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Phoberus villeti Strümpher, new species (Figs. 1–5) Type locality. “ SOUth AfrIca, Eastern Cape PrOvInce, HOgsback [AmathOle DIstrIct], 32°35′51.15″S 26°57′31.87″E ”. Type material examined. South Africa, Eastern Cape Province. Holotype ♀ (ALBM, specImen cleaned) (Figs. 1–4): “ SOUTH AFRICA: Eastern Cape | PrOv., HOgsback, 02.v.2010, | 32°35′51.15″S 26°57′31.87″E | leg. G. van Heerden ” || “HOLOTYPE | Phoberus villeti | det. W.P. Strümpher 2023 ”. Description of holotype ( ♀ ). Size. Length: 11.9 mm. Width: 5.0 mm. Body shape. Strongly convex, oval; elytral profile convex, maximum height in the middle. Colour (Figs. 1–3). Matt black. Surface of head, parts of pronotum, and ventral aspects with light brown tomentose coating. Body setae mostly yellowish brown to brown. Head (Figs. 1–3). Clypeus obtusely triangular, disc slightly elevated, feebly convex, tubercles or ridges absent, apex of clypeus with apical portion downward and rounded, margin slightly reflexed, entire; surface coarsely sculpted with discrete round to irregularly shaped punctures; punctures close, large, each with a short stout seta, surface between punctures tomentose. Clypeogenal and frontoclypeal sutures distinctly raised; frontoclypeal suture complete, with a low indistinct clypeogenal tubercle on either side, tubercles punctate. Frons raised, with paired, setose carinae, separated by a distinct, raised median ridge, setose carinae on frons marked by row of long, erect, stout seta; surface coarsely sculpted with distinct round punctures. Genal angles obtuse; genae shape rounded, narrow; genal surface depressed with discrete and irregularly shaped punctures; genal margin with distinct row of long erect setae. Vertex densely and evenly covered with round punctures, punctures placed close together, central vertex distinctly elevated, marked by two central tubercles, tubercles fused forming a broad and almost V-shaped ridge. Eyes reduced, barely visible in dorsal view.Antennal scape rounded, longer than wide, dorsally punctate, with long straight brown setae; pedicel attached apically; antennal club dark brown, velutinous. Mouthparts with dense, brown setae. Labrum shape asymmetric with a transversal carina parallel to the posterior margin; surface of labrum punctate, punctures evenly spaced, each puncture bearing a long stout seta. Pronotum (Figs. 1–2). Strongly attenuated anteriorly, pronotal length almost half of elytral length, pronotal width only slightly narrower than elytron width. Sides of pronotum broad and flat, lateral margin smooth, tomentose with fringes of stout setae, anterior angle acute, posterior angles obtuse; base of pronotum obliquely truncate laterally behind posterior angles, base evenly rounded, marginate, margin setose, weakly notch medially (at scutellum). Pronotal surface with discrete, round punctures, uniformly spaced, rather close, space between punctures less than diameter of punctures, exception on apex where punctures are more widely spaced, surfaces between punctures dull with superficial fine microreticulation present between punctures. Median discal area of pronotum distinctly raised, broad, stretching over length of pronotum, somewhat flattened anteriorly and strongly declivous on sides; median pronotal ridges distinct, extending almost to pronotal base with deep depression between, partly obstructed in the middle by partial transverse ridge; with another distinct ridge present laterad of and almost parallel to the anterior half of the median discal ridges, with shallow depression between; laterad of median pronotal ridges in posterior half a single broadly-flattened tubercle, partly fused with discal ridges; median basal tubercles distinct, slightly laterally offset, not fused to discal ridges; distinct antero-lateral tubercle and lateral basal tubercle present on either side, basal one larger and elongate, apical one smaller and round; distinct lateral carina present below lateral tubercles; all tubercles and ridges