Systematic revision of the Afrotropical genus BolbaOEer Vulcano, Martinez and Pereira (Coleoptera: Scarabaeoidea: Bolboceratidae: Bolboceratinae), with descriptions of eight new species
Author
Gussmann, S. M. V.
Author
Scholtz, C. H.
text
Journal of Natural History
2001
2001-07-31
35
7
1013
1084
http://www.tandfonline.com/doi/abs/10.1080/002229301300323910
journal article
10.1080/002229301300323910
1464-5262
5277129
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
FIG. 19.
BolbaOEer princeps
male: abdomen, posterior view (a), posterior part of abdomen, ventral view (b), aedeagus, dorsal (c), ventral (d), lateral view (e);
B. princeps
female: abdomen, posterior view (f);
B. gigas
male: posterior part of abdomen, ventral view (g), aedeagus, dorsal (h), ventral (i), lateral view (j);
B. gigas
female: abdomen, posterior view (k).
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
.
FIG. 20. Distributional records:
BolbaOEer princeps
(
E
).
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.