFA619C1FB95BB11.taxon	etymology	Etymology. The name “ allonasutus ” comes from the Greek allós (other or different) and the name nasutus referring to the species from which the new species was separated (see taxonomic remarks). Taxonomic remarks. The new species is purposed here to place populations from western Mexico to Panama, which have been traditionally known as Ligyrus nasutus. The species was described by Burmeister (1847) with two females from Yucatán and Guatemala. Endrödi (1969) choose the female from Yucatán (in bad conditions) as the lectotype for L. nasutus, but his descriptions and illustrations are based only on populations from western Mexico and Guatemala to Panama. After his work, the name L. nasutus was widely used for supposedly a unique species with wide distribution, but there was confusion since the original description as Burmeister (1847) included two female specimens from different species as types for L. nasutus. However, as the lectotype of L. nasutus is from Yucatán, only populations of this region correspond to L. nasutus, and L. allonasutus is proposed here for the remaining populations (western Mexico and Central America). Geographic distribution. Ligyrus allonasutus is widely distributed from western Mexico to Panama, between sea level and 1,850 m. Locality records (Fig. 34). 383 examined specimens from CAFESI, CIIDIR, CZUG, EMAC, IEXA, MN / UFRJ, UNSM, USNM. Some data from Ratcliffe (2003), Ratcliffe & Cave (2006), and Ratcliffe et al. (2013). COSTA RICA (20). Alajuela (1): Caño Negro. Guanacaste (16): Cañas; Barra Honda; Estación Enrique Jiménez Nuñes; Estación Almendros; Estación Murciélagos; Estación Palo Verde; Finca Jenny; Finca La Pacífica; Lomas Barbudal; Playa Naranjo; Reserva Carara; Reserva Monteverde; Río San Lorenzo; Santa Rosa; Tierras Morenas. EL SALVADOR (68). Ahuachapán (6): Guaymango; Izcaquillo; Parque Nacional El Imposible. Cabañas (33): Cinquera; Ilobasco. Cuscatlán (2): El Playón (7 km NE Coctepeque); San Pedro Perulapán; La Libertad (14): Canton El Barillo; La Libertad; Parque Natural Walter Thilo Deininger; San Diego; San Andrés; Santa Tecla; Valle Zapotitán; Zaragoza. La Paz (5): Campo San Luís Talpa; Caprex; Zacatecoluca. La Unión (1): La Unión. Morazán (1): Arambala; Río Sapo. San Miguel (1): Laguna El Jocotal. San Salvador (3): Parque Saburo Hirao; San Salvador. Santa Ana (3): Los Planes; San José Ingenio. Sonsonate (1): Sonsonate. GUATEMALA (104). Baja Verapaz (5): Niño Perdido; Rabinal; Salama; San Jerónimo. Chiquimula (1): Quetzaltepeque. El Progreso (3): El Rancho; San Agustín Acasaguastlán. Escuintla (17): Chulamar; Likin; Peñón de Siquinalá; Puerto de San José; Santa Lucía Cotzumalguapa; Santa Rosa; Taxisco (86 km N); Tiquisate. Guatemala (10): Amatitlán; Ciudad de Guatemala; Puerto Parada; San Jerónimo; Villa Canales. Huehuetenango (4): Jacaltenango; La Laguna; Nenton. Jalapa (1): Jalapa. Jutiapa (6): Cañón ed Monjoy; Comapa; Jutiapa (4 mi. E); Quezada. Quetzaltenango (1): Coatepeque. Retalhuleu (8): Retalhuleu. Sacatepéquez (2): San Lucas de Sacatepéquez; Santo Domingo. San Marcos (4): Ayutla; Ixbén. Santa Rosa (8): Chiquimulilla; Cuilapa; Monterrico; Taxisco. Suchitepéquez (13): Cuyotenango; Finca Reforma; Ixtacapa; Mazatenango; Reserva Los Tarrales; San Juan Bautista; Santa Bárbara. Zacapa (21): Río Hondo; Río Teculután; San José (3 km SE); San Lorenzo; Santa Cruz; Tres Marías. HONDURAS (555). Atlántida (302): Carretera a Cortés; Curla (8 km NW La Ceiba); Islita (60 km La Ceiba); La Ceiba; Tela. Choluteca (33): Choluteca; Duyure; Pespire; Tierra Blanca. Comayagua (49): Comayagua; Siguatepeque; Taladro. Cortés (39): San Pedro Sula. El Paraiso (101): Barrio Las Flores; Capire (8.3 km SE); Danlí, El Barro; Morocelí (10 km W). Francisco Morazán (25): Casa Blanca; El Zamorano; La Lima; Santa Lucía; Tatumbla; Villa San Francisco. Intibucá (1): Jesús de Otoro. Isla Roatán (1): Jicaro Galán. Lempira (1): Erandique. Ocotepeque (1): Reserva Güisayote. Valle (1): Nacaome. Yoro (1): Parque Nacional Pico Pijol. MEXICO (327). Chiapas (96): Aguacero; Albino Corzo; Berriozabal; Cahuare; Chiapa de Corzo; Cinco Cerros; Cintalapa; Comitán de Domínguez; El Chorreadero; El Jardín; Laguna Bélgica; Musté; Ocosingo; Ocozocoautla; Pijijiapan; Rizo de Oro; Suchiapa; Tapachula; Tuxtla Gutiérrez; Villaflores. Colima (5): Colima; Comala. Guerrero (86): Acapulco; Acahuizotla; Aserradero; Caletilla; Chilpancingo; Juan R. Escudero, El Terrero; Mochitlán; Quechultenango; Rodecia; San Luís Acatlán, Tlaxcalixtlahuaca; Tecpán; Zihuatajeno. Hidalgo (2): Mixquiahuala. Jalisco (54): Casimiro Castillo; Chamela; Guadalajara; La Cañada; Melaque; Nevado del Colima; Puerto Vallarta; Tomatlán. México (4): Soyaniquilpan de Juárez; Valle de Bravo. Michoacán (12): Apatzingan; Chinicuila; Coahuayana; San Telmo, Ojo de Agua. Morelos (16): Acamilpa; Jantetelco; Jojutla de Juárez; Tlaltizapan. Nayarit (4): San Blas (9 mi. E); Sayulita; Volcán Ceboruco. Oaxaca (41): Almoloya; Juchitán de Zaragoza; Matías Romero (23 mi. S); Oaxaca; Puerto Ángel; Puerto Escondido; Salina Cruz; Tapanatepec (8 mi. W); Tehuantepec. Puebla (1): Tehuixtla. San Luís Potosí (1): El Salto. Tamaulipas (2): Ciudad Victoria. Veracruz (3): Córdoba; Santiago Tuxtla. NICARAGUA (618). Carazo (4): San Francisco; San Marcos. Chinandega (8): Chichigalpa; Chinandega; San Benito; El Viejo. Granada (46): Nandaime; Volcán Mombacho. León (111): La Paz; Laguna de Asososca; Las Marías; Lechecuago; León; Mina El Limón; Posoltega; Santa Clara; Valle de las Zapatas. Managua (34): La Calera; La Chinampa; Laguna de Xiloa; Managua; Matano; Quinta Salvatierra; Tipitapa. Masaya (412): Las Flores; Masatepe. Matagalpa (3): Las Pilas; Matagalpa; San Isidro. PANAMA (3). Colón (2): Madden Dam; Pipeline Road (2 km W Gamboa); Chiriquí (1): Finca La Suiza (5.3 km N Los Planes). Natural history. One examined specimen was collected in detritus of Atta mexicana (Smith) (Hymenoptera: Formicidae). The larval stages were described by Ramírez-Salinas et al. (2010) (as L. nasutus), who found larvae in agricultural soils where cattle grazed. It is a common species in tropical deciduous forests between sea level and 1800 m, but mainly below 500 m (Ratcliffe & Morón 1997; Ratcliffe et al. 2013).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA7AFA619C1FAA1BA1B.taxon	description	(Figs. 1 G, 13 F, 15 J, 18 B, 25 G; 35)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA7AFA619C1FAA1BA1B.taxon	materials_examined	Male neotype designated by López-García & Deloya (2019: 128) (USNM) “ La Rioja / Tinogasta / v. 965 Kohler // ARGENTINA / 1968 Colln. / J. Daguerre // Ligyrus bidentulus / Fairmaire, 1892 / NEOTYPE ”. Type locality: La Rioja, Argentina.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA7AFA619C1FAA1BA1B.taxon	description	Description. Habitus as in Fig. 25 G. Length 12.1 – 19.3 mm; humeral width 5.8 – 10.7 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and with only sparse, small punctures. Frontoclypeal region with a carina, narrowly interrupted at middle (Fig. 1 G). Ocular canthus acute; with 8 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex (Fig. 1 G). Clypeal teeth triangular, slightly separated (Fig. 1 G). Mandible with 2 apical and 1 lateral, rounded tooth, tooth 1 longer and narrower than tooth 2. Maxilla subrectangular; galea with 3 dorsal and 2 ventral teeth. Apex of labrum straight to rounded. Interocular distance equal to 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex with small tubercle and shallow fovea. Elytra: First interval punctate, punctures similar to the 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 10 transverse, parallel, short carinae. Pygidium: 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. Protarsus of male simple, inner claw simple. Metatibia triangular. Apex of metatibia entire, with 25 – 29 spinules. 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 (Fig. 13 F). Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with apical 4 th narrowed, without minute spines; lateral ventral teeth short, triangular (Figs. 15 J, 18 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA7AFA619C1FAA1BA1B.taxon	diagnosis	Diagnosis. Ligyrus bidentulus can be recognized by the very long antennal club (2 times longer than antennomeres 2 – 7); clypeus long, strongly contracted towards apex (Fig. 1 G); clypeal teeth triangular, long, and not separated; frontal carinae narrowly interrupted at middle (Fig. 1 G); pronotum with small tubercle and narrow fovea (Fig. 25 G); parameres with apical 4 th narrowed, without minute spines; lateral teeth of parameres short, triangular (Figs. 15 J, 18 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA7AFA619C1FAA1BA1B.taxon	distribution	Distribution. Argentina (Endrödi 1985) and Uruguay (Neita-Moreno & Ratcliffe 2017). Locality records (Fig. 35). 58 examined specimens from MERKB, MNHM, MNHN, USNM, ZMUB. Some records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (101). Buenos Aires (9): Bahía Blanca; Buenos Aires; Carmen de Patagones; Necochea; Olavarria; San Blas; Villa Gesell. Catamarca (3): Ancafri; Sinquil. Chaco (1): Resistencia. Chubut (6): Dolavon; Estación de Esquel; Golfo de San Jorge; Península Valdés. Entre Ríos (3): Gualeguaychu. Jujuy (1): Posta de Lozano. La Pampa (1): Chacharramendi. La Rioja (9): Jaguel; La Unión; Mascasin; Patquía; Tinogasta; Villa Unión. Mendoza (19): Bardas Blancas; Cerro Guadal; Mendoza; El Nihuil (60 km S); Las Heras; La Valle. Misiones (1): Posadas. Neuquén (14): Añelo; Bajada del Marucho; Bajada del Agrio; Collón-Curá; Covunco; Huncal; La Negra; La Pintada; Picún Leufú; Piedra del Águila; Pilmatué; Planicie Banderita; Río Salado; Zalapa-Catuto. Río Negro (15): Allen; Catriel; Cipolletti; Conesa; El Bolsón; Fray Luís Beltrán; General Gómez; General Roca; Paso Flores; Piedra del Águila; Río Negro; San Antonio; Valcheta; Villa Regina. Salta (14): Cafayate (desierto); Coronel Olleros; Las Lajitas; Salta. San Luís (1): Paso Patria. Tucumán (1): Tafi de Valle. No data (3). URUGUAY (1). Canelones (1): Banda Oriental. Natural history. Adults are attracted to lights at night. Most records are from arid and desert areas.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA6AFA61F97FB3DBBF4.taxon	description	(Figs. 4 D, 5 D, 6 A, 7 E, 8 B, E, 15 K, 18 C, 25 H; 36)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA6AFA61F97FB3DBBF4.taxon	materials_examined	Male lectotype designated by Endrödi (1969: 65) (MNHN) “ Buenos Aires // Burmeisteri / 5 ♀ Steinh // Ex-Musaeo / E. Steinheil // MUSEUM PARIS / 1952 / COLL. OBERTHUR // Lectotypus / Ligyrus / burmeisteri / Steinh. / Endrody // LECTOTYPE / Tomarus / burmeisteri (Steinheil / 1872) // MNHN / EC 4415 ” Type locality: Buenos Aires, Argentina. Ligyrus distinctus Fairmaire, 1892: 244. Synonym. Female lectotype designated by Endrödi (1969: 65) (MNHN) “ Ligyrus / distinctus / Jan / Rioja // MUSEUM PARIS / Collection León Fairmaire / 1906 // TYPE // Lectotype / Ligyrus / distinctus / Fairm / Endrody // MNHN / EC 6757 ”. Type locality: La Rioja, Argentina.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA6AFA61F97FB3DBBF4.taxon	description	Description. Habitus as in Fig. 25 H. Length 9.9 – 13.5 mm; humeral width 5.7 – 7.5 mm. Color reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and only with sparse, small punctures. Frontoclypeal region with a carina, narrowly interrupted at middle. Ocular canthus acute; with 8 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3 times wider than apex. Clypeal teeth triangular, separated by 1 diameter tooth. Mandible with 2 apical and 1 lateral widely rounded tooth, tooth 1 longer and narrower than tooth 2 (Fig. 5 D). Maxilla subrectangular; galea with 5 small teeth (Fig. 4 D). Apex of labrum bilobed (Fig. 6 A). Interocular distance equal to 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex without tubercle or fovea. Elytra: First interval punctate, punctures similar to the other intervals. Inner surface of apex with small transverse tubercles (Fig. 8 E) forming about 69 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 10 transverse, parallel, short carinae (Fig. 8 B). 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. Protarsus of male simple; inner claw simple. Metatibia slightly contracted at apex. Apex of metatibia entire, with 19 – 25 spinules. 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. Parameres with apical 4 th narrowed, without minute spines; lateral ventral teeth long (Figs. 15 K, 18 C). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thin and fold.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA6AFA61F97FB3DBBF4.taxon	diagnosis	Diagnosis. Ligyrus burmeisteri can be recognized by the antennal club very long (2 times longer than antennomeres 2 – 7); clypeus long, strongly contracted towards apex; clypeal teeth triangular, long, and not separated; frontal carina narrowly interrupted at middle; pronotum without tubercle or fovea (Fig. 25 H); parameres with apical 4 th narrowed, without minute spines; lateral teeth of parameres long (Figs. 15 K, 18 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD79FFA6AFA61F97FB3DBBF4.taxon	distribution	Distribution. Argentina, Uruguay, Paraguay, Brazil, Bolivia, Chile, and Peru (Endrödi 1985). Locality records (Fig. 36). 333 examined specimens from CEIOC, CERPE, FSCA, MNHN, MN / UFRJ, MZUSP, UNSM, USNM, ZMUB. Some records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (768). Buenos Aires (283): Algarrobo; Azul; Bahía Blanca; Buenos Aires; Felipe Sola; Goyena; General Madariaga; Guamini; La Plata; Las Flores Ciudad; Coronel Rosales; Mar del Plata; Mercedes; Olivos; Quequén-Saladón; San Blas; San Fernando; San Isidro; Seis de Septiembre; Sierra de la Ventana, Sauce Grande; Tandil; Trenque Lauquen; Villa Iris. Catamarca (94): Catamarca; Guayamba; La Viña. Chaco (8): Chaco de Santiago; La Represa de los Indios; Resistencia. Chubut (6): Las Chapas; Puerto Madryn. Córdoba (42): Alpa Corral; Alta Gracia; Córdoba; Cosquín; Cruz del Eje; Espinillo; La Carlota; Los Olmos; Mar Chiquita; Río Primero; Santa María; Villa Dolores; Villa Soto. Corrientes (3): Corrientes; San Roque; Santo Tomé. Entre Ríos (1): Conconrdia. Formosa (1): Clorinda. Jujuy (52): Cosquin; El Volcán; Jujuy; Ledesma; Palmar; Parque Nacional Calilegua; San Pedro de Jujuy; Santa Bárbara; Volcán Tilcara; Yuto. La Pampa (20): General Pico; Guatraché; Santa Rosa; Winfredo. La Rioja (15): Jaguey; La Rioja; Patquía; Pozo de Piedra (4 km E Chepes). Mendoza (69): Cerro Guadal; Chacras de Coria-Luján de Cuyo; Mendoza; Moliches; Guaymallen; Rodeo de la Cruz-Guaymallen; San Martín; Santa Rosa. Misiones (10): Concepción Santa María; Loreto; Posadas; San Pedro. Salta (57): Cafayate; Campo Quijano; El Carril; El Corralito; El Naranjo; General Güemes; Guachipas; La Rioja; Rosario de Lerma; Salta; San Lorenzo; San Martín. San Juan (3): Desamparados; San Juan; Sarmiento. San Luís (15): Arizona; San Luís. Santa Fé (12): Reconquista; Rosario; Sancti Spiritu; San Javier; Santo Tomé, Río Salado. Santiago del Estero (16): Río Salado; Robles; Santiago del Estero. Tucumán (61): Estación Gendarmería; San Pedro de Colalao; Tucumán. BOLIVIA (14). Cochabamba (10): Cochabamba. La Paz (1): Murillo. Santa Cruz (1): Ichilo, Calacoto. Imprecise data (2): Bolivia. BRAZIL (77). Distrito Federal (1): Brasilia. Mina Gerais (18): Lambary; São Joao da Lagoa; Vila Monteverde. Rio de Janeiro (24): Itatiaia; Miguel Pereira; Rio de Janeiro. Santa Catarina (6): Corupa; Mafra; Rio Vermelho; São Bento. São Paulo (28): Alto de Pirra; Barueri; Cantareira; Itu, Fazenda Pau d’ Alho; Lageado; Porto Flor; Santo Amaro; São Paulo. PARAGUAY (1). Caaguazú (1): Estancia Primera. URUGUAY (22). Maldonado (1): Punta Ballena. Montevideo (8): Colón; Montevideo; Río Uruguay. Natural history. In Bolivia, L. burmeisteri larvae are abundant in humus and peaty soils in the wetter regions, such as the highlands of Cochabamba, where they infest potatoes (Squire 1972). The adults are attracted to lights during the summer and are found in dry to wet habitats (Neita-Moreno & Ratcliffe 2017).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD77FFA8AFA619C1FBFDBE5C.taxon	description	(Figs. 9 D, 15 L, 18 D, 26 A; 37)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD77FFA8AFA619C1FBFDBE5C.taxon	materials_examined	Female lectotype (MNHN) “ [unreadable text] // MUS. LA PERTÉ / 671 // this specimen is probably Latreille’s type of Scar. foss-or // Ex-Musaeo / D. Sharp 1890 // Museum Paris / ex Coll. / R. Oberthur // HOLOTYPE // HOLOTYPE / Scarabaeus / fossor Latreille, 1833 // HOLOTYPE / Tomarus / fossor (Latreille, 1833) // MNHN / EC 7078 ” here designated. Type locality: America (it was collected during Humboldt and Bonpland’s voyage).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD77FFA8AFA619C1FBFDBE5C.taxon	description	Description. Habitus as in Fig. 26 A. Length 13.5 – 15.0 mm; humeral width 6.5 – 7.0 mm. Color reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and only with sparse, small punctures. Frontoclypeal region with a carina, narrowly interrupted at middle. Ocular canthus acute; with 8 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3 times wider than apex. Clypeal teeth triangular, separated by 1 tooth diameter. Mandible with 2 apical and 1 lateral widely rounded tooth; tooth 1 truncate at apex, longer and narrower than tooth 2. Maxilla subrectangular; galea with 4 well-developed teeth, with 2 basal reduced teeth (Fig. 3 G). Apex of labrum bilobed. Interocular distance equal to 3.2 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex without tubercle or fovea. Elytra: First interval punctate, punctures similar to the other intervals. Inner surface of apex with small transverse tubercles forming about 70 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 10 transverse parallel short carinae. Pygidial surface with large punctures, evenly distributed (Fig. 9 D). Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate; basal tooth slightly distant from others. Protarsus of male simple; inner claw simple. Metatibia slightly contracted at apex. Apex of metatibia entire, with 18 – 20 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subrectangular (wider than long); coxite strongly concave. Subcoxite wider and longer than coxite. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with apical 4 th narrowed, without minute spines; lateral ventral teeth long and curved (Figs. 15 L, 18 D). Internal sac with lamellar spiny belt very long (about 12 times longer than copulatory lamellae); accessory lamella thickened.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD77FFA8AFA619C1FBFDBE5C.taxon	diagnosis	Diagnosis. Ligyrus fossor can be recognized by the antennal club very long (2 times longer than antennomeres 2 – 7); maxilla with 6 teeth (Fig. 3 G); clypeus long, strongly contracted towards apex; clypeal teeth triangular, long, and not separated; frontal carina narrowly interrupted at middle; pronotum without tubercle or fovea (Fig. 26 A); parameres with long, dorsal teeth, without ventral teeth (Figs. 15 L, 18 D); female coxite strongly concave.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD77FFA8AFA619C1FBFDBE5C.taxon	distribution	Distribution. El Salvador, Panama, Colombia, Venezuela, Curaçao, Antigua, Puerto Rico, Jamaica, Cuba, Bahamas (Endrödi 1969; Ratcliffe & Cave 2015). Locality records (Fig. 37). 