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Sisyphus alveatus Boucomont 1935 (Figs. 2, 3G). Boucomont 1935: 280; Haaf 1955: 344; Ferreira 1972: 792; Schaefer & Fischer 2001: 49; Montreuil 2015c: 2. Type locality: Kenya, Uasin Gishu, Size: Male: length: 5.0��� 2.7 mm, width: 2.5��� 2.1 mm. Female: length: 5.1��� 2.7 mm, width: 2.5���2.0 mm. Diagnosis: S. alveatus is similar to S. perissinottoi and S. manni. However, it is distinguished by the straight edge between the medial clypeal teeth, unlike in S. perissinottoi and S. manni where this feature is concave. S. alveatus also bears bare patches of setae on the pronotum, whereas in S. perissinottoi and S. manni the setae are uniformly distributed. Examined type material Holotype: (MHNH, through photograph without locality data). Examined non-type material: See Supplementary information. Distribution: In southern Africa, S. alveatus is known from forest at Popa Falls in Namibia and from open woodland in Central Mozambique (Fig. 4). It has also been recorded from miombo woodland and moist highland forest in East Africa (Montreuil 2015c). It is unlikely to occur in South Africa where records for S. alveatus (Paschalidis 1974, Montreuil 2015c) probably relate to other species, presumably, S. manni., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 17, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Boucomont, A. (1935) Coleoptera. VI. Scarabaeidae Scarabaeini. In: Jeannel, R. (Ed.), Mission Scientifique de l'Omo. Tome 2 Zoologie, Fascicule 16. Paul Chevalier & Fils, Paris, France, pp. 279 - 290.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp."]}

Sisyphus swazi Daniel & Davis sp. n. (Fig. 2F) ZooBank: http://zoobank.org/ C42533D7-5760-4187-8415-45608ED98DD0 Diagnosis: Sisyphus swazi sp. n is similar to S. alveatus in that both bear tufts of setae on the elytra. However, it differs by having a concave anterior margin between the medial clypeal teeth whereas in S. alveatus the margin is straight. In S. swazi sp. n the anterior margin of the pronotum is wider than the posterior margin, unlike in S. alveatus where the width is almost uniform. Pronotal setae arise from the centre of ocellate punctures in the new species, while in S. alveatus they are inserted between ocellate points. The interior edges of the metatibiae are weakly serrated in S. swazi sp. n whereas in S. alveatus they bear strong dentition. Examined type material Holotype: (♂ SANC): SWAZILAND (Mbabane) Ekuvinjelweni, 26.09353°S 31.30082°E, baited pitfalls, leg: Cebisile N. Magagula. Description: Male holotype: Size. 5.0 mm, width: 2.7 mm. Colour. body and setae brown; meso- and metasternum black. Head. Margin between clypeal medial teeth concave; dorsal surface of the clypeus setose and punctate; clypeo-frontal suture visible; vertex setigerous with ocellate points. Pronotum. Complete lateral prothoracic margin between the prothoracic disc and prothoracic episternum. Strong anterolateral emargination; anterior edge wider than posterior margin. Setae arise from the centre of ocellate punctures on the pronotal disc. Setae not uniform, but randomly arranged in patches; forming distinct bald spots on pronotal disc. Elytra. Short; narrow posteriorly; elytral striae minutely punctate and crenulate; double line of striae non-continuous, interrupted by fine ocellate strial puctures. Interstrial setae arranged in tufts. Hind wing and venation reduced to small sclerites (Fig. 2J). Pygidium. Ushaped and setigerous with ocellate punctures. Sternites. Abdominal sternites finely crenulate, setigerous with ocellate punctation; setae arranged in rows laterally; mesometasternal suture visible; punctate depression on the postero-medial surface of the metasternum. Legs. In ventral view, antero-lateral carina of profemur punctate, with a lateral row of fine and well developed setae; meso- and metafemur very small anteriorly, enlarged mid-posteriorly, densely punctate and setose; protibia with three teeth and a single terminal spine; mesotibia punctate and setose with two terminal spines; metatibia setose with weak indentation anteriorly, serrated laterally with two spurs; pro- meso- and metatarsi five segmented with two claws, setose laterally; first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the external edge. Aedeagus. Parameres symmetrical, somewhat truncated towards the apex. In ventral view, a knob-like structure present basally (Fig. 2F) Female: unknown. Distribution: The new species has only been recorded in shaded vegetation along the escarpment of the Lebombo Mountains in Swaziland (Fig. 4). Remarks: By showing a concave edge between the medial teeth of the clypeus and setae on the pronotum arising from the centres of ocellate points, Sisyphus swazi sp. n. is superficially similar to two other southern African species of the muricatus species-group: namely, S. manni and S. perissinottoi. However, both species differ from S. swazi sp. n. by having uniformly arranged setae on the pronotum, which lacks bald patches. Furthermore, both species bear a pronotum of uniform width, in contrast to S. swazi sp. n in which the latero-anterior margins of the pronotum are expanded. Etymology: Patronym is a noun in apposition, which reflects the name of the country (Swaziland) in which the holotype specimen was collected.

Sisyphus inconspicuus Daniel & Davis sp. n. (Figs. 11, 12D). ZooBank: http://zoobank.org/ A8C95127-D39B-4DC7-A564-0F77CCA4AB14 Diagnosis: Sisyphus inconspicuus sp. n. is similar to S. costatus; both bear inconspicuous ocellate punctation on the dorsal surface of the clypeus and frons, however, the vertex bears scattered ocellate punctures in S. costatus and dense ocellate points in the new species. Sisyphus inconspicuus sp. n. has a straight margin between the medial clypeal teeth and the anterior margins of the genae are arcuate. In S. costatus, the dorsal clypeal margin is distinctly concave and curved upwards between medial teeth whereas the genal margin is virtually straight (Figs. 11G, I). Additionally, the new species bears setae that are uniformly arranged on the elytra whereas, in S. costatus, the setae on the elytral interstriae alternate between rows that are dense or less dense. Description: Male holotype: Size. Length: 7 mm; width: 3.9 mm. Colour. Body and setae brown; meso- and metasternum black; antennae brown. Head. Medial teeth on the clypeus separated by a straight margin; shallowly notched edge between medial and lateral teeth; genal margin arcuate anteriorly. Dorsal surface of the clypeus and frons inconspicuously punctate (Fig. 11I); epicranial suture clearly visible; vertex setigerous with dense ocellate punctation. Pronotum. Convex; maximum length equal to maximum width; dorsal surface finely setose with ocellate punctation; setae on the antero-lateral projection well-developed. Complete lateral prothoracic ridge between the prothoracic disc and prothoracic episternum. Elytra. Wider and ovoid proximally, narrow posteriorly; elytral striae minutely punctate and crenulate, characterized by a distinct crenulate double line, which is interrupted by fine, ocellate strial puctures. Granulation and notches basally on elytral striae 1���4; setae evenly distributed on the interstriae. Hind wings and venation fully developed. Pygidium. Setigerous with ocellate punctation arranged in a U-shape. Sternites. Abdominal sternites finely crenulate, setigerous with ocellate punctation; setae arranged in rows laterally; meso-metasternal suture visible, acute laterally; meso- and metasternum densely punctate and setose; a punctate depression present on the postero-medial surface of the metasternum. Legs. In ventral view, antero-lateral carina of profemur punctate, internally pubescent with a lateral row of fine and well developed setae; meso- and metafemur with granulation, densely punctate and setose; meso- and metatrochanter contiguous with femur, metatrochanter slightly projected backwards; metacoxa punctate and shagreened ventroanteriorly; protibia with three teeth and single terminal spine; mesotibia punctate and setose with two terminal spines; metatibia densely setose, serrated laterally with two spurs; pro- meso- and metatarsi five segmented with two claws, setose laterally; first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the external edge. Aedeagus. Paremeres simple, symmetric; truncated basally (Fig. 12D). Morphological variation Size: Male: length: 8.2���6.0 mm; width: 3.8���4.1 mm; Female: 8.0���6.0 mm; width: 3.0��� 3.6 mm. Head: In some populations, mainly from the Eastern Cape (The Haven): lateral side of the clypeus is completely smooth as far as the clypeo-genal suture in some specimens; the lateral margin of the genae tends to be less convex; the surface of the clypeus is less setigerous. Colour: Inland populations are black and brown whilst the majority of the coastal population is black although two specimens from Pomene (Mozambique) are brown. Male: Meta- and mesotibia curved (Fig. 1H); last abdominal sternite narrowed medially (Fig. 1A). Female: Meta- and mesotibia almost straight (Fig. 1G); last visible abdominal sternite not constricted medially (Fig. 1B). Examined type material Holotype: (♂ TMSA): SOUTH AFRICA, KZN, Ithala Game Reserve, Ngubu; Dense woodland 27��32���55���S 31��13���32���E. 13.i.1999, dung baited pitfall; leg: Chown, McGeogh & Davis. Paratypes: (6♂, 2♀ SANC): with the same data as the holotype. (12♂, 9♀ UPSA) SOUTH AFRICA, KZN, Ithala Game Reserve 27��31���S 31��14���E, 12���14.i.1999, dung baited pitfall, leg: Davis. (3♂, 3♀ TMSA) SOUTH AFRICA, E. Transvaal, B Blyde River Canyon 24��35���S 30�� 48���E, 28.xi.1991, dung baited pitfall, leg: Klimaszewski. (3♂, 3♀ TMSA) SOUTH AFRICA, Transvaal, 20 km NE Thabazimbi 24.32��S 27.24��E, 23���24.iii.1985, leg: Vans. (1♂, 1♀ TMSA) SOUTH AFRICA, N. Transvaal, Waterberg, Farm 223; 24.11��S 27.50��E, 28.xi.1991, leg: Strydom. (1♂, 1♀ TMSA) SOUTH AFRICA, E. Transvaal, Barberton 16 km N, 25.44��S 30.59��E, 28.xi.1991, dung baited pitfall, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, Transvaal, Nelspruit, Nat. Res. Dry valley, 25.29��S 30.55��E, 09.ii.1987, dung baited pitfall, groundtraps 53 days, leg: Endr��dy-Younga. (1♂ TMSA; 1♂ SANC) SOUTH AFRICA, Transvaal, 20 km NE of Pretoria, Farm Roodeplaat, dung baited pitfall, 14���15.xi.1983, leg: Davis. (1♂, 1♀ TMSA) SOUTH AFRICA, E. Transvaal, Klaserie, 23.59��S 31.02��E. 3.v.1981, dung baited pitfall, leg: Endr��dy- Younga. (2♂, 2♀ TMSA) SOUTH AFRICA, Transvaal, Nelspruit, 18 km 25.37��S 30.58��E. 24.ix.1986, dung baited pitfall, groundtraps 31 days, leg: Endr��dy-Younga. (1♂, ♀ TMSA) SOUTH AFRICA, E. Transvaal, Nelspruit district, Farm de Hoop E, 24.ix.1986, dung baited pitfall, groundtraps, leg: Endr��dy-Younga. (1♂, 1♀ TMSA) SOUTH AFRICA, E. Transvaal, Mariepskop, 24.35��S 30.50��E, 2.v.1986, dung baited pitfall, groundtraps 5days, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, E. Transvaal, Nerina Nat. Res. 23.42��S 30.16��E, 2.v.1986, dung baited pitfall, groundtraps 50 days, leg: Breytenbanch. (1♂ TMSA) SOUTH AFRICA, Transvaal, Langjan Nature Res, 22.52��S 29.14��E, 10���20.i.1980, leg: Prinsloo. (13♂, 17♀ TMSA) SOUTH AFRICA, Kruger National Park, Sukuza, 1 km NW, 24.59��S 31.37��E, 22.i.1995, dung baited pitfall, groundtraps, leg: Endr��dy-Younga. (3♂, 5♀ TMSA) SOUTH AFRICA, Kruger National Park, Pafuri res. Camp, 22.25��S 31.12��E, 31.i.1994, dung baited pitfall, groundtraps 10 days, leg: Endr��dy-Younga. (1♂, 2♀ SANC) SOUTH AFRICA, Kruger National Park, Sukuza, 1 km NW, 24.59��S 31.37��E. 22.i.1999, cattle dung, leg: Inward. (1♂, 2♀ TMSA) SOUTH AFRICA, Kruger National Park, Punda Maria, 22.38��S 30.59��E, 11.ii.1994, elephant dung, leg: Endr��dy-Younga. (1♂ UPSA) SOUTH AFRICA, Kruger National Park, Sukuza, 24.00��S, 31.00��E, xii.1977, cattle dung, leg: Scholtz. (10♂, 13♀ TMSA) SOUTH AFRICA, N. Transvaal, Mmabolela Estates, 22.40��S 28.15��E, 10.iii.1973, groundtraps, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, Zululand, Sodwana Bay 10 km NW, 27.32��S 32.37��E, 24.xi.1992, groundtraps 5days, leg: Endr��dy-Younga. (7♂, 8♀ TMSA) SOUTH AF- RICA, Zululand, Ndumu Game Reserve, 26.54��S 32.17��E. 24.xi.1992, white rhino dung, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, KwaZulu Natal, Ngome State Forest, 27.48��S 31.25��E, 12���17.xi.1995, white rhino dung, leg: Endr��dy-Younga. (4♂, 6♀ SANC) SOUTH AFRICA, KZN, Mkuze Game Reserve, 24.xi.1981, legs: Doube, MacQueen, Davis & Flanagan. (4♂, 5♀ SANC) SOUTH AFRICA, KZN, St. Lucia Estuary, 26.viii.1978, leg: Bornemissza. (1♂, 3♀ SANC) SOUTH AFRICA, KZN, Hluhluwe Game Reserve (Inzimane), 24.xi.1982, leg: Doube. (4♂, 3♀ SANC) SOUTH AFRICA, KZN, St. Lucia Est. Nat. Res, forest, 23.i.1979, leg: Aschenborn. (9♂, 7♀ UPSA) SOUTH AFRICA, KZN, Ntshondwe, 27��43���S 31��15���E, 24.i.1999, leg: Davis. (2♂, 2♀ TMSA) SOUTH AFRICA, Limpopo, 17 km N Thabazimbi, Kransberg, 20.xii.2009, leg: Beyers. (1♂, 1♀ TMSA) MOZAMBIQUE, Pomene, 22.59��S 35.35��E, 02.v.1974, leg: Strydom. (1♂ UPSA) SOUTH AFRICA, Gauteng, Roodeplaat Nat. Res, 25��37-39���S 28��20-21���E. 13���14.xii.2001, leg: Davis & Deschodt. (1♂ UPSA) SOUTH AFRICA, Northwest Province, Rustenburg, 25��43���S 27��10���E. 27.ii.1999, leg: Wagemaker. (1♂ TMSA) SOUTH AFRICA, Zululand, Mtubatuba, 28.22��S 32.19��E, 3.iv.1974, fruit trap, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, Zululand, S. Natal, Weza, Ingeri Forest, 30.32��S, 29.41��E, 23.xi.1989, leg: Endr��dy-Younga & Klimaszew. (1♀ SANC) SOUTH AFRICA, Limpopo, Murmelende Waters Farm, Kampersrus, Mariepskop 24��32���S 30��17���E, 08���11.iv.1992, leg: Stals. (1♂, 1♀ TMSA) SOUTH AFRICA, Limpopo Prov, Lajuma, southern slope, 900 m, 16.i.2016, leg: Colin Schoeman. (1♂, 1♀ SANC) SOUTH AFRICA, Natal, Umfolozi, 1���7.x.1970, legs: Bornemissza & Aschenborn. (1♂, 1♀ SANC) SOUTH AFRICA, Cape Province, Avontuur (20 km S), 2.ii.1977, leg: Bornemissza. (3♂, 6♀ TMSA): SOUTH AFRICA, Eastern Cape, Transkei, The Haven, 32.15��S 28.55��E, 09.xii.1979, E-Y: 1696, groundtraps, 7 days, leg: Endr��dy-Younga. (2♂, 2♀ TMSA): SOUTH AFRICA, Eastern Cape, Transkei, Coast Dwesa forest, 32.17��S 28.50��E, 5.iii.1985, zebra dung, leg: Endr��dy-Younga. (1♂ TMSA) SOUTH AFRICA, Zululand, Lake Bangazi, 28.07��S, 32.31��E, 12.xii.1975, groundtraps, 5 days, leg: Endr��dy- Younga. (12♂, 8♀ UPSA): SOUTH AFRICA, NW Province, Mooinooi, Maretlwane Bush Camp (near to river) 25��47���07���S 27��34���37���E, 1274 m, 12���14.ii.2016, leg: Daniel. (1♂, 1♀ SANC) MOZAMBIQUE, Sofala, Beira (15 mi N), 19.i.1972, leg: Bornemissza & Kirk. (2♂, 2♀ UPSA) MOZAMBIQUE, Sofala, Gorongosa National Park (Cheringoma), near Claud���s waterfall Camp 19��01���43���S; 34��40���24���W, 1���3.v.2013, leg: B de Medeiros. (5♂, 6♀ UPSA) MOZAMBIQUE, Sofala, Gorongosa National Park (near Chitengo) Sandforest 18��57���24���S 34��20���24���E, 26 m, 14.vi.2014, pitfall, h. dung, leg: GM Daniel (1♂, 1♀ SANC): MOZAMBIQUE, Sofala, Dondo 19��37���S 34��45���E. 19.i.1972, leg: Bornemissza & Kirk. Etymology: The species name reflects the inconspicuous punctures on the frons. Distribution: Sisyphus inconspicuus sp. n. occurs in South Africa and Mozambique. It has been recorded from moist savanna or dense woodland and riverine vegetation. It has also been collected in dry shaded vegetation along the coastline, in sand forest, and in dry, dense savanna from KwaZulu Natal (South Africa) to Sofala Bay (Mozambique) (Fig. 13)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 32-35, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629

