Ligyrus (Ligyrus) peninsularis ( Casey, 1915 ) 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 : 48-49

Ligyrus (Ligyrus) peninsularis ( Casey, 1915) new combination

( Figs. 15Q , 18I , 26F ; 39 )

Oxygrylius peninsularis Casey, 1915: 209 . Original combination. Male syntype (USNM). Not examined. Type locality: San José del Cabo, Baja California Sur, Mexico.

Description. Habitus as in Fig. 26F . 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. 15Q , 18I ). Internal sac with lamellar spiny belt short (2.5 times longer than copulatory lamellae); accessory lamella thickened, elongate.

Diagnosis. Ligyrus peninsularis is recognized by the triangular clypeus, with 1 central tooth (as in L. ruginasus , Fig. 1E ); frontal carina straight, not interrupted; pronotum with prominent tubercle and deep, rounded fovea; pronotal punctation deep and large ( Fig. 26F ); metatibia elongate with apex strongly narrowed; parameres with sides nearly parallel, lateral ventral teeth short, apex slightly expanded ( Figs. 15Q , 18I ); 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).

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).

Figures

FIGURE 1. Head. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Ligyrodes) peruvianus, D) L. (Anagrylius) moroni, E) L. (Ligyrus) ruginasus, F) L. (Ligyrus) nasutus, G) L. (Ligyrus) bidentulus, H) Tomarus adoceteus, I) T. maimon, J) T. pullus, K) T. pumilus.

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 26. Habitus of the species of Ligyrus. Subgenus Ligyrus: A) Ligyrus fossor, B) L. gibbosus, C) L. nasutus, D) L. neglectus, E) L. paranaensis, F) L. peninsularis, G) L. rubripes, H) L. ruginasus, I) L. villosus. Scale bar: 5 mm.

FIGURE 39. Distributional records of Ligyrus (Ligyrus) ruginasus and L. peninsularis.