164 examined specimens from FSCA, ICN, MEFLG, MZUSP, UNAB, UNSM, USNM. Some data from Ratcliffe (2003), Escalona & Joly (2006), and Ratcliffe & Cave (2015). ANTIGUA (1). St. Paul (1): English Harbor. BAHAMAS (1). Andros (1): N Andros Island. COLOMBIA (36). Antioquia (1): Turbaco. Atlántico (7): Barranquilla; Juan de Acosta; Repelón; Bolívar (3): Carmen; Cartagena; Isla Barú. Córdoba (3): Cereté. Cesar (6): Agustín Codazzi. Guajira (6): Campamento Mushaisa. Magdalena (5): Parque Nacional Tayrona. Ambiguous data (2): Manizales; Medellín. No data (2). CUBA (21). Cienfuegos (1): Cayamas. Ciego de Avila (1): Chambas. Guantanamo (1): Tortuguilla. Holguin (1): Guardalavaca. La Habana (9): La Habana; Santiago de las Vegas. Matanzas (2): Colón; Matanzas. Pinar del Río (4): Consolación del Sur; Pinar del Río; San Diego de los Baños; San Juan y Martínez. Santiago (2): Dos Caminos; Segundo Frente. CURAÇAO (3). Willemstad (3): Hato; Piscadera. EL SALVADOR (2). Ahuachapan (1): El Imposible. La Unión (1): Isla Martín Pérez. JAMAICA (95). Clarendon (2): Alligator Hole; Milk River Bath. St. Andrew (70): Bull Bay; Cockburn Peninsula; Cross Roads; Gordon Town; Halfway Tree; Irish Town; Jack’s Hill; Kingston; Liguanea; Seaview Gardens. St. Ann (3): Baron Hill Forest Reserve; Ocho Ríos. St. Catherine (2): Worthy Park Estate. St. Elizabeth (3): Maggotty. St. James (11): Greenwood; Montego Bay. St. Mary (1): Oracabessa. St. Thomas (1): Bath. Trelawny (4): Ducans; Martha Brae. PANAMA (28). Coclé (3): Interamerican Highway at Rio Grande. Colón (2): Madden Dam; Parque Nacional Soberanía. Los Santos (2): Corozal; Paraíso. Panamá (12): Alhajuela; Chame; Panamá; Taboga Island. Veraguas (3): Soná. No data (5). PUERTO RICO (2). Ponce (1): Mercedita. Mayagüez (1): Mayagüez. VENEZUELA (135). Anzoátegui (1): Clarines; El Hatillo. Aragua (1): Cardoncito; Cata; Chaparral; Choroní; El Limón; La Cabrera; La Victoria; Maracay; Parque Nacional Henri Pittier; Rancho Grande; San Mateo; Tasajeras; Turiamo; Turmero. Barinas (1). Reserva Forestal Ticoporo. Bolívar (10). Ciudad Bolívar; El Pao; Guasipi; Moitaco. Carabobo (19). Bejuma; Cachinche; Las Quiguas; Las Trincheras; Los Colorados; Nirgua; San Joaquín; Tacarigua; Yeramalis. Cojedes (10). El Baúl; El Pao; Girardot; San Carlos. Distrito Capital (14). Caracas; El Valle; Las Adjuntas. Falcón (7). Boca de Aroa; Cerro Santa Ana; Chichiriviche; Sanare; San José de Cocodite; San Juan; Santa Cruz de Bucaral. Guarico (6). El Sombrero; La Smith; Llanos de Cruz; Valle de la Pascua. Lara (13). Barquisimeto; Carora. Crespo; El Cuji; La Pastora (4 km NW); Parapara; Quibor. Mérida (1). Pueblo Nuevo. Miranda (2). Los Teques; Tacarigua de Mamporal. Monagas (4). El Piñal; San Antonio de Maturin; Santa Bárbara; Uverito. Nueva Esparta (4). Cerro Cimarrón; Guatamare; Salamanca; Tacarigua. Portuguesa (3). Acarigua; Ospino. Sucre (2). Cariaco; Cumanacoa. Táchira (2). La Pediera; San Cristóbal. Trujillo (3). El Cenizo; Escuque; San Pablo de Mendoza. Vargas (2). Caracaya; Punta de Tarma. Yaracuy (3). La Hoya; Sabana de Parra; Yaritagua. Zulia (6). Maracaibo. Natural history. There were some records of adults in roots and inflorescences of sunflower (Helianthus annus L.), and roots of maize. Most records are from tropical dry forest between 2 – 269 m.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD76FFADAFA61BD9FD94BBF4.taxon	description	(Figs. 2 C, 15 M, 18 E, 26 B; 38)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD76FFADAFA61BD9FD94BBF4.taxon	materials_examined	Type lost. The specimen labeled “ Scarabaeus / gibbosus / DeGeer / Holotype // Cyclocephala / verticalis Burm / det. Dr. Endrödi 1966 // Nicht die Type / v. L. gibbosus / Deg.! // NHRS-JLKB / 000027136 ” is a female of Cyclocephala as indicated by Endrödi and is not part of the type series. Type locality: Pennsylvania. Scarabaeus juvencus Fabricius, 1775: 18. Synonym. Not examined. According to Endrödi (1969) a syntype was at ZMK, but only three specimens without type labels were found by the curator. Type locality: America. Bothynus morio LeConte, 1847: 87. Synonym. Female lectotype (MCZ) “ (pink disc = Middle States (Md., Del., N. Y., N. J., Pa., Conn.?, R. I.?) // type 3716 // Ligyrus morio Lec. // gibbosus 19 ” Type locality: “ Provinciis mediis ” here designated. Bothynus obsoletus LeConte, 1847: 87. Synonym. Male holotype (MCZ) “ (blue disc) // type / 3715 // var.? obsoletus / Lec. // gibbosus 12 ” Type locality: Long`s Peak, Colorado. Podalgus variolosus Burmeister, 1847: 121. Synonym. Male lectotype designated by Endrödi (1969: 62) (MLUH) “ variolosus / Dej. / Am. Bor. Dp. // Lectotypus / Ligyrus / (Podalgus / variolosus Burm / Endrody’ ’ Male paralectotype (MLUH) “ Paratypus / Podalgus / variolosus / Burm ” Type locality: North America. Ligyrus californicus Casey, 1909: 283. Synonym. Female lectotype (USNM) “ Cal. / CASEY / bequest / 1925 // TYPE USNM / 35698 / californicus / Csy ” here designated. One male and three female paralectotypes “ Cal. / CASEY / bequest / 1925 // PARATYPE USNM / 35698 / californicus / Csy ”. Male and female paralectotypes (USNM) “ S. Bernandino / CASEY / bequest / 1925 // PARATYPE USNM / 35698 / californicus / Csy ”. Male syntype (USNM) “ S. Diego / CASEY / bequest / 1925 // PARATYPE USNM / 35698 / californicus / Csy ”. Ligyrus spissipes Casey, 1909: 283. Synonym. Male lectotype (USNM) “ Tex // CASEY / bequest / 1925 // NEOTYPE USNM / 35697 // spissipes / Csy ” here designated. One male and two female paralectotypes (USNM) “ Tex // CASEY / bequest / 1925 // PARATYPE USNM / 35697 // spissipes / Csy ”. Ligyrus longulus Casey, 1915: 193. Synonym. Male syntype (USNM) “ L. I. / CASEY / bequest / 1925 // TYPE USNM / 35677 // Ligyrus / longulus / Csy ”. Ligyrus virginicus Casey, 1915: 193. Synonym. Male syntype (USNM) “ Va. / CASEY / bequest / 1925 // TYPE USNM / 35678 // virginicus / Csy ”. Ligyrus parallelus Casey, 1915: 194. Synonym. Male syntype (USNM) “ Waco, Tex / Mou. // CASEY / bequest / 1925 // TYPE USNM / 35679 // parallelus / Csy ”. Male syntype (USNM) “ Waco, Tex / Mou. // CASEY / bequest / 1925 // PARATYPE USNM / 35679 // parallelus / Csy ”. Ligyrus remotus Casey, 1915: 194. Synonym. Female syntype (USNM) “ N. J. // CASEY / bequest / 1925 // TYPE USNM / 35681 // remotus / Csy ”. Male syntype (USNM) “ N. J. // CASEY / bequest / 1925 // PARATYPE USNM / 35681 // remotus / Csy ”. Male syntype (USNM) “ Atlantic City / N. J. // CASEY / bequest / 1925 // PARATYPE USNM / 35681 // remotus / Csy ”. Ligyrus puncticauda Casey, 1915: 195. Synonym. Male syntype (USNM) “ Miss // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35686 // puncticauda / Csy ”. Ligyrus texanus Casey, 1915: 195. Synonym. Male syntype (USNM) “ Lee Co. / Tex // CASEY / bequest / 1925 // TYPE USNM / 35687 // texanus / Csy ”. Eight male and three female syntypes (USNM) “ Bollas Tex // CASEY / bequest / 1925 // PARATYPE USNM / 35687 // texanus / Csy ”. Ligyrus breviusculus Casey, 1915: 196. Synonym. Male syntype (USNM) “ N. J. / ♂ // CASEY / bequest / 1925 // TYPE USNM / 35682 // breviusculus / Csy. One male and two female syntypes (USNM) “ N. J. // CASEY / bequest / 1925 // PARATYPE USNM / 35682 // breviusculus / Csy ”. Ligyrus lacustris Casey, 1915: 196. Synonym. Male syntype (USNM) “ N. I 11 // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35683 // lacustris / Csy ”. Ligyrus laetulus Casey, 1915: 197. Synonym. Male syntype (USNM) “ Tex // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35688 // laetulus / Csy ”. Two male and one female syntypes (USNM) “ Tex // ♂ // CASEY / bequest / 1925 // PARATYPE USNM / 35688 // laetulus / Csy ”. Ligyrus laticauda Casey, 1915: 197. Synonym. Male syntype (USNM) “ N. E. / U. S. // CASEY / bequest / 1925 // TYPE USNM / 35685 // laticauda / Csy ”. Ligyrus bicorniculatus Casey, 1915: 198. Synonym. Female syntype (USNM) “ Ks. // CASE / beques / 1925 // TYPE USNM / 35690 // bicorniculatus / Csy ”. Ligyrus rubidus Casey, 1915: 198. Synonym. Male holotype (USNM) “ Colo / IIII // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35692 // rubidus / Csy. ”. Ligyrus lucublandus Casey, 1915: 199. Synonym. Male syntype (USNM) “ Denver / vi. 24.11 Col. / C. A. Frost // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35691 // lucublandus / Csy ”. Ligyrus curtipennis Casey, 1915: 199. Synonym. Male syntype (USNM) “ N. Mex // CASEY / bequest / 1925 // TYPE USNM / 35680 // curtipennis / Csy. ”. Ligyrus effetus Casey, 1915: 200. Synonym. Male syntype (USNM) “ N. M. // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35694 // effetus / Csy. ”. Ligyrus farctus Casey, 1915: 200. Synonym. Male syntype (USNM) “ mex // CASEY / bequest / 1925 // TYPE USNM / 35693 // farctus / Csy. ”. Ligyrus arizonensis Casey, 1915: 201. Synonym. Female syntype (USNM) “ Ari. // CASEY / bequest / 1925 // TYPE USNM / 35696 // arizonensis / Csy. ”. Ligyrus brevipes Casey, 1915: 202. Synonym. Male syntype (USNM) “ Tex // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35684 // brevipes / Csy. ”. Ligyrus laevicauda Casey, 1915: 202. Synonym. Male syntype (USNM) “ N. M. // CASEY / bequest / 1925 // TYPE USNM / 35689 // laevicauda / Csy. ”. Ligyrus laticollis Casey, 1915: 203. Synonym. Male syntype (USNM) “ TYPE USNM / 35695 // laticollis / Csy ”. Ligyrus scitulus Casey, 1915: 203. Synonym. Male syntype (USNM) “ Cal // ♂ // CASEY / bequest / 1925 // TYPE USNM / 35699 // scitulus / Csy. ”.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD76FFADAFA61BD9FD94BBF4.taxon	description	Description. Habitus as in Fig. 26 B. Length 10.9 – 16.8 mm; humeral width 5.7 – 8.7 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex, with only sparse, small punctures. Frontoclypeal region with a complete, straight carina. Ocular canthus acute, with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex. Clypeal teeth triangular, separated by 1 tooth diameter. Mandible with 2 apical and 1 lateral, widely rounded tooth, tooth 1 longer and narrower than tooth 2. Maxilla subrectangular; galea with 3 dorsal teeth. Apex of labrum straight. Interocular distance equal to 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex with prominent tubercle and rounded, deep fovea. Elytra: First interval punctate, punctures similar in other intervals. Inner surface of apex with small, transverse tubercles forming about 66 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 10 transverse, parallel, short carinae. Pygidial surface with small to large punctures, denser on base and anterior corners. Surface in lateral view strongly convex in male, flat basally and slightly convex apically in female. Legs: Protibia tridentate; basal tooth slightly distant from others. Protibial surface sparsely punctate. Protarsus of male simple; inner claw simple. Metatibia triangular to slightly contracted at apex. Apex of metatibia entire, with 20 – 35 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subrectangular; surface strongly concave. Subcoxite wider and longer than coxite. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with apical 4 th narrowed, without minute spines; lateral ventral teeth short, triangular (Figs. 15 M, 18 E). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thickened, elongate.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD76FFADAFA61BD9FD94BBF4.taxon	diagnosis	Diagnosis. Ligyrus gibbosus can be recognized by the frontal carina straight, not interrupted (as in L. ruginasus, Fig. 1 E); pronotum with tubercle and a deep, rounded fovea (Fig. 26 B); protibial surface sparsely punctate; parameres with apical 4 th narrowed, without minute spines; lateral teeth of parameres triangular and short (Figs. 15 M, 18 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD76FFADAFA61BD9FD94BBF4.taxon	distribution	Distribution. Canada, United States, and Mexico (Ratcliffe & Cave 2017). Locality records (Fig. 37). 2,592 examined specimens from CERPE, CNIN, FSCA, IEXA, MCZ, MN / UFRJ, MZUSP, UNSM, USNM. Localities from Canada and Mexico, and United States of America counties are listed from Ratcliffe et al. (2013) and Ratcliffe & Cave (2017). CANADA (49). Ontario (48): Aldershot, Burlington; Chatham Laboratory; Delhi; Dunnville; Erieau; Grand Bend; Harrow; Ottawa; Point Pelee; Port Rowan; Ruthven; Toronto; Waterloo. Quebec (1): Sherbrooke. MEXICO (442). Aguascalientes (13): Aguascalientes; Pabellón de Arteaga. Baja California (270): Algodones; Bahía de los Ángeles; Bahía San Luís Gonzaga; Camalú; Colonet; Colonia Guerrero; El Rosario; El Socorro; Ensenada; La Zapotita; Laguna Salada; Meling Rach; Mexicali; Misión Santo Domingo; Santo Tomás; Rancho Agua San Matías; San Felipe; San José del Castillo; San Telmo; Tijuana; Valle de Trinidad. Chihuahua (34): Ahumada; Ciudad Juárez; Matamoros; Ojo de Lucero; Samalayuca; Santa Clara. Coahuila (3): Cuatro Ciénagas; Coyote; Santa Fé; Torreón. Distrito Federal (1): Villa Obregón. Durango (52): Durango; El Salto. Guanajuato (1): Dolores. Morelos (1): Cuernavaca. Nuevo León (1): Linares. San Luís Potosí (11): Matehuala; San Luís Potosí; Santa María del Río. Sinaloa (24): Culiacán; Mazatlán. Sonora (21): Agua Zarca; Bahía Kino; Colla Bay; El Seguro; Guaymas; Navajoa; Puerto Peñasco; Santa Ana; Sonoita. Tamaulipas (5): Tula; Playa Altamira. Zacatecas (2): Calera; Fresnillo. UNITED STATES OF AMERICA (16,496). Alabama (385): Autauga; Baldwin; Blount; Chilton; Covington; Dale; Dallas; DeKalb; Elmore; Etowah; Fayette; Greene; Hale; Henry; Houston; Jackson; Jefferson; Lauderdale; Lee; Limestone; Madison; Marion; Mobile; Monroe; Morgan; Randolph; Tallapoosa; Tuscaloosa; Winston. Arizona (857): Apache; Cochise; Coconino; Gila; Graham; La Paz; Maricopa; Mohave; Navajo; Pima; Santa Cruz; Yavapai; Yuma. Arkansas (95): Benton; Boone; Chicot; Crawford; Cross; Franklin; Independence; Johnson; Lee; Mississippi; Monroe; Sevier; St. Francis; Union; Washington. California (2,639): Butte; Contra Costa; Fresno; Humboldt; Imperial; Inyo; Kern; Kings; Los Angeles; Merced; Mono; Monterey; Orange; Plumas; Riverside; Sacramento; San Bernardino; San Diego; Santa Barbara; Shasta; Sonoma; Stanislaus; Trinity; Tuolumne; Ventura; Yolo; Yuba. Colorado (512): Alamosa; Arapahoe; Baca; Boulder; Denver; El Paso; Elbert; Fremont; Jefferson; Larimer; Mesa; Montezuma; Morgan; Otero; Prowers; Pueblo; Saguache; Washington; Weld; Yuma. Connecticut (8): Fairfield; New Haven. Delaware (106): Kent; New Castle; Sussex. District of Columbia (50): Washington. Florida (105): Alachua; Bay; Brevard; Broward; Collier; Dade; Dixie; Duval; Highlands; Liberty; Marion; Monroe; Nassau; Okaloosa; Orange; Palm Beach; Pinellas; Santa Rosa; Sarasota; Union; Volusia. Georgia (734): Baldwin; Bibb; Blakely; Butts; Candler; Catoosa; Chatham; Chattooga; Clarke; Cook; Crawford; Crisp; Dade; Decatur; DeKalb; Dougherty; Floyd; Fulton; Glascock; Hart; Henry; Houston; Jackson; Johnson; Liberty; Lowndes; Madison; Montgomery; Morgan; Muscogee; Newton; Oconee; Peach; Pike; Polk; Putnam; Richmond; Rockdale; Screven; Spalding; Sumter; Talbot; Taylor; Telfair; Thomas; Tift; Toombs; Upson; Walton; Wheeler; Worth. Idaho (484): Ada; Bannock; Benewah; Canyon; Caribou; Elmore; Fremont; Gooding; Idaho; Latah; Minidoka; Nez Perce; Owyhee; Payette; Power; Twin Falls; Valley; Washington. Illinois (470): Adams; Alexander; Calhoun; Champaign; Cook; Franklin; Hancock; Hardin; Iroquois; Jackson; Jefferson; Livingston; Macon; Marshall; Mason; McDonough; Peoria; Pike; Pope; Pulaski; Putnam; Randolph; Saline; St. Clair; Vermilion; Winnebago. Indiana (335): Allen; Bartholomew; Brown; Crawford; Dubois; Greene; Hendricks; Knox; La Porte; Marion; Morgan; Newton; Orange; Parke; Posey; Tippecanoe; Vigo. Iowa (135): Adair; Boone; Bremer; Cherokee; Crawford; Dallas; Decatur; Dickinson; Fremont; Guthrie; Harrison; Henry; Jefferson; Johnson; Lee; Linn; Louisa; Mahaska; Marion; Monona; Monroe; Muscatine; Osceola; Page; Plymouth; Polk; Poweshiek; Scott; Story; Taylor; Van Buren; Wapello; Winnebago; Woodbury. Kansas (432): Atchison; Barber; Chase; Cheyenne; Clay; Crawford; Douglas; Ellis; Finney; Ford; Greeley; Harvey; Jefferson; Jewell; Kearny; Kingman; Kiowa; Labette; Leavenworth; Logan; Morton; Pottawatomie; Reno; Russell; Saline; Scott; Seward; Shawnee; Sumner; Trego; Wichita; Woodson. Kentucky (299): Ballard; Caldwell; Clinton; Fayette; Green; Hardin; Henderson; Jefferson; Kenton; Knox; Lincoln; Nelson; Ohio; Scott; Shelby; Spencer; Warren. Louisiana (208): Avoyelles; Beauregard; Caddo; Clairborne; De Soto; East Feliciana; Evangeline; Franklin; Grant; Iberia; Jefferson; Livingston; Morehouse; Rapides; St; Bernard; St. John the Baptist; St. Landry; St. Mary; St. Tammany; Tangipahoa; Vermilion; Webster. Maine (7): Aroostook; Kennebec; Lincoln; Penobscot; York. Maryland (157): Anne Arundel; Baltimore; Calvert; Caroline; Carroll; Charles; Dorchester; Frederick; Montgomery; Prince George’s; Queen Anne’s; Somerset; St. Mary’s; Washington; Wicomico; Worcester. Massachusetts (15): Barnstable; Dukes; Essex; Hampden; Nantucket; Suffolk. Michigan (301): Allegan; Alpena; Arenac; Barry; Bay; Berrien; Branch; Calhoun; Clinton; Genesee; Green; Ingham; Jackson; Kalamazoo; Kent; Lake; Lenawee; Livingston; Macomb; Midland; Oakland; Oceana; Ottawa; St. Joseph; Van Buren; Washtenaw; Wayne; Wexford. Minnesota (137): Anoka; Clearwater; Hennepin; Houston; Isanti; Olmsted; Ramsey; Sherburne; Wabasha; Winona. Mississippi (138): Adams; Alcorn; Attala; Bolivar; Calhoun; Coahoma; Covington; DeSoto; Franklin; George; Greene; Grenada; Hancock; Harrison; Hinds; Issaquena; Jackson; Jefferson Davis; Jones; Kemper; Lafayette; Lamar; Lauderdale; Leflore; Lincoln; Lowndes; Madison; Marshall; Montgomery; Newton; Oktibbeha; Panola; Pearl River; Perry; Pontotoc; Quitman; Rankin; Sharkey; Simpson; Tallahatchie; Tishomingo; Tunica; Union; Warren; Washington; Wilkinson; Winston; Yalobusha. Missouri (737): Barry; Barton; Bollinger; Boone; Buchanan; Carter; Cass; Clay; Cole; Crawford; Holt; Jackson; Jasper; Jefferson; Lincoln; New Madrid; Pemiscot; Platte; Randolph; Rankin; Reynolds; Saline; Shannon; St. Louis; Stoddard; Vernon; Wayne. Montana (2): Gallatin. Nebraska (630): Antelope; Arthur; Box Butte; Brown; Buffalo; Cherry; Cheyenne; Colfax; Cuming; Custer; Dakota; Dawes; Dixon; Douglas; Dundy; Fillmore; Franklin; Frontier; Furnas; Gage; Grant; Greeley; Hall; Hamilton; Holt; Howard; Jefferson; Johnson; Kearney; Keith; Keya Paha; Kimball; Lancaster; Lincoln; Madison; McPherson; Otoe; Pierce; Red Willow; Richardson; Sarpy; Saunders; Scotts Bluff; Sheridan; Thomas; Wayne; Wheeler; York. Nevada (84): Benton; Clark; Churchill; Esmeralda; Humboldt; Lincoln; Nye; Washoe. New Hampshire (23): Strafford; Rockingham. New Jersey (98): Atlantic; Bergen; Burlington; Camden; Cape May; Cumberland; Mercer; Morris; Ocean; Salem. New Mexico (822): Bernalillo; Catron; Chaves; Curry; Doña Ana; Eddy; Grants; Guadalupe; Hidalgo; Lea; Lincoln; Luna; McKinley; Otero; San Juan; Sierra; Socorro; Taos; Torrance; Valencia. New York (78): Erie; Dutchess; Kings; Nassau; Queens; Dutchess. North Carolina (133): Burke; Carteret; Catawba; Cleveland; Columbus; Craven; Cumberland; Currituck; Dare; Durham; Edgecomb; Granville; Mecklenburg; Moore; Onslow; Person; Polk; Robeson; Stokes; Wake; Watauga; Wayne. North Dakota (44): Burleigh; Cass; Morton; Ransom; Richland. Ohio (82): Adams; Ashtabula; Champaign; Erie; Franklin; Guernsey; Hocking; Jefferson; Licking; Lucas; Ross; Scioto; Warren; Washington; Wayne; Wood. Oklahoma (686): Adair; Alfalfa; Beckham; Bryan; Caddo; Canadian; Cleveland; Comanche; Cotton; Custer; Deleware; Garfield; Greer; Hughes; Latimer; Lincoln; Marietta; Marshall; Mayes; McCurtain; Noble; Oklahoma; Osage; Rogers; Seminole; Texas; Tulsa; Wagoner; Washita; Wayne; Woods; Woodward. Oregon (281): Baker; Benton; Gilliam; Malheur; Morrow; Multnomah; Sherman; Umatilla; Union; Wasco; York. Pennsylvania (45): Allegheny; Berks; Chester; Delaware; Erie; Fulton; Huntingdon; Lancaster; Perry; Philadelphia. Rhode Island (21): Kent; Providence; Washington. South Carolina (202): Aiken; Anderson; Barnwell; Beaufort; Charleston; Chester; Clarendon; Edgefield; Florence; Hampton; Horry; Kershaw; Lexington; Marion; Newberry; Ocean; Oconee; Pickens; Richland; Sumter. South Dakota (722): Aurora; Beadle; Bennett; Bon Homme; Brookings; Brown; Brule; Clay; Codington; Davison; Fall River; Grant; Gregory; Haakon; Hughes; Hyde; Jackson; Jones; Lincoln; Marshall; Oglala; Pennington; Shannon, Stanley; Tripp; Union. Tennessee (70): Blount; Clay; Davidson; Dyer; Hardeman; Haywood; Henderson; Knox; Madison; Morgan; Rhea; Roane; Sevier; Shelby; Stewart; Warren. Texas (1,352): Anderson; Andrews; Atascosa; Bailey; Bastrop; Bexar; Bosque; Bowie; Brazos; Brewster; Brooks; Burnet; Cameron; Carson; Cass; Castro; Cherokee; Childress; Collingsworth; Comanche; Cooke; Crane; Crosby; Culberson; Dallam; Dallas; Deaf Smith; Denton; Dickens; Dimmit; Duval; Eastland; El Paso Erath; Frio; Galveston; Garza; Gillespie; Goliad; Gonzales; Gray; Hansford; Harris; Harrison; Hemphill; Hidalgo; Hopkins; Howard; Jasper; Jeff Davis; Jim Wells; Johnson; Karnes; Kaufman; Kenedy; Kleberg; Leon; Live Oak; Llano; Lubbock; Martin; Mason; Maverick; McMullen; Medina; Midland; Montgomery; Nacogdoches; Nueces; Ochiltree; Odessa; Panola; Parker; Parmer; Pecos; Potter; Presidio; Randall; Red River; Reeves; Rusk; San Patricio; Shelby; Smith; Tarrant; Trinity; Upton; Uvalde; Van Zandt; Victoria; Waco; Ward; Wheeler; Wichita; Wilbarger; Williamson; Winkler; Wood; Yoakum. Utah (533): Bannock; Clark; Emery; Garfield; Grand; Iron; Juab; Kane; Lincoln; Millard; Salt Lake; San Juan; Tooele; Washington; Wayne; Weber. Virginia (256): Accomack; Amelia; Arlington; Bedford; Campbell; Caroline; Charles City; Charlotte; Chesterfield; Dinwiddie; Essex; Fairfax; Fall Church; Fauquier; Fredericksburg; Goochland; Greensville; Halifax; Isle of Wight; James City; Lancaster; Lee; Mecklenburg; Nelson; New Kent; Norfolk; North Hampton; Northumberland; Orange; Pittsylvania; Prince Edward; Richmond; Suffolk; Virginia Beach; York. Washington (750): Adams; Asotin; Benton; Chelan; Columbia; Douglas; Ferry; Franklin; Garfield; Grant; Kittitas; Klickatat; Okanogan; Walla Walla; Whitman; Yakima. West Virginia (106): Berkeley; Clay; Grant; Hampshire; Jefferson; Mason; Mineral; Morgan; Randolph. Wisconsin (75): Columbia; Dane; Grant; Green; Green Lake; Jackson; Jefferson; La Crosse; Marquette; Milwaukee; Monroe; Richland; Sauk; Waushara; Wood. Wyoming (5): Platte. Natural history. Some examined specimens were collected in roots of Xanthium saccharatum Wallr. & Widder. (Asteraceae) in the state of New Mexico, and several in tunnels and mounds formed by pocket gophers (Geomys Rafinesque; Mammalia: Geomyidae) in Arkansas and Illinois. It is a species of economic concern known as carrot beetle that can affect different crops, such as the carrot, celery, potato (Ritcher 1958), and sunflower (Bottrell et al. 1973). They can be also found feeding on roots of Asteraceae such as Amaranthus (Davis 1916), Ambrosia L., Xanthium strumarium L., and Zinnia L. (Ratcliffe & Cave 2017). Its larval stages were described by Ritcher (1966). The life cycle is one year, the adults appear above ground in fall and reenter the soil to pass winter, waiting to put eggs until next spring (Davis 1916).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD72FFAFAFA619C1FF55BDB0.taxon	description	(Figs. 1 F, 15 N, 18 F, 26 C; 34)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD72FFAFAFA619C1FF55BDB0.taxon	materials_examined	Female lectotype designated by Endrödi (1969: 59) (MLUH) “ nasutus * / Yucat. / Berg // Lectotypus / Ligyrus / (Podalgus) / nasutus Burm / Endrody ” Type locality: Guatemala and Yucatan. Although Guatemala is mentioned in the original description no paralectotype was found. Ligyrus pygidialis Bates, 1888: 317. Original combination. Male lectotype designated by Endrödi (1969: 59) (BMNH) “ LECTO- / TYPE // Type // B. C. A., Col., II (2) / Ligyrus / pygidialis. // N. Yucatan. / Gaumer. // Ligyrus / pygidialis / Bates ♂ // Lectotypus / Ligyrus / pygidialis / Bat. / Endrody ”. Female paralectotype (BMNH) “ PARA- / LECTO- / TYPE // Type // B. C. A., Col., II (2) / Ligyrus / pygidialis. // N. Yucatan. / Gaumer. // Ligyrus / pygidialis / Bates ♀ ”. Male paralectotype (BMNH) “ PARA- / LECTO- / TYPE // B. C. A., Col., II (2) / Ligyrus / pygidialis. // N. Yucatan. / Gaumer. ”. There are 13 syntypes which should be labeled as paralectotypes at MNHN (EC 7079 - 7091). The following are considered invalid types as the species was not originally described with paratypes: Two male paratypes and two female paratypes (USNM) “ Yucatan / Temax / Gaumer // USNM / PARATYPE / 49710 // Ligyrus / pygidialis / Bates / Paratype ”.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD72FFAFAFA619C1FF55BDB0.taxon	description	Description. Habitus as in Fig. 26 C. Length 14.8 – 17.4 mm; humeral width 8.31 – 9.2 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex, with only sparse, small punctures. Frontoclypeal region with a bilobed, complete carina (Fig. 1 F). Ocular canthus acute, with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex (Fig. 1 F). Clypeal teeth triangular, separated by 1 diameter tooth (Fig. 1 F). Mandible with 2 apical and 1 lateral, widely rounded tooth, tooth 1 longer and narrower than tooth 2. Maxilla rectangular; galea with 3 dorsal teeth. Apex of labrum straight. Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, dense 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 70 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 11 transverse, parallel, short carinae. Pygidial surface with small punctures, denser on base and anterior corners. Surface in lateral view strongly convex (male) to flat (female). Legs: Protibia tridentate, basal tooth slightly distant from others. Protibial surface sparsely punctate. Protarsus of male simple, inner claw simple. Metatibia strongly contracted at apex. Apex of metatibia entire, with 15 – 20 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subrectangular (wider than long); surface strongly concave. Subcoxite wider and shorter than coxite. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with sides nearly parallel, lateral ventral teeth long (Figs. 15 N, 18 F). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thickened, irregular.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD72FFAFAFA619C1FF55BDB0.taxon	diagnosis	Diagnosis. Ligyrus nasutus can be recognized by the frontal carina bilobed, not interrupted (Fig. 1 F); pronotum without tubercle or fovea, with a basal small swelling (Fig. 26 C); pronotal punctation deep and large; metatibia elongate with apex strongly narrowed; parameres with sides nearly parallel; lateral ventral teeth long (Figs. 15 N, 18 F); internal sac with accessory lamella thick and irregular (as in L. allonasutus, Fig. 21 E); coxite of female strongly concave. Taxonomic remarks. Ligyrus nasutus was described with two females, one from Yucatán and the other from Guatemala. Ligyrus pygidialis was described by Bates (1888) based on a large series of males and females from Yucatán (Mexico), which according to his description were different to L. nasutus by the shape of the frontoclypeal carina divided at middle. It is not clear if Bates (1888) reviewed the type specimen of L. nasutus of Yucatán, but he cited additional specimens from Guerrero (Mexico) and San Jerónimo (Guatemala). Endrödi (1969) chose the female from Yucatán (in poor condition) as the lectotype for L. nasutus and considered L. pygidialis as an “ aberration ”. He only described and illustrated the parameres from the populations from western Mexico and Guatemala to Costa Rica but did not illustrate the parameres of the Yucatán populations, even though the lectotype of L. nasutus was from this region. After his work, the name L. nasutus was widely used for supposedly a unique species with wide distribution, but there was confusion since the original description of Burmeister included two female specimens from different species as types for L. nasutus. In the present work, the lectotypes of L. nasutus and L. pygidialis were analyzed in comparison with additional specimens from the region of Yucatán and compared with specimens from other regions of Mexico and Guatemala to Panama. Clear differences were found between the specimens from Yucatán region and the other populations, mainly in the shape of the frontoclypeal carina, pronotum, and parameres, which is supported by the biogeographic identity of the region of Yucatán. Following the International Commission on Zoological Nomenclature (1999), as the lectotype of L. nasutus is from Yucatán, only the populations of this region correspond to this species, and the remaining populations from western Mexico to Panama are named in the present work as L. allonasutus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD72FFAFAFA619C1FF55BDB0.taxon	distribution	Distribution. Ligyrus nasutus is restricted to the Yucatán region of Mexico, in the states of Campeche, Yucatán, Quintana Roo, and eastern Chiapas at elevations below 50 m. Locality records (Fig. 34). 62 examined specimens from BMNH, CNIN, EMAC, IEXA, UNSM, USNM. Some data from Ratcliffe et al. (2013). MEXICO (81). Campeche (16): Calkini; Campeche (5 mi. N); Grutas Xtacambilxunán. Chiapas (1): Palenque. Quintana Roo (12): Cancún; Carrillo Puerto (20 – 24 km N); Cobá; Nuevo X-Can; Playa del Carmen (10.9 km S). Yucatán (51): Chichén Itzá; Conkal; Mérida; Piste (12 km N); Santa Helena; Temax.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD71FFAFAFA61B75FB23BB97.taxon	description	(Figs. 13 G, 15 O, 18 G, 26 D; 37)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD71FFAFAFA61B75FB23BB97.taxon	materials_examined	Female lectotype (MCZ) “ [green disc] / type 3714 // L. juvencus / (Oliv.) Burm. / neglectus Lec. // gibossus 4 ” here designated. Type locality: Provinciis australibus	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD71FFAFAFA61B75FB23BB97.taxon	description	Description. Habitus as in Fig. 26 D. Length 13.2 – 16.0 mm; humeral width 7.3 – 8.9 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and only with sparse, small punctures. Frontoclypeal region with straight, complete carina. Ocular canthus acute; with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, base 3.5 times wider than apex. Clypeal teeth triangular, separated by 1 tooth diameter. Mandible with 2 apical and 1 lateral, widely rounded tooth, tooth 1 longer and narrower than tooth 2. Maxilla subrectangular; galea with 3 dorsal teeth. Apex of labrum straight. Interocular distance equal to 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, dense punctures. Apex with prominent tubercle and rounded, deep fovea. Elytra: First interval punctate, punctures similar to other intervals. Inner surface of apex with small, transverse tubercles forming about 66 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 10 transverse parallel short carinae. Pygidial surface with small to large punctures, denser on base and anterior corners. Surface in lateral view strongly convex (male) to flat (female). Legs: Protibia tridentate; basal tooth slightly distant from others. Protibial surface densely punctate. Protarsus of males simple, inner claw simple. Metatibia triangular to slightly contracted at apex. Apex of metatibia entire, with 19 – 25 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subquadrate (as long as wide); surface strongly concave. Subcoxite wider and subequal in length to coxite (Fig. 13 G). Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with apical 4 th narrowed, without minute spines; lateral ventral teeth short, triangular (Figs. 15 O, 18 G). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thickened, elongate.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD71FFAFAFA61B75FB23BB97.taxon	diagnosis	Diagnosis. Ligyrus neglectus can be recognized by the frontal carina straight, not interrupted (as in L. ruginasus, Fig. 1 E); pronotum with tubercle and deep, rounded fovea (Fig. 26 D); pronotal punctation deep and large; protibial surface densely punctate; parameres with sides nearly parallel, ventral teeth short and triangular (Figs. 15 O, 18 G).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD71FFAFAFA61B75FB23BB97.taxon	distribution	Distribution. The species occurs along the Atlantic coastal plain from North Carolina to Florida in the United States (Ratcliffe & Cave 2017). Locality records (Fig. 37). 588 examined specimens from FSCA, IEXA, UNSM, USNM. Counties listed from Ratcliffe & Cave (2017). UNITED STATES OF AMERICA (1,177). Alabama (18): Baldwin; Butler; Coffee; Geneva; Mobile; Monroe. Florida (840): Alachua; Baker; Bay; Brevard; Broward; Charlotte; Citrus; Clay; Collier; Columbia; Dade; Dixie; Duval; Escambia; Flager; Gadsden; Gilchrist; Highlands; Hillsborough; Indian River; Lake; Leon; Levy; Liberty; Manatee; Marion; Orange; Pinellas; Polk; Putnam; Santa Rosa; Sarasota; Seminole; Suwannee; Union. Georgia (162): Bibb; Bryan; Camden; Clarke; Dougherty; Grady; Hart; Johnson; Lowndes; Macon; McIntosh; Seminole; Thomas; Tift; Ware; Wheeler. Louisiana (9): East Feliciana; St. Mary; St. Tammany; Washington. Mississippi (19): Forrest; George; Greene; Harrison; Jackson; Lamar; Pearl River; Stone. North Carolina (58): Bladen; Brunswick; Craven; Dare; Moore; Richmond; Robeson. South Carolina (71): Aiken; Allendale; Barnwell; Beaufort; Charleston; Chesterfield; Colleton; Georgetown; Hampton; Jasper; Kershaw; Lexington; Richland; Sumter. Natural history. Some examined specimens were collected in pocket gopher mounds (Geomys; “ tuzas ” in Spanish) in Florida and Alabama. This was previously reported by Ratcliffe & Cave (2017).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFAEAFA619C1FA3FB8B8.taxon	description	(Figs. 9 E, 15 P, 18 H, 26 E; 36)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFAEAFA619C1FA3FB8B8.taxon	materials_examined	Male holotype (USNM) “ BRASIL / Halik 1966 / Collection // Halik / det. 1965 // Ligyrus / gibbosus / DeGeer // Ponta Grossa / Parana / 3. II 1965 / Moses / 26438 // Tomarus paranaensis / López-García & Deloya / HOLOTYPE ”	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFAEAFA619C1FA3FB8B8.taxon	description	Description. Habitus as in Fig. 26 E. Length 15.5 – 16.1 mm; humeral width 8.7 – 9.1 mm. Color dark reddish brown. Head: Frons coarsely punctate, area between eyes nearly impunctate. Frontoclypeal region with a carina, narrowly interrupted at middle. Clypeus slightly narrowed towards apex, base 2 times as wide as apex. Clypeal teeth triangular, separated by a tooth diameter. Mandible with 2 apical, acute teeth and a basal, widely rounded lobe. Maxilla subrectangular; galea with 3 dorsal and 2 ventral teeth. Apex of labrum bilobed. Interocular distance 5 times an eye width. Ocular canthus acute, with ventral setae. Antennal club long, 1.5 times longer than antennomeres 2 – 7. Pronotum: Surface with small, dense punctures. Apical tubercle and fovea absent. Elytra: First interval punctate, punctures similar to the other intervals. Pygidial surface without rugosity (Fig. 9 E). Punctures small, evenly distributed. Strongly convex in lateral view. Apex regularly rounded. Legs: Protibia tridentate without a denticle; basal tooth slightly distant from others. Protarsus subcylindrical, not enlarged, inner claw entire. Metatibia strongly narrowed before apex. Apex of metatibia entire, with 17 – 19 spinules. 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. Parameres with a medial tooth on each side; apex widened but ending in an acute, triangular point (Figs. 15 P, 18 H). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae), accessory lamella thin and folded.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFAEAFA619C1FA3FB8B8.taxon	diagnosis	Diagnosis. Ligyrus paranaensis and L. burmeisteri are similar in their external morphology, but the metatibia is strongly narrowed before apex and has 17 – 19 spinules on the apical margin in L. paranaensis, while it has the sides nearly parallel and the apical margin with 25 – 27 spinules in L. burmeisteri. The parameres have the apex acute in L. paranaensis (Figs. 15 P, 18 G) and rounded to subquadrate in L. burmeisteri (Fig. 15 K; see also fig. 18 in NeitaMoreno and Ratcliffe 2017).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFAEAFA619C1FA3FB8B8.taxon	distribution	Distribution. It is known only from the type locality, Ponta Grossa (Paraná, Brazil). Locality records (Fig. 36). 3 examined specimens from USNM and MN / UFRJ. BRAZIL (3). Paraná (3): Ponta Grossa. Natural history. Specimens known are from the Atlantic Forest biome and they were collected in February.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFB1AFA61C76FAA2B96C.taxon	description	(Figs. 15 Q, 18 I, 26 F; 39)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFB1AFA61C76FAA2B96C.taxon	description	Description. Habitus as in Fig. 26 F. Length 14.5 – 18.2 mm; humeral width 7.2 – 9.7 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 straight, complete carina. Ocular canthus acute, with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, ending in an acute tooth. Mandible with 2 apical and 1 lateral acute tooth, tooth 1 longer and narrower than tooth 2. Maxilla rectangular; galea with 3 dorsal teeth. Apex of labrum straight. Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, dense punctures. Apex with a prominent tubercle and a deep, rounded fovea. Elytra: First interval punctate, punctures similar to other intervals. Inner surface of apex with small, transverse tubercles forming about 46 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 8 – 9 transverse parallel short carinae. Pygidial surface with small punctures, denser on base and anterior corners. Surface in lateral view strongly convex (male) to flat (female). Legs: Protibia tridentate, basal tooth slightly distant from others. Protibial surface sparsely punctate. Protarsus of male simple, inner claw simple. Metatibia strongly contracted at apex. Apex of metatibia entire, with 19 – 20 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subquadrate (as long as wide); surface strongly concave. Subcoxite wider and subequal in length to coxite. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with sides nearly parallel; lateral ventral teeth short; apex slightly expanded (Figs. 15 Q, 18 I). Internal sac with lamellar spiny belt short (2.5 times longer than copulatory lamellae); accessory lamella thickened, elongate.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFB1AFA61C76FAA2B96C.taxon	diagnosis	Diagnosis. Ligyrus peninsularis is recognized by the triangular clypeus, with 1 central tooth (as in L. ruginasus, Fig. 1 E); frontal carina straight, not interrupted; pronotum with prominent tubercle and deep, rounded fovea; pronotal punctation deep and large (Fig. 26 F); metatibia elongate with apex strongly narrowed; parameres with sides nearly parallel, lateral ventral teeth short, apex slightly expanded (Figs. 15 Q, 18 I); internal sac with accessory lamella thick and elongate; coxite of female strongly concave. Taxonomic remarks. This species was combined in the genus Bothynus by Hardy (1964), who considered Oxygrylius as a synonym of Bothynus (name erroneously used at that time for the genus Ligyrus). According to Hardy (1964), the difference in the shape of the clypeus was not enough to differentiate the two known species of Oxygrylius with those of Ligyrus because he had found a specimen of L. gibbosus with triangular apex as in Oxygrylius, and so this shape could be caused by a mutation in a single character. The taxonomic change was not followed by Endrödi 1969 or later authors, but the results of the present phylogenetic analysis support the hypothesis raised by Hardy (1964).