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Sarophorus bidentatus Frolov & Scholtz, 2003: 187 Type locality: Namaqualand, Kamieskroon [30°12’S 17°56’E, Northern Cape, South Africa]. Distribution: South Africa (Frolov & Scholtz 2003; Davis et al. 2020)., Published as part of Strümpher, Werner P., Daniel, Gimo M. & Davis, Adrian L. V., 2022, Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 396-400 in Zootaxa 5188 (4) on page 399, DOI: 10.11646/zootaxa.5188.4.7, http://zenodo.org/record/7095688, {"references":["Frolov, A. V. & Scholtz, C. H. (2003) Revision of the Afrotropical dung beetle genus Sarophorus Erichson (Coleoptera: Scarabaeidae). African Entomology, 11, 183 - 198.","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeinae dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute (SANBI), Pretoria, 800 pp. [http: // opus. sanbi. org / handle / 20.500.12143 / 7672]"]}

Sarophorus tuberculatus (Laporte de Castelnau, 1840: 88) Type locality: Afrique [Africa]. Distribution: South Africa (Frolov & Scholtz 2003; Davis et al. 2020)., Published as part of Strümpher, Werner P., Daniel, Gimo M. & Davis, Adrian L. V., 2022, Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 396-400 in Zootaxa 5188 (4) on page 399, DOI: 10.11646/zootaxa.5188.4.7, http://zenodo.org/record/7095688, {"references":["Frolov, A. V. & Scholtz, C. H. (2003) Revision of the Afrotropical dung beetle genus Sarophorus Erichson (Coleoptera: Scarabaeidae). African Entomology, 11, 183 - 198.","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeinae dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute (SANBI), Pretoria, 800 pp. [http: // opus. sanbi. org / handle / 20.500.12143 / 7672]"]}

Sarophorus frolovi Roets, 2017: 268 Type locality: Midlands, Weza forestry plantations, –30.61680[°] 29.683626[°] (KwaZulu-Natal, South Africa). Distribution: South Africa (Roets et al. 2017; Davis et al. 2020)., Published as part of Strümpher, Werner P., Daniel, Gimo M. & Davis, Adrian L. V., 2022, Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 396-400 in Zootaxa 5188 (4) on page 399, DOI: 10.11646/zootaxa.5188.4.7, http://zenodo.org/record/7095688, {"references":["Roets, F., Crous, C. & Pryke, J. (2017) Sarophorus diabolus sp. n. and Sarophorus frolovi sp. n. (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology, 25 (1), 264 - 270. https: // doi. org / 10.4001 / 003.025.0264","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeinae dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute (SANBI), Pretoria, 800 pp. [http: // opus. sanbi. org / handle / 20.500.12143 / 7672]"]}

Sarophorus carinatus Frolov & Scholtz, 2003: 187 Type locality: Lydenburg Distr., Ohrigstad [24°45’S 30°34’E, Mpumalanga, South Africa]. Distribution: South Africa (Frolov & Scholtz 2003; Davis et al. 2020)., Published as part of Strümpher, Werner P., Daniel, Gimo M. & Davis, Adrian L. V., 2022, Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 396-400 in Zootaxa 5188 (4) on page 399, DOI: 10.11646/zootaxa.5188.4.7, http://zenodo.org/record/7095688, {"references":["Frolov, A. V. & Scholtz, C. H. (2003) Revision of the Afrotropical dung beetle genus Sarophorus Erichson (Coleoptera: Scarabaeidae). African Entomology, 11, 183 - 198.","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeinae dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute (SANBI), Pretoria, 800 pp. [http: // opus. sanbi. org / handle / 20.500.12143 / 7672]"]}

Sarophorus diabolus Roets, 2017: 266 Type locality: Riebeeck-Kasteel, Porseleinberg, –33.45995[°], 18.88627[°] (Western Cape, South Africa). Distribution: South Africa (Roets et al. 2017; Davis et al. 2020)., Published as part of Strümpher, Werner P., Daniel, Gimo M. & Davis, Adrian L. V., 2022, Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 396-400 in Zootaxa 5188 (4) on page 399, DOI: 10.11646/zootaxa.5188.4.7, http://zenodo.org/record/7095688, {"references":["Roets, F., Crous, C. & Pryke, J. (2017) Sarophorus diabolus sp. n. and Sarophorus frolovi sp. n. (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology, 25 (1), 264 - 270. https: // doi. org / 10.4001 / 003.025.0264","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeinae dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute (SANBI), Pretoria, 800 pp. [http: // opus. sanbi. org / handle / 20.500.12143 / 7672]"]}

Following its original description, Kheper namibicus Krajcik, 2006, was synonymized with Kheper cupreus (Laporte de Castelnau, 1840) by Deschodt et al. (2011) but later re-erected as a valid species or subspecies (Krajcik 2020). This paper discusses the evidence for validity versus synonymy and provides further support that Kheper namibicus is, indeed, a junior subjective synonym of Kheper cupreus. The description of Kheper namibicus was based on three specimens, the holotype and a paratype from near Gobabis [S22.45° E18.97°] and a paratype from near Otavi [S19.63° E17.33°] in Namibia (Fig. 1). According to Krajcik (2006), K. namibicus is close to K. cupreus (Fig. 2) but differs from it in (1) geographical distribution, (2) the “non-metallic colour of [the] dorsal side”, (3) the “non-lustrous elytral suture (in day light)”, (4) the darker colour than K. cupreus, (5) denticles of the front tibiae that are “not projected, rather obtuse” and (6) the shape of the parameres. Although the present authors have been unable to examine the holotype of K. namibicus, the photographs in Krajcik (2006) are sufficiently clear for comparisons with other specimens. The synonymy of K. namibicus with K. cupreus (Deschodt et al. 2011) was based on a reference collection containing individuals identified as K. cupreus from more than 15 localities across northern Namibia and more than 30 localities to the east (Figs 1A, 1B). Deschodt et al. (2011) observed that Gobabis and Otavi fell well within the known range of K. cupreus and that northern Namibian specimens were darker than the cupreous individuals found further east. With regards to type material of K. namibicus, they also explained that the “not projected” or “obtuse” condition of denticles on the front tibiae resulted from wear during soil excavation. Furthermore, the aedeagus of the holotype was comparable to a cupreous specimen from Kariba, Zimbabwe (16°31’ S, 28°46’ E) determined as K. cupreus by Richard zur Strassen, a specialist in the tribe Scarabaeini. Subsequently, at the end of a paper dealing primarily with insects in famous artworks of ancient Egypt, Krajcik (2020) provided a seven line re-evaluation of K. namibicus and, again, elevated it to valid species or subspecies status. As regards this re-evaluation, an English translation from Czech follows in inverted commas (Google translation services, 27 May 2022): “In 2006, a description of a new species of the genus Kheper from Namibia was published, which was named Kheper namibicus (Krajcik 2006, Animma.x, 14: 21). Deschodt et al. (2011) synonymized this species with cupreus. Apparently K. cupreus, as imagined by Deschodt and his colleagues, is in fact different from what was described by Laporte de Castelnau in 1840, and therefore K. namibicus should be recognized by a valid taxon (species or subspecies)”. Notably, Deschodt et al. (2011) was published without our having examined type specimens of Kheper cupreus or photographs of types. This has now been rectified by an examination of photographs of a syntype habitus sent from the Muséum national d’Histoire naturelle in Paris. The specimen should be considered a syntype (Fig. 2A) as it is unclear if Laporte de Castelnau’s original description was based on a single or multiple individuals. However, staff of the museum have been unable to locate other specimens that might belong to a type series. As Kheper species show little, external, sexual dimorphism, dissection by museum staff has shown that the syntype is a female. We have compared the habitus image of the female Caffraria syntype (cupreous with worn tibial and clypeal dentition) with that of the male K. namibicus holotype from Namibia (dark with a cupreous sheen and worn tibial/clypeal dentition) (Krajcik 2006, 2020), that of the male K. cupreus specimen from Kariba in Zimbabwe (cupreous with unworn fore tibial/ clypeal dentition) (Deschodt et al. 2011) and those of male specimens that we determined as K. cupreus comprising a dark specimen with unworn fore tibial/clypeal dentition from 20km south of Opuwo (S18.2027° E13.81305°) in Namibia as well as a cupreous specimen with unworn tibial/clypeal dentition from Orapa to the east in Botswana. All images show close similarities. Therefore, this comparison suggests that, contrary to Krajcik (2020), we have correctly understood the identity of K. cupreus as originally described in only twelve, half-column lines (78 words) by Laporte de Castelnau (1840). Furthermore, images of the aedeagi extracted from the Opuwo (dark), Orapa (cupreous), Kariba (cupreous) and K. namibicus holotype (dark with cupreous sheen) specimens (Krajcik 2006; Deschodt et al. 2011, Fig. 3) are clearly identical. Therefore, after a consideration of (1) the close similarity between the habitus of the five compared specimens representing the two colour varieties (K. cupreus syntype, K. namibicus holotype, Namibia, Botswana and Zimbabwe K. cupreus), (2) the identical aedeagi from K. namibicus and three western and eastern specimens identified as K. cupreus and (3) the identical aedeagi from dark worn (K. namibicus) and dark unworn (K. cupreus) specimens from Namibia, we confirm the synonymy of Kheper namibicus Krajcik, 2006, with Kheper cupreus (Laporte de Castelnau, 1840)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2022, Comparison with a syntype of Kheper cupreus (Laporte de Castelnau, 1840) confirms that Kheper namibicus Krajcik, 2006 is a synonym, pp. 193-196 in Zootaxa 5169 (2) on pages 193-195, DOI: 10.11646/zootaxa.5169.2.8, http://zenodo.org/record/6920241, {"references":["Krajcik, M. (2006) New species of Kheper Janssens from Namibia. Animma. x, 14, 19 - 23.","Laporte de Castelnau, F. L. (1840) Histoire naturelle des Insectes Coleopteres. Histoire naturelle des Animaux articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Vol. 2. P. Dumenil, Paris, 565 pp. https: // doi. org / 10.5962 / bhl. title. 35290","Deschodt, C. M., Davis, A. L. V. & Scholtz, C. H. (2011) New dung beetle taxa from arid south-western Africa. Insect Systematics and Evolution, 42, 277 - 294. https: // doi. org / 10.1163 / 187631211 X 568173","Krajcik, M. (2020) Hmyz jako predloha vyznacnych umeleckych del (specialni vyber). I. Posvatni brouci starovekeho Egypta. Insects in famous artworks (special selection). I. Sacred beetles of ancient Egypt. Animma. x, 82, 1 - 20.","Ferreira, M. C. (1972) Os escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11, 5 - 1088.","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086"]}

Stiptopodius singularis species group Differential diagnosis. The Stiptopodius singularis species group differs from the Stiptopodius doriae species by having the sides of the pronotum strongly sinuate before the postero-lateral angle in lateral view., Published as part of Daniel, Gimo M., Deschodt, Christian M., Davis, Adrian L. V. & Sole, Catherine L., 2020, The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa, pp. 394-404 in Zootaxa 4763 (3) on page 399, DOI: 10.11646/zootaxa.4763.3.4, http://zenodo.org/record/3759527

