Bolbaoeer bremeri, Nikolajev, 1979

Gussmann, S. M. V. & Scholtz, C. H., 2001, Systematic revision of the Afrotropical genus BolbaOEer Vulcano, Martinez and Pereira (Coleoptera: Scarabaeoidea: Bolboceratidae: Bolboceratinae), with descriptions of eight new species, Journal of Natural History 35 (7), pp. 1013-1084 : 1039-1041

BolbaOEer bremeri Nikolajev

(®gures 3a±c, 7a±e, 8a±c, h, 11)

BolbaOEer bremeri Nikolajev, 1982: 35 , ®gures 1, 2, 15±17; Krikken, 1984: 39.

Description male

Body length 20.5±27.0 mm (15 specimens). Colour light brown to dark brown, head and pronotum rarely darker than elytra. Outer margin of mandible very feebly sinuate. Clypeus with bituberculate, feebly arcuate or feebly bisinuate transverse posterior carina; position of tubercles variable, from in line with antennal insertions to in line with clypeo-frontal transition; length of posterior carina variable, from slightly shorter to slightly longer than anterior carina. Frons feebly depressed posterior to posterior clypeal carina. Antennal club with glabrous area of basal segment more than one-third but less than one-half of exposed surface. Pronotum with anterior margin medially feebly raised, in dorsal view arcuate and not projecting over head; with four protrusions along posterior margin (®gure 3a, b); in major males outer horns distinctly longer and more strongly inward curved than inner horns (®gure 3a); in minor males all four horns much reduced in size, less curved, more or less equal in length and more approximated (®gure 3b); disc with ®ne, wellspaced punctures, these increasingly interspersed by some medium-sized but mostly large punctures, often conūent towards sides and sometimes along anterior margin of pronotum. Scutellum moderately punctate. Elytron with strial punctures separated by three and a half to four and a half puncture diameters; intervals with approximately four to ®ve punctures between two striae. Protibia ®ve-dentate. Protibial spur longer than ®fth tarsomere, apex shaped as in ®gure 7a. Metatrochanter of unmodi- ®ed shape, entire surface with moderately long and fairly dense setation. Metafemur in ventral view in posterior third with line of densely spaced short and moderately long setae; with fairly dense, long setation in anterior third; with very dense, short setation in basal area and along posterior margin; remaining areas asetose. Metatibia in lateral view with subapical carina bilobed (as in ®gure 1f); with outer spur hooked (®gure 7b). Underside with posterior margin of sternite 4 medially produced, apex of process truncate but medially with very small triangular incision (®gure 7d); sternite 5 medially almost completely hidden by medially produced sternite 4 (®gure 7d); sternite 6 postero-medially deeply divided, with elongate, apically rounded process on each side of division (®gure 7d, e); pygidium with feebly bisinuate apex (®gure 7c). Genitalia with aedeagus as in ®gure 8a±c; genital capsule apically with very long setation.

Description female

Body length 22.3±26.7 mm (13 specimens). Colour as in male. Outer margin of mandible rounded. Frons with strongly raised bituberculate, transverse carina between eye-canthi; carina straight and shorter than anterior clypeal carina. Antennal club as in male. Pronotum with anterior margin as in male; with transverse, feebly carinate swelling at median portion of disc; carina bisinuate in frontal view (®gure 3c); swelling with ®ne, well-spaced punctures, towards anterior margin interspersed with some large punctures; with large, mostly conūent punctures lateral and posterior to carinate swelling. Scutellum and elytron as in male. Protibia as in male. Protibial spur longer than ®fth tarsomere, acuminate. Metatrochanter of unmodi®ed shape; surface in basal third and along posterior margin with moderately long and fairly dense setation, remaining surface asetose. Metafemur in ventral view in posterior third with line of densely spaced long setae; with fairly dense, long setation in anterior third, in basal area and along posterior margin; remaining areas asetose. Metatibia as in male but outer spur unmodi®ed. Underside with posterior margin of sternite 3 medially bisinuate (®gure 8h); posterior margin of sternite 4 medially with small, strongly sclerotized, densely setose process (®gure 8h); sternite 5 unmodi®ed; sternite 6 hidden by sternite 5, unmodi®ed; pygidium unmodi®ed.

