Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae)
Génier, François
Moretto, Philippe
Zootaxa
2017
4248
1
1
110
363WQ
Genier
Genier
[151,628,1230,1256]
Insecta
Scarabaeidae
Digitonthophagus
Animalia
Coleoptera
69
70
Arthropoda
species
namaquensis
sp. nov.
Typelocality. Farm Klipkoppies (6) ( 28.87137 S 21.30944 E), 957 m, NorthernCape, South Africa.
Description. Holotype ♂( Fig. 22). Measurements. Length 10.5 mm, width 6.0 mm. Head( Figs. 53–54). Anterior clypeal edge straight on median fifth in dorsal view; clypeofrontal carina broadly arcuate and interrupted at gena; vertex lacking median tubercle, surface with punctures fine to small, separated by one to four diameters, scabrous. Horns rather short, slightly arcuate in frontal view, gradually tapering from base to apex; posterointernal edge unmodified basally, apicointernal surface with few scabrous punctures; genal edge slightly upturned and broadly arcuate, forming a broad angle with clypeal edge. Pronotum( Fig. 70). Surface with weakly-defined granulate punctures on anterior half only, with scattered punctures on posterior half of disc; punctures scattered and small posterolaterally, with distinct minute punctures throughout; anteromedian tubercle atrophied, forming an obtuse angle in lateral view, median longitudinal sulcus rather wide and moderately deep; surface behind the eyes with a transverse oval and shallow depression, surface of anterior angles slightly concave; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles slightly upturned and simply arcuate in dorsal view; anterior hypomeral ridge broadly arcuate anteriorly, anterior hypomeral depression surface light in color. Elytra( Fig. 22). Interval 2 with one to three fine granules on apical declivity, interval 4 lacking fine granules from base to apex. Legs. Protibial apicointernal tooth enlarged, with dorsal ridge extending to apex. Aedeagus( Fig. 134). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites( Figs. 180–182). Axial sclerite arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to apex, extending straight and interrupted before apex of frontolateral peripheral sclerite apical portion, with a brush of villi on apical fourth. Frontolateral peripheral sclerite basoventral apophysis large; lacking medioventral carinae; right lateral fold produced into a rather small everted and open apically conical process with irregular apical edge located behind oblique right lateral edge of apical lobe; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous, small and obliquely projecting dorsally, interrupted well before anterior edge of apical lobe, apex set on left side in dorsal view; apical lobe truncate and slightly directed obliquely on left side, lateral edges folded ventrally, with a small additional longitudinal fold in line with subapicoventral lobe; apical villi becoming longer toward apex; subapicoventral lobe acute in lateral view, interrupted before apex of apical lobe. Variation. Measurements ( 97 ♂♂, 74 ♀♀). Length: male 7.0–12.0 mm (10.0 ± 0.9 mm), female 8.0– 11.5 mm(9.9 ± 0.8 mm). Females allotype. Cephalic outline in dorsal view as in Fig. 86; vertex with a high and broadly arcuate, transverse carina, dorsal edge subangular medially in frontal view, lateral portion gradually sloping down posteriorly; anterior pronotal tubercles well developed, external lateral edges distinctly widening toward apex in dorsal view, anterolateral surface obliquely excavated, anterosuperior edge slightly arcuate in dorsal view (Fig. 117), lateral portion of anterosuperior edge slightly upturned (Fig. 118). Anterior tibia short, with external teeth more robust. Primary type data. Holotype ♂ (CMNC): [SOUTH AFRICA: N. CAPE / Farm Klipkoppies (6) / S28.87137° E21°30.944’ [ sic] / 957 m; 12–14 March 2004/ A.L.V. Davis & C. Deschodt]; [WORLD / SCARAB. / DATABASE / WSD00029107] barcode label; [HOLOTYPE ♂ / Digitonthophagus/ namaquensis n.sp./ des. F. Génier, 2016] red card. MAP 17.Distribution of Digitonthophagus namaquensis.