Stiptopodius savuti Daniel & Deschodt, new species (Figs. 1B, 4) Type locality: Savuti, North West, Botswana. Type material. Holotype, female: Savuti [Botswana, North West], Site 4 | S18.53092 [°]; E24.08378 [°] | 15.xii. 2005 | Deschodt & Tshikae | Carrion pitfall trap || HOLOTYPE: Stiptopodius | savuti Daniel & Deschodt | 2020 [printed on the red card] (TMSA). Description. Holotype, female. Length: 4.3 mm, width: 1.9 mm. Colour. Dorsally black; lateral edge of clypeus, legs, and last abdominal ventrite brown; setae yellowish. Head. Incision between medial clypeal teeth deeply concave; latero-clypeal edges not sinuate; genae with external protrusion sharply triangular; surface of the head smooth and punctate; ocellate punctures sparse on the middle of vertex; some large punctures with outer edges moderately raised on clypeus, otherwise, finely punctate; clypeo-frontal carina not visible; clypeal-genal suture present; vertex with carina occupying half the space between eyes, carina interrupted in the middle clearly forming two small cephalic tubercles. Antennae with eight antennomeres, labium setose. Pronotum. Convex, wider than long; base of pronotum marginate; anterior margin wider than posterior edge; in lateral view, lateral edge strongly sinuate before postero-lateral angles, central disc glabrous; punctation ocellate with sparse, short setae antero-laterally that emanate from centres of punctures. Elytra. Laterally straight, somewhat convex distally and concave proximally; sparsely and finely punctate in external-most four striae, the rest apunctate; interstriae with sparse, short setae visible basally, only interstriae V–VIII punctate. Pygidium. Convex with dense ocellate punctation bearing setae emanating from centres of punctures. Abdominal ventrites. Prosternum with anterior angles slanted forward and inwards, surface somewhat excavated. Mesoventrite dull, setose and punctate. Metaventrite shiny with long setae anteriorly, glabrous mid-posteriorly. Abdominal ventrites glabrous medially, shiny laterally, setigerous with ocellate punctures, setae arranged in lateral rows; last ventrites wider medially than others (female character). Legs. External and internal margins of profemora punctate, with well-developed setae, central area apunctate and glabrous; external and internal edges strongly and slightly marginate, respectively. Meso- and metafemora with external and internal edges sparsely punctate, all punctures bearing setae (shorter than those in profemora), surface between external and internal edges glabrous, strongly marginate internally. Protibiae crenulate with four teeth on external margin, a line of setae around teeth in dorsal view; in ventral view, protibiae with two carinae, internally punctate and pilose. Protibial spur long and inward curving. Meso- and metatibiae triangular and strongly widened apically. Protarsi broken off in both legs. Meso- and metatarsomeres I–IV triangular, widened apically, wider than long, strongly concave medially, external and internal edges with distal lobe; meso- and metatarsomere V rectangular and longer than others, with pair of very tiny apical claws. Male. Unknown. Distribution. This species is only known from the type locality Savuti in Botswana. Etymology. The specific name, a noun in apposition, refers to the name of the type locality. Diagnosis. Stiptopodius savuti new species is morphologically similar to S. granulosus. However, S. savuti new species can be distinguished by having the dorsal cephalic and pronotal surface smooth while in S. granulosus they are densely rugose. The new species bears a carina which is interrupted medially and clearly forms two small cephalic tubercles, unlike S. granulosus, whose cephalic carina does not form tubercles. Furthermore, the head of the new species is glabrous, whereas in S. granulosus it is setigerous. Remarks: The Afrotropical Stiptopodius savuti new species seems to be similar to the brown Somalian species of Stiptopodius glabricollis Müller, 1942. Our knowledge on Afrotropical dung beetle biogeographical distribution patterns suggests that it is very unlikely that the inland species from Botswana could be a sub-population of the coastal eastern horn of Africa species, S. glabricollis (see also a similar case in Génier & Josso 2017: 338). As such a wide and disjunct distribution is very rare among Afrotropical Scarabaeinae, we therefore consider the specimen from Savuti (Botswana) as a new species. Stiptopodius granulosus (from Zambia) is morphologically similar to the new species. Since both species occur in neighbour countries, there is a huge potential of misidentification between them, that’s why the diagnosis of the new species is compared to the closest species in southern Africa. Conservation status and potential threats. This species was recorded from the well-protected Savuti area of Chobe National Park in northern Botswana. The vegetation consists of somewhat open to dense stands of Philenoptera nelsii (Schinz) Schrire. The trees are about one to two meters tall with about 30% canopy cover (CMD, personal observation). Although it is protected at Savuti, the extent of occurrence of this species is not known and it should thus be assessed as Data Deficient.

Stiptopodius muellerae Daniel & Deschodt, new species (Figs. 1C, 2 A���B, 3A, 3C, 4) Type locality: Caconda, Huila, Angola. Type material. Holotype, male: Angola, Huila Prov. | 75 km S Kacopnda [Caconda] | 13.24 S ��� 15. 28 E || 8.11.2011;E-Y: 3916 | sandy forest, 1643m | leg. Ruth M��ller || TM SOUTH AFRICA | TMSC13817 || HOLOTYPE: Stiptopodius | muellerae Daniel & Deschodt | 2020 (printed on red card) (TMSA). Description. Holotype, male. Length: 7.6 mm, width: 3.8 mm. Colour. Black. Head. Incision between medial clypeal teeth deeply concave, lateral clypeal edges not sinuate; external protrusion of genae sharply triangular; surface of head with green sheen; punctation varying from fine to large; setae well-developed except on glabrous anterior clypeal edge; clypeo-frontal carina visible; clypeo-genal suture present; vertex with small medial carina. Antennae with eight antennomeres; labium bears a few well-developed setae. Pronotum. Convex, wider than long; base of pronotum marginate; anterior margin wider than posterior; in lateral view, lateral edge strongly sinuate before postero-lateral angles, surface with green metallic sheen, ocellate punctation varying from fine to large, each side with small longitudinal tubercle; well-developed setae, except on glabrous central disc. Elytra. Lateral margin straight, somewhat convex distally and concave proximally, well-developed humeral umbone; striae sparsely and finely punctate. Interstriae punctate, fully setose. Pygidium. Convex and setose, punctation ocellate. Abdominal ventrites. Prosternum surface with well-developed setae, anterior angles of prosternum slanted forward and inward. Meso- and metaventrite dull, setose and densely punctate; metaventrite with long setae, surface impressed longitudinally at middle. Abdominal ventrites with ocellate punctures and strong setae arranged in rows laterally, glabrous medially; last abdominal ventrite narrowed medially (male character). Legs. Profemora sparsely punctate, bearing well-developed setae, external edge strongly marginate, internal margin slightly marginate. Meso- and metafemora with external and internal edges sparsely setigerous and punctate; surface between external and internal margin with few setigerous punctures, strongly marginate in both edges (external and internal). Protibiae crenulate with four teeth on external margin, line of setae around teeth in dorsal view; in ventral view, with two carinae, internally punctate and pilose. Protibial spur long and inward curving. Meso- and metatibiae triangular and strongly widened apically. Protarsi with five tarsomeres and clearly visible pair of claws; fifth protarsomere at least 2x longer than others (each individually). Meso- and metatarsomeres I���IV triangular, longer than wide, with internal line of strong setae on tarsomere I, strongly concave medially, bilobate distally, external lobe strong, internal lobe weak; tarsomere V rectangular and longer than others, followed by pair of very tiny apical claws. Aedeagus. Parameres symmetrical, elongate with tips acute (Figs. 2 A���B). Female. Unknown Distribution. The species is only known from the type locality, Caconda, Province of Huila, Angola. It was hand collected in sandy forest. Etymology. The specific name honours Ruth M��ller, a friend and curator in charge of Coleoptera at the Ditsong National Museum of Natural History, Gauteng, Pretoria, South Africa, who collected the holotype. Diagnosis. The new species is somewhat similar to S. singularis by having the basal half of the pronotal disc glabrous and frontal carina obsolete. However, S. singularis bears uniform punctation on the pronotum, with a brown metallic sheen, whereas S. muellerae new species has a distinctive green metallic sheen with punctures of variable sizes ranging from fine to large. The elytra of this new species are fully setose, while those of S. singularis are slightly setose posteriorly and glabrous anteriorly. The mesotibiae are sinuate externally in S. muellerae new species but not in S. singularis. The male genitalia differ between the two species by having the tips of the parameres sharp and projecting forward in the new species (Figs. 2 A-B), whereas in S. singularis they are short and blunt (Figs. 2 E-F). S. muellerae new species is also much larger than S. singularis. Remarks. This is the first record of a Stiptopodius species whose meso- and metatarsomeres I���IV are longer than wide (Figs. 3A, C). The first tarsomere is the longest, which is in contrast to the other 5-articulated species, where tarsomere V is longer than others. Furthermore, in S muellerae new species, meso- and metatarsomeres I���IV bear a weak lobe externally. Conservation status and potential threats. The forest at the type locality is surrounded by lands supporting agricultural and pastoral farming activities (R. M��ller, personal communication). It is therefore probably under severe pressure. As this species is only known from this locality, it may warrant the threat category of at least Vulnerable., Published as part of Daniel, Gimo M., Deschodt, Christian M., Davis, Adrian L. V. & Sole, Catherine L., 2020, The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa, pp. 394-404 in Zootaxa 4763 (3) on pages 400-402, DOI: 10.11646/zootaxa.4763.3.4, http://zenodo.org/record/3759527, {"references":["Harold, E. von (1871) Verzeichniss der von Dr Beccari in Bogos gesammelten Coprophagen Lamellicornien. Coleopterologische Hefte, 8, 1 - 28. https: // doi. org / 10.1002 / mmnd. 47918710229","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa (Lucanidae and Scarabaeidae). Transactions of the South African Philosophical Society, 12, 1 - 920. https: // doi. org / 10.1080 / 21560382.1901.9525977"]}

Identification key to Stiptopodius species modified from Branco (2009) 1 Sides of pronotum (lateral view) weakly sinuate before postero-lateral angle (S. doriae species group).................. 2 ��� Sides of pronotum (lateral view) strongly sinuate before postero-lateral angle (S. singularis species group).............. 4 2 Body elongate, parallel-sided. Pronotum quadrate, much shorter than elytra. Head with two oblique tubercles on the vertex, often obsolete. Length 3.9 - 5.7 mm. Ethiopia, Somalia, Kenya, Sudan, Uganda, Chad, Niger, Nigeria, Mali, Burkina Faso........................................................................... Stiptopodius doriae Harold, 1871 ��� Body short and wide. Pronotum with sides arcuate, only slightly shorter than elytra Head with two transversely-aligned weak tubercles on vertex.................................................................................... 3 3 Pronotal disc brown with metallic sheen. Meso- and metatarsi with five tarsomeres and apical pair of claws, meso- and metatarsomeres I to IV triangular and wider than long. (male; female unknown). Length 4.0 mm. South Africa (KwaZulu-Natal, Richards Bay)........................................... Stiptopodius peringueyi Daniel & Deschodt, new species ��� Pronotal disc dull. Meso- and metatarsi clawless with three tarsomeres, meso and metatarsomeres I and II much longer than wide, tarsomere III small and not bilobate. Length 4.5���5.2 mm. Kenya, Tanzania.... Stiptopodius longipedis Branco, 1991 4 Head with two clearly separated tubercles on vertex. Clypeo-frontal carina present................................. 5 ��� Head with carina on vertex, entire or slightly interrupted in the middle, but never divided into two separated small tubercles. Clypeo-frontal carina absent............................................................................ 6 5 Cephalic tubercles approximately equidistant from each other and the inner edge of eyes; space between tubercles flat or very slightly depressed. Length 5.4���7.6 mm. Eritrea, Ethiopia, Somalia, Kenya, Tanzania..................................................................................................... Stiptopodius patrizii Boucomont, 1923 ��� Cephalic tubercles much closer to inner edge of eyes than to each other; space between tubercles strongly depressed (male; female unknown). Length 7.3 mm. Kenya...................................... Stiptopodius krausei Branco, 1991 6 Clypeal surface not granular, punctation varies from small, simple punctures to large and ocellate..................... 7 ��� Clypeal surface at least partly granular and punctate........................................................ 12 7 Second elytral interstria with punctation aligned alongside the striae, middle of interstria apunctate. Head and pronotum of metallic colour or with a metallic sheen................................................................... 8 ��� Second elytral interstria with punctation irregularly distributed over entire surface. Head and pronotum black or brown, without metallic sheen...................................................................................... 10 8 Disc of pronotum glabrous at basal half, densely punctate. Frontal carina most often obsolete........................ 9 ��� Disc of pronotum pilose at basal half, with sparsely set double punctation, the larger punctures setiferous. Frontal carina most often strong. Length 5.3���6.6 mm. Kenya, Tanzania, Malawi................... Stiptopodius nitidus (Boucomont, 1923) 9 Pronotum with uniform, dense and medium-sized punctation. Elytra sparsely setose posteriorly. Mesotibiae not sinuate externally. Parameres short, with blunt tips. Length 4.5���6.0 mm. Tanzania, Malawi, Mozambique, Zambia, Zimbabwe, South Africa (Limpopo, Gauteng and Natal), Angola................................... Stiptopodius singularis (P��ringuey, 1901) ��� Pronotum with punctures of mixed size from fine to large. Elytra fully setose. Mesotibiae sinuate externally. Parameres with tips sharp and projecting forward (male; female unknown). Length 7.6. mm. Angola....................................................................................... Stiptopodius muellerae Daniel & Deschodt, new species, 10 Disc of pronotum glabrous, with uniform punctation, punctures of fine to medium-size. Elytral interstriae either pilose or glabrous............................................................................................. 11 ��� Disc of pronotum pilose, with double punctation, the larger punctures fairly strong, ocellate and setigerous. Elytra always entirely pilose. Length 6.5���7.3 mm. Niger, Nigeria, Cameroon.................. Stiptopodius gaillardi (Boucomont, 1923) 11 Frontal carina strong. Upper surface of the body entirely glabrous, except sometimes on sides of pronotum and on the eighth elytral interstria. Length 6.0��� 7.2 mm. Sudan..................................... Stiptopodius glaber Branco, 1992 ��� Frontal carina obsolete. Upper surface of the body either pilose or glabrous. Pronotum always pilose on the sides; the eighth elytral interstriae varies from pilose or glabrous. Length 4.8���6.5 mm. Nigeria, Benin, Burkina Faso, Ghana, Senegal.......................................................................... Stiptopodius nodieri (Boucomont, 1923) 12 Clypeal surface only partly granular; genae and vertex not granulose. Integument of pronotum not rugose.............. 13 ��� Head entirely granular. Integument of pronotum strongly rugose laterally. Length 4.9���5.0 mm. Zambia....................................................................................... Stiptopodius granulosus Branco, 2009 13 Head and pronotum of bronze colour. Length 4.5���5.9 mm. Somalia............... Stiptopodius glabricollis M��ller, 1942 ��� Head and pronotum of black colour (female; male unknown). Length 4.3 mm. Botswana (Savuti)................................................................................ Stiptopodius savuti Daniel & Deschodt, new species, Published as part of Daniel, Gimo M., Deschodt, Christian M., Davis, Adrian L. V. & Sole, Catherine L., 2020, The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa, pp. 394-404 in Zootaxa 4763 (3) on pages 403-404, DOI: 10.11646/zootaxa.4763.3.4, http://zenodo.org/record/3759527, {"references":["Branco, T. (2009) Notes on \" Stiptopodius \" Harold, 1871: number of tarsal segments of S. longipedis Branco, 1991, and description of a new species from Zambia (Coleoptera, Scarabaeidae). Boletin Sociedad Entomologica Aragonesa, 44, 87 - 91.","Harold, E. von (1871) Verzeichniss der von Dr Beccari in Bogos gesammelten Coprophagen Lamellicornien. Coleopterologische Hefte, 8, 1 - 28. https: // doi. org / 10.1002 / mmnd. 47918710229","Branco, T. (1991) Revision des genres du \" groupe \" Stiptopodius: Les genres Stiptopodius Harold et Stiptotarsus Branco (Coleoptera: Scarabaeidae). Annales de la Societe Entomologique de France, Nouvelle Serie, 27, 265 - 285.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa (Lucanidae and Scarabaeidae). Transactions of the South African Philosophical Society, 12, 1 - 920. https: // doi. org / 10.1080 / 21560382.1901.9525977","Branco, T. (1992) Revision des genres du \" groupe \" Stiptopodius: Donnes complementaires (Coleoptera, Scarabaeidae). Annales de la Societe Entomologique de France, Nouvelle Serie, 28, 101 - 108."]}