Sisyphus bicuariensis Daniel & Davis sp. n. (Fig. 3A). ZooBank: http://zoobank.org/ E02CDE50-8BB1-45D9-87A9-610A7C3713A0 Diagnosis: S. bicuariensis sp. n. resembles S. trichodichromicus Montreuil, 2015. Both bear bicoloured setae. However, S. trichodichromicus bears a shallow notch separating the medial teeth from the blunt lateral teeth; the clypeo-genal suture shows an obtusely angled cleft; and the genae are simple and arcuate. This differs to S. bicuariensis sp. n., where a markedly deep notch separates the medial teeth from the sharp lateral teeth; the clypeo-genal suture shows a rectangular cleft; and the genae are long and virtually parallel. Additionally, the shapes of the parameres are different in both species. Description: Male holotype: Size. Length: 4.2 mm; width: 2.2 mm. Colour. Black body with grey and black setae (Fig. 5G); clypeus shiny brown antero-laterally; meso- and metasternum black with brown and white setae; antennal club brown. Head. Clypeus upcurved antero-laterally; margin between medial teeth straight; a deep notch between medial and lateral teeth. Clypeo-genal suture with a rectangular cleft; genae long and virtually straight. Frons and vertex with dorsal ocellate punctation and setation. Frons declivous dorso-medially; epicranial suture visible; vertex with a few granules antero-medially. Pronotum. Convex with maximum width longer than maximum length; disc bearing three depressions and ocellate punctation; setae black and grey medially on the central disc, grey laterally; setae on pro-episternum grey, well-developed antero-laterally. A complete lateral prothoracic ridge between the pronotal disc and pro-episternum. Elytra. Narrow posteriorly bearing thicker black setae alternating with thin and thick grey setae either in equal proportion (1:1) or with less grey setae; weakly developed tufts of setae protrude from the epipleurae below the edge of the elytra (Hind wing. Wing and venation fully developed. Pygidium. Bearing scattered setae and ocellate punctation; narrow basally. Sternites. Margin between abdominal sternites finely crenulate; setation well-developed laterally, less so medially; punctation dense laterally and sparse medially; meso-metasternal suture visible; meso- and metasternum densely punctate with matted setae antero-laterally; metasternum with a punctate depression postero-medially. Mesepimerum and metepisternum finely punctate and setose. Legs. In ventral view, profemur punctate and pubescent, carinate antero-laterally with a lateral row of fine and well-developed setae; in lateral view, meso- and metafemur with granulation, also densely punctate and setose; meso- and metatrochanter contiguous with femur, metatrochanter larger and slightly projected backwards; metacoxa punctate and shagreened ventro-anteriorly; protibia with three teeth and a single terminal spine; mesotibia punctate and setose with two terminal spines; metatibia densely setose, serrated laterally with two spurs; pro-, meso- and metatarsus five-segmented, with two claws, setose laterally; first tarsal segment of meso- and metathoracic legs with a row of dense setae on the external margin. Aedeagus. Phallobase slightly curved. Median lobe clearly visible. Parameres (Fig. 6A). Morphological variation Size: Male: length: 5.0–4.0 mm; width: 2.6–2.0 mm; Female: length: 4.9– 4.1 mm, width 2.7–2.0 mm. Male. Meta- and mesotibia curved; with a carina on the dorso-posterior edge of the meta- and mesofemur; last abdominal sternite narrowed medially (Fig. 1A). Female. Meta- and mesotibia almost straight; lacking a carina on the dorso-posterior margin of the meta- and mesofemur; last visible abdominal sternite two times wider than in males (Fig. 1B). Examined type material Holotype: (♂ SANC) ANGOLA, Bicuari NP (15.375173°S 14.752320°E) (2 km N Camp); 18.xii.1974; leg: Davis & Temby /1671/, / Ex coll. CSIRO, Div. Entomology, South Africa Station /. Paratypes: (20♂, 16♀ SANC) with the same data as the holotype. (10♂, 6♀ SANC) ANGOLA, Caconda (37 & 30 km SW); 21.xii.1974; leg: Davis & Temby; / 1678/, / Ex coll. CSIRO, Div. Entomology, South Africa Station /. (2♂, 3♀ SANC) ANGOLA, Sá da Bandeira (11 & 3 km W), 29.xii.1974; leg: Davis & Temby /1691/, / Ex coll. CSIRO, Div. Entomology, South Africa Station /. (2♂, 2♀ SANC) ANGOLA, Paiva Coucero (50 & 45 km E); 18.xii.1974; leg: Davis & Temby, /1671/, / Ex coll. CSIRO, Div. Entomology, South Africa Station /. (1♂, 1♀ SANC) ANGOLA, Huila; 22.xii.1974, leg: Davis & Temby /1683/, / Ex coll. CSIRO, Div. Entomology, South Africa Station /. Additional examined type and non-type material: S. trichodichromicus (Holotype: MHNH, through photograph without locality data); (3♂, 4♀ TMSA) ZAMBIA, Liuwa Plains; 14.643259°S 22.626423°E, 29–30.xi.2003, 1050 m, leg: Deschodt & Groenewald. (2♂, 4♀ SANC) ZAMBIA, idem; (2♂, 2♀ UPSA) ZAMBIA, with the same data. Etymology: Patronym is a noun in apposition, which reflects the name of the area in which the majority of type specimens were collected: Bicuari National Park, South Angola. Distribution: The new species may be associated with unshaded vegetation and open woodland in southern Angola (Fig. 7).

Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L., Scholtz, Clarke H. (2020): Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions. Insect Systematics & Evolution 51 (1): 1-61, DOI: 10.1163/1876312X-00002195

Neosisyphus tembyi Daniel & Davis sp. n. (Figs. 7, 8C) ZooBank: http://zoobank.org/ 476EBD87-13C2-4DE0-A31E-883EEC69DA66 Diagnosis: N. tembyi sp. n. is similar to Neosisyphus jossoi Montreuil, 2015. Both species are roughly the same size and lack sharp spines on the posterior margin of the metafemur. However, N. tembyi sp. n. bears a small angled protuberance on the midposterior margin of the metafemur in males. The metatrochanter projection in major males of the new species is ½ of the total length of the metafemur and is somewhat inwardly projected at the tip. Unlike in N. jossoi, in which the metatrochanter projection is ¾ of the total length of the metafemur in major males and strongly inwardly projected. Southern African specimens of N. setiger Roth (1851) differ from the new species by bearing a shorter metatrochanter in males, which is ¼ of the total length of the metafemur. Furthermore, the metatrochanter in Southern African N. setiger projects straight and is never bent on the tip. The phallobase and parameres of all of the above species are distinctly different. Description: Male Holotype: Size. Male: length: 7.3 mm; width: 4.5 mm. Colour. Black body with brown setae; meso- and metasternum black; antennal club brown. Head. Anterior edge of clypeus with shallow sinuosity between medial teeth; lateral extreme of clypeal margin simply arcuate. Clypeo-genal suture markedly cleft; genae elongate. Frons and vertex dorsally granular, setose and minutely punctate, epicranial suture clearly visible. Pronotum. Convex with a sheen on dorsal surface; maximum width as long as the maximum length; setae inserted between ocellate punctures. Incomplete lateral prothoracic ridge between pronotal disc and prothoracic episternum. Elytra. Broad proximally and narrow posteriorly; well-spaced setae on interstriae; elytral surface shiny, distinct antero-lateral humeral callus; striae fine with double marginal lines interrupted by smooth ocellate strial punctures; granulation basally on elytral striae 4–7. Pygidium. Bearing U-shaped, ocellate punctation, a metallic sheen and setation. Sternites. Margins between abdominal sternites setigerous laterally, and glabrous medially with minute ocellate punctation; meso-metasternal suture visible; meso- and metasternum finely punctate and setose; metasternum with a punctate depression on the postero-medial edge. Mesepimeron and metepisternum finely punctate and setose. Legs. In ventral view, profemur carinate antero-laterally, punctate and internally pubescent with a lateral row of fine and well-developed setae; mesofemur finely punctate and setose, lacking any projection posteriorly; metafemur with an angled-protuberance mid-posteriorly with fine punctation and setae; metatrochanter projected backwards almost reaching top of angled protuberance; tip of metatrochanter slightly project- ed inward, but always parallel with metafemur. Metacoxa punctate and shagreened ventro-anteriorly. Protibia with three teeth and single terminal spine; meso-metatibia densely setose, serrated laterally, with two terminal spines on mesotibia, and two terminal spurs on metatibia; pro-, meso- and metatarsus five-segmented, with two claws; mostly setose. Aedeagus. Parameres separated by visible dorsal membranous portion extending nearly to the middle; arciform ridge bordering the membrane laterally, deeply notched medially; inwardly curved on the apex (Fig. 18C). Morphological variation Males bear a backwardly projected metatrochanter that is curved inwardly at the tip; this projection is absent in females where the metatrochanter is contiguous with the metafemur. The pronotum in males is larger than in females. Examined type material Holotype: (1♂ SANC) / Angola, Paiva Coucero (14.816667°S 15.55°E) (50 & 45 km E), 18.xii.1974, leg: Davis & Temby /, / Ex coll CSIRO, Div. Entomology S. African Station /. Paratypes: (1♀ SANC) same data as holotype. Distribution: The species is only known from woodland vegetation in southern Angola (Paiva Coucero) (Fig. 19). Etymology: The patronym is named after Ian Temby, a former researcher at the CSIRO, Division of Entomology, South African station; who was one of the collectors of the type material.

Sisyphus manni Montreuil, 2015 (Fig. 2E). Montreuil 2015c: 2���3 Type locality: Limpopo, Kruger National Park. Size: Male: length: 5.5���4.0 mm; width: 2.5���2.0 mm. Female: length: 5.5���4.0 mm, width 2.5���2.0 mm. Diagnosis: In S. manni, proximal elytral setae are primarily distributed uniformly becoming arranged in sparse tufts mid-basally and posteriorly, unlike in S. perissinottoi where dense tufts of setae occur across the entire elytra. S. manni bears three depressions on the pronotal disc, which are lacking in S. perissinottoi. Examined type material Holotype: (MHNH, through photograph without locality data). Examined non-type material: See Supplementary information. Distribution: S. manni is only known from South Africa. The species is associated with upland to highland grassland and open woodland in Gauteng (cited as S. alveatus by Davis et al. 2005), KwaZulu Natal, Mpumalanga and Northwest Provinces. It has also been collected in unshaded riverine vegetation in Limpopo (Fig. 4)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 15-16, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Davis, A. L. V., Scholtz, C. H., & Deschodt, C. (2005) A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna. African entomology, 13 (1): 1 - 16."]}

Sisyphus oralensis Daniel & Davis, 2016 (Figs. 14B, E; 15A). Daniel et al. 2016: 70 Size: Male: length: 3.4���4.8 mm, width: 2.1���2.5 mm. Female: length: 3.9���4.7 mm, width: 2.1���2.5 mm. Type locality: Richard���s Bay (KwaZulu Natal, South Africa) Diagnosis: S. oralensis resembles Sisyphus auricomus sp. n. However, it differs by the less dense cover of long yellow setae on the pronotal disc that are not distributed uniformly, but, mostly arranged in a linear pattern separated by bare patches (Fig. 14E). Furthermore, parameres of S. oralensis are notched dorso-transversally (Fig. 15A). Examined type material Holotype: (♂ TMSA): SOUTH AFRICA, KZN, Richard���s Bay 28��39���29.1���S 32��15���19.2���E, 27.i.2000, leg: Davis & Delport. Paratypes: (5♂, 8♀ UPSA): SOUTH AFRICA, KZN Richard���s Bay 28��39���S 32��15���E, 26���27.i.2000, leg: Davis & Delport. (4♀ UPSA; 3♂, 2♀ SANC): SOUTH AFRICA, KZN, Thembe Elephant Park 27��01���S 32��24���E, 17.xii.1996, leg: B.J. Van Rensburg. (9♂, 8♀ UPSA): SOUTH AFRICA, KZN, Thembe Elephant Park 27��01���S 32��24���E, 17.vii.1995, leg: B.J. Van Rensburg. (2♂, 3♀ UPSA): SOUTH AFRICA, KZN Thembe Elephant Park 26��55���S 32��23���E, 15���30.x.2008, leg: B.J. Van Rensburg. (2♂ SANC, 1♀ TMSA): SOUTH AFRICA, KZN, Umfolozi 28��15���S 31�� 55���E, 25.viii.1971, leg: Bornemissza & Insley. (1♂, 10♀ SANC): SOUTH AFRICA, KZN St Lucia Est. Nat. Reserve, Dune Forest 28��13���S 32��21���E, 27.x.1978, leg: Bornemissza. (1♀ SANC): SOUTH AFRICA, KZN Ntl Kosi Bay Nature Res 26��34���S 32��28���E, 8���11.ii.1990, leg: B. Grobbelaar. (1♀ SANC): SOUTH AFRICA, KZN, Near Richards Bay, 28��36���52���S 32��17���34���E, 27.x.1978, leg: Bornemissza. (1♂ SANC): SOUTH AFRICA, KZN, Thembe Elephant Park 27��01���S 32��24���E, 01���49. ii.1996, leg: Stals. (5♂, 6♀ TMSA): SOUTH AFRICA, KZN, Ntl Kosi Bay Nature Res 26��34���S 32��28���E, 14.xi.2002, leg: Burger, Harrison & Muller. (7♂, 8♀ TMSA): SOUTH AFRICA, Zululand, Sodwana Bay 5km, 27��21���S 32��23���E, 23.xi.1992, leg: Endr��dy-Younga. (8♂, 8♀ TMSA): SOUTH AFRICA, N Zululand, Ndumu Game Reserve 26��32���S 32��10���E, 1.xii.1992, leg: Endr��dy- Younga. (30♂, 42♀ TMSA): SOUTH AFRICA, N Zululand, Lake Bangazi 28��04���S 32��18���E, 12.xii.1992, leg: Endrodi-Younga. (1♂, 2♀ TMSA): SOUTH AFRICA, Zululand, St. L��cia. Mission Rock 28��13���S 32��21���E, 23.xi.1992, leg: Endr��dy-Younga. (1♂, 2♀ TMSA): SOUTH AFRICA, Natal, Cape Vidal, Forest litter 28��8���S 32��33���E, 23.i.1990, leg: J. Klimaszewski. (1♂, 2♀ TMSA): SOUTH AFRICA, N Zululand, Hluhluwe Game Reserve. 28��03���S 32��02���E, 20.xi.1992. leg: Endr��dy-Younga. (12♂, 30♀ SANC); SOUTH AFRICA, KZN St Lucia Est. Nat. Reserve, Forest 28��13���S 32��21���E, 24���23.i.1979, leg: NH Aschenborn. (1♂, 2♀ TMSA): MOZAMBIQUE, Inhambane, Pomene, 22��35���S 35��21���E, 04.v.1974, leg: A. Strydom. (5♂, 5♀ UPSA): MOZAMBIQUE, Maputo Elephant Reserve 26��39���S 32��43���E, 10���16.xi.2007, leg: W. Str��mpher & C. Deschodt. Examined non-type material: See Supplementary information. Distribution: S. oralensis is a coastal and sandforest endemic (Jacobs et al. 2010; Daniel et al. 2016). It has been observed in dense coastal woodlands and forest from Maputaland, north-east South Africa, to south-east Mozambique (Maputo Elephant Reserve and Pomene) (Fig. 16). Furthermore, recently, we found new distribution records from uMkhuze Game Reserve in South Africa (see Supplementary information). The species was collected in shaded deep sand vegetation, under pitfalls baited by dead diplopods. Remarks: Revision of S. oralensis type material has revealed that specimens from Mozambique (Dondo, Beira and Gorongosa) with distinct golden hair on the pronotum belong to a new species of the seminulum species-group that is described below. In addition, it should be noted that in the original description of S. oralensis (Daniel et al. 2016), three specimens (a male and two females) from Ithala Game Reserve (Kwa- Zulu Natal, South Africa) were cited in error as paratypes of S. oralensis. These same three specimens were also cited correctly as paratypes of S. umbraphilus in the same publication., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 37-38, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Jacobs, C. T., Scholtz, C. H., Escobar, F., & Davis, A. L. V. (2010) How might intensification of farming influence dung beetle diversity (Coleoptera: Scarabaeidae) in Maputo Special Reserve (Mozambique)? Journal of Insect Conservation, 14 (4): 389 - 399."]}