Materialexamined( 100 ♂♂, 75 ♀♀), distribution (Map 17): ANGOLA: BENGUELA, Benguela( 12°35'S, 13°25'E), [no date], [illegible]— 1 ♀, 1 ♂( paratypes) ( IRSNB); NAMIBIA: ERONGO, Ameib Farm, 19 kmNW Karibib( 21°47'S, 15°38'E), 31.i–2.ii.1972, SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH); Gobabeb( 23°34'S, 15°2'E), 10.ii.1978, O. Lomhodlt— 1 ♂( paratype) ( ZMUC); Kahn River, 5 mi. N Usakos( 22°0'S, 15°36'E), 30–31.i.1972, SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH); Kuiseb Canyon( 23°18'S, 15°45'E), 22–23.i.1972, SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH); HARDAP, Farm Lovedale, N Tiras Mountains, 1600 m( 25°32'24''S, 16°23'24''E), 16–17.ii.2010, W. Schawaller— 1 ♀, 5 ♂♂( 6 paratypes) ( SMNS); Sesriem Farm, Maltahoe District( 24°11'S, 15°59'E), 19–20.i.1972, SouthernAfrica Expedition— 1 ♀( paratype) ( BMNH); Voigtgrund( 24°49'S, 17°26'E), [no date], K. Jordan— 1 ♀, 2 ♂♂( 3 paratypes) ( BMNH); Malthahöhe district, Zais6 ( 24°2'58''S, 16°9'14''E), 16.i.2000, Mann& Marais— 1 ♀( paratype) ( OUMNH); Rehoboth district, on road verge, Kambes498 ( 24°4'32''S, 16°28'21''E), 17.i.2000, Mann& Marais— 1 ♀, 3 ♂♂( 4 paratypes) ( OUMNH); KARAS, Barby Farm, 25 mi. W Helmeringhausen( 25°53'S, 16°49'E), 17–18.i.1972, SouthernAfrica Expedition— 11 ♀♀, 16 ♂♂( 27 paratypes) ( BMNH); Keetmannshoop( 26°34'S, 18°8'E), 1913, S. Hardt& G. Lotz— 3 ♀♀, 2 ♂♂( 5 paratypes) ( SMF); Mesosaurus Fossil Farm, NE Keetmanshoop, 1100 m(28°14'24''S, 18°16'48''E), 18–19.ii.2010, W. Schawaller—1 ♂ (paratype) (SMNS); Noachabeb, 27 mi. NNE Grunau( 27°23'S, 18°28'E), 10–12.i.1972, SouthernAfrica Expedition— 3 ♀♀, 5 ♂♂( 8 paratypes) ( BMNH); near Onseepkans, Orange RIVERbanks ( 28°44'S, 19°18'E), 8–10.i.1972, SouthernAfrica Expedition— 1 ♂( paratype) ( BMNH); PlateauFarm, 22 mi. E Aus( 26°38'S, 16°34'E), 14–17.i.1972, SouthernAfrica Expedition— 1 ♀, 1 ♂( paratypes) ( BMNH); Seeheim( 26°49'S, 17°47'E), 16.ii.1934, J. Ogilvie— 1 ♀( paratype) ( BMNH); KAVANGO, Mayana Lodge, Rundu District( 17°51'55''S, 19°55'48''E), 19.xii.1999, Mann& Marais— 6 ♀♀, 7 ♂♂( 13 paratypes) ( OUMNH); Mile46 Agricultutal Station, Sovo area( 18°19'14''S, 19°15'28''E), 23.iii.2002, Mann, Marais, & Kasch— 1 ♀, 1 ♂( paratypes) ( OUMNH); KHOMAS, Farm Namibgrens, Spreetshoogte, 1800 m( 23°22'12''S, 16°8'24''E), 3–6.ii.2010, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS); Solitaire( 23°53'S, 16°0'E), 22.i.1972, SouthernAfrica Expedition— 2 ♀♀, 8 ♂♂( 10 paratypes) ( BMNH); Windhoek( 22°34'12''S, 17°5'1''E), 1909, Külz— 2 ♀♀( 2 paratypes) ( SMF); KUNENE, 87 kmW Kamanjab( 19°49'S, 14°10'E), 2.iv.1977, J. Boomker— 1 ♂( paratype) ( BMNH); Khorixas( 20°22'S, 14°58'E), iii.2004, O. Aschenborn— 3 ♀♀, 2 ♂♂( 5 paratypes) (CMD); Orupembe( 