Stiptopodius doriae species group Differential diagnosis. The Stiptopodius doriae species group differs from the Stiptopodius singularis species group by having the sides of the pronotum weakly sinuate before the postero-lateral angle in lateral view., Published as part of Daniel, Gimo M., Deschodt, Christian M., Davis, Adrian L. V. & Sole, Catherine L., 2020, The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa, pp. 394-404 in Zootaxa 4763 (3) on page 395, DOI: 10.11646/zootaxa.4763.3.4, http://zenodo.org/record/3759527

Revalidation of Gyronotus marginatus P��ringuey, 1888 (Figs. 1E, 3) Although Scholtz & Howden (1987: 82) did not see the type of G. marginatus, they stated that ��� Gyronotus marginatus P��ringuey is without doubt a synonym of G. pumilus. Although the type is missing from the P��ringuey collection it is clear from the description and the type locality that the two names refer to one species. Consequently we synonymize G. marginatus with G. pumilus ���. The type of Gyronotus marginatus has subsequently been located in the Iziko South African Museum (ISAM). It is clearly larger than any specimens of Gyronotus pumilus, as are the paratype specimens (examined by ALVD) that are housed in BMNH. Consequently, we revalidate Gyronotus marginatus from the junior subjective synonymy with Gyronotus pumilus and re-elevate it to species rank. Diagnosis. Gyronotus marginatus is clearly much bigger than Gyronotus pumilus. Also, the elytra of G. marginatus are shagreened and fairly even while the elytra of G. pumilus are shagreened and somewhat rugose., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2019, New southern African species and a revalidation in the dung beetle genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an updated key, pp. 275-282 in Zootaxa 4624 (2) on page 281, DOI: 10.11646/zootaxa.4624.2.10, http://zenodo.org/record/3259222, {"references":["Scholtz, C. H. & Howden, H. F. (1987) A revision of the southern African Canthonina (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of the Entomological Society of Southern Africa, 50, 75 - 119."]}

Gyronotus ovalis Deschodt & Davis, new species (Figs. 1A, 2 A���D, 3) Type material. Holotype, male: ��� Nkandla Forest Reserve, Grassland | S28.744514�� E31.140557��, 1070m | 10- 12.xii.2014, Deschodt and Davis | pig dung baited pitfall traps��� (TMSA). Paratypes: 2 males and 8 females, same data as holotype (5 in TMSA, 3 in UPSA, 2 in ISAM). Description of the holotype, male. Size. Length 12.3 mm and width 8.0 mm with head extended (not deflexed). Habitus. Body shape ovoid. Dark brown in appearance. Head. Tip of clypeus bidentate, separated by a wide ���U��� shaped depression. Surface, densely punctate, punctures closer together than their diameter. Clypeogenal sutures fairly distinct and nearly straight. Frontogenal suture clearly visible only at outside tenth, middle four fifths not visible. No tubercle present. Antennae dark orange. Thorax. Pronotal disc densely punctate, punctures medium sized and round to slightly ovoid, almost touching each other. Lateral margin rounded. Prosternum with large faint punctures, surface dull. Mesosternum with large almost contiguous punctures. Metasternum with small sized punctures up to two times further apart than their diameter. Surface of prosternum and metasternum otherwise shagreened. Meso-metasternal suture clearly visible and somewhat curved. Elytra. Striae faint and narrow with punctures spaced at intervals of two to three puncture diameters. Interstriae punctate, punctures small separated by one to two puncture diameters. Surface otherwise shagreened. Pygidium with large shallow punctures about two puncture diameters apart, otherwise shagreened giving a dull appearance. Aedeagus. Parameres bilaterally asymmetrical (Fig. 2). Female. As with all Gyronotus species the tip of the front tibiae strongly curves downward with a shorter spur whereas the tip of the front tibiae of females is straight with a longer spur. Also, the mesotibiae of males strongly widen apically whereas the mesotibiae of females widen gradually. Distribution. Gyronotus ovalis Deschodt & Davis, new species occurs in the grassland around Nkandla forest. Etymology. The generic name and specific epithet is a pleonasm (Gyro=round and ovalis=oval) accentuating its ovoid shape. Diagnosis. This species bears short setae on the elytra as opposed to almost glabrous elytra in the morphologically similar, Gyronotus glabrosus and Gyronotus kearneyorum Deschodt & Davis, new species., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2019, New southern African species and a revalidation in the dung beetle genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an updated key, pp. 275-282 in Zootaxa 4624 (2) on pages 277-278, DOI: 10.11646/zootaxa.4624.2.10, http://zenodo.org/record/3259222

Gyronotus dracomontanus Deschodt & Davis, new species (Figs. 1C, 3) Type material. Holotype, female: ���Z.A.29 | Bergville Dist. | Drakensberg || Prairie subalpine | Alt. 2,400m., x-1960 || N. Leleup Leg. ��� (TMSA). Paratypes: two females, no locality data: ���ex collection of | Pietermaritzburg | Museum ��� (UPSA). Description of the holotype, female. Size. Length 12.0 mm and width 8.0 mm with head extended (not deflexed). Habitus. Body shape ovoid. Black in appearance. Head. Tip of clypeus bidentate, separated by a wide ���U��� shaped depression. Surface with dense punctation bearing very short fine setae almost touching each other. Clypeogenal sutures fairly distinct and nearly straight. Frontogenal suture clearly visible only at external one tenth; middle four fifths not visible. Small round tubercle medially on frontogenal suture. Antennae dark orange. Thorax. Pronotal disc densely punctate, punctures round to slightly ovoid, mostly touching one another. Lateral margin with obtuse angle almost halfway, otherwise relatively straight. Surfaces of all thoracic sternites shagreened between punctation. Prosternum with medium sized punctures that are separated by about half a puncture diameter. Mesosternum with medium sized punctures that are close together frontally and further apart anteriorly; row of overlapping setigerous punctures on mesometasternal suture. Metasternum with small sized punctures spaced up to three diameters apart. Mesometasternal suture clear, evenly slightly curved. Elytra. Interstriae punctate, punctures small and setigerous separated by half up to two puncture diameters. Surface otherwise shagreened appearing slightly rugose under low magnification (x10). Striae indistinct, visible as slight impressions bearing small punctures that are spaced one to two puncture diameters apart. Pygidium. Faintly punctate, punctures small and setigerous about one to three puncture diameters apart, laying closer together dorsally and further apart ventrally, surface otherwise shagreened giving a dull appearance. Distribution. Gyronotus dracomontanus Deschodt & Davis, new species most probably occurs in the high grassland above 2000 m between Royal Natal National Park [S28.702�� E28.915��] and Cathedral Peak [S28.925�� E29.134��]. Etymology. This species is named for the Drakensberg mountain range where it occurs. Diagnosis. This species can be separated from most other Gyronotus species by having prominent, dense setae on the elytra. The highland grassland species, Gyronotus dracomontanus Deschodt & Davis, new species, has six or less adjacent setae on the second interval of the interstriae compared to eight or more in the coastal forest species, Gyronotus pumilus., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2019, New southern African species and a revalidation in the dung beetle genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an updated key, pp. 275-282 in Zootaxa 4624 (2) on pages 276-277, DOI: 10.11646/zootaxa.4624.2.10, http://zenodo.org/record/3259222

Tribe Endroedyolini Davis, Deschodt & Scholtz, new tribe Type genus. Endroedyolus Scholtz & Howden, 1987, here designated (Figs. 3, 6). Head. Clypeus bidentate with a notch to the side of each denticle. Antennae nine segmented. Pronotum. Very convex with lateral margins explanate, sometimes carinate laterally. Elytra. Strongly convex; setae in straight and slightly converging lines; metathoracic wings always absent. Sterna. Smooth to punctate with punctures of varying density and size; prosternum fairly well developed; metasternum smooth to punctate, sometimes with deep fovea extending from anterior to sometimes half way between mesocoxae. Protibia. With three small denticles on outside margin. Meso- and metatibia. Unmodified with setae. Pygidium. With small setiferous punctures; with or without depression or fovea. Aedeagus. Parameres symmetrical. Body size. Very small to small: 1.5mm to 4.8 mm. Diagnosis and known distribution. The combination of extreme small size, very convex body, unique clypeal shape and explanate lateral pronotal margins, distinguishes members of this tribe from any other. They are restricted to forest patches of south east South Africa where they are associated with forest litter.

