Bolbaoeer princeps, (Kolbe)

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 : 1060-1065

BolbaOEer princeps (Kolbe)

(®gures 1d±f, 4a±f, 19a±f, 20)

Bolboceras princeps Kolbe, 1894: 167 (Descr. male); Boucomont, 1902: 5; 1912: 12; Paulian, 1941: 24; Ferreira, 1965: 1226 (Cat.); Petrovitz, 1969: 318, ®gure 6 (aedeagus, nec Kolbe).

BolbaOEer princeps (Kolbe) : Nikolajev, 1982: 39, ®gures 7, 12, 20; Krikken, 1984: 39.

Bolboceras atavus Kolbe, 1894: 167 (Descr. females); Boucomont, 1902: 4; 1910 (1911): 347 (syn. of Bolbaceras princeps, Syn. ); 1912: 12 (syn. of Bolboceras princeps ); PeÂringuey, 1908: 646 ( Bolboceras atavus ); Paulian, 1941: 24 (syn. of Bolboceras princeps ); Ferreira, 1965: 1226 (syn. of Bolboceras princeps, Cat. ).

Bolboceras panza PeÂringuey, 1896: 159 ; 1901: 480 (Redescr.); Felsche, 1901: 154 (syn. of Bolboceras atavus, Syn. ); Boucomont, 1902: 5; PeÂringuey, 1908: 646 (syn. of Bolboceras atavus ); Boucomont, 1912: 12 (syn. of B. princeps ); Paulian, 1941: 24 (syn. of Bolboceras princeps ); Ferreira, 1965: 1226 (syn. of Bolboceras princeps, Cat. ); Nikolajev, 1982: 35 (incertae sedis).

Description male

Body length 20.9±25.4 mm (116 specimens). Colour light to dark brown. Outer margin of mandible feebly sinuate. Clypeus with bituberculate, feebly arcuate transverse posterior carina; position of tubercles variable, from in line with antennal insertions to in line with clypeo-frontal transition; posterior carina same length as but mostly shorter than anterior carina. Frons feebly concave behind posterior clypeal carina. Antennal club with glabrous area of basal segment about one-half of exposed surface. Pronotum with anterior margin medially raised, in dorsal view usually projecting over frons and with feebly to distinctly bisinuate curvature; with three to ®ve protrusions along posterior margin (®gure 4a±e): major males with pair of long, inward curved outer horns and transverse, non-tuberculat e swellings between horns and one median horn (®gure 4d); or major males with one pair of long, inward curved outer horns, an inner pair of smaller horns and with one additional median horn (®gure 4a); intermediate males with reduced outer and inner horns and reduced median horn (®gure 4b, e); minor males with ®ve tubercles of equal size (®gure 4c); disc with small and medium-sized punctures of variable density, these increasingly interspersed by large, often conūent punctures towards sides and along anterior margin of pronotum. Scutellum with surface coarsely punctate. Elytron with strial punctures separated by three to ®ve puncture diameters; intervals with approximately ®ve to six punctures between two striae. Protibia ®ve-, rarely six-dentate. Protibial spur longer than ®fth tarsomere, acuminate. Metatrochanter of unmodi®ed shape; with moderately long setae on entire surface, very dense in apical area, dense in other areas. Metafemur in ventral view in posterior part with line of densely spaced moderately long and long setae; with very dense, short and moderately long setae in basal third along posterior margin; with fairly dense, long setae in anterior third; remaining areas with sparse, moderately long setation. Metatibia in lateral view with subapical carina bilobed; with unmodi®ed spurs (®gure 1f). Underside with sternite 4 unmodi®ed; posterior margin of sternite 5 medially very feebly concave (®gure 19b); posterior margin of sternite 6 medially deeply divided, the two lateral lobes elongate (®gure 19a, b), their apices densely setose and in ventral view angulate (®gure 19b); apex of pygidium roundedly emarginate (®gure 19a). Genitalia

with aedeagus as in ®gure 19c±e; genital capsule apically with moderately long setation.