18°10'S, 12°34'E), 14.iv.2005, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS); Palmwag Lodge( 19°53'10''S, 13°56'13''E), 18.iv.2005, W. Schawaller— 1 ♀, 2 ♂♂( 3 paratypes) ( SMNS); Purros( Hoaruzsib Valley) ( 18°46'S, 12°57'E), 15–16.iv.2005, W. Schawaller— 1 ♀( paratype) ( SMNS); Khorixas District, Ae-ams/ Khoanibconfluence ( 19°14'44''S, 13°20'34''E), 28.xii.1999, Mann, Marais, & Newman— 1 ♀, 1 ♂( paratypes) ( OUMNH); Hoanib rivernear Dubis( 19°13'36''S, 13°22'E), 28.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH); Palm708, adjacent to spring ( 19°52'6''S, 14°1'13''E), 4.i.2000, Mann& Marais— 1 ♂( paratype) ( OUMNH); Spaarwater711, near water hole ( 20°3'24''S, 14°2'55''E), 5.i.2000, Mann& Marais— 3 ♀♀, 1 ♂( paratypes) ( OUMNH); Opuwo District, Lower Hoanib( 19°23'7''S, 13°4'11''E), 30.xii.1999, Mann& Marais— 1 ♀, 1 ♂( paratypes) ( OUMNH); OMUSATI, Narawanduroad, Etosha National Park( 18°51'4''S, 15°32'55''E), 24–25.xii.1999, Mann& Marais— 1 ♀( paratype) ( OUMNH); OSHIKOTO, Mushara area, sandy road ( 18°36'24''S, 16°53'23''E), 21.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH); Mushara Waterhole( 18°35'19''S, 16°55'11''E), 23.xii.1999, Mann& Marais— 1 ♂( paratype) ( OUMNH); OTJOZONDJUPA, Epupa Falls, 660 m( 17°0'3''S, 13°14'36''E), 11–12.iv.2005, W. Schawaller— 3 ♀♀, 4 ♂♂( 7 paratypes) ( SMNS); SOUTH AFRICA: EASTERNCAPE, Willowmore( 33°18'S, 23°29'E), ii.1916, Dr. Brauns— 1 ♀, 1 ♂( paratypes) ( NMPC); NORTHERNCAPE, Farm Klipkoppies, 957 m( 28°52'16.93''S, 21°18'33.98''E), 12–14.iii.2004, A.L.V. Davis & C. Deschodt— 11 ♀♀, 12 ♂♂( holotype, allotype, 21 paratypes) ( BDGC, CMNC, FGIC, OUMNH, PMOC); Groblershoop, Farm Brulpan( 28°53'56''S, 21°46'49''E), 23–26.iii.2001, A. Davis— 2 ♂♂( 2 paratypes) ( BMNH); Keograbie Farm(1), 1038 m( 29°4'33.6''S, 21°46'5.63''E), 6–8.iii.2004, A.L.V. Davis & C. Deschodt— 8 ♀♀, 7 ♂♂( 15 paratypes) ( FGIC, PMOC); Rooiplato( 31°26'S, 19°24'E), 14.i.1980, [anonymous]—1 ♂ (paratype) (BMNH); Upington( 28°27'S, 21°15'E), 7.xii.1933, L. Ogilvie— 1 ♀( paratype) ( BMNH).
Etymology. Namaquensis(from Namaqualand) a Latin adjective pertaining to the distribution of this species. Natural history. Davis & Scholtz (2004), collected this species at the Brulpan Farm near Groblershoop, in Bushmanland Nama Karoo, in the south-west arid late summer rainfall region of NorthernCape. Specimens were collected in cow dung baited traps and the species is recorded in this work as a soil generalist. The current data indicates a Namibian savanna woodlands and a Nama karoo centered distribution. Some specimens collected in cow and elephant dung. Asingle specimen collected under old elephant dung. This species is attracted to mercury vapor and ultraviolet light traps.
