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

Strümpher, Werner P., Daniel, Gimo M., Davis, Adrian L.V. (2022): Woody plant communities of southern South Africa and new distribution records for the rare dung beetle species Sarophorus punctatus Frolov & Scholtz, 2003 (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 5188 (4): 396-400, DOI: https://doi.org/10.11646/zootaxa.5188.4.7

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

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

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

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

Genus Silvaphilus Roets & Oberlander, 2010 Type species: Silvaphilus oubosiensis Roets & Oberlander, 2010, by original designation. Silvaphilus Roets & Oberlander, 2010: 369; Davis et al. 2019: 139; Davis et al. 2020: 159., Published as part of Daniel, Gimo M., Str��mpher, Werner P. & Deschodt, Christian M., 2022, An unexpected new flightless dung beetle species (Coleoptera: Scarabaeidae: Scarabaeinae: Endroedyolini) from the Cederberg Mountains, South Africa, pp. 414-420 in Zootaxa 5093 (4) on page 415, DOI: 10.11646/zootaxa.5093.4.2, http://zenodo.org/record/5912678, {"references":["Roets, F. & Oberlander, K. C. (2010) Silvaphilus: A new relict forest-endemic Canthonini dung beetle genus from the Western Cape Province of South Africa (Coleoptera: Scarabaeidae: Scarabaeinae). African Entomology, 18, 369 - 373. https: // doi. org / 10.4001 / 003.018.0213","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2019) Defining new dung beetle tribes to resolve discrepancies between phylogeny and tribal classification in the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Zootaxa, 4608 (1), 131 - 144. https: // doi. org / 10.11646 / zootaxa. 4608.1.7","Davis, A. L. V., Deschodt, C. M. & Scholtz, C. H. (2020) Conservation assessment of Scarabaeine dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological notes. Suricata 6. South African National Biodiversity Institute, Pretoria, 800 pp."]}

