Digitonthophagus gazella (Fabricius, 1787), conservation of name in prevailing usage

(Figs. 14, 37–38, 62, 78, 101–102, 126, 156–158, 185, 188, 190; Map 9)

Scarabaeus Gazella Fabricius, 1787b: 377 (original description) Scarabaeus Dorcas Olivier, 1789: 121 (original description) proposed new synonymy Scarabaeus Gazella — Gmelin 1790: 1537 (diagnosis)

Scarabaeus gazella — Fabricius 1792: 56 (diagnosis)

Copris Dorcas — Fabricius 1798: 31 (diagnosis)

Scarabaeus Dorcas — Illiger 1800: 228 (monograph)

Scarabaeus Gazella — Illiger 1800: 228 (mentioned as synonym) Copris Dorcas — Fabricius 1801: 44 (diagnosis)

Copris Gazella — Fabricius 1801: 47 (diagnosis)

Copris Gazella — Illiger 1804: 149 (mentioned as synonym) Copris Gazella — Schönherr 1806: 48 (catalog)

Onthophagus Gazella — Dejean 1821: 53 (catalog)

Onthophagus Gazella — Sturm 1826: 178 (catalog)

Onthophagus bonasus — Latreille 1827: 282 (diagnosis, distribution, comment) Onthophagus Gazella — LePeletier & Audinet-Serville 1828: 354 (comment taxonomy) Onthopagus [sic] Gazella — Klug 1833: 163 (diagnosis)

Scarabaeus Dorcas — Klug 1833: 163 (synonymy)

Onthophagus Gazella — Klug 1835: 33 (distribution)

Onthophagus Gazella — Dejean 1836: 157 (catalog)

Onthophagus Gazella — Hope 1837: 33 (catalog)

Ontophagus gazella — Reiche 1840: 243 (comment)

Ontophagus gazella — Laporte 1840: 85 (diagnosis, distribution) Onthophagus Gazella — Sturm 1843: 107 (catalog)

Ontophagus gazella — Erichson 1843: 208 (mention)

Onthophagus Gazella — Bertoloni 1849: 415 (diagnosis, distribution) Onthophagus gazella — Fåhraeus 1857: 275 (redescription)

Onthophagus gazella — Motschulsky 1860: 153 (mention)

Onthophagus gazella — Klug 1862: 232 (comment, distribution) Onthophagus gazella — Harold 1867: 49 (comment)

Onthophagus gazella — Harold 1869: 1029 (catalog)

[Scarabaeus] dorcas — Harold 1869: 1029 (mentioned as synonym) Copris gazella — Gerstaecker 1871: 50 (synonymy)

Onthophagus gazella — Harold 1871: 119 (comment)

Scarabaeus Dorcas — Gerstaecker 1873: 130 (mentioned as synonym) Scarabaeus gazella — Gerstaecker 1873: 130 (mentioned as synonym) Onthophagus gazella — Wallengren 1881: 21 (comment)

Onthophagus gazella — Lansberge 1882: XXII (mention)

Onthophagus gazella — Reitter 1892: 183 (identification key, distribution) Onthophagus gazella — Péringuey 1892: 29 (distribution)

Onthophagus gazella — Fairmaire 1893: 10 (distribution)

Onthophagus gazella — Brancsik 1893: 223 (distribution)

Onthophagus gazella — Reitter 1893: 62 (identification key, distribution) Onthophagus gazella — Kolbe 1897a: 150 (distribution)

Onthophagus gazella — Kolbe 1897b: 83 (distribution)

Onthophagus gazella — Gahan & Arrow 1900: 24 (distribution) Onthophagus (O.) gazella — d’Orbigny 1900a: 218 (monograph) Onthophagus gazella — d’Orbigny 1900b: 299 (comment, distribution) Onthophagus gazella — Péringuey 1901: 181 (monograph)

Onthophagus (O.) gazella — d’Orbigny 1904: 304 (distribution) Onthophagus (O.) gazella — d’Orbigny 1905a: 494 (diagnosis, distribution) Onthophagus gazella — d’Orbigny 1905b: 445 (distribution) Onthophagus gazella — Kolbe 1908: 128 (distribution)

Onthophagus gazella — d’Orbigny 1909: 525 (distribution)

Onthophagus gazella — d’Orbigny 1911: 20 (distribution)

Onthophagus (O.) gazella — d’Orbigny 1913a: 249 (monograph) Onthophagus (O.) gazella — d’Orbigny 1913b: 96 (comment, distribution) Onthophagus (O.) gazella — Kolbe 1914: 297 (distribution)

Onthophagus (O.) gazella — d’Orbigny 1914: 654 (catalog)

Onthophagus (O.) gazella — d’Orbigny 1916: 11 (distribution) Onthophagus gazella — Boucomont 1923: 96 (distribution)

Onthophagus gazella — Gillet & Boucomont 1927: 138 (catalog) [Scarabaeus] dorcas — Gillet & Boucomont 1927: 138 (mentioned as synonym) Scarabaeus gazella — Arrow 1931: 230 (mentioned as synonym) Onthophagus gazella — Boucomont 1934: 26 (mentioned as synonym) Onthophagus gazella — Paulian 1936: 132 (identification key) Onthophagus gazella — Boucomont 1937: 131 (distribution) Onthophagus gazella — Balthasar 1941: 110 (distribution)

Onthophagus gazella — Müller 1942: 96 (comment, distribution) Onthophagus gazella — Gomes Alves 1944: 3 (distribution)

Onthophagus gazella — Paulian 1948: 151 (mentioned as synonym) Onthophagus (O.) gazella — Gomes Alves 1956: 94 (distribution) Onthophagus gazella — Frey 1956: 654 (distribution)

Onthophagus gazella — Ferreira 1958: 512 (faunistic)

Onthophagus gazella — Frey 1958: 73 (distribution)

Onthophagus gazella — Ferreira 1959: 84 (distribution)

Onthophagus gazella — Frey 1961: 90 (distribution, ecology) Onthophagus gazella — Petrovitz 1961: 102 (distribution)

Onthophagus gazella — Ferreira 1962: 153 (faunistic)

Onthophagus (Digitonthophagus) gazella — Balthasar 1963: 365 (monograph) [Scarabaeus] dorcas — Balthasar 1963: 365 (mentioned as synonym) Onthophagus gazella — Ferreira 1967: 1169 (catalog)

Onthophagus dorcas — Ferreira 1968: 593 (mentioned as synonym) Onthophagus gazella — Ferreira 1968: 593 (monograph)

Onthophagus (Digitonthophagus) gazella — Balthasar 1969: 68 (faunistic) Onthophagus gazella — Bornemissza 1970: 31 (natural history)

Onthophagus (O.) gazella — Endrödi 1971: 294 (faunistic)

Onthophagus (Digitonthophagus) gazella — Cambefort 1972: 246 (distribution, ecology) Onthophagus (Digitonthophagus) gazella — Ferreira 1972: 780 (catalog) Onthophagus dorcas — Ferreira 1972: 780 (mentioned as synonym) Onthophagus gazella — Balthasar 1972: 22 (faunistic)

Onthophagus gazella — Endrödi 1973a: 125 (faunistic)

Onthophagus (O.) gazella — Endrödi 1973b: 198 (distribution)

Onthophagus gazella — Endrödi 1974: 3 (faunistic)

Onthophagus gazella — Blume & Aga 1975: 735 (natural history)

Onthophagus gazella — Endrödi 1976: 156 (distribution)

Onthophagus gazella — Alfieri 1976: 205 (faunistic)

Onthophagus gazella — Bornemissza 1976: 7 (natural history)

Onthophagus gazella — Blume & Aga 1978: 190 (distribution, ecology) Onthophagus gazella — Rougon & Rougon 1980: 381 (natural history, description of larvae) Digitonthophagus gazella — Zunino 1981: 414 (new combination, faunistic) Onthophagus gazella — Rougon & Rougon 1981: 6 (description of larvae) Onthophagus gazella — Rougon & Rougon 1983: 499 (natural history) Onthophagus gazella — Fincher et al. 1986: 2 (natural history)

Onthophagus gazella — Paulian 1986: 107 (mentioned as synonym) Onthophagus gazella — Gutierrez et al 1988: 39 (natural history)

Digitonthophagus gazella — Zunino & Halffter 1988: 17 (distribution) Digitonthophagus gazella — Kohlmann 1994: 35 (distribution)

Onthophagus gazella — Ripa et al. 1995: 438 (distribution)

Digitonthophagus gazella — Barbero et al. 1998: 242 (distribution) Onthophagus (Digitonthophagus) gazella — Kim & Lumaret, 1989: 336 (description of larvae) Digitonthophagus gazella — Deloya 2000: 125 (faunistic)

Onthophagus gazella — MacRae & Penn 2001: 49 (distribution)

Digitonthophagus gazella — Noriega 2002: 213 (distribution)

Digitonthophagus gazella —Barbero & Palestrini 2003: 316 (distribution) Digitonthophagus gazella —Barbero et al. 2003: 333 (phylogeny)

Digitonthophagus gazella — Cambefort & Bordat 2003: 560 (distribution, ecology) Digitonthophagus gazella — Noriega et al. 2006: 379 (distribution) Digitonthophagus gazella — Moretto & Bordat 2007: 122 (distribution, ecology) Digitonthophagus gazella — Monaghan et al. 2007: 678 (phylogeny) Digitonthophagus gazella — Ivie & Philips 2008: 10 (distribution)

Digitonthophagus gazella —Cabral-de-Mello et al. 2008: 1244 (karyotype) Onthophagus gazella — Wirta et al. 2008: 1079 (phylogeny)

Digitonthophagus gazella — Noriega et al. 2010: 451 (distribution) Digitonthophagus gazella — Noriega et al. 2011: 35 (distribution)

Digitonthophagus gazella — Tarasov & Solodovnikov 2011: 9 (phylogeny) Digitonthophagus gazella — Whipple et al. 2012: 45 (DNA analysis) Digitonthophagus gazella — Mlambo et al. 2015: 318 (phylogeny)

Digitonthophagus gazella — Roggero et al. 2016a: [2] (phylogeny)

Digitonthophagus gazella — Breeschoten et al. 2016: 90 (phylogeny) Digitonthophagus gazella — Génier & Davis 2017: 497 (comment taxonomy, distribution) Scarabaeus dorcas — Génier & Davis 2017: 498 (comment taxonomy)

Type locality. Unspecified.

Diagnosis. Pronotum rather glossy (Fig. 62); FLP sclerite subapicodorsal lobe short anteriorly (Figs. 156– 157); right lateral fold produced into a longitudinal spoon-shape process with apical portion open ventrally, ventral edge bordered with a conspicuous brush of villi (Fig. 185 A), lacking well-sclerotized projection on apicoventral edge (Fig. 185 B); axial sclerite shorter than subaxial sclerite, acute apically (Fig. 156); subaxial sclerite moderately long and robust (Fig. 153).