1455384277
[151,1278,1373,1398]
South Africa
957
-28.87137
Northern Cape
1
21.30944
69
70
1
Northern Cape
holotype
1455384287
[199,1206,1506,1531]
Angola
Material
70
71
175
75
100
BENGUELA
1455384279
IRSNB
Angola
-12.583333
Benguela
1293
13.416667
70
71
2
1
1
Benguela
paratype
1455384286
1972-01-31
1972-02-02
1972-01-31
Ameib Farm
Namibia
-21.783333
Karibib
1262
15.633333
70
71
1
ERONGO
paratype
1455384283
[666,1317,1578,1603]
BMNH
Namibia
Southern
70
71
1
1
Southern
paratype
1455384291
1978-02-10
ZMUC
Namibia
-23.566668
Gobabeb
1255
15.033334
70
71
1
1
Southern
paratype
1455384295
1972-01-30
1972-01-31
1972-01-30
N Usakos
Namibia
-22.0
River
1262
15.6
70
71
1
River
paratype
1455384292
1972-01-22
1972-01-23
1972-01-22
BMNH
Kuiseb Canyon
Namibia
-23.3
Southern
1256
15.75
70
71
2
1
1
Southern
paratype
1455384273
[428,1118,1686,1711]
BMNH
Namibia
Southern
70
71
2
1
1
Southern
paratype
1455384275
2010-02-16
2010-02-17
2010-02-16
SMNS
Farm Lovedale, N
Namibia
1600
-25.539999
Tiras Mountains
21
16.39
70
71
12
1
5
HARDAP
paratype
1455384284
1972-01-19
1972-01-20
1972-01-19
BMNH
Namibia
-24.183332
Maltahoe District
1252
15.983334
Sesriem Farm
70
71
1
1
Southern
paratype
1455384294
[397,1432,1794,1819]
BMNH
Namibia
-24.816668
Voigtgrund
1249
17.433332
70
71
6
1
2
Southern
paratype
1455384297
[151,1432,1830,1855]
2000-01-16
OUMNH
Namibia
Malthahohe district
-24.049444
Mann
21
16.153889
Zais
70
71
1
1
Southern
paratype
1455384280
2000-01-17
OUMNH
Namibia
Rehoboth district
-24.075556
Mann
21
16.4725
Kambes
70
71
8
1
3
Southern
paratype
1455384281
[426,1431,1902,1927]
1972-01-17
1972-01-18
1972-01-17
Barby Farm & W Helmeringhausen
Namibia
-25.883333
KARAS
1244
16.816668
70
71
1
KARAS
paratype
1455384276
[151,935,1938,1963]
BMNH
Namibia
Southern
70
71
54
11
16
Southern
paratype
1455384290
SMF
S. Hardt
Namibia
-26.566668
Keetmannshoop
1241
18.133333
70
71
10
3
2
Southern
paratype
1455384293
1972-01-10
1972-01-12
1972-01-10
BMNH
Namibia
71
72
-27.383333
Grunau
1237
18.466667
Noachabeb
70
71
16
3
5
Southern
paratype
1455384282
1972-01-08
1972-01-10
1972-01-08
Onseepkans
Namibia
-28.733334
RIVER
1230
19.3
71
72
1
RIVER
paratype
1455384278
BMNH
Namibia
Southern
71
72
1
1
Southern
paratype
1455384288
[270,977,223,248]
1972-01-14
1972-01-17
1972-01-14
E Aus
Namibia
-26.633333
Plateau
1241
16.566668
71
72
1
Plateau
paratype
1455384289
BMNH
Namibia
Southern
71
72
2
1
1
Southern
paratype
1455384285
[404,1282,260,285]
1934-02-16
BMNH
Namibia
-26.816668
Seeheim
1240
17.783333
71
72
1
1
Southern
paratype
1455384311
1999-12-19
OUMNH
Namibia
Mayana Lodge
-17.865278
Mann
21
19.93
Rundu District
71
72
26
6
7
KAVANGO
paratype
1455384300
2002-03-23
OUMNH