Silvaphilus joselmae Daniel, Str��mpher & Deschodt, new species (Figs 1A���F, 2���3) Nomenclatural act: LSIDurn:lsid:zoobank.org:act: F290EEB4-901D-4A68-AD23-6F35EF90E7AC Type locality. 4 km SE of Algeria Forest Station (32��24���30���S 19��05���03���E), Cederberg Wilderness Area, Western Cape Province, South Africa (Figs 2���3). Type material. HOLOTYPE, ♂ (TMSA) (Figs 1A���F): ��� SOUTH AFRICA: Western Cape Prov., | Cederberg Wilderness Area, | 4 km SE of Algeria Forest Station,��� || ��� 7-9.viii.2021, altitude 790 m, | 32��24���30���S 19��05���03���E, | GM Daniel & WP Str��mpher��� || " SITE IV: 790 m, Shaded vegetation, | near stream, pitfall trap | baited with pig dung.��� || ��� HOLOTYPE | Silvaphilus joselmae | Daniel, Str��mpher & | Deschodt, 2021 [red cardstock]���. PARATYPES, 10♂♂, 18♀♀ (BMSA 4♂ 6♀; TMSA 5♂ 9♀; SANC 1♂; 3♀♀); same collection data as holotype. Description of holotype ( ♂). Size: Length: 3.4 mm. Width: 1.7 mm. Colour (Figs 1A���B): Black; mouthparts and antenna yellowish. Body surface covered with recurved tan setae. Body shape (Fig. 1A): Round (in dorsal view) and strongly convex (in lateral view). Head (Figs 1A���B): Rounded, surface densely and coarsely punctate with few scattered tan setae. Clypeus broadly rounded, somewhat depressed anterolaterally. Anterior clypeal margin bidentate; edge between denticles broadly W-shaped with small triangular medial denticle present; lateral clypeal margin continuous; fronto-clypeal suture indistinct. Clypeo-genal margin continuously rounded. Dorsal part of eyes elongated and curved. Antennae with nine antennomeres; mouthparts with few setae. Pronotum (Fig. 1A): Strongly convex, wider than long, coarsely punctate, some punctures bearing recurved tan setae; basal margin with single clear line of uniform elliptical punctures arranged in parallel; basal margin wider than apical; lateral margin somewhat explanate. Elytra (Figs 1A, D): Strongly convex, fused, with nine striae. Striae 1���7 formed by continuous row of double semi-circles; stria 8 with a carina on either side, modified as deep and obvious groove; stria 9 adjacent to epipleuron. Epipleuron surface smooth, well-defined, wider basally and narrower apically. Elytral interstriae with single row of recurved tan setae paralleling outer edge of each interval; seta spaced every four or five punctures. Medial edges impunctate; interstriae surface densely punctate on first basal half, while on second apical half punctate only on lateral edges. Elytral humerus and apical callosity absent. Metathoracic wings absent. Prosternum and propleuron (Fig. 1B): Surface dull, with few irregular sets of ocellate punctures. Ventrites (Fig. 1B): Mesepimere with few punctures; surface dull. Mesoventrite with surface shining; virtually glabrous; punctures almost touching and about twice larger than those on the metaventrite. Mesometaventral suture very slightly curving anteriorly. Metaventrite surface shining; surface completely punctate with punctures separated by almost one puncture diameter, glabrous anteriorly, setose posteriorly. Six abdominal ventrites visible, punctate laterally, last abdominal ventrite twice wider than others with single row of recurved setae. Pygidium (Fig. 1C): About twice as wide as high, surface shagreened with punctures, lateral edges with few punctures, each bearing tan seta. Complete and strongly marginate, marginal bead obtusely angulate at basal midline. Legs (Figs 1A���B): Profemora ventrally punctate, with some punctures bearing seta, surface micro-shagreened, shining; upper anterior and posterior edges strongly marginate; lower anterior edge weakly marginate; distinct pit on trochanterofemoral segments present. Protibiae, slender basally and stouter apically, truncate apically with three prominent teeth on outer margin; entire margin distinctly serrated, area between teeth serrate; apical tooth largest, medial tooth somewhat smaller than basal one. In dorsal view, a row of six-aligned setae running parallel to outer margin visible; outer margin carina present, extending from basal edge of tibia to basal tooth. In ventral view, median row of seven-aligned setae running in between inner and outer margin; shallowly carinate on basal half of outer margin. Protibial spur present. Protarsi with four visible tarsomeres and tiny pair of claws; protarsomeres 1���3 triangular and short; protarsomere 4 cylindrical at least twice longer than others, with pair of tiny apical claws. Meso- and metafemora ventrally sparsely, finely punctate hind edges, punctures bearing setae, surfaces microreticulate, shiny; anterior and posterior edges marginate. Meso- and metatibiae flattened, gradually expanded nearly to apex, with prominent spur on inner margin. Meso- and metatarsomeres with five tarsomeres, protarsomeres 1���4 triangular, short, covered with few stiff setae; protarsomere 5 cylindrical at least twice longer than others, with pair of very tiny apical claws. Aedeagus (Fig. 1E): Symmetrical, parameres with the apex tapered and inwardly curving. Morphological variation: Length: 3.3���3.8 mm. Width: 1.6���1. 9 mm. There is no clear sexual dimorphism, but females are slightly larger than males. Diagnosis: Silvaphilus joselmae new species can be distinguished from S. oubosiensis by the following combination of characters: males of the new species lack the lateral elytral spine which is present in males of S. oubosiensis. Silvaphilus oubosiensis shows a pronotal depression on the basal-medial edge, which is not present in the new species. Silvaphilus joselmae new species is much smaller in size than S. oubosiensis; in the former the medial width of its mesoventrite is twice wider and with two rows of punctures, in the latter the mesoventrite is narrower and with one row of punctures. Furthermore, the mesometaventral suture curves very slightly anteriorly in the new species, whereas it is rounded in S. oubosiensis. Lastly, the eighth stria in S. joselmae new species is modified to form a deep and obvious groove, but in S. oubosiensis it is formed of deep and clear elongated and connected punctures. Etymology: The new species is named after Joselma G. Pereira, the wife of the first author, Gimo M. Daniel, for her unconditional love and support in his entomological career. Distribution (Figs 2���3): Silvaphilus joselmae new species is known from a single locality in the Cederberg Mountains where it was recorded in Fynbos riparian vegetation (Aza 1; Mucina & Rutherford 2006) (Fig. 3). In the Cederberg area there are less than ten severely fragmented patches of the Fynbos riparian vegetation covering a total area of less than 50 km 2. Although the vegetation unit is well protected in the Cederberg Wilderness Area dedicated collecting in the area is required to determine if this species occurs in any of the other patches of riparian scrub vegetation or if it exhibits broader vegetation specialisation., Published as part of Daniel, Gimo M., Str��mpher, Werner P. & Deschodt, Christian M., 2022, An unexpected new flightless dung beetle species (Coleoptera: Scarabaeidae: Scarabaeinae: Endroedyolini) from the Cederberg Mountains, South Africa, pp. 414-420 in Zootaxa 5093 (4) on pages 415-417, DOI: 10.11646/zootaxa.5093.4.2, http://zenodo.org/record/5912678, {"references":["Mucina, L. & Rutherford, M. C. (Eds.) (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzi. Vol. 19. South African National Biodiversity Institute, Pretoria, 807 pp."]}