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD70FFB1AFA61C76FAA2B96C.taxon	distribution	Distribution. United States and Mexico, southern California and peninsula of Baja California (Hardy 1964; Ratcliffe & Cave 2017). Locality records (Fig. 39). Specimens examined from CERPE and IEXA. Mexican localities listed from Ratcliffe et al. (2013) and United States of America counties from Ratcliffe & Cave (2017). MEXICO (1,910). Baja California (150): Alfonsina; Bahía de San Rafael; Caño Tajo; Cataviña; Cerro del Palmarito; Ejido Bonfil; El Rosario; Isla Ángel de la Guarda; La Huerta; Las Ánimas; Laguna Chapala; Punta Prieta; Punta San Francisquito; Mexicali; San Borja; San Felipe; San Pedro; Valle de la Trinidad. Baja California Sur (1,760): Arroyo Comondú; Boca de la Sierra; Cabo San Lucas; Caduaño; Canipole; Ciudad Constitución; Colonia Fernando de la Toba; El Cien; El Coro; El Pescador; El Triunfo; Isla Cerralvo; Isla del Carmen; Isla Espírito Santo; Isla San José; Isla Santa Catalina; La Burrera; La Paz; La Purísima; Las Barrancas; Las Cruces; Las Cuevas; Las Tinajitas; Las Vírgenes; Loma Escondida; Loreto; Los Barriles; Los Naranjos; Miraflores; Misión de Nuestra Señora de La Paz Airapí; Palo Blanco; Playa El Coyote; Playa los Cerritos; Puerto Escondido; Rosarito; San Antonio; San Bartolo; San Hilario; San Ignacio; San Isidro; San José de Camondú; San José del Cabo; San Julio; San Lucas; San Pedro; Santa Águeda; Santa Catarina; Santa Rita; Santa Rosalía; Santa Victoria; Santiago; Sierra de La Laguna; Todos Los Santos; Valle Perdido; Villa Insurgentes. UNITED STATES OF AMERICA (303). California (279): Imperial; Inyo; Los Angeles; Riverside; San Bernardino; San Diego. Nevada (27): Clark; Lincoln. Natural history. The adults are active and frequently collected at lights from late June to late December, most abundantly in August and September, and overwintering is apparently passed as an adult (Hardy 1964).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6FFFB0AFA61C09FAF3BF3C.taxon	description	(Figs. 15 R, 18 J, 26 G; 35)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6FFFB0AFA61C09FAF3BF3C.taxon	materials_examined	Male lectotype designated by Endrödi (1969: 54) (NHRS) “ rubripes. Blm. Eug. / Bohem. // Lectotypus / (Ligyrus) / rubripes / Boh. / Endrody // 8840 / E 91 + // 336 / 66 // Kinb. // Monte- / video // NHRS-JLKB / 000023334 ” Ligyrus patagonus Steinheil, 1874: 560. New synonym. Original combination. Female holotype (MNHN) “ Patagones // patagonus / G. Steinh // ExMusaeo / E. Steinheil // Holotypus ♀ / Ligyrus / patagonus / Steinh. // MNHN / EC 7092 ” Type locality: Patagones, Argentina. According to Endrödi (1969), the holotype is a “ monstrous ” specimen of L. villosus but the shape of the pronotal tubercle and metafemur and its type locality are evidence that it is a female of L. rubripes. Ligyrus gianucai Dechambre & Lumaret, 1985: 107. Synonym. Male holotype (MNHN) “ Sand dunes / 28 Km S. Casino Beach / Brésil 25 II 1982 // N. M. gianuca Leg. // Ligyrus gianucai n. sp. inilú / HOLOTYPE / R. - P. Dechambre det. 1984 // HOLOTYPE // MNHN / EC 6731 ”.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6FFFB0AFA61C09FAF3BF3C.taxon	description	Description. Habitus as in Fig. 26 G. Length 12.1 – 16.5 mm; humeral width 5.8 – 9.2 mm. Color dark reddish brown. Head: Frons deeply rugopunctate, with large and sparse punctures, nearly smooth on vertex and only with sparse, small punctures. Frontoclypeal region with a carina, narrowly interrupted at middle. Ocular canthus acute, with 8 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus short, with lateral margins perpendicularly elevated, 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. Maxilla subrectangular; galea with 3 dorsal and 2 ventral teeth. Apex of labrum straight to rounded. Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, sparse punctures. Apex with prominent tubercle and shallow to deep 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. Protarsus of male simple, inner claw simple. Metatibia triangular. Apex of metatibia entire, with 28 – 30 spinules. 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. Parameres with apical 4 th narrowed, without minute spines; lateral, ventral teeth short, triangular, and closer to parameral base (Figs. 15 R, 18 J).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6FFFB0AFA61C09FAF3BF3C.taxon	diagnosis	Diagnosis. Ligyrus rubripes can be recognized by a short clypeus, with lateral margins perpendicularly elevated, strongly contracted towards apex; clypeal teeth triangular, long, and widely separated; frontal carina narrowly interrupted at middle; pronotum with prominent, acute tubercle and deep (Fig. 26 G), wide fovea (about 1.2 times interocular width); metatibia strongly triangular, without lateral constrictions; coxite surface flat; parameres with apical 4 th narrowed, without minute spines; lateral teeth of parameres short, triangular (Figs. 15 R, 18 J). Geographic distribution. Ligyrus rubripes is found in southern Brazil, Uruguay, and Argentina. Locality records (Fig. 35). 10 specimens examined from MN / UFRJ, MNHN, MZUSP, NHRS. ARGENTINA (2). Buenos Aires (2): Carmen de Patagones; Felipe Solá. BRAZIL (7). São Paulo (6): Guarujá; Itanhaemi; Peruíbe. Rio Grande do Sul (1): Praia do Cassino (28 km S). URUGUAY (1). Montevideo (1): Montevideo.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6EFFB3AFA61AF9FE30B855.taxon	description	(Figs. 1 E, 15 S, 18 K, 26 H; 39)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6EFFB3AFA61AF9FE30B855.taxon	materials_examined	Female holotype (MCZ) “ Type / 3717 // L. ruginasus / Rg. Barracks / Hald. / Lec. ” Type locality: Ringgold Barracks, Texas (near Rio Grande City, Starr County) Oxygrylius pimalis Casey, 1915: 209. Synonym. Syntypes (USNM). Not examined.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6EFFB3AFA61AF9FE30B855.taxon	description	Description. Habitus as in Fig. 26 H. Length 14.2 – 18.7 mm; humeral width 7.2 – 10.0 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 straight, complete carina (Fig. 1 E). Ocular canthus acute, with 10 ventral setae. Clypeal surface rugopunctate to transversely rugose, striae denser than those on frons. Clypeus triangular, ending in an acute tooth (Fig. 1 E). Mandible with 2 apical and 1 lateral, acute tooth, tooth 1 longer and narrower than tooth 2. Maxilla rectangular; galea with 3 dorsal teeth. Apex of labrum straight. Interocular distance 3.5 times an eye width. Antennal club very long. Pronotum: Surface with large, dense punctures. Apex with a prominent tubercle and a deep, rounded fovea. Elytra: First interval punctate, punctures similar to other intervals. Inner surface of apex with small, transverse tubercles forming about 66 parallel lines. Abdomen: Apex of tergite IV with a diagonal line of 7 transverse, parallel, short carinae. Pygidial surface with small punctures, denser on base and anterior corners. Surface in lateral view strongly convex (male) to flat (female). Legs: Protibia tridentate; basal tooth slightly distant from others. Protibial surface sparsely punctate. Protarsus of male simple, inner claw simple. Metatibia strongly contracted at apex. Apex of metatibia entire, with 25 spinules. Female genitalia: Subcoxite subrectangular (2 times wider than long). Coxite subquadrate (as long as wide); surface strongly concave. Subcoxite wider and subequal in length to coxite. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres without ventral teeth; apex widely expanded (Figs. 15 S, 18 K). Internal sac with lamellar spiny belt short (2.5 times longer than copulatory lamellae); accessory lamella thickened, elongate.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6EFFB3AFA61AF9FE30B855.taxon	diagnosis	Diagnosis. Ligyrus ruginasus can be recognized by the triangular clypeus, with 1 central tooth; frontal carina straight, not interrupted (Fig. 1 E); pronotum with prominent tubercle and deep, rounded fovea (Fig. 26 H); pronotal punctation deep and large; metatibia elongate with apex strongly narrowed; parameres without ventral teeth; apex widely expanded (Figs. 15 S, 18 K); internal sac with accessory lamella thick and elongate; coxite of female strongly concave. Taxonomic remarks. The species was originally described as Ligyrus ruginasus but then transferred in the genus Oxygrylius (Casey 1915). More details are provided in taxonomic remarks of L. peninsularis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6EFFB3AFA61AF9FE30B855.taxon	distribution	Distribution. This species inhabits the Sonoran Desert and drier areas of the southwestern United States (except most of California) and northern and western Mexico (Hardy 1964). Locality records. 21 specimens examined from CERPE, FSCA, IEXA, MN / UFRJ, MZUSP, UNSM. Mexican localities from Ratcliffe et al. (2013) and United States of America counties from Ratcliffe & Cave (2017). MEXICO (1,801). Chihuahua (81): Chihuahua; Ciudad Camargo; Ciudad Jiménez; Ciudad Juárez; Crell; Nueva Casas Grandes; Ojo del Lucero; San Pedro; Santa Eulalia. Coahuila (26): Cuatro Ciénagas; Dolores; Saltillo; San José de la Niña; San Pedro de las Colonias; Torreón. Durango (85): Ciudad Lerdo; Cuencamé; El Entroque; Reserva de la Biosfera Mapimí; San Ignacio. Estado de México (4): San José Villa de Allende. Jalisco (16): Ameca; Guadalajara; Hostotipaquillo; Zapopan. Nayarit (73): Jesús María; Los Sabinos; Pochotitlan; Tepic; Volcán Ceboruco. Nuevo León (89): Apodaca; Cañón de la Huasteca; El Álamo; Linares; Monterrey; Punta de la Loma; San Nicolás de los Garza; Parque Ecoturístico Cola de Caballo. San Luís Potosí (1): Ciudad Valles. Sinaloa (405): Choix; Concordia; Culiacán; El Dorado; El Zapotillo; Elota; Escuinapa; Guamúchil; Isla La Chiva; Isla Macapule; Isla Mazocahui; Isla Niscoco; Isla Pájaros II; Isla San Ignacio; Isla Tesobiare; Isla Vinorama; Los Mochis; Mazatlán; Mesa El Carrizal; Río Piaxta; Rosario; San Blas; Tierra Blanca; Vado Hondo; Venados; Villa Unión. Sonora (963): Agua Caliente; Agua Pietra; Álamos; Bahía Kino; Benjamin Hill; Caborca; Cananea; Carbó; Cerro Basura; Cerro Masiaca; Ciudad Obregón; Esperanza; Estación Don; Estación Llano; Estación Luís; Guaymas; Hermosillo; Imuris; Isla Tiburón; La Sandía; Magdalena; Moctezuma; Navajoa; Nogales; Pitiquito; Poza; Puerto Peñasco; Rosario del Tesopaco; Santa Ana; Saric; Sonoita; Tónichi; Valle de Yaqui; Yécora. Tamaulipas (53): Ciudad Mante; Ciudad Victoria; Güemez; Llera; Nuevo Laredo; Rancho San Felipe; Santander Jiménez. Zacatecas (10): Apazol; Jalpa. UNITED STATES OF AMERICA (3,807). Arizona (3,017): Apache; Cochise; Coconino; Gila; Graham; Greenlee; La Paz; Maricopa; Mohave; Pima; Pinal; Santa Cruz; Yavapi; Yuma. California (105): Imperial; Riverside. Colorado (8): La Plata. Kansas (2): Morton. New Mexico (108): Doña Ana; Eddy; Grant; Hidalgo; Luna; Otero; Sierra; Socorro. Texas (554): Bexar; Brewster; Cameron; Culberson; Dimmit; Duval; El Paso; Hidalgo; Hudspeth; Jeff Davis; Kimble; Kleberg; Maverick; Nueces; Presidio; Starr; Terrell; Uvalde; Val Verde; Webb; Zapata. Natural history. The adults are attracted to lights and found from May to December, but are most abundant in August (Hardy 1964).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6DFFB3AFA61DD0FBF2BA27.taxon	diagnosis	Diagnosis. For complete description see Neita-Moreno & Ratcliffe (2017). The species can be recognized by the subtriangular clypeus with 2 apical teeth; frontoclypeal carina incomplete at middle; pronotum without fovea or tubercle; apex of metatibia with about 29 spinules; coxite surface flat; and apical half of parameres with small spinules. Geographic distribution. Ligyrus spinipenis is known from Argentina (Neita-Moreno & Ratcliffe 2017). Locality records (Fig. 36). Records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (9). La Rioja (2): La Rioja. San Luís (1): San Antonio. Santiago del Estero (6): Parque Nacional Del Copo; Santiago del Estero. Natural history. Adults are attracted to lights (Neita-Moreno & Ratcliffe 2017).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6DFFB2AFA61FC1FD33B918.taxon	description	(Figs. 2 D, 4 E, 5 E, 6 B, 11 G, 12 F, L, 14 D, 15 T, 18 L, 26 I; 35)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6DFFB2AFA61FC1FD33B918.taxon	materials_examined	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6DFFB2AFA61FC1FD33B918.taxon	description	Description. Habitus as in Fig. 26 I. 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. 5 E). Maxilla subrectangular; galea with 4 teeth (Fig. 4 E). Apex of labrum rounded (Fig. 6 B). 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. 11 G). Protarsus of male simple, inner claw simple. Metatibia triangular. Apex of metatibia entire to slightly crenulate, with 17 – 20 spinules (Fig. 12 L). 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. 14 D). Parameres with apical 4 th narrowed, without minute spines; latero-ventral teeth short, triangular (Figs. 15 T, 18 L).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6DFFB2AFA61FC1FD33B918.taxon	diagnosis	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. 26 I); pronotal punctation absent or minute; metatibia strongly triangular (Fig. 12 L); coxite surface flat; parameres with apical 4 th narrowed, without minute spines; lateral teeth of parameres short, triangular (Figs. 15 T, 18 L). 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).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	type_taxon	Type species: Tomarus maimon Erichson, 1847: 96 by monotypy.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	materials_examined	Type species: Ligyrus laevicollis Bates, 1888: 316 here designated.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	description	Description. Length 17.7 – 30.0 mm. Humeral width 8.5 – 16.4 mm. Head: Clypeus subtriangular (Fig. 1 H – K). Apical clypeal teeth triangular, slightly to widely separated. Ocular canthus rounded; without ventral or dorsal setae. Frons with 2 transverse to conical tubercles. Mentum subtriangular to subrectangular (Fig. 2 I – K), with a strong constriction before apex; surface strongly concave at base. Mentum and ligula fused. Ligula well developed; covered with long, dense setae along lateral margins; ligular lobes fused or separated by a narrow emargination. Maxillary apical palpomere 1.3 times longer than the second palpomere. Maxilla with galea subquadrate (1.5 times longer than wide) (Fig. 4 F – I); with 6 well-developed teeth (rarely reduced); maxillary teeth 3 and 4 in parallel position. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth (Fig. 5 F – H) or lateral tooth not well developed (Fig. 1 J); incisor subequal (Fig. 5 F) or longer than tooth 2 (Figs. 1 J, 5 H). Labrum subrectangular (Fig. 6 D – G); apex straight; with long, dense setae. Antenna with 10 antennomeres, club short (1.3 times longer than antennomeres 2 – 7). Pronotum: Apical marginal bead complete. Apex with or without tubercle; without fovea or with deep, wide fovea. Scutellum: Surface with minute sparse punctures or with deep punctures forming a line parallel to lateral margins. Elytra: Inner surface of apex with rounded small to large tubercles forming 14 – 30 parallel lines (rarely not forming defined lines) (Fig. 8 K – N). Wings: RA with sparse, small to minute peg-like setae on medial ventral region (Fig. 7 F – H); without peg-like setae on dorsal surface. Edge of RA 3 with dense setae. Abdomen: Apex of tergite IV with a triangular wide area with tubercles forming diagonal parallel lines (Fig. 8 G – J). Sternite VIII emarginate in male; entire in female. Pygidium with rugosity (Fig. 9 F – L), slightly convex in male, nearly flat in female. Propygidium without stridulatory area. Venter: Prosternum wide, regularly rounded. Propleura with short, sparse setae on anterior region, medial region glabrous, posterior region with 5 – 10 sparse setae. Apex of prosternal process flat; without dorsal setae. Metepisternum with complete carina; inner surface with deep, large punctures, without setae; outer surface homogeneously rugose, without setae. Metasternum with large, deep, confluent punctures; with minute to short, sparse setae. Legs: Male protarsus simple, inner claw simple. Protibia tridentate, sometimes with an additional, basal denticle, teeth equidistant. Apical margin of meso- and metatibia entire to slightly crenulate. Metafemur without well-defined anterior row of punctures. Female genitalia: Subcoxite as long as wide. Coxite subrectangular (longer than wide); surface slightly concave. Subcoxite as wide as and shorter than coxite. Male genitalia: Phallobase 1.0 – 1.2 times longer than parameres. Parameres with 1 – 2 dorsal teeth, without ventral teeth. Internal sac with copulatory lamella and short to long lamellar spiny belt; with a complex of 3 – 8 (26 in T. pullus) spine-like accessory lamellae, with or without granules at base (Figs. 22 – 23).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	diagnosis	Diagnosis. Tomarus can be differentiated from other Pentodontini genera by the following: Subtriangular clypeus, clypeal teeth triangular to transverse; frontoclypeus with 2 tubercles; antennal club short; pronotum usually with tubercle and with or without fovea; protibia tridentate (sometimes with a basal denticle) (Fig. 11 H – J).; protarsus of male simple, inner claw simple; metafemur without well-defined anterior row of punctures; pygidium of male with transverse thin rugosity at base (Fig. 9 F – L); prosternal process flat, without dorsal setae; internal sac with a complex of spine-like, accessory lamellae (Figs. 22 – 23); stridulatory apparatus consisting of a triangular wide area with tubercles forming 13 – 20 oblique, parallel lines on apex of sternite IV (Fig. 8 G – J), and apical internal surface of elytra with rounded small to large tubercles forming 14 – 30 parallel lines (rarely not forming defined lines) (Fig. 8 K – N).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	distribution	Distribution. It is a Neotropical group probably originated in South America and dispersed along Central America, with few species reaching the tropical regions of Mexico, the Caribbean islands, and southern Florida in the United States of America.