Namibia
Mile
-18.320557
Sovo area
21
19.257778
Agricultutal Station
71
72
2
1
1
KAVANGO
paratype
1455384305
2010-02-03
2010-02-06
2010-02-03
SMNS
Namibia
1800
-23.37
Spreetshoogte
21
16.14
Farm Namibgrens
71
72
6
1
2
KHOMAS
paratype
1455384308
1972-01-22
BMNH
Namibia
-23.883333
Solitaire
1254
16.0
71
72
20
2
8
Southern
paratype
1455384304
SMF
Namibia
-22.570002
Windhoek
21
17.083612
71
72
4
2
Southern
paratype
1455384318
1977-04-02
BMNH
Namibia
-19.816668
Kamanjab
1270
14.166667
71
72
1
1
KUNENE
paratype
1455384314
2004-03
2005-04-14
2004-03
SMNS
Namibia
-18.166666
Orupembe
1276
12.566667
Khorixas
71
72
16
4
4
KUNENE
paratype
1455384312
[151,1351,583,608]
2005-04-18
SMNS
Namibia
-19.88611
Palmwag Lodge
21
13.936945
71
72
6
1
2
KUNENE
paratype
1455384313
2005-04-15
2005-04-16
2005-04-15
SMNS
Namibia
-18.766666
Hoaruzsib Valley
1274
12.95
Purros
71
72
1
1
KUNENE
paratype
1455384299
1999-12-28
OUMNH
Namibia
Khorixas District
-19.245556
Khoanib
21
13.342777
Ae-ams
71
72
2
1
1
KUNENE
paratype
1455384321
1999-12-28
OUMNH
Namibia
Hoanib river
-19.226667
Mann
873
13.366667
Dubis
71
72
1
1
KUNENE
paratype
1455384315
2000-01-04
OUMNH
Namibia
-19.868334
Mann
21
14.020278
Palm
71
72
1
1
KUNENE
paratype
1455384303
2000-01-05
OUMNH
Namibia
-20.056665
Mann
21
14.048612
Spaarwater
71
72
4
3
1
KUNENE
paratype
1455384316
1999-12-30
OUMNH
Namibia
Opuwo District
-19.385277
Mann
21
13.069722
Lower Hoanib
71
72
2
1
1
KUNENE
paratype
1455384306
1999-12-24
1999-12-25
1999-12-24
OUMNH
Namibia
Narawandu
-18.851112
Mann
21
15.548612
Etosha National Park
71
72
1
1
OMUSATI
paratype
1455384307
1999-12-21
OUMNH
Namibia
-18.606667
Mann
21
16.889723
Mushara area
71
72
1
1
OSHIKOTO
paratype
1455384317
1999-12-23
OUMNH
Namibia
-18.588612
Mann
21
16.919722
Mushara Waterhole
71
72
1
1
OSHIKOTO
paratype
1455384310
2005-04-11
2005-04-12
2005-04-11
SMNS
Namibia
660
-17.000834
Epupa Falls
21
13.243334
71
72
14
3
4
OTJOZONDJUPA
paratype
1455384322
1916-02
NMPC
Namibia
-33.3
Willowmore
1205
23.483334
71
72
2
1
1
EASTERN CAPE
paratype
1455384301
[265,1348,1052,1077]
2004-03-12
2004-03-14
2004-03-12
Farm Klipkoppies
Namibia
957
-28.87137
NORTHERN CAPE
1
21.309439
71
72
1
NORTHERN CAPE
paratype
1455384309
2001-03-23
2001-03-26
2001-03-23
BDGC, CMNC, FGIC, OUMNH, PMOC
Groblershoop & Farm Brulpan
Namibia
-28.89889
A
20
21.780277
71
72
44
11
12
A
holotype
1455384319
2004-03-06
2004-03-08
2004-03-06
BMNH
Keograbie Farm
Namibia
1038
-29.076
A
1
21.76823
71
72
4
2
A
holotype
1455384320
1980-01-14
FGIC, PMOC
Rooiplato
Namibia
-31.433332
A
1216
19.4
71
72
30
8
7
A
holotype
1455384302
[268,1153,1231,1256]
1933-12-07
BMNH
Namibia
-28.45
Upington
1232
21.25
71
72
1
1
A
paratype