Distribution (®gure 11). West Africa, from Senegal to the easternmost distributional record from the Sudan.

Type material. HOLOTYPE l[diss.]:` Sudan, Prov. Darfur, El Geneina [13.29N 22.28E], 7.8.1977, H.J.Bremer leg. ad lucem’ /` Holotypus, BolbaOEer bremeri Nikolaijev 1979 ’ [printed on red label] ( MRAC); 4 PARATYPES: 2mm ( GNCK), 1m ( MRAC), same data as holotype except `Paratypus, BolbaOEer bremeri Nikolaijev 1979 ’; 1m, same data except`... 12.8.1977 ...’ ( MRAC) .

Additional material examined (25 specimens). Burkina Faso : ALLOTYPE m, Bolboceras splendidum Petrovitz ,` Allotypus m’ [printed on pink label] /`Coll. Mus. Tervuren, Haute Volta: Bobo-Dioulasso [11.12N 04.18W], vii. / viii.1964, R.SiOEointe’ /`Allotypus’ [printed on red label] /` Bolboceras splendidum n.sp. Petrovitz’ [printed on red label] ( MRAC) ; 1m, same locality label ( MRAC) ; 1, Haut-Dahomey [Benin], Gourma et Mossi [two regions in Burkina Faso not Benin, S and SE of Ougadougou], 1910, A. Chevalier ( MNHN) ; 1, Haute Volta, Itua [not traced], 1881, Coll. A. Argod ( MNHN) . Central African Republic: 1 l[diss.], Rep. Centrafricaine, Hte Sangha [04.30N 16.00E], Carnot ( ISNB) . Mali: 1, 1m, Bore [BoreÂ, 15.08N 03.29W], R. Demange ( MNHN) ; 1, Mourdiah , 14.28N 07.28W, 3 September 1989, J. Passerini ( HAHC) . Mauritania: 1 l[diss.], Selibaby [15.10N 12.11W], 12 August 1982, don Ch. Verstraeten ( MRAC) . Niger: 1, Sadore [13.14N 02.17E], 1987, M.J. Lukefahr ( HAHC) . Nigeria: 3 ll[diss.], 2mm, Kano Dist. [ca 12.00N 09.00E], July 1954, W.E.S. Merrett, BM. 1962±270. ( BMNH) ; 3 ll[diss.], 4mm, same data but: June 1954 ( BMNH) ; 1 l[diss.], 1m, same data ( TMSA) ; 1, Kano, 26 July 1921, Crottin ( TMSA) .

Comments. The holotype of B. bremeri Ðan intermediate maleÐand two of the four female paratypes were kindly sent to us from GNCK and are now, by mutual agreement, deposited in MRAC.

Males of B. bremeri are easily distinguished from males of the closely related B. splendidus by the presence of four pronotal protrusions (versus two), the diOEerent modi®cations on the abdominal sternites 4, 6 and the pygidium (compare ®gure 7c±e with 7b, f, g) as well as the shape of the aedeagus which is characterized by elongate, slender parameres with broadly rounded apices (versus slender parameres distinctly narrowing towards apices). Males of both species can immediately be separated from males of all other congeneric species by the outer metatibial spur which is hooked instead of feebly arcuate. A slightly modi®ed shape of the apex of the protibial spur is shared by male B. bremeri and male B. splendidus but a similar modi®ed shape is also found in male B. dudleyi . Nikolajev (1982) stated that the curved tip of the protibial spur is blunt in B. bremeri males but acuminate in B. splendidus males. We found that in both species intraspeci®c variation exists and that in most specimens the tip of the spur is shaped as in ®gure 7a but may be more blunt or more acuminate in other specimens.

The small, strongly sclerotized process at the posterior margin of sternite 4 is a unique character in females of B. bremeri and absent in females of all other species. Sternites 5 and 6 and the pygidium are unmodi®ed in females of B. bremeri .