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD6CFFB4AFA61C9CFA22BB85.taxon	discussion	Phylogenetic relationships. The morphological phylogeny of Tomarus (sensu lato) evidenced that this group, as previously considered, was polyphyletic and contained four different genera. Taxonomic remarks. Erichson (1847) described the genus Tomarus to place a new species from Peru (T. maimon), characterized by having a maxilla with 6 teeth, elytra with stridulatory area, and mentum “ oblongus ”. A year later (Erichson 1848) he described T. gyas from Guiana. Lacordaire (1855) synonymized Tomarus with Ligyrus, and the synonymy was maintained for several years. Endrödi (1969) recognized Tomarus as a subgenus of Ligyrus. In more recent years, Ratcliffe (2003) revalidated the name Tomarus based on the priority principle, which was widely accepted. Although Escalona & Joly (2006) suggested maintaining the name Ligyrus due to the wide use and agricultural importance, they never submitted an application to the International Commission on Zoological Nomenclature. Morón & Grossi (2015) separated the genus into Ligyrus and Tomarus based on the number of teeth on the outer margin of mandible. Tomarus (sensu Morón & Grossi 2015) coincides only in part with the classification purposed in the present work. The species E. ebenus and E. similis are excluded, while T. adoceteus and T. pullus are included in Tomarus, and the species T. colombianus and T. rostratus (not considered in Morón & Grossi 2015) are also in Tomarus as well as other species described later: T. roigjunenti, T. laticaudus, T. maracaiboensis, and T. pilcopataensis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB7AFA619C1FEE3BA9F.taxon	description	(Figs. 1 H, 16 A, 19 A, 27 A; 41)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB7AFA619C1FEE3BA9F.taxon	materials_examined	Male holotype (BMNH) “ Roy Soc-CIG Expdn. / Little Cayman, BWI / Pirates’ Point / House light / 30.7.1975. R. R. Askew // BM 1981 / 342 // TOMARUS / ADOCETEUS / RATCLIFFE & CAVE / HOLOTYPE ” Female paratype (BMNH) “ GRAND CAYMAN IS. B. W. I. / Georgetown / 16 sept 1973 / E. J. Gerberg // At black light // Ratcliffe & Cave / db Dynastine / West Indies // TOMARUS / ADOCETEUS / RATCLIFFE & CAVE / PARATYPE. Female paratype (BMNH) “ WEST INDIES: Cayman Is. / Grand Cayman / VI- 1992 coll. P. Fitzgerald / blacklight trap // Ratcliffe & Cave / db Dynastine / West Indies // TOMARUS / ADOCETEUS / RATCLIFFE & CAVE / PARATYPE. Female paratype (BCRC) “ Roy Soc-CIG Expdn. / Little Cayman / Pirates’ Point / 9. viii. 1975 // TOMARUS / ADOCETEUS / RATCLIFFE & CAVE / PARATYPE ”. Type locality: Pirates’ Point, Little Cayman Island.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB7AFA619C1FEE3BA9F.taxon	description	Description. Habitus as in Fig. 27 A. Length 20.0 – 22.5 mm; humeral width 10.3 – 12.4 mm. Color dark reddish brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 transverse, low tubercles separated by about 7 tubercle diameters. Clypeus narrowed towards apex, base 3 times as wide as apex (Fig. 1 H). Clypeal teeth small, transverse, separated by a tooth diameter. Mandible with 2 apical teeth and a lateral, slightly rounded tooth. Mentum abruptly constricted at apical 3 rd. Galea of maxilla with teeth 5 and 6 vestigial. Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures; denser and deeper on anterior and lateral angles. 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: With deep, large punctures forming 2 lines parallel to margins. Elytra: Punctures on first interval as large as those on other intervals; sutural stria complete. Inner surface of apex with rounded, large tubercles forming 13 – 14 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with small, irregular tubercles forming 12 – 14 diagonal, nearly parallel lines. Pygidium with dense rugosity on basal 3 rd; apex rounded. Venter: Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate, without basal denticle. Apex of metatibia crenulate, with 10 – 11 spinules. Male genitalia: Phallobase 1.2 times longer than parameres. Parameres with 2 dorsal teeth on each side; basal tooth acute, longer than apical tooth that is inconspicuous (Figs. 16 A, 19 A). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB7AFA619C1FEE3BA9F.taxon	diagnosis	Diagnosis. Tomarus adoceteus is differentiated by the following character combination: frontoclypeal tubercles transverse and low (Fig. 1 H); galea of maxilla with teeth 5 and 6 vestigial (as in T. subtropicus, Fig. 4 I); pronotal surface with small punctures; pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1 / 3 as wide as interocular distance) (Fig. 27 A); protibia tridentate, without an additional basal denticle; apex of metatibia with 10 – 11 spinules; parameres with 2 dorsal teeth on each side, main teeth long, widely separate from secondary; apex of parameres slender (Figs. 16 A, 19 A). Taxonomic remarks. Tomarus adoceteus was described originally by Ratcliffe & Cave (2010) in the genus Tomarus, but Morón & Grossi (2015) placed it in Ligyrus, while their most similar species were remained in Tomarus. The phylogenetic analysis evidences that this species does not have a close relationship with the members of Ligyrus, so it is maintained in its original combination.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB7AFA619C1FEE3BA9F.taxon	distribution	Distribution. Bahamas and Cayman Islands (Ratcliffe & Cave 2015). A single male specimen from Yucatán (Mexico) was examined but more evidence is necessary to confirm that the species occurs in that country. Locality records (Fig. 41). 7 specimens examined from BCRC, BMNH, and IEXA. Some records from Ratcliffe & Cave (2015). BAHAMAS (6). Bimini (4): Allen Cay; North Bimini; South Bimini. CAYMAN ISLANDS (7). Grand Cayman (3): Georgetown. Little Cayman (4): Pirate’s Point. MEXICO (1). Yucatán (1): Reserva Estatal El Palmar. Natural history. Little is known about T. adoceteus. Specimens have been found between June and September using blacklights.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB6AFA61E07FE43BB43.taxon	description	(Figs. 8 G, K, 16 B, 19 B, 23 A, 27 B; 42)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB6AFA61E07FE43BB43.taxon	materials_examined	Female lectotype (BMNH) “ SYNTYPE // Ega / Braz. // Ligyrus ♀ / amazonicus Arr. / M. E. Bacchus det. 1970 ” here designated. Female paralectotype (BMNH) “ SYNTYPE // ♀ // Amazon / Bates // Fry Coll. / 19051100. // 19194 // Ligyrus / amazonicus Arr. / M. E. Bacchus det. 1970 ”. Female paralectotype (BMNH) “ SYNTYPE // Obydos. / Lower Amazon. / 2. II. 9 – / E. E. Austen. / 96 -- 80 // Ligyrus ♀ / amazonicus Arr. / M. E. Bacchus det. 1970 ”.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB6AFA61E07FE43BB43.taxon	description	Description. Habitus as in Fig. 27 B. Length 22.0 – 25.0 mm; humeral width 11.0 – 12.5 mm. Color dark brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 transverse tubercles separated by about 4.5 tubercle diameters. Clypeus narrowed towards apex, base 3 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures, denser and deeper on anterior and posterior angles. Apical tubercle small, rounded. Subapical fovea shallow, narrow (1 / 3 as wide as interocular distance), equal in both sexes, surface transversely rugose. Scutellum: 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 11 – 12 parallel lines (Fig. 8 K). Abdomen: Apex of tergite IV with a triangular wide area with large, rounded tubercles forming 18 diagonal parallel lines (Fig. 8 G). Pygidium with dense rugosity on basal 3 rd; apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate, with a basal small denticle. Apex of metatibia crenulate, with 23 – 28 spinules. Male genitalia: Phallobase as long as parameres. Parameres with 1 dorsal tooth each side; tooth long, acute; apices roundly expanded (Figs. 16 B, 19 B). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 8 spine-like accessory lamellae, without or with reduced granules at base (Fig. 23 A). Taxonomic remarks. This species was described by Arrow (1914) and then synonymized by Endrödi (1969) under T. gyas based on the approximate shape of the parameres and difficulty to delimitate the differences on the shape of pronotal fovea and protibia. Despite the convergence in the shape of parameres, the phylogenetic analysis evidenced that this species is not close to T. gyas, and they are different in several characters, such as the shape of the frontal tubercles, setation on the metasternum, the stridulatory apparatus, and the internal sac of the aedeagus. Therefore, T. amazonicus is treated here as a valid species and classified in the genus Tomarus.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB6AFA61E07FE43BB43.taxon	diagnosis	Diagnosis. Tomarus amazonicus is recognized by the following character combination: frontoclypeal tubercles transverse; pronotal tubercle small and rounded (Fig. 27 B); subapical fovea shallow, narrow (1 / 3 as wide as interocular distance), equal in both sexes; metasternum with minute setae on anterior angles; protibia tridentate and with an additional basal denticle; apex of metatibia with 23 – 28 spinules; parameres with a long, acute tooth on each side, apical half strongly contracted, apices roundly expanded (Figs. 16 B, 19 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD69FFB6AFA61E07FE43BB43.taxon	distribution	Distribution. Amazon region of Brazil, Bolivia, Peru, Ecuador, and Colombia. Locality records (Fig. 42). 462 specimens examined from BMNH, CEIOC, CERPE, FSCA, IAvH, ICN, MEKRB, MN / UFRJ, MZUSP, SENASA, UNAB, UNSM, and USNM. Some records from CMNC and Endrödi (1969). BOLIVIA (11). Beni (10): Riberalta; Rurrenabaque; Trinidad; Vaca Diez. Buena Vista (1): Tacaraundi. BRAZIL (319). Acre (3): Cruzeiro do Sul; Porto Walter; Vila Fejo. Amazonas (118): Benjamin Constant; Boca do Rio Madeira; Borba; Coari; Codota’s; Curucuri; Guajará; Ipixuna, Rio Gregorio; Itacoatiára; Lago Januaca; Manaus; Rio Japula; Rio Liberdade, Estirao da Preta; Rio Salimoes, entre Lago Jacaré e Purus; São Sebastian do Uatuma; Tabatinga; Tefé (Ega). Pará (125): Boca do Lago de Terra Santa; Ilha do Itanduba; Maicurú; Obidos; Oriximiná; Paraná de Alenquer; Prainha; Rio Paru; Santarém. Rondônia (73): Guapore; Porto Velho, Rio Madeira; Samuel. Tocantins (1): Buriti do Tocantis, Assentamento Ouro Verde, Chacára da Bisa. No data (6). COLOMBIA (41). Amazonas (22): Leticia; Leticia (20 km N); Purité, San Juan. Caquetá (19): Florencia; Río Orteguaza near Río Peneya. ECUADOR (18). Napo (15): Jatun Sacha Biological Station; Lago Agrio (18 km E); Misahualli Jungle Lodge. Imprecise data (3): Quito. PERU (72). Junín (1): Satipo, Huancayo. Loreto (55): Iquitos; Explorama Inn; Jungle Amazon Inn; Maynas; Yanamono. Huánuco (7): Marañón; Tingo María. Madre de Dios (6): Reserva Nacional Tambopata. Ucayali (1): Laguna Yarinacocha. No data (2). IMPRECISE DATA (2). Amazonas. Natural history. This species is restricted to the Amazon rainforest between 300 – 1050 m. Some specimens were collected using light traps.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD68FFB8AFA61E22FC8CBBD8.taxon	description	(Figs. 9 F, 11 H, 16 C, 19 C, 27 C; 43)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD68FFB8AFA61E22FC8CBBD8.taxon	materials_examined	Type series lost. Type locality: Santo Domingo, Dominican Republic. Ligyrus latifovea Bates, 1888: 315. Synonym. Female lectotype designated by Cartwright (1959: 518) (BMNH) “ LECTO- / TYPE // TYPE // Mexico. / Salle Coll. / Teapa // Sp. figured. // B. C. A. Col. II. (2) // Ligyrus / latifovea / Bates // Lectotype / B. latifovea Bates / Cartwright 1959 ”. Type locality: Teapa, Veracruz, Mexico. Ligyrus maximus Arrow, 1913: 466. Synonym. Male lectotype (BMNH) “ TYPE // SYN- / TYPE // ♂ // Brazil. / Stanford Exped. / 1913 - 56. // Porto Velho, Rio / Madeira Brazil // Mann & Baker. // Ligyrus maximus / type / Arrow ” here designated. Male paralectotype (BMNH) “ SYN- / TYPE // Brazil / Serpa / 99 – 195 ”. Female syntype / Madeira Brazil // Mann & Baker. ” Type locality: Porto Velho, Amazonas, Brazil. Ligyrus latus Arrow, 1914: 273. Synonym. Male lectotype (BMNH) “ Type / H. T. // SYN- / TYPE // 19695 // Venez. a // Fry Coll. / 1905.100. // ♂ // Ligyrus / latus / type Arrow ” here designated. Male paralectotype (BMNH) “ SYN- / TYPE // Fry Coll. 1905.100. // Columbia // ♂ // Ligyrus ♂ / latus Arrow / M. E. Bacchus det. 1970 ”. Type locality: Venezuela.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD68FFB8AFA61E22FC8CBBD8.taxon	description	Description. Habitus as in Fig. 27 C. Length 25.0 – 30.0 mm; humeral width 13.0 – 15.5 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures; denser and deeper on anterior and posterior angles. Apical tubercle prominent, acute; slightly recurved. Subapical fovea deep, transverse, wide (> 1 / 2 the interocular distance); 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 14 – 15 diagonal parallel lines. Pygidium with dense rugosity on basal 3 rd (Fig. 9 F). Apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate (Fig. 11 H), sometimes with a basal, small denticle (populations from Mexico and Central America). Apex of metatibia crenulate, with 15 – 17 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with 1 dorsal tooth each side; tooth small, acute (Figs. 16 C, 19 C). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD68FFB8AFA61E22FC8CBBD8.taxon	diagnosis	Diagnosis. Tomarus bituberculatus can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent (Fig. 27 C); pronotal fovea transversely oval (as wide as interocular distance); protibia tridentate, rarely with an additional basal denticle (Fig. 11 H); apex of metatibia with 15 – 17 spinules; parameres with a small, acute tooth on each side, apical half strongly contracted and slender, apices triangularly expanded (Figs. 16 C, 19 C). Taxonomic remarks. The species was described by Palisot de Beauvois (1811) from Santo Domingo (Dominican Republic), even though there is not any known record for the species there and the type series was lost at sea. Since many localities in Palisot de Beauvois’s descriptions have proven to be incorrect this record should be considered erroneous. Burmeister (1847: 100) redescribed Heteronychus scarabaeinus, originally described as Cyclocephala scarabaeina Perty, 1830 and placed it in Ligyrus. As it was already recognized by Bates (1888), Burmeister (1847) confounded two very different species under L. scarabaeinus, which were L. scarabaeinus and L. ebenus, so Bates (1888) used the new name L. latifovea instead of L. scarabaeinus (sensu Burmeister 1847) and listed Cyclocephala scarabaeina Perty, 1830 as a synonym for L. ebenus. Endrödi (1969) also noticed the Burmeister’s error, and attempted to designate a male lectotype (deposited at MLUH) for L. scarabaeinus (Burmeister, 1847). However, this lectotype designation is invalid since Burmeister (1847) did not describe Heteronychus scarabaeinus as a new species; therefore, “ L. scarabaeinus Burmeister 1847 ” is not an available name and there is no type series.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD68FFB8AFA61E22FC8CBBD8.taxon	distribution	Distribution. Brazil, Bolivia, Peru, Trinidad (Endrödi 1985), Ecuador (Carvajal et al. 2011), Colombia (LópezGarcía et al. 2015), Venezuela (Escalona & Joly 2006), Costa Rica, Panama (Ratcliffe 2003), El Salvador, Nicaragua, Honduras (Ratcliffe & Cave 2006), Guatemala, and Mexico (Ratcliffe et al. 2013). Locality records (Fig. 43). 815 examined specimens from ANDES, BCRC, BMNH, CAFESI, CEIOC, CEUA, CEUN, CIUQ, CNIN, CTI, EMAC, FSCA, ICN, IEXA, LEUC, MEFLG, MEKRB, MLS, MN / UFRJ, MPUJ, MZUSP, SENASA, UNAB, UNSM, UPN, USNM. Some records from CMNC, Endrödi (1969), Escalona & Joly (2006), Ratcliffe & Cave (2006), and Ratcliffe et al. (2013). BOLIVIA (79). Beni (5): Rio Bent, Cavinas; Reyes; Trinidad; Vaca Diez. Cochabamba (1): Chaparé. La Paz (1): Río Madidi. Santa Cruz (72): Buena Vista, Fauna & Flora Hotel; Saavedra; Potrerillos del Guenda Reserva Natural. BRAZIL (52). Amazonas (34): Benjamin Constant, Rio Javari; Borba, Rio Madeira; Codajas; Manaos; Olivença, Rio Solimões; São Paulo de Olivença. Mato Grosso (1): Barra do Tapirapé. Rondônia (14): Porto Velho, Rio Madeira; Porto Velho, San Antonio. Imprecise data (3). Brazil; Río Inga-Putumayo; Serpa. COLOMBIA (252). Amazonas (65): Leticia; Leticia, Macedonia, Resguardo Indígena Ticuna; Tarapacá, Río Tacana. Antioquia (16): Apartadó; El Carmen de Viboral; Medellín; Puerto Berrío; Santa Elena, Vereda El Mazo. Arauca (3): Alred; Río Arauca. Boyacá (2): Villa de Leyva, Santuario de Fauna y Flora Iguaque. Caldas (6): La Dorada; Manizales; Manizales, Vereda Hoyo Frío, Reserva Planalto Cenicafé; Manizales, Vereda Maltería, Granja Tesorito; Palestina, Vereda Santágueda, Granja Montelindo, Valle de Santágueda. Caquetá (2): Florencia. Casanare (20): Agua Azul (1 km W); Tauramena, Vereda Villa Rosa, Finca Barcelona; Yopal, Agua Azul, Hacienda Los Rosales; Yopal. Cauca (2): Popayán, Munchique; Santander de Chilichao. Córdoba (2): Ayapel. Cundinamarca (12): Bogotá; Fusagasugá, Cuenca Río Jordán; Guasca; La Calera, Vereda La Jagada; La Mesa, Vereda Acacías; Tocaima. Guaviare (2): Caño Mocuaré; San José del Guaviare, Finca La Floresta. Huila (2): San Agustín. Magdalena (3): Río Frío; Santa Ana. Meta (15): La Macarena; Puerto López; Puerto López, Vda. Menegua; Puerto Gaitán; Puerto Gaitán, Altamira, Los Llaneros; Villavicencio, Correg; Remolinos. Putumayo (1): Puerto Asís. Risaralda (1): Pereira. Santander (5): Carare-Opón; Cimitarra, Correg; Puerto Olaya; Vélez, Campo Capote. Tolima (46): Honda; Valencia. Valle del Cauca (32): Buenaventura; Buga; Bugalagrande; Cali; Palmira; Roldanillo. Vichada (6): Puerto Carreño, Finca El Toro; Puerto Carreño, Barrio El Centro; Santa Rosalia. Imprecise data (8): Colombia. COSTA RICA (85). Alajuela (2): Caño Negro; Parque Nacional Guanacaste. Guanacaste (18): Barra Honda; Cerro El Hacha; Estaciones Los Almendros, Los Murciélagos, Maritza, Pitilla y Santa Rosa; Finca Jenny; Finca Montezuma (SW slope Volcán Tenorio); La Pacífica; Liberia; Playas del Coco; Tierras Morenas. Heredia (34): Barra Colorado; Chilamate; La Selva; La Virgen de Sarapiqui; Puerto Viejo; Río Frío. Limón (15): Estación Esquinas; Estación Las Mellizas; Hamburg Farm; Hitoy Cerere; Parque Nacional Tortuguero; Salvadora Farm. Puntarenas (8): Estación Sirena; Fila Guerra; La Escuadra; Monteverde; Quepos; Rancho Quemado; Reserva Carará; San Vito. San José (4): San José; Estación Bijagual. Santa Clara (2): Gaulipes. Imprecise data (1). Costa Rica. ECUADOR (4). Napo (4): Coca; Estación Biologica Jatun Sacha; Misahualli. EL SALVADOR (2). San Salvador (2): San Salvador. GUATEMALA (33). Alta Verapaz (11): Cahabon; Finca Chisiguan; Finca Chulac; La Tinta; Panzos; Teleman; Valle de Polochic. Baja Verapaz (2): Purulha. El Progreso (1): No data. Escuintla (1): Puerto San José. Izabal (12): Finca Firmeza; Morales; Puerto Barrios; Río Dulce; Santo Tomás. Petén (11): Sayachyl; Yaxhá. Suchitepéquez (2): Finca Parraxe. Zacapa (2): Gualan; Río Hondo. HONDURAS. Atlántida (18): La Ceiba; San Alejo; Tela. Comayagua (3): Siguatepeque. Copán (1): Lancetilla. Cortés (24): La Lima; San Pedro Sula. Francisco Morazán (1): Tegucigalpa. Gracias a Dios (1): Morocon. Olancho (3): Culmi. Imprecise data (1): Honduras. MEXICO (129): Chiapas (18): Boca de Chajul; Cinco Cerros; Ocosingo, Corozal; Ocosingo, Montes Azules; Palenque. Oaxaca (8): Cárdenas; Río Jaltepec; Temascal; Tuxtepec; Uxpanapa. Tabasco (12): Cárdenas; Emiliano Zapata; La Venta; Macultepec; Teapa; Villa Hermosa. Veracruz (91): Alvarado; Catemaco; Cerro del Vigía; Chocamán; Ciudad Alemán; Córdoba; Cosamaloapan; Cujuliapan; Dos Amates; Francita; Huatusco; Las Choapas; Los Tuxtlas; Minatitlán; Naolinco; San Andrés Tuxtla; Sontecomapan; Tinajas; Tlacotalpan; Río Uxpanapa; Veracruz; Xalapa; Zapoapan de Cabañas. NICARAGUA (52). Río San Juan (53): Bartola; Los Gatuzos; Reserva Indio Maíz. Zelaya (1): Bonanza. PANAMA (12). Chiriquí (1): Finca La Suiza. Colón (3): Coco Solo; Gamboa; Santa Rita. Panamá (8): Ancón; Barro Colorado; Ipeti; Isla de Majé; Isla San José; Panamá. PERU (263). Ayacucho (1): La Mar, Santa Rosa. Huánuco (14): Tingo María. Ica (1): Ica. Junín (3): Chanchamayo, La Merced. Loreto (69): Alto Amazonas, Yurimaguas; Explorama Inn, 25 mi. NE Iquitos; Explorama Lodge, 50 mi. NE Iquitos; Iquitos. Pasco (1): Oxapampa, Villa Rica. San Martín (33): Nuevo Progreso; San Agustín; Tocache; Uchiza. Ucayali (11): Laguna Yarinacocha; Pucallpa. Imprecise data (4). Peru. TRINIDAD (1). Port of Spain (1): Port of Spain. VENEZUELA (23). Amazonas (4): Puerto Ayacucho; San Fernando de Atabapo; Suromoni. Barinas (21): Barintas; Barrancas; Río Caparo. Bolívar (15): La Urbana. Cojedes (1): El Tinaco. Guárico (1): Hacienda Nicolasito. Portuguesa (8): San Nicolás. Táchira (9): Complejo Hidroeléctrico Leonardo Ruíz Pineda; Colón; Río Frío; San Cristóbal. Trujillo (1): Valera. Zulia (92): Baralt; Colón; Perijá; Tres Bocas. Natural history. Tomarus bituberculatus is one of the most common species of the genus and is found between 30 – 2800 m, mainly in tropical humid forests. There is one record of an adult in Monstera deliciosa Liebm. (Araceae) in Costa Rica, two in banana debris, and 28 in oil palm. Couturier et al. (1996) reported T. bituberculatus as a pest of rice in the Peruvian Amazonia (Loreto Region).