setose. Scutellum (Fig. 1). Small, narrow, longer than wide. Elytra (Figs. 1–2). Elytral shape ovate; elytral length 2.1 of length of pronotum; surface matt, feebly shiny, finely alutaceous. Humeral angles rounded, calli absent, elytral sides flattened, lateral margins smooth to weakly undulate with regularly spaced tufts of setae along elytral length, margin laterally and ventrally tomentose; basal margin slightly concave with row of long setae. Sutural margin slightly raised, margin with tufts of setae; tufts roughly of similar size, separated, evenly distributed along margin. Elytral costae distinct, even-numbered costae more prominent than odd-numbered. Even-numbered costae with low oval to round tubercles (especially on intervals 2, 4 and 6); tubercles near base coalesced into an indistinct ridge, thereafter separated, distributed along elytral length, each with a large tuft of setae, tufts roughly of similar size. Odd-numbered costae marked by row of short setae; setae separate, irregularly spaced along elytral length. Intercostae marked by row of large round punctures; punctures deep, of similar size, regularly spaced over length of elytra. Metathoracic wings. Completely reduce, only wing rudiments visible. Legs (Figs. 1–3). Protibiae straight, slender, with single prominent tooth on outer margin, ending with apical comb of short straight setae, apex narrow, semi flattened, squarely rounded; protibial spur stout, pointed forward, nearly as long as four basal tarsomeres combined; tarsal segments smooth; tarsomeres 1–4 subequal in length, tarsomere 5 as long as tarsomeres 2–4 combined; tarsomeres 1 and 2 without setae, tarsomere 3 ventrally with single fine seta, tarsomeres 4 and 5 ventrally each with single long stout seta; tarsomere 5 with two claws, simple, equal, small apicoventral process present on tarsomere below claws. Protibia dorsally with apical half marked by a series of irregular circular punctures, thereafter, modified into groove or series deep connected punctures; each puncture with a short erect seta; inner top edge of protibia marginate. Outer surface of protibia rugose with row of large round irregular punctures, some bearing single stout seta, outer margin distad of protibial tooth smooth, without indentations, margin proximad of tooth round, edge marked by row of straight stout setae. Inner protibial surface smooth, glabrous, proximad third with round punctures, sparsely setose; distad two thirds with rows of long setae. Protibia ventrally with double row of stout setae running closely parallel to outer edge, surface distad of lateral tooth rugose, surface proximad of lateral tooth semi-flattened, mostly smooth, glabrous. Profemur robust, widest basally, marginate on front face with row of straight forward pointing setae; ventral face with regularly spaced round punctures, each with single seta, surface in parts indumentose; dorsal surfaces on front face sparsely punctate, mostly smooth, glabrous, remainder of dorsal surface with long dense straight thin setae. Procoxa narrow, elongate, marginate on front face; surface rugose, punctuate, some punctures with stout seta, inner half indumetose. Protochanter indumentose, with small cluster of long erect setae. Mesotibial and metatibial surfaces smooth, feebly covered with fine indument; superomedial longitudinal carinae with row of stout setae; inner and outer carinae with row of long slender setae; outer margin on apex of meso- and metatibia with row of closely spaced stout backward pointing setae; meso- and metatibia with two apical spurs on inner margin; upper spur slightly longer than tarsomere 2, lower spur shorter than tarsomere 1. Mesotibia with three distinct separate teeth on lateral margin. Mesotarsi and metatarsi smooth, with long setae, tarsomeres 1–4 subequal in length, tarsomere 5 as long as tarsomers 3 and 4 combined; distal tarsomere with two claws simple, equal. Ventral aspects (Fig. 3). Proepisterna: Surface smooth, glabrous, with long thin setae, anterior margin distinctly raised, rounded, margin