Sisyphus costatus (Thunberg, 1818) (Figs. 11, 12B). Peringu��y 1901:100; Gillet 1912:22; Arrow 1927:458; Haaf 1955:355���6; Ferreira 1972: 794���5; Harold 1869:984 Ateuchus costatus Thunberg, 1818 Thunberg 1818:412 Size: Male: length: 5.3��� 4.8 mm, width: 3.0���2.0 mm. Female: length: 5.3��� 4.2 mm, width: 2.6��� 2 mm. Type locality: (Cape of Good Hope) South Africa Diagnosis: S. costatus is close in appearance to S. australis sp. n. In both species, the elytral interstriae bear alternating rows of well-developed or less-developed setae (Fig. 11F). However, S. costatus is distinguished by an upcurved anterior clypeal edge and a concavity between the medial clypeal teeth. In addition, it lacks a carina between the clypeus and the frons (Fig. 11G). Examined type material (1♀ SMNH) Lectotype (designated here): Red label / Uppsala University Zoology Museum; Thunberg, saml.nr. 3206., Ateuchus costatus, Cap. minutus. F. TYP/ / Lectotype, det. GM Daniel, 2016 /. Examined non-type material: See Supplementary information. Distribution: S. costatus has been recorded in moist upland savanna and highland grassland (Davis et al. 1999, Davis et al. 2005) from the Eastern Cape to the north of South Africa (Fig. 13). Remarks: The species was described by Thunberg (1818) as Ateuchus costatus. Gillet (1912) transferred Thunberg���s species to the genus Sisyphus., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 29-30, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Thunberg, C. P. (1818) Coleoptera Capensia, Antennis Lamellatis, sive, clava fissili instructa, descripta, memoires de l'Acad. T. VI.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Davis, A. L. V., Scholtz, C. H., & Chown, S. L. (1999) Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa. Journal of Biogeography, 26 (5): 1039 - 1055.","Davis, A. L. V., Scholtz, C. H., & Deschodt, C. (2005) A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna. African entomology, 13 (1): 1 - 16."]}

Sisyphus genierorum Montreuil, 2015 (Figs. 5, 6C). Montreuil 2015a: 91, 93, 99���100 Type locality: Quirimbas National Park, Mareja (Cabo Delgado, Mo��ambique) Size: Male: length: 10���8.0 mm, width: 5.5��� 5 mm. Female: length: 9.5��� 8.5 mm, width: 5.0��� 4.7 mm. Diagnosis: S. genierorum resembles S. sordidus. In both species, tufts of epipleural setae are visible from above at the outer margin of the elytra (Fig. 5H); also, pronotal setae arise from the centre of ocellate punctures. However, S. genierorum differs by the much finer and uniformly arranged setae on the elytral interstriae. It is also much larger than S. sordidus. Examined type material Paratypes: (2♀ CMN) MOZAMBIQUE: / Cabo Delgado, Taratibu (site 3), P.N. Quirimbas, 340 m; 12��48���02���S 39��41���49���E, 8.i.2013, eastern Miombo woodlands, dung trap 24 h. F & S. G��nier. 2013-47 /; /World Scarab database WSD00024494/, Sisyphus genierorum n. sp.; O. Montreuil d��t. 2014. (1♀, 3♂ CMN) MOZAMBIQUE: / Cabo Delgado, Mareja (site 2), P.N. Quirimbas, 200 m; 12��51���41���S 40��09���31���E, 24.xii.2012, eastern Miombo woodlands, dung trap (shaded), leg: F & S. G��nier. 2012-08 /; /World Scarab database WSD00024355/, / Sisyphus genierorum n. sp.; O. Montreuil d��t. 2014/. Distribution: S. genierorum is currently only known from Quirimbas National Park (Cabo Delgado, Mozambique) (Fig. 7), where it is associated with the Eastern Miombo Woodlands ecoregion of Olson et al. (2001)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 22, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V., Underwood, E. C., & Loucks, C. J. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth: A new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience, 51 (11): 933 - 938."]}

Sisyphus caffer Boheman, 1857 (Figs. 5, 6B). Boheman 1857: 195���6; Harold 1869: 984; P��ringuey 1902: 106���7; Haaf 1955: 362���4; Ferreira 1972: 82, Sch��fer & Fischer 1992:121; Sch��fer & Fischer 2001:53 Size: Male: length: 7.8��� 6.9 mm, width: 4.4��� 3.5 mm. Female: length: 7.6��� 7.1 mm, width: 4.5���3.0 mm. Type locality: Caffraria (South Africa). Diagnosis: S. caffer is close in appearance to S. sordidus Boheman, 1857. However, they can be separated morphologically by the following features: S. caffer bears much finer elytral setae, which are distributed in single rows; S. sordidus bears thicker elytral setae, which are distributed in non-single rows forming bunches. In S. caffer, pronotal setae are inserted between ocellate punctures (Fig. 5F), whereas in S. sordidus, the setae originate from the centre of ocellate punctures (Fig. 5H). Furthermore, S. sordidus occurs in unshaded lowland vegetation associated with warm temperatures whereas S. caffer is associated with highland grassland under cooler conditions. Examined type material Holotype (photograph): (1♂ NHRS): /type/; /Caffraria/; /leg: J.Wahlb /; /309; 72/; /244; 73/; / S. caffer Bohm /; red label: /typus/; white label (handwritten): /TYPE Sisyphus caffer BOHEMAN, det. J. Ferrer /; /Naturhistoriska, Riksmuseet Stockholm; loan 37/90/; /9385 E92+/; /NHRS-SRAH 000000221/. Examined non-type material: See Supplementary information. Distribution: S. caffer occurs in South Africa, Namibia, Lesotho and Bostwana. The species has been recorded in upland and highland grasslands under cooler conditions (Davis et. al. 2005) in both South Africa and Lesotho (29.7��S 27.4��E). In South Africa, the localities extend northwards from the Eastern Cape and eastern Free State to the latitude of Pretoria (28.19��E 25.80��S) and also along the eastern escarpment of KwaZulu Natal and Mpumalanga. The lowlands beyond the northern Gauteng Mountains of the Witwatersrand (24.54��S 27.08��E) may act as barriers, although the species is also found in isolated uplands further to the northwest, such as the Blouberg. S. caffer has also been recorded in unshaded vegetation at the margins of its range in Kneneng (Botswana) (23.78��S 25.27��E) and in a dry upland outlier in the north of Namibia (Kaoko Otavi) (Fig. 7). Kaoko Otavi represents a disjunction in the distribution of S. caffer from its main range on the Highveld of South Africa. It is noteworthy that N. macrorubrus and Proagoderus lanista (Castelnau, 1840) also show similar disjunct occurrences on the South African Highveld and as rarities in north Namibia (Davis 1997). Remarks: We cannot confirm or refute the validity of S. crispatus because the type specimen has been lost. Furthermore, the original description and image presented by Gory (1833) are imprecise and insufficient to determine the identity of the species. S. crispatus was described from the same imprecise type locality (Caffraria) as two other species (S. caffer and S. sordidus). Under these circumstances, we are not confident to state which of the two species might be a junior synonym of S. crispatus. According to the ICZN, article 75.5 ���when an author considers that the taxonomic identity of a nominal species-group taxon cannot be determined from its existing name-bearing type, its name is a nomen dubium ���. Therefore, we propose S. crispatus as a nomen dubium., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 20-21, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Schafer, P. & Fischer, E. (2001) Zur Scarabaeidenfauna Rwandas, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini, Mitteilungen des internationalen entomologischen Vereins supplement 9: 1 - 93.","Davis, A. L. V. (1997) Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.). african Journal of ecology, 35 (1): 10 - 38.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p."]}

Sisyphus Latreille 1807 Latreille 1807: 79; Gory 1833: 1���15; Kolbe 1914: 317; Harold 1869: 984���5; P��ringuey 1901: 98���108; Gillet 1911: 22���25; Arrow 1909: 517���519; Arrow 1927: 456���465; Haaf 1955: 341���381; Balthasar 1968: 954, Ferreira 1972: 789���844; Paschalidis 1974b: 299���303; Endr��di 1983: 207���210; Sch��fer & Fischer 1992: 119���137; Sch��fer & Fischer 2001: 1���93; Montreuil 2014: 177���180; Montreuil 2015a: 91���100; Montreuil 2015b: 1���2; Montreuil 2015c: 1���13; Montreuil 2016: 167���174; Daniel et al. 2016: 67���74; Montreuil 2017: 1���22, Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 11, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Latreille, P. A. (1807) Genera Crustaceorum et Insectorum; Secundum ordinem Naturalem in familias disposita, inconubus exemplisque Plurimis Explicata. Tomus secundus Parisiis at Argentorati, apud Amand Konig, Bibliopolam, 1 - 280 pp.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Kolbe, H. J. (1914) Lieferung 3: Scarabaeiden. In: Shubotz, H. (Ed.), Wissenschaftliches Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907 unter Fuhrung Adolf Friedrichs, Herzog zu Mecklenburg. Zoologie III, 5. Klinkhardt & Biermann, Leipzig, Germany, pp. 261 - 416.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Arrow, G. J. (1909) On some new species of Coleoptera from Rhodesia and adjacent territories. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London 8 (3): 517 - 523.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Balthasar, V. (1968) Neue Scarabaeiden-Arten, 129. Beitrag zur kenntnis der Scarabaeoidea (Coleoptera). Beitrage zur Entomologie, 18: 953 - 958.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Paschalidis, K. M. (1974 b) The identity of two species in the genus Sisyphus (Coleoptera: Scarbaeidae). Journal of Entomology of Southern Africa. 37 (2): 299 - 303.","Endrodi, S. (1983) Sisyphus bornemisszanus sp. n. from South Africa (Coleoptera, Scarabaeidae): Annales Historico-Naturales Musei Nationali S Hungarici (74): 207 - 210.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Schafer, P. & Fischer, E. (2001) Zur Scarabaeidenfauna Rwandas, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini, Mitteilungen des internationalen entomologischen Vereins supplement 9: 1 - 93.","Montreuil, O. (2014) Deux nouveaux Sisyphus Latreille de Tanzanie (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 119 (2): 177 - 180.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Montreuil, O. (2016) Nouveaux Sisyphus Latreille, 1807, du groupe seminulum: le complexe arboreus (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 121 (2): 167 - 174.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Montreuil, O. (2017) Nouveaux Sisyphus Latreille africains du groupe seminulum: les complexes seminulum, desaegeri et ocellatus (Coleoptera, Scarabaeidae, Sisyphini), Catharsius La Revue, 14: 1 - 22."]}

Sisyphus neobornemisszanus Daniel & Davis, 2016 (Fig. 2, 3B). Daniel et. al. 2016: 67, 71, 73 Type locality: Zululand, St. Lucia (South Africa). Size: Male: length: 9.7– 7.6 mm, width: 5.6– 4.6 mm. Female: length: 10.0–8.0 mm, width 5.62– 4.4 mm. Diagnosis: S. neobornemisszanus resembles S. muricatus. However, it may be distinguished by the following combination of features: inconspicuous punctation on the pronotum; interstriae 1, 3, 5, and 7 with rows of black setae arranged in tufts; interstriae 2, 4, and 6 with sparsely arranged single setae. Examined type material (♂ TMSA) Holotype: SOUTH AFRICA, Zululand, St. Lucia 28°13’12”S 32°15’00”E, 7.xii.1975, leg: Endrödy-Younga. Paratypes: (1♀ TMSA) SOUTH AFRICA, Zululand, Mission Rock, 28°15’36”S 32°17’24”E, 7.xii.1975, legs: Endrödy-Younga. (2♂ TMSA) SOUTH AFRICA, Zululand, St. Lucia 28°13’12”S 32°15’00”E, 28.viii.1971. leg. GF. Bornemissza. (2♂ TMSA) SOUTH AFRICA, Zululand, St. Lucia, Estuary. 28.viii.1971, leg: GF. Bornemissza. (15♂, 13♀ TMSA) SOUTH AFRICA, Zululand, Sodwana Bay, 5 km, 27°21’0”S 32°23’24”E, 2.xi.1992, leg. Endrödy-Younga. (1♀ UPSA) SOUTH AFRICA, KZN. Richard’s Bay, 28°37’39”S 32°17’24”E, 27.i.2000, leg: Davis & Delport. (2♀, 6♂ UPSA) SOUTH AFRICA, KZN, Richard’s Bay, 28°43’28”S 32°10’52”E, 27.i.2000, leg: Davis & Delport. (5♀, 2♂ UPSA) SOUTH AFRICA, KZN. Sileza Nature Reserve 27°06’S 32°36’E, 03.iv.1996, leg: Van Rensburg. (1♀ UPSA) SOUTH AFRICA, KZN, Sodwana, 27°32’S 32°41’E, 07.iv.1989, leg: Mansfield. (2♀, 5♂ UPSA) MOZAMBIQUE, Maputo, Elephant Maputo Reserve, 26°22’30”S 33°47’50”E, 14–16.i.2003, leg: Scholtz & Holter. (2♀ UPSA) MOZAM- BIQUE, Maputo, Elephant Maputo Reserve, 26°39’S 32°43’E, 10–16.xi.2007, pitfall trap, leg: Strümpher & Deschodt. (2♂ TMSA) MOZAMBIQUE, Inhambane, Inharrime, 24°28’37”S 35°01’49”E, 1974, leg: Moor. Examined non-type material: See Supplementary information. Distribution: S. neobornemisszanus is an endemic, east coastal, dune forest species (Davis et al. 2002b). It has been recorded in the Maputaland Coastal Forest Mosaic (Olson et al. 2001) from KwaZulu Natal to southern Mozambique (Endrödi, 1983, Davis et al. 2002; Daniel et al. 2016) (Fig. 4).