Tribe Odontolomini Davis, Deschodt & Scholtz, new tribe Type genus. Odontoloma Boheman, 1857, here designated (Figs. 2, 5). Head. Clypeus with a small median tooth flanked by two others; frons and vertex without horns; antenna nine segmented. Pronotum. More or less flat dorsally; strongly curving downward both anteriorly and laterally; anterio-lateral margins often explanate. Elytra. Seventh stria or eighth interstria interval partly carinate anteriorly; inflexed pseudepipleuron, epipleuron narrow; metathoracic wings well developed to greatly reduced. Sterna. Prosternum fairly well developed; metasternum mostly with clear punctures; sutures clearly visible. Protibia. Front tibia truncate and obtuse, widening apically with three teeth on outside margin. Meso- and metatibia. Widening apically, with setae. Pygidium. Basal transverse groove or depression present, groove terminating in a fovea on each side near margin. Aedeagus. Parameres symmetrical and short. Body size. All species are small, less than 5 mm in length with some size variation observed within species. Diagnosis and known distribution. Body oval and moderately convex; black; densely punctate with scattered semi-erect setae; clypeus five-dentate; all tibiae widening anteriorly, anterior tibiae with three denticles, truncated and obtuse. Widespread in the Afrotropical region but with a strong bias in species diversity centred on southern Africa., Published as part of Davis, Adrian L. V., Deschodt, Christian M. & Scholtz, Clarke H., 2019, Defining new dung beetle tribes to resolve discrepancies between phylogeny and tribal classification in the subfamily Scarabaeinae (Coleoptera: Scarabaeidae), pp. 131-144 in Zootaxa 4608 (1) on page 136, DOI: 10.11646/zootaxa.4608.1.7, http://zenodo.org/record/3993625

Tribe Byrrhidiini Davis, Deschodt & Scholtz, new tribe Type genus. Byrrhidium Harold, 1869, here designated (Figs. 1, 4). Head. Punctate; anterior clypeal margin often with forklike small to large teeth protruding anteriorly; single very small median tooth on lower margin of clypeus; never with a horn on the frons or vertex; antenna nine segmented; mouthparts varying between genera but labrum always strongly sclerotized apically, inside margins of glossae with sclerotized denticles. Pronotum. Pronotum convex, with sub-parallel sides; punctation indistinct. Elytra. Convex; fused without humeral umbones; elytra with seven to eight feebly visible striae followed by an acute sub-lateral pseudoepipleural carina and inflexed pseudepipleuron; interstriae intervals flat; metathoracic wings absent; scutellum not visible from above. Sterna. Smooth to punctate; sutures well defined. Protibia. With a terminal spur; two or three denticles on anterior outside margin; posterior outside margin serrated or smooth; tarsi short. Meso- and metatibia. Unmodified, long and thin; each meso-and metatarsus of uniform width, often setate. Pygidium. Somewhat convex; punctate. Aedeagus. Phallobase unmodified; parameres symmetrical and extremely variable. Body size. Small to medium with size varying between 4.9 mm �� 3.8 mm for the smallest and 11.8 mm �� 7.5 mm for the largest known species. Diagnosis and known distribution. The Byrrhidiini new tribe, can be distinguished from all other dung beetle tribes by the combination of the following characters: flightless with body medium sized, sparsely setate and strongly convex; antennae nine segmented; labrum sclerotized apically, inside margins of glossae with sclerotized denticles; punctures on pronotum visible but indistinct; an acute sub-lateral pseudoepipleural carina and inflexed pseudepipleuron; striae and punctures on the elytra indistinctly visible; no humeral umbone visible; scutellum not visible from above; unmodified meso- and metatibiae of the same width; distributed in the arid to hyper-arid region along the southwest seaboard of Namibia and South Africa between about 21 and 31 degrees south with most known species occurring in Namibia (Fig. 4)., Published as part of Davis, Adrian L. V., Deschodt, Christian M. & Scholtz, Clarke H., 2019, Defining new dung beetle tribes to resolve discrepancies between phylogeny and tribal classification in the subfamily Scarabaeinae (Coleoptera: Scarabaeidae), pp. 131-144 in Zootaxa 4608 (1) on pages 134-136, DOI: 10.11646/zootaxa.4608.1.7, http://zenodo.org/record/3993625

Epirinus schoolmeestersi Deschodt & Davis, new species Figs. 1D, 3B, 5. Type material: Holotype. Male: S.Afr., SE CapeMts | Baviaanskloofberg | 33.36 S - 24.23 E || 15.11.1982, at 595m. | Groundtraps, 30 days | leg. W.Breytenbach || groundtraps with faeces bait WB240 (TMSA). Paratypes. 14 males and 12 females with same label as holotype and 2 males and 2 females with same label as holotype except the altitude is “at 885m ” and the collection number is “ WB242 ”. All specimens with two additional determination labels: EPIRINUS | silvestris | Cambefort | det. C.H. Scholtz || EPIRINUS | hilaris Péringuey | det. Claudia A. Medina. 22 TMSA, 2 UPSA. Description of holotype, male. Body: 4.1 x 2.1 mm (length with head extended x maximum width) Small and convex. Head: Clypeal teeth small, pointing upwards and separated by a rounded indentation about twice their length; very small notch on lateral margin where genal suture terminates; frontal suture clearly effaced medially; clypeal surface densely punctate, no associated setae; eyes narrowing posteriorly; antennae dark. Pronotum: Surface shagreened, densely punctate with round punctures; shallow medial depression at base of the disc extending in a feebly defined, puncture-free line that extends halfway over pronotum; no lateral foveae. Elytra: Striae distinct, with clear double margins; interstriae with oval granules arranged in a clear single line and separated by one and a half to two granule widths with a single recurved seta on each granule; elytral margin raised for apical two thirds; single apical tubercle present on third and fifth interstriae. Pygidium: Shagreened, with large punctures and two horizontal basal costae. Venter: Ventrites finely setose with single irregular row of punctures; metaventrite punctures large and almost touching; mesoventrite punctures smaller and separate. Male genitalia: Asymmetrical (Fig. 3B). Etymology: This species is named after Paul Schoolmeesters, who is the tireless custodian of the World Scarabaeidae Database. Diagnosis: In appearance, this species is very similar to E. hilaris Péringuey, 1901. However, E. schoolmeestersi new species can be readily separated by the presence of granules and setae on its elytra clearly arranged in a single line. In comparison, E. hilaris has almost no granules, somewhat irregular lines of setae and a larger, clear, puncture-free line on the pronotum. Distribution, ecology, conservation status, and potential threats: This species occurs on the Baviaanskloof mountains and probably extends into the nearby Kouga mountain range. It is probably restricted to the Fynbos Biome that stretches over these mountain ranges. Based on limited distribution data, this species is assessed as data deficient (DD). It is well protected in the Baviaanskloof Nature Reserve.

Epirinus davisi Scholtz & Howden, 1987 Figs. 2A, 4A, 5. Epirinus hluhluwensis Medina & Scholtz, 2005 new subjective synonymy Epirinus ngomae Medina & Scholtz, 2005 new subjective synonymy In their revision of Epirinus, Medina & Scholtz (2005) describe Epirinus hluhluwensis Medina & Scholtz, 2005 and Epirinus ngomae Medina & Scholtz, 2005, as new species that are separated from other Epirinus by the presence of two pronounced lateral foveae and from one another by the shape of the clypeal teeth. With reference to characters given in the species descriptions and the accompanying identification key (Medina & Scholtz 2005), we compared paratype specimens of E. hluhluwensis and E. ngomae with paratype and newly collected specimens of Epirinus davisi Scholtz & Howden, 1987. Only minor external morphological differences were observed between those specimens, which originated from five localities (Table 1, Figs. 2, 4, 5). With regards to external morphology of the aedeagi, there was very little difference between the specimens. Most notably we observed only slight variation in the depth of the median depression at the base of the pronotum. The observed minor differences are interpreted as intraspecific variability. We could not see the differences noted by Medina & Scholtz (2005) concerning sclerites X and Y. Hence, we propose that E. hluhluwensis and E. ngomae should be regarded as new junior subjective synonyms of E. davisi. As a result, E. davisi would now be known from widely-separated coastal and lower escarpment forest localities of northeast KwaZulu-Natal and eSwatini that represent an extent of occurrence (EOO) of ~ 11700 km 2., Published as part of Deschodt, Christian M., Davis, Adrian L. V. & Daniel, Gimo M., 2019, The endemic Southern African genus Epirinus Dejean, 1833 (Coleoptera: Scarabaeidae: Scarabaeinae) revisited: five new species and two new synonyms, pp. 327-340 in Zootaxa 4603 (2) on page 335, DOI: 10.11646/zootaxa.4603.2.6, http://zenodo.org/record/2682436, {"references":["Scholtz, C. H. & Howden, H. F. (1987) A revision of the southern African genus Epirinus. Journal of the entomological Society of Southern Africa, 50, 121 - 154.","Medina, C. A. & Scholtz, C. H. (2005) Systematics of the southern African genus Epirinus Reiche Coleoptera: Scarabaeinae: Canthonini: descriptions of new species and phylogeny. Insect Systematics and Evolution, 36 (2), 145 - 160. https: // doi. org / 10.1163 / 187631205788838500"]}

Epirinus jacobsae Deschodt & Davis, new species Figs. 1E, 3A, 5. Type material: Holotype. Dissected male: Mapelane, Pig dung. | -28��24'30.18" 32��25'13.83" | 16-18.x.2011 Deschodt (TMSA). Paratypes. 5 females, same data as holotype, 2 TMSA, 2 UPSA, 1 SANC. Description of holotype, male. Body: 4.1 x 2.3 mm (length with head extended x maximum width) Head: Clypeal teeth prominent, pointing upwards with a pair of setae at base of each tooth; no distinct median indentation; very small notch on lateral margin where genal suture terminates; frontal suture clearly effaced medially; clypeal surface densely punctate, with associated setae; eyes narrowing posteriorly. Pronotum: Surface densely punctate with round punctures associated with recurved setae; medial depression distinct with no median longitudinal line and no lateral foveae. Elytra: Striae distinct, with large, round punctures; interstriae shiny and smooth; first and seventh interstriae with one row of setae, all other interstriae (up to the eighth) with two well-defined rows of setae (some setae abraded); setae golden brown with recurved points; elytral margin raised for apical two thirds; apical tubercles absent. Pygidium: With two large tubercles on surface. Venter: Hypomeral carina reaching anterior pronotal margin; ventrites fairly rugose with single row of punctures, setae fine; metaventrite punctures small, separated by approximately 2x their diameter. Male genitalia: Asymmetrical (Fig. 3A). Etymology: This species is named after Carmen Jacobs who tirelessly administered the bureaucracy of the Scarab Research Group. Diagnosis: In appearance, this species is very similar to E. davisi. However, E. jacobsae new species can be readily separated by its smaller metaventrite punctures that are further apart. Remarks: Males with slightly more sinuous hind legs than females. Distribution, ecology, conservation status, and potential threats: Epirinus jacobsae new species is the first flightless Epirinus recorded from Northern Coastal Forest as defined by Mucina and Rutherford (2006). This forest type is categorised as ���least threatened in general��� and ���well protected��� within the iSimangaliso Wetland Park (Mucina & Rutherford 2006). However, it is uncertain if E. jacobsae new species occurs to the north of the type locality in Maphelane Nature Reserve, which is isolated from the much larger northern part of the iSimangaliso park by the Umfolozi River and St Lucia Estuary. Epirinus jacobsae new species is currently only known from Maphelane, which lies at the northernmost extent of the single forest patch bordered to the south by the Sokhulu State Forest (S 28.50�� E 32.40��). No specimens were recorded during two previous extensive sampling efforts (Davis et al. 2003, 2013) in Sokhulu State Forest. Should E. jacobsae new species occur in Sokhulu, it is noteworthy that mining for Titanium-bearing sands within the state forest are projected to continue northwards terminating just to the south of Maphelane. This will reduce the potential habitat for E. jacobsae new species from 4300 hectares (Ha) to about 800 Ha (area sizes from Mucina & Rutherford 2006). The mining company is actively engaged in the rehabilitation of the forest (van Aarde et al. 1996). However, Davis et al. (2003) noted that another flightless dung beetle species, Gyronotus carinatus Felsche, 1911, is notably absent from formerly disturbed forest patches whereas it is still present in the relatively undisturbed Sokhulu Forest Reserve. The reason for this absence is unclear but could be due to unsuitability of habitat, disturbance, or isolation from the relatively undisturbed dune forest at Sokhulu. We believe that it would be even more difficult for the much smaller E. jacobsae new species to colonize any of the rehabilitated habitat. According to the IUCN red list categories, E. jacobsae new species should be classified as endangered B2ab (iii). See IUCN Standards and Petitions Subcommittee (2012) for an explanation of the criteria., Published as part of Deschodt, Christian M., Davis, Adrian L. V. & Daniel, Gimo M., 2019, The endemic Southern African genus Epirinus Dejean, 1833 (Coleoptera: Scarabaeidae: Scarabaeinae) revisited: five new species and two new synonyms, pp. 327-340 in Zootaxa 4603 (2) on pages 328-330, DOI: 10.11646/zootaxa.4603.2.6, http://zenodo.org/record/2682436, {"references":["Mucina, L. & Rutherford, M. C. (Eds.) (2006) The Fegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, 807 pp. https: // doi. org / 10.5962 / bhl. title. 61905","Davis, A. L. V., van Aarde, R. J., Scholtz, C. H. & Delport, J. H. (2003) Convergence between dung beetle assemblages of a postmining chronosequence and unmined dune forest. Restoration Ecology, 11, 29 - 42. https: // doi. org / 10.1046 / j. 1526 - 100 X. 2003.00133. x","Davis, A. L. V., van Aarde, R. J., Scholtz, C. H., Guldemond, R. A. R., Fourie, J. & Deschodt, C. M. (2013) Is microclimate-driven turnover of dung beetle assemblage structure in regenerating coastal vegetation a precursor to re-establishment of a forest fauna? Journal of Insect Conservation, 17, 565 - 576. https: // doi. org / 10.1007 / s 10841 - 012 - 9542 - 8","van Aarde, R. J., Ferreira, S. M. & Kritzinger, J. J. (1996) Successional changes in rehabilitating coastal dune communities in northern KwaZulu / Natal, South Africa. Landscape and Urban Planning, 34, 277 - 286. https: // doi. org / 10.1016 / 0169 - 2046 (95) 00237 - 5","Felsche, C. (1911) Coprophage Scarabaeiden. Deutsche Entomologische Zeitschrift, 133 - 141."]}

