Tomarus burmeisteri ( Steinheil, 1872 )

Neita-Moreno, Jhon César, Alexander, Recursos Biológicos, Agustín, San, de, Villa, Leyva, Boyacá, Ratcliffe, Brett C., Collections, Systematics Research, of, University, Museum, Nebraska State, Hall, Nebraska, of, University, Nebraska, Lincoln, Ne & Usa, 2017, The genus Tomarus Erichson (Coleoptera: Scarabaeidae: Dynastinae: Pentodontini) in Argentina, Chile, and Uruguay, Insecta Mundi 2017 (547), pp. 1-36 : 5-6

Tomarus burmeisteri ( Steinheil, 1872)

( Fig. 1b , 2d , 3b–f , 4b , 5b , 6c–d , 7b , 8b , 10c–d , 11b , 12b , 13b , 14b , 15b , 16b , 18 )

Ligyrus burmeisteri Steinheil 1872: 560 (original combination).

Ligyrus distinctus Fairmaire 1892: 245 (synonym).

Redescription. Male. Length 10.9–13.6 mm; width 5.7–7.3 mm. Color light to dark reddish brown. Head: Frons mostly impunctate between eyes and coarsely rugopunctate just behind frontoclypeal carina. Frontoclypeal region with transverse carina slightly separated at middle ( Fig. 4b ). Clypeus strongly rugopunctate to transversely rugose, tapering to broad, slightly emarginate apex with 2 reflexed teeth. Mandibles with 2 teeth, apical tooth subacute, subapical notch moderately deep ( Fig. 6c–d ); labium densely setose, paraglossa undeveloped, converging to apex ( Fig. 7b ); maxilla with galea developed, with 5 teeth. Pronotum: Surface crazed, sparsely punctate; punctures moderately large, umbilicate. Anterior margin with small, median tubercle, fovea absent ( Fig. 8b ). Elytra: Surface with micropunctures and large punctures mixed, large punctures moderately dense, ocellate, and in 3 pairs of double rows. First broad interval with or without large punctures. Pygidium: Surface sparsely punctate, punctures becoming denser in basal angles; punctures small to moderate in size, weakly ocellate. In lateral view, surface in males convex, weakly convex in females. Legs: Protibia tridentate, basal tooth slightly removed. Protarsus in male simple, not enlarged. Mesotibia relatively short, transverse carinae with stout, dense spinules ( Fig. 10c–d ). Metatibia with 1 basal, transverse carina interrupted at middle and with spine-like setae and 1 medial, transverse carina; apex truncate with about 25 spinules ( Fig. 10d ). Venter: Prosternal process tall, thick; apex small, suboval, often with slender, long setae. Mesosternal surface sparsely setose, slightly concave at middle. Metasternal surface densely punctate, punctures ocellate, setose; setae slender and long. Male Genitalia: Parameres ( Fig. 11b , 12b , 13b ) subtriangular, rounded apices curving outwards, strong tooth projecting laterally at mid-shaft. Internal sac ( Fig. 15b ) lacking copulatory lamellae.

Female. Length 12.0– 13.9 mm; width 7.0– 7.3 mm. Similar to male, but pronotal disc smooth. Abdominal ventrite VIII entire as in Fig. 1h . Pygidium slightly concave. Gonocoxite smaller than gonocoxal sternite, lacking a semicircular furrow ( Fig. 14b ).

Distribution. Tomarus burmeisteri is known from Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay ( Endrödi 1985).

Locality Records ( Fig. 16b ). 535 specimens examined from BCRC, CMNC, FSCA, IADIZA, MACN, MLPA, NMW, UMSP, USNM.

ARGENTINA (517). BUENOS AIRES (143): Algarrobo, Azul, La Plata, Mar del Plata, Capital Federal, Mercedes, Olivos, Quequén-Saladón, San Blas, San Isidro, Hogar Agrícola-San Cayetano, Puán, Seis de Septiembre, Tandil, Villa Iris (30 km SW). CATAMARCA (94): Catamarca-Capital, Guayamba, La Viña. CHACO (6): La Represa de los Indios, Resistencia. CHUBUT (6): Las Chapas, Puerto Madryn. CÓRDOBA (23): Alta Gracia, Alpa Corral, Campamento Balnea-San Javier, Córdoba, Cosquín, Cruz del Eje, Espinillo, Los Olmos, La Carlota, Mar Chiquita, Rió Primera, Santa Maria, Villa de Soto. COR- RIENTES (3): Corrientes-Capital, San Roque, San Tolomé. ENTRE RÍOS (1): Concordia. FORMOSA (1): Clarinda (50 km NW), JUJUY (49): Jujuy-Capital, El Volcán, Ledesma, Parque Nacionla Calilegua, Santa Barabara, Volcán a Tilcara, Yuto. LA PAMPA (12): General Pico, Guatiache, Santa Rosa, Winfredo. LA RIOJA (13): Jaguey, La Rioja-Capital, La Rioja-Paquis, Patiquía, Pozo de Piedra. MENDOZA (36): Cerro Quadral, Chacras de Coria-Luján de Cuyo, Mendoza-Capital, Moliches, Rodeo de la Cruz- Guaymallen, San Martín, Santa Rosa. MISIONES (10): Departamento de Concepción Santa María, Loreto, Posadas, San Pedro. SALTA (31): Cafayate- 1 km Norte, Campo Quijano, El Carril (15 km S), El Corralito, El Naranjo, General Güemes, La Rioja, Rosario de Lerma, Salta-Capital, San Loenzo, San Martín. SAN JUAN (3): Desamparada, San Juan-Capital, Sarmiento. SAN LUIS (15): Arizona, Arizona (13 km S), San Luis-Capital. SANTA FÉ (8): Reconquista, Rosario, San Espiritu, San Javier, Santo Tomé. SANTIAGO DEL ESTERO (16): Río Salado, Robles, Santiago del Estero-Capital. TUCUMÁN (53): Lamadrid-Arboles-Gendarmes Camping, RN 38-Estación, Estación-Gendarmería, San Pedro de Colalao, Tucumán-Capital. PARAGUAY (1). CAAGUAZÚ (1): Estancia Primera. URUGUAY (17). MALDONADO (4): Punta Bellena. MONTEVIDEO (13): Colón, Montevideo-Capital, Río del Uruguay.