Neosisyphus fortuitus (P��ringuey, 1901) (Figs. 20, 21A) Sisyphus fortuitus P��ringuey, 1901 P��ringuey 1901: 103; Arrow 1927: 464; Gillet 1911: 23; Haaf 1955: 371; Ferreira 1972: 819; Montreuil 2015b: 9. Type locality: Durban, Natal (South Africa) Size: Male: length: 11.1���9.0 mm; width: 5.7���5.0 mm. Female: length: 11.5���9.0 mm; width: 6.0��� 4.4 mm. Diagnosis: Neosisyphus fortuitus is distinguished from other members of the spinipes species group in southern Africa by lacking a spine or projection on the mid-posterior edge of the metafemur. In addition, the projecting metatrochanter is almost �� of the length of the metafemur in major males. The parameres are distinctly different to those of the other two species within the group (Fig. 21A). Examined type material Lectotype: (1♂ SAM) /Natal, Durban/ (red label) / S. fortuitus P��r, Lectotype, det. E. Haaf, 1954/. Examined non-type material: See Supplementary information. Distribution: N. fortuitus occurs in shaded vegetation in southern (South Africa, Mozambique, Zimbabwe) and East Africa (Kenya) (Fig. 22)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 47-48, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36."]}

Sisyphus gazanus Arrow, 1909 (Figs. 11, 12C). Arrow 1909: 518; Gillet 1911: 23; Arrow 1927: 461; Haaf 1955: 361; Ferreira 1972: 799; Sch��fer & Fischer 1992: 129. Type locality: Chirinda, Mashonaland (Zimbabwe) Size: Male: length: 5��� 4.5 mm; width: 3.1��� 2.5 mm; Female: length: 5��� 4.4 mm, width 3.1��� 2.4 mm. Diagnosis: S. gazanus is relatively close in appearance to S. inconspicuus sp. n. However, it is easily distinguished by a combination of features as follows: S. gazanus has a somewhat shallow depression dorsally on the genae; the dorsal surface of the head (vertex and frons) is entirely covered by distinct ocellate punctures (Fig. 11J); the pronotal disc shows a patchwork of bald areas and sparse short setae. In addition, the parameres of both species are distinctly different (Figs. 12 C���D). Examined type material Lectotype: (♂ NHML): ZIMBABWE, Mashonaland, Chirinda, 05.x.1908. Leg: Marshall (handwritten Sisyphus gazanus Arrow 1909), lectotype det: Bacchus, 1976 / violet label /. Paralectotype: (♂ NHML): ZIMBABWE, Mashonaland, Mt. Chirinda, 01.xii.1908, leg: Marshall (handwritten Sisyphus gazanus Arrow 1909), lectotype det: Bacchus, 1976; /blue label/. Examined non-type material: See Supplementary information. Distribution: Sisyphus gazanus is associated with cooler conditions in highland forest in Zimbabwe and Malawi. It was described from Mount Chirinda (Zimbabwe) at the southern end of the Inyanga Highlands and Chimanimani Mountains. This range may have acted as a corridor into Malawi where the species has also been recorded from Mount Mulanje and Mount Gomoloti (Fig. 13). So far, it has not been recorded under cooler, shaded conditions on the intervening Muchinga Mountains (Manica, Mozambique) or Mount Binga (19.78��S 33.30��E)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 30-31, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Arrow, G. J. (1909) On some new species of Coleoptera from Rhodesia and adjacent territories. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London 8 (3): 517 - 523.","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137."]}

Neosisyphus calcaratus (Klug, 1855) (Figs. 17, 18B) Montreuil 2015b: 15 Sisyphus calcaratus Klug, 1855 Klug 1855: 651; Harold 1869: 984; Arrow 1927: 464; Boucomont 1933: 3; Haaf 1955: 377, Ferreira 1972: 829. Sisyphus rubripes Boheman, 1857 Boheman 1857:193 Size: Male: length: 7.1��� 6.6 mm; width: 4.5���3.0 mm. Female: length: 7.0��� 6.5 mm; width: 4.0��� 3.8 mm. Type locality: Sena (Sofala, Mozambique). Diagnosis: N. calcaratus is similar to N. barbarossa. Males of both species bear a sharp pointed spine on the mid-posterior edge of the metafemur, which is absent in the female. However, the elytra of N. calcaratus bear black setae that are bent over at the tip whilst the elytra of N. barbarossa are covered by brown setae. Furthermore, compared to N. barbarossa, major males of N. calcaratus are smaller but the projection of the metatrochanter is twice as long. Examined type material Sisyphus rubripes Boheman, 1857 : (1♀ NHRS): / Caffraria /, / Typus /; / rubripes Bohem /; / Sisyphus rubripes Boh =armatus Gory, det. E. Haaf, 1954/; / Sisyphus calcarutus Klug, det. KM, Paschalidis 1975/; /NHRS_JLKB: 000025221. Examined non-type material: See Supplementary information. Distribution: N. calcaratus has been recorded in dry lowland savanna and mixed woodland in the lowveld of South Africa, Botswana, Zimbabwe and Swaziland. It has also been observed on the coastal plain and interior of southern central Mozambique (Fig. 19)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 44, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Klug, J. C. F. (1855) Diagnosen neuer Coleoptera aus Mossambique. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 643 - 660.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1."]}

Sisyphus goryi Harold 1859 (Figs. 5, 6D). Harold 1859:224; Harold 1869:984; P��ringuey 1902:106���7; Kolbe 1914: 317; Arrow 1927:460; Haaf 1955:362���3; Ferreira 1972:797; Sch��fer & Fischer 2001:53���4; Sch��fer & Fischer 1992:121���2. Sisyphus hirtus Wiedeman, 1823 Wiedeman 1823:21; Gory 1833:14; Harold 1869:984; Arrow 1909:518���9; 1927:460; Haaf 1995:362���3; Sch��fer & Fischer 2001:53���4; Sch��fer & Fischer 1992:121���2. S. pygmaeus Dejean, 1837 Dejean 1837: 151; Klug, 1862:219; Harold 1869:984; Haaf 1955:362���3. Type locality: Senegal Size: Male: length: 7���4.0 mm, width: 3.4��� 2.2 mm. Female: Length: 7.0��� 3.9 mm, width: 3.1��� 2.4 mm. Diagnosis: S. goryi is similar to Sisyphus bicuariensis sp. n. However, it differs by the uniform brown colour of the setae compared to the bicoloured setae in S. bicuariensis sp. n. The male genitalia of both species differ (Figs. 6 A, D). Furthermore, it should be noted that larger S. goryi specimens resemble those of S. caffer. However, S. goryi bears well-developed tufts of setae, which emanate from below the abdominal epipleurae (Fig. 5H) (>10 single setae), which differ to those in S. caffer where the tufts consist of few single setae (Examined non-type material: See Supplementary information. Morphological variation Head. Antennal club varies in colour between brown or black. Clypeus. In some specimens, especially old ones, the lateral edge of the clypeus is worn completely smooth; the third indentation is absent; and the medial surface bears a metallic sheen. Sternites. Some populations are completely hairless on the latero-posterior edge of the mesosternum. Pygidium. Setae are evenly distributed in some individuals whereas in others they are arranged in a trident pattern. Populations from highlands and areas of shaded vegetation, such as Inyanga (Zimbabwe), Manyara (Tanzania) and Meru (Kenya) are distinctly larger in size and may, in some cases, resemble S. caffer. Distribution: S. goryi is widespread in the Afrotropical region and is associated with a great range of habitats from shaded to open vegetation. Moreover, it has been recorded at a range of different altitudes (Fig. 7). The geographical distribution and morphological variation suggest a species complex. However, as we could not access the type material of S. goryi, the putative species complex currently remains undetermined pending further study. Remarks: The original name of Sisyphus hirtus Gory, 1833, is preoccupied by Sisyphus hirtus Wiedmann, 1825 (Indian species). Gory���s junior homonym was, therefore, renamed as Sisyphus goryi Harold, 1859. Subsequently, P��ringuey (1901) synonymized S. caffer Boheman, 1857, and S. sordidus Boheman, 1857, with S. goryi, whereas, Haaf (1955) and Ferreira (1972) synonymized S. goryi with S. crispatus Gory, 1833. Based on morphological and biogeographical evidence we have, here, revalidated S. caffer, S. sordidus and S. goryi following the unpublished revision of Paschalidis (1974)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 23, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Harold, E. von (1859) Beitrage zur Kenntniss einiger coprophagen Lamellicornien. Deutsche Entomologische Zeitschrift, 3 (2 - 3), 193 - 224.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Kolbe, H. J. (1914) Lieferung 3: Scarabaeiden. In: Shubotz, H. (Ed.), Wissenschaftliches Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907 unter Fuhrung Adolf Friedrichs, Herzog zu Mecklenburg. Zoologie III, 5. Klinkhardt & Biermann, Leipzig, Germany, pp. 261 - 416.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Schafer, P. & Fischer, E. (2001) Zur Scarabaeidenfauna Rwandas, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini, Mitteilungen des internationalen entomologischen Vereins supplement 9: 1 - 93.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Wiedemann, C. K. W. (1823) Zweihundert neue Kafer von Java, Bengalen und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin, 2 (1): 3 - 133.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Arrow, G. J. (1909) On some new species of Coleoptera from Rhodesia and adjacent territories. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London 8 (3): 517 - 523.","Dejean, P. F. (1837) Catalogue des Coleopteres de la collection de M. le Comte Dejean. Mequignon-Marvis.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp."]}

Sisyphus fasciculatus Boheman 1857 (Figs. 2, 3C) Boheman 1857: 192; Harold 1869: 984; P��ringuey 1902: 101���106, Arrow 1927: 465; Haaf 1955: 376; Ferreira 1972: 799; Davis et al, 2008: 154; Montreuil 2015c: 3, 11. Type locality: Caffraria Size: Male: length: 9.4��� 7 mm, width: 5.7��� 3.5 mm. Female: length: 9.0���7.0 mm, width 5.6��� 3.5 mm. Diagnosis: S. fasciculatus is similar to S. muricatus and S. neobornemisszanus, but differs from these two species by bearing tufts of dense setae on the pronotum, which alternate with bare patches of short setae. Examined type material Holotype (photograph): (♀ NHRS): / Caffraria /, / J. Wahlb /, / type /, brown label, handwritten / fasciculatus Bohem /; red label / Typus: Sisyphus fasciculatus, det. Dr. E. Haaf, 1954/, /311:72/, /247:73/; /9415 E92 +/, brown label, handwritten / fasciculatus Boh /, /NHRS-SRAH 000000228/. Examined non-type material: See Supplementary information. Distribution: S. fasciculatus occurs in South Africa and Swaziland where it is a shade specialist (Davis et al. 1999). It is associated with riverine and lower hillside forest to dense partially shaded woodland thickets, particularly, on the eastern escarpment of Limpopo, Mpumalanga, west KwaZulu Natal and Swaziland. However, it also occurs in the warmer inland parts of the Maputaland Coastal Forest Mosaic ecoregion of Olson et al. (2001) southwards to the northernmost extent of the KwaZulu-Pondoland Coastal Forest Mosaic (Fig. 4). Conservation status: Although currently assessed as Least Concern (LC), S. fasciculatus is a potentially threatened species, due to the loss of forest and shaded woodland across its fairly restricted range (Davis 2013a). However, the species has been recorded in reserves where shaded vegetation is protected, such as in Hluhluwe, Umfolozi and Ithala Game Reserves (KwaZulu Natal) as well as Mlawula Nature Reserve (Swaziland). Because of this protection, it can be locally abundant in both coastal reserves and along the lower part of the northern escarpment of the Drakensberg., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 13-15, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Davis, A. L. V., Frolov, A. V., & Scholtz, C. H. (2008) The African dung beetle genera. Protea Book House. 148 - 155 pp.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Davis, A. L. V., Scholtz, C. H., & Chown, S. L. (1999) Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa. Journal of Biogeography, 26 (5): 1039 - 1055.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V., Underwood, E. C., & Loucks, C. J. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth: A new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience, 51 (11): 933 - 938.","Davis, A. L. V. (2013 a) Sisyphus fasciculatus. The IUCN Red List of Threatened Species 2013: e. T 137250 A 522146. http: // dx. doi. org / 10.2305 / IUCN. UK. 20132. RLTS. T 137250 A 522146. en. (accessed 07 September 2016)."]}

Neosisyphus rubrus (Paschalidis, 1974) (Figs. 29, 30B) Montreuil 2015b: 20, 28 Sisyphus rubrus Paschalidis 1974 Paschalidis, 1974b: 299 Type locality: Castle Gorge (South Africa) Size: Male: length: 9.0���6.0 mm; width: 4.1���3.0 mm. Female: length: 8.7���6.0 mm; width: 4.0���3.0 mm. Diagnosis: N. rubrus closely resembles N. macrorubrus (Paschalidis 1974). However, the species can be distinguished by the lateral protrusions at the apices of the parameres in N. rubrus (Fig. 30B), which are lacking in N. macrorubrus. Examined type material Holotype: (♂ SANC) / SOUTH AFRICA, TVL, Castle Gorge (42 mi W Pretoria), 15.v.1971, leg: Bornemissza & Insley /; /283a/, /ex coll. CSIRO, Div. Entomology; S. African Station /; red label / Holotype ♂, Sisyphus rubrus spec. nov, det. KM Paschalidis, 1974 /, white label / SANC TYPH 00723/. Paratypes: (5 ♂, 1 ♀ SAM) / RHODESIA, Sebakwe /, / Sisyphus rubripes P��r, det. Dr. Haaf, 1954/. (15♂, 15 ♀ SANC) SOUTH AFRICA, TVL, Castle Gorge (42 mi W Pretoria), 15.v.1971, leg: Bornemissza & Insley. (2♂, 1 ♀ SANC) SOUTH AFRICA, OFS, Vredefort (20 mi S), 03.xi.1971, leg: Bornemissza & Kirk. (3♂, 5 ♀ SANC) SOUTH AFRICA, CP, Mafeking, 22.v.1972, leg: Davis. (2♂, 1 ♀ SANC) SOUTH AFRICA, SE. TVL, Bergen, 1.v.1972, leg: Kirk. (1♂, 5 ♀ SANC) SOUTH AFRI- CA, N. TVL, Louis Trichardt (11 mi N), 14.xii.1971, leg: Olsen. (2♂, 1 ♀ SANC) SOUTH AFRICA, CP, Aliwal North (22 mi S), 4.iii.1971, leg: Aschenborn, (50♂, 47 ♀ SANC) SOUTH AFRICA, E. TVL, Bosbokrand (5 mi N), 6.ii.1971, leg: Olsen. (2♂, 3♀ SANC) SOUTH AFRICA, Transkei, Umtata (10 mi N), 19.v.1971, leg: Aschenborn. (1♂, 2♀ SANC) MOZAMBIQUE, Vila Pery, (5 mi E), 10.iv.1971, leg: Bornemissza & Aschenborn. (1♂ SANC) SOUTH AFRICA, Vryburg, N. CP, (15 mi E, Leeuwrand Farm), 7.x.1972, leg: Kluge. Sisyphus rubripes P��ringuey, 1901 : Lectotype: (♂ SAM) /Natal, Durban/; / Sisyphus rubripes P��r, det. Dr. Haaf, 1954/; red label / Sisyphus rubripes P��r, Lectotype, det: E. Haaf, 1954/. Examined non-type material: See Supplementary information. Distribution: N. rubrus is distributed in upland and coastal, moist grassland and savanna (Davis et al. 1999) in southeast Africa (Fig. 31). Conservation status: N. macrorubrus and N. rubrus are listed as Least Concern species on the IUCN Red Data List (Davis 2013 e, f). Remarks: It has been suggested that on grounds of gender, the spelling of N. rubrus and N. macrorubrus should be changed to N. ruber and N. macroruber respectively (Davis et. al. 1999; Davis et al. 2008, Davis 2013 e, f). It should be noted that; ��� ruber ��� is a masculine classical adjective, which has been used in a rare case as an alternative form of ��� rubrus ��� (Brown, 1954). However, the original description of each species does not specify the etymological meaning of the specific name ��� rubrus ��� (Paschalidis, 1974 b). According to ICZN (article: 31.2.2) ���Where the author of a species-group name did not indicate whether he or she regarded it as a noun or as an adjective, and where it may be regarded as either and the evidence of usage is not decisive, it is to be treated as a noun in apposition to the name of its genus���. Therefore, rubrus and macrorubrus must be treated as indeclinable names., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 56-57, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Paschalidis, K. M. (1974 b) The identity of two species in the genus Sisyphus (Coleoptera: Scarbaeidae). Journal of Entomology of Southern Africa. 37 (2): 299 - 303.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Davis, A. L. V., Scholtz, C. H., & Chown, S. L. (1999) Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa. Journal of Biogeography, 26 (5): 1039 - 1055.","Davis, A. L. V. (2013 e) Neosisyphus rubrus. The IUCN Red List of Threatened Species 2013: e. T 137480 A 525709. http: // dx. doi. org / 10.2305 / IUCN. UK. 20132. RLTS. T 137480 A 525709. en. (accessed 07 September 2016).","Davis, A. L. V., Frolov, A. V., & Scholtz, C. H. (2008) The African dung beetle genera. Protea Book House. 148 - 155 pp."]}