Description. Male specimen from Central Province, Zambia (Fig. 14). Measurements. Length 12.5 mm, width 7.0 mm. Head (Figs. 37–38). 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. Horns rather short, slightly divergent in frontal view, parallel sided on basal four-fifths and abruptly tapering externally and narrower on apical sixth, posterointernal edge produced into low, angular projection basally, apicointernal surface lacking granules, with few scabrous punctures; genal edge slightly upturned and feebly angulate on anterior third, forming a broad angle with clypeal edge. Pronotum (Fig. 62). Surface with granulate punctures extending on posterior half, with distinct umbilicate punctures on posterior half of disc; punctures smaller posterolaterally, with distinct, minute punctures throughout. Anteromedian tubercle atrophied, simply round in lateral view, median longitudinal sulcus narrow and moderately deep; surface behind the eyes with a simple, round, shallow depression, surface of anterior angles flat; anterior half of lateral edge arcuate in dorsal and lateral view; posterior angles slightly upturned and simply arcuate in dorsal view; anterior hypomeral ridge arcuate anteriorly, anterior hypomeral depression surface slightly darker in color medially. Elytra (Fig. 14). Interval 2 with few scattered 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. 126). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites (Figs. 156–158). Axial sclerite broadly arcuate, with apex acute. Subaxial sclerite, gradually tapering from base to apex, slightly angled on apical third, extending straight approximately in line with apex of frontolateral peripheral sclerite apical portion, with a thick brush of villi on apical fifth. Frontolateral peripheral sclerite basoventral apophysis well developed; lacking medioventral carinae; right lateral fold produced into longitudinal spoon-shape process with apical portion open ventrally, ventral edge bordered with a brush of villi; left lateral lobe membranous, slightly developed; subapicodorsal lobe membranous, not reaching anterior edge, apex set medially in dorsal view; apical lobe round and directed obliquely on left side, left edge emarginate, apical villi irregular in shape; subapicoventral lobe round and interrupted in line with apical lobe, with few villi along apical edge only.

Variation. Measurements (237 ♂♂, 169 ♀♀). Length: male 7.0–13.0 mm (10.6 ± 1.1), female 7.5–13.0 mm (10.5 ± 1.0 mm). Female specimen. Cephalic outline in dorsal view as in Fig. 78; vertex with a slightly sinuate transverse carina, dorsal edge slightly sinuous in frontal view, lateral portion abruptly sloping down; anterior pronotal tubercles well developed, external lateral edges slightly widening toward apex in dorsal view, anterolateral surface obliquely concave, anterosuperior edge slightly arcuate in dorsal view (Fig. 101), lateral portion of anterosuperior edge slightly upturned (Fig. 102). Protibia short, with external teeth more robust.

Primary type data. Scarabaeus gazella Fabricius: Syntype (ZMUK): [] small green square; [ZMUC / 00508509]; [TYPE] red card; [WORLD / SCARAB. / DATABASE / WSD00029479] barcode label; [SYNTYPE ♂ / Scarabaeus / gazella / Fabricius, 1787 / vid. F. Génier, 2016] red card; [Guinea / Isert / Mus. T. Lund / Copris / gazella / ♂ / F.] handwritten [in error] see Nomenclature and taxonomy below for more details.

Note. The first label, a small green square without inscription, confirms this specimen as a Fabrician type (Zimsen 1964). The last label was added after Fabricius’s time and is erroneously cited as African. See nomenclature and taxonomy below section for more details.

Scarabaeus dorcas Olivier: Neotype ♂ (MNHN) present designation: [Madagascar / Diego-Suarez 7 / Ch. Alluaud, 1893] partly handwritten; [MUSEUM PARIS / MADAGASCAR / COLL. CH. ALLUAUD 1904] green card; [WORLD / SCARAB. / DATABASE / WSD00030025] barcode label; [NEOTYPE / Scarabaeus / dorcas / Olivier, 1789 / des. F. Génier, 2016] red card. Note. This neotype designation is made with the express purpose of clarifying the taxonomic status and the type locality of Scarabaeus dorcas Olivier. The specimen designated agrees in morphology to the description presented above. The original type material used for Olivier’s description is lost or destroyed (Olivier Montreuil, MNHN, personal communication). The neotype has been selected to match Olivier’s description and type locality, in addition to possessing the best diagnostic character for the species. The neotype is deposited in the collection of the Museum national d’Histoire naturelle de Paris along with the remains of Olivier’s collection.

MAP 9. Distribution of Digitonthophagus gazella (native range in continental Africa only).

Material examined (715 ♂♂, 407 ♀♀), distribution (Map 9): ANGOLA: CUANDO CUBANGO, Menongue (14°39'S, 17°41'E), iii.1914, Mission Rohan-Chabot— 1 ♂ (MNHN); HUILA, 10 km S Caconda (13°50'S, 15°6'E), 5.xi.2011, G. Werner— 2 ♀♀, 2 ♂♂ (FGIC, GWPC); 10 km S Cacula (14°35'S, 14°3'E), 4.xi.2011, G. Werner— 1 ♂ (GWPC); 30 km W Quipungo (14°48.02'S, 14°15.32'E), 26.xi.2013, G. Werner— 3 ♀♀, 2 ♂♂ (FGIC, GWPC); 33 km S Cacula (14°41'S, 13°54'E), 13.xi.2011, G. Werner— 1 ♀, 1 ♂ (GWPC); Capelongo près Luceque (14°39'S, 15°4'E), xii.1914, Mission Rohan-Chabot— 1 ♂ (MNHN); Capelongo-Dongo (14°31'S, 16°1'E), 1914, Mission Rohan-Chabot— 1 ♀ (MNHN); Capelongo-Dongo (14°31'S, 16°1'E), xii.1914, Mission Rohan-Chabot— 2 ♂♂ (MNHN); AUSTRALIA: NEW SOUTH WALES, Camp Blackman, Warrumbungle National Park (31°16'37''S, 148°59'51''E), 5–6.xii.1988, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Gilgandra (31°43'S, 148°40'E), 12.iv.1988, H. & A. Howden— 1 ♂ (CMNC); Narrabri (30°20'S, 149°47'E), 5.xii.1977, G.G. Burns— 2 ♀♀ (CMNC); NORTHERN TERRITORY, CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 1–10.xii.1993, S. & J. Peck (93-85)— 1 ♀, 1 ♂ (CMNC); CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 5.i.1994, S. & J. Peck (93-166)— 1 ♀, 1 ♂ (CMNC); CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 9.i.1994, S. & J. Peck (93-167)— 1 ♂ (CMNC); Elsey National Park, Mataranka (14°58'S, 133°11'E), 9.xii.1993, S. & J. Peck (93- 115)— 1 ♀, 1 ♂ (CMNC); Elsey National Park, Mataranka (14°58'S, 133°11'E), 9.xii.1993, S. & J. Peck (93- 116)— 1 ♀, 1 ♂ (CMNC); Gungaree Rainforest, South Alligator River, Kakadu National Park (12°40'46''S, 132°29'9''E), 22.xii.1993, S. & J. Peck (93-133)— 1 ♂ (CMNC); Magela Creek (12°40'S, 133°1'E), 4.xi.1989, C.W. & L.B. O'Brien— 1 ♂ (CMNC); Tenat Creek (19°39'S, 134°11'E), 22.ix.1988, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); QUEENSLAND, 12 mi. S Ayr (19°44'S, 147°28'E), 26.ii.1975, H. & A. Howden— 1 ♂ (CMNC); 16 km W Ravenshoe (17°39'S, 145°21'E), 2.i.1975, M.S. & B.J. Moulds— 1 ♂ (CMNC); 4–8 km S Atherton (17°20'S, 145°30'E), 26.xii.1988, H. & A. Howden— 1 ♂ (CMNC); 40 km N Lynd (18°36'S, 144°43'E), ii.1993, H. Demarz— 1 ♂ (CEMT); 8 km N Mareeba (16°55'S, 145°25'E), 28.i.1997, H. & A. Howden— 1 ♀ (CMNC); 9 km NE Julatten (16°33'S, 145°23'E), 13.xii.1986, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Atherton (17°16'S, 145°29'E), 7.ii.1975, H. & A. Howden— 1 ♀ (CMNC); Atherton (17°16'S, 145°29'E), 10.ii.1975, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Black Beach Reserve, near Mackay (21°4'39''S, 149°11'44''E), 28.i.1975, H. & A. Howden— 2 ♂♂ (CMNC); Burketown (17°44'S, 139°33'E), 20.ix.1975, Walford-Huggins— 1 ♀, 1 ♂ (CMNC); Cairns (16°55'S, 145°46'E), 10.v.1971, A. & M. Walford-Huggins— 1 ♀, 1 ♂ (CMNC); Cairns (16°55'S, 145°46'E), 24.viii.1971, A. & M. Walford-Huggins— 1 ♀, 1 ♂ (CMNC); Cairns (16°55'S, 145°46'E), 30.i.1975, H.

