Digitonthophagus gazella (Fabricius, 1787)

Génier, François & Moretto, Philippe, 2017, Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae), Zootaxa 4248 (1), pp. 1-110: 39-52

publication ID

10.5281/zenodo.439444

publication LSID

lsid:zoobank.org:pub:71F0AC03-C1FB-40AB-8532-99A638FC91E9

persistent identifier

http://treatment.plazi.org/id/3D5987B7-0759-FFA0-FF17-A71F033CFCF1

treatment provided by

Plazi

scientific name

Digitonthophagus gazella (Fabricius, 1787)
status

 

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

( Figs. 14View FIGURES 11 – 14, 37–38View FIGURES 31 – 38. 31, 33, 35, 37, 62View FIGURES 55 – 62, 78View FIGURES 71 – 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 ( DNAAbout 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. 62View FIGURES 55 – 62); FLP sclerite subapicodorsal lobe short anteriorly ( Figs. 156– 157View FIGURES 147 – 158); 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. 185View FIGURES 183 – 190 A), lacking well-sclerotized projection on apicoventral edge ( Fig. 185View FIGURES 183 – 190 B); axial sclerite shorter than subaxial sclerite, acute apically ( Fig. 156View FIGURES 147 – 158); subaxial sclerite moderately long and robust ( Fig. 153View FIGURES 147 – 158).

Description. Male specimen from Central Province, Zambia ( Fig. 14View FIGURES 11 – 14). Measurements. Length 12.5 mm, width 7.0 mm. Head ( Figs. 37–38View FIGURES 31 – 38. 31, 33, 35, 37). 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. 62View FIGURES 55 – 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. 14View FIGURES 11 – 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. 126View FIGURES 119 – 134). Parameres with dorsal and ventral edges slightly diverging toward apex in lateral view. Internal sac sclerites ( Figs. 156–158View FIGURES 147 – 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. 78View FIGURES 71 – 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 ♂ ( MNHNAbout 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, MNHNAbout 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 ♂ ( MNHNAbout MNHN)GoogleMaps  ; HUILA, 10 km S Caconda (13°50'S, 15°6'E), 5.xi.2011, G. Werner— 2 ♀♀, 2 ♂♂ ( FGICAbout FGIC, GWPC)GoogleMaps  ; 10 km S Cacula (14°35'S, 14°3'E), 4.xi.2011, G. Werner— 1 ♂ ( GWPC)GoogleMaps  ; 30 km W Quipungo (14°48.02'S, 14°15.32'E), 26.xi.2013, G. Werner— 3 ♀♀, 2 ♂♂ ( FGICAbout FGIC, GWPC)GoogleMaps  ; 33 km S Cacula (14°41'S, 13°54'E), 13.xi.2011, G. Werner— 1 ♀, 1 ♂ ( GWPC)GoogleMaps  ; Capelongo près Luceque (14°39'S, 15°4'E), xii.1914, Mission Rohan-Chabot— 1 ♂ ( MNHNAbout MNHN)GoogleMaps  ; Capelongo-Dongo (14°31'S, 16°1'E), 1914, Mission Rohan-Chabot— 1 ♀ ( MNHNAbout MNHN)GoogleMaps  ; Capelongo-Dongo (14°31'S, 16°1'E), xii.1914, Mission Rohan-Chabot— 2 ♂♂ ( MNHNAbout MNHN)GoogleMaps  ; 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)GoogleMaps  ; Gilgandra (31°43'S, 148°40'E), 12.iv.1988, H. & A. Howden— 1 ♂ ( CMNC)GoogleMaps  ; Narrabri (30°20'S, 149°47'E), 5.xii.1977, G.G. Burns— 2 ♀♀ ( CMNC)GoogleMaps  ; NORTHERN TERRITORY, CSIROAbout CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 1–10.xii.1993, S. & J. Peck (93-85)— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; CSIROAbout CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 5.i.1994, S. & J. Peck (93-166)— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; CSIROAbout CSIRO McMillans Road, Darwin (12°24'42''S, 130°55'19''E), 9.i.1994, S. & J. Peck (93-167)— 1 ♂ ( CMNC)GoogleMaps  ; Elsey National Park, Mataranka (14°58'S, 133°11'E), 9.xii.1993, S. & J. Peck (93- 115)— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Elsey National Park, Mataranka (14°58'S, 133°11'E), 9.xii.1993, S. & J. Peck (93- 116)— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; 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)GoogleMaps  ; Magela Creek (12°40'S, 133°1'E), 4.xi.1989, C.W. & L.B. O'Brien— 1 ♂ ( CMNC)GoogleMaps  ; Tenat Creek (19°39'S, 134°11'E), 22.ix.1988, H. & A. Howden— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; QUEENSLAND, 12 mi. S Ayr (19°44'S, 147°28'E), 26.ii.1975, H. & A. Howden— 1 ♂ ( CMNC)GoogleMaps  ; 16 km W Ravenshoe (17°39'S, 145°21'E), 2.i.1975, M.S. & B.J. Moulds— 1 ♂ ( CMNC)GoogleMaps  ; 4–8 km S Atherton (17°20'S, 145°30'E), 26.xii.1988, H. & A. Howden— 1 ♂ ( CMNC)GoogleMaps  ; 40 km N Lynd (18°36'S, 144°43'E), ii.1993, H. Demarz— 1 ♂ ( CEMT)GoogleMaps  ; 8 km N Mareeba (16°55'S, 145°25'E), 28.i.1997, H. & A. Howden— 1 ♀ ( CMNC)GoogleMaps  ; 9 km NE Julatten (16°33'S, 145°23'E), 13.xii.1986, H. & A. Howden— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Atherton (17°16'S, 145°29'E), 7.ii.1975, H. & A. Howden— 1 ♀ ( CMNC)GoogleMaps  ; Atherton (17°16'S, 145°29'E), 10.ii.1975, H. & A. Howden— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Black Beach Reserve, near Mackay (21°4'39''S, 149°11'44''E), 28.i.1975, H. & A. Howden— 2 ♂♂ ( CMNC)GoogleMaps  ; Burketown (17°44'S, 139°33'E), 20.ix.1975, Walford-Huggins— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Cairns (16°55'S, 145°46'E), 10.v.1971, A. & M. Walford-Huggins— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Cairns (16°55'S, 145°46'E), 24.viii.1971, A. & M. Walford-Huggins— 1 ♀, 1 ♂ ( CMNC)GoogleMaps  ; Cairns (16°55'S, 145°46'E), 30.i.1975, H.GoogleMaps 