Identification key for males of southern African sisyphine species 1. Lateral ridge on the pronotal disc extends from the posterior margin without reaching the lateral margin (Fig. 1F); strong setae absent on the external margin of the first meso- and metatarsal segments (Fig. 1D)........................ Neosisyphus (2) ��� Lateral prothoracic ridge between pronotal disc and prothoracic episternum complete (Fig. 1E); first tarsal segments of the meso- and meta-thoracic legs with a row of dense setae in a straight line on the external edge (Fig. 1C)........................................................................................................................... Sisyphus (13) 2. Mesotibia strongly modified by indentations and protrusions; a large spine distally on the posterior margin of the mesofemur. Completely black, pygidium simple and lacking a pair of distinct tubercles. Eastern coastal forests of South Africa and South Mozambique................................................................................................................(tibialis species-group; Neosisyphus mirabilis (Arrow,1927) (Figs. 23, 24A) ��� Mesotibia simple, without any spine projected distally on the posterior margin of the mesofemur.................................................................................................(3) 3. Femur and tibia of mesolegs without cuticular projections on the posterior margin............................................. barbarossa species-group (Figs. 17 A���D) (4) ��� Femur and tibia of mesolegs with cuticular projections on the posterior margin...(7) 4. Metafemur with sharp pointed spine on the mid-posterior margin (Figs. 17 A���B)........................................................................................................................(5) ��� Metafemur with angled-protuberance on the mid-posterior margin (Figs. 17 C���D)........................................................................................................................(6) 5. Projecting metatrochanter longer than �� of the metafemur length in major males and �� of the metafemur length in minor males. In major males the metatrochant- er is slightly inwardly-curved at the tip and straight in minor males; setae on the elytra black, recurved at the apex. Lowland dry savannas in southern Africa.................................................. Neosisyphus calcaratus (Klug, 1855) (Figs. 17, 18B) ��� Projecting metatrochanter shorter in minor males ( Neosisyphus barbarossa (Wiedemann, 1823) (Figs. 17, 18A) 6. Projecting metatrochanter relatively short; four times shorter than total length of metafemur. Lowland and upland moist vegetation from northeast to southern Africa..................................... Neosisyphus setiger (Roth, 1851) (Figs. 17, 18D) ��� Metatrochanter slighty inwardly-projected at the apex, metatrochanter projection is �� the length of the metafemur. Upland woodland in southern Angola............................................. Neosisyphus tembyi Daniel & Davis sp. n (Figs. 17, 18C) 7. Posterior margin of the mesoleg with spine or projection distally on mesofemur and basally on mesotibia; first quarter of mesotibia from femoro-tibial joint abruptly narrowed................................ spinipes species-group (Figs. 20 A���C) (8) ��� Mesotibial spine projecting posteriorly and gradually narrowing distally........(10) 8. Spine present on the mid-posterior margin of the metafemur; metatrochanter relatively short, �� of the metafemur length........................................................... (9) ��� Spine absent on posterior margin of the metafemur; metatrochanter is �� the length of the metafemur in major males. Shaded vegetation and thickets from east to southern Africa........ Neosisyphus fortuitus (P��ringuey, 1901) (Figs. 20, 21A) 9. Distinct triangular shaped projection present on the basal posterior margin of the mesotibia. Parameres not curved at the apex (Fig. 21C). Moist upland and coastal grassland from east to southern Africa............................................................................................................. Neosisyphus spinipes (Thunberg, 1818) (Fig. 20C) ��� Lacking a triangular protuberance on the basal posterior margin of the mesotibia in minor males. There is a slight sinuosity in major males. Parameres curved downward apically (Fig. 21B). Lowland dry savanna from east to southern Africa................................................. Neosisyphus infuscatus (Klug, 1855) (Fig. 20B) 10. Basal posterior spine on mesotibia projecting perpendicularly; body uniformly brown; size of beetles 6.0��� 10 mm......... rubrus species-group (Figs. 29 A���B) (11) ��� Basal posterior spine on mesotibia not projecting perpendicularly to the ventral apical margin of the femur; colour is not uniform overall, size of beetles 11.0��� 13.5 mm...................................... quadricollis species-group (Figs. 25 A���B) (12) 11. Parameres with protrusion apical-laterally (Fig. 30B). Upland and coastal moist grassland and savanna in southern Africa......................................................................................................... Neosisyphus rubrus (Paschalidis, 1974) (Fig. 29B) ��� Parameres simple (Fig. 30A). Cool, dry savanna, karoo and grassland in South Africa (Northern Cape and Free State) and Namibia (Etosha Pan, Okaukuejo)..................................... Neosisyphus macrorubrus (Paschalidis, 1974) (Fig. 29A) 12. Lateral margin of elytra emarginate; concavity of margin distinctly deep in the middle; short projecting metatrochanter in males (1/3 of metafemur length); metafemur lacking spines. Deep sands and shrubland of southwest coast, and karoo in the Eastern Cape (South Africa)................................................................................................... Neosisyphus quadricollis (Gory, 1833) (Figs. 26, 27A) ��� Margin of elytra with regular shape; a protuberance distally on the posterior margin of the metafemur, the metatrochanter exceeds �� the length of the metafemur in males. Moist highland grassland in South Africa............................................................................................. Neosisyphus kuehni (Haaf, 1955) (Figs. 26, 27B) 13. Setae on the elytral interstriae arranged in tufts............................................................................................................. muricatus species-group (Figs. 2 A���F) (14) ��� Setae on the elytral interstriae evenly distributed, not grouped in tufts.......... (20) 14. Setae arranged in tufts on the pronotum, which alternate with depressed bare patches.......................................................................................................... (15) ��� Setae on the pronotum not arranged in tufts................................................. (17) 15. Concave and upwardly curved margin between the anterior medial teeth of the clypeus; macropterous (Fig. 2I). Woodland thickets and forest in north South Africa and Swaziland.......... Sisyphus fasciculatus Boheman, 1857 (Figs. 2, 3C) ��� Anterior margin between medial clypeal teeth not upcurved; hind wings reduced or completely brachypterous (Figs. 2 H���J).....................................................(16) 16. Anterior margin between medial clypeal teeth concave; pronotum wider anteriorly than posteriorly. Parameres (Fig. 3F). Only known from Swaziland........................................................................ Sisyphus swazi Daniel & Davis sp. n (Fig. 2F) ��� Straight edge between medial teeth of clypeus; width of pronotum uniform from anterior to posterior margins. Parameres (Fig. 3G). Popa Falls forest (Namibia), Central Mozambique and East Africa.................................................................................................................... Sisyphus alveatus Boucomont, 1935 (Fig. 2G) 17. Very strong lateral emargination and distinct shiny punctation on pronotum. Cooler southern forest and montane grassland in South Africa............................................................................ Sisyphus muricatus (Olivier, 1789) (Figs. 2, 3A) ��� No distinct lateral emargination on the pronotum.........................................(18) 18. Punctation on the pronotum unclear; interstriae 1, 3, 5, and 7 with rows of black setae arranged in tufts, interstriae 2, 4, and 6 with sparsely arranged single setae. Coastal sand forest of northeast South Africa and southeast Mozambique................................ Sisyphus neobornemisszanus Daniel & Davis, 2016 (Figs. 2, 3B) ��� Punctation on the pronotum conspicuous; setae on pronotal disc arise from the centre of ocellate punctures (Fig. 5H)............................................................(19) 19. Proximal elytral setae evenly arranged; mid-basal elytral setae arranged in tufts of scattered individual setae. Pronotal disc with three prominent round depressions. Parameres (Fig. 3E). Primarily upland and highland grassland in Gauteng, Kwa- Zulu Natal, Limpopo, Mpumalanga and Northwest Province..................................................................................... Sisyphus manni Montreuil, 2015 (Fig. 2E) ��� Elytra entirely covered in tufts of setae, each tuft dense and compact; depressions on the pronotal disc absent; body larger than S. manni and oval. Parameres (Fig. 3D). Unshaded vegetation in the Eastern Cape......................................................................................... Sisyphus perissinottoi Montreuil, 2015 (Fig. 2D) 20. Well-developed tufts of epipleural setae (Fig. 5H); edge between medial teeth almost straight; setae on the pronotum evenly distributed........................................................................................................ goryi species-group (Figs. 5 A���E) (22) ��� Weak tufts of epipleural setae; margin between medial teeth V-shaped)................................................................................................. umbraphilus species-group 21. Pronotal disc with setae interspersed with bald patches. Dense woodland and thicket in KwaZulu Natal, Gauteng and North West Provinces (South Africa)......................................... Sisyphus umbraphilus Daniel & Davis, 2016 (Fig. 8A) ��� Pronotal disc without clear setae interspersed with bald patches.................... (26) 22. Setae on the pronotum inserted between or on the posterior margin of ocellate punctures (Fig. 5F)........................................................................................ (23) ��� Setae on the pronotum inserted centrally on ocellate punctures.................................................................................................................................. (Fig. 5G) (25) 23. Fine setae on elytra arranged in single rows, epipleural tufts containing few setae ( Sisyphus caffer Boheman, 1857 (Figs. 5, 6B) ��� Much finer setae on elytra, distributed in single rows; epipleural tufts containing many setae (>10); elongate body shape..........................................................(24) 24. Setae on the pronotum and elytra of uniform colour and regularly arranged. Parameres (Fig. 6D). Widespread woodland species in sub-Saharan Africa................................................................................ Sisyphus goryi Harold, 1859 (Fig. 5D) ��� Setae on pronotum and elytra bicoloured (black and gray). Parameres (Fig. 6A). Open woodland in southern Angola........................................................................................................... Sisyphus bicuariensis Daniel & Davis, sp. n (Fig. 5A) 25. Elytral setae thick, arranged in non-single rows and forming bunches; body size: 6.6��� 4.4 mm. Open lowland vegetation in South Africa, Mozambique, and Zimbabwe....................................... Sisyphus sordidus Boheman, 1857 (Figs. 5, 6E) ��� Elytral setae fine, regularly arranged on interstriae; body size: 10���8.0 mm. East African miombo woodland; known only from Quirimbas National Park (Northern Mozambique)..................... Sisyphus genierorum Montreuil, 2015 (Figs. 5, 6C) 26. Setae on outer margin of elytra not arranged in tufts; margin between medial teeth concave or straight; very large ocellate punctures on dorsal surface of pronotal disc...................................................... costatus species-group (Figs. 11 A���E) (27) ��� Setae on outer margin of elytra not arranged in tufts; margin between medial teeth narrowed (almost V-shaped); very fine ocellate punctures on dorsal surface of pronotal disc......................................... seminulum species-group (Figs. 6 A���C) (30) 27. Dorsal surface of the clypeus and frons with inconspicuous ocellate punctation, but more visible on the vertex (Figs. 11 G���I).................................................. (28) ��� Dorsal surface of the head (clypeus, frons and vertex) with distinct ocellate punctation (Figure11J).......................................................................................... (30) 28. Vertex with dense ocellate punctation, margin between medial teeth straight; genae convex anteriorly (Fig. 11I); elytral setae fine and uniformly arranged. Moist savanna or dense woodland and riverine vegetation in the interior of South Africa. Shaded vegetation on coastline from Sofala Bay (Mozambique) to Eastern Cape (South Africa)................ Sisyphus inconspicuus Daniel & Davis, sp. n (Fig. 5B) ��� Vertex with scattered ocellate punctation (Fig. 11 G���H), elytral setae thicker, alternating on the interstriae between rows of well-developed and less developed setae (Fig. 11F)......................................................................................................(29) 29. Margin between medial teeth distinctly concave and upcurved; lacking carinae on the frons; genae virtually straight laterally (Fig. 11G). Parameres (Fig. 12B). Highland grassland in South Africa....................................................................................................................... Sisyphus costatus (Thunberg, 1818) (Fig. 11B) ��� Edge between medial teeth straight; well-defined clypeo-frontal carina; genal margin virtually convex (Fig. 11H). Parameres (Fig. 12E). Shaded vegetation from Western and Eastern Cape Provinces, South Africa.......................................................................................... Sisyphus australis Daniel & Davis sp. n (Fig. 11E) 30. Small-bodied; genae with shallow depression dorsally; pronotal disc showing distinct pattern of bare patches amongst sparse short setae; metafemur simple. Highland forest under cooler conditions in Zimbabwe and Malawi........................................................................ Sisyphus gazanus Arrow, 1907 (Figs. 11, 12C) ��� Large-bodied; genal depression absent; setae arranged regularly on pronotal disc; anvil- shaped projection on mid-posterior margin of metafemur. Lowland shaded Savanna from east to southern Africa..................................................................................................... Sisyphus impressipennis Lansberge, 1886 (Figs. 11, 12A) 31. Relatively large-bodied; pronotal disc with conspicuous metallic sheen; open woodland in north of Namibia and south of Angola............................................................................................ Sisyphus splendidus Montreuil, 2015 (Fig. 14A) ��� Relatively small to medium-bodied................................................................ (32) 32. Very small-sized body (3.0��� 3.8 mm); pronotal disc dull bearing some depressions. Shaded vegetation in eastern and southern Africa....................................................................................... Sisyphus nanniscus P��ringuey, 1901 (Figs. 14C, 15B) ��� Medium-sized body (3.9���5.0 mm)................................................................ (33) 33. Pronotal disc bears sparse, long and fine yellow setae, separated by linear bare patches (Fig. 14E); parameres notched dorsally (Fig. 15A). Dense coastal woodlands and forest from northeast South Africa to southern Mozambique........................................................ Sisyphus oralensis Daniel & Davis, 2016 (Fig. 14B) ��� Pronotum bears thick and short golden setae; directional projection of the setae well-defined (Fig. 14F); parameres without excavation dorsally, attenuating sharply at the apex (Fig. 15C). Forest and dense woodland species in central Mozambique................... Sisyphus auricomus Daniel & Davis sp. n (Fig. 14D), Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 5-9, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Klug, J. C. F. (1855) Diagnosen neuer Coleoptera aus Mossambique. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 643 - 660.","Wiedemann, C. K. W. (1823) Zweihundert neue Kafer von Java, Bengalen und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin, 2 (1): 3 - 133.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Thunberg, C. P. (1818) Coleoptera Capensia, Antennis Lamellatis, sive, clava fissili instructa, descripta, memoires de l'Acad. T. VI.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Boucomont, A. (1935) Coleoptera. VI. Scarabaeidae Scarabaeini. In: Jeannel, R. (Ed.), Mission Scientifique de l'Omo. Tome 2 Zoologie, Fascicule 16. Paul Chevalier & Fils, Paris, France, pp. 279 - 290.","Olivier, M. (1789) Entomologie, Histoire Naturelle des Insectes, Avec leurs caracteres generiques et specifiques, leur description, leur synonymie, et leur figure enluminee. Coleopteres, Tome premier, Paris, 1 - 455 pp.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Harold, E. von (1859) Beitrage zur Kenntniss einiger coprophagen Lamellicornien. Deutsche Entomologische Zeitschrift, 3 (2 - 3), 193 - 224.","Lansberge, J. W. (1886) Scarabaeides, Buprestides et Cerambycides de l'afrique occidentale. Notes from Leyden Museum, 8 (2): 69 - 120."]}

Neosisyphus macrorubrus (Paschalidis, 1974) (Figs. 29, 30A) Montreuil 2015b: 20, 28 Sisyphus macrorubrus Paschalidis 1974 Paschalidis, 1974b: 300 Size: Males: length: 10.2��� 7.5 mm; width: 4.5���3.0 mm. Females: length: 10.5���7.0 mm; width: 4.5���3.0 mm. Type locality: Vanstandensrus, OFS (Free State, South Africa) Diagnosis: N. macrorubrus is very close in appearance to N. rubrus. However, it shows simple parameres (Fig. 30A) that lack the lateral protrusions, which typify N. rubrus. It also shows a different geographical distribution pattern and minimal overlap with N. rubrus. Examined type material Holotype: (♂ SANC) / SOUTH AFRICA, OFS, Vanstadensrus, 10.xii.1971, leg: Kirk /, /457/1/, /ex coll. CSIRO, Div. Entomology; S. African Station /; red label / Holotype ♂, Sisyphus marcorubrus spec. nov, det. KM Paschalidis, 1974 /, white label /SANC TYPH 00724/ Paratypes: (4♂, 4♀ SANC; 1♂, 1♀ SAM) SOUTH AFRICA, OFS, Ladybrand (Maseru Rd), 27.xi.1971, leg: Bornemissza & Aschenborn. (3♂, 3♀ SANC; 1♂, 1♀ SAM) SOUTH AFRICA, OFS, Zastron (21 mi N), 12.v.1972, leg: Olsen. (2♂, 1♀ SANC; 1♂, 1♀ SAM) SOUTH AFRICA, OFS, Caledon River (3 mi W Maseru), 2.iii.1971, leg: Aschenborn. (9♂, 6♀ SANC) SOUTH AFRICA, CP, Aliwal North (North), 3.iii.1971, leg: Aschenborn. (3♂, 3♀ SANC) SOUTH AFRICA, OFS, Wepener (22 mi N), 11.v.1971, leg: Aschenborn. (4♂, 3♀ SANC) SOUTH AFRICA, CP, Kimberley (25 mi S), 14.iii.1971, leg: Bornemissza & Kirk. (2♂, 1♀ SANC) SOUTH AFRICA, OFS, Rouxville, 01.xi.1971, leg: Bornemissza & Kirk. (1♂, 2♀ SANC) SOUTH AFRICA, OFS, Bloemfontein (11 mi N), 03.xi.1971, leg: Bornemissza & Kirk. (1♂, 3♀ SANC) SW.AFRICA (NAMIBIA), Outjo (15 kms N), 10.iv.1974, leg: Aschenborn. Examined non-type material: See Supplementary information. Distribution: N. macrorubrus is distributed in dry savanna, karoo and grassland, un- der cooler conditions in South Africa (Northern Cape and Free State), Namibia (Etosha Pan Okaukuejo) and Botswana (near Kanye) (Fig. 31). Remarks: The type series of S. rubripes Boheman 1857 contained specimens belonging to two different species. P��ringuey (1901) described one of Boheman���s type specimens under the name S. rubripes. However, the characters which P��ringuey discussed did not match those of the holotype of S. rubripes (Arrow 1927), which means that, P��ringuey was describing an unnamed species. The P��ringuey species was recognized as new by Haaf (1955), Ferreira (1972), and later, Paschalidis (1974b) who proposed the current official name for the species., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 54-55, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Paschalidis, K. M. (1974 b) The identity of two species in the genus Sisyphus (Coleoptera: Scarbaeidae). Journal of Entomology of Southern Africa. 37 (2): 299 - 303.","Boheman, C. H. (1857) Insecta Caffrariae annis 1838 - 1845 a JA Wahlberg collecta. Pars. II. Coleoptera (Scarabaeides). Holmiae: Offieina Nordstedtiana. - 395 pp, 1.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088."]}

