Tomarus bituberculatus ( Palisot de Beauvois, 1811 )

López-García, Margarita M. & Deloya, Cuauhtémoc, 2022, Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision, Zootaxa 5211 (1), pp. 1-119 : 56-58

Tomarus bituberculatus ( Palisot de Beauvois, 1811)

( Figs. 9F , 11H , 16C , 19C , 27C ; 43 )

Scarabaeus bituberculatus Palisot de Beauvois 1811: 103 . Original combination.

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.

Description. Habitus as in Fig. 27C . 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 3rd. 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 3rd ( Fig. 9F ). Apex rounded. Venter: Apex of prosternal process flat, longitudinally oval to subrectangular. Metasternum with minute setae on anterior angles. Legs: Protibia tridentate ( Fig. 11H ), 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. 16C , 19C ). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 5 spine-like accessory lamellae, with dense granules at base.

Diagnosis. Tomarus bituberculatus can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal tubercle acute and prominent ( Fig. 27C ); pronotal fovea transversely oval (as wide as interocular distance); protibia tridentate, rarely with an additional basal denticle ( Fig. 11H ); 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. 16C , 19C ).

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.

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).

Figures

FIGURE 9. Pygidium of male. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) cuniculus, D) L. (Ligyrus) fossor, E) L. (Ligyrus) paranaensis, F) Tomarus bituberculatus, G) T. colombianus, H) T. maracaiboensis, I) T. pilcopataensis, J) T. pullus, K) T. roigjunenti, L) T. rostratus.

FIGURE 11. Protibia in frontal view.A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) relictus, E) L. (Ligyrodes) sallaei, F) L. (Ligyrus) allonasutus, G) L. (Ligyrus) villosus, H) Tomarus bituberculatus, I) T. colombianus, J) T. pullus.

FIGURE 16. Parameres in frontal view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon, J) T. maracaiboensis, K) T. maternus, L) T. pilcopataensis, M) T. pullus, N) T. pumilus, O) T. roigjunenti, P) T. rostratus, Q) T. selanderi, R) T. subtropicus.

FIGURE 19. Parameres in lateral view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon.

FIGURE 27. Habitus of the species of Tomarus: A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon. Scale bar: 5 mm.

FIGURE 43. Distributional records of Tomarus bituberculatus and T. maimon.