& A. Howden— 2 ♂♂ (CMNC); Cairns (16°55'S, 145°46'E), 3.ii.1975, H. & A. Howden— 1 ♂ (CMNC); Cairns (16°55'S, 145°46'E), 5.ii.1975, H. & A. Howden— 1 ♀ (CMNC); Charters Towers (20°5'S, 146°16'E), 16– 23.xii.1988, C. Lockett— 1 ♂ (CMNC); Granite Gorge Nature Park, 12 km SW Mareeba (17°2'32''S, 145°21'4''E), 8.i.1989, H. & A. Howden— 1 ♂ (CMNC); Holloways Beach (16°50'40''S, 145°44'20''E), 30.ix.1973, R.E. Parrott— 2 ♂♂ (CNC); Lockerbie, Cape York Peninsula (10°47'19''S, 142°27'33''E), 11.iv.1975, M. Walford- Huggins— 1 ♀, 1 ♂ (CMNC); Mareeba (16°59'S, 145°25'E), 19–28.xii.1988, H. & A. Howden— 1 ♀ (CMNC); Mornington Island (16°31'S, 139°24'E), 21.ix.1975, A. & M. Walford-Huggins— 1 ♂ (CMNC); Petford (17°20'34''S, 144°55'45''E), 5.ii.1989, H. & A. Howden— 1 ♂ (CMNC); Station Creek, 10 mi. S Mount Carbine (16°37'S, 145°13'E), 11.vi.1974, A. & M. Walford-Huggins— 1 ♂ (CMNC); Toowoomba (27°34'S, 151°57'E), 26– 27.ii.1983, Munroe & Common— 1 ♀, 1 ♂ (CMNC); Yeppoon (23°7'40''S, 150°44'43''E), 25.i.1975, H. & A. Howden— 1 ♀ (CMNC); Yeppoon (23°7'40''S, 150°44'43''E), 26.i.1975, H. & A. Howden— 1 ♂ (CMNC); Yeppoon (23°7'40''S, 150°44'43''E), 27.i.1975, H. & A. Howden— 1 ♂ (CMNC); BOTSWANA: CENTRAL, 120 km N Nata (19°12'58''S, 25°47'38''E), 29.xi.2003, Werner & Smrz— 1 ♂ (CMD); 35 km N Nata (19°55'S, 26°16'E), 26.xi.2001, B.D. Gill— 2 ♀♀, 2 ♂♂ (BDGC); 5 km SE Serowe (22°22'30''S, 26°52'30''E), 22.i.1985, Johnson— 1 ♂ (BMNH); Stevensford Game Reserve, 13 km E Sherwood (22°54.56'S, 28°2.36'E), 27–29.xi.2001, B. & J. Gill— 2 ♀♀, 2 ♂♂ (BDGC); NORTH-WEST, 20 km SW Sehitwa (20°22'30''S, 22°37'30''E), 22.i.1978, Holm, Jacobs, Kirsten, Scholtz— 1 ♂ (BMNH); Chobe River, 5 km E Kasane (17°48'S, 25°9'E), 18.i.2002, Holm, Marais, Nel, Saieva— 1 ♀, 3 ♂♂ (BMNH); Chobe, 20 km WSW Kasane, 938 m (17°50.2'S, 24°52.93'E), 25–26.xi.2001, B. & J. Gill— 1 ♀, 2 ♂♂ (BDGC); Kasane (17°47'S, 25°10'E), 2.xii.2012, M. Boulétreau— 2 ♀♀, 1 ♂ (CMD); Kuke Pan (20°59'S, 22°25'E), 15.iv.1972, Southern Africa Expedition— 1 ♀, 1 ♂ (BMNH); Maun (19°59'S, 23°25'E), ii.1973, [anonymous]— 1 ♂ (OMOC); SOUTH-EAST, Otse (25°2'S, 25°45'E), 16.i.1980, D. Levin— 1 ♂ (CMNC); BRAZIL: BAHIA, Cruz das Almas (12°40'27''S, 39°6'8''W), ii.1995, J.D.B. Macedo— 1 ♂ (CEMT); Mercado, Brumado (14°12'44''S, 41°40'18''W), 20.i.2012, B.N. de Souza— 1 ♂ (CEMT); Nazaré das Farinhas (13°2'17''S, 39°0'10''W), iii.2005, C. Ribeiro & A. Fonseca— 1 ♂ (CEMT); GOIÁS, Fazenda Piratininga, São Miguel do Araguaia (13°16'23''S, 50°9'49''W), 6–9.i.2000, V. Canhedo— 1 ♂ (CEMT); MARANHÃO, Sitio Melancia, Zona Rural, Mirador (6°22'21''S, 44°21'55''W), 19–20.xii.2006, F. Limeira-de-Oliveira— 1 ♂ (CEMT); MATO GROSSO, Base UFMT, Poconé (16°29'58''S, 56°24'49''W), iii.2012, M. Rossini— 1 ♂ (CEMT); Cáceres, Curvelândia (15°36'43''S, 57°55'19''W), ii–iii.2010, L.S. Silva— 1 ♂ (CEMT); Fazenda Conceição, Poconé (16°19'39''S, 56°30'14''W), 20.v.2012, M.B. Pessoa— 1 ♂ (CEMT); Fazenda Curitiba, Tangará da Serra, 320 m (14°21'48''S, 57°29'29''W), 31.iii.2012, R.J. Silva— 1 ♂ (CEMT); Fazenda São Luiz, Querêcia (12°38.611'S, 52°22.637'W), ii.2009, R. Andrade— 1 ♂ (CEMT); Fazenda São Nicolau, Cotriguaçu (9°51'23''S, 58°14'57''W), 14.x.2007, F. Vaz de Mello— 1 ♂ (CEMT); Fazenda São Nicolau, Cotriguaçu (9°51'23''S, 58°14'57''W), 31.x.2007, O. Peres Filho— 1 ♂ (CEMT); Fazenda São Nicolau, Cotriguaçu (9°51'23''S, 58°14'57''W), 14.xi.2007, O. Peres Filho— 2 ♂♂ (CEMT); Fazenda São Nicolau, Cotriguaçu (9°51'23''S, 58°14'57''W), 15.xii.2007, O. Peres Filho— 1 ♂ (CEMT); UNEMAT, Tangará da Serra, 442 m (14°39'58''S, 57°25'58''W), 26.vii.2008, R.J. Silva— 1 ♂ (CEMT); Vila Rica (10°1'S, 51°7'W), ii.2000, D.C. Souza— 1 ♀, 1 ♂ (CEMT); MATO GROSSO DO SUL, Fazenda São Bento, Proximo Porto Jofre, Corumbá (17°21'S, 56°44'W), 6.xii.2010, F.R. Tortato— 1 ♂ (CEMT); Pôrto Murtinho (21°42'53''S, 57°53'3''W), 26.i.2008, F.R. Fernandes— 1 ♀, 1 ♂ (CEMT); PARÁ, Alter do Chão (2°31'S, 54°57'W), 5.viii.2003, R. A. Matavelli— 1 ♂ (CEMT); Fazenda Marajoara, Pau d’Arco (7°50'S, 50°16'W), 16.x.1998, P.Y. Scheffler— 1 ♂ (CEMT); PARAÍBA, Fazenda Almas, São José dos Cordeiros, 650 m (7°28'S, 36°53'W), iii.2003, M.I. M. Hernández— 1 ♂ (CEMT); PERNAMBUCO, Floresta (8°35'S, 38°14'W), vii.2012, R.V. Nunes— 1 ♂ (CEMT); Japecanga (8°14'S, 36°8'W), 1.iv.2007, C. E.B. Nobre— 1 ♂ (CEMT); RONDÔNIA, Guajará-Mirim (10°22'16''S, 64°44'47''W), 25–27.i.2010, F. Coletti— 1 ♂ (CEMT); Guajará-Mirim (10°22'16''S, 64°44'47''W), 27.i.2010, F. Coletti— 1 ♂ (CEMT); Guajará-Mirim (10°22'16''S, 64°44'47''W), 15–18.ii.2010, F. Coletti— 1 ♀, 1 ♂ (CEMT); COMOROS: MWALI, Bandamalé, 420 m, ix.1958, Raharizonina— 4 ♀♀, 3 ♂♂ (MNHN); Fomboni, 10 m (12°17'2''S, 43°44'34''E), ix.1958, Raharizonina— 4 ♀♀, 3 ♂♂ (MNHN); Miringoni, 40 m (12°18'10''S, 43°38'11''E), ix.1958, Raharizonina— 4 ♀♀, 4 ♂♂ (MNHN); Mohéli Island (12°19'S, 43°43'E), 21.viii.1973, R. Mourglia— 1 ♀, 1 ♂ (FETC); Nioumachoua (12°22'S, 43°43'E), [no date], Pupier— 1 ♂ (MNHN); NZWANI, M’Remani, Anjouan (12°18'47''S, 44°29'51''E), ix.1958, Raharizonina— 1 ♀, 6 ♂♂ (MNHN); Mutsamudu, Anjouan, 50 m (12°10'S, 44°24'E), x.1958, Raharizonina— 1 ♀, 1 ♂ (MNHN); DEMOCRATIC REPUBLIC OF THE CONGO: KASAI ORIENTAL, Pastorale Lomami (6°11'S, 23°56'E), 12.iv.1941, H.-J. Brédo— 1 ♀, 2 ♂♂ (FGIC, IRSNB); KATANGA, Kaswabilenga, Parc National de l’Upemba, 700 m (8°51'S, 26°43'12''E), 11– 16.x.1947, G.F. de Witte (817a)— 1 ♂ (IRSNB); Kaswabilenga, Parc National de l’Upemba, 700 m (8°51'S, 26°43'12''E), 29–30.x.1947, G.F. de Witte (889a)— 2 ♀♀, 2 ♂♂ (IRSNB); Kaswabilenga, Parc National de l’Upemba, 700 m (8°51'S, 26°43'12''E), 30.x.1947, G.F. de Witte (902bis a)— 2 ♀♀, 2 ♂♂ (IRSNB); Kaswabilenga, Parc National de l’Upemba, 700 m (8°51'S, 26°43'12''E), 3–8.xi.1947, G.F. de Witte (915a)— 2 ♀♀, 2 ♂♂ (IRSNB); Kazofu (10°3'S, 24°46'E), xii.2004, T. Garnier— 1 ♀, 1 ♂ (PMOC); Lubumbashi (11°40'S, 27°29'E), 19.ii.1941, H.-J. Brédo— 1 ♂ (IRSNB); Mabwe, Parc National de l’Upemba, 585 m (8°39'12''S, 26°30'38''E), 17–22.xi.1948, G.F. de Witte (1970a)— 2 ♀♀, 2 ♂♂ (IRSNB); SOUTH-KIVU, Kabunambo (3°6'36''S, 29°8'18''E), 1956–1957, [anonymous]— 2 ♀♀, 2 ♂♂ (MNHN); Uvira (3°24'S, 29°8'E), [no date], [anonymous]— 2 ♀♀, 2 ♂♂ (MNHN); Uvira (3°24'S, 29°8'E), 1927, G. Babault— 1 ♂ (MNHN); DOMINICAN REPUBLIC: SAN PEDRO DE MACORÍS, Juan Dolio (18°26'37''N, 69°24'51''W), 26.x.2000, D. Prunier— 2 ♀♀, 2 ♂♂ (OMOC); FIJI: NORTHERN DIVISION, Nadi (16°55'55''S, 178°46'7''E), 27.viii.2004, H. & A. Howden— 1 ♂ (CMNC); WESTERN DIVISION, Tavua (17°26'30''S, 177°51'48''E), 20–23.viii.1978, S. & J. Peck— 1 ♀, 1 ♂ (CMNC); FRANCE: MAYOTTE, Chingoni, 70 m (12°48'S, 45°7'E), x.1958, Raharizonina— 1 ♂ (MNHN); Dembeni (12°50'40''S, 45°11'5''E), 1971, Brunhes— 1 ♀, 1 ♂ (MNHN); [unspecified locality] (12°46'S, 45°13'E), [no date], [anonymous]—2 ♂♂ (BMNH, IRSNB); GUADELOUPE: GRANDE-TERRE, Le Gosier (16°13'N, 61°29'W), ii.1992, O. Montreuil— 1 ♂ (OMOC); JAMAICA: ST. ANN PARISH, Discovery Bay (18°27'30''N, 77°23'55''W), 25.v.1990, G. Haghebaert— 1 ♂ (IRSNB); Priory (18°26'27''N, 77°13'26''W), 4.vi.1990, G. Haghebaert— 1 ♂ (IRSNB); KENYA: KILIFI, Kikambala (3°49'S, 39°48'E), v.1993, [anonymous]— 5 ♀♀, 6 ♂♂ (JFJC); KWALE, de Tiwi à Gazi (4°20'S, 39°33'E), xi.1911, Alluaud & Jeannel— 2 ♀♀, 1 ♂ (MNHN); MOMBASSA, Mombassa (4°3'S, 39°40'E), xi.1985, [anonymous]— 2 ♀♀, 2 ♂♂ (CMD); Mombassa (4°3'S, 39°40'E), [no date], C. Alluaud— 1 ♂ (MNHN); Mombassa (4°3'S, 39°40'E), 10.i.1939, A.H.N.— 1 ♂ (BMNH); Mombassa (4°3'S, 39°40'E), 10.iii.1939, A.H.N.— 3 ♀♀, 3 ♂♂ (BMNH); TAITA TAVETA, Voi (3°23'S, 38°34'E), xi.1997, M. Snižek— 1 ♂ (CEMT); MADAGASCAR: ANTANANARIVO, Ambatolampy (19°23'S, 47°26'E), 1931, Lasère— 1 ♀, 1 ♂ (MNHN); Antananarivo (18°53'S, 47°30'E), 1931, Lasère— 1 ♀, 1 ♂ (MNHN); DIEGO- SUAREZ, Amber Gebirge [= Montagne d’Ambre] (12°35'S, 49°9'E), [no date], H. Rolle— 2 ♂♂ (IRSNB); Diego- Suarez [= Antisiranana] (12°16'S, 49°17'E), 1893, C. Alluaud— 2 ♀♀, 3 ♂♂ (IRSNB, MNHN); Diego-Suarez [= Antisiranana] (12°16'S, 49°17'E), 1939–1941, [anonymous]— 1 ♀ (MNHN); Marovato, 30 km E Ambanja (13°56'51''S, 48°33'19''E), 1.xii.2001, I. Pljushtch— 1 ♂ (OUMNH); Nosy Be (13°19'24''S, 48°15'34''E), 1885, Stumpff— 3 ♀♀, 3 ♂♂ (SMF); Nosy Be (13°19'24''S, 48°15'34''E), 23.xi.1898, P. Frey— 2 ♀♀ (SMF); Nosy Be (13°19'24''S, 48°15'34''E), 30.xii.1998, G. Ferrer— 3 ♂♂ (OMOC); FIANARANTSOA, Ambalavao (21°50'S, 46°56'E), [no date], [anonymous]— 1 ♀, 1 ♂ (MNHN); Camp Catta, 40 km S Ambalavao (22°4'48''S, 46°46'29''E), 26.xi–2.xii.2003, S. Murzin & A. Shamaev— 1 ♀ (CMD); Fianarantsoa (21°27'S, 47°5'E), [no date], [anonymous]— 1 ♀, 2 ♂♂ (BMNH); km. P. 713, Isalo, 1000 m (22°38'S, 45°20'E), 18.iii.1968, K.M. G. & P.D. — 1 ♂ (BMNH); Vondrozo (22°49'S, 47°19'E), 1921, [anonymous]— 1 ♀, 1 ♂ (MNHN); MAHAJANGA, Ananalava (14°36'S, 47°44'E), 13.v.1973, P. Calabri— 1 ♂ (CMNC); Environs Ambodimanga (16°41'S, 45°50'E), 14– 16.xi.1995, J. Stolarczyk— 2 ♂♂ (OUMNH); Mandritsara (15°50'S, 48°49'E), [no date], [anonymous]— 1 ♀, 1 ♂ (MNHN); Nosy Béraphia (14°0'S, 47°48'E), 1934, [anonymous]— 1 ♂ (MNHN); MAJUNGA, Majunga (15°43'S, 46°20'E), v.1968, S. Key— 2 ♂♂ (BMNH); Majunga (15°43'S, 46°20'E), 14.v.1968, K.M. Guichard— 1 ♂ (BMNH); TAMATAVE, Ambatondrazaka (17°50'S, 48°25'E), xi.1983, P. Arnaud— 1 ♂ (BDGC); Andasibe- Mantadia National Park, Périnet (18°53'3''S, 48°25'44''E), 21.iv.1993, J.P. & S. Lignon— 1 ♀, 1 ♂ (CMD); Antanambe (16°26'S, 49°51'E), x.1972, [anonymous]— 2 ♂♂ (CEMT); Antanambe (16°26'S, 49°51'E), x.1970, [anonymous]— 1 ♂ (CEMT); Antanambe (16°26'S, 49°51'E), x.1971, [anonymous]— 1 ♂ (CEMT); Périnet (18°55'24''S, 48°25'11''E), xi.2003, [anonymous]— 1 ♀, 1 ♂ (CMD); Tamatave [= Toamasina] (18°8'50''S, 49°23'43''E), [no date], [anonymous]— 1 ♂ (IRSNB); TANARIVE, Tsiroanomandidy (18°46'11''S, 46°2'58''E), xi.1983, P. Arnaud— 1 ♀, 1 ♂ (BDGC); TOAMASINA, Maroantsetra (15°26'S, 49°44'E), 7.i.1999, J. Moravec— 1 ♂ (FETC); Station agronomique du lac Alaotra (17°48'S, 48°24'E), 18.ii.1988, Donskoff— 1 ♀, 1 ♂ (MNHN); TOLIARA, Ankilimary (23°26'S, 45°33'E), 25.v.1948, [anonymous]— 2 ♂♂ (CMNC); Beloha (25°10'26''S, 45°3'36''E), [no date], [anonymous]— 1 ♀, 1 ♂ (MNHN); Betioky, 275 m (23°43'S, 44°23'E), 4.iv.1968, K.M. G. & P.D. — 1 ♀, 2 ♂♂ (BMNH); Betroka (23°16'2''S, 46°5'50''E), [no date], [anonymous]— 2 ♀♀, 2 ♂♂ (IRSNB); Fort Dauphin [= Taolagnaro] (25°2'S, 46°59'E), x.2006, Bouletreau— 1 ♀, 1 ♂ (CMD); Lavanono (25°25'47''S, 44°57'16''E), 20.x.2009, G. Prelle— 5 ♀♀, 5 ♂♂ (PMOC); Sakaraha (22°54'41''S, 44°31'55''E), [no date], [anonymous]— 3 ♀♀, 3 ♂♂ (SMF); Sakaraha (22°54'41''S, 44°31'55''E), 2.iii.1963, R. Vieu— 2 ♂♂ (PMOC); MALAWI: CENTRAL REGION, 20 km NW Dedza (14°16'S, 34°9'E), 28.xi.1983, [anonymous]— 1 ♂ (BMNH); Lilongwe, Sani (13°55'S, 33°25'E), 5.xii.1983, [anonymous]— 1 ♂ (BMNH); SOUTHERN REGION, Chikwawa (16°2'14''S, 34°47'55''E), 14.xii.1993, C.R. Owen— 1 ♀, 1 ♂ (PMOC); Monkey Bay (14°5'S, 34°55'E), iii.1989, C.R. Owen— 1 ♀, 1 ♂ (PMOC); Balaka, near Phalula (15°14'S, 34°53'E), 27.i.2001, J.