Neosisyphus infuscatus (Klug, 1855) (Figs. 20, 21B) Montreuil 2015b: 10; Sisyphus infuscatus Klug, 1855 Klug 1855: 651; Harold 1869: 984; Gillet 1911: 23; Arrow 1927: 464; Haaf 1955: 344; Ferreira 1972: 811. Size: Male: length: 8.1��� 7.1 mm; width: 4.8���3.6. Female: length: 8.0���7.0 mm; width: 4.5���3.0 mm Type locality: Sena, Sofala, Mozambique Diagnosis: N. infuscatus bears a sharp mid-posterior metafemoral spine as opposed to the more obtuse spine in N. spinipes. N. infuscatus also differs by lacking the prominent sharp-angled protrusion on the basal-posterior edge of the mesotibia. Parameres of N. infuscatus are distinctly bent inwards at the apex (Fig. 21B). Examined type material Paralectotype (designated here): (♂ SAM) /Sena/, Sisyphus fortuitus, PJ/, red label / Type SAM, 2677 / spinipes Thunb, det. Haaf 1954/, / Sisyphus infuscatus Klug, det. KM. Paschalidis 1975/, / Paralectotype, det: GM, Daniel, 2016 /. Examined non-type material: See Supplementary information. Distribution: N. infuscatus is an East to southern African species, associated with lowland dry savanna (Fig. 22)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 49, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Klug, J. C. F. (1855) Diagnosen neuer Coleoptera aus Mossambique. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 643 - 660.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74."]}

Neosisyphus kuehni (Haaf, 1955) (Figs. 26, 27B) Montreuil 2015: 28. Sisyphus kuehni Haaf, 1955 Haaf 1955: 345; Montreuil 2015: 28. Type locality: Natal (Griqualand), South Africa. Size: Male: length: 13.5���10; width: 6.5���5.5; Female: length: 13.0���11; width: 6.5���5.6. Diagnosis: N. kuehni bears a gibbosity apically on the metafemur, which is lacking in the closely-related N. quadricollis. The projecting metatrochanter in males of N. kuehni is also longer, comprising over half of the total length of the metafemur. Unlike in N. quadricollis, the elytra in N. kuehni are not laterally emarginate. Examined non-type material: See Supplementary information. Distribution: N. kuehni occurs in highland grassland under the moist to wet conditions of the eastern escarpment of South Africa (Eastern Cape, KwaZulu Natal and Mpumalanga) (Fig. 28). Conservation status: N. kuehni was assessed as a Data Deficient species for the IUCN Red Date List (Davis 2013g). Further collecting and monitoring is required to accurately assess the conservation status of this species., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 53-54, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Davis, A. L. V. (2013 g) Neosisyphus kuhni. The IUCN Red List of Threatened Species 2013: e. T 138156 A 537143. http: // dx. doi. org / 10.2305 / IUCN. UK. 20132. RLTS. T 138156 A 537143. en. (accessed 14 October 2016)."]}

Sisyphus umbraphilus Daniel & Davis, 2016 (Figs. 8, 9A) Daniel et al. 2016: 67���69 Size: Male: length: 5.1���7.1 mm, width: 2.9���4.2 mm; Female: length: 5.4���7.4 mm, width 3.2���4.2 mm. Type locality: Umfolozi, KwaZulu Natal (South Africa) Diagnosis: S. umbraphilus belongs to its own species-group. However, it is somewhat similar to S. oralensis. In this case, S. umbraphilus is distinguished by having tufts of epipleural setae. It bears relatively small ocellate punctures on the pronotum. Furthemore, In S. umbraphilus, the parameres are simple and attenuated towards the apex (Fig. 9A) whereas, in S. oralensis they are somewhat obtuse and truncated towards the apex (Fig. 14A). Examined type material Holotype: (♂ SANC): SOUTH AFRICA, KZN, Umfolozi 28��15���S 31��55���E, 1���7.x. 1970, leg: Bornemissza & Aschenborn. Paratypes: (3♂, 9♀ SANC) with the same data as holotype. (1♂, 2♀ UPSA): SOUTH AFRICA, KZN, Ithala Game Reserve 27��27���53.6���S 31��15���36.1���E, 13.i.1999, leg: Chown, McGeogh & Davis. (2♂ SANC): SOUTH AFRICA, KZN, Jozini 27��26���0���S 32��4���0���E, 15.x.1978, leg: Bornemissza & Aschenborn. (7♂, 14♀ SANC): SOUTH AFRICA, Gauteng, Pretoria, 35 km NE of Pretoria Farm 25��42���0���S 28��13���0���E, 12���13.xii.1983, leg: Davis. (1♂ SANC): SOUTH AFRICA, KZN, Weke-Weke Stream, Shongweni Farm 29��48���S 30��43���E, 11.xi.2010, leg: AJ Armstrong & G Van Bassouw. Examined non-type material: See Supplementary information. Distribution: S. umbraphilus has been recorded in patches of dense woodland and thicket in KwaZulu Natal (Daniel et al. 2016) as well as in dense riverine woodland in North West Province (South Africa). In Gauteng, it has been collected from shaded vegetation on deep sands (Davis 1996) (Fig. 10)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 26-27, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Davis, A. L. V. (1996) Habitat associations in a South African, summer rainfall, dung beetle community (Coleoptera: Scarabaeidae, Aphodiidae, Staphylinidae, Histeridae, Hydrophilidae). Pedobiologia, 40 (3): 260 - 280."]}

Neosisyphus M��ller, 1942 M��ller 1942: 86; Haaf 1955: 346; Ferreira 1972: 810���844; Schaefer & Fisher 1992: 130; Montreuil 2015b: 1���44 Type species: Neosisyphus spinipes (Thunberg, 1818), Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 42, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Muller, G. (1942) Nuovi coleotteri dell'Africa Orientale (2 a serie) Atti del Museo Civico di Storia Naturale di Trieste, 15 (3): 63 - 86.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Thunberg, C. P. (1818) Coleoptera Capensia, Antennis Lamellatis, sive, clava fissili instructa, descripta, memoires de l'Acad. T. VI."]}

seminulum species-group (Figs. 14 A���D, 15A���C). Sisyphus splendidus Montreuil 2015 (Fig. 14A). Montreuil 2015a: 91���102 Type locality: Otavi mts, Farm Gauss (Namibia) Size: Female: length: 4.5 mm, width: 2.5 mm. Diagnosis: S. splendidus may be separated from other members of the seminulum species-group by the distinct metallic sheen on the pronotal disc. Examined non-type material: See Supplementary information. Distribution: Sisyphus splendidus is found in unshaded vegetation of northern Namibia and southern Angola (Fig. 16)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 37, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102."]}

Sisyphus perissinottoi Montreuil, 2015 (Fig. 2, 3D). Montreuil 2015:10 Type locality: Amsterdamhoek (South Africa, Eastern Cape) Size: Male: length: 7.0���5.0 mm; width: 3.0��� 2.5 mm. Female: length: 7.3���5.0 mm, width 3.0��� 2.5 mm. Diagnosis: S. perissinottoi resembles S. manni. Setae on the pronotal discs of both species arise from the centre of ocellate punctures (Fig. 5H). However, S. perissinottoi differs by bearing tufts of dense and compact setae on the elytra and, lacking depressions on the latero-posterior edge of the pronotal disc. S. perissinottoi is also larger than S. manni. Examined type material Holotype: (MHNH, through photograph without locality data). Examined non-type material: See Supplementary information. Distribution: S. perissinottoi is only known from the Eastern Cape (South Africa) (Montreuil 2015c). It has been recorded from unshaded vegetation. (Fig. 4)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 15, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293."]}

Sisyphus australis Daniel & Davis sp. n. (Fig. 11, 12E). ZooBank: http://zoobank.org/ 762856F2-064A-4B6C-9A1C-BB22BA70A271 Diagnosis: Sisyphus australis sp. n. is similar to S. costatus. However, the new species is distinguished by the straight margin between the medial clypeal teeth. It also bears a strong clypeo-frontal carina (Fig. 11H). In addition, the parameres of S. costatus and S. australis sp. n. are distinctly different (Fig. 12 B, E) Description: Male holotype: Size. Length: 5.7 mm; width: 3.2 mm. Colour. Black body; setae brown; meso- and metasternum black; antennae grey. Head. Medial teeth not sharp apically; edge between medial teeth separated by a straight margin; a rectangular margin between the medial tooth and lateral indentation; genal margin straight. Dorsal surface of the clypeus setose, without punctation; clypeo-frontal carina distinctly defined; vertex setigerous with scattered ocellate punctures (Fig. 11H). Pronotum. Convex; maximum length longer than maximum width; dorsal surface with setation and ocellate punctures of different sizes; setae on the antero-lateral projection well-developed. Complete lateral prothoracic margin between the prothoracic disc and prothoracic episternum. Elytra. Elongate, narrow posteriorly; elytral striae minutely punctate and crenulate, characterized by a distinct crenulate double line, which is interrupted by fine ocellate strial puctures; interstriae with alternating rows of well or weakly developed setae; hind wing and venation are fully developed. Pygidium. Setigerous with ocellate punctation. Sternites. Abdominal sternites finely crenulate, setigerous with ocellate punctation; setae arranged in rows laterally; meso-metasternal suture visible; punctate depression on the postero-medial surface of the metasternum. Legs. In ventral view, antero-lateral carina of profemur punctate, with a lateral row of fine and well developed setae; meso- and metafemur with granulation, densely punctate and setose; meso- and metatrochanter contiguous with femur, metatrochanter projected somewhat backwards; protibia with three teeth and single terminal spine; mesotibia punctate and setose with two terminal spines; metatibia densely setose, serrated laterally with two spurs; pro- meso- and metatarsi five segmented with two claws, setose laterally; first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the external edge. Aedeagus. Parameres simple and symmetrical (Fig. 12E) Morphological variation Size: Male: length: 8.2– 5.5 mm; width: 3.2–4.0 mm; Female: 8.0–6.0 mm; width: 3.0– 3.6 mm. Head: Mainly in populations from the Eastern Cape (The Haven), the surface of the clypeus is less setigerous. Male: Meta- and mesotibia curved; last abdominal sternite narrowed medially (Fig. 1A). Female: Meta- and mesotibia almost straight; last visible abdominal sternite not constricted medially (Fig. 1B). Examined type material Holotype: (♂ TMSA) SOUTH AFRICA: /S. Cape Prov (Western Cape), Outeniekwaberg, 33.50°S; 23.11°E /; 13.12.1977; E-Y: 1422, dung collecting, leg: Endrödy-Younga. Paratypes: (3♂, 3♀ TMSA) same data as holotype. (1♂, 1♀ TMSA) SOUTH AFRI- CA: Cape Prov. (Western Cape) George, Saasveld, 205, leg: Breytenbach. (1♀ SANC) SOUTH AFRICA: Cape Prov. (Western Cape) George, Saasveld, 205, leg: Breytenbach. (1♂ SANC) SOUTH AFRICA: C.P. (Western Cape) (10 km SW), 3.iii.1973, leg: Davis. (3♂, 5♀ TMSA) SOUTH AFRICA: Transkei (Eastern Cape), The Haven, 32.15°S 28.55°E, 09.xii.1979, E-Y: 1696, groundtraps, 7 days, leg: Endrödy-Younga. (2♂, 2♀ TMSA): SOUTH AFRICA: Transkei (Eastern Cape), Coast Dwesa forest, 32.17°S 28.50°E, 5.iii.1985, zebra dung, leg: Endrödy-Younga. (1♂ TMSA): SOUTH AFRICA: Transkei (Eastern Cape), Alexandria For. St., 33.43°S 26.23°E, 5.xii.1987, groundtraps with faeces bait, 2 days, leg: Endrödy-Younga. (1♀ SANC) Transkei (Eastern Cape), Wavecrest, 25.xi.1981, leg: Doube. Etymology: The species name reflects the geographical distribution of the species, which is restricted to the southern part of South Africa. Distribution: Sisyphus australis sp. n., has been recorded along the coastline in association with shaded vegetation of the Western and Eastern Cape Provinces, South Africa (Fig. 13). Remarks: Detailed study of type and non-type material of S. gazanus and S. costatus have revealed a species complex composed of S. costatus; S. australis sp. n.; S. inconspicuous sp. n. and S. gazanus. The main distinguishing features for the species complex are present- ed in Table 1. Furthermore, the shape of the parameres (Figs. 12 A–E) and the distributions (Fig. 13) are useful to separate the four species belonging to the gazanus complex.

Sisyphus muricatus (Olivier, 1789) (Figs. 2, 3A) Scarabaeus muricatus Olivier, 1789 Olivier 1789: 188���9, Gory 1833: 7���8; Harold 1869: 985; P��ringuey 1902: 105���6; Arrow, 1927: 458; Haaf 1955: 367���8; Ferreira 1972: 801; Paschalidis 1974: 7���9, Montreuil 2015c: 3, 11; Daniel et al. 2016: 71. Type locality: Cape of Good Hope (South Africa). Size: Male: length: 12.5���11.0 mm, width: 8.0��� 6.5 mm; Female: length: 12��� 9.6 mm, width 6.5���5.0 mm. Diagnosis: S. muricatus is similar to S. neobornemisszanus. However, it differs by bearing, mostly, more strongly developed clypeo-frontal carinae; also, very strong, deep, lateral emargination and distinct shiny punctation on the pronotum. Examined non-type material: See Supplementary information. Distribution: S. muricatus shows an Afrotemperate forest and montane grassland distribution in South Africa (Davis et al., 2008, 2002 a, 1999) (Fig. 4). It has been recorded from grassland in the Golden Gate Highlands National Park and the foothills of the Drakensberg. It has also been collected in grassland at Mount Sheba. There are records of this species in coastal afrotemperate forest (Davis et al. 1999) in the Cape (Knysna Forest, Garden of Eden, and George). However, these qualitative data are insufficient to support a significant association with shaded vegetation on the south coast of South Africa. Remarks: The species was originally described as Scarabaeus muricatus by Olivier (1789) from an unspecified type locality in ���la Am��rique m��ridionale���. Gory (1833) transferred the species to the genus Sisyphus citing the Cape of Good Hope (South Africa) as the type locality., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 11, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Olivier, M. (1789) Entomologie, Histoire Naturelle des Insectes, Avec leurs caracteres generiques et specifiques, leur description, leur synonymie, et leur figure enluminee. Coleopteres, Tome premier, Paris, 1 - 455 pp.","Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Montreuil, O. (2015 c) Premiers cas de brachypterisme dans le genre Sisyphus Latreille (Coleoptera, Scarabaeidae, Sisyphini). Annales de la Societe entomologique de France, 51 (4): 281 - 293.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Davis, A. L. V., Frolov, A. V., & Scholtz, C. H. (2008) The African dung beetle genera. Protea Book House. 148 - 155 pp.","Davis, A. L. V., Scholtz, C. H., & Philips, T. K. (2002 a) Historical biogeography of scarabaeine dung beetles. Journal of Biogeography, 29 (9): 1217 - 1256.","Davis, A. L. V., Scholtz, C. H., & Chown, S. L. (1999) Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa. Journal of Biogeography, 26 (5): 1039 - 1055."]}