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBBAFA619C1FCE6BA76.taxon	description	(Figs. 9 G, 11 I, 16 D, 19 D, 27 D; 44)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBBAFA619C1FCE6BA76.taxon	description	Description. Habitus as in Fig. 27 D. 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 3 rd. 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 3 rd (Fig. 9 G), 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. 11 I). 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. 16 D, 19 D). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBBAFA619C1FCE6BA76.taxon	diagnosis	Diagnosis. Tomarus colombianus can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal surface nearly smooth (Fig. 27 D); 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. 11 I); apex of metatibia with 9 – 11 spinules; parameres with 2 dorsal teeth on each side, main teeth long, widely separate from secondary teeth (Figs. 16 D, 19 D).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBBAFA619C1FCE6BA76.taxon	distribution	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBAAFA61F3FFA24BADC.taxon	description	(Figs. 16 E, 19 E, 23 B, 27 E; 45)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBAAFA61F3FFA24BADC.taxon	description	Description. Habitus as in Fig. 27 E. Length 21.0 – 29.5 mm; humeral width 11.0 – 14.7 mm. Color dark brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles separated by about 2.5 tubercle diameters. Clypeus narrowed towards apex, base 3.0 – 3.5 times as wide as 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures, denser and deeper on anterior and posterior angles. Apical tubercle prominent, acute; slightly recurved. Subapical fovea deep, rounded, very wide (as wide as the interocular distance), equal in both sexes. Foveal 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 25 – 30 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, rounded tubercles forming 15 – 17 diagonal parallel lines. Pygidium with dense rugosity on basal 3 rd, apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with short, sparse setae. Legs: Protibia tridentate, with a basal small denticle. Apex of metatibia crenulate; with 21 – 25 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase as long as parameres. Parameres with 1 dorsal tooth each side; tooth long, acute (Figs. 16 E, 19 E). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like accessory lamellae, with dense granules at base (Fig. 23 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBAAFA61F3FFA24BADC.taxon	diagnosis	Diagnosis. Tomarus discrepans is recognized by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent (Fig. 27 E); pronotal fovea rounded (as wide as interocular distance); metasternum with short setae; protibia tridentate and with an additional basal denticle; apex of metatibia with 21 – 25 spinules; parameres with a long, acute tooth on each side, apical half strongly contracted, apices widely expanded (Figs. 16 E, 19 E). Taxonomic remarks. Tomarus discrepans was described by Escalona & Joly (2006) to include specimens from Venezuela similar in external morphology to T. gyas but with the characteristic parameres of the male. The authors said that the species was also in Panama and Costa Rica based on the illustrations of the parameres presented for T. gyas in Ratcliffe (2003). Although in the present revision the presence of the species is confirmed in the countries from Mexico to Panama, T. gyas is also distributed in Costa Rica and Panama, and so examination of parameres is necessary for correct identification of both species.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD65FFBAAFA61F3FFA24BADC.taxon	distribution	Distribution. Venezuela, Panama, Costa Rica, El Salvador, Honduras, Nicaragua, Guatemala, and Mexico. It is probably found along the Caribbean coast of Colombia. Locality records (Fig. 45). 184 specimens examined from CNIN, EMAC, FSCA, IEXA, UNSM, and USNM. Some records from Escalona & Joly (2006). COSTA RICA (15). Cartago (2): Turrialba. Limón (4): Hamburg Farm; Salvadora Farm; Sixaola. San José (9): Estación La Selva; Heredia; Santa Tecla; Tena. EL SALVADOR (7). La Libertad (2): San Andrés; Santa Tecla. San Salvador (3): San Salvador. Santa Ana (1): Santa Ana. Usulután (1): Usulután. GUATEMALA (48). Alta Verapaz (6): Chulac; La Tinta; Panzos; Senahú. El Progreso (1): El Jicaro. Izabal (35): Cerro Negro Norte; Finca Firmeza; Morales. Retalhuleu (1): Retalhuleu. San Marcos (1): Puente Talismán. Santa Rosa (1): Mal País. Suchitepéquez (3): Santa Bárbara; Finca Parraxe; Finca Moca, Volcán Atitlán. HONDURAS (62). Atlántida (31): La Ceiba; Tela; Parque Nacional Cuero y Salado; Parque Nacional Pico Bonito. Comayagua (1): Siguatepeque. Copán (1): Lancetilla. Cortés (8): La Lima; San Pedro Sula. El Paraiso (2): Capiré (8 km SE). Gracias a Dios (3): Reserva Río Plátano; Wampusirpi. Olancho (10): Catamacas; Montaña de Melacate. Yoro (6): Cuevitas; Parque Nacional Pico Pijol. MEXICO (9). Chiapas (8): Boca de Chajul; Las Guacamayas; Ocosingo. Veracruz (1): El Plan. NICARAGUA (7). Granada (1): La Calera. Río San Juan (4): Bartola; Río Papaturro. Zelaya (2): Bonanza; Cerro Saslaya. PANAMA (28). Colón (3): Madden Dam. Panamá (24): Altos de Maje; Barro Colorado; La Chorrera; Llano-Carti Road; Panamá. VENEZUELA (6). Carabobo (1): Nirgua. Cojedes (1): San Carlos. Portuguesa (1): Estación Experimental San Nicolás, 56 km de Guanare. Táchira (1): Complejo Hidroeléctrico Leonardo Ruíz Pineda, Las Cuevas. Zulia (2): Perijá, El Tucuco; Perijá, Kasmera, Río Yasa. Natural history. Adults can be collected using lights at night. The species is found mainly in tropical dry to humid forests below 650 m and infrequently in montane forest at 1,200 – 1,400 m (Ratcliffe et al. 2013, as T. gyas).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD64FFBCAFA61E59FCC3BFE0.taxon	description	(Figs. 2 I, 3 F, 4 F, 5 G, 6 D, 7 F, 8 J, N, 16 F, 19 F, 27 F; 42)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD64FFBCAFA61E59FCC3BFE0.taxon	materials_examined	Female lectotype here designated (ZMHB) “ 27452 / Brit. Guyana / Moritz // Tomarus / Gyas Er. // SYNTYPUS / Tomarus / gyas Erichson, 1848 / labelled by MNHUB 2016 // Tomarus gyas / Erichson, 1848 / LECTOTYPE / Det: M. M. López-García 2016 ”. Female paralectotype (ZMHB) “ Guyana Moritz // HistColl. (Coleoptera) Nr. 27452 / Tomarus / Gyas Erichs. / Brit. Guyan. / Zool. Mus. Berlin // SYNTYPUS / Tomarus / gyas Erichson, 1848 / labelled by MNHUB 2016 ”. Female paralectotype (ZMHB) “ Hist-Coll. (Coleoptera) Nr. 27452 / Tomarus / Gyas Erichs. / Brit. Guyan. / Zool. Mus. Berlin // SYNTYPUS / Tomarus / gyas Erichson, 1848 / labelled by MNHUB 2016 ”. Female paralectotype (ZMHB) “ Hist-Coll. (Coleoptera) Nr. 27452 / Tomarus / Gyas Erichs. / Brit. Guyan. / Zool. Mus. Berlin // SYNTYPUS / Tomarus / gyas Erichson, 1848 / labelled by MNHUB 2016 ”. Type locality not explicity mentioned in the original description. Lacordaire (1855) attributed the name of the species to Erichson (1848), who gave the first description even though he had cited it as “ T. gyas Hoffgg. ” (Hoffmannsegg).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD64FFBCAFA61E59FCC3BFE0.taxon	description	Description. Habitus as in Fig. 27 F. Length 25.0 – 29.0 mm; humeral width 13.0 – 15.5 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 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth triangular, separated by a tooth diameter. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth (Fig. 5 G). Mentum abruptly constricted at apical 3 rd. Galea of maxilla with teeth 5 and 6 well developed (Fig. 4 F). Interocular distance 3.5 times an eye width. Pronotum: Surface with minute to small punctures, denser and deeper on anterior and posterior angles. Apical tubercle prominent, acute; slightly recurved. Subapical fovea deep, rounded, wide (1 / 2 – 2 / 3 as wide as interocular distance), narrower and shallower in females, 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 25 – 30 parallel lines (Fig. 8 N). Abdomen: Apex of tergite IV with a triangular wide area with large, rounded tubercles forming 15 – 17 diagonal parallel lines (Fig. 8 J). Pygidium with dense rugosity on basal 3 rd, apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with short, sparse setae. Legs: Protibia tridentate, with a basal, small denticle. Apex of metatibia crenulate, with 21 – 30 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase as long as parameres. Parameres with 1 dorsal tooth each side; tooth long, acute; apices triangularly expanded (Figs. 16 F, 19 F). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 or 7 spine-like accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD64FFBCAFA61E59FCC3BFE0.taxon	diagnosis	Diagnosis. Tomarus gyas is recognized by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent (Fig. 27 F); pronotal fovea rounded (as wide as interocular distance), smaller in female; metasternum with short setae; protibia tridentate and with an additional basal denticle; apex of metatibia with 21 – 30 spinules; parameres with a long, acute tooth on each side, apical half strongly narrowed, apices triangularly expanded (Figs. 16 F, 19 F). Taxonomic remarks. Tomarus gyas is one of the most problematic species of the genus because of variable taxonomic characters of external morphology and similar shape of the parameres that sometimes make difficult the differentiation with close species, which have been proposed as synonyms by some authors. After the original description by Erichson (1848), Arrow (1914) described T. amazonicus to include specimens from Brazilian Amazonia. However, Endrödi (1969) synonymized it under T. gyas based on the approximate shape of the parameres and difficulty to delimitate the differences on the shape of the pronotal fovea and protibiae. More recently, Escalona & Joly (2006) described T. discrepans, which was differentiated only by the shape of the parameres, but it was not recognized in later works such as Ratcliffe et al. (2013). The present phylogenetic analysis and taxonomic revision shows that T. amazonicus is a valid species not close to T. gyas and differing in several characters, such as the shape of the frontal tubercles, setation on the metasternum, stridulatory apparatus, and internal sac of the aedeagus. Tomarus discrepans, although close to T. gyas, is also a different, valid species.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD64FFBCAFA61E59FCC3BFE0.taxon	distribution	Distribution. Argentina to Costa Rica. No specimens were examined from Ecuador, but specimens were cited by Endrödi (1969, p. 74) from Guayaquil. Natural history. Specimens were collected with light traps between 140 – 2,530 m. Locality records (Fig. 42). 296 specimens examined from BMNH, CEIOC, CEUN, CTI, FSCA, HJG, ICN, LEUC, MEKRB, MEFLG, MLS, MN / UFRJ, MPUJ, MZUSP, SENASA, UPN, UPTC, UNSM, and USNM. Some records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (40). Buenos Aires (2): Quilmes; San Jacinto. Catamarca (2): Catamarca; El Rodeo. Corrientes (5): Bella Vista; Corrientes; Goya; Rio Paraná. Entre Ríos (2): Concordia; Paraná. Formosa (2): Pilcomayo. Jujuy (2): Jujuy; Parque Nacional Calilegua. La Rioja (1): La Rioja. Río Negro (1): General Roca. Salta (12): Campinchuelo; Coronel Moldes; General Güemes; Hipólito Yrigoyen; Parque Nacional El Rey; Pocitos; Rosario de Lerma; Salta; Vespucio. Santa Fé (6): Rosario de Santa Fé; Santo Tomé. Santiago del Estero (2): Amigo; Santiago del Estero. Tucumán (2): Tucumán. BOLIVIA (19). Beni (1): Cavinas. Chuquisaca (1): Camatindi. La Paz (2): Chulumani; Río Madidi. Santa Cruz (14): Buena Vista; Flor de Oro, Parque Nacional Noel Kempff Mercado; Boyuibe (11 km N); Flora & Fauna Hotel (3.7 km S). Tarija (1): Yacuiba. BRAZIL (115). Bahía (1): Salvador, Canela. Espírito Santo (9): Ribeirão do Engano, Vale do Itauna. Mato Grosso (5): Corumba; Porto Velho, Río Tapirapé; Tres Lagoas. Mina Gerais (4): Cambuquira; Caxambu; Lambary; Passa Quatro. Pará (1): Cachimbo; Mangabeira; Mocajuba. Rio de Janeiro (16): Belem; Campo Grande; Itatiaia; Manguinhos; Passa Tres; Serra dos Orgaos; Teresópolis. Rio Grande do Sul (7): Pelotas; Puerto Alegre; São Leopoldo. Rondônia (16): Ariquemes, near Fazenda Rancho Grande (62 km SW); Cacaulandia, Fazenda Rancho Grande (7 km NE); Ouro Preto do Oeste. São Paulo (51): Angatuba; Atibaia; Barueri; Bofete; Itatiba; Jardim America; Monte Alegre, Fazenda Santa Maria; Piracicaba; Pirassununga; Riberao Preto, Fazenda da Pedra; Riberao Preto; Tieté. Imprecise data (2). South Eastern Part. GUIANA (4). No data (4). COLOMBIA (40). Amazonas (2): Puerto Nariño. Antioquia (7): Medellín. Arauca (7): Arauquita, Campamento Caño Limón. Boyacá (1): Puerto Boyacá. Caldas (2): Manizales, Ecoparque Alcazares. Casanare (2): Aguazul; Yopal. Guaviare (1): Caño Mocuaré. Huila (1): Acevedo. Magdalena (1): Río Frío. Meta (3): Granada; Restrepo; Villavicencio. Nariño (2): Pasto. Norte de Santander (1): Cúcuta. Quindío (1): Quimbaya, Reserva Natural La Montaña del Ocaso. Santander (1): Río Cararé. Tolima (1): Espinal. Vichada (1): Gaviotas. No data (3). COSTA RICA (4). Cartago (1): Turrialba. Heredia (2): La Selva. No data (1). PANAMA (1). Panamá (1): Bayano (2.5 km W Ipiti). PARAGUAY (1). Asunción (1): Asunción. PERU (48). Ayacucho (3): La Mar, Santa Rosa. Cuzco (7): La Convención, Kinteroni. Pilcopata; Estación Biológica Villa Carmen; Quince Mil. Huánuco (3): Tingo María. La Libertad (2): Trujillo, Hacienda Casa Grande. Lima (1): Lima. Madre de Dios (28): Amazonas Lodge (N of Atalaya); Reserva Nacional Tambopata, 30 air km SW Puerto Maldonado. San Martín (3): Huallaga; Moyobamba. No data (1). URUGUAY (51). Colonia (3): Colonia; La Floresta. Maldonado (42): Maldonado; Punta Ballena. Montevideo (6): Colón; Montevideo. VENEZUELA (8). Aragua (3): Maracay. Distrito Capital (5): Caracas. Natural history. Adults can be collected using lights at night.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD62FFBFAFA61A90FDBAB81D.taxon	description	(Figs. 16 G, 19 G, 27 G; 44)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD62FFBFAFA61A90FDBAB81D.taxon	materials_examined	Male lectotype designated by Cartwright (1959: 518) (BMNH) “ LECTO- / TYPE // TYPE // Sp. figured. // Acapulco, / Guerrero. / Höge. // B. C. A. Col. II (2) / Ligyrus / laevicollis, Bates // Ligyrus / laevicollis / Bates ♂ // Lectotype / B. laevicollis Bates // Cartwright 1959 ”. Male paralectotype (BMNH) “ PARA- / LECTO- / TYPE // Chilpancingo, / Guerrero. / Höge. // B. C. A. Col. II (2) / Ligyrus / laevicollis, Bates ”. Female paralectotype (BMNH) “ PARA- / LECTO- / TYPE // Belize. / Blancaneaux. // Ligyrus / laevicollis / Bates ♂ // B. C. A. Col. II (2) / Ligyrus / laevicollis, Bates ”. Male paralectotype (MNHN) “ Cordova, / Vera Cruz / Höge. // ♂. // H. W. Bates / Biol. Cent. Amer. // Museum Paris / ex Coll. / R. Oberthur // Tomarus laevicollis / (Bates 188) [(Bates 1888)] / det F. Dupuis 2015 // PARALECTOTYPE // MNHN / EC 6758 ” Type locality: Acapulco, Guerrero, Mexico. Ligyrus bryanti Rivers, 1891: 97. Synonym. Male lectotype (CASC) “ San Jose / del Cabo. // RIVERS / Type. // 10275 // LECTOTYPE / bryanti // California Academy / of Sciences / Type / No. / 61 ” here designated. Type locality: San José del Cabo, Lower California, Mexico.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD62FFBFAFA61A90FDBAB81D.taxon	description	Description. Habitus as in Fig. 27 G. Length 18.0 – 24.5 mm; humeral width 9.8 – 12.2 mm. Color dark reddish brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles separated by about 9 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth triangular, separated by a tooth diameter. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth. Galea of maxilla with teeth 5 and 6 well developed. Mentum abruptly constricted at apical 3 rd. Interocular distance 3 times an eye width. Pronotum: Surface with minute punctures; denser and deeper on anterior and lateral angles, disc impunctate. Apical tubercle small, rounded, visible in lateral view. Subapical fovea very shallow, elongate, narrow (1 / 2 the interocular distance), equal in both sexes, surface rugopunctate. Scutellum: Surface smooth, without punctures. Elytra: Punctures on first interval minute to absent; sutural stria incomplete, present only on apical 1 / 4. 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 3 rd, apex rounded. Venter: Apex of prosternal process flat, subquadrate to longitudinally oval. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate, with a basal small denticle. Apex of metatibia crenulate; with 9 – 11 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 2 dorsal teeth on each side; basal tooth acute, longer than apical tooth that is small and joined to the basal tooth (Figs. 16 G, 19 G). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD62FFBFAFA61A90FDBAB81D.taxon	diagnosis	Diagnosis. Tomarus laevicollis is differentiated by the following character combination: frontoclypeal tubercles conical; pronotal surface nearly smooth (Fig. 27 G); pronotal tubercle small but visible in lateral view; pronotal fovea shallow and narrow (1 / 3 as wide as interocular distance); elytral punctures on first interval minute to absent; sutural stria incomplete, present only on apical 1 / 4; protibia tridentate, with an additional basal denticle; apex of metatibia with 9 – 11 spinules; parameres with 2 dorsal teeth on each side, main teeth long, close to the secondary, small teeth (Figs. 16 G, 19 G).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD62FFBFAFA61A90FDBAB81D.taxon	distribution	Distribution. Mexico, Guatemala, Belize, Honduras, Nicaragua, El Salvador, and Costa Rica (Endrödi 1985; Ratcliffe et al. 2013). Locality records (Fig. 44). 