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

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 ZMUCAbout 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).

DNA

Department of Natural Resources, Environment, The Arts and Sport

MNHN

Museum National d'Histoire Naturelle

FGIC

Francois Genier

CSIRO

Australian National Fish Collection

CNC

Canadian National Collection of Insects, Arachnids, and Nematodes

UFMT

Universidade Federal de Mato Grosso

IRSNB

Institut Royal des Sciences Naturelles de Belgique

SMF

Forschungsinstitut und Natur-Museum Senckenberg

SMNS

Staatliches Museum fuer Naturkund Stuttgart

CVM

City Museum

ZMUC

Zoological Museum, University of Copenhagen

MHNL

Musee Guimet d'Histoire Naturelle de Lyon

NMPC

National Museum Prague

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Coleoptera

Family

Scarabaeidae

Genus

Digitonthophagus

Loc

Digitonthophagus gazella (Fabricius, 1787)

Génier, François & Moretto, Philippe 2017

2017
Loc

Digitonthophagus gazella

Genier 2017: 497Genier 2017: 498
Breeschoten 2016: 90

2016
Loc

Digitonthophagus gazella

Mlambo 2015: 318
Whipple 2012: 45
Tarasov 2011: 9

2011
Loc

Digitonthophagus gazella

Noriega 2011: 35
Noriega 2010: 451

2010
Loc

Digitonthophagus gazella

Wirta 2008: 1079

2008
Loc

Digitonthophagus gazella

Ivie 2008: 10
Moretto 2007: 122
Monaghan 2007: 678
Noriega 2006: 379
Cambefort 2003: 560

2003
Loc

Digitonthophagus gazella

Noriega 2002: 213

2002
Loc

Onthophagus gazella

MacRae 2001: 49

2001
Loc

Digitonthophagus gazella

Deloya 2000: 125
Barbero 1998: 242
Kim 1989: 336

1998
Loc

Onthophagus gazella

Ripa 1995: 438

1995
Loc

Digitonthophagus gazella

Kohlmann 1994: 35
Zunino 1988: 17

1988
Loc

Onthophagus gazella

Gutierrez 1988: 39
Paulian 1986: 107

1986
Loc

Onthophagus gazella

Fincher 1986: 2
Rougon 1983: 499
Zunino 1981: 414
Rougon 1981: 6
Rougon 1980: 381
Blume 1978: 190