-F. Josso— 1 ♀, 1 ♂ (JFJC); Mangochi, Nkopola Forest Reserve (14°19'39''S, 35°8'57''E), 20–21.i.1985, A.V. Evans & C.H. Scholtz— 1 ♀, 1 ♂ (BMNH); MEXICO: SAN LUIS POTOSÍ, San Nicolás Tolentino (22°15'N, 100°33'W), viii.1996, S. Boucher— 2 ♀♀, 2 ♂♂ (PMOC); SINALOA, Guamúchil (25°27'31''N, 108°4'40''W), 1.viii.2002, [anonymous]— 1 ♀ (CMD); SONORA, Yécora (28°22'N, 108°55'W), 5.ix.2012, [anonymous]— 1 ♀, 1 ♂ (CMD); VERACRUZ, Corral Nuevo (18°6'46''N, 95°6'54''W), 17.viii.2002, S. Prepsl— 1 ♂ (CMD); MOZAMBIQUE: CABO DELGADO, Mareja (site 1), Parque Nacional Quirimbas, 100 m (12°50'56''S, 40°10'53''E), 22.xii.2012, F. & S. Génier & M. Denja (2012-02)— 1 ♀ (FGIC); Parque Nacional Quirimbas, 100 m (12°50'56''S, 40°10'53''E), 23.xii.2012, F. & S. Génier & M. Denja (2012-05)— 2 ♀♀, 2 ♂♂ (FGIC); Pemba (12°58'S, 40°30'E), [no date], F.V. Beste— 1 ♂ (BMNH); INHAMBANE, Zavala, Quissico (24°40'58''S, 34°40'7''E), 12.xi.1967, M.I. Russell— 1 ♂ (BMNH); MANICA, Casa Msika Lodge / Camping, 20 km E Chimoio (19°2'25''S, 33°3'55''E), 24.xii.2005, P. Schüle— 1 ♂ (SMNS); Chimoio (19°7'S, 33°27'E), ix.1928, P. Lesne— 3 ♂♂ (FGIC, MNHN); Chimoio (19°7'S, 33°27'E), iii.1928, P. Lesne— 1 ♀, 1 ♂ (MNHN); environs de Tambara (16°43'S, 34°15'E), iv.1926, J. Surcouf—1 ♂ (MNHN); MAPUTO, Lourenço Marques [= Maputo] (25°57'55''S, 32°35'21''E), 10.viii.1900, F. Muir— 2 ♂♂ (BMNH); Mission Rikatla (25°38'S, 32°31'E), [no date], H. A. Junod— 2 ♀♀, 2 ♂♂ (MNHN); NAMPULA, Lumbo (15°2'S, 40°40'E), ii.1983, F. Králík— 1 ♀ (CVM); SOFALA, 10 mi. NW Beira (19°41'S, 34°46'E), 16.iv.1971, Bornemissza & Aschenborn— 1 ♂ (CMNC); Caia (17°49'13''S, 35°20'39''E), xii.1926, J. Surcouf— 1 ♂ (MNHN); Caia (17°49'13''S, 35°20'39''E), 23.x.1910, H. Swale— 1 ♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 25.x.1910, H. Swale— 1 ♀, 1 ♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 5.xii.1910, H. Swale— 1 ♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 25.xii.1910, H. Swale— 1 ♀, 2 ♂♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 23.ix.1911, H. Swale— 1 ♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 29.ii.1912, H. Swale— 1 ♂ (BMNH); Caia (17°49'13''S, 35°20'39''E), 18.iii.1912, H. Swale— 1 ♂ (BMNH); Canxixe (17°35'S, 34°19'E), xii.1926, J. Surcouf— 2 ♀♀ (MNHN); Chemba (17°10'S, 34°53'E), xi.1928, J. Surcouf— 1 ♀, 1 ♂ (MNHN); Chemba (17°10'S, 34°53'E), xii.1928, J. Surcouf— 1 ♀, 1 ♂ (MNHN); Chemba (17°10'S, 34°53'E), 1928, J. Surcouf— 2 ♂♂ (MNHN); Gorongoza National Park (18°49'S, 34°29'E), 15.xii.1972, C. Besnard— 1 ♀, 2 ♂♂ (PMOC); Nova Chupanda (18°2'S, 35°36'E), xii.1928 – i.1929, J. Surcouf— 2 ♀♀, 2 ♂♂ (FGIC, MNHN); NAMIBIA: CAPRIVI, 4 km W Immelman runway, West Caprivi game Park (17°46'56''S, 23°16'31''E), 14.xii.1999, Mann & Marais— 1 ♀ (OUMNH); KAVANGO, 2 km W Xawasha Pan, Tsunkwe District (19°9'57''S, 20°52'55''E), 26.xii.1998, D.J. Mann & E. Marais— 1 ♀ (OUMNH); 5 km N Nova, Caprivi Game Park (18°9'56''S, 21°44'31''E), 16.xii.1999, Mann & Marais— 3 ♂♂ (OUMNH); 5 km N Nova, Caprivi Game Park (18°9'56''S, 21°44'31''E), 17.xii.1999, Mann, Marais, & Newman— 4 ♀♀, 5 ♂♂ (OUMNH); 5 km N Nova, Caprivi Game Park (18°9'56''S, 21°44'31''E), 17.xii.1999, Mann & Marais— 2 ♀♀, 2 ♂♂ (OUMNH); Divundu, West Caprivi Game Park (18°4'4''S, 21°28'51''E), 1.i.1999, D.J. Mann & E. Marais— 1 ♀, 1 ♂ (OUMNH); Khaudom Game Reserve, 5 km N Cwiba (18°23'4''S, 20°43'1''E), 30.xii.1998, D.J. Mann & E. Marais— 2 ♂♂ (OUMNH); Mayana Lodge, Rundu District (17°51'55''S, 19°55'48''E), 19.xii.1999, Mann & Marais— 1 ♀ (OUMNH); Mile 46 Agricultural Station, Novo area (18°15'52''S, 19°15'18''E), 25.iii.2002, Mann, Marais, & Kasch— 4 ♀♀, 5 ♂♂ (OUMNH); Shamvura Camp (18°2'3''S, 20°51'40''E), 30–31.xii.2006, R. Perissinoto & L. Clennell— 1 ♀, 1 ♂ (PMOC); OTJOZONDJUPA, Aha Hills, Tsumkwe District (19°47'36''S, 20°59'51''E), 22.xii.1998, Mann & Marais— 1 ♀ (OUMNH); Nama Pan area, Tsumkwe district (19°54'34''S, 20°44'8''E), 21.xii.1998, D.J. Mann & E. Marais— 1 ♀ (OUMNH); NEW CALEDONIA: NORTH PROVINCE, Île Art (19°43'S, 163°40'E), 7–8.iii.1995, M. Boulard— 1 ♂ (MNHN); SOUTH PROVINCE, La Foa (21°42'36''S, 165°49'40''E), 16.xi.2007, T. Thery— 1 ♀, 1 ♂ (CTT); La Foa (21°42'36''S, 165°49'40''E), 3.xii.2007, T. Thery— 1 ♀, 1 ♂ (CTT); Route des Nordistes, Bourail (21°35.706'S, 165°24.593'E), 21.ii.2015, M. Dierkens— 1 ♀, 1 ♂ (CMD); Sentier du Pic de Ouitchambo, Bouloupari (21°52'S, 166°3'E), 15.iii.2013, M. Dierkens— 1 ♀, 1 ♂ (CMD); [NO DATA]:, [no date], [anonymous]— 1 ♂ (ZMUC); PAPUA NEW GUINEA: MADANG, Nagada Mission (5°9'24''S, 145°47'24''E), 6.v.1992, I. Lansbury— 1 ♂ (OUMNH); Nagada Mission (5°9'24''S, 145°47'24''E), 18.v.1992, I. Lansbury— 1 ♂ (OUMNH); MOROBE, Lae (6°44'S, 147°0'E), 3.iii.1973, K.W. Stroeder— 1 ♂ (CNC); SOUTH AFRICA: EASTERN CAPE, Black Rock (32°1'S, 29°6'E), 15.ii.1997, C.R. Owen— 1 ♀, 1 ♂ (CEMT); East London (32°59'S, 27°52'E), [no date], Dr. Martin— 1 ♂ (MNHN); East London (32°59'S, 27°52'E), 13.i.1976, R.E. Parrott— 1 ♂ (CNC); East London (32°59'S, 27°52'E), 15.ii.1976, R.E. Parrott— 2 ♀♀, 1 ♂ (CNC); Grahamstown (33°18'S, 26°32'E), [no date], A. Vogt— 1 ♂ (IRSNB); Mbotyi Coast and Forest (31°27'47''S, 29°43'58''E), 29.xi–3. xii.2 0 0 3, W. Schawaller— 2 ♀♀, 2 ♂♂ (SMNS); Port St. John, Pondoland (31°38'S, 29°32'E), x.1923, R.E. Turner— 1 ♂ (BMNH); Port St. John, Pondoland (31°38'S, 29°32'E), 6–25.ii.1924, R.E. Turner— 1 ♂ (BMNH); Silaka Wildlife Reserve, Port St. John (31°39'15''S, 29°30'20''E), 8.i.2009, P. Schüle— 1 ♂ (PMOC); The Haven Hotel, Dwesa-Cwebe Wildlife Reserve & Marine Sanctuary (32°14'26''S, 28°54'38''E), 4–6.xii.2003, P. Schüle— 1 ♂ (SMNS); Umtata [= Mthatha] (31°35'S, 28°47'E), 18.ii–18.iii.1923, R.E. Turner— 2 ♂♂ (BMNH); FREE STATE, 20 km W Boshof (28°39'S, 25°4'E), 12– 14.xii.1984, H. & A. Howden— 2 ♂♂ (CMNC); Bloemfontein (29°8'S, 26°12'E), [no date], E. Eckersley— 1 ♂ (BMNH); Florisbad (28°46'S, 26°5'E), 9–12.xii.1984, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Kimberley (28°44'S, 26°46'E), [no date], [anonymous]— 1 ♀, 2 ♂♂ (BMNH); Kimberley (28°44'S, 26°46'E), [no date], Rombach— 1 ♂ (IRSNB); Vredefort (27°0'27''S, 27°22'6''E), 17.xii.2009, Perissinoto & Clennell— 1 ♀, 1 ♂ (PMOC); GAUTENG, 15 mi. SW Johannesburg (26°25'S, 27°52'E), 28.xii.1971, Southern Africa Expedition ((3))— 1 ♂ (BMNH); Boekenhoutskloof, 30 km NE Pretoria (25°32'S, 28°30'E), 4.i.1977, G. Bernon— 2 ♀♀, 2 ♂♂ (BDGC); Boekenhoutskloof, 30 km NE Pretoria (25°32'S, 28°30'E), 7.i.1977, G. Bernon— 1 ♀, 2 ♂♂ (BDGC); Boekenhoutskloof, 30 km NE Pretoria (25°32'S, 28°30'E), 7.xii.1977, G. Bernon— 1 ♂ (CMNC); Boekenhoutskloof, 30 km NE Pretoria (25°32'S, 28°30'E), 5.ii.1978, G. Bernon— 2 ♂♂ (BDGC); Castel Gorge, 42 mi. W Pretoria (25°49'S, 27°37'E), 27.ii.1972, Bornemissza & Insley— 1 ♂ (MNHN); Die Wilgers Silverton (25°37'30''S, 28°7'30''E), iii.1986, C.H. Scholtz— 4 ♂♂ (BMNH); Heidelberg (26°29'30''S, 28°23'0''E), 25.xii.1978, P. Walter— 1 ♀ (PHWC); Johannesburg (26°12'S, 28°2'E), 28.xii.1971, Southern Africa Expedition— 2 ♀♀, 2 ♂♂ (BMNH); Magalieskruin, Pretoria (25°41'9''S, 28°14'2''E), 31.x.1977, J. Boomker— 1 ♀, 1 ♂ (BMNH); Nagana Research Laboratory, Onderstepoort (25°39'S, 28°11'E), 1922, H.H. Curson— 3 ♂♂ (BMNH); Nagana Research Laboratory, Onderstepoort (25°39'S, 28°11'E), xii.1922, H.H. Curson— 1 ♂ (BMNH); Pretoria (25°42'25''S, 28°13'46''E), [no date], W.L. Distant— 2 ♀♀, 2 ♂♂ (BMNH); Pretoria (25°42'25''S, 28°13'46''E), iv.1962, S.C. K.— 1 ♂ (BMNH); Pretoria (25°42'25''S, 28°13'46''E), iii.1963, W. R.