149 specimens examined from BMNH, CAFESI, CNIN, FSCA, EMAC, IEXA, UNSM, and USNM. Some records from Ratcliffe (2003), Ratcliffe & Cave (2006), and Ratcliffe et al. 2013. BE-LIZE (1). Cayo (1): Blancaneaux. COSTA RICA (2). Limón (2): Amubri; Hitoy Cerere. EL SALVADOR (1). San Miguel (1): Laguna el Jocotal. GUATEMALA (37). Chimaltenango (1): San Martín Jilotepequez. Escuintla (15): Chulamar; La Gomera; Puerto de Ixtapa; San José. Guatemala (2): Ciudad de Guatemala. Huehuetenango (1): No data. Petén (1): No data. San Marcos (2): Puerto de Ocos; Tilapa. Santa Rosa (14): Chiquimulilla; La Avellana; Monterrico; Playa Hawai; Taxisco. Zacapa (1): San Lorenzo, Cerro de los Monos. HONDURAS (69). Atlántida (51): Curla; Danto; Ceiba; La Montañita; Jujiapa; Reserva Cuero y Salado. Gracias a Dios (5): Reserva Río Plátano. Isla Roatán (10): Sandy Bay; Spyglass Hill. MEXICO (209). Baja California (28): San Juan del Cabo; San Lucas; Santa Rosa; Santiago; Todos Santos (25 mi. SE). Chiapas (10): Acapetahua; Cinco Cerros; Pipijiapán; Puerto Arista; Tapachula; Villa Flores. Guerrero (19): Acapulco; Chilpancingo; Ixtapa-Zihuatanejo; Jochuchutla; Jocutla; Rodecia. Jalisco (48): Careyes; Chamela; Melaque; Puerto Vallarta; Tenacatita. Michoacán (3): Cohuayana; Playa Azul; San Telmo. Nayarit (58): Acaponeta; San Blas; Ruíz; Tepic. Oaxaca (5): La Ventosa; Río Grande; Salina Cruz (5 km N); Tehuantepec. Puebla (1): Huachinango. Quintana Roo (1): Cancún. San Luíz Potosí (1): Tamuin. Sinaloa (15): El Zapotillo; Malpica; Mazatlán; Venado; Villa Unión. Tabasco (2): Centla, Manglar de la Barra San Pedro; Ruinas de Comalcalco. Veracruz (18): Alvarado; Córdoba; La Tomata; Palma Sola; Tecolutla; Tuxpan; Veracruz. NICARAGUA (2). Río San Juan (1): Los Guatuzos. Zelaya (1): Campus Uracan Buffields. Natural history. The species inhabits tropical moist forest areas between 0 to 2,100 m, and many specimens have been collected using lights.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD61FFBEAFA61D99FEC0BF43.taxon	description	(Figs. 13 H, 16 H, 19 H, 23 C, 27 H; 44)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD61FFBEAFA61D99FEC0BF43.taxon	description	Description. Habitus as in Fig. 27 H. Length 18.8 – 24.4 mm; humeral width 9.9 – 12.2 mm. Color reddish brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 conical tubercles separated by about 7 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.3 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 without or with sparse, small punctures, not forming lines parallel to margins. Elytra: First interval punctate, punctures smaller than on other intervals, sutural stria complete. 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. Pygidial surface rugose on basal 4 th and lateral angles, disc with minute punctures; apex regularly rounded in male, strongly dilated in female. Venter: Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae. Legs: Protibia tridentate, with an additional small, basal denticle. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate; with 10 – 12 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 2 pairs of dorso-lateral teeth; basal tooth long, wide, and upwardly arched, apical tooth short and joined to the basal tooth (Figs. 16 H, 19 H). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like, accessory lamellae, with dense granules at base (Fig. 23 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD61FFBEAFA61D99FEC0BF43.taxon	diagnosis	Diagnosis. Tomarus laticaudus can be recognized by the nearly impunctate pronotum; small pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1 / 2 as wide as interocular distance); and protibia tridentate with an additional basal 4 th tooth. It is similar to T. laevicollis but can be differentiated by the strongly dilated pygidial apex in female, and the parameres with the main teeth wider and joined to secondary teeth (Figs. 16 H, 19 H).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD61FFBEAFA61D99FEC0BF43.taxon	distribution	Distribution. Panama (López-García & Deloya 2019). Locality records (Fig. 44). 21 examined specimens from UNSM and USNM. PANAMA (21). Los Santos (8): Corozal, Paraíso. Panamá (12): Ancon; Diablo Heights. No data (1). Natural history. The species inhabits tropical moist forest areas between 10 to 80 m. It has been found mainly in April using lights.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD60FFC1AFA61A23FE03B845.taxon	description	(Figs. 1 I, 8 H, L, 16 I, 19 I, 23 D, 27 I; 43)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD60FFC1AFA61A23FE03B845.taxon	materials_examined	Male lectotype (ZMHB) “ Hist-Coll (Coleoptera) / Nr. 27450 / Tomarus / Maimon Erichs. / Peru, v. Tschudi / Zool. Mus. Berlin // SYNTYPUS / Tomarus / maimon Erichson, 1847 / labelled by MNHUB 2013 // TOMARUS / MAIMON / ERICHSON / LECTOTYPE / B. C. Ratcliffe ”, here designated. Ratcliffe labeled the specimen but did not publish the lectotype designation. Female paralectotype (ZMHB) same labels as lectotype except “ TOMARUS / MAIMON / ERICHSON / LECTOALLOTYPE / B. C. Ratcliffe ”. Male paralectotype (ZMHB) same labels as lectotype except “ TOMARUS / MAIMON / ERICHSON / PARALECTOTYPE / B. C. Ratcliffe 2014 ”. Male paralectotype (ZMHB) “ Maimon / Er / Peru v. Tschudi / Lima // SYNTYPUS / Tomarus / maimon Erichson, 1847 / labelled by MNHUB 2013 // TOMARUS / MAIMON / ERICHSON / PARALECTOTYPE / B. C. Ratcliffe 2014 ”. Type locality: Peru. Heteronychus fossator Burmeister, 1847: 101. Synonym. Female lectotype designated by Dupuis (2016: 31) (MLUH) “ fossator / Dej. / Cayenne Buq // Pronotum + kopf fraglich // Lectotypus / Ligyrus / (Heteronychus) fossator Burm. / Endrödy ” Head and pronotum glued are a different species of another genus with rounded clypeus and bituberculate pronotum. Dupuis (2016) chose the abdomen as the lectotype, which is valid based on Article 73.1.5 of the International Commission on Zoological Nomenclature (1999). Type locality: Cayenne, French Guiana.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD60FFC1AFA61A23FE03B845.taxon	description	Description. Habitus as in Fig. 27 I. Length 21.9 – 28.0 mm; humeral width 11.0 – 13.5 mm. Color dark reddish brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles (Fig. 1 I), separated by about 4.5 – 5.0 tubercle diameters. Clypeus narrowed towards apex (Fig. 1 I), base 3 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures; denser and deeper on anterior and posterior angles. Apical tubercle prominent, rounded, slightly recurved. Subapical fovea deep, rounded, narrow (> 1 / 2 the interocular distance), 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 30 diagonal parallel lines (Fig. 8 L). Abdomen: Apex of tergite IV with a triangular wide area with large, rounded tubercles forming 14 – 15 parallel lines (Fig. 8 H). Pygidium with dense rugosity on basal 3 rd, apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with short, sparse setae. Legs: Protibia tridentate, without basal denticle. Apex of metatibia crenulate, with 15 – 19 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 1 dorsal small tooth each side, rarely without teeth, apices widely dilated (Figs. 16 I, 19 I). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like accessory lamellae, with dense granules at base (Fig. 23 D).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD60FFC1AFA61A23FE03B845.taxon	diagnosis	Diagnosis. Tomarus maimon can be differentiated by the following character combination: frontoclypeal tubercles conical (Fig. 1 I); pronotal tubercle acute and prominent; pronotal fovea transversely oval (as wide as interocular distance); protibia tridentate, without basal denticle; apex of metatibia with 15 – 19 spinules; parameres with 1 dorsal small tooth each side, rarely without teeth; apices roundly dilated (Figs. 16 I, 19 I).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD60FFC1AFA61A23FE03B845.taxon	distribution	Distribution. Brazil, French Guiana, Guyana, Chile, Peru, and Ecuador. Locality records (Fig. 43). 195 examined specimens from IEXA, FSCA, MERKB, MN / UFRJ, SENASA, UNSM, USAAC, USNM, and ZMHB. Some records from CMNC and Endrödi (1969) and Dupuis (2016). BRAZIL (5). Mato Grosso (2): Parque Nacional Xingú; Rosario Oeste. Pará (3): Carindé, Río Guripi; Utinga. CHILE (6). Arica y Parinacota (6): Arica; Valle de Azapa. ECUADOR (5). Guayas (1): Guayaquil. Manabí (2): Bahía de Caraquez; Manta. Los Ríos (2): Los Ríos. FRENCH GUIANA (1). Cayenne (1): Cayenne. GUYANA (1). Georgetown (1): Georgetown. PERU (212). Ancash (1): Santa, Moro. Arequipa (1): Camaná, Quilca. Cajamarca (1): Baños del Inca. Ica (36): Chavín, Cerro Lindo; Chincha; Ica; Paracas, Santa Cruz; Topará; Villacurí. Junín (12): Chanchamayo; La Merced; Pichanaqui; Satipo, Mazamari. La Libertad (15): Ascope, Chócope; Libertad; Pacasmayo; Trujillo; Virú. Lambayeque (2): Chiclayo, Lambayeque. Huánuco (7): Aucayacu, Tingo María. Lima (87): Canta, Santa Rosa de Quives; Cañete; Huaura, Lachay; Huaura, Sayán / Andahuasi; Huaral, Huando; Huarochirí, Matucana; Huarochirí, San Pedro de Casta; Huarochirí, Santa Eulalia; Lima, La Molina; Lurín; Quebrada Verde. Loreto (11): Alto Amazonas, Yurimaguas; Yanamono. Moquegua (1): Ilo, Río Osmore. Piura (11): Chira; Morropón, Chulucanas; Paita; Piura. Tambogrande / Progreso Bajo. Tacna (6): Jorge Basadre, Locumba; Tacna. San Martín (1): Tocache. Imprecise data (20): Peru. Natural history. Tomarus maimon is a relative common species in Peru where it is known as “ gusano blanco ” and has been cited as occasional pest of sugar cane feeding on roots and stumps (Carrasco 1967; Ayquipa & Cueva 1979); it can also attack the roots of the sweet potato that is known as camote (Daza & Rincón 1993). Adults are attracted to lights at night.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1FFFC0AFA61D3FFBD8BDCC.taxon	description	(Figs. 9 H, 16 J, 20 A, 28 A; 44)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1FFFC0AFA61D3FFBD8BDCC.taxon	materials_examined	Male holotype (USNM) “ maracaibo // BB // Tomarus / maracaiboensis / López-García / & Deloya / HOLOTYPE ”. Two male paratypes (USNM, UNSM) “ maracaibo // Tomarus / maracaiboensis / López-García / & Deloya / PARATYPE ” Type locality: Maracaibo, Venezuela.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1FFFC0AFA61D3FFBD8BDCC.taxon	description	Description. Habitus as in Fig. 28 A. Length 20.9 – 23.3 mm; humeral width 10.0 – 12.3 mm. Color brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 conical tubercles separated by about 7 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.3 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, 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. Pygidial surface rugose on basal 4 th and on lateral angles, disc with minute punctures (Fig. 9 H), apex regularly rounded. Venter: Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae. Legs: Protibia tridentate, with an additional small, basal undulation. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate, with 8 – 9 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 2 pairs of dorso-lateral teeth, both equal in length and upwardly arched (Figs. 16 J, 20 A), apices short. Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 4 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1FFFC0AFA61D3FFBD8BDCC.taxon	diagnosis	Diagnosis. Tomarus maracaiboensis 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); protibia tridentate with a small basal undulation; and apex of metatibia with 8 – 9 spinules. It is similar to T. laevicollis, T. colombianus, and T. laticaudus but it can be differentiated by the parameres with the apex very short, the secondary teeth similar in shape and as long as the main teeth (Figs. 16 J, 20 A), and the internal sac with a complex of 4 spinelike accessory lamellae.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1FFFC0AFA61D3FFBD8BDCC.taxon	distribution	Distribution. Maracaibo in Venezuela. Locality records (Fig. 44). Three examined specimens from UNSM and USNM. VENEZUELA (3). Zulia (3): Maracaibo. Natural history. Nothing is known about the natural history of T. maracaiboensis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1EFFC3AFA618A9FE6ABCB5.taxon	description	(Figs. 4 G, 5 H, 6 E, 7 G, 8 I, M, 13 I, 16 K, 20 B, 22 C, 28 B; 46)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1EFFC3AFA618A9FE6ABCB5.taxon	materials_examined	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).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1EFFC3AFA618A9FE6ABCB5.taxon	description	Description. Habitus as in Fig. 28 B. 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. 5 H). Mentum abruptly constricted at apical 3 rd. 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. 8 M). Abdomen: Apex of tergite IV with a triangular wide area with small and large, irregular tubercles not forming defined lines (Fig. 8 I). Pygidial surface rugose on basal 3 rd 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. 16 K, 20 B). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 8 spine-like accessory lamellae, without granules at base (Fig. 22 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1EFFC3AFA618A9FE6ABCB5.taxon	diagnosis	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. 16 K, 20 B); spine-like accessory lamellae of internal sac without granules at base (Fig. 22 C).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1EFFC3AFA618A9FE6ABCB5.taxon	distribution	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC3AFA61871FBE8B9B4.taxon	description	(Figs. 9 I, 16 L, 20 C, 28 C; 47)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC3AFA61871FBE8B9B4.taxon	materials_examined	Male holotype (USNM) “ PERU: Cuzco; / Pilcopata, 600 meters / 11 - 14 XII 1976 / J. B. Heppner, premontane / moist forest // Tomarus / pilcopataensis / López-García & / Deloya / HOLOTYPE ” Type locality: Pilcopata, Cuzco, Peru.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC3AFA61871FBE8B9B4.taxon	description	Description. Habitus as in Fig. 28 C. Length 27.7 mm; humeral width 14.3 mm. Color dark reddish brown. Head: Frons coarsely rugose. Frontoclypeal region with 2 transverse tubercles separated by about 3 tubercle diameters. Clypeus narrowed towards apex, base 3.8 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 4.1 times an eye width. Pronotum: Surface with dense, large punctures. Apical tubercle small, rounded. Fovea shallow, striate, and about a 1 / 5 as wide as interocular distance. Scutellum: Surface with minute, sparse punctures. Elytra: First interval punctate, punctures similar in size to other intervals; sutural stria complete. Inner surface of apex with rounded, small tubercles forming 40 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, elongate tubercles forming 16 diagonal parallel lines. Pygidial surface coarsely and densely rugose (Fig. 9 I); apex regularly rounded. Venter: Apex of prosternal process flat, longitudinally oval. Metasternum with short, sparse setae on anterior angles. Legs: Protibia tridentate with an additional inconspicuous, basal convexity. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia slightly crenulate, with 33 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with lateral teeth short, widely triangular; apical 4 th narrowed, apices outwards (Figs. 16 L, 20 C). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 3 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC3AFA61871FBE8B9B4.taxon	diagnosis	Diagnosis. Tomarus pilcopataensis can be recognized by the following character combination: frontal tubercles transverse; pronotal fovea and tubercle inconspicuous; pronotum deeply and densely punctate (Fig. 28 C); metasternum with sparse, short setae on anterior angles; and internal sac with a complex of 3 spine-like accessory lamellae. Tomarus pilcopataensis is similar to T. pumilus (Prell) and T. roigjunenti Neita and Ratcliffe, 2017 but differs from them by the completely rugose surface of the pygidium, Fig. 9 I (rugose only on the basal 3 rd in the other 2 species). The general shape of the parameres of T. pumilus (Fig. 16 N) is similar, but in T. pilcopataensis the apical 4 th is strongly narrowed, and the lateral medial teeth are wider and not directed backwards (Fig. 16 L).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC3AFA61871FBE8B9B4.taxon	distribution	Distribution. This species is known only from its type locality (Pilcopata, Cuzco) in the moist premontane forest of Peru at 600 m. Locality records (Fig. 47). One examined specimen from USNM. PERU (1). Cuzco (1): Pilcopata. Natural history. Nothing is known about the natural history of T. pilcopataensis.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC2AFA61F71FCE7B870.taxon	description	(Figs. 1 J, 9 J, 11 J, 14 E, 16 M, 20 D, 22 A, 28 D; 48)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC2AFA61F71FCE7B870.taxon	materials_examined	Male lectotype designated by Endrödi (1969: 64) (ZMHB) “ Venezuela / San Fernando de Apure / L. Laglaize 5 - 10 1897 // Ligyrus / pullus Prell / ♂ - Type // Lectotypus / Ligyrus / pullus / Prell / Endrody ”. Female paralectotype (ZMHB) “ Venezuela / San Fernando de Apure / L. Laglaize 5 - 10 1897 // Ligyrus / pullus Prell / ♀ - Type // Paratypus / Ligyrus / pullus / Prell ”. Type locality: San Fernando de Apuré, Apuré, Venezuela.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC2AFA61F71FCE7B870.taxon	description	Description. Habitus as in Fig. 28 D. Length 13.6 – 23.3 mm; humeral width 9.0 – 15.6 mm. Color dark reddish brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 transverse, inconspicuous tubercles (Fig. 1 J). Clypeus short (3.2 times wider than long) and narrowed towards apex (base 3.8 times wider than apex). Clypeal teeth transverse separated by 1.5 tooth diameters. Mandible with incisor longer than second tooth, lateral tooth not well developed. Mentum subtriangular. Galea of maxilla with vestigial teeth, teeth 3 and 5 absent. Interocular distance 3.0 – 3.5 times an eye width. Pronotum: Surface with dense, large punctures. Apical tubercle absent. Fovea shallow, punctate, narrow (about 1 / 10 as wide as interocular distance) (Fig 1 J). Scutellum: Surface with small, sparse punctures. Elytra: First interval punctate, punctures similar in size to other intervals; sutural stria complete. Inner surface of apex with rounded, small tubercles forming parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, elongate tubercles forming diagonal parallel lines. Pygidial surface coarsely and densely rugose on basal half, apical half with sparse, deep punctures (Fig. 9 J), apex regularly rounded. Venter: Apex of prosternal process flat, rounded. Metasternum with long, sparse setae on anterior angles. Legs: Protibia tridentate, without basal denticles (Fig. 11 J). Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia slightly crenulate, with 26 – 29 spinules. Male genitalia: Spiculum gastrale without basal plates (Fig. 14 E). Phallobase 1.25 times longer than parameres. Parameres with lateral teeth short, widely triangular; apical 4 th narrowed, apices rectangularly expanded (Figs. 16 M, 20 D). Internal sac with copulatory lamella and long lamellar spiny belt; with a complex of 26 spine-like, accessory lamellae, without dense granules at base (Fig. 22 A).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC2AFA61F71FCE7B870.taxon	diagnosis	Diagnosis. Tomarus pullus can be recognized by the following character combination: pronotal fovea elongate (Fig. 1 J); pronotum without tubercle, deeply and densely punctate (Fig. 1 J); clypeus short (3.2 times wider than long); mandible with incisor longer than second tooth, lateral tooth not well developed (Fig. 1 J); mentum subtriangular; galea of maxilla with vestigial teeth, teeth 3 and 5 absent; metasternum with sparse, long setae on anterior angles; parameres with lateral teeth short, widely triangular; apical 4 th narrowed, apices rectangularly expanded (Figs. 16 M, 20 D); and internal sac with lamellar spiny belt long and a complex of 26 spine-like accessory lamellae, without granules at base (Fig. 22 A).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1DFFC2AFA61F71FCE7B870.taxon	distribution	Distribution. The species is restricted to the Orinoquia region of Colombia and Venezuela. Locality records (Fig. 48). 9 examined specimens from MPUJ, UPN, USNM, ZMHB. Some data from Escalona & Joly (2006). COLOMBIA (2). Meta (1): Remolinos, Cafam Llanos. Vichada (1): Puerto Carreño. VENEZUELA (11). Amazonas (3): Puerto. Ayacucho; Samariapo. Apuré (2): San Fernando de Apuré. Aragua (2): Maracay. Bolívar (3): La Urbana, Río Orinoco; Pararuma. Monagas (1): Uverito. Natural history. Some specimens were collected at lights.