with row of long fine setae; prosternal apophysis broadly triangular; disc with short medial longitudinal ridge. Epipleura: Epipleura very broad basally, becoming slightly narrower distally, slightly concave along entire length, surface mostly smooth, feebly tomentose, weakly undulating, sparsely setose, setae fine; inner margin with thick tomentosity. Mesoventrum: Sunken below metaventrum; disc raised anteriorly, sunken posteriorly; mesoventrum apophysis prominent between mesocoxae, narrow, raised, extending almost the length of mesocoxa. Surfaces of mesoventrum rugose and coarsely punctate, punctures close, thickly indumentose, round to oval. Metaventrum: Metaventrum strongly convex, shortened, 2.8 times as wide as long, surface rugose and coarsely punctate, setose; lateral and anterior margin with tomentosity; plate sunken along mesocoxal margin forming narrow basinlike depression; metaventral disc rhomboidal, visibly depressed medially, with costate edges that converge outwards to distinct transverse carinae on either side of median depression, carinae setose, with covering of indument; plate strongly declivous behind without covering of indument. Abdominal ventrites: five visible ventrites (4–8); ventrite 4 with rugose surface, coarsely punctate, sparsely setose; ventrite 5 rugose in apical half, relatively smooth on distal half, medially with single transverse row of short stout setae; distal margins of ventrite 5 subparallel with apical margin of ventrite 6; ventrite 6 with band of coarse punctures along apical margin, remainder of surface smooth; distal third marked by transverse row of short stout setae arranged parallel to distal margin; distal margin concave; ventrite 7 only feebly punctate, surface mostly smooth, transverse row of short stout setae along distal margin, distal margin concave, curvature subparallel with distal margin of ventrite 6; ventrite 8 with distal margin convex, surface smooth, distal half with with short, evenly spaced setae. Male. Unknown. Differential diagnosis. Phoberus villeti Strümpher, new species, is quite different from all previously described flightless, and also alate, species of Phoberus and bear only superficial resemblance to the following species, P. aculeatus (Harold, 1872) (Figs. 6–7) and P. horridus (Fabricius, 1775) (Figs. 8–9). Phoberus villeti Strümpher, new species, can be easily distinguished at a glance from both by the presence of the very distinctive tufts of yellow setae present on the even-numbered costae and lateral margin of the elytra (Figs. 1–3). In contrast, P. aculeatus and P. horridus have straight, and almost spine-like black setae on the even-numbered costae and lateral margin, giving them a prickly appearance (Figs. 6, 8). Comments. I am well aware of the undesirability of basing a description on a single female specimen, especially in Phoberus, where male genitalia are important for species recognition since shape and associated aedeagal characters, with few exceptions (Strümpher & Scholtz 2017, 2019), are usually unique to each species. However, in this case, the description of the new species can be justified for the following reasons. First, the new species is sufficiently morphologically diagnosable and can hardly be confused with any of the other species in the genus. Second, due to the limited amount of sexual dimorphism in Phoberus descriptions and diagnoses of species largely applies equally to males and females. Lastly, flightless species of Phoberus are rare in collections, and in nature. They have restricted distributions and because of their elusive nature are difficult to collect—thus, it is not surprising that this new species has gone unnoticed, till now—and many more flightless species await discovery in South Africa. The species is described herein to bring it to the attention of coleopterists working in the region with the hope that additional material including males will be discovered in other localities. Distribution. Known only from the type locality (Fig. 5). Etymology. The specific