Bothynus entellus ( LePeletier & Serville, 1828 )

Duarte, Paulo R. M. & Grossi, Paschoal C., 2020, Bothynus entellus (LePeletier & Serville) (Coleoptera: Scarabaeidae: Dynastinae) species group: taxonomic revision and description of two new species, Zootaxa 4750 (1), pp. 101-121 : 102-105

Bothynus entellus ( LePeletier & Serville, 1828)

( Figs. 1 A–E; 7A; 8A; 9A; 10A, E; 11A; 12A)

Scarabaeus entellus LePeletier & Serville, 1828: 347 (original description).

Scarabaeus glaucon Perty, 1830: 45 (original description, synonym).

Scarabaeus glycon Perty, 1830: 214 (original description, synonym).

Strategus montesuma Hope, 1837: 88 (original description, synonym).

Diagnosis. Bothynus entellus is distinguished from other species of the group by the following combination of characters: major and median males with two pronotal horns directed forward ( Fig. 1 A–B); two or one lateral tubercles instead of horns in minor males ( Fig. 1C ). Minor males of B. entellus without lateral tubercles are similar to B. araya but B. entellus differs by the stridulatory apparatus with weakly marked carinae ( Fig 7A ); basal half of parameres with outer sides gradually constricted toward middle area; apical half of parameres with lateral “flaps” rounded and broader than basal half in caudal view ( Fig. 8A ). Female of B. entellus differs from other females of the group by the pronotal surface nearly completely punctate, with the concavity moderately and disc finely punctate ( Fig. 10A ); proventrite with two small furrows at anterolateral angles ( Fig. 10E ); tergite VIII with weakly emarginate and transverse apical invagination (about 4.4 times wider than long) ( Fig. 11A ).

Type material. Not examined.