Temporal Distribution. January (58), February (151), March (19), April (5), July (5), August (2), October (16), November (37), December (35).

Diagnosis. Tomarus burmeisteri superficially resembles T. spinipenis because both species are similar in size, color, and lack a pronotal tubercle and fovea. Tomarus burmeisteri can be distinguished by a mentum that lacks slender, long setae at its base ( Fig. 7b ); metatibial surface on the external edge rugopunctate; and different parameres (compare Fig. 11a , 12a , 13a and 11g , 12g , 13g ). The genital plate of the females is also different (compare Fig. 14a and 14g ). This species has a broad distribution that is reflected in slight differences in the parameres; compare, for example, the parameres of the Guaymallen (Mendoza, Argentina) population with those from the Argentine provinces of Catamarca, Mendoza and La Pampa ( Fig.18 ).

Tomarus burmeisteri also resembles T. bidentulus but is easily distinguished by the absence of the pronotal tubercle, which is present in T. bidentulus .

Life History. Adults are attracted to lights during the southern summer. This species occupies a wide diversity of habitats ranging from dry to wet.

Figures

Figure 1. Habitus of Tomarus spp. a) Tomarus bidentulus (Fairmaire) dorsal view. b) Tomarus burmeisteri (Steinheil) dorsal view. c) Tomarus gyas Erichson dorsal view. d) Tomarus rosettae (Endrödi) dorsal view. e) Tomarus roigjunenti n. sp. dorsal view, holotype. f) Tomarus roigjunenti n. sp. dorsal view, allotype.

Figure 2. Habitus of Tomarus spp. a) Tomarus rosettae (Endrödi) dorsal view. b) Tomarus spinipenis n. sp. dorsal view, holotype. c) Tomarus spinipenis dorsal view, allotype. d) Tomarus burmeisteri (Steinheil) dorsal view. e) Tomarus villosus (Burmeister) ventral view. f) Tomarus gyas ventral view. g) Tomarus bidentulus (Fairmaire) abdomen segments, ventral view (female). h) Tomarus bidentulus abdomen segments, ventral view (male).

Figure 3. Types of Tomarus spp. a) T. bidentulus female holotype, dorsal view. b) T. burmeisteri male holotype, dorsal view. c) T. burmeisteri male holotype, lateral view. d) T. burmeisteri male holotype parameres, caudal view. e) T. burmeisteri male holotype parameres (lateral view). f) Label for T. burmeisteri holotype. g) T. gyas male holotype, dorsal view and labels and parameres, caudal view). h) T. maimon holotype, dorsal view. i) T. rosettae male holotype, dorsal view. j) T. villosus male holotype, dorsal view. k) T. villosus male holotype, dorsal view of head. l) T. villosus male holotype, parameres, caudal view. m) Label for T. villosus holotype.

Figure 4. Head, dorsal view of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 5. Maxilla, ventral view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 6. Mandibles, ventral view, of Tomarus spp. a–b) T. bidentulus. c–d) T. burmeisteri. e–f) T. gyas. g–h) T. maimon. i–j. T. roigjunenti n. sp. k–l. T. rosettae. m–n. T. spinipenis n. sp. o–p. T. villosus.

Figure 7. Labium, dorsal view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 8. Pronotum, dorsal view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 10. Meso- and metatibia, dorsal view, of Tomarus spp. a–b) T. bidentulus. c–d) T. burmeisteri. e–f) T. gyas. g–h) T. maimon. i–j. T. roigjunenti n. sp. k–l. T. rosettae. m–n. T. spinipenis n. sp. o–p. T. villosus.

Figure 11. Parameres, caudal view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 12. Parameres, lateral view, of: a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 13. Parameres, anterior view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus.

Figure 14. Gonocoxite, ventral view, of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus. Components are: (Gx) gonocoxite), (Gs) gonocoxal sternite.

Figure 15. Internal sac of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. gyas. d) T. maimon. e) T. roigjunenti n. sp. f) T. rosettae. g) T. spinipenis n. sp. h) T. villosus. Components are: (AL) accessory lamella, (CL) copulatory lamella (CL), (Ra) raspula, (Esp) speculum, and (Fg) a complex, spike-like flagellum that varies in length and thickness.

Figure 16. Distribution of Tomarus spp. a) T. bidentulus. b) T. burmeisteri. c) T. maimon. d) T. gyas.

Figure 18. Variation of the parameres, anterior view, of T. burmeisteri in Argentina.