epithet honours Martin H. Villet, professor of Entomology at Rhodes University (Makhanda, South Africa), the author’s doctorate supervisor and mentor. It is formed in the genitive singular case., Published as part of Strümpher, Werner P., 2023, A new flightless species of Phoberus Macleay, 1819 (Coleoptera: Trogidae) from the Eastern Cape Province in South Africa, pp. 143-150 in Zootaxa 5264 (1) on pages 144-146, DOI: 10.11646/zootaxa.5264.1.11, http://zenodo.org/record/7836072, {"references":["Strumpher, W. P. & Kalawate, A. S. (2023) Omorgus (Omorgus) khandesh (Coleoptera: Scarabaeoidea: Trogidae), a new species from India, along with an annotated catalogue of the Oriental and Palearctic species of the subgenus Omorgus Erichson, 1847. Zootaxa, 5231 (5), 501 - 522. https: // doi. org / 10.11646 / zootaxa. 5231.5.1","Strumpher, W. P. & Scholtz, C. H. (2017) A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group. Zootaxa, 4216 (3), 292 - 298. https: // doi. org / 10.11646 / zootaxa. 4216.3.6","Strumpher, W. P. & Scholtz, C. H. (2019) A new Phoberus MacLeay species (Coleoptera: Scarabaeoidea: Trogidae) from the Maloti-Drakensberg Mountains of southern Africa and the Phoberus caffer species group reviewed. Zootaxa, 4674 (5), 551 - 563. https: // doi. org / 10.11646 / zootaxa. 4674.5.4"]}

Phoberus MacLeay, 1819 Phoberus MacLeay, 1819: 137���138 (original description); Str��mpher et al. 2014: 557���558 (phylogenetics); Str��mpher et al. 2016b (reinstated as genus); Z��dek 2017: 96���113 (checklist). A bibliography of Phoberus was provided by Str��mpher et al. (2016b: 79���80). Type species: Trox horridus Fabricius, 1775 (by monotypy)., Published as part of Str��mpher, Werner P. & Stals, Riaan, 2021, Proper cleaning of keratin beetles reveals a cryptic species: Phoberus fumarius (Haaf, 1953) from southern Africa is reinstated as valid (Coleoptera: Trogidae), pp. 543-557 in Zootaxa 4942 (4) on page 545, DOI: 10.11646/zootaxa.4942.4.3, http://zenodo.org/record/4612451, {"references":["MacLeay, W. S. (1819) Horae entomologicae: or essays on the annulose animals. Volume I. Part I. Containing general observations on the geography, manners, and natural affinities of the insects which compose the genus Scarabaeus of Linnaeus; to which are added a few incidental remarks on the genera Lucanus and Hister of the same author. With an appendix and plates. S. Bagster, London, xxx + [2] + 160 pp., 3 pls. https: // doi. org / 10.5962 / bhl. title. 48636","Strumpher, W. P., Sole, C. L., Villet, M. H. & Scholtz, C. H. (2014) Phylogeny of the family Trogidae (Coleoptera: Scarabaeoidea) inferred from mitochondrial and nuclear ribosomal DNA sequence data. Systematic Entomology, 39 (3), 548 - 562. https: // doi. org / 10.1111 / syen. 12074","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016 b) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics & Evolution, 47 (1), 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133","Zidek, J. (2017) Updated checklist and bibliography of family Trogidae (Coleoptera: Scarabaeoidea). Folia Heyrovskyana, Series A, 25 (1), 93 - 127.","Fabricius, J. C. (1775) Systema entomologiae, sistens insectorvm classes, ordines, genera, species, adiectis synonymis, locis, descriptionibvs, observationibvs. Kortii, Flensburg and Leipzig, [32] + 832 pp. https: // doi. org / 10.5962 / bhl. title. 36510"]}

Phoberus cyrtus (Haaf, 1953) (Figs 1���7, 15, 21) Trox cyrtus Haaf, 1953: 333 (original description). Trox (Trox) cyrtus: Scholtz 1980a: 66, 68, 87, 97 (redescription, illustrations, distribution); Scholtz 1982: 3 (catalogue). Trox (Phoberus) cyrtus: Zidek 2013: 9 (checklist). Phoberus cyrtus: Str��mpher et al. 2016b: 78 (new combination); Z��dek 2017: 99 (checklist). Type locality. Vall��e du Haute Orange [Valley of high Orange River] (border of Mohale���s Hoek and Quthing Districts, Lesotho) Type material examined. All type specimens were examined through detailed photographs. HOLOTYPE �� (MNHN, aedeagus extracted) (Figs 1���3): Rectangle, faded blue cardstock: ���MUSEUM