Ligyrus (Anagrylius) cuniculus ( Fabricius, 1801 ) López-García, Margarita M. & Deloya, Cuauhtémoc, 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 : 25-27

Ligyrus (Anagrylius) cuniculus ( Fabricius, 1801) new combination

( Figs. 7B, D , 9C , 15D , 17D , 25A ; 32 )

Geotrupes cuniculus Fabricius, 1801: 20 . Original combination.

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.

Description. Habitus as in Fig. 25A . 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. 2D ). Clypeus trapezoidal, base 1.5 times as wide as apex; apical angles rounded ( Fig. 2D ). 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. 25A ). 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. 9C ). 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 4th, apices projecting outwards, ventral margin without teeth ( Figs. 15D , 17D ).

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. 15D , 17D ) (present in L. moroni ) ( Figs. 15E , 17E ).

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.

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 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 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 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 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 32. Distributional records of the subgenus Ligyrus (Anagrylius).