Neosisyphus quadricollis (Gory, 1833) (Figs. 26, 27A) Montreuil 2015b: 20. Sisyphus quadricollis Gory, 1833 Gory 1833: 9; Harold 1869: 985; P��ringuey 1901: 87; Arrow 1927: 459; Haaf 1955: 345; Ferreira 1972: 829. Type locality: Cape of Good Hope. Size: Male: length: 13.5���11; width: 6.5���5.4; female: length: 13.8���11; width: 6.5���5.5. Diagnosis: N. quadricollis is close in appearance to N. kuehni. However, N. quadricollis may be separated by a combination of the following characters: elytra emarginated laterally with the concavity of the margin distinctly deep in the middle; the projecting metatrochanter in males is one-third of the total length of the metafemur; the metafemur lacks spines. Examined non-type material: See Supplementary information. Distribution: N. quadricollis has been recorded from deep sands and natural shrubland under drier, warmer winter rainfall climate on the southwest coast of South Africa (Western Cape) (Davis 1987, 1997; Davis et. al. 2008). In this region, it is active for a short period between late October to late December. It also has been collected between November and March at Willowmore and Lemoenkloof (Eastern Cape) in karoo vegetation of the late summer rainfall region (Fig. 28)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 52-53, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Gory, M. (1833) Monographie du genre Sisyphe. Mequignon-Marvis pere et fils, Paris. 1 - 15 p.","Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Harold, E. von (1869). Scarabaeidae. Gemminger, M. and Harold, E. Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus, 4: 976 - 1346. http: // www. iucnredlist. org / static / categories _ criteria _ 3 _ 1 access on 2017.03. 23.","Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Davis, A. L. V. (1987) Geographical distribution of dung beetles (Coleoptera: Scarabaeidae) and their seasonal activity in south-western Cape Province. Journal of the Entomological Society of Southern Africa, 50 (2): 275 - 285.","Davis, A. L. V. (1997) Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.). african Journal of ecology, 35 (1): 10 - 38."]}

Sisyphus nanniscus P��ringuey, 1901 (Figs. 14C, 15B). P��ringuey 1901: 107���8; Gillet 1911:23; Arrow, 1909: 519; Arrow 1927: 465; Haaf: 355���6; Montreuil 2017: 3 Sisyphus ocellatus nanniscus P��ringuey, 1901 Haaf 1995: 357���8; Ferreira: 1972: 807; Type locality: Durban (South Africa) Size: Male: length: 3.0��� 3.8 mm; width: 1.7��� 1.5 mm; Female: length: 3.0���4.0 mm, width 1.8��� 1.5 mm. Diagnosis: S. nanniscus resembles S. seminulum. However, it can be easily distinguished as it bears large, dense ocellate punctures on the pronotum, unlike in S. seminulum where the punctures are relatively smaller and clearly separated from each other. Additionally, in S. nanniscus the internal margin of the metatibia is weakly dentate whilst, in S. seminulum, it is distinctly serrated. Examined type material Sisyphus nanniscus P��ringuey, 1901 : (♂ SAM): Lectotype (designated here): handwritten on brown label / Sisyphus nanniscus, type /, red label /type, SAM, ento: 2672/, / Lectotype, det. GM Daniel, 2016 /. (♂ SAM): Paralectotype (designated here): / Durban, x. 1896 leg: undetermined/, (white label) / Sisyphus seminulum Gerstaecker, det. K.M. Paschalidis, 1975/, (handwritten on brown label) / Sisyphus nanniscus, type/, (handwritten on brown label bordered by blue colour) / Ssp. nanniscus P��r /, red label / type /, / Lectotype, det. GM Daniel, 2016 /. Sisyphus seminulum Gerstaecker, 1871 : Lectotype (designated here): (1♀ MN- HUB): / Zanzibar /, (brown label) / typus/, /56381/; handwritten on blue label / seminulum Gerst /; (red label) / Syntypus Sisyphus seminulum Gerstaecker 1871, labelled by MNHUB 2016/, /leg: Cooke/; / Lectotype, det. GM Daniel, 2016 /. Paralectotype (designated here): (1♂ MNHUB): / Zanzibar /; brown label / paratypus /, / 56381 /; handwritten on blue label / seminulum Gerst, det: Muller/, red label / Syntypus Sisyphus seminulum Gerstaecker 1871, labelled by MNHUB 2016/, leg: Cooke, / Paralectotype, det. GM Daniel, 2016 /. (3♂, 2♀ MCZ): / Zanzibar / C. Cooke/, (red label written) / type/ S. seminulum /, costatus, det. PFS Pereira/ Paralectotype, det. GM Daniel, 2017/. Examined non-type material: See Supplementary information. Distribution: Sisyphus nanniscus is found primarily under shaded vegetation in southern Africa (Fig. 16). Remarks: Type material has been studied of both S. nanniscus and S. seminulum. External morphology and parameres are sufficiently different for separation into two species. Therefore, we support the revalidation and new status of S. nanniscus proposed by Montreuil (2017)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 39-40, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Peringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African philosophical Society. Cape Town, 12: 1 - 563 (13 - 368).","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Arrow, G. J. (1909) On some new species of Coleoptera from Rhodesia and adjacent territories. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London 8 (3): 517 - 523.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Montreuil, O. (2017) Nouveaux Sisyphus Latreille africains du groupe seminulum: les complexes seminulum, desaegeri et ocellatus (Coleoptera, Scarabaeidae, Sisyphini), Catharsius La Revue, 14: 1 - 22.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Gerstaecker, A. (1871) Beitrag zur Insektenfauna von Zanzibar III. Coleoptera. Archiv fur Naturgeschichte, 37: 42 - 86."]}

Sisyphus impressipennis Lansberge, 1886 (Figs. 11, 12A). Lansberge 1886: 73���74, Gillet 1911: 23; Arrow 1927: 462���5; Haaf 1955: 362; Schefer-Fischer 1992: 124, Schaefer-Fischer, 2001:58, Montreuil 2015a: 96. Sisyphus transvaalensis P��ringuey, 1908 P��ringuey 1908: 557; Arrow 1927: 462; Haaf 1955: 364; Balthasar 1968: 954; Ferreira 1972: 807; Montreuil 2015a: 96 Sisyphus callosipes Arrow, 1909 Arrow 1909: 517; Arrow 1927: 462; Haaf 1955: 365; Montreuil 2015a: 96. Type locality: Humpata (Angola). Size: Male: length: 10.0��� 7.1 mm; width: 5.6���4.0 mm; Female: length: 10.1��� 7.5 mm, width 5.1���4.0 mm. Diagnosis: S. impressipennis is easily distinguished from other species in the costatus species-group by bearing an anvil-shaped projection on the mid-ventral edge of the metafemur, mainly in males. Examined type material Sisyphus callosipes Arrow, 1909 : Lectotype: (1♂ NHML), / MOZAMBIQUE, Nyassa /, /coll: Simons & Fry, 1905/, handwritten on brown label / Sisyphus callopsis, Arrow type /, / Sisyphus callopsis det. ME. Bacchus, 1975; Lectotype /. Paralectotype: (1♀ NHML) / Mozambique, Nyassa /, /coll: Simons & Fry, 1905/, handwritten on brown label/, / Sisyphus callopsis det. ME. Bacchus, 1975; Paralectotype /. (1♂, 1♀ NHML), / RDC, 150���200 miles, W. of Kambove (3500���4500 ft)/, /coll: Neave, 18.10.1907 /, / Sisyphus callopsis det. ME. Bacchus, 1975. Sisyphus transvaalensis P��ringuey, 1908 : Lectotype (designated here): (1♂ SAM): / SOUTH AFRICA, Transvaal, Pietersburg /, handwritten on brown label, / Sisyphus transvaalensis P��ringuey /, /red label, /TYPE/ (Lectotype det. GM. Daniel 2016). Examined non-type material: See Supplementary information. Distribution: S. impresssipennis was described from south-west Angola. It is distributed in east-central and southern Africa (Davis et al. 2005) where it is associated with shaded areas in savanna and dense miombo woodland. It has been recorded from lowlands in northern South Africa, in upland Chirinda Forest (Zimbabwe), Mt Mulanje (Malawi), Central Mozambique, Manyara National Park (Tanzania) and Kambove (Republic Democratic of Congo) (Fig. 13C). Remarks: Examination of S. callosipes and S. transvaalensis type material showed that the external morphology and genitalia are similar to S. impressipennis. Therefore, we regard both species as junior synonyms of S. impressipennis, as previously proposed by Paschalidis (1974) and Montreuil (2015a)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on pages 28-29, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Lansberge, J. W. (1886) Scarabaeides, Buprestides et Cerambycides de l'afrique occidentale. Notes from Leyden Museum, 8 (2): 69 - 120.","Gillet, J. J. E. (1911) Scarabaeidae: Coprinae I. Coleopterorum Catalogus, 38: 1 - 263.","Arrow, G. J. (1927) Notes on the Coleopterous genus Sisyphus. Annals and Magazine of natural History, 9 (19): 456 - 465.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Schafer, P., & Fischer, E. (1992) Zur Scarabaeiden-Fauna Rwandas, Zentralafrika. Sisyphini (Insecta: Coleoptera: Lamellicornia). Senckenbergiana biologica, 72 (1 - 3), 119 - 137.","Schafer, P. & Fischer, E. (2001) Zur Scarabaeidenfauna Rwandas, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini, Mitteilungen des internationalen entomologischen Vereins supplement 9: 1 - 93.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102.","Balthasar, V. (1968) Neue Scarabaeiden-Arten, 129. Beitrag zur kenntnis der Scarabaeoidea (Coleoptera). Beitrage zur Entomologie, 18: 953 - 958.","Ferreira, M. C. (1972) Os Escarabideos de Africa (sul do Saara). Revista de Entomologia de Mocambique, 11 (1968 - 1969): 5 - 1088.","Arrow, G. J. (1909) On some new species of Coleoptera from Rhodesia and adjacent territories. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London 8 (3): 517 - 523.","Daniel, G. M., Davis, A. L. V, & Scholtz, C. H. (2016) Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa. Zootaxa, 4147 (1): 67 - 74.","Davis, A. L. V., Scholtz, C. H., & Deschodt, C. (2005) A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna. African entomology, 13 (1): 1 - 16.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp."]}

Neosisyphus setiger Roth (1851) (Figs. 7, 8D) Montreuil 2015b: 11 Sisyphus setiger Roth, 1851 Roth 1851: 124; Montreuil 2015b: 11 Neosisyphus confrater (Kolbe, 1914) Montreuil 2015b: 11 Sisyphus confrater Kolbe, 1914 Kolbe 1914: 317; Haaf 1955: 376; Type local: Bukoba (Tanzania) Size: Male: length: 9.0���5.0 mm; width: 4.8���3.0 mm. Female: length: 9.2���5.0 mm; width: 4.5���3.0 mm Diagnosis: N. setiger is distinguished from other members of the barbarossa speciesgroup, by a combination of the following features: a triangular protrusion on the midposterior edge of the metafemur in males. Additionally, the projected metatrochanter is relatively short, comprising �� of the total length of the metafemur in males. Examined non-type material: See Supplementary information. Distribution: As defined by Montreuil (2015b), N. setiger is distributed in moist lowland and upland vegetation from northeast to southern Africa (Fig. 19). Remarks: We have compared Kenyan specimens identified as N. setiger with southeast African specimens identified as N. confrater by Paschalidis (1974). Females of N. confrater and N. setiger are close morphologically making it difficult to separate them into species. However, we have observed that males of southern Africa ��� N. confrater ��� bear a short blunt triangular projection on the mid-posterior edge of the metafemur and a short projecting metatrochanter. By contrast, males of east African ��� N. setiger ��� have a sharp projecting spine on the mid-posterior edge of the metafemur and a metatrochanter as large as in Neosisyphus jossoi Montreuil, 2015. Montreuil (2015b) synonymized N. confrater with Neosisyphus setiger (Roth 1851). The synonymy seems to be based on the holotype female of N. setiger from Ethiopia and a lectotype male of N. confrater from Tanzania. This is questionable as females are often difficult to separate into species as stated above. Furthermore, although the photograph of a major male of N. setiger (Montreuil 2015b) closely resembles ��� N. confrater ���, there is no indication of where the individual was collected. Nevertheless, we have retained the name N. setiger for southern African material as we have not been able to compare it with the type and non-type material used in Montreuil���s study (2015b)., Published as part of Daniel, Gimo M., Davis, Adrian Lv., Sole, Catherine L. & Scholtz, Clarke H., 2020, Taxonomic review of the tribe Sisyphini sensu stricto Mulsant, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern Africa, including new species descriptions, pp. 1-61 in Insect Systematics & Evolution 51 (1) on page 47, DOI: 10.1163/1876312X-00002195, http://zenodo.org/record/3786629, {"references":["Montreuil, O. (2015 b) Le genre Neosisyphus Muller em Afrique (Coleoptera, Scarabaeidae, Sisyphini). Catharsius La Revue, 12 (1): 1 - 36.","Kolbe, H. J. (1914) Lieferung 3: Scarabaeiden. In: Shubotz, H. (Ed.), Wissenschaftliches Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907 unter Fuhrung Adolf Friedrichs, Herzog zu Mecklenburg. Zoologie III, 5. Klinkhardt & Biermann, Leipzig, Germany, pp. 261 - 416.","Haaf, E. (1955) Uber die Gattung Sisyphus Latr. (Col. Scarab.). Entomologische Arbeitenaus dem Museum G. Frey, 6 (1): 341 - 381.","Paschalidis, K. M. (1974 a) The genus Sisyphus Latr. (Coleoptera: Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes University, Grahamstown, South Africa, 183 pp.","Montreuil, O. (2015 a) Nouveaux Sisyphus Latreille d'afrique (Coleoptera, Scarabaeidae, Sisyphini). Bulletin de la Societe entomologique de France, 120 (1): 91 - 102."]}