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	description	(Figs. 1 K, 16 N, 20 E, 28 E; 47)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	materials_examined	Male lectotype designated by Endrödi (1969: 71) (NHMB) “ Corralitos // Ligyrus / pumilus / ♂ - Type // Lectotypus / Ligyrus / pumilus / Prell / Endrody // TOMARUS / PUMILUS / (PRELL) / det. B. C. Ratcliffe 2011 ”. Type locality: Corralito, Córdoba, Argentina. Ligyrus rosettae Endrödi, 1968: 68. Synonym.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	materials_examined	Male holotype (HNHM) “ Holotypus / Ligyrus / rosettae / Endr. // Proc. San Clemente del Tuyu Arg. / Prov. Bs. Aires, xii. 49 // Museum Frey / München // KEYENCE 105 ”. Male paratype (HNHM) “ Proc. San Clemente / del Tuyu Arg. / Prov. Bs. Aires. xii. 49 // Paratypus / Ligyrus / rosettae / Endr. ”. Female paratype (HNHM) “ Mendoza / Rep. Arg. / v. 26 // Chili / Faz ded. // Paratypus / Ligyrus / rosettae / Endr. ” Male paratype (HNHM) “ Mendoza / Rep. Arg. / v. 26 // Chili / Faz ded. // Ligyrus / scarabaeinus Pertÿ // Paratypus / Ligyrus / rosettae / Endr. ” Type locality: San Clemente del Tuyu, Buenos Aires, Argentina.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	description	Description. Habitus as in Fig. 28 E. Length 19.2 – 23.6 mm; humeral width 9.1 – 12.3 mm. Color dark reddish brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 transverse tubercles (Fig. 1 K), separated by about 3 tubercle diameters. Clypeus narrowed towards apex, base 3.8 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance equal to 4.1 times an eye width. Pronotum: Surface with dense, large punctures. Apical tubercle small, rounded. Fovea shallow, striate, narrow (1 / 5 as wide as interocular distance). Scutellum: Surface with minute, sparse punctures. Elytra: First interval punctate, punctures similar in size to the other intervals; sutural stria complete. Inner surface of apex with rounded, small tubercles forming 38 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, elongate tubercles forming about 16 diagonal parallel lines. Pygidium with dense rugosity on basal 3 rd; apex regularly rounded. Venter: Apex of prosternal process flat, longitudinally oval. Metasternum with short, sparse setae on anterior angles. Legs: Protibia tridentate with an additional inconspicuous, basal convexity. Metatibia triangular; apex slightly crenulate, with 21 – 27 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with lateral teeth short, acute; apical half narrowed, apices outwards (Figs. 16 N, 20 E). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 1 – 3 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	diagnosis	Diagnosis. Tomarus pumilus can be recognized by the following character combination: frontal tubercles transverse (Fig. 1 K); pronotal fovea and tubercle inconspicuous (Fig. 28 E); pronotum deeply and densely punctate; metasternum with sparse, minute setae on anterior angles; internal sac with a complex of 1 – 3 spine-like, accessory lamellae; and parameres with lateral teeth short, triangular, acute; apical half narrowed, apices outwards (Figs. 16 N, 20 E).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1CFFC5AFA61D35FE47BEC8.taxon	distribution	Distribution. Argentina. Locality records (Fig. 47). 20 examined specimens from HNHM, NHUMB, and USNM. Some records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (33). Buenos Aires (4): Buenos Aires; San Clemente del Tuyu; Villa Gesell. Catamarca (1): Catamarca. Córdoba (4): Corralito; Tanti. Mendoza (16): Mendoza; Pedregal; Potrerillos; Puente del Inca-Las Heras; San Rafael; Telteca-La Valle. Río Negro (5): Allen; General Roca. Salta (1): Urundel. San Juan (2): Jachal; San Juan. Natural history. Adults are attracted to lights at night (Neita-Moreno & Ratcliffe 2017) and are mainly found between November and March.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1BFFC4AFA61BADFE7BBCB4.taxon	description	(Figs. 9 K, 16 O, 20 F, 28 F; 47)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1BFFC4AFA61BADFE7BBCB4.taxon	materials_examined	Male holotype not examined. Male paratype (USNM) “ Neuquen / Capital Federal / 1. iii. 1961 / Dafuern // TOMARUS / ROIGJUNENTI / NEITA & RATCLIFFE / PARATYPE ”. Female paratype (USNM) “ Neuquen / Capital / 1. iii. 1961 // TOMARUS / ROIGJUNENTI / NEITA & RATCLIFFE / PARATYPE ”. Two female paratypes (USNM) “ Mendoza / C. S. Reed // TOMARUS / ROIGJUNENTI / NEITA & RATCLIFFE / PARATYPE ” Type locality: Neuquén, Argentina.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1BFFC4AFA61BADFE7BBCB4.taxon	description	Description. Habitus as in Fig. 28 F. Length 26.3 – 27.6 mm; humeral width 12.6 – 13.9 mm. Color dark reddish brown to black. Head: Frons coarsely rugose. Frontoclypeal region with 2 transverse tubercles separated by about 3 tubercle diameters. Clypeus narrowed towards apex, base 3.8 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 4 times an eye width. Pronotum: Surface with dense, large punctures. Apical tubercle small, rounded. Fovea shallow, striate, narrow (1 / 5 as wide as interocular distance). Scutellum: Surface with minute, sparse punctures. Elytra: First interval punctate, punctures similar in size to other intervals; sutural stria complete. Inner surface of apex with rounded, small tubercles forming 38 – 40 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with large, elongate tubercles forming about 18 diagonal, parallel lines. Pygidium with dense rugosity on basal 3 rd (Fig. 9 K), apex regularly rounded. Venter: Apex of prosternal process flat, longitudinally oval. Metasternum with short, sparse setae on anterior angles. Legs: Protibia tridentate with an additional inconspicuous, basal convexity. Metatibia triangular; apex slightly crenulate, with 21 – 25 spinules. Male genitalia: Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with lateral teeth large, acute; apical half narrowed, apices outwards, rounded (Figs. 16 O, 20 F). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 1 – 3 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1BFFC4AFA61BADFE7BBCB4.taxon	diagnosis	Diagnosis. Tomarus roigjunenti can be recognized by the following character combination: frontal tubercles transverse; pronotal fovea and tubercle inconspicuous (Fig. 28 F); pronotum deeply and densely punctate; metasternum with sparse, minute setae on anterior angles; internal sac with a complex of 1 – 3 spine-like, accessory lamellae; and parameres with lateral teeth large and long, arcuate; apical half narrowed, apices outwards, rounded (Figs. 16 O, 20 F).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1BFFC4AFA61BADFE7BBCB4.taxon	distribution	Distribution. Argentina (Neita-Moreno & Ratcliffe 2017). Locality records (Fig. 47). 4 examined specimens from USNM. Records from Neita-Moreno & Ratcliffe (2017). ARGENTINA (20). Chubut (1): Chubut. Buenos Aires (1). Buenos Aires. Mendoza (4). Tunuyan; Uspallata. Neuquén (9). Capital Federal; Barrancas; Collón Cura; Covunco; Isla Victoria; Laguna Blanca; Vatra Lauquen. Río Negro (5). Allen; Cipolletti; General Roca; Río Negro; Villa Regina. Natural history. Some specimens were found in sand dunes and adults are attracted to lights at night (NeitaMoreno & Ratcliffe 2017).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC4AFA61871FA3EBAE8.taxon	description	(Figs. 5 F, 7 H, 9 L, 13 J, 14 F, 16 P, 20 G, 22 B, 28 G; 49)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC4AFA61871FA3EBAE8.taxon	materials_examined	Male holotype (MNHN) “ Peru. / Tomarus / Er. / Maimon // MUSÉUM PARIS / 1952 / coll. R. OBERTHÜR // Tomarus rostratus n. sp. / HOLOTYPE / F. Dupuis 2014 // HOLOTYPE // MNHN / EC 4187 ”. Type locality: Peru.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC4AFA61871FA3EBAE8.taxon	description	Description. Habitus as in Fig. 28 G. Length 17.4 – 27.5 mm; humeral width 8.3 – 10.0 mm. Color dark reddish brown. Head: Frons coarsely and irregularly rugose. Frontoclypeal region with 2 transverse tubercles separated by about 3 tubercle diameters. Clypeus trapezoidal, not strongly narrowed toward apex; apex with 2 triangular teeth separated by about 1.5 tooth diameters. Mandible with 2 apical teeth and a lateral, widely rounded tooth (Fig. 5 F). Mentum abruptly constricted at apical 3 rd. Galea of maxilla with teeth 5 and 6 well developed. Pronotum: Surface punctate, punctures rounded, smaller than punctures on elytra; densely distributed on apical and lateral margins. Apical margin with a small tubercle, visible in lateral view. Pronotum without rugose fovea, with small to large punctures on each side of tubercle. Scutellum: Surface without punctures. Elytra: Sutural stria complete. Elytral punctures dense and deep, punctures mainly ocellate, 3 pairs of distinct double rows. Abdomen: Pygidium convex (male) to flat (female) in lateral view; surface deeply rugopunctate on basal 3 rd with disc smooth (male) (Fig. 9 L), or completely rugopunctate (female). Apical margin of sternite VII straight in male, slightly emarginate with notch broad and shallow in female. Apex of sternite VIII emarginate, notch wide in male and narrow and deep in female. Venter: Prosternal process long, thick, apex transversely oval to rounded, and bordered with long, pale yellow setae. Metasternum with short, sparse setae on anterior angles. Legs: Protibia tridentate with an additional small, basal denticle. Apex of metatibia slightly crenulate, with 8 – 15 spinules. Female genitalia: Coxite surface strongly concave (Fig. 13 J). Male genitalia: Spiculum gastrale with basal plates (Fig. 14 F). Parameres widened at apical 4 th, elongate from basal 4 th to apex; sides parallel, without teeth or lateral projections (Fig. 16 P), regularly curved at lateral view (Fig. 20 G). Internal sac with small copulatory lamella, lamellar spiny belt reduced, complex of accessory lamellae with 6 spines joined at the base by dense granules (Fig. 22 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC4AFA61871FA3EBAE8.taxon	diagnosis	Diagnosis. Tomarus rostratus is unique in the genus because of its extremely elongate parameres, without lateral teeth. It can be also differentiated by the following combination of characters: clypeus trapezoidal; pronotum with small tubercle and without rugose fovea, with small to large punctures on each side of tubercle (Fig. 28 G); protibia tridentate with a small denticle; apical border of pygidium strongly truncate in male; apical margin of sternite VIII emarginated in female; coxite surface strongly concave (Fig. 13 J); spiculum gastrale with basal plates (Fig. 14 F); internal sac of aedeagus with small copulatory lamella and reduced lamellar spiny belt (Fig. 22 B).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC4AFA61871FA3EBAE8.taxon	distribution	Distribution. Tomarus rostratus is an endemic species from the coastal desert of Peru (Dupuis 2014) and Chile (López-García & Deloya 2018). Locality records (Fig. 49). 39 examined specimens from FSCA, MEKRB, MNHN, USNM, SENASA, UGBC. CHILE (12). Arica y Parinacota: Cuya; Puente Chapa (50 km S Arica); Valle de Azapa (6 km E Arica). PERU (27). Arequipa (3): Camaná, Quilca; Quiroz. Lima (19): Canta, Santa Rosa de Quives; Cañete; Huarochirí, Santa Eulalia; Huaura, Végueta, Medio Mundo; Lima. No data (5): Peru. Natural history. Specimens have been found between 12 – 1,036 m elevation between August and February.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC7AFA61F8DFE16B9EA.taxon	description	(Figs. 2 J, 4 H, 6 F, 14 G, 16 Q, 20 H, 28 H; 45)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC7AFA61F8DFE16B9EA.taxon	materials_examined	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 / EC 6732 ”. Male paratype (MNHN) “ Los Mochis / Sinaloa, Mex / 22 - VII- 1955 // R & J Selander // PARATYPE / Bothynus / selanderi / Cartwright // PARATYPE // MNHN / EC 6733 ”. Type locality: Los Mochis, Sinaloa, Mexico.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC7AFA61F8DFE16B9EA.taxon	description	Description. Habitus as in Fig. 28 H. 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 3 rd. Galea of maxilla with teeth 5 and 6 well developed (Fig. 4 H). 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 3 rd. 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. 14 G). Phallobase 1.1 times longer than parameres. Parameres without dorsal teeth, apex wide, triangularly dilated (Figs. 16 Q, 20 H). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC7AFA61F8DFE16B9EA.taxon	diagnosis	Diagnosis. Tomarus selanderi can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent (Fig. 28 H); 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. 16 Q, 20 H).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD1AFFC7AFA61F8DFE16B9EA.taxon	distribution	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.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	description	(Figs. 2 K, 4 I, 6 G, 13 K, 14 H, 16 R, 20 I, 28 I; 41)	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	synonymic_list	Female holotype (PERC) “ Dunedin, Fla. / W. S. B. Coll. / 6 - 4 /. 1913 // Purdue / Blatchley / collection // TYPE // Ligyrus / subtropicus / sp. nov. // PERC / 0066475 ”. Type locality: Dunedin, Florida. Ligyrus blatchleyi Cartwright, 1944: 34. Synonym.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	materials_examined	Male holotype (USNM) not examined. Allotype (USNM) “ Miami, Florida / 20. Jun. 1933 / Frank N. Young // ALLOTYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Allotype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 // Tomarus subtropicus / (Blatchley) / Det. B. C. Ratcliffe 2015 ”. Two female paratypes (USNM) “ Miami / 25. Jun. 1933 / Frank N. Young // PARATYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Paratype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 ”. Two female paratypes (USNM) “ Miami / 22. Jun. 1933 / Frank N. Young // PARATYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Paratype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 ”. Female paratype (USNM) “ Miami / 16. May. 1933 / Frank N. Young // PARATYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Paratype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 ”. Male paratype (USNM) “ Miami / 30. Jun. 1933 / Frank N. Young // PARATYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Paratype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 ”. Male paratype (USNM) “ Savannah Ga. / 13. Jul. 1937 / P. W. Fatting // PARATYPE / USNM / 56936 // Ligyrus / blatchleyi / Cartwright / Paratype // Bothynus subtropicus (Blatchley) / Det. Cartwright 1958 ”. Another five paratypes not examined. Type locality: Miami, Florida, United States of America.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	description	Description. Habitus as in Fig. 28 I. Length 17.5 – 27.5 mm; humeral width 8.8 – 14.0 mm. Color dark reddish brown to black. Head: Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 transverse, low tubercles separated by about 7 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth small, transverse, separated by a tooth diameter. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth. Mentum abruptly constricted at apical 3 rd (Fig. 2 K). Galea of maxilla with teeth 5 and 6 vestigial (Fig. 4 I). Interocular distance 3.5 times an eye width. Pronotum: Surface with small punctures, denser and deeper on anterior and lateral angles, disc impunctate. Apical tubercle small, rounded, visible in lateral view. Subapical fovea shallow, elongate, narrow (1 / 3 the interocular distance); equal in both sexes, surface rugopunctate. Scutellum: Surface with sparse, minute punctures not forming lines parallel to margins. Elytra: Punctures on first interval as large as those on other intervals, sutural stria complete. Inner surface of apex with rounded, large tubercles forming 17 parallel lines. Abdomen: Apex of tergite IV with a triangular wide area with small, irregular tubercles forming 12 – 14 diagonal, nearly parallel lines. Pygidium with dense rugosity on basal 3 rd, apex rounded. Venter: Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae. Legs: Protibia tridentate, rarely with basal denticle. Apex of metatibia crenulate, with 11 – 14 spinules. Female genitalia: Coxite concave near the inner margin (Fig. 13 K). Male genitalia: Spiculum gastrale without basal plates (Fig. 14 H). Phallobase 1.2 times longer than parameres. Parameres with 2 dorsal teeth each side; basal tooth large, strongly curved backwards, longer than apical tooth that is short (Figs. 16 R, 20 I). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like, accessory lamellae, with dense granules at base.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	diagnosis	Diagnosis. Tomarus subtropicus can be differentiated by the following character combination: frontoclypeal tubercles transverse and low (as in T. adoceteus, Fig. 1 H); galea of maxilla with teeth 5 and 6 vestigial (Fig. 4 I); pronotal surface with small punctures (Fig. 28 I); pronotal tubercle small but visible in lateral view; pronotal fovea shallow and narrow (1 / 3 as wide as interocular distance); protibial tridentate, rarely with an additional basal denticle; apex of metatibia with 11 – 14 spinules; parameres with 2 dorsal teeth on each side, main teeth long and strongly curved backwards, secondary teeth short (Figs. 16 R, 20 I).	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
A209493BFD19FFC6AFA61C8BFEA3BAD4.taxon	distribution	Distribution. Southeastern United States of America (Endrödi 1985; Ratcliffe & Cave 2017). Locality records (Fig. 41). 125 examined specimens from FSCA, UNSM, USNM. Counties listed from Ratcliffe & Cave (2017). UNITED STATES OF AMERICA (492). Alabama (1): Mobile. Florida (459): Alachua; Baker; Broward; Calhoun; Charlotte; Collier; Dade; Dixie; Duval; Franklin; Hendry; Hillsborough; Indian River; Lee; Manatee; Monroe; Palm Beach; Pinellas; Sarasota; Seminole; St. Lucie; Taylor; Volusia. Georgia (6): Chatham; Glynn. Lousiana (18): Lafourche. South Carolina (8): Beaufort; Charleston. Natural history. The species is univoltine. Adults emerge between June and August, eggs are laid from late June to early August, and larvae are active from July to February (Kostromytska & Buss 2008); winter is passed as a larva. The larva was described by Gordon & Anderson (1981). It is considered the species of Scarabaeidae of primary economic importance in sugar cane fields in the United States of America (Cartwright 1959; Gordon & Anderson 1981) but also attacks the roots of other grasses (Stenotaphrum secundatum (Walt. Kuntze) and Cynodon dactylon (L.) (Poaceae) (Kostromytska & Buss 2011). The females of T. subtropicus lay their eggs at the base of the plant, and then the larvae feed on the roots until the plant lacks support. Cherry et al. (2017) sampled 20 fields of sugar cane in Florida during 2015 and 2016 and found that although T. subtropicus was once the most abundant white grub in this region, no specimens were found. They believed that the most logical explanation for this drastic population decline is the increased flooding of sugarcane fields as this species is less flood tolerant compared with wireworms.	en	López-García, Margarita M., Deloya, Cuauhtémoc (2022): Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision. Zootaxa 5211 (1): 1-119, DOI: https://doi.org/10.11646/zootaxa.5211.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5211.1.1