Epirinus muellerae Deschodt & Davis, new species Figs. 1A, 3C, 5. Type material: Holotype. Male: LESOTHO | Thaba Tseka | 22.11.1997; R Müller (TMSA). Paratypes. 3 males and 4 females with same label as holotype. All specimens with single additional determination label: EPIRINUS | striatus | Scholtz & Howden | det. C.A.Medina 2002 (TMSA). 3 males and 6 females: SANI PASS, DRAKENSBERG | KWAZULU/NATAL, 2800 m | 29°35’ S 29°17’ E | 14–15.I.1997, | SCHOLTZ & NORDSTROM (UPSA). 1 male and 1 female: LESOTHO | Katse | (3 km SW of) | alt 2300 m | 22.I.1991 | H. Geertsema | LHDA-0010 and second specimen same except LHDA ♂ (TMSA). 1 male: LESOTHO | Laitsoka Pass | alt. 2650m | 3.X.1991 | H. Geertsema | LHDA-416 (TMSA). Description of the holotype, male. Body: 9.7 x 5.8 mm (length with head extended x maximum width) Medium sized and somewhat convex. Head: Clypeal teeth small, pointing upwards and separated by a rounded indentation more than twice their length; frontal suture clearly effaced medially; clypeal surface densely and irregularly punctate, punctures touching giving rugose appearance, no associated setae; eyes narrowing posteriorly; antennae dark. Pronotum: Surface shagreened with sparsely irregularly placed short black setae, densely punctate with round punctures, punctures almost touching; shallow, medial, basal depression terminating in a puncture-free line that extends over one-third the length of the pronotal disc anteriorly; no lateral foveae. Elytra: Striae distinct and punctate with clear double margins; interstriae with small granules, single recurved brown setae on some granules; setae arranged in a single row on sutural interval and in double rows from second interval becoming more irregular after fifth interval. Pygidium: Shagreened, with small punctures, short irregular black setae present. Venter: Ventrites very finely setose with single irregular row of punctures; metaventrite punctures irregular in size and shape; mesoventrite rugose. Male genitalia: Asymmetrical (Fig. 3C). Etymology: This new species is named in honour of Ruth Müller (TMSA), long-time curator at the Ditsong National Museum of Natural History, and who collected half of the type series. Diagnosis: This species is very similar to E. striatus Scholtz & Howden, 1987. However, E. muellerae new species can be readily separated by it having a median basal depression on the pronotal disc which is absent in E. striatus. Remarks: Males with slightly more sinuous hind legs than females. Distribution, ecology, conservation status, and potential threats: This species occurs in the highlands of Lesotho with a range that may extend onto the eastern escarpment of South Africa, although observations are lacking. It is centred on the Drakensberg Grassland bioregion in the Grassland biome. Although the protection of grassland is limited in Lesotho, there is little anthropic development across its known extent of occurrence, which is about 2000 km 2. Therefore, it is assessed as least concern (LC).

Garreta australugens Davis & Deschodt, new species Figs 1–4 Description of holotype. Holotype ♂: 20.3 x 11.6 mm (length with head extended x maximum width). Habitus uniformly charcoal black. Head. Clypeus with four well defined denticles; middle two denticles larger and separated by right-angled gap; clypeal surface with large, irregular granules that are close together but clearly separated by dense microgranulation. Pronotal disc. Surface of pronotum with small more or less rounded and irregularly spaced granules that are separated by less than one or two granule diameters; area between granules, micro-granular giving a shagreened appearance. Longitudinal midline almost indiscernible. Two very short diagonal grooves (basal impressions) on each side of midline at the base of pronotum. Elytra. Surface sculpture of elytral interstriae similar to pronotum, i.e. large granules separated by one to three granule diameters; area between granules very finely microgranular with a shagreened appearance. First elytral striae next to midline with widely spaced and very faint punctures; all other striae apunctate, narrow, unclear and shagreened. Anterior legs. Narrow and slightly curved inwards. Aedeagus. Similar to other species of Garreta (Fig. 3). Sexual dimorphism. Inward curving front legs of male type material narrower than those of females. Terminal spur of front legs acute in females and more blunt in males. Variability. The colour of paratype specimens is uniform with no variation; size of paratype specimens varies from slightly smaller to slightly larger than the holotype. Etymology. The name is a combination of the specific name of its closest relative Garreta lugens and its southern distribution. Type specimens examined. Holotype ♂: "Wildlife College, Site 3, S24°32’27” E31°20’23”, 23-25.xi.2009, C. Du Toit & C. Jacobs. ", in SANC. Paratypes: 2 spec. same data as holotype, in UPSA; 1 spec. "S. Afr.; Transvaal, Pafuri, 29.3.73., Potg.&Scholtz", in TMSA; 1 spec. " Bubye River S. Rhodesia 29/10/73, N.J.Duke ", in TMSA; 2 spec. "2553a. Satara, KNP, 19.I.82, Various soil and vegetation types, Doube & Macqueen, COLS01270 ", in SANC; 1 spec. "329, GORONGOSA N.P., Mozam. 11-28.I.72, Bornemissza & Kirk, COLS08747 ", in SANC; 1 ♂ spec. "BIRCHENOUGH BRIDGE, Rhod. (60 kms E) 18.I.74, H.H. Aschenborn, COLS08748 ", in SANC. All SANC specimens (except holotype) have two additional determination labels: Garreta lugens (Fairmaire, 1891), det. CSIRO, DBRU and Garreta lugens (Fairmaire, 1891), det. ALV Davis, 2013. Differential diagnosis. This new species can be easily separated from Garreta lugens by pronotal granulation; granules are clearly separate and more or less round in Garreta australugens new species, whereas those of G. lugens are often connected forming irregular shapes.

Garreta namalugens Davis & Deschodt, new species Figs 1���6 Description of holotype. Holotype ♂: 14.1 x 8.7 mm (length with head extended x maximum width). Habitus uniformly charcoal with dark blue or green undertone; undertone occasionally cupreous. Head. Clypeus with four denticles; right angled gap between middle two denticles; outside denticles rounded; clypeal surface with irregular granules, which are close together but clearly separated from one another. Pronotum. Pronotum surface very finely shagreened with small, discrete granules that are separated by more than one granule diameter. Sculpture-free longitudinal midline extends from the basal margin halfway up the disc of the pronotum. Elytra. Elytral interstriae with similar surface sculpture to the pronotum, i.e. very finely shagreened with small granules that are separated by more than one granule diameter. Striae are narrow, unclear, shagreened and apunctate. Anterior legs. Narrow and strongly curved inwards. Aedeagus. Similar to other species of Garreta (Fig. 3). Sexual dimorphism. Inward curving front legs of male type material are narrower than those of females. Terminal spur of front legs acute in females and more blunt in males. Variability. Very little size variation between the paratype specimens; colour varies from dark blue or green undertone to cupreous. Etymology. The name is a combination of the specific name of its closest relative Garreta lugens and the Nama people of Namibia. Type specimens examined. Holotype ♂: " NamibRand Nature Reserve, S 24.885184�� E16.065035 ��, 14.iii.2017, 1184m, Deschodt & Davis, Fresh Hyrax dung", in SANC. Paratypes: 18 spec. same as holotype, 10 in UPSA [9 kept in 99% Ethanol to allow for possible future molecular work], 5 in TMSA, 3 in SANC; 6 spec. " Spitzkoppe S21.829723�� E15.181811��, 17-19.iii.2017, Deschodt & Davis ", 2 in UPSA, 2 in TMSA, 2 in SANC; 1 spec. " Losberg, NamibRand Nature Reserve, S25.073295�� E16.048717��, 15.iii.2017 Deschodt & Davis, Hyrax dung midden", in UPSA; 1 spec. " NAMIBIA: KAOKOLAND, Otjihipa Mountains, base camp at 780 m, 17�� 16.4'S 12�� 28.8'E, 16-19.iii.2005 ��� E.Holm et al. ", in SANC. Differential diagnosis. G. namalugens new species, shows slightly smaller body-size than Garreta australugens new species and Garreta lugens; it also lacks the basal impressions on each side of the midline of the pronotum that are present in G. australugens new species and G. lugens. Habitat and distribution. The holotype and two paratype specimens of Garreta australugens, new species were sampled using a composite bait of pig, cattle and elephant dung on finer-grained, gabbro-derived soils in dry, open woodland at the South African Wildlife College on the west border of the Kruger National Park (Davis et al. 2012). Further paratype specimens are known from lowland localities in dry areas of northeast South Africa and southeast Zimbabwe as well as moist areas of central Mozambique. Specimens cited as G. lugens from South Africa, Mozambique and Botswana (Moretto & G��nier 2015) are probably also G. australugens new species, although reexamination is required to validate their identity. The authors recorded Garreta namalugens new species, flying abundantly on March 13-15th 2017, circa two weeks after substantial rainfall in Namibrand Nature Reserve. Activity was observed throughout the day. Live individuals were seen flying into crevices between huge granite boulders at the edge of the Nubib Mountains, presumably, in search of fresh dung of the Rock Hyrax (Procavia capensis (Pallas, 1766)). Bodies of dead individuals were excavated from silt under the accumulated dung in Hyrax middens. Although G. namalugens new species, was also attracted to pitfall traps baited with pig dung in the mountains, none was observed to fly into the adjacent sandy plains in order to feed on the dung of large herbivorous mammals, comprising either substantial, coarse-fibred, zebra droppings, or, dung pellets dropped by gemsbok or springbok. As regards species in the tribe Gymnopleurini, only Gymnopleurus andreaei Ferreira, 1954, and a few G. humanus Macleay, 1821, were observed on herbivore dung on the plains (ALVD and CMD, pers. observ.). So far, records for G. namalugens new species, have only been validated for west central Namibia with a single specimen from the Kaokoveld in northwest Namibia. The three species comprising Garreta australugens new species, G. lugens and G. namalugens new species, appear to represent a radiation between, now, geographically isolated, mostly dry lowlands in southwest, southeast and northeast Africa. However, confirmation of close relationships needs to be made using a molecular systematic analysis., Published as part of Davis, Adrian L. V. & Deschodt, Christian M., 2018, Two new species of Garreta Janssens, 1940 (Coleoptera: Scarabaeidae: Scarabaeinae) from Southern Africa, pp. 242-248 in Zootaxa 4450 (2) on pages 243-246, DOI: 10.11646/zootaxa.4450.2.4, http://zenodo.org/record/1444704

Namakwanus kamfferi Deschodt & Davis, new species (Figs. 1D, 3) Type material. Holotype female: Bloedkoppe, Namibia, S22.843909° E15.376124°, 17.iii.2017 739m, C. Deschodt & A. Davis (TMSA). One female paratype with same data as holotype (UPSA). Additional material examined. Various disarticulated body parts of different specimens from the same site as holotype, mounted on a single card; two elytra and two pronota from different beetles, Bloedkoppe, Namibia, S22.844586° E15.376308°, 745m 13.06.2013, Deschodt Jacobs Kamffer, collected dead from hyrax dung (UPSA). Description: Holotype female: Medium sized beetle. Measurements with head inflexed, 7.50 mm (length) × 5.01 mm (width) × 3.86 mm (height). Head. Clypeus bidentate, with two well-developed slightly upturned teeth, clypeal margin between teeth more or less straight. Teeth longer than distance between them. One very small tooth on lower clypeal margin, between dorsal teeth, while a second even smaller median tooth on the upper clypeal margin. Clypeogenal suture distinct and straight. Clypeal surface clearly punctate. Surface shagreened and clearly punctate. Pronotum. Convex, small but clear punctures separated by about one and a half to two lengths of their own diameter. Pronotal surface shagreened with sculpture free midline extending across pronotum. Elytra. Convex. Elytra fused. Striae very shallow and faint, faintly punctate. Irregular and very faint punctures on interstriae. Sterna. Prosternum shagreened with large shallow punctures. Meso- and metasternum fused, mesometasternal suture straight. Surface of mesosternum shagreened with clear, medium-sized punctures. Surface of metasternum shagreened with clear medium sized punctures. Protibia. Inner margin fairly straight with no denticles. Tibial apex blunt with short, spatulate downwardcurving spur close to inside corner. Tarsi attached under spur. Setae on inner margins of tarsi short and tan. Three teeth at apical outside margin of second half. First half of outer margin from joint smooth. Mesotibia. Fairly straight with smooth margins. Apically with two short spurs. Tarsi with short tan setae, ending with two claws. Metatibia. Fairly straight with smooth margins. Tarsi with short tan setae, ending with two claws. Pygidium. Surface shagreened with large, shallow and closely spaced punctures. Male. Unknown. Etymology. This species is named after Mr Dewald Kamffer to remind him of the day of its discovery when he was present in the flesh but very absent in other faculties. Diagnosis. The pronotal punctures of Namakwanus kamfferi new species are smaller than those of N. scholtzi Deschodt & Davis, 2017 and only of a single size. The most proximal denticle of the front tibia is much larger in Namakwanus kamfferi new species than in N. scholtzi. Distributions, ecology, conservation status and potential threats. This species may be restricted to the isolated Blutkoppe and Langer Heinrich Mountains in the Namib Desert, Namibia. As hyrax colonies remain protected in the area of collection within the Namib-Naukluft Park, N. kamfferi new species is probably not facing any current threat although the Langer-Heinrich Uranium mine lies within its predicted EOO and AOO. This species should be assessed as Data Deficient (DD) as the EOO is unknown. Its distribution area could be as small as 100 km 2 (combined area of the Blutkoppe and Langer Heinrich Mountains).

