Euligyrus similis ( Endrödi, 1968 ) 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 : 19-21

Euligyrus similis ( Endrödi, 1968) new combination

( Figs. 2H , 3D , 4A , 5A , 7A, C , 10A–B , 11A , 12A, 12G , 15B , 17B , 24B ; 31 )

Ligyrus similis Endrödi, 1968: 166 . Original combination.

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.

Description. Habitus as in Fig. 24B . 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. 1A ). 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. 24B ). 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 3rd. Strongly (male) to slightly (female) convex in lateral view. Legs: Protibia tridentate, basal tooth distant from others ( Fig. 11A ). Protarsus enlarged; inner claw bifid ( Fig. 10A–B ). Metatibia not narrowed before apex, sides nearly parallel ( Fig. 12G ). 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. 15B , 17B ).

Diagnosis. See the Diagnosis of Euligyrus ebenus .

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

Type species. Ligyrus cicatricosus Prell, 1937 , here designated

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. 8D ). 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. 8A ). 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. 11B ). Apical margin of mesotibia and metatibia dentate ( Fig. 12B, 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. 13B ). 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. 21B ).

Diagnosis. Clypeus trapezoidal, apex with 2 transverse, small teeth ( Fig. 1B ). Mentum subquadrate ( Fig. 2F ). Galea with 6 teeth (3 dorsal, 3 ventral) ( Fig. 4B ). Pronotum with apical marginal bead incomplete on middle 1/5 ( Fig. 1B ). Apex without tubercle, with a small swelling and shallow, depression. Surface of pronotum, elytra, and pygidium with large, dense punctures ( Fig. 24C ). Metepisternum glabrous. Male protarsus simple. Protibia tridentate, basal tooth distant from others ( Fig. 11B ). Apical margin of mesotibia and metatibia dentate ( Fig. 12C, I ). Sternite VIII of female narrowly emarginate. Coxite surface strongly concave ( Fig. 13B ). Internal sac of aedeagus without copulatory lamella; accessory lamellae with a flat, oval base (comma-like lamellae) ( Fig. 21B ).

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.

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 .

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 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 4. Maxilla. A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Ligyrodes) relictus, D) L. (Ligyrus) burmeisteri, E) L. (Ligyrus) villosus, F) Tomarus gyas, G) Tomarus maternus, H) T. selanderi, I) T. subtropicus.

FIGURE 5. Mandible. A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Ligyrodes) relictus, D) L. (Ligyrus) burmeisteri, E) L. (Ligyrus) villosus, F) Tomarus rostratus, G) T. gyas, H) T. maternus.

FIGURE 7. Left hindwing in ventral view. A, C) Euligyrus similis, B, D) Ligyrus (Anagrylius) cuniculus, E) L. (Ligyrus) burmeisteri, F) Tomarus gyas, G) T. maternus, H) T. rostratus.

FIGURE 8. Stridulatory apparatus. A, D) Proculigyrus cicatricosus, B, E) Ligyrus (Ligyrus) burmeisteri, C, F) L. (Ligyrodes) relictus, G, K) Tomarus amazonicus, H, L) T. maimon, I, M) T. maternus, J, N) T. gyas.

FIGURE 10. Protarsus and inner claw. A–B) Euligyrus similis, C–E) Ligyrus (Ligyrodes) sallaei.

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 17. Parameres in lateral view. A) Euligyrus ebenus, B) E. similis, C) Proculigyrus cicatricosus, D) Ligyrus (Anagrylius) cuniculus; E) L. (Anagrylius) moroni, F) L. (Ligyrodes) relictus, G) L. (Ligyrodes) sallaei.

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 24. Habitus. A) Euligyrus ebenus, B) E. similis, C) Proculigyrus cicatricosus. Scale bar: 5 mm.

FIGURE 31. Distributional records of the genus Euligyrus.