Ligyrus (Ligyrus) allonasutus López-García & Deloya, 2022

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 : 37-38

Ligyrus (Ligyrus) allonasutus López-García & Deloya new species

( Figs. 2B , 3H , 11F , 12E, K , 13E , 15I , 21E , 25F ; 34 )

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 ”.

Description. Holotype. Habitus as in Fig. 25F . 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. 11F ). Protibial surface sparsely punctate. Protarsus of male simple, inner claw simple. Metatibia strongly contracted at apex ( Fig. 12K ). Apex of metatibia entire, with 13 spinules. Male genitalia: Spiculum gastrale thin, apex truncate. Parameres with sides nearly parallel, lateral ventral teeth short ( Figs. 15I , 18A ). Internal sac with lamellar spiny belt long (5 times longer than copulatory lamellae); accessory lamella thickened, elongate ( Fig. 21E ).

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. 13E ). 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.

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. 15I , 18A ); internal sac with accessory lamella thick and elongate ( Fig. 21E ); coxite of female strongly concave ( Fig. 13E ).

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).

Figures

FIGURE 2. Labium. A) Ligyrus (Anagrylius) moroni, B) Ligyrus (Ligyrus) allonasutus, C) L. (Ligyrus) gibbosus, D) L. (Ligyrus) villosus, E) L. (Ligyrodes) relictus, F) Proculigyrus cicatricosus, G) Euligyrus ebenus, H) E. similis, I) Tomarus gyas, J) Tomarus selanderi, K) Tomarus subtropicus. Abbreviations: LL: Ligular lobes, LP: Labial palpus, M: Mentum, S: Suture.

FIGURE 3. Maxilla. A) Ligyrus (Anagrylius) moroni, B) L. (Ligyrodes) sallaei, C) Ligyrus (Ligyrus) allonasutus. Numeration of teeth of galea: D) Euligyrus similis, E) L. (Ligyrodes) sallaei, F) Tomarus gyas, G) L. (Ligyrus) fossor, H) L. (Ligyrus) allonasutus.

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 12. Mesotibia (A–F). A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) sallaei; E) L. (Ligyrus) allonasutus, F) L. (Ligyrus) villosus. Metatibia (G–L). G) Euligyrus similis, H) Proculigyrus cicatricosus, I) Ligyrus (Anagrylius) moroni, J) L. (Ligyrodes) sallaei, K) L. (Ligyrus) allonasutus, L) L. (Ligyrus) villosus.

FIGURE 13. Female genital plates. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) sallaei, E) L. (Ligyrus) allonasutus, F) L. (Ligyrus) bidentulus, G) L. (Ligyrus) neglectus, H) Tomarus laticaudus, I) T. maternus, J) T. rostratus, K) T. subtropicus. Abbreviations: SCx: Subcoxite, Cx: Coxite.

FIGURE 15. Parameres in frontal view. A) Euligyrus ebenus, B) E. similis, C) Proculigyrus cicatricosus, D) Ligyrus (Anagrylius) cuniculus, E) L. (Anagrylius) moroni, F) L. (Ligyrodes) peruvianus, G) L. (Ligyrodes) relictus, H) L. (Ligyrodes) sallaei, I) L. (Ligyrus) allonasutus, J) L. (Ligyrus) bidentulus, K) L. (Ligyrus) burmeisteri, L) Ligyrus (Ligyrus) fossor, M) L. (Ligyrus) gibbosus, N) L. (Ligyrus) nasutus, O) L. (Ligyrus) neglectus, P) L. (Ligyrus) paranaensis, Q) L. (Ligyrus) peninsularis, R) L. (Ligyrus) rubripes, S) L. (Ligyrus) ruginasus, T) L. (Ligyrus) villosus.

FIGURE 18. Parameres in lateral view.A) L. (Ligyrus.) allonasutus, B) L. (Ligyrus) bidentulus, C) L. (Ligyrus) burmeisteri, D) Ligyrus (Ligyrus) fossor, E) L. (Ligyrus) gibbosus, F) L. (Ligyrus) nasutus, G) L. (Ligyrus) neglectus, H) L. (Ligyrus) paranaensis, I) L. (Ligyrus) peninsularis, J) L. (Ligyrus) rubripes, K) L. (Ligyrus) ruginasus, L) L. (Ligyrus) villosus.

FIGURE 21. Internal sac. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) sallaei, E) L. (Ligyrus) allonasutus. Abbreviations:AL:Accessory lamellae, Br: Brush of setae, CL: Copulatory lamella, LSB: Lamellar spiny belt, Ra: Raspula.

FIGURE 25. Habitus of the species of Ligyrus. Subgenus Anagrylius: A) Ligyrus cuniculus, B) L. moroni. Subgenus Ligyrodes: C) L. peruvianus, D) L. relictus, E) L. sallaei. Subgenus Ligyrus: F) L. allonasutus, G) L. bidentulus, H) L. burmeisteri. Scale bar: 5 mm.

FIGURE 34. Distributional records of Ligyrus (Ligyrus) allonasutus and L. nasutus.