Namaphilus nubibmontanus Deschodt & Davis, new species (Figs. 1C, 2C���D, 3���4) Type material. Holotype male: Nubib Mountain, Namibia, S24.885184�� E16.065035��, 14.iii.2017 1184m, Deschodt, Jacobs & Davis (TMSA). Paratypes: Three specimens (unsexed), same data as holotype, (2 SANC, 1 UPSA); one female: Losberg, Namibia, S25.073295�� E16.048717��, 15.iii.2017, C. Deschodt & A. Davis (UPSA). Description: Holotype male: Medium sized beetle. Measurements with head inflexed, 5.22 mm (length) �� 3.90 mm (width) �� 2.80 mm (height). Head. Clypeus bidentate and surface shagreened; clypeal margin between teeth slightly concave. One very small tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture indistinct and fairly straight. Clypeal surface faintly punctate. Pronotum. Convex, and completely covered by small faint punctures, separated by about one and a half lengths of their own diameter. About half of the punctures associated with very short tan setae. Elytra. Convex with surface shagreened. Elytra fused. Striae unclear. Very faint punctures associated with short and recurved setae on interstriae, all distributed in double rows, except for interstriae adjoining elytral suture, where the punctures are aligned in a single row. Sterna. Prosternum shagreened with large shallow indistinct punctures, almost touching. Meso- and metasternum fused, mesometasternal suture slightly curved anteriorly. Surface of mesosternum shagreened with clear, medium-sized punctures, punctures separated by at least one puncture diameter. Surface of metasternum shagreened with a few very faint punctures. Protibia. Inside margin forming bulbous widening distal half of protibia and with no denticles, short tan setae present along the margin. Dorsal surface shagreened, irregularly punctate and with carina extending lengthwise. Tibial apex blunt with small spatulate spur. Tarsi attached under spur close to inner margin of tibia. Ventral surface shagreened with sinuous carina from inside margin of second half to middle of apex. Three teeth at apical third of outer margin, distal two thirds serrated. Mesotibia. Fairly straight, with smooth margins and with five rows of short tan setae. Apically with two spurs. Tarsi with short tan setae, ending with two claws. Metatibia. Fairly straight, with smooth margins and five rows of short tan setae. Tarsi with short tan setae, ending with two claws. Pygidium. Surface shagreened with large shallow punctures. Aedeagus. Parameres symmetrical, see Fig. 2. Female. There is no variation between males and females of this species. Etymology. This species is named after the Nubib Mountain range, the inselberg where the type series was collected. Diagnosis. This new species is similar in appearance to Namaphilus endroedyi (Deschodt, Davis & Scholtz, 2011) but the aedeagi (Fig. 2) are clearly different. Distribution, conservation status and potential threats. This species may be restricted to the isolated Nubib Mountains in Namibia. As hyrax colonies remain protected in the area of collection within the Namibrand Nature Reserve, N. nubibmontanus new species is probably not facing any current threats. However, it should be assessed as Data Deficient (DD) as the EOO is unknown. This species could encompass a geographic area as small as 1100 km 2 (area of the Nubib Mountains)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2018, New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia in Zootaxa 4450 (1), DOI: 10.11646/zootaxa.4450.1.4, http://zenodo.org/record/1444612, {"references":["Deschodt, C. M., Davis, A. L. V. & Scholtz, C. H. (2011) New dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) taxa from arid south-western Africa. Insect Systematics & Evolution, 42 (3), 277 - 294. https: // doi. org / 10.1163 / 187631211 X 568173"]}

Namakwanus minutus Deschodt & Davis, new species (Figs. 1E, 2E���F, 3���4) Type material. Holotype male: Nubib Mountain, Namibia S24.885184�� E16.065035��, 14.iii.2017 1184m, Deschodt, Jacobs & Davis (TMSA). Two female paratypes with same data as holotype (UPSA). Description: Holotype male: Medium sized beetle. Measurements with head inflexed, 4.93 mm (length) �� 3.86 mm (width) �� 2.75 mm (height). Head. Clypeus bidentate, with two well-developed upturned teeth, clypeal margin between teeth curved. One very small tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture fairly distinct and straight. Clypeal surface clearly punctate. Pronotum. Convex, surface shagreened, small and oval punctures covering the pronotal surface, punctures separated by about one to one and a half the length of their own diameter. Elytra. Convex, surface shagreened. Striae very shallow but visible, faintly punctate. Irregular and very faint punctures associated with short setae on interstriae. Sterna. Prosternum with large and very close punctures. Meso- and metasternum fused, mesometasternal suture straight. Surface of mesosternum shagreened with faint, medium-sized shallow punctures. Surface of metasternum shagreened with a few faint punctures. Protibia. Inside margin more or less straight with no denticles. Tibial apex blunt with a small spatulate spur. Tarsi attached under spur close to inner corner of tibia. Setae on inner margins of tarsi short. Apical outer margin of protibiae with three teeth. Distal third of outer margin somewhat serrated, middle third smooth. Mesotibia. Fairly straight, with smooth margins and five rows of short tan setae. Apically with two short spurs. Tarsi with short tan setae, ending with two claws. Metatibia. Fairly straight, with smooth margins and five rows of short tan setae. Tarsi with short tan setae, ending with two claws. Pygidium. Surface shagreened with large shallow punctures. Female. As for all Namakwanus species the female clypeus is slightly smaller than the male. Etymology. The specific epithet refers to its small size. Diagnosis. Namakwanus minutus new species is the smallest known Namakwanus species. Thus, it can easily be separated from these species as the other constituent species of Namakwanus are more or less the same larger size. Distributions, ecology, conservation status, and potential threats. This species may be restricted to the isolated Nubib Mountains in Namibia. As hyrax colonies remain protected in the area of collection within the Namibrand Nature Reserve, N. minutus new species is probably not facing any current threats. However, it should be assessed as Data Deficient (DD) as the EOO is unknown. Its distribution area could be as small as 1100 km 2 (area of the Nubib Mountains)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2018, New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia in Zootaxa 4450 (1), DOI: 10.11646/zootaxa.4450.1.4, http://zenodo.org/record/1444612

Ausmontins Deschodt & Davis, new genus (Figs. 1A, 2A���B, 3) Type species: Ausmontins jacobsi Deschodt & Davis, new species, here designated Head. Clypeus bidentate, with two fairly well-developed and forward-pointing teeth. Single minute tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture fairly distinct and straight. Pronotum. Highly convex. Antero-lateral margin somewhat flattened. Elytra. Convex and fused. Striae visible. Sterna. Meso- and metasternum fused, mesometasternal suture well defined, straight. Densely punctate with small punctures. Protibia. Three teeth at apical outside margin. Pygidium. Parameres surface shagreened with medium sized faint punctures. Aedeagus. Symmetrical. Phallobase with normal shape, parameres complexly shaped with three extending spines (see Fig. 2). Etymology. The gender is masculine. This genus is named after the Aus Mountains, Namibia, an isolated inselberg where the type species of the genus was collected. Diagnosis. Ausmontins new genus clearly falls within the tribe Deltochilini by being dorsally strongly convex, bearing three protibial teeth, the lateral margin of elytra being entire and by fore tarsi being present. It keys out to the genus Namaphilus in the key provided by Deschodt & Davis (2017). This new genus can be separated from Namaphilus with its standard aedeagus shape, by the very peculiar denticulate and angulate shaped parameres of the aedeagus and by its highly convex habitus., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2018, New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia in Zootaxa 4450 (1), DOI: 10.11646/zootaxa.4450.1.4, http://zenodo.org/record/1444612, {"references":["Deschodt, C. M. & Davis, A. L. V. (2017) Transfer of three species of Namakwanus Scholtz & Howden to Uersicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae). Annals of the Ditsong National Museum of Natural History, 7, 109 - 114."]}

Namaphilus tirasmontanus Deschodt & Davis, new species (Figs. 1C, 3���4) Type material. Holotype female: Tirasberg Conservancy, S26.231242�� E16.571429��, 13.iii.2017 1363m, C. Deschodt & A. Davis (TMSA). Additional material examined. Various disarticulated body parts of different specimens from the same site mounted on a single card (UPSA). Description: Holotype female: Medium sized beetle. Measurements with head inflexed, 5.84 mm (length) �� 4.44 mm (width) �� 3.32 mm (height). Head. Clypeus bidentate, with two small teeth that are well separated; clypeal margin between teeth straight with minute irregularities. Surface somewhat dull. One very small tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture indistinct but straight. Clypeal surface shagreened and punctate. Pronotum. Convex, coarsely shagreened, medium ovate punctures with associated short setae covering pronotum, punctures separated by less than one puncture diameter. Elytra. Convex. Elytra fused. Striae very shallow but visible, faintly punctate. Irregular and faint punctures with medium length associated setae in double rows on all interstriae except on those adjoining elytral sutures bearing a single row of punctures. Edges of punctures forming low undulations. Sterna. Prosternum with large shallow round punctures. Meso- and metasternum fused, mesometasternal suture straight. Surface of mesosternum shagreened with clear, medium-sized shallow punctures. Surface of metasternum shagreened with punctures slightly smaller and closer together than on mesosternum. Protibia. Inside margin slightly curved inward with no denticles. Tibial apex blunt with small spatulate spur next to inside corner. Tarsi attached under spur and close to inner margin of tibia. Three teeth at apical outer margin. Proximal two thirds of outer margin from joint serrated. Dorsal and ventral surface shagreened and punctate. Mesotibia. Fairly straight, with smooth margins and with five rows of short tan setae. Apically with two short spurs. Tarsi with short tan setae, ending with two claws. Metatibia. Fairly straight, with smooth margins and with five rows of short tan setae. Tarsi with short tan setae, ending with two claws. Pygidium. Surface shagreened with medium shallow punctures, some with associated setae. Male. Unknown. Etymology. This species is named after the Tiras Mountain range, the inselberg mountain complex where the holotype type specimen was collected. Diagnosis. The longer setae on the elytra and slight undulations on the elytra will distinguish this species from Namaphilus endroedyi and Namaphilus nubibmontanus new species. Distributions, ecology, conservation status and potential threats. This species may be restricted to the isolated Tiras Mountains in Namibia. As hyrax colonies remain protected in the area of collection within the Tiras Conservancy, N. tirasmontanus new species is probably not facing any current threats. However, it should be assessed as Data Deficient (DD) as the EOO is unknown. Its distribution area could be as small as 150 km 2 (area of the Tiras Mountains)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2018, New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia in Zootaxa 4450 (1), DOI: 10.11646/zootaxa.4450.1.4, http://zenodo.org/record/1444612