Identification key to Endroedyolini, adapted and modified from Deschodt et al. (2020). 1. Metaventrite anteriorly with deep fovea on each side of midline, pronotal disc on each side near posterior angle with short elevated carina................................................ Endroedyolus paradoxus Scholtz & Howden, 1987 - Metaventrite anteriorly without deep fovea on each side of midline, and pronotal disc with or without short elevated carina on each side near posterior angle........................................................................... 2 2. Pronotal disc with short elevated carina on each side near posterior angle................................................................................................... Aliuscanthoniola similaris Deschodt & Scholtz, 2008 - Pronotal disc without short, elevated carina on each side near posterior angle...................................... 3 3. Mesoventrite medially wide, or wide enough for, more than three irregular rows of punctures......................... 4 - Mesoventrite medially narrow, with at most two irregular rows of punctures (punctures sometimes absent in this area)..... 5 4. Mesometaventral suture angulate at anterior apex.................. Outenikwanus tomentosus Scholtz & Howden, 1987 - Mesometaventral suture rounded at anterior apex...................... Upsa centennial Deschodt, Sole & Scholtz, 2020 5. Elytron in lateral view with depression on lateral side......................................................... 6 - Elytron in lateral view without depression on posterolateral side................................................. 7 6. Pronotal depression on mid-basal edge present, mesometaventral suture rounded; lateral elytral spine in males present.................................................................. Silvaphilus oubosiensis Roets & Oberlander 2010 - Pronotal depression on mid-basal edge absent, mesometaventral suture curving very slightly anteriorly; lateral elytral spine in males absent....................................... Silvaphilus joselmae Daniel, Strümpher & Deschodt new species 7. Mesometaventral suture rounded at anterior apex............................................................ 8 - Mesometaventral suture angulate at anterior apex........................................................... 10 8. Protibial teeth 2 and 3 small, almost half the size of protibial tooth 1........... Peckolus parvus Scholtz & Howden, 1987 - Protibial teeth 2 and 3 minute, much less than half the size of protibial tooth 1..................................... 9 9. The recurved setae on the body almost half the length of the straight setae...... Peckolus alpinus Howden & Scholtz, 1988 - The recurved setae on the body barely discernible..................... Peckolus poenskopius Deschodt & Scholtz, 2008 10. Lateral part of mesometaventral suture (between mesocoxae) converging anteriorly........................................................................................... Parvuhowdenius harrisoni Deschodt & Scholtz, 2008 - Lateral part of mesometaventral suture (between mesocoxae) parallel........................................... 11 11. Punctures on mesoventrite large and clear, separated by less than one puncture diameter.......................................................................................... Nebulasilvius insularis Deschodt & Scholtz, 2008 - Punctures on mesoventrite small and separated by at least one puncture diameter.................................................................................................. Nebulasilvius johani Deschodt & Scholtz, 2008

Daniel, Gimo M., Strümpher, Werner P., Deschodt, Christian M. (2022): An unexpected new flightless dung beetle species (Coleoptera: Scarabaeidae: Scarabaeinae: Endroedyolini) from the Cederberg Mountains, South Africa. Zootaxa 5093 (4): 414-420, DOI: https://doi.org/10.11646/zootaxa.5093.4.2

Daniel, Gimo M., Strümpher, Werner P., Snäll, Stanislav (2022): ERRATUM A new species and new distributional records of Haroldius Boucomont, 1914 (Coleoptera: Scarabaeidae: Scarabaeinae) from southern Africa. Zootaxa 5091 (4): 600-600, DOI: https://doi.org/10.11646/zootaxa.5091.4.9