Description female

Body length 21.1±26.2 mm (109 specimens). Colour as in male. Outer margin of mandible rounded. Clypeus unarmed. Frons with strongly raised bituberculate, transverse carina, positioned between eye-canthi; carina straight and variable in length, from slightly to distinctly shorter than anterior clypeal carina. Antennal club as in male. Pronotum with anterior margin medially feebly raised, in dorsal view arcuate; with transverse, distinctly carinate and strongly developed swelling at median portion of disc; carina in dorsal view straight, in frontal view more or less straight and laterally arcuate (®gure 4f); swelling with a few ®ne punctures, interspersed with large punctures; with large, mostly conūent punctures anterior, lateral and posterior to carinate swelling. Scutellum and elytron as in male. Protibia and protibial spur as in male. Metatrochanter of unmodi®ed shape; with fairly dense and moderately long setation in basal third and along posterior margin; remaining surface asetose. Metafemur in ventral view in posterior third with line of densely spaced, moderately long and long setae; with fairly dense and long setae in anterior third and apical area; with fairly dense, moderately long setae along posterior margin; remaining areas with few isolated setae. Metatibia as in male. Underside with sternites 4, 5 and 6 unmodi®ed; apex of pygidium very feebly roundedly emarginate (®gure 19f).

Distribution (®gure 20). BolbaOEer princeps is by far the most widely distributed species within the genus. It occurs in most parts of subsaharan Africa except in arid regions. The lack of distributional records from countries such as Angola, Botswana, Mozambique, Uganda, Sudan, Central African Republic, Cameroon etc. is obvious but quite clearly due to historically poor collecting in those countries.

Type material examined. HOLOTYPE l[diss.], Bolboceras princeps Kolbe :`Tanga, O. Kickel V.’ /` Bolboceras princeps n. sp. Kolbe’ [handwritten by Kolbe] /`5 atavus m.’ [handwritten by Kolbe] /`66128’ /` princeps Kolbe *’ [handwritten by unknown curator] /`Type’ [black print on red label] (ZMHB); LECTOTYPE m (here designated), Bolboceras atavus Kolbe :`N.Nyassa-See, Conde-Gebiet, Linnaea V.’ /` Bolboceras atavus n.sp. Kolbe’ [handwritten by Kolbe] /`66129’ /`Type’ [printed on red label] (ZMHB); PARALECTOTYP E m,`N.Nyassa-See, Conde-Gebiet, Linnaea V.’ /`Type’ [printed on red label] (ZMHB); LECTOTYPE l(here designated), Bolboceras panza PeÂringuey :`D’Urban [Durban], Bowker’ /` Bolboceras panza ltyp. Py’ [handwritten by PeÂringuey] /`Type SAM / Ent 5084’ [printed and handwritten on red label] /`SAM’ [printed on white label] (SAMC); PARALECTOTYPE m,`D’Urban, Bowker, 1887’ /` Bolboceras panza typ. Py’ [handwritten by PeÂringuey] /`Type SAM / Ent 5084’ [printed and handwritten on red label] /`SAM’ [printed on white label] (SAMC).

Additional material examined (218 specimens from AMMM, AGCZ, AKCS, BMNH, CCIT, COCS, DMSA, EBCI, HAHC, HNHM, ISNB, MCST, MNHN, MMCM, MSCJ, MKCS, MRAC, NSCJ, RMCI, SAMC, SANC, TMSA, UPSA, ZMHB). Burundi: 1m, Kitega [03.26S 29.56E]; 1, Usumbura [Bujumbura, 03.22S 29.19E]; 1m, Usumbura (plaine); 1m, Uzumbura . Cameroon: 1m, Kamerun int., Mao Rei [not traced] . Chad: 1, Bekao [BeÂkao, 07.54N 16.02E]. D.R.C.: 2 ll, Kivou [Kivu Prov.], Ouvira [Uvira, 03.24S 29.05E]; 1 l[diss.], 1m, Lomami , Mwene-Ditu [07.00S 23.27E]; 1 l[diss.], Lulua , Sandoa [09.41S 22.52E]; 1, 1m, Katanga , Kafakumba [09.41S 23.44E]; 2 ll, 1m [diss. 26 January 1995 by Riaan Stals but no genitalia found], 3mm, Katanga, Elisabethville; 1 l[diss.], Katanga, Lubumbashi; 1m, LeÂopoldville; 1m, Kapanga [06.48S 22.32E]; 1m, Kiambi [07.20S 28.01E]; 1m, Nioka [many places by this name]; 2mm, Congo belge; allotype m, Bolboceras coriaceum Petrovitz :`Coll. Mus. Tervuren, Katanga: Kipopo (E’ville), 3.xii.1961, Don R .