Additional material examined. 147 males and 50 females. Brasil: Bahia: Camacan, RPPN Serra Bonita , 27–29.II.2012 , M.L. Monné— 1♂ ( MNRJ); Una, (10 km SE San José), Mata Atlântica , luz, 7–25.X.1986 — 2♂ 2♀ ( MNRJ). Espírito Santo: Santa Teresa , 830 m, 8.I.1966 — 1♂ ( DZUP), 12.XI.1966 — 1♀ ( DZUP), 15.XI.1966 — 1♂ 1♀ ( DZUP), 20.XI.1966 — 2♂ ( DZUP), 9.XI.1967 — 2♀ ( DZUP), 15.XI.1967 — 8♂ 1♀ ( DZUP), 16.XI.1967 — 49♂ 11♀ ( DZUP), 30.XI.1967 — 1♀ ( DZUP), 9.XII.1967 — 2♂ ( DZUP), 22.XII.1967 — 1♂ ( DZUP), 16.VII.1967 — 1♂ ( DZUP), 17.VII.1967 — 2♀ ( DZUP), 8.I.1966 — 1♂ ( DZUP), 2.XI.1967 — 1♂ ( DZUP), 13.XI.1955 , Santos, Mach- ado, & Barros— 1♂ ( MNRJ), XII.1965 , Paviote— 3♂ ( MNRJ). Rio Bonito , X–XI.1965 — 6♂ 1♀ ( DZUP). Tijuco Preto , XI.1931 , Campos Seabra— 13♂ ( MNRJ), X–XI.1937, M. Alvarenga— 3♂ 1♀ ( DZUP). Minas Gerais: Ibiá , X.1965 — 1♂ ( DZUP). Belo Horizonte , XI.1951 , F. Justus Jorge— 1♂ ( DZUP). Lavras , 15.II.1999 — 1♂ ( DZUP). Águas Vermelhas , XI.1991 — 1♂ ( DZUP). Maria da Fé , XI.1980 , G.S. Andrade— 1♂ ( MNRJ). Paraná: Colombo, Embrapa, BR 476— Km 20, (25°19’19.92”S, 49°9’25.60”W), 952 m, 31.X.1986 — 1♂ ( DZUP), (25°19’20.52”S, 49°9’24.42”W), 952 m, 4.XI.1986 — 1♂ 4♀ ( DZUP), XI.2014 , M. Savaris— 1♂ ( CERPE). Morretes, Marumbí , 15–16.X.1966 — 1♂ ( DZUP). Piraquara , X.2010 — 3♂ ( CERPE), XI.2009 — 2♂ ( CERPE). Ponta Grossa , IV.1943 , F. Justus Jorge— 1♂ ( DZUP). São José dos Pinhais, Serra do Mar, BR 277— Km 54, (25°34’2.74”S, 48°59’2.27”W), 966 m, 4.XI.1987 — 1♀ ( DZUP). Três Portões , 5.XI.1967 — 1♂ 1♀ ( MZUSP). Londrina , XII.1936 — 1♂ ( MZUSP). Tapejara , I.1953 , F. Justus Jorge— 1♂ ( DZUP). Pernambuco: São Lourenço da Mata, Estação Ecológica de Tapacurá , 6.X.1982 , J.C. Barros— 2♂ ( CERPE). Rio de Janeiro: Angra dos Reis, Jussaral , IX.1935 , D. Mendes— 1♀ ( MNRJ). Cabo Frio— 1♂ ( MNRJ). Rio de Janeiro, Corcovado , X.1961 — 1♂ ( DZUP), IX.1934 , Travassos— 1♂ ( MNRJ), 5.XI.1959 , Zajciw— 1♂ ( MNRJ), 9.X.1975 , Campos Seabra— 1♂ ( MNRJ). Itatiaia, Instituto Biológi- co, Entomologia Agrícola , XI.1929 , Dario Mendes— 1♂ ( MNRJ). Parque Nacional do Itatiaia , (22°27’56.95”S, 44°33’52.86”W), (409 m), 6–8.XI.2009 , M.L. & M.A. Monné— 1♂ ( MNRJ), (22°29’9.46”S, 44°36’52.79”W), 568 m, 11–13.XI.2011 , Cupelo, Machado, Souza, Simões, & Carell— 1♂ ( MNRJ), 21.XI.1929 , Zikán— 1♂ ( MNRJ), 7.XI.1957 , W. Zikán— 1♀ ( MNRJ). Nova Friburgo, Sans Souci , (22°16’50.61”S, 42°30’45.49”W), 1039 m, 6.XI.2005 , E. & P. Grossi— 1♂ ( EPGC), (22°16’50.06”S, 42°30’45.20”W), 1068 m, I.1995 , E. & P. Grossi— 2♂ ( EPGC), (22°16’50.84”S, 42°30’45.17”W), 1068, XII.1999 , E. & P. Grossi— 1♀ ( EPGC), XI. 2009 — 2♀ ( CERPE), XI.2014 — 1♂ 2♀ ( CERPE). Petrópolis , 23. VIII.1962 — 1♂ ( DZUP); Petrópolis. Independência , X.1920 , Mario Rosa— 4♀ ( MNRJ). Rio de Janeiro, 8.II.1950 , 1♀ ( DZUP), XI.1965 , 1♀ ( DZUP). Teresópolis , IX.1956 , Campos Seabra— 1♂ ( DZUP), XI.1946 , V.M. de Ribeiro— 1♂ 1♀ ( MNRJ), IX.1956 , P.R. Teles— 1♂ ( MNRJ). Vassouras , 12.XII.1955 , Santos Barros— 1♀ ( MNRJ). Santa Catarina: Água Clara, Rio Negrinho , XI.1925 , A. Maller— 1♀ ( MNRJ). Campos Novos , (27°13’50.10”S, 51° 7’14.53”W), 940 m, II.2011 – 4♂ ( DZUP). Corupá , X.1952 — 1♀ ( DZUP), 14.XII.1956 — 1♂ ( DZUP), XII.1937 , A. Maller— 1♂ ( MNRJ). Rio Vermelho , 14.XII.1960 — 1♂ ( DZUP), 14.X.1958 — 2♂ 1♀ ( DZUP), (26°16’31.38”S, 49°22’31.89”W), X.1952 — 1♀ ( DZUP). São Paulo: Barueri , 16.XI.1955 , K. Lenko— 1♂ ( MNRJ). Cantareira , XII.1923 — 1♂ ( MZUSP). Itú , Fazenda Pau d’ Alho, 28–29.X.1965 , M. Alvarenga— 1♀ ( DZUP). Parque Mathias , II.1936 — 1♂ ( MZUSP). Rio Grande , X.1900 — 1♂ ( MZUSP). Salesópolis, Estrada Biológica da Boracéia , (23°35’9.32”S, 45°51’24.37”W), 984 m, 23–29.XI.2008 , E. & P. Grossi— 1♀ ( EPGC), I.1950 , M. Alvarenga— 1♂ ( DZUP) .