Haroldius lyleae Daniel, Str��mpher & Sn��ll, new species (Figs 1���13, 15) Type locality: Soetvlakte Farm [26��45���S 22��50���E], Northern Cape Province, South Africa (Fig. 15). Type material. HOLOTYPE (♂, TMSA, aedeagus extracted) (Figs 1���9): ��� S.Af. [South Africa]; Northern Cape [Province] | Soetvlakte Farm, 1027m | 26[��].45[�����]S ��� 22[��].50[���]E��� || ���17-19.2[February].2010; E-Y: 3873 | pitfall;Vaalbos in grassl [grassland] | leg. Robin Lyle ��� || ��� HOLOTYPE | Haroldius lyleae | Daniel, Str��mpher & | Sn��ll 2021 (red cardstock)���. PARATYPE (female, ESRC; https://www.inaturalist.org/observations/20160703) (Figs 10���13): ��� RSA: Gauteng pr [Province] | Springs 2.IX.2006 | 26��14���30���S, 28��30���22���E | Stanislav Sn��ll leg.��� || ��� PARATYPE | Haroldius lyleae | Daniel, Str��mpher & | Sn��ll, 2021 (red cardstock)���., Published as part of Daniel, Gimo M., Str��mpher, Werner P. & Sn��ll, Stanislav, 2021, A new species and new distributional records of Haroldius Boucomont, 1914 (Coleoptera: Scarabaeidae: Scarabaeinae) from southern Africa, pp. 34-42 in Zootaxa 5072 (1) on page 35, DOI: 10.11646/zootaxa.5072.1.4, http://zenodo.org/record/5728961

Genus Haroldius Boucomont, 1914 Haroldius Boucomont, 1914: 253. Type species: Haroldius rugatulus Boucomont, 1914, by subsequent designation (Arrow 1931: 413). = Cyclotrogus Wasmann, 1918: 2. Type species: Cyclotrogus heimi Wasmann, 1918, by original designation. Synonymised by Arrow (1931: 413). = Ponerotrogus Silvestri, 1924: 583. Type species: Ponerotrogus annandalei Silvestri, 1924, by original designation. Synonymised by Paulian (1985: 197). = Afroharoldius Janssens, 1949: 183. Type species: Afroharoldius ennearthrus Janssens, 1949, by original designation. Synonymised by Paulian (1985: 196). = Formicdubius Philips & Scholtz, 2000: 227. Type species: Formicdubius convexus Philips & Scholtz, 2000, by original designation. Synonymised by Krell & Philips (2010: 44)., Published as part of Daniel, Gimo M., Str��mpher, Werner P. & Sn��ll, Stanislav, 2021, A new species and new distributional records of Haroldius Boucomont, 1914 (Coleoptera: Scarabaeidae: Scarabaeinae) from southern Africa, pp. 34-42 in Zootaxa 5072 (1) on page 35, DOI: 10.11646/zootaxa.5072.1.4, http://zenodo.org/record/5728961, {"references":["Arrow, G. J. (1931) The Fauna of British India including Ceylon and Burma, Coleoptera, Lamellicornia. Part III. (Coprinae).","Wasmann, E. (1918) Myrmecophile und termitophile Coleopteren aus Ostindien, hauptsachlich gesammelt von P. J. Assmuth S. J. II. Scarabaeidae. Wiener Entomologische Zeitung, 37, 1 - 33.","Silvestri, F. (1924) Description of a new genus of myrmecophilous Scarabaeidae (Coleoptera) of India. Records of the Indian Museum, 26, 583 - 586.","Paulian, R. (1985) Notes sur les coleopteres Scarabaeidae du Museum de Geneve. II. Revue Suisse de Zoologie, 92, 189 - 193.","Philips, T. K & Scholtz, C. H. (2000) A new genus and species of trichome-bearing dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology, 8, 227 - 231.","Krell, F-T. & Philips, T. K (2010) Formicdubius Philips & Scholtz from South Africa,"]}