MareÂchal’ ( MRAC); 12 paratypes, Bolboceras coriaceum Petrovitz : 3mm,`MuseÂe du Congo, Elisabethville, 1935, Dr. Richard’ ( MRAC); 1m,` MuseÂe du Congo, Kivu: Luvungi [02.52S 29.02E], iii.1935 , Wylocke’ ( NHMG); 1m,`MuseÂe du Congo, Elisabethville , 2.iii.1924 , Ch.Seydel’ ( NHMG); 1, same data except`... 26.i.1924 ...’ ( MRAC); 1m,`MuseÂe du Congo, Lulua : Kapanga [06.48S 22.32E] xii.1933 , F.G.

Overlaet’ ( MRAC); 1m,`MuseÂe du Congo, Elisabethville, xii.1932, Mlle Kerkvoorde’ ( MRAC); 1m,`MuseÂe du Congo, Kivu: Futwe, xii.1928, Ch. Seydel’ ( MRAC); 1m,`Coll. Mus. Tervuren, Katanga: Kipopo (E’ville), 6.xii.1961, Don R. MareÂchal’ ( MRAC); 1m, same data but`... 21.i.1962 ...’ ( MRAC); 1m,`MuseÂe du Congo, Katanga: Lubumbashi, v.1911, Buttgenbach’ ( MRAC). Ethiopia: 1, Begemdir [Bagemder Prov.], AzozoÁ [Azoza Å, 12.33N 37.26E]; 1, Sidamo [Sõ Å damo] Province, 10 km NW of Neghelli [Negele Å, ca 05.22N 39.35E]; 1, Sidamo Prov. , Kebri [Kibre] Mengist [05.54N 38.59E]ÐBitata [not traced]; 1 l[diss.], Metahara [Mataha Å ra Å, 08.53N 39.52E]; 1, Shoa Prov. , Lake Langano [ca 07.40N 38.50E]. Ghana: 1 l [diss.], Bawku [11.05N 00.11W]. Guinea: 1, Sattelberg [not traced]. Kenya: 1, Nairobi [01.17S 36.50E]; 1m, Nairobi , Ngang Forest [Ngong, 01.22S 36.39E]; 1m, Kenya ; 1m, Afrique Orient. Angl., sud du lac Rodolphe , entre le chemin de fer at le lac [not traced, too vague]. Malawi: 2 ll, 2mm, Chintheche [Stream, taken at 11.47S 34.13E], 1 December 1975, R. JocqueÂ; 1 l[partially diss.], Euthini [Eutini, 11.28S 33.25E]; 2 ll[diss.], North Senga Forest Res. , 13.43S 34.37E ; 1, Machinga Dist., Chikala Forest Reserve [ca 15.07S 35.25E]; 1, M 1541 [notes stolen but collected in the southern regions, probably in the Shire Highlands near Blantyre / Limbe± personal comment by C.O. Dudley]; 1m, Lilongwe Dist., Dzalanyama Forest Reserve [14.15S 33.25E]; 1m, Zomba [15.22S 35.22E]; 1m, Mlanje [16.02S 35.30E]; 1, 1m, Dedza [14.20S 34.24E]; 2mm, Mzimba [11.54S 33.36E]; 1, Ituri [not traced] NW of Nk [h]ata Bay [11.36S 34.18E], 21 February 1958; 2mm from`Nyassaland’ .

Namibia: 1 l[diss.], Abachaus [19.47S 16.44E] . Nigeria: 2 ll[diss.], 1m, Kano Dist.