— 1 ♂ (BMNH); Pretoria (25°42'25''S, 28°13'46''E), [no date], L.M. Bucknill— 2 ♂♂ (BMNH); Pretoria (25°42'25''S, 28°13'46''E), 1893, E. Simon— 1 ♂ (MNHN); Pretoria (25°42'25''S, 28°13'46''E), 24.x.1914, [anonymous]— 1 ♂ (BMNH); Pumulani (25°37'30''S, 28°7'30''E), 9.x.1981, T. J.H. Bernaro— 1 ♂ (BMNH); Rietondale Research Station, Pretoria (25°43'49''S, 28°14'7''E), 5.ii.1978, G. Bernon— 1 ♂ (BDGC); Rivonia (26°3'S, 28°3'E), 28.x.1967, M.I. Russell— 1 ♂ (BMNH); Soetvelde (26°45'S, 28°1'E), 27.xii.1989, L.G. Le Roux— 1 ♂ (BMNH); Soutpan near Pretoria (25°36'S, 28°10'E), 10.iii.1992, J. Klimaszewski— 1 ♂ (CMNC); University of Pretoria Experimental Farm (25°44'49''S, 28°15'31''E), 3.vi.1985, D. D'Hotman— 1 ♂ (BMNH); KWAZULU-NATAL, 30 km NW Nongoma, 808 m (27°41'S, 31°29'E), 28.i.2008, P. Schüle— 1 ♀, 1 ♂ (PMOC); 5 mi. S Bergville (28°48'S, 29°23'E), 4– 5.xii.1971, A. A. Kirk— 1 ♂ (CMNC); Boschberg Farm, Ladysmith (28°14'S, 29°46'E), 9–11.i.2001, A.L.V. Davis— 2 ♀♀, 2 ♂♂ (BMNH); Busland’s Halt (28°0'S, 32°0'E), 15.i.1986, H.H. Aschenborn— 1 ♀, 1 ♂ (BMNH); Cathedral Peak State Forest, 75 km WSW Estcourt (29°0'S, 29°15'E), 7–31.xii.1979, S. & J. Peck— 2 ♂♂ (CMNC); Colenso (28°44'S, 29°49'E), ii–iii.1900, H.B. Marley— 2 ♂♂ (BMNH); Durban (29°51'S, 31°1'E), 1902, F. Muir— 1 ♂ (BMNH); Enseleni [= Mseleni] (27°22'13''S, 32°31'32''E), 15.x.2005, Perissinoto & Clennell— 2 ♂♂ (PMOC); Enseleni [= Mseleni] (27°22'13''S, 32°31'32''E), 19.xi.2009, Perissinoto & Clennell— 1 ♀ (PMOC); Estcourt (29°1'S, 29°53'E), [no date], [anonymous]— 1 ♂ (BMNH); Estcourt (29°1'S, 29°53'E), 24.x.2009, Perissinoto & Clennell— 1 ♂ (PMOC); Frere, 1158 m (28°53'35''S, 29°46'24''E), x.1891, G. A.K. Marshall— 1 ♀, 1 ♂ (BMNH); Giants Castle, Drakensberg (29°16'S, 29°31'E), 27.xii.1978, P. Walter— 2 ♀♀, 2 ♂♂ (PHWC); Hluhluwe Game Reserve (28°0'S, 32°0'E), i.1986, Aschenborn— 1 ♂ (BDGC); Hluhluwe Game Reserve (28°0'S, 32°0'E), 3.i.1978, J.C. & E. Willame— 1 ♂ (PHWC); Hluhluwe Game Reserve (28°0'S, 32°0'E), 30.xii.1978, P. Walter— 1 ♀, 4 ♂♂ (PHWC); Hluhluwe Game Reserve (28°0'S, 32°0'E), 2.x.1985, E. Jane Wright— 3 ♂♂ (BMNH); Hluhluwe Game Reserve (28°0'S, 32°0'E), 16.xi.1985, H.H. Aschenborn— 6 ♀♀, 3 ♂♂ (BMNH); Hluhluwe Game Reserve (28°0'S, 32°0'E), 19–22.ii.1986, DBRU [= Dung Beetle Research Unit]— 2 ♀♀, 2 ♂♂ (BDGC); Ithala Game Reserve (27°30'S, 31°14'E), 13–14.i.1999, A.L.V. Davis— 3 ♀♀, 1 ♂ (BMNH); Ladysmith (28°33'S, 29°47'E), [no date], [anonymous]— 1 ♀, 1 ♂ (BMNH); Ladysmith (28°33'S, 29°47'E), [no date], Dr. Martin— 1 ♂ (MNHN); Malvern (29°53'S, 30°55'E), iii.1896, G. A.K. Marshall— 1 ♂ (BMNH); Maritzburg (29°37'S, 30°23'E), xi.1896, [anonymous]— 1 ♀, 1 ♂ (BMNH); Maritzburg (29°37'S, 30°23'E), ii.1896, J. Cregoe— 2 ♀♀, 2 ♂♂ (BMNH); Mkuze (27°37'S, 32°2'E), iii.1984, O. Schmitt— 1 ♂ (PMOC); Mtunzini (28°57'35''S, 31°45'0''E), 28.ix.2007, Perissinoto & Clennell— 1 ♂ (PMOC); Ndumu Game Reserve (26°53'S, 32°16'E), 5–7.xi.1984, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Newcastle (27°45'S, 29°56'E), [no date], J.P. Cregoe— 2 ♀♀, 2 ♂♂ (BMNH); Nongoma (27°53'40''S, 31°38'34''E), 28.i.2008, Perissinoto & Clennell— 2 ♀♀, 2 ♂♂ (PMOC); Nongoma (27°53'40''S, 31°38'34''E), 28.i.2008, P. Schüle— 1 ♀, 1 ♂ (PMOC); Port Natal [= Durban] (29°51'S, 31°1'E), [no date], [anonymous]— 2 ♂♂ (BMNH); Port Natal [= Durban] (29°51'S, 31°1'E), [no date], V. van De Poll— 1 ♀, 1 ♂ (ZMUC); Royal Natal Park (28°42'S, 28°56'E), 3.i.1978, J. & E. Willame— 1 ♀, 2 ♂♂ (PHWC); SW Magudu (27°34'S, 31°35'E), 4–5.i.2009, P. Schüle—2 ♀♀, 2 ♂♂ (SMNS); The Caverns Drakensberg (28°38'8''S, 28°57'43''E), 26.xii.1978, P. Walter— 1 ♀ (PHWC); Umfolozi Game Reserve (28°7'S, 32°3'E), 11.xi.1970, H. & M. Townes— 1 ♂ (CMNC); Umfolozi Game Reserve (28°7'S, 32°3'E), 2.i.1978, J. & E. Willame— 1 ♀ (PHWC); Umfolozi Game Reserve (28°7'S, 32°3'E), 30.xii.1978, P. Walter— 2 ♂♂ (PHWC); Umfolozi Game Reserve (28°7'S, 32°3'E), 5.x.2000, W. Grosser— 2 ♀♀, 2 ♂♂ (OUMNH); uMkhuze Game Reserve (27°40'S, 32°15'E), 22–24.iii.1984, P.B. Edwards & H.H. Aschenborn— 1 ♀, 1 ♂ (CMD); Weenen (28°51'S, 30°4'E), 1925, H.P. Thomasset— 1 ♀, 2 ♂♂ (BMNH); Weenen (28°51'S, 30°4'E), iii.1928, H.P. Thomasset— 1 ♂ (BMNH); Weenen (28°51'S, 30°4'E), xi.1923, H.P. Thomasset— 1 ♂ (BMNH); LIMPOPO, Anthrax camp, Pafuri, Kruger National Park (22°25'S, 31°14'E), 9–24.xi.1985, M. Brigham— 1 ♀, 1 ♂ (CMNC); Ellisras [= Lephalale] (23°40'S, 27°45'E), iv.1909, R. H.— 1 ♂ (BMNH); Farm Sericea, Nylsvaley (24°25'48''S, 28°23'24''E), 27.xi.1985, S.L. Chown— 5 ♂♂ (BDGC); Farm Sericea, Nylsvaley (24°25'48''S, 28°23'24''E), 11– 12.xii.1985, A.V. Evans— 1 ♂ (BMNH); Guernsey Farm, 15 km E Klaserie, 500 m (24°33'S, 31°9'E), 19– 31.xii.1985, H. & A. Howden— 1 ♂ (CMNC); Guernsey Farm, 15 km NE Klaserie (24°33'S, 31°9'E), 18– 30.xii.1985, S. & J. Peck— 1 ♀, 1 ♂ (CMNC); Guernsey Farm, 15 km NE Klaserie (24°33'S, 31°9'E), 18– 31.xii.1985, S. & J. Peck— 2 ♀♀, 2 ♂♂ (CMNC); Kruger National Park North (22°44'S, 31°11'E), 25.xi.2012, P. Richoux— 1 ♀, 2 ♂♂ (CMD); Langjan (22°57'S, 28°15'E), 27.ii.1988, M. Vogt— 1 ♂ (BMNH); Langjan Nature Reserve (22°52'30''S, 29°7'30''E), 21.ii.1987, W.Z. Schultz— 1 ♂ (BMNH); Makapan (23°11'S, 28°36'E), 1893, E. Simon— 1 ♂ (MNHN); Mmabolela estate (22°40'S, 28°15'E), 20–24.xi.1991, F. Génier— 2 ♀♀, 1 ♂ (CMNC); Musina (22°21'S, 30°2'E), 28.ii.1998, F.M. Brits— 1 ♂ (BMNH); Nwanedzi (23°45'28''S, 30°27'17''E), 28.xii.2009, Perissinoto & Clennell— 1 ♀, 2 ♂♂ (PMOC); Radium (25°5'S, 28°17'E), 3.iii.1989, P. Snyman— 1 ♂ (BMNH); Sentrum (24°18'S, 27°20'E), 11.iii.1989, E. Swanepoel— 1 ♂ (BMNH); Shiluvane (24°3'S, 30°17'E), [no date], H. A. Junod— 7 ♂♂ (IRSNB, MNHN); Steenbokpan (23°43'S, 27°15'E), 4.v.1988, E. Swanepoel— 1 ♂ (BMNH); Thornybush Lodge, Hoedspruit (24°21'S, 30°57'E), 7.xii.1992, Jansen & Klimaszewski— 1 ♂ (CMNC); Vaalwater (24°7'30''S, 28°22'30''E), 2.iii.1980, T.L. Prinsloo— 1 ♂ (BMNH); Waterpoort (22°52'30''S, 29°22'30''E), iv.1978, I. Bruner— 1 ♂ (BMNH); Woodside, Letsitele (23°53'25''S, 30°23'10''E), 15.iii.1976, R.E. Parrott— 2 ♀♀, 3 ♂♂ (CMNC, CNC); MPUMALANGA, 10 km S Skukuza, Kruger National Park (25°4'6''S, 31°32'56''E), 12– 14.xii.1985, E.G. Monroe— 2 ♂♂ (CMNC); Barberton (25°47'S, 31°3'E), xi–xii.1908, L.C. de Beer— 1 ♀, 2 ♂♂ (MNHN); Berg-en-Dahl Camp, Kruger National Park (25°25'24''S, 31°27'0''E), 15.xii.1984, M. Davis— 1 ♀, 4 ♂♂ (CNC); Kaapmuiden (25°32'S, 31°20'E), 4–7.xii.1984, H. & A. Howden— 2 ♂♂ (CMNC); Klaserie Nature Reserve (24°12'20''S, 31°12'19''E), 28.xi.1998, J.-F. Josso— 1 ♂ (JFJC); Klaserie Nature Reserve (24°12'20''S, 31°12'19''E), 25–27.xi.2002, P. Schüle— 2 ♂♂ (SMNS); Leeuwfontein (25°13'S, 28°53'E), 8.iv.1998, Jiya— 1 ♂ (BMNH); Mariepskop, Blyde River, 800–1000 m (24°35'32''S, 30°52'56''E), 23–26.xi.2008, W. Schawaller— 1 ♀, 1 ♂ (SMNS); Near Satara, Kruger National Park (24°24'S, 31°47'E), 15–18.xii.1985, H. & A. Howden— 1 ♀, 1 ♂ (CMNC); Satara, Kruger N. Pk. (24°23'S, 31°43'E), 15–18.xii.1985, S. & J. Peck— 1 ♀, 1 ♂ (CMNC); SE 25 31 Ad (25°22'30''S, 31°22'30''E), 17.ii.1981, S. Mostert— 1 ♂ (BMNH); Skukuza Camp (24°59'40''S, 31°35'14''E), i.1999, D. Inward— 5 ♀♀, 5 ♂♂ (BMNH); Skukuza, Kruger National Park (24°59'40''S, 31°35'14''E), 12– 15.xii.1985, S. & J. Peck— 2 ♀♀, 1 ♂ (CMNC); Skukuza, Kruger National Park (24°59'40''S, 31°35'14''E), 12– 14.xii.1985, S. & J. Peck— 1 ♂ (CMNC); Skukuza, Kruger National Park (24°59'40''S, 31°35'14''E), 12– 14.xii.1985, H. & A. Howden— 1 ♂ (CMNC); Skukuza, Kruger National Park (24°59'40''S, 31°35'14''E), 12.xii.1986, M. Sanborne— 1 ♂ (CNC); Timbavati Game Reserve (24°28'S, 31°17'E), 23.i.1996, P. Holter— 1 ♀, 3 ♂♂ (ZMUC); Timbavati river, Kruger National Park (24°5'S, 31°40'E), i.1999, D. Inward— 2 ♀♀, 3 ♂♂ (BMNH); Tshokwana-Nwanedzi Road, Kruger National Park (24°37'S, 31°57'E), i.1999, D. Inward— 2 ♀♀, 2 ♂♂ (BMNH); NORTH WEST, 16 km NW Brits (25°22'30''S, 27°37'30''E), xii.1977, E.E. Zocke— 1 ♂ (BMNH); 20 km W Bothaville (27°19'48''S, 26°27'12''E), 31.i.2001, M. Snižek— 1 ♀, 1 ♂ (CEMT); Baberspan (25°7'S, 26°5'E), 14– 21.xii.1993, T. Joffe— 1 ♂ (BMNH); Dinokana (25°27'S, 25°52'E), [no date], [anonymous]— 1 ♀, 2 ♂♂ (SMF); Nieverdiend (25°7'30''S, 26°7'30''E), 8.i.1978, Holm, Jacobs, Kirsten, Scholtz— 1 ♀, 1 ♂ (BMNH); Potchefstroom (26°43'S, 27°6'E), 6.xii.1913, [anonymous]— 1 ♀, 2 ♂♂ (BMNH); Vryburg (26°57'S, 24°44'E), 1893, E. Simon— 1 ♀, 1 ♂ (MNHN); Zeerust (25°32'S, 26°5'E), [no date], P. Born— 1 ♂ (IRSNB); NORTHERN CAPE, Farm Klipkoppies, 957 m (28°52'16.93''S, 21°18'33.98''E), 12–14.iii.2004, A.L.V. Davis & C. Deschodt— 1 ♂ (CMNC); J. Kempdorp (27°52'30''S, 24°52'30''E), 5.xi.1983, F. van Eeden—1 ♂ (BMNH); Koegrabie Farm (1) (29°4'33.6''S, 21°46'5.63''E), 6–8.iii.2004, A.L.V. Davis & C. Deschodt— 2 ♂♂ (PMOC); Pneil Road, 35 km W Kimberley (28°43'S, 24°21'E), 12–13.xii.1984, H. & A. Howden— 1 ♂ (CMNC); SE 29 23 Bb (29°7'30''S, 23°52'30''E), iv.1978, J.P. Krygsman— 1 ♂ (BMNH); WESTERN CAPE, Mossel Bay (34°11'S, 22°8'E), xii.1921, [anonymous]— 1 ♂ (BMNH); SWAZILAND: MANZINI, Mlilwane Wildlife Sanctuary (26°29'S, 31°9'E), 13.xii.1984, M. Davis— 1 ♀, 7 ♂♂ (CNC, FGIC); TANZANIA: Zanzibar, [no date], Raffray— 1 ♂ (IRSNB); ARUSHA, Arusha (3°23'S, 36°41'E), 11.iii.1995, [anonymous]— 1 ♀, 1 ♂ (OMOC); DAR ES SALAAM, Dar-es- Salaam (6°48'S, 39°17'E), iv.1924, W. A. Cutler— 1 ♂ (BMNH); Dar-es-Salaam (6°48'S, 39°17'E), [no date], R. v. Bennigsen— 1 ♂ (SMF); Dar-es-Salaam (6°48'S, 39°17'E), 2.iii.1925, F.W.H. Migeod— 1 ♂ (BMNH); Dar-es- Salaam (6°48'S, 39°17'E), 14.ix.1961, G. Heinrich— 1 ♀, 3 ♂♂ (CNC); Yombo (6°51'S, 39°14'E), 28.x.1992, W. R.B. Hynd— 2 ♂♂ (BMNH); IRINGA, Mkimbizi Mountain (7°46'12''S, 35°41'24''E), 28.i.2006, K. Angelus— 1 ♀, 1 ♂ (PMOC); Mkimbizi-Bima Mountain, 20.ii.2006, K. Angelus— 1 ♂ (PMOC); LINDI, Nachingwea, Mtokokuyana (10°23'51''S, 38°53'55''E), 21–24.xii.2009, Local collectors— 1 ♀, 1 ♂ (PMOC); MOROGO, Morogoro area (6°49'20''S, 37°39'40''E), 21.xi.1989, P. Holter & C. Sommer— 2 ♀♀, 1 ♂ (ZMUC); MOROGORO, Beho Beho Camp, Selous Game reserve (7°36'44''S, 37°50'23''E), 8–11.i.2001, T. Bouyer— 2 ♀♀, 2 ♂♂ (PMOC); Kisaki (7°29'S, 37°36'E), i–v.2011, Local collectors— 4 ♀♀, 5 ♂♂ (BMNH); Magubike, 787 m (8°18.049'S, 37°13.005'E), 20.ii.2010, P. Darge— 5 ♀♀, 5 ♂♂ (MHNL, PMOC); Mhonda-Ouzingua [= Station du Sacré-Coeur de Mhonda] (6°3'36''S, 37°21'36''E), 1879– i–iii.1880, A. Hacquard— 1 ♀, 1 ♂ (MNHN); Savane à l’O. de Mikesse, 525 m (6°43.752'S, 37°51.395'E), 29.iv.2005, R. Minetti— 