Tomarus villosus ( Burmeister, 1847 )

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 : 12

Tomarus villosus ( Burmeister, 1847)

( Fig. 2e , 3j–m , 4h , 5h , 6o–p , 7h , 8h , 10o–p , 11h , 12h , 13h , 14h , 15h , 17d )

Podalgus villosus Burmeister 1847: 120 (original combination).

Oryctes nitidicollis Solier 1851: 79 (synonym).

Ligyrus patagonus Steinheil 1872: 560 (synonym).

Redescription: Male. Length 14.9–18.4 mm; width 7.8–9.9 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. 4h ). Clypeus strongly rugopunctate to transversely rugose, tapering to slightly emarginate, broad apex with 2 reflexed teeth. Mandibles with 2 teeth, apical tooth subacute, subapical notch broad, deep ( Fig. 6o–p ); labium densely setose, paraglossa undeveloped, apex tapering to narrow apex ( Fig. 7h ); maxilla with galea developed and with 4 teeth ( Fig. 5h ). Pronotum: Surface smooth or with small, sparse punctures except for lateral margins where punctures small to moderate in size and density. Anterior margin with small, median tubercle, fovea absent ( Fig. 8h ). Elytra: Surface with micropunctures and large punctures mixed; large punctures moderately dense, ocellate, and with 3 pairs of double rows. First broad interval with or without large punctures. Pygidium: Surface densely punctate, punctures becoming denser in basal and lateral angles; punctures small to moderate in size, weakly ocellate, with minute and tawny setae. In lateral view, male with convex surface, females with weakly convex surface. Legs: Protibia tridentate, basal tooth slightly removed. Protarsus in male simple, not enlarged. Mesotibial transverse carinae vary from short to long, arcuate to nearly straight ( Fig. 10o–p ). Metatibia with 1 basal, transverse carina interrupted at middle (carina with spine-like setae) and 1 medial, transverse carina; apex with about 18 spinules ( Fig. 10p ). 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, long ( Fig. 2e ). Male Genitalia: Parameres ( Fig. 11h , 12h , 13h ) subtriangular, apices curving outwards, mid-shaft with large tooth extending laterally. Internal sac ( Fig. 15h ) lacking copulatory lamellae.

Female: Length 17.1–19.1 mm; width 8.5–9.6 mm. Similar to males, but pronotal surface with punctures slightly less dense, and elytra with punctures slightly denser. Gonocoxite smaller than gonocoxal sternite and lacking a semicircular furrow ( Fig. 14h ).

Figures

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 17. Distribution of Tomarus spp. a) T. roigjunenti n. sp. b) T. rosettae. c) T. spinipenis n. sp. d) T. villosus.