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

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

Stiptopodius savuti Daniel & Deschodt, new species (Figs. 1B, 4) Type locality: Savuti, North West, Botswana. Type material. Holotype, female: Savuti [Botswana, North West], Site 4 | S18.53092 [��]; E24.08378 [��] | 15.xii. 2005 | Deschodt & Tshikae | Carrion pitfall trap || HOLOTYPE: Stiptopodius | savuti Daniel & Deschodt | 2020 [printed on the red card] (TMSA). Description. Holotype, female. Length: 4.3 mm, width: 1.9 mm. Colour. Dorsally black; lateral edge of clypeus, legs, and last abdominal ventrite brown; setae yellowish. Head. Incision between medial clypeal teeth deeply concave; latero-clypeal edges not sinuate; genae with external protrusion sharply triangular; surface of the head smooth and punctate; ocellate punctures sparse on the middle of vertex; some large punctures with outer edges moderately raised on clypeus, otherwise, finely punctate; clypeo-frontal carina not visible; clypeal-genal suture present; vertex with carina occupying half the space between eyes, carina interrupted in the middle clearly forming two small cephalic tubercles. Antennae with eight antennomeres, labium setose. Pronotum. Convex, wider than long; base of pronotum marginate; anterior margin wider than posterior edge; in lateral view, lateral edge strongly sinuate before postero-lateral angles, central disc glabrous; punctation ocellate with sparse, short setae antero-laterally that emanate from centres of punctures. Elytra. Laterally straight, somewhat convex distally and concave proximally; sparsely and finely punctate in external-most four striae, the rest apunctate; interstriae with sparse, short setae visible basally, only interstriae V���VIII punctate. Pygidium. Convex with dense ocellate punctation bearing setae emanating from centres of punctures. Abdominal ventrites. Prosternum with anterior angles slanted forward and inwards, surface somewhat excavated. Mesoventrite dull, setose and punctate. Metaventrite shiny with long setae anteriorly, glabrous mid-posteriorly. Abdominal ventrites glabrous medially, shiny laterally, setigerous with ocellate punctures, setae arranged in lateral rows; last ventrites wider medially than others (female character). Legs. External and internal margins of profemora punctate, with well-developed setae, central area apunctate and glabrous; external and internal edges strongly and slightly marginate, respectively. Meso- and metafemora with external and internal edges sparsely punctate, all punctures bearing setae (shorter than those in profemora), surface between external and internal edges glabrous, strongly marginate internally. Protibiae crenulate with four teeth on external margin, a line of setae around teeth in dorsal view; in ventral view, protibiae with two carinae, internally punctate and pilose. Protibial spur long and inward curving. Meso- and metatibiae triangular and strongly widened apically. Protarsi broken off in both legs. Meso- and metatarsomeres I���IV triangular, widened apically, wider than long, strongly concave medially, external and internal edges with distal lobe; meso- and metatarsomere V rectangular and longer than others, with pair of very tiny apical claws. Male. Unknown. Distribution. This species is only known from the type locality Savuti in Botswana. Etymology. The specific name, a noun in apposition, refers to the name of the type locality. Diagnosis. Stiptopodius savuti new species is morphologically similar to S. granulosus. However, S. savuti new species can be distinguished by having the dorsal cephalic and pronotal surface smooth while in S. granulosus they are densely rugose. The new species bears a carina which is interrupted medially and clearly forms two small cephalic tubercles, unlike S. granulosus, whose cephalic carina does not form tubercles. Furthermore, the head of the new species is glabrous, whereas in S. granulosus it is setigerous. Remarks: The Afrotropical Stiptopodius savuti new species seems to be similar to the brown Somalian species of Stiptopodius glabricollis M��ller, 1942. Our knowledge on Afrotropical dung beetle biogeographical distribution patterns suggests that it is very unlikely that the inland species from Botswana could be a sub-population of the coastal eastern horn of Africa species, S. glabricollis (see also a similar case in G��nier & Josso 2017: 338). As such a wide and disjunct distribution is very rare among Afrotropical Scarabaeinae, we therefore consider the specimen from Savuti (Botswana) as a new species. Stiptopodius granulosus (from Zambia) is morphologically similar to the new species. Since both species occur in neighbour countries, there is a huge potential of misidentification between them, that���s why the diagnosis of the new species is compared to the closest species in southern Africa. Conservation status and potential threats. This species was recorded from the well-protected Savuti area of Chobe National Park in northern Botswana. The vegetation consists of somewhat open to dense stands of Philenoptera nelsii (Schinz) Schrire. The trees are about one to two meters tall with about 30% canopy cover (CMD, personal observation). Although it is protected at Savuti, the extent of occurrence of this species is not known and it should thus be assessed as Data Deficient., Published as part of Daniel, Gimo M., Deschodt, Christian M., Davis, Adrian L. V. & Sole, Catherine L., 2020, The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa, pp. 394-404 in Zootaxa 4763 (3) on pages 399-400, DOI: 10.11646/zootaxa.4763.3.4, http://zenodo.org/record/3759527, {"references":["Genier, F. & Josso, J. F. (2016) Notes on Afrotropical Catharsius species described by Edgar von Harold (Coleoptera: Scarabaeidae, Scarabaeinae). Zootaxa, 4072 (3), 333 - 342. https: // doi. org / 10.11646 / zootaxa. 4072.3.3"]}

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

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

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."]}