[ 12.00N 09.00E]; 1 l[diss.], 1m, W.E.S. Merrett, BM. 1962-270; 1 l[diss.], Yola [many localities by that name]; 1 l[diss.], 2mm, Samaru [09.45N 08.23E or 11.10N 07.38E]; 1m, Zaria, Samaru. Rwanda: 1, Gabiro [01.33S 30.25E]. South Africa: 3 ll[diss.], 2mm, Pretoria [25.45S 28.12E]; 1 l[diss.], Krugersdorp [26.06S 27.46E]; 1, 2mm, Transvaal, Lydenburg [25.06S 30.27E]; 1m, Mpumalanga, Kopeeskraal, Lydenburg , 25.07S 30.12E ; 1m, Lydenburg, Madoki [not traced]; 1 l[diss.], Waterval Onder [25.39S 30.20E]; 1 l[diss.], 7 ll, 1m [diss.], 3mm, Transvaal, Barberton [25.47S 31.03E]; 1m, Pilgrims Rest, Selati [24.03S 30.38E]; 1m, Branddraai [24.38S 30.37E]; 2 ll[diss.], White River [Whitrivier, 25.20S 31.01E]; 1 l[diss.], Natal , Krantzkloof [too vague, several localities by that name]; 1, Natal , Eshove , 28.54S 31.28E ; 1 l [diss.], Modderpoort [Ladybrand, O., 29.07S 27.28E]; 1 l[diss.], Greytown [in Kwazulu-Natal Prov., 29.04S 30.35E]; 1 l[diss.], Maritzburg [Pietermaritzburg, 29.36S 30.24E]; 1 l[diss.], 2 ll, 1m, Natal, Malvern [29.53S 30.56E]; 3 ll, 1m, Durban [29.52S 31.00E]; 1m, Umkomas N. [Umkomaas, 30.13S 30.48E]; 1 l[diss.], 1 l[partially diss.], 1, 3mm from untraced / vague localities. Tanzania: 1 l[diss.], Chunya District, Chunya; 1 l[diss.], Ndala Mission , 04.45S 33.15E ; 1, Wanga [05.12S 39.04E]; 1 l[diss., but genitalia missing from card], Mts Uluguru, Morogoro Campus Fac. Agric. [ca 06.49S 37.40E]; 1, Morogoro [distr., 07.43S 35.52E]; 1, Manow [Manow mission, 09.15S 33.48E]; 1, near Mikumi [Mikumi town, 07.22S 37.00E]; 1, Mikumi [Mikusi] N.P. [too vague]; 1m, near Babati [04.12S 35.45E]; 1, Tanga [05.40S 39.04E]; 1m, (Iringa) Ma ®nga [08.18S 35.13E], December 1997 ; 1m, (Iringa) Sadani [08.14S 35.01E], December 1997 ; 1 l[diss.], Tanganyika; 1, Ost-Afrika, Jpiama b. Langenburg [not traced]; 1 l[diss.], Massai. Zambia: 1 l [diss.], Fwambo [08.52S 31.32E]; 1 l[diss.], El Dorado [farm, 12.30S 28.18E]; 1 l[diss.], 2mm, Broken Hill [Kabwe, 14.27S 28.27E]; 1 l[diss.], Congo-Zambesi Watershed [not traced, too vague]; 3 ll, 3mm, S. Prov., Choma Mansai Farm, 16.45S 27.00E ; 1m, Kashitu [13.45S 28.40E]; 3 ll, 6mm, Kasama [10.10S 31.11E]; 8 ll, 6mm, Lusaka, Kafue [15.44S 28.10E]; 1m, Monze [16.16S 27.28E]; 1, Bhano [not traced] .