PARIS | Basoutoland, Lessouto [Lesotho] | Vall��e du Ht Orange [Valley of high Orange River, 30��17���S 27��46���E], 1500 m. | (legit R. Ellenberger) | E. HAUG 1906��� || Rectangle, red cardstock: ��� HOLOTYPE ��� || Rectangle, minimally foxing white: ��� Trox | cyrtus Haaf | det. Dr.E. Haaf 1953 ��� || Rectangular curatorial label, white: ��� MNHN | EC10025 ���. PARATYPES 2���� (MNHN): Blue label identical || Rectangle, red cardstock: ��� PARATYPE ��� || Foxy white label identical || White curatorial labels: ��� MNHN | EC10027 ��� and ��� MNHN | EC10029 ���. PARATYPES 3♀♀ (MNHN): Labels identical to those of ��� paratypes, but each specimen with an additional foxy white square: ��� ♀ ��� || ��� MNHN | EC10028 ���; ��� MNHN | EC10026 ��� with red label: ��� ALLOTYPE ���; ��� MNHN | 10030��� with different determination label, white rectangle: ��� PARATYPE | Trox | cyrtus Haaf, 1953 ���. PARATYPE 1�� (MNHB): Blue label identical || Rectangle, white, preprinted red text interspersed with black handwriting: ��� PARATYPUS | Trox | cyrtus sp.n. | det. Dr.E. Haaf 1953 ���. Haaf (1953) described this species from six males and four females, all with the same label data. We studied four males and three females of the type series, including the holotype. Haaf (1953) indicated that all ten type specimens would be lodged in the MNHN, but we located one male in the MNHB. Additional material examined. SOUTH AFRICA. Eastern Cape Province. 1��� Aliwal North, [30��42���S 26��42���E], 15���25.ii.1971, Snyman & Jones (TMSA). Free State Province. 13��� Nova 667 [farm], near Ladybrand, SE 2927 Ab [29��07���S 27��22���E], 19���22.i.1976, GW Ferreira, S van Ee (BMSA: BMSA (C)-21032���21044) (Figs 4���7). 3���, 3♀ idem but 01���12.ii.1977, GW & MC Ferreira, A Strydom (BMSA: BMSA (C)-21045���21046; BMSA (C)- 21975; BMSA (C)-21978; BMSA (C)-22031; BMSA (C)-22293). 4���, 4♀ Kerkenberg, near Harrismith, 2829 Ac /d [28��30���14���S 29��06���30���E], 08���18.xi.1976, Museum staff (BMSA: BMSA (C)-21020���21027). KwaZulu-Natal Province. 1��� [uKhahlamba-Drakensberg Park World Heritage Site], Mdedelelo Forestry Area, 29��07���S 29��26���E, 13���14.xi.1981, SJ van Tonder, C Kok (SANC). 1♀ [uKhahlamba-Drakensberg Park World Heritage Site], Giant���s Castle Game Reserve, at Bannerman Hut, 29��15���S 29��26���E, 2,300 m, 24.iv.1994, J duG Harrison, R Stals (TMSA). Mpumalanga Province. 1��� Kruger National Park, Skukuza Rest Camp, 24��59���S 31��36���E, 28.i.1994, E-Y:2952, UV light trap, S Endr��dy-Younga (TMSA). 1��� Winkelhaak [mine], Ermelo district, [26��30���S 29��07���E], x.1970, L Schulze (TMSA). North West Province. 2���, 7 unsexed: Rustenburg, [25��39���S 27��14���E], 28.x.1956 (NHMB, through photographs). 7 unsexed: idem but 13.viii.1961 (NHMB, through photographs). BOTSWANA . South-East District. 1��� Otse, [25��02���S 25��44���20���E], 02.ix.1963, JA Nagle (TMSA). Redescription. Size: Length: 9���13 mm. Width across humeri: 5���8 mm. Colour (Figs 1, 4���5): Matt black, with dull reflection. Body setae yellow unless stated otherwise below. Head (Figs 4���5): Surface sculpted with discrete punctures, surfaces between punctures with dull reflection. Clypeus triangular, straight, apically deflexed and rounded; anterior clypeal margin entire, bare; clypeogenal and frontoclypeal sutures distinctly raised; frontoclypeal suture complete, with a low clypeogenal tubercle on either side, tubercles punctate. Frons raised, with paired, arched, setose frontoclypeal carinae delimiting two semicircular depressions, separated by a distinct, setose median carina. Genal angles obtuse, wide, recurved; genal margin with straight setae. Eyes large and bulbous, barely visible in dorsal view. Antennal scape rounded, longer than wide, punctate, with long light brown setae; pedicel attached apically; antennal club dark brown, velutinous. Setae around mouthparts dense, yellow to rust-brown. Pronotum (Figs 4���5, 15): Slightly narrower than elytra. Surface with discrete, round punctures, surfaces between punctures dull to shiny. Lateral