2 ♀♀, 2 ♂♂ (PMOC); Uluguru Mountains (7°6'S, 37°39'E), 27–29.xi.2005, [anonymous]— 2 ♀♀, 2 ♂♂ (PMOC); PEMBA NORD, Ngezi Forest (4°55'35''S, 39°42'4''E), viii.1994, [anonymous]— 1 ♀ (JFJC); PWANI, 70 km E Morogoro, 300 m (6°38'S, 38°8.2'E), 12.xii.2006, F. Kantner— 2 ♀♀, 1 ♂ (SMNS); Bagamoyo (6°26'S, 38°54'E), [no date], H. Schaedle— 2 ♀♀, 2 ♂♂ (BMNH, MNHN); Mandera, 166 m (6°14.3'S, 38°23.197'E), 7.xii.2008, P. Darge— 2 ♀♀, 2 ♂♂ (MHNL); Mwetemo, 191 m (6°21.808'S, 38°30.852'E), 9.xii.2008, P. Darge— 2 ♀♀, 1 ♂ (MHNL); Route Dar-es-Salam- Chalinze, 40 m (6°40.739'S, 38°40.188'E), 1.v.2005, R. Minetti— 2 ♀♀, 2 ♂♂ (PMOC); Utende Base Camp, Mafia Island, 15 m (7°58'46''S, 39°44'31''E), 3.xii.1992, [anonymous]— 2 ♀♀, 3 ♂♂ (BMNH); TANGA, Tanga (5°5'30''S, 39°6'0''E), iv.1912, Alluaud & Jeannel— 1 ♂ (MNHN); Tanga (5°5'30''S, 39°6'0''E), [no date], Gierra— 2 ♂♂ (MNHN); Tanga (5°5'30''S, 39°6'0''E), iii.1916, Methner— 1 ♂ (NMPC); UGANDA: CENTRAL, Tero Forest (0°50'S, 31°40'E), 26–30.ix.1911, S. A. Neave— 1 ♂ (BMNH); UNITED STATES OF AMERICA: ARIZONA, Santa Cruz County, Harshaw Creek Road (31°30'53''N, 110°41'32''W), 10–18.viii.1990, L. Desaulniers— 1 ♂ (CMNC); FLORIDA, Alachua County, Paynes Prairies Preserve State Park (29°34'8''N, 82°18'3''W), 25.iv.1996, R. Vigneault— 1 ♀, 1 ♂ (PMOC); HAWAII, Honolulu County, Ewa, O’ahu (21°20'29''N, 158°2'14''W), 6.i.1961, D.F. Hardwick— 1 ♂ (CNC); Honolulu County, Ewa, O’ahu (21°20'29''N, 158°2'14''W), 10.ii.1961, D.F. Hardwick— 2 ♀♀, 3 ♂♂ (CNC); Honolulu (21°18'N, 157°51'W), 30.ix.1966, J.R. Vockeroth— 1 ♂ (CNC); Honolulu (21°18'N, 157°51'W), 1.xii.1966, J.R. Vockeroth— 2 ♀♀, 2 ♂♂ (CNC); Honolulu (21°18'N, 157°51'W), 27.xii.1966, J.R. Vockeroth— 2 ♀♀, 2 ♂♂ (CNC); Honolulu (21°18'N, 157°51'W), 20.i.1967, J.R. Vockeroth— 1 ♀, 1 ♂ (CNC); Ka’Ena Point, O’ahu (21°34'6''N, 158°14'53''W), 11.i.1960, [anonymous]— 2 ♂♂ (CNC); Makapuu Point, O’ahu (21°18'30''N, 157°39'5''W), 16.i.1961, D.F. Hardwick— 3 ♀♀ (CNC); Manoa, Honolulu (21°18'57''N, 157°48'12''W), 27.vii.1963, D.M. Tsuda & R.M. Young— 1 ♂ (MNHN); South end Waianae Mountains, 244 m (21°28'19''N, 158°7'36''W), 18.i.1961, D.F. Hardwick— 3 ♀♀, 3 ♂♂ (CNC); Wailupe Valley, 305 m (21°18'24''N, 157°45'12''W), 14.iii.1961, D.F. Hardwick— 1 ♀, 1 ♂ (CNC); Waipio, O’ahu (21°24'52''N, 157°59'44''W), 11.i.1961, D.F. Hardwick— 3 ♀♀ (CNC); Waipio, O’ahu (21°24'52''N, 157°59'44''W), 18.i.1961, D.F. Hardwick— 2 ♂♂ (CNC); West side Ka’Ena Point (21°34'28''N, 158°16'37''W), 14.i.1961, D.F. Hardwick— 1 ♀, 2 ♂♂ (CNC); West side Ka’Ena Point (21°34'28''N, 158°16'37''W), 21.i.1961, D.F. Hardwick— 4 ♀♀, 2 ♂♂ (CNC); Maui County, Hana (20°45'27''N, 155°59'18''W), 22.ii.2004, B. & M. Holme— 1 ♂ (CMNC); SOUTH CAROLINA, Chesterfield County, Hwy 151, approx. 10.7 mi. NW McBee (34°33'57''N, 80°22'1''W), 11.v.2002, J.P. Gruber— 1 ♂ (CMD); TEXAS, Medina County, Hondo (29°20'49''N, 99°8'44''W), 26.x.1982, B.F. & J.L. Carr— 1 ♂ (CNC); Presidio County, 10 km S Ruidosa, 800 m (29°45'45''N, 104°33'17''W), 16.viii.2007, Mann & Edmonds— 1 ♂ (OUMNH); Humphrey Ranch, approximately 30 km SE Marfa (hwy 169), 1250 m (30°2'34''N, 104°1'18''W), 10.viii.2007, Mann & Edmonds— 1 ♀ (OUMNH); Miller Ranch, approximately 20 km W Valentine, 1390 m (30°32'50''N, 104°39'40''W), 13.viii.2007, Mann & Edmonds— 1 ♀ (OUMNH); Presidio (29°34'N, 104°22'W), 2.x.1982, B.F. & J.L. Carr— 1 ♂ (CNC); Ruidosa (29°59'37''N, 104°40'42''W), 16.viii.2007, Mann & Edmonds— 1 ♂ (OUMNH); VANUATU: SHEFA, Epi (16°44'S, 168°16'E), 12.xii.2007, P. Moulin— 2 ♀♀, 2 ♂♂ (PMOC); ZAMBIA: CENTRAL PROVINCE, 25 km NE Lilemone, 1250 m (15°13'14''S, 26°19'41''E), 11.xii.2009, S. Rojkoff— 1 ♂ (PMOC); 6 km N Chunga, 1075 m (14°59'39''S, 26°1'11''E), 4.xii.2010, S. Rojkoff— 1 ♀, 1 ♂ (OMOC); 6.2 km W Mukambi Lodge Junction on M9, 1100 m (14°57'2''S, 25°56'21''E), 18.xi.2010, F. Génier (2010-02)— 10 ♀♀, 12 ♂♂ (FGIC); 6.2 km W Mukambi Lodge Junction on M9, 1100 m (14°57'2''S, 25°56'21''E), 4.xii.2010, F. Génier (2010-42)— 1 ♀, 2 ♂♂ (FGIC); 6.5 km N Chunga, 1100 m (14°59'40''S, 26°1'11''E), 4.xii.2010, F. Génier (2010-43)— 12 ♀♀, 12 ♂♂ (FGIC); Huntley Farm, Chisamba, 1122 m (15°0'33.57''S, 28°8'41.91''E), 26–27.xi.2005, S. Beynon & M. Jubb— 1 ♂ (OUMNH); Mayukuyuku Camp (14°54'58''S, 26°3'55''E), 13.xii.2009, Josso, Juhel, & Minetti— 1 ♀, 1 ♂ (JFJC); Mukambi Safari Lodge, 1250 m (14°58'32''S, 25°59'29''E), 8.xii.2010, F. Génier (2010-50)— 10 ♀♀, 10 ♂♂ (FGIC); Mukambi Safari Lodge, 1250 m (14°58'32''S, 25°59'29''E), 9.xii.2010, F. Génier (2010-52)— 14 ♀♀, 14 ♂♂ (FGIC); route principale, Kafue National Park (14°57'S, 25°52'E), 8.xii.2008, J.F. Josso & R. Minetti— 1 ♀, 1 ♂ (JFJC); route principale, Kafue National Park (14°57'S, 25°52'E), 16–17.xii.2008, J.F. Josso & R. Minetti— 1 ♂ (JFJC); Zambeef Farming Limited, 1122 m (15°0'33.57''S, 28°8'41.91''E), 25–26.xi.2005, S. Beynon & M. Jubb— 1 ♂ (OUMNH); Zambeef Farming Limited, 1122 m (15°0'33.57''S, 28°8'41.91''E), 26–27.xi.2005, S. Beynon & M. Jubb— 1 ♂ (OUMNH); Zambeef Farming Limited, 1122 m (15°0'33.57''S, 28°8'41.91''E), 28–29.xi.2005, S. Beynon & M. Jubb— 1 ♀, 1 ♂ (OUMNH); Zambeef Farming Limited, 1122 m (15°0'33.57''S, 28°8'41.91''E), 29– 30.xi.2005, S. Beynon & M. Jubb— 1 ♀, 1 ♂ (OUMNH); COPPERBELT, Chingola (12°33'S, 27°52'E), 20.xi.1989, [anonymous]— 1 ♀, 2 ♂♂ (CVM, OUMNH); Chingola (12°33'S, 27°52'E), 22.xi.1989, [anonymous]— 1 ♂ (OUMNH); Kishikishi, Rural Ndola (12°59'S, 28°39'E), [no date], Samukole— 1 ♂ (FETC); LUSAKA PROVINCE, 30 km E Lusaka (15°16'S, 28°56'E), 20.iii.1991, P. Schüle— 1 ♂ (SMNS); Kafue City - Kafue River, 1200 m (16°3'S, 28°30'E), 22.xi–2.xii.1987, R. Mourglia— 2 ♀♀, 2 ♂♂ (CVM); NORTH- WESTERN PROVINCE, Chimfunzi Nature Reserve, 1312 m (12°21.95'S, 27°28.93'E), 9–11.xi.2001, B. & J. Gill— 1 ♂ (BDGC); NORTHERN PROVINCE, Chocha (8°26'S, 29°48'E), 21.i.1944, H.-J. Brédo— 1 ♂ (IRSNB); Chocha (8°26'S, 29°48'E), 1.ii.1944, H.-J. Brédo— 3 ♂♂ (FGIC, IRSNB); Kaputa (8°28'13''S, 29°40'4''E), 3.ii.1944, H.-J. Brédo— 2 ♂♂ (IRSNB); Lake Mweru Mantipa (8°40'S, 29°36'E), [no date], H.-J. Brédo— 1 ♂ (IRSNB); Sumbu (8°31'30''S, 30°28'30''E), [no date], J. Duvivier— 1 ♂ (IRSNB); SOUTHERN PROVINCE, 12 km NE Choma (16°38'32''S, 27°1'47''E), 16.i.2010, P. Schüle— 2 ♀♀, 2 ♂♂ (PMOC); Bruce-Miller Game Farm, 18 km NNW Choma, 1170 m (16°38.22'S, 27°1.51'E), 6–8.xi.2001, B. & J. Gill— 4 ♀♀, 2 ♂♂ (BDGC); Bruce- Miller Game Farm, 18 km NNW Choma, 1170 m (16°38.22'S, 27°1.51'E), 22–24.xi.2001, B. & J. Gill— 2 ♀♀, 2 ♂♂ (BDGC); Bruce-Miller Game Farm, 18 km NNW Choma, 1170 m (16°38.22'S, 27°1.51'E), 26.xi–7.xii.2013, Josso, Juhel, & Minetti— 1 ♂ (FGIC); Livingstone, Zambesi River (17°51'S, 25°52'E), ii.1913, H.C. Dollman— 1 ♂ (BMNH); Livingstone, Zambesi River (17°51'S, 25°52'E), i.1913, H.C. Dollman— 1 ♂ (BMNH); Livingstone, Zambesi River (17°51'S, 25°52'E), 30.i.1913, H.C. Dollman— 1 ♂ (BMNH); Namwazi (15°57'S, 28°4'E), 1.vi.1916, H.C. Dollman— 1 ♀, 1 ♂ (BMNH); Namwazi (15°57'S, 28°4'E), 4.vi.1916, H.C. Dollman— 3 ♂♂ (BMNH); Nkwali Camp, South Luangwa GMA (13°7'3''S, 31°44'16''E), 10–18.xi.2012, R. Smith & H. Takano— 3 ♀♀, 2 ♂♂ (BMNH); WESTERN PROVINCE, 13 km W Sesheke, 1000 m (17°26'55''S, 24°10'15''E), 7.xii.2013, S. Rojkoff— 1 ♂ (CMD); Lealui (15°14'S, 23°1'E), 1919, V. Ellenberger— 3 ♀♀, 3 ♂♂ (MNHN); Lealui (15°14'S, 23°1'E), 1915, V. Ellenberger— 2 ♂♂ (MNHN); ZIMBABWE: BULAWAYO, Bulawayo (20°9'S, 28°35'E), [no date], [anonymous]— 1 ♀, 4 ♂♂ (BMNH, FGIC, IRSNB); HARARE, 15 km N Harare (17°37'8''S, 30°57'59''E), 24.xii.1995, A. Thilliez— 1 ♂ (PMOC); Harare (17°49'4''S, 31°2'41''E), 1989, G. Nazaret— 1 ♀, 1 ♂ (CMD); Salisbury, 1524 m (17°49'S, 31°3'E), xi.1898 – i.1899, G. A.K. Marshall— 3 ♀♀, 3 ♂♂ (OUMNH); Salisbury [= Harare] (17°49'4''S, 31°2'41''E), i.1906, G. A.K. Marshall— 1 ♀, 1 ♂ (BMNH); Salisbury [= Harare] (17°49'4''S, 31°2'41''E), [no date], G. A.K. Marshall— 1 ♀, 1 ♂ (BMNH); MANICALAND, 15 mi. N Umtali [= Mutare] (18°47'S, 32°41'E), 9.iv.1971, Bornemissza & Aschenborn— 1 ♂ (CMNC); Umtali [= Mutare] (18°58'S, 32°40'E), xii.1901, G. A.K. Marshall— 1 ♂ (BMNH); Umtali [= Mutare] (18°58'S, 32°40'E), iii.1957, N.L.H. Krauss— 5 ♂♂ (neotype) (ZMUK); MASHONALAND EAST, Enkeldoorn [= Chivhu] (19°1'S, 30°54'E), [no date], E.S. Youngs— 5 ♀♀, 6 ♂♂ (BMNH); Marandellas (18°11'S, 31°33'E), ii.1962, J.S. Weir— 2 ♀♀, 2 ♂♂ (BMNH); Marandellas (18°11'S, 31°33'E), 8.iv.1971, Bornemissza & Aschenborn— 1 ♂ (CMNC); MASHONALAND WEST, 20 km SE Karoi (16°58'39''S, 29°49'3''E), 10.xii.2005, S. Rojkoff & K. Werner— 2 ♂♂ (CMD, PMOC); Atlantica, 16 mi. W Harare (17°51'S, 30°47'E), i.1974, M.B. Fenton— 1 ♀, 1 ♂ (CMNC); MASVINGO, Bubi River, 95 km NE Beitbridge (21°42'26''S, 30°30'0''E), 12–27.xii.2005, P. Schüle— 1 ♀ (PMOC); Lake Kyle, Lake Mutirikwe (20°12'36''S, 31°2'29''E), 27.xi.2006, P. Schüle— 1 ♂ (PMOC); MATEBELELAND NORTH, Hwange National Park, Main Camp near pan 5 (18°44'S, 26°57'E), 10.xi.1961, J.S. Weir— 2 ♀♀, 3 ♂♂ (BMNH); Inyati (19°41'S, 28°51'E), i.1969, R. Nielson— 1 ♂ (BMNH); Lonely Mine (19°30'S, 28°45'E), [no date], A. Swale— 1 ♂ (BMNH); Lonely Mine (19°30'S, 28°45'E), 13.i.1913, H. Swale— 1 ♂ (BMNH); Lonely Mine (19°30'S, 28°45'E), 15.i.1914, H. Swale— 1 ♂ (BMNH); Lonely Mine (19°30'S, 28°45'E), 3.xii.1914, H. Swale— 2 ♂♂ (BMNH); Wankie Nat. Park, Pan 0.8 (18°45'S, 26°0'E), xi.1961, J.S. Weir— 1 ♂ (BMNH); MATEBELELAND SOUTH, 20 km W Gwanda, 120 km SE Bulawayo (20°58'19''S, 28°50'13''E), 6.xii.1999, Kantner— 2 ♀♀, 2 ♂♂ (SMNS); Matopos, 35 km S Bulawayo (20°30'20''S, 28°26'11''E), 31.i.1996, F. Kantner— 2 ♀♀, 2 ♂♂ (SMNS); MIDLANDS, Ngezi Recreational Park, 60 km E Kwekwe (18°41'S, 30°24'E), 1.xii.1998, F. Kantner— 2 ♀♀, 1 ♂ (SMNS).