Redescription. Male. Body length: 24.0– 35.1 mm. Body width: 13.0–20.0 mm. Color: From black to dark reddish brown ( Fig. 1 A–D). Head: Clypeus subpentagonal, 1.6 times wider than long, transversely rugose or rugopunctate, slightly arched at middle, weakly setose on sides; apex bidentate, teeth conic, reflexed; apical half constricted laterally, basal half with subparallel sides. Frontoclypeal suture with 2 strong tubercles in major male, or a weak ridge in medium and minor males. Interocular width equals 2.0 transverse eye diameters, frontal surface rugose, with few setae confined on sides, posterior area between eyes usually smooth. Eye canthus triangular with dorsoventrally scattered setae. Mouthparts: Mandibles bidentate, teeth conic. Maxillae usually with 5 teeth on galea; 2 teeth at apex (1 strong, 1 weak), 1 medial tooth (strong), 2 basal teeth (weak). Mentum subtriangular, moderately convex, with setose punctures confined to sides, basal half subparallel laterally, sometimes with a triangular pit. Pronotum: Horned, anteriorly with a small, conical tubercle followed by a posterior, wide concavity. Horns variable in length; major male ( Fig. 1A ) with 1 long horn on each side, directed forward, exceeding head length, apex bifurcated, with 2 upcurved prolongations, inner edge irregularly crenulate Medium male ( Fig. 2A ) with horns not exceeding head length, internal edge not crenulated. Minor male with or without small tubercles ( Fig. 1 C–D). Concavity broad, deep, usually U-shaped, occupying nearly 1/2 of pronotal area in major and medium males ( Fig. 1 A–B), and 1/ 4 in minor males ( Fig. 1 C–D). Pronotal surface inconspicuously punctate, with few scattered setae on anterior area. Scutellar shield: Subtriangular, smooth. Elytra: Surface nearly smooth, with fine punctures on disc, better observed under 90X magnification; lateral area below humerus with 3 or 4 incomplete rows of punctures observed under 25X magnification in major and median males, indistinct in minor males; striae usually barely marked. Legs: Protarsus thickened; tarsomere V wider than previous ones, slightly curved, with a ventro-basal tooth; inner claw enlarged, incised, parabolic on external edge; outer claw simple. Mesotibiae with 2 transverse carinae on external surface; external surface slightly concave between carinae, strongly concave just behind second carina; apex with outer spur obliquely truncated, inner spur acuminate; carinae and apex bordered by spinules. Metatibiae with spatulate inner spur. Abdomen: Ventrites I–II completelly setose, III–V with setose punctures confined to sides, VI bordered by setae on posterior margin, apex emarginated. Tergite VIII convex; surface with setae scattered laterally and anteriorly, sides rugopunctate, punctures usually absent on disc. Tergite VII with stridulatory apparatus situated on disc; stridulatory apparatus with barely marked, transverse carinae, only observed under high magnifications (90X) ( Fig. 7A ). Aedeagus: Parameres in caudal view ( Fig. 8A ), symmetric, basal half with outer sides constricted gradually toward to middle area, apical half expanded in shape of lateral “flaps”, lateral flaps with rounded outer margins, broader than basal half. Parameres in lateral view, apex downcurved ( Fig. 9A ).

Female ( Fig. 1E ). Body length: 22.5–32.0 mm. Body width: 12.0–18.0 mm. Differ from male in the following aspects: Pronotum: Simply convex, horns absent, concavity small, shallow, confined to anterior area near apex; surface almost completely punctate; anterior angles with dense, coalescent punctures, distinctly wider in size compared to punctures of concavity and disc; concavity moderately punctate; disc finely punctate ( Fig. 10A ). Legs: Inner proclaw similar to outer claw. Venter: Proventrite with 2 small furrows at lateroanterior angles ( Fig. 10E ). Abdomen: Ventrite VI triangularly shaped, not emarginate at apex. Tergite VIII apically invaginated on posterior margin; invagination about 4.4 times wider than long, weakly emarginate ( Fig. 11A ).

Geographic distribution. Brazil: Pernambuco, Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro, São Paulo, Paraná, Santa Catarina ( Fig. 12A ). Bothynus entellus has the broadest distribution of species in the group although restricted to the Brazilian Atlantic coastal rainforest. The records cited by Endrödi (1969, 1985) for B. entellus from Honduras, Panama, Peru, and Pará state (northern Brazil) are erroneous. This mistake probably happened due to confusion with the females of B. complanus or B. quadridens , which are common species in the Central America and that have similarities with B. entellus (smooth elytra). Moreover, these dubious records could be due to mislabeling.

Figures

FIGURE 1. Bothynus entellus. A, Major male, dorsal view; B, median male, dorsal view; C–D, minor males, dorsal view; E, female, dorsal view. Scale bars: 2 mm.

FIGURE 2. Bothynus stenelus. A, Major male, dorsal view; B, major male, lateral view; C, minor male, dorsal view; D, female, dorsal view; E, male protarsus, lateral view (ventral apex elongate, arrow); F, metatrochanter, ventral view (rounded apex, arrow). Scale bars: 2 mm.

FIGURE 7. Tergite VII, detail of the stridulatory area (carinae, arrow). A, Bothynus entellus; B, B. araya; C, B. lancifer. Scale bars: 2 mm.

FIGURE 8. Parameres in caudal view. A, Bothynus entellus; B, B. stenelus; C, B. tricornis; D, B. lancifer; E, B. araya; F, B. condacki (inner apical contraction, arrow). Scale bars: 2 mm.

FIGURE 9. Aedeagus in lateral view. A, Bothynus entellus; B, B. stenelus; C, B. tricornis; D, B. lancifer (lateroventral contraction, arrow); E, B. araya; F, B. condacki. Scale bars: 2 mm.

FIGURE 10. Pronotal surface of females in dorsal view. A, Bothynus entellus; B, B. stenelus; C, B. tricornis; D, B. araya. Proventrite of females in ventral view. E, Bothynus entellus (small furrows, arrow); F, B. araya (long furrow, arrow). Scale bars: 2 mm.

FIGURE 11. Tergite VIII, detail of the apical invagination (arrow). A, Bothynus entellus; B, B. stenelus; C, B. tricornis. Scale bars: 2 mm.

FIGURE 12. Distribution map of the Bothynus entellus species group. A, Bothynus entellus; B, B. stenelus; C, B. tricornis; D, B. lancifer; E, B. araya; F, B. condacki.