1978
Loc

Onthophagus gazella

Endrodi 1976: 156

1976
Loc

Onthophagus gazella

Alfieri 1976: 205

1976
Loc

Onthophagus gazella

Bornemissza 1976: 7

1976
Loc

Onthophagus gazella

Blume 1975: 735

1975
Loc

Onthophagus gazella

Endrodi 1974: 3

1974
Loc

Onthophagus gazella

Endrodi 1973: 125

1973
Loc

Onthophagus (O.) gazella

Endrodi 1973: 198

1973
Loc

Onthophagus (Digitonthophagus) gazella

Cambefort 1972: 246Ferreira 1972: 780Ferreira 1972: 780Balthasar 1972: 22

1972
Loc

Onthophagus (O.) gazella

Endrodi 1971: 294

1971
Loc

Onthophagus (Digitonthophagus) gazella

Bornemissza 1970: 31
Balthasar 1969: 68

1969
Loc

Onthophagus dorcas

Ferreira 1968: 593Ferreira 1968: 593

1968
Loc

Onthophagus (Digitonthophagus) gazella

Ferreira 1967: 1169
Balthasar 1963: 365
Balthasar 1963: 365

1963
Loc

Onthophagus gazella

Ferreira 1962: 153

1962
Loc

Onthophagus gazella

Frey 1961: 90Petrovitz 1961: 102

1961
Loc

Onthophagus gazella

Ferreira 1959: 84

1959
Loc

Onthophagus gazella

Ferreira 1958: 512

1958
Loc

Onthophagus gazella

Frey 1958: 73

1958
Loc

Onthophagus gazella

Gomes 1956: 94Frey 1956: 654
Paulian 1948: 151

1948
Loc

Onthophagus gazella

Gomes 1944: 3
Muller 1942: 96

1942
Loc

Onthophagus gazella

Balthasar 1941: 110
Boucomont 1937: 131
Paulian 1936: 132
Boucomont 1934: 26
Arrow 1931: 230
Gillet 1927: 138
Gillet 1927: 138

1927
Loc

Onthophagus (O.) gazella

Boucomont 1923: 96
d'Orbigny 1916: 11

1916
Loc

Onthophagus (O.) gazella

d'Orbigny 1914: 654

1914
Loc

Onthophagus (O.) gazella

Kolbe 1914: 297
d'Orbigny 1913: 249
d'Orbigny 1913: 96

1913
Loc

Onthophagus gazella

d'Orbigny 1911: 20

1911
Loc

Onthophagus gazella

d'Orbigny 1909: 525

1909
Loc

Onthophagus (O.) gazella

Kolbe 1908: 128
d'Orbigny 1905: 494
d'Orbigny 1905: 445
d'Orbigny 1904: 304

1904
Loc

Onthophagus gazella

Peringuey 1901: 181
Gahan 1900: 24
d'Orbigny 1900: 218
d'Orbigny 1900: 299

1900
Loc

Onthophagus gazella

Kolbe 1897: 83

1897
Loc

Onthophagus gazella

Fairmaire 1893: 10

1893
Loc

Onthophagus gazella

Brancsik 1893: 223

1893
Loc

Onthophagus gazella

Kolbe 1897: 150
Reitter 1893: 62

1893
Loc

Onthophagus gazella

Reitter 1892: 183Peringuey 1892: 29

1892
Loc

Scarabaeus

Wallengren 1881: 21
Gerstaecker 1873: 130
Gerstaecker 1873: 130

1873
Loc

Onthophagus gazella

Harold 1871: 119

1871
Loc

Onthophagus gazella

Harold 1869: 1029

1869
Loc

Scarabaeus

Gerstaecker 1871: 50
Harold 1869: 1029

1869
Loc

Onthophagus gazella

Harold 1867: 49
Klug 1862: 232

1862
Loc

Onthophagus gazella

Motschulsky 1860: 153

1860
Loc

Onthophagus

Fahraeus 1857: 275
Bertoloni 1849: 415

1849
Loc

Ontophagus gazella

Erichson 1843: 208

1843
Loc

Ontophagus gazella

Reiche 1840: 243

1840
Loc

Ontophagus gazella

Sturm 1843: 107
Laporte 1840: 85

1840
Loc

Onthophagus

Hope 1837: 33

1837
Loc

Onthophagus

Dejean 1836: 157

1836
Loc

Onthophagus

Klug 1835: 33

1835
Loc

Scarabaeus

Klug 1833: 163

1833
Loc

Onthophagus bonasus

Klug 1833: 163
LePeletier 1828: 354
Latreille 1827: 282

1827
Loc

Onthophagus

Sturm 1826: 178

1826
Loc

Onthophagus

Dejean 1821: 53

1821
Loc

Copris

Schonherr 1806: 48
Illiger 1804: 149

1804
Loc

Copris

Fabricius 1801: 47

1801
Loc

Scarabaeus

Illiger 1800: 228

1800
Loc

Scarabaeus

Fabricius 1801: 44
Illiger 1800: 228

1800
Loc

Copris

Fabricius 1798: 31

1798
Loc

Scarabaeus gazella

Fabricius 1792: 56

1792
Loc

Scarabaeus Gazella Fabricius, 1787b : 377

Gmelin 1790: 1537Olivier 1789: 121Fabricius 1787: 377

1790