Natural history. In fact, very little is known about the natural history and ecology of D. gazella in the field within its native distribution. The main contributions were performed in controlled environments and were part of “The Australian dung beetle project” (Bornemissza 1976) initiated to introduce D. gazella in Australia. A second series of studies were done for a subsequent introduction in continental United States of America. For more information, Bornemissza (1970, 1976), Blume & Aga (1975, 1978), Fincher et al. (1986), Gutierrez et al. (1988) are among more than 200 applied entomology publications related to “ O. gazella ”. The Rougon & Rougon (1980, 1983) studies on the nesting behavior and the morphology of the larvae applies to a different Sahelian species, presumably D. eucatta.

Nomenclature and taxonomy. An application was submitted to the International Commission of Zoological Nomenclature on 20 October 2016 (Génier & Krell 2017) and was assigned case number 3722. The application is made to maintain prevailing usage of the species name D. gazella for the widely introduced species through the process of setting aside all previous type fixations for the nominal species Scarabaeus gazella Fabricius and the designation of a neotype.

The circumstances with the type material of Scarabaeus gazella are more intricate. In this case we are in presence of three putative syntypes. Unlike the syntypes of S. catta, these three specimens all bear a small green square typical of Fabrician types (Zimsen 1964). There is a well-developed male (10.5 mm), a small male (8.0 mm) with secondary sexual characters resembling a female, and a moderately large female (10.0 mm). The large male and the female bear a handwritten label with the following information: “ Guinea, Isert, Mus. T. Lund, Copris gazella F.” These labels were added at least 14 years after the species was originally described, as the generic name used on these labels is Copris, a genus Fabricius used for the first time in Fabricius (1801). After a careful study of these three specimens, which include the dissection and examination of the internal sac sclerites of the two males, we conclude that this series is a mixture of two species. The large male is a representative of the oriental species D. catta and the small male and female are representative of the West African species D. falciger. At this point, there is no realistic possibility of finding out the true historical details about these specimens. The most likely chain of events is that Fabricius had only the large male specimen lacking geographic information when he described the species in the appendix of the second volume of Mantissa Insectorum published in December 1787 (Evenhuis 1997; Bousquet 2016). The original description is clear: “occipite cornubus duobus arcuatis”, and can only apply to the specimen with two well-developed cephalic horns. We can only speculate on the number of specimens seen by Fabricius at the time of the description. Subsequently, Fabricius had received two additional specimens from Tønder Lund collected in Africa (Guinea), which he identified as his S. gazella and added the new information on the distribution five years later in the Entomologia systematica (Fabricius 1792).