Zimbabwe: 1 l[diss.], Mazoe [Mazowe, 17.30S 30.59E]; 5 ll[diss], 4 ll, 9mm, Salisbury [Harare, 17.50S 31.02E]; 1 l[diss.], Macheke [18.07S 31.50E]; 1 l[diss.], Nyanga Nat. Park, Nyangombe Camp, 18.16,6S 32.43,2E; 1 l[diss.], Umtali [Mutare, 19.00S 32.40E]; 2 ll, Kadoma, 15 km S [18.26S 29.55E]; 2mm, Mashonaland, Hanyani R. [Hunjani R., 15.38S 30.40E]; 1m, Zambezi, Victoria Falls [17.55S 25.45E]; 1m, Umwinsidale [17.34S 31.12E]; 1m, Que Que [junction Sebakwe Riv., 18.46S 29.44E]; 1m, Bulawayo [20.10S 28.35E]; 1, Chipinge [20.12S 32.37E], February 1990 ; 3 ll[diss.], Mashonaland; 1m, Rhodesia; 1m, Zambezi; 1, Burnside [not traced]. Not traced/vague: 4 ll, 8mm .

Comments. BolbaOEer princeps is the most widely distributed and most commonly collected BolbaOEer species. It is the only species that extends its range as far south as the coastline of Kwazulu-Natal in South Africa. As could be expected from a species with a large distribution, certain characters vary more than in other species although some characters remain very uniform and enable a secure identi®cation of the species in both sexes. In males, pronotal armature is very variable and two types of transformations are found, randomly spread over the entire distribution range: major and intermediate males either possess ®ve tubercles / horns along the posterior pronotal margin as in ®gure 4a, b or three as in ®gure 4d, e. Only minor males consistently possess ®ve, equally sized tubercles as in ®gure 4c. On the other hand, sternal and pygidial modi®cations as well as the aedeagal shape are stable throughou t the populations. The very feebly concave posterior margin of sternite 4 and the two elongate, apically angulate lateral lobes of sternite 6 (®gure 19a, b) are invariably visible and su ce for an easy identi®cation. The small incision on the apex of the pygidium, however, is not always visible ®rstly because of the dense setation on the pygidium or because the pygidium is often retracted under the elytra.

All females of the B. princeps species-group are characterized by a general lack of modi®cations making it easy to separate them from females of other speciesgroups but making it more di cult to distinguish them from one another. DiOEerences between the females of B. princeps and B. gigas are discussed under the latter. BolbaOEer princeps females are separated from B. nikolajevi females as follows: the bituberculate, transverse carina on the frons is positioned more anteriad, i.e. right between the eye-canthi in the former while the carina is positioned at most in line with the postero-lateral genal angle but mostly posteriad to the postero-genal angle in the latter. Other diOEerences are the feeble but discernible postero-median swelling on sternite 5 and the somewhat pointed posterior margins of the otherwise unmodi- ®ed sternite 6 and pygidium in B. nikolajevi as opposed to unmodi®ed sternites 5, 6 and pygidium in B. princeps . The feeble emargination on the apex of the pygidium in females of B. princeps as illustrated in ®gure 19f is not always discernible, thus not a very reliable character state. In addition, the pygidium is very often retracted under the elytra. More obvious, if not retracted, is the broader shape of the pygidium (compare ®gure 19f with 17f) in B. princeps .

The holotype of Bolboceras princeps Kolbe represents a major male with pronotal armature as in ®gure 4a but with the median horn broken oOE. The lectotype of Bolboceras panza PeÂringuey is an intermediate male as illustrated in ®gure 4b. One male specimen was found in MNHN, labelled as follows:`Transvaal, Lydenburg, F.Wilms S.V.’ [printed] /` Bolboceras atavus Kolbe ex typ.’ [handwritten by Kolbe] /`MuseÂe de Berlin’ [printed] /`Type ’ [black print on red label] /`Museu m Paris ex Coll. R. Oberthur’ [printed on blue label]. This specimen is not considered a type because Kolbe’s description (1894) of Bolboceras atavus clearly refers to two female specimens with locality data as found on the labels of the lecto- and paralectotype of Bolboceras atavus (see above). It seems, however, that Kolbe subsequent to the descriptions of Bolboceras princeps and Bolboceras atavus discovered that the latter represents the matching sex to the former, i.e. the latter name is a synonym of the former. This is evident from a label in Kolbe’s handwriting, attached to the holotype of Bolboceras princeps and saying:`5 atavus m.’ but this was not published.