Sisyphini Mulsant 1842

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 ¾ of the metafemur length in major males and ½ 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 (<¼ the length of the metafemur) and longer in major males (½ the length of the metafemur), elytral setae yellow. Moist highveld and coastal upland grassland of South Africa............................................... 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 ½ 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, ¼ of the metafemur length........................................................... (9)

– Spine absent on posterior margin of the metafemur; metatrochanter is ½ the length of the metafemur in major males. Shaded vegetation and thickets from east to southern Africa........ Neosisyphus fortuitus (Péringuey, 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 ½ 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 (<10); larger-bodied species. Highland grassland under cooler conditions in South Africa, Botswana, and Namibia....................................................................................................................... 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 Péringuey, 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)