Ausmontins jacobsi Deschodt & Davis, new species (Figs. 1A, 2A���B, 3) Type material. Holotype male: Klein Aus Vista, Namibia, S26.660429�� E16.238527��, 12.iii.2017 1482m, D.H. Jacobs (TMSA). Paratypes: four specimens (one male and three unsexed), same data as holotype (1 SANC; 1 TMSA; 2 UPSA). Description: Holotype male: Medium sized beetle. Measurements with head inflexed, 7.37 mm (long) �� 5.57 mm (wide) �� 3.98 mm (high). Head. Clypeus bidentate, with two well-developed forward pointing teeth, area between teeth straight. One very small tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture fairly distinct and straight. Clypeal surface clearly and densely punctate with small punctures which are separated by about one puncture diameter. Pronotum. Convex; surface shagreened (or sharkskin like) with two distinct puncture types; (1) small, clear, regular ovoid punctures somewhat organised in lines each separated by about one puncture length and two to three puncture widths; (2) larger, clear round and deeper set punctures, each separated by about three to four puncture diameters. Elytra. Fused together, strongly convex with surface shagreened. Striae shallow but clearly visible, punctate with punctures about three to four puncture diameters apart. Punctures on interstriae forming somewhat regular rows, punctures about four puncture diameters apart, each with a single very short and thin recurved seta. Sterna. Prosternum surface shagreened and covered with medium punctures, punctures separated by less than one puncture diameter. Meso- and metasternum fused, mesometasternal suture more or less straight. Surface of mesosternum shagreened with small punctures, punctures separated by less than one puncture diameter. Metasternum same as mesosternum but punctures slightly smaller. Protibia. Inner margin for first two thirds more or less straight with no denticles, last third bulbous. Very short, fairly regular spaced setae on inner margin. Tibial apex blunt with small spatulate spur above tarsus. Underside of tibial apex with slightly downward curving spatulate process. Three teeth at apical third of outer margin. First two thirds of outer margin from joint irregularly serrated. Mesotibia. Slightly curving inward. Outer and inside margin smooth with single row of closely arranged tan setae. Dorsal surface flat and shagreened with single row of densely arranges tan setae, ventral surface with two rows of densely arranged tan setae. Metatibia. Slightly curving inward. Outside and inside margin smooth with single row of closely arranged tan setae, row of setae on inside less dense in first half. Dorsal and ventral surface same as for mesotibiae. Pygidium. Surface shagreened with medium sized faint punctures. Aedeagus. Parameres symmetrical, see Fig. 2. Female. No significant differences were noted. Etymology. This species is named after Dr Dawid Jacobs who collected the type series. Diagnosis. This species can easily be separated from all other hyrax associated deltochiline species by the following suite of characters: three teeth on outside fore tibial margin, short teeth on the clypeus, angulate and denticulate aedeagus and very convex body. It is larger than any known Namaphilus species with which it could otherwise be confused. Distribution, conservation status and potential threats. Because of flightlessness and probable low vagility, this species may be restricted to the isolated Aus Mountains just southwest of Aus in Namibia, an area that is used primarily for grazing of livestock and recreation. Although it may be dependent on the middens of hyrax colonies, a number remain in the area of collection. Therefore, A. jacobsi new species is probably not facing any current threats. However, it should be assessed as Data Deficient (DD) as the EOO is unknown. It could be as small as 120 km 2 (area of the Aus Mountains)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2018, New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia in Zootaxa 4450 (1), DOI: 10.11646/zootaxa.4450.1.4, http://zenodo.org/record/1444612

Byrrhidium convexum Scholtz & Howden, 1987. Widespread in Namaqualand from the environs of Nuwerus [31.149°S 18.358°E] in the south to Steinkopf [29.266°S 17.734°E] in the north. More or less confined to the Namaqualand Klipkoppe Shrubland vegetation unit as defined by Mucina and Rutherford (2006). Collected some distance from rock hyrax middens among Namaqualand heuweltjies (Mima-like earth mounds, Lovegrove and Siegfried, 1986), but probably also associated with rock hyraxes., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 113, DOI: 10.5281/zenodo.5562912, {"references":["SCHOLTZ, C. H. and HOWDEN, H. F., 1987. A revision of the southern African Cathonina [sic] (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of the Entomological Society of Southern Africa 50: 75 - 119.","MUCINA, L. and RUTHERFORD, M. C., eds, 2006. The Vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria, viii + 1 - 807.","LOVEGROVE, B. G. and SIEGFRIED, W. R., 1986. Distribution and formation of Mima-like earth mounds in the Western Cape Province of South Africa. South African Journal of Science 82: 432 - 436."]}

Namakwanus scholtzi spec. nov., Figs 1–2 SPECIMEN EXAMINED. Holotype ♂: [NAMIBIA]: Gobabeb, Kuiseb River, Central Namib, S. W. Afr., 11.v.1959, H. Dick Brown.– (TMSA). ETYMOLOGY. This species is named in honour of Professor Clarke H. Scholtz, for his extensive and lifelong contribution to entomology. DESCRIPTION. Dark brown, convex. Head: two long, distinct, slightly upturned clypeal teeth and one much smaller tooth medially on lower margin of clypeus. Slight red tinge laterally. Pronotum: punctate with round punctures of variable size, surface shagreened. Laterally clearly flattenedwith ridge in median one-third justinside of lateral margin. Ridge parallel with outside margin anteriorly and curving inward posteriorly. Lateral sides with slight red tinge. Protibia: surface shagreened, outside margin anteriorly tridentate in last one-third and slightly sinuous to dentate in first two-thirds. Outside two denticles larger than median denticle. Median dorsal ridge present, angulate at posterior denticle of outside margin. Golden brown setae present, longer terminally.Ventral surface withmedian ridge. Elytra: convex. Surface shagreened, punctures on second to fourth intervals more or less in straight double lines, punctures on all other intervals forming more or less straight single lines. Punctures and striae not well defined. Sterna: prosternum with clear round and adjacent punctures. Mesometasternal suture distinct and very slightly curved, punctures on mesosternum larger and further apart than on metasternum, surface shagreened. Fig. 1 Plate showing the habitus and aedeagus of one new species and habitus of all other species treated in this paper: A: Namakwanus irishi Scholtz & Howden,1987; B: Namakwanus scholtzi spec.nov.; C: Aedeagus of Namakwanus scholtzi spec.nov.; D: Namaphilus davisi (Deschodt & Scholtz,2007) comb.nov.; E: Namaphilus endroedyi (Deschodt,Davis & Scholtz,2011) comb.nov.; F: Namaphilus ameibensis spec.nov.; G: Versicorpus streyi (Frolov,2005) comb.nov.; H: Versicorpus erongoense Deschodt,Davis & Scholtz,2011. Fig. 2 Map showing the collecting localities of all the known species of the Byrrhidium group.The black outline indicates the distribution of rock hyraxes (shapefile supplied by IUCN). See online version for map in colour. Pygidium: surface shagreened and punctate, puncturessmalland further apartthantheirdiameter. DISCUSSION. This new species can be separated from N. irishi by the tips of the clypeal horns being slightly further apart, more upturned and slightly pointing inward at the apex., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on pages 110-112, DOI: 10.5281/zenodo.5562912, {"references":["SCHOLTZ, C. H. and HOWDEN, H. F., 1987. A revision of the southern African Cathonina [sic] (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of the Entomological Society of Southern Africa 50: 75 - 119.","DESCHODT, C. M., KRYGER, U. and SCHOLTZ, C. H., 2007. New taxa of relictual Canthonini dung beetles (Scarabaeidae: Scarabaeinae) utilizing rock hyrax middens as refuges in south-western Africa. Insect Systematics & Evolution 38: 361 - 376.","DESCHODT, C. M., DAVIS, A. L. V. and SCHOLTZ, C. H., 2011. New dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) taxa from arid south-western Africa. Insect Systematics & Evolution 42 (3): 277 - 294.","FROLOV, A. V., 2005. A new species of the dung beetle genus Namakwanus (Coleoptera, Scarabaeidae) from Namibia. In: KONSTANTINOV, A., TISHECHKIN, A. and PENEV, L., eds, Contributions to systematics and biology of beetles, papers celebrating the 80 th birthday of Igor Konstantinovich Lopatin, pp. 245 - 247. Pensoft, Sofia."]}

Dicranocara deschodti Frolov & Scholtz, 2003. Confined to the lower Boom River [28.013°S 17.063°E] in southern Namibia in close association with rock hyrax dung middens.

Dicranocara tatasensis Deschodt & Scholtz, 2007. Only recorded from the Tatasberg Mountains [28.323°S 17.241°E], Richtersveld National Park, South Africa, in close association with rock hyrax dung middens. Asingle female was collected live in the shade of boulders six months after the previous rains (C.M.D. pers. obs.)., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 113, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., KRYGER, U. and SCHOLTZ, C. H., 2007. New taxa of relictual Canthonini dung beetles (Scarabaeidae: Scarabaeinae) utilizing rock hyrax middens as refuges in south-western Africa. Insect Systematics & Evolution 38: 361 - 376."]}

Namakwanus irishi Scholtz & Howden, 1987. Known only from the four specimens in the type series. Not collected since its description in 1987. Most probably occurs in the mountains around Windhoek [c. 22.571°S 17.086°E]., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 114, DOI: 10.5281/zenodo.5562912, {"references":["SCHOLTZ, C. H. and HOWDEN, H. F., 1987. A revision of the southern African Cathonina [sic] (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of the Entomological Society of Southern Africa 50: 75 - 119."]}

Namaphilus ameibensi spec. nov. Known only from a single female specimen collected under a rock in the Erongo Mountains, Namibia.

Namaphilus endroedyi (Deschodt, Davis & Scholtz, 2011). Collected live in the Naukluft Mountains, Namibia,at least six months after the previous rain (C.M.D. pers. obs.). The holotype is currently curated by the South African National Collection of Insects [SANC] in Pretoria., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 114, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., DAVIS, A. L. V. and SCHOLTZ, C. H., 2011. New dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) taxa from arid south-western Africa. Insect Systematics & Evolution 42 (3): 277 - 294."]}

Namakwanus scholtzi spec. nov. Known only from a single specimen collected in 1959, most probably from the Kuiseb River Canyon near Barrowberg Mountain [23.658°S 15.477°E] close to Gobabeb in the central Namib Desert., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 114, DOI: 10.5281/zenodo.5562912

Versicorpus erongoense Deschodt, Davis & Scholtz, 2011. Known only from a single female specimen collected in 1975 from under a rock in the Erongo Mountains, Namibia., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 114, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., DAVIS, A. L. V. and SCHOLTZ, C. H., 2011. New dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) taxa from arid south-western Africa. Insect Systematics & Evolution 42 (3): 277 - 294."]}

Drogo stalsi Deschodt, Davis & Scholtz, 2016. Known from a single male specimen collected at Zebrafontein [27.750°S 16.883°E], about 30 km NNE of Rosh Pinah., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 113, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., DAVIS, A. L. V. and SCHOLTZ, C. H., 2016. A new monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa. Zootaxa 4147 (4): 490 - 500."]}

Namaphilus ameibensis spec. nov., Figs 1–2 TYPE MATERIAL. Holotype ♀: [NAMIBIA]: S. W. Afr[ica], Erongo Mt., Farm Ameib, 21.45S – 15.39E, 23.2.1975; E-Y:687, from under stones, leg. Endrödy & Schulze.– (TMSA). ETYMOLOGY. This species is named after its collecting locality. DESCRIPTION. Dark brown, convex. Head: two short and small slightly upturned clypeal teeth and one much smaller tooth medially on lower margin of clypeus. Clypeal teeth shorter than half the distance between them. Pronotum: punctate with punctures almost round medially becoming ovoid posteriorly and laterally, surface shagreened. Short, golden brown setae present at outside lateral margin. Protibia: surface shagreened, outside margin anteriorly tridentate in last one-third and smooth in first two-thirds. Median dorsal ridge present, almost parallel with outside margin. Golden brown setae present, longer and more numerous terminally. Ventral surface more or less flat, punctate. Elytra: convex. Surface shagreened, punctures on elytral interval more or less in single line, punctures on other intervals random. Punctures and striae well defined. Sterna: prosternum with clear, round punctures placed in close proximity. Mesometasternal suture distinct and straight, punctures on mesosternum larger and further apart than on metasternum. Metasternum with faint and shallow anterior pointing curve between coxae. Pygidium: surface shagreened and punctate, puncturessmalland furtherapart thantheirdiameter. DISCUSSION. This new species can be separated from all other species in the genus by being slightly larger and the edge between the clypeal teeth being curved and not straight as in the other species., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 110, DOI: 10.5281/zenodo.5562912