The currently recognized synonymy for D. gazella was established by d’Orbigny (1913). The first species listed is Scarabaeus catta Fabricius, the entry is preceded by a question mark indicating that this synonymy is dubious. There is, however, no indication as to why d’Orbigny considered this synonymy dubious. This synonymy is here resolved on the principle of priority and the name D. catta is valid for an oriental species. The second synonym listed is Scarabaeus dorcas Olivier, described from Madagascar. This name is as a synonym of D. gazella assuming that the International Commission on Zoological Nomenclature rules on the conservation of the name D. gazella for the widely introduced species. The third species listed is Copris metallicus Fabricius, described from India. The holotype (by monotypy) of the species has been studied and despite the fact that it is missing the head and prothorax, it can be stated here that it does not belong to the genus Digitonthophagus. It is an Onthophagus of the Onthophagus orientalis species group and most likely belonging to the subgenus Sinonthophagus Kabakov.

The species is therefore here removed from synonymy with D. gazella and considered valid with the following combination Onthophagus metallicus (Fabricius). It should be noted here that a junior homonym, Onthophagus metallicus Fåhraeus, exists as a synonym of Proagoderus bicallosus (Klug). The fourth species listed is Copris antilope Fabricius, also described from India. Three syntypes are in ZMUC and an additional specimen in the ZMUK. The examination of the specimens revealed that they do not belong to the genus Digitonthophagus but rather belong to the genus Onthophagus and are similar to Onthophagus (Colobonthophagus) dama (Fabricius). The species is therefore here removed from synonymy with D. gazella and considered valid with the following combination Onthophagus antilope (Fabricius). Again, it should be noted here that a junior homonym, Onthophagus antilope Motschulsky exists, the latter is a synonym of Onthophagus (Palaeonthophagus) vacca Linnaeus. It is beyond the scope of this work to determine if O. antilope is synonymous with O. dama. The fifth name listed as synonyms is Onthophagus intermedius Reiche with a type locality “ Sénégal.” This name was first listed as “ O. gazella var. intermedius Dupont ” a variety of Onthophagus gazella in Dejean (1836) and is a nomen nudum. A specimen of D. sahelicus with a small manuscript label stating “1395” and a second manuscript label stating, “ intermedius Reiche. gazella? Fab. Senegal ” was found in the BMNH collection. It is not clear if this is a specimen acquired from Dupont’s collection by The Natural History Museum of London. Reiche (1840) stated that Dupont, who was a prominent natural history specimen merchant in the 19th century (Cambefort 2006), was the first to use the name O. intermedius for specimens in his collection. The name O. intermedius Reiche is, however, unavailable as it was first published in an available work as a synonym of “ Onthophagus gazella ” (Article 11.6; International Commission on Zoological Nomenclature 1999).