Sisyphus neobornemisszanus Daniel & Davis sp. nov. (Fig. 3a) Sisyphus bornemisszanus Endr��di, 1983 (pars) Diagnosis. This species is close to Sisyphus muricatus (Olivier, 1789) and Sisyphus fasciculatus Boheman, 1857. Like S. neobornemisszanus, S. muricatus also has tufts of setae only on the elytra with uniformly distributed setae on the pronotum. However, the carinae on the head of S. muricatus are mostly more developed with deeper and very strong lateral emargination on the pronotum. S. fasciculatus is easily distinguished from S. neobornemisszanus sp. nov. by bearing black tufts of setae on both the pronotum and elytra (Fig. 3 b). Description. Size: Male: Length: 7.6���9.7 mm, width: 4.6���5.6 mm; Female: Length: 8.0��� 10 mm, width 4.4��� 5.62 mm. Colour: Black, opaque; except mouth parts, and tarsi which are reddish brown; Antennal club dark gray or dark brown. Head: Apex of clypeus broadly and deeply emarginate, bordered by two acute medial teeth and two more obtuse lateral teeth, genae angular anteriorly and curved laterally, surface of clypeus pubescent and finely punctate. Pronotum: Convex with dense cover of setae, much broader than long, lateral edges curved anteriorly, narrowed posteriorly, complete lateral prothoracic ridge separating prothoracic disc and prothoracic episternum. Elytra: narrowed posteriorly, interstriae 1, 3, 5, and 7 with rows of black setae arranged in tufts, interstriae 2, 4, and 6 with sparsely arranged single setae, a rudimentary row of setae along outer margin of abdominal pleurae. Pygidium: V-shaped, with indistinct rows of brown setae and ocellate punctation. Sternites: Six visible abdominal sternites with setae and ocellate punctures; meso- and metasternum black, shiny and pubescent with ocellate punctation; meso-metasternal suture clearly visible. Legs: Profemur punctate ventrally and setose laterally; meso- and metafemur pubescent with ocellate punctation; mesotrochanter short and contiguous with femur; metatrochanter finely punctate, slightly angular and projected away from femur; Protibia with three teeth and a single terminal spurs; meso- and metatibia serrated, setose with two terminal spurs; pro- meso- and metatarsus five segmented with two claws; the first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the ventral edge. Aedeagus: Phallobase slightly curved dorsally as is common in the genus Sisyphus; parameres symmetrical, short, triangular, separated by membranous area dorsally, sides almost straight and slightly impressed above (Fig. 3 c). Sexual dimorphism. Female: Lateral carina of pronotum somewhat stronger and more curved than in male; protibial teeth longer; meta- and mesotibia almost straight; last visible abdominal sternite not constricted medially. Male: Protibial teeth strong and short with rudimentary lateral setae; meta- and mesotibia curved; last abdominal sternite narrowed medially. Etymology. Species named as S. neobornemisszanus, because the designated holotype was in the mixed type series of S. bornemisszanus. Examined material. ( ♂ TMSA) Holotype: SOUTH AFRICA, Zululand, St. Lucia 28[���] 13[���] 12[������] S; 32[���] 15[���] 00[������] E, 7.xii.1975, leg: Endrody-Younga. Paratypes: (1♀ TMSA) SOUTH AFRICA, Zululand, Mission Rock, 28[���] 15[���] 36[������] S; 32[���] 17[���] 24[������] E, 7.xii.1975, legs: Endrody-Younga. (2♂ TMSA) SOUTH AFRICA, Zululand, St. Lucia 28[���] 13[���] 12[������] S; 32[���] 15[���] 00[������] E, 28.viii.1971. leg. GF. Bornemissza. (2♂ TMSA) SOUTH AFRICA, Zululand, St. Lucia, Estuary. 28.viii.1971, leg: GF. Bornemissza. (15♂, 13♀ TMSA) SOUTH AFRICA, Zululand, Sodwana Bay, 5km, 27[���] 21[���] 0[������] S; 32[���] 23[���] 24[������] E, 2.xi.1992, leg. Endrody-Younga. (1♀ UPSA) SOUTH AFRICA, KZN. Richard���s Bay, 28[���] 37[���] 39[������] S, 32[���] 17[���] 24[������] E, 27.i.2000, leg: Davis & Delport. (2♀, 6♂ UPSA) SOUTH AFRICA, KZN. Richard���s Bay, 28[���] 43[���] 28.5[������] S; 32[���] 10[���] 52[������] E, 27.i.2000, leg: Davis & Delport. (5♀, 2♂ UPSA) SOUTH AFRICA, KZN, Sileza Nature Reserve 27[���] 06[���] S; 32[���] 36[���] E, 03.iv.1996, leg: Van Resnburg. (1♀ UPSA) SOUTH AFRICA, KZN, Sodwana, 27[���] 32[���] S; 32[���] 41[���] E, 07.iv.1989, leg: Mansfield. (2♀, 5♂ UPSA) MOZAMBIQUE, Maputo, Elephant Maputo Reserve, 26[���] 22[���] 30[������] S; 33[���] 47[���] 50[������] E, 14-16.i.2003, leg: Scholtz & Holter. (2♀ UPSA) MOZAMBIQUE, Maputo, Elephant Maputo Reserve, 26[���] 39[���] S; 32[���] 43[���] E, 10-16.xi.2007, pitfall trap, leg: Strumpher & Deschodt. (1♂ TMSA) MO��AMBIQUE, Inhambane, Inharrime, 24[���] 28[���] 37[������] S; 35[���] 01[���] 49[������], 1974, leg: Moor. (1♂ TMSA) MO��AMBIQUE, Inhambane, Pomene 22[���] 35[���] 24[������] S; 35[���] 21[���] 00[������], 2.v.1974, leg. A. Strydom. Habitat. This species has been recorded in KwaZulu-Natal Province (South Africa) as well as Maputo and Inhambane Provinces (Mozambique), although records from Pomene suggest a wider distribution northwards along the Mozambique coastline (25.11 S; 33.73 E). In Maputaland, it is found within coastal sand and dune forests where the canopy is closed and the floor is open., Published as part of Daniel, Gimo M., Davis, Adrian Lv. & Scholtz, Clarke H., 2016, Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa, pp. 67-74 in Zootaxa 4147 (1) on pages 71-73, DOI: 10.11646/zootaxa.4147.1.4, http://zenodo.org/record/266251

Sisyphus oralensis Daniel & Davis sp. nov. (Fig. 2a) Diagnosis. Sisyphus oralensis sp. nov. is similar to S. umbraphilus sp. nov., but can be distinguished by the following combination of characters: genae rounded forming obtuse angle with genal suture; less dense cover of long setae on pronotal disc, not distributed uniformly but mostly arranged in a linear pattern separated by bare patches; S. oralensis sp. nov. is smaller than S. umbraphilus sp. nov. (Fig. 2 bi-ii). Description. Size: Male: Length: 3.4–4.8 mm, width: 2.1–2.5 mm. Female: Length: 3.9–4.7 mm, width: 2.1– 2.5 mm. Colour: Dark brown to black. Head: Clypeus with a strong cleft between two medial teeth, deep excavation lateral to each tooth, lateral convex sinuosity between excavation and genal suture, genae rounded forming obtuse angle with genal suture; frons finely punctate and setose, post-occipital suture fully rounded and arciform, eyes reniforme with dark ocular stains, antennae brown. Pronotum: Convex, dark brown, coarsely punctate; anteriorly inclined; central diameter wider than elytra; complete lateral prothoracic ridge separating prothoracic disc and prothoracic episternum; long setae of pronotal disc not uniformly spaced but forming a clear, mostly linear pattern separated by bare patches (Fig. 2 bii). Elytra: Dark brown; narrowed posteriorly; finely punctate; well-spaced; fine curled setae on interstriae. Pygidium: Dark brown; strongly narrowed and "v" shaped towards apical base; finely punctate and setose. Sternites: Abdominal sternites punctate with fine setae arranged in rows; meso- and metasternum black, metallic shiny, and finely punctate; mesepimerum and metepisternum: punctate with fine setae. Legs: Profemur with well-developed setae laterally, punctate ventrally; meso- and metafemur brown, punctate and setose; meso- and metatrochanter contiguous with femur, forming a divergent spine at the tip in latter; promeso- and metatibia setose with single terminal spur, protibia with three teeth; pro- meso- and metatarsus five segmented with two claws; the first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the ventral edge. Aedeagus: Phallobase curved dorsally, forming a strong angle with parameres, visible medial line ventrally; parameres nearly symmetrical, separated by membranous area visible dorsally on apical edge; basal edges slightly excavated longitudinally both ventrally and dorsally. (Fig. 2 c) Etymology. We named this species S. oralensis because it is found on the Indian Ocean coastline of South Africa and Mozambique where it is associated with dense woodland and forest. Examined material. Holotype: (♂ TMSA): SOUTH AFRICA, KZN, Richard’s Bay 28[◦] 39[’] 29.1[”] S; 32[◦] 15[’] 19.2[”] E, 27.i.2000, leg: Davis & Delport. Paratypes: (5♂, 8♀ UPSA): SOUTH AFRICA, KZN Richard’s Bay 28[◦] 39[’] S; 32[◦] 15[’] E, 26-27.i.2000, leg: Davis & Delport. (1♂, 2♀ UPSA): SOUTH AFRICA, KZN, Itala Game Reserve 27[◦] 27[’] 53.6[”] S; 31[◦] 15[’] 36.1[”] E, 13.i.1999, leg: Chown, McGeogh & Davis. (4♀ UPSA; 3♂, 2♀ SANC): SOUTH AFRICA, KZN, Thembe Elephant Park 27[◦] 01[’] S; 32[◦] 24[’] E, 17.xii.1996, leg: B.J. Van Rensburg. (9♂, 8♀ UPSA): SOUTH AFRICA, KZN, Thembe Elephant Park 27[◦] 01[’] S; 32[◦] 24[’] E, 17.vii.1995, leg: B.J. Van Rensburg. (2♂, 3♀ UPSA): SOUTH AFRICA, KZN Thembe Elephant Park 26[◦] 55[’] S; 32[◦] 23[’] E, 15-30.x.2008, leg: B.J. Van Rensburg. (2♂ SANC, 1 ♀ TMSA): SOUTH AFRICA, KZN, Umfolozi 28[◦] 15[’] S; 31[◦] 55[’] E, 25.viii.1971, leg: Bornemissza & Insley. ( 1♂, 10♀ SANC): SOUTH AFRICA, KZN St Lucia Est. Nat. Reserve, Dune Forest 28[◦] 13[’] S; 32[◦] 21[’] E, 27.x.1978, leg: Bornemissza. ( 1♀ SANC): SOUTH AFRICA, KZN Ntl Kosi Bay Nature Res 26[◦] 34[’] S; 32[◦] 28[’] E, 8- 11.ii.1990, leg: B. Grobbelaar. ( 1♀ SANC): SOUTH AFRICA, KZN, Near Richards Bay, 28[◦] 36[’] 52[’’] S; 32[◦] 17[’] 34[’’] E, 27.x.1978, leg: Bornemissza. (1♂ SANC): SOUTH AFRICA, KZN, Thembe Elephant Park 27[◦] 01[’] S; 32[◦] 24[’] E, 01-49. ii.1996, leg: R Stals. ( 5♂, 6♀ TMSA): SOUTH AFRICA, KZN, Ntl Kosi Bay Nature Res 26[◦] 34[’] S; 32[◦] 28[’] E, 14.xi.2002, leg: Burguer, Harrison & Muller. ( 7♂, 8♀ TMSA): SOUTH AFRICA, Zululand, Sodwana Bay 5km, 27[◦] 21[’] S; 32[◦] 23[’] E, 23.xi.1992, leg: Endrody-Younga. ( 8♂, 8♀ TMSA): SOUTH AFRICA, N Zululand, Ndumu Game Reserve 26[◦] 32[’] S; 32[◦] 10[’] E, 1.xii.1992, leg: Endrody-Younga. ( 30♂, 42♀ TMSA): SOUTH AFRICA, N Zululand, Lake Bangazi 28[◦] 04[’] S; 32[◦] 18[’] E, 12.xii.1992, leg: Endrodi-Younga. ( 1♂, 2♀ TMSA): SOUTH AFRICA, Zululand, St. Lúcia. Mission Rock 28[◦] 13[’] S; 32[◦] 21[’] E, 23.xi.1992, leg: Endrody-Younga. ( 1♂, 2♀ TMSA): SOUTH AFRICA, Natal, Cape Vidal, Forest litter 28[◦] 8[’] S; 32[◦] 33[’] E, 23.i.1990, leg: J. Klimaszewski. ( 1♂, 2♀ TMSA): SOUTH AFRICA, N Zululand, Hluhluwe Game Reserve. 28[◦] 03[’] S; 32[◦] 02[’] E, 20.xi.1992. leg: Endrody-Younga. (12♂, 30♀ SANC); SOUTH AFRICA, KZN St Lucia Est. Nat. Reserve, Forest 28[◦] 13[’] S; 32[◦] 21[’] E, 24-23.i.1979, leg: NH Aschenborn. ( 1♂, 2♀ TMSA): MOZAMBIQUE, Inhambane, Pomene, 22[◦] 35[’] S; 35[◦] 21[’] E, 04.v.1974, leg: A. Strydom. (5♂, 5♀ UPSA): MOZAMBIQUE, Maputo Elephant Reserve 26[◦] 39[’] S; 32[◦] 43[’] E, 10- 16.xi.2007, leg: W. Strümpher & C. Deschodt. (7♂, 10♀ SANC): MOZAMBIQUE, Sofala, Dondo 19[◦] 37[’] S; 34[◦] 45[’] E. 19.i.1972, leg: Bornemissza & Kirk. (1♂, 1♀ SANC): MOZAMBIQUE, Sofala, Beira 19[◦] 50[’] S; 34[◦] 51[’] E 19.i.1972, leg: Bornemissza & Kirk. (1♂ SANC); MOZAMBIQUE, Sofala, Gorongosa 18[◦] 57[’] S; 34[◦] 10[’] E, 11-28.i.1972, leg: Bornemissza & Kirk. Habitat. This new species is confined to dense coastal woodlands and forest from northeast South Africa to at least Central Mozambique (Inhambane, Pomene, Gorongosa, Dondo, Beira) (Fig. 4). Across a vegetation restoration gradient at Richards Bay, KwaZulu-Natal, South Africa, S. oralensis was absent from grassland cleared of forest, but present in younger and older regenerating woodland (19, 6) plus natural dune forest (14) (Davis et al. 2002). Remarks. Both species (S. umbraphilus sp. nov. and S. oralensis sp. nov.) exhibit little sexual dimorphism, but may be separated by meta- and mesotibia curved; the mid-ventral narrowing of the last abdominal sternite in males; compared to the lack of narrowing, and meta- and mesotibia almost straight in females

Sisyphus umbraphilus Daniel & Davis sp. nov. (Fig. 1a) Diagnosis. This new species is similar to S. oralensis sp. nov. but can be distinguished from the latter by the following characters: lateral edges of genae virtually parallel; second and third abdominal epipleurae with welldeveloped tufts of setae; pygidium with uniformly scattered setae, except for a distinct non-setose line medially (Fig. 1 bi-iii). Description. Size: Male: Length: 5.1���7.1 mm, width: 2.9���4.2 mm; Female: Length: 5.4���7.4 mm, width 3.2��� 4.2 mm. Colour: Dark brown to black. Head: Clypeus dark and punctate; anterior margin with two medial teeth and a convex lateral sinuosity between teeth and genal sutures; lateral margins of genae almost parallel, post-occipital suture not fully rounded; antennae brown. Pronotum: Convex, shiny black, densely punctate, lateral edges slightly compressed, maximum length shorter than maximum width, complete lateral prothoracic ridge between prothoracic disc and prothoracic episternum, disc showing an indistinct pattern of bare patches, some with sparse short setae, arranged within a matrix of dense, long setae (Fig. 1a). Elytra: Dark brown, shiny in some individuals, ovoid posteriorly, interstriae finely punctate bearing lightcoloured setae with recurved tips, epipleurae of the second and third abdominal segments each bearing a tuft of yellow setae. Pygidium: Narrow with ocellate punctation and scattered setae but with a distinct medial line lacking setae. Sternites: Setae arranged in rows laterally, except on first sternite where setae form a ventral row; visible meso-metasternal suture; meso- and metasternum black shiny, punctate with lateral setae; mesepimerum and metepisternum punctate and setose. Legs: Profemur black, punctate, internally pubescent with a lateral row of well-developed setae in ventral view; meso- and metafemur densely punctate and setose; meso- and metatrochanter contiguous with femur, sharp tip of metatrochanter projecting away from femur; metacoxa punctate and shagreened ventro-anteriorly; protibia with three teeth and single terminal spine; mesotibia punctate and setose with two terminal spines; metatibia densely setose, serrated laterally with two spurs; pro- meso- and metatarsus five segmented with two claws, setose laterally; first tarsal segment of the meso- and meta-thoracic legs with a row of strong setae on the ventral edge. Aedeagus: Phallobase almost straight, slight angular depression immediately distal from the phallobaseparamere suture; parameres symetrical, sligthly curved inward at the tip, separated from each other by fine membranous area visible dorsally on apical edge (Fig. 1 c). Etymology. The species name reflects the shaded vegetation with which the species is associated. Examined material. Holotype: (♂ SANC): SOUTH AFRICA, KZN, Umfolozi 28[���] 15[���] S; 31[���] 55[���] E, 1- 7.x. 1970, leg: Bornemissza & Aschenborn. Paratypes: (3♂, 9♀ SANC) with the same data as holotype. (1♂, 2♀ UPSA): SOUTH AFRICA, KZN, Itala Game Reserve 27[���] 27[���] 53.6[���] S; 31[���] 15[���] 36.1[���] E, 13.i.1999, leg: Chown, McGeogh & Davis. (2♂ SANC): SOUTH AFRICA, KZN, Jozini 27[���] 26[���] 0[���] S; 32[���] 4[���] 0[���] E, 15.x.1978, leg: Bornemissza & Aschenborn. (7♂, 14♀ SANC): SOUTH AFRICA, Gauteng, Pretoria, 35km NE of Pretoria Farm 25[���] 42[���] 0[���] S; 28[���] 13[���] 0[���] E, 12-13.xii.1983, leg: Davis. (1♂ SANC): SOUTH AFRICA, KZN, Weke-Weke Stream, Shongweni Farm 29[���] 48[���] S; 30[���] 43[���] E, 11.xi.2010, leg: AJ Armstrong & G Van Bassouw. Habitat. This new species has been recorded in patches of dense woodland and thicket. On the bushveld near Pretoria, it was sampled only in thickets on deep sand and not in open woodland or grassland (Davis 1996). It has been recorded only from South Africa (Fig. 4)., Published as part of Daniel, Gimo M., Davis, Adrian Lv. & Scholtz, Clarke H., 2016, Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa, pp. 67-74 in Zootaxa 4147 (1) on pages 68-69, DOI: 10.11646/zootaxa.4147.1.4, http://zenodo.org/record/266251, {"references":["Davis, A. L. V. (1996) Habitat associations in a South Africa, Summer rainfall, dung beetle community (Coleoptera: Scarabaeidae, Aphodiidae, Staphylinidae, Histeridae, Hydrophilidae), Pedologia, 40, 260 - 280."]}