Deschodt, Christian M., Davis, Adrian L. V. (2017): Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae). Annals of the Ditsong National Museum of Natural History 7 (7): 109-114, DOI: http://doi.org/10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., DAVIS, A. L. V. andSCHOLTZ, C. H., 2011. New dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) taxa from arid south-western Africa. Insect Systematics & Evolution 42(3): 277-294.","DESCHODT, C. M., DAVIS, A. L. V. andSCHOLTZ, C. H., 2016.Anew monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa. Zootaxa 4147(4): 490-500.","DESCHODT,C.M., KRYGER,U.andSCHOLTZ,C.H., 2007. New taxa of relictual Canthonini dung beetles (Scarabaeidae: Scarabaeinae) utilizing rock hyrax middens as refuges in south-western Africa. Insect Systematics & Evolution 38: 361-376.","FROLOV, A. V., 2005. A new species of the dung beetle genus Namakwanus (Coleoptera, Scarabaeidae) from Namibia.In: KONSTANTINOV, A., TISHECHKIN, A. and PENEV, L., eds, Contributions to systematics and biology of beetles, papers celebrating the 80th birthday of Igor Konstantinovich Lopatin, pp. 245-247. Pensoft, Sofia.","FROLOV,A.V.and SCHOLTZ,C.H., 2003.A new genus and species of dung beetle from southern Namibia. African Entomology 11(2): 297-299.","IUCN (International Union for Conservation of Nature) In: IUCN 2012. IUCN Red List of threatened species. Version 2012.1. http://www.iucnredlist.org. Downloaded on 26 May 2015.","LOVEGROVE, B.G.and SIEGFRIED, W.R., 1986.Distribution and formation of Mima-like earth mounds in the Western Cape Province of South Africa. South African Journal of Science 82: 432-436.","MEDINA, C. A., 2015. Canthonini (Scarabaeidae: Scarabaeinae) of Gondwana: comparison of male genitalia, evolutionary trends and phylogeny.Ph.D. thesis, University of Pretoria, Pretoria, South Africa.xv + 1-125.","MORETTO, P., 2016. Description d'un nouveau Canthonini de Namibie appartenant au genre Dicranocara Frolov & Scholtz, 2003 (Coleoptera, Scarabaeidae).Entomologia Africana 21(2): 47-49.","MUCINA,L.and RUTHERFORD, M.C., eds,2006.The Vegetationof South Africa,Lesotho and Swaziland.Strelitzia 19.South African National Biodiversity Institute,Pretoria, viii + 1-807.","PALLAS, P.S., 1766.Miscellanea Zoologica. A.P.van Cleef, La Haye, 224 pp.","SCHOLTZ,C.H.and HOWDEN,H .F., 1987.A revisionof the southern African Cathonina [sic] (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of the Entomological Society of Southern Africa 50: 75-119."]}

Dicranocara inexpectata Deschodt & Scholtz, 2007. Confined to the Witputs area [27.573°S 17.147°E] in southern Namibia in close association with rock hyrax dung middens., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on page 113, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., KRYGER, U. and SCHOLTZ, C. H., 2007. New taxa of relictual Canthonini dung beetles (Scarabaeidae: Scarabaeinae) utilizing rock hyrax middens as refuges in south-western Africa. Insect Systematics & Evolution 38: 361 - 376."]}

NEW KEY TO THE GENERA AND SPECIES IN THE BYRRHIDIUM GROUP (adapted from Deschodt et al., 2016) 1 Foretibiawiththreeexternal denticles..... 2 — Foretibiawithtwoexternaldenticles...... 8 2 Habitus relatively large, longer than 10 mm and elongate Versicorpus Deschodt, Davis dScholtz......................... 3 — Habitus smaller than 10 mm long and convex................................ 4 3 Mesometasternal suture curving forward; punctures on prosternum large; Erongo Mountain.... V. erongoense Deschodt, Davis & Scholtz — Mesometasternal suture straight; punctures on prosternum small; Tsondab Mountain........................ V.streyi (Frolov) 4 Clypeal horns as long as, or longer than, distance between them....... Namakwanus Scholtz & Howden … 5 — Clypeal horns shorter than distance between them.. Namaphilus gen. nov. Deschodt & Davis … 6 5 Space between clypeal horns about one and a half times more at base than the width of a single horn at apex; clypeal horns not upturned and slightly pointing inward at the apex; Windhoek area................ N. iriSsi Scholtz & Howden — Spacebetween clypealhornsabouttwotimes greater at base than the width of a single horn at apex; clypeal horns more upturned and slightly pointing inward at the apex; Gobabebarea........ N. scholtzi spec. nov. 6 Area between clypeal teeth somewhat arcuate.......... N. ameibensis spec. nov. — Areabetweenclypealteethstraight...... 7 7 Tips of parameres with two notches in frontal view; Naukluft Mountains to the south of the Tsondab River canyon.... N. endroedyi (Deschodt,Davis & Scholtz) — Tips of parameres without notches in frontal view; HardapDam.... N. davisi (Deschodt & Scholtz) 8 Clypeus clearly bidentate with clypealhorns long...... Dicranocara Frolov & Scholtz … 9 — Clypeus quadridentate, with short medial clypealhorns....................... 12 9 Aedeagus without lateral horns on parameres............................. 10 — Aedeagus with lateral horns on parameres.................................. 11 10 Apex of parameresbulky; northern Richtersveld...... D. inexpectata Deschodt & Scholtz — Apex of parameres slender; southern Richtersveld, Boom River............. D. deschodti Frolov & Scholtz 11 Side horns on aedeagus extending somewhat sideways, tip curved downwards; Richtersveld National Park in South Africa.......... D. tatasensis Deschodt & Scholtz — Side horns on aedeagus slightly forward, tip not curved downwards; Fish River Canyon......... D. vandersmisseni Moretto 12 Clypealhorns short, mesometasternal suture not well defined; south of Orange River............. Byrrhidium Harold … 13 — Clypealhorns of medium length, mesometasternal suture well defined; north of Orange River...... Drogo stalsi Deschodt, Davis & Scholtz 13 Aedeagus with side protrusions, fore tibia with outer teeth parallel; Richtersveld in South Africa........... B. convexum Scholtz & Howden — Aedeagus plain with no side protrusions, fore tibia with outer teeth diverging; Namaqualand.................. B. ovale Harold NOTES ON THE CURRENTLY KNOWN SPECIES IN THE BYRRHIDIUM GROUP The Byrrhidium group is confined to the arid west of Namibia and South Africa (Fig. 2). It currently consists of six genera and 14 described species. These species (Deschodt et al., 2007, 2011, 2016) may mostly be associated with dung middens of rock hyraxes (Procavia capensis (Pallas, 1766)). As there are gaps in the known range of this group relative to that of rock hyraxes (Fig. 2), new related dung beetle species and genera may await discovery., Published as part of Deschodt, Christian M. & Davis, Adrian L. V., 2017, Transfer of three species of Namakwanus Scholtz & Howden to Versicorpus Deschodt, Davis & Scholtz or to Namaphilus gen. nov., with descriptions of two new species (Coleoptera: Scarabaeidae: Scarabaeinae), pp. 109-114 in Annals of the Ditsong National Museum of Natural History 7 (7) on pages 112-113, DOI: 10.5281/zenodo.5562912, {"references":["DESCHODT, C. M., DAVIS, A. L. V. and SCHOLTZ, C. H., 2016. A new monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa. Zootaxa 4147 (4): 490 - 500.","PALLAS, P. S., 1766. Miscellanea Zoologica. A. P. van Cleef, La Haye, 224 pp."]}

Drogo Deschodt, Davis & Scholtz, gen. n. Figs. 1, 2 and 4 Type species: Drogo stalsi Deschodt, Davis & Scholtz, sp. n. Head. Clypeus quadridentate, with two relatively well-developed slender forward-pointing teeth, and two outside teeth that are right angled at ends. One very small tooth on lower clypeal margin between dorsal teeth. Clypeogenal suture distinct and inward curving. Pronotum. Convex. Elytra. Convex. Striae visible. Sterna. Meso- and metasternum fused, mesometasternal suture well defined, somewhat curved anteriorly. Protibia. Two teeth at apical outside margin. Pygidium. Surface shagreened with big shallow punctures. Aedeagus. Symmetrical. Medial sclerotized plate serrated ventrally (Fig. 1 B, C). Diagnosis. Drogo clearly falls within the tribe Canthonini by being strongly convex, having no more than three protibial teeth, the lateral margin of elytra being entire and by having fore tarsi present. It falls within the Byrrhidium group as described above. This genus and Byrrhidium are the only genera in the group that have a quadridentate clypeus. However Drogo can be separated from Byrrhidium by having longer clypeal horns and the mesometasternal suture being well defined. The body shape of Drogo is also more rounded whereas it is slightly elongate in Byrrhidium. Drogo can be separated from Namakwanus by having two protibial teeth instead of three and having the clypeus quadridentate instead of bidentate as in Namakwanus. None of the other species in the Byrrhidium group has the saw-like serrations on the medial sclerotized plate of the aedeagus (note that the aedeagus of Versicorpus is unknown). Etymology. The gender is masculine. The aedeagus of Drogo reminds us of a sword thus the name is from contemporary fantasy after a powerful warlord known for his swordsmanship., Published as part of Deschodt, Christian M., Davis, Adrian L. V. & Scholtz, Clarke H., 2016, A new monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa, pp. 490-500 in Zootaxa 4147 (4) on pages 492-493, DOI: 10.11646/zootaxa.4147.4.9, http://zenodo.org/record/255041

Key to the genera and species in the Byrrhidium group Note: Some species can only reliably be separated by examination of the male genitalia. The females and fragmented body parts may thus not be identifiable. In those cases, identity may be obtained from geographical distribution, which is included in the key, and may suggest an identity. 1 Fore tibia with three external denticles................................................................... 2 - Fore tibia with two external denticles..................................................................... 6 2 Habitus fairly elongate; Erongo Mountain........................ Versicorpus erongoense Deschodt, Davis and Scholtz - Habitus convex........................................................ Namakwanus Scholtz and Howden... 3 3 Clypeal horns as long as, or longer than, distance between them; Windhoek and Gobabeb..... N. irishi Scholtz and Howden - Clypeal horns shorter than distance between them............................................................ 4 4 Apices of aedeagus not curving outwards terminally, with rows of long setae; Remhoogte Mountains to the south of the Tsondab River canyon...................................................................... N. streyi Frolov - Apices of aedeagus curving outwards terminally, without rows of long setae...................................... 5 5 Tips of parameres with two notches in frontal view; Naukluft Mountains to the south of the Tsondab River canyon........................................................................... N. endroedyi Deschodt, Davis and Scholtz - Tips of parameres without notches in frontal view; Hardap Dam........................ N. davisi Deschodt and Scholtz 6 Clypeus clearly bidentate with clypeal horns long.............................. Dicranocara Frolov and Scholtz ��� 7 - Clypeus quadridentate, with short medial clypeal horns....................................................... 9 7 Aedeagus with lateral clypeal horns on parameres that extend forward; punctures on pronotum fewer, faint and shallow; Rich- tersveld south of Orange River, in South Africa................................... D. tatasensis Deschodt and Scholtz - Aedeagus without lateral clypeal horns on parameres; punctures on pronotum numerous and obvious; Richtersveld north of Orange River, in Namibia.............................................................................. 8 8 Apex of parameres bulky; northern Richtersveld and Fish River Canyon............. D. inexpectata Deschodt and Scholtz - Apex of parameres slender; southern Richtersveld: Boom River........................ D. deschodti Frolov and Scholtz 9 Clypeal horns short, mesometasternal suture not well defined; south of Orange River Byrrhidium Harold.............. 10 - Clypeal horns medium length, mesometasternal suture well defined; north of Orange River....... Drogo stalsi gen. et. sp. n. 10 Aedeagus with side protrusions, fore tibia with outer teeth parallel; Richtersveld in South Africa................................................................................................. B. convexum Scholtz and Howden - Aedeagus plain with no side protrusions, fore tibia with outer teeth diverging; Namaqualand.............. B. ovale Harold, Published as part of Deschodt, Christian M., Davis, Adrian L. V. & Scholtz, Clarke H., 2016, A new monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa, pp. 490-500 in Zootaxa 4147 (4) on pages 497-498, DOI: 10.11646/zootaxa.4147.4.9, http://zenodo.org/record/255041