margins broad and flat, somewhat explanate, entire to irregularly, coarsely dentate, strongly attenuate anteriad, with fringes of long, straight setae. Apex with row of long, recurved, evenly spaced setae. Base with dense fringe of straight setae. Median discal area only slightly raised, broad, narrowing towards base to form a distinct triangular disc; dorsal outline evenly arched in anterior view; median discal depression shallow; median basal tubercles distinct; lateral basal tubercles fused to those in front of them to form a low, broad carina; all tubercles and ridges with setal tufts. Scutellum (Figs 4���5): Longer than wide; surface smooth; apicolaterally rounded, lateral margins straight; mediobasally depressed, depression weakly sculpted to punctate. Elytra (Figs 1, 4���5): Elongate, length 2.7���2.8 pronotum length, lateral margins subparallel, sides narrow, lateral margin finely dentate and bearing long setae; widest at c. 0.6 length; profile slightly convex, attaining maximum height approximately in the middle, strongly declivous posteriorly. Humeral calli prominent, with short setae. Sutural margin raised, surface smooth, with low (sometimes flat) round to oval tubercles, widely separated and irregularly distributed along margin, roughly of similar size for 0.75 elytral length, thereafter decreasing in size until barely discernible; these tubercles smooth, dull shiny, each with a posterior setal tuft. Elytral costae distinct, evennumbered costae more prominent than odd-numbered. Even-numbered costae with raised oval tubercles widely and fairly evenly spaced along elytral length; these tubercles smooth, shiny, each with a posterior setal tuft. Apical callosity present on fourth costal interval at top of elytral declivity. Elytral costae 2 and 4 with 5���8 tubercles for 0.75 elytral length (up to apical callosity); odd-numbered costae marked by small round to oval, shiny tubercles, each with one or two short setae, these tubercles regularly spaced, roughly of equal size for 0.75 elytral length, thereafter decreasing in size until barely discernible. Dorsally costae 1���4 separated from intercostae as slightly elevated bands; laterally costae 5���9 barely discernible as bands. Intercostae with very shallow, undulating depressions separated by transverse ridges; dorsally, margins of intercostae 1���4 marked by weak narrow longitudinal ridges; intercostal carinae laterally vestigial, barely discernible or lacking. Legs (Figs 4���5): Surfaces rugose and coarsely punctate. Protibia dorsally keeled, with small apical process and row of short, stout transverse setae; outer margin with 2 distinct median teeth and 3���5 smaller basal teeth; inner margin with long, dense setae; protibial apex divided into two spines, distinct in males, females with spines partly fused; protibial spur as long as, or longer than, third tarsomere, flattened laterally, apex pointed and recurved. Mesotibia and metatibia with outer margins dentate and with long, dense yellow setae; with 2 apical spurs on inner margin, spurs as long as, or slightly longer than, second tarsomere. Tarsomeres ventrally with sparse setae. Tarsal claws simple, equal. Metathoracic wings: Complete. Male genitalia (Figs 2, 6): Symmetrical, slender; phallobase arched, slightly longer than parameres. Parameres long, extending beyond median lobe, apices pointed, long and decurved. Median lobe divided into two symmetrical parts, these narrow, subparallel, in dorsal view concave upward; apices acutely rounded; two bladelike projections and hooks visible between parameres and median lobe. Diagnosis. Phoberus cyrtus and Phoberus fumarius are very similar, but the former can be distinguished from the latter in having a less coarsely punctate pronotum (Figs 15���16). Additionally, the arrangement of the tubercles on elytral costae 2 and 4 differs between the species: Phoberus cyrtus has these tubercles widely separated, with 5���8 tubercles between the elytral base and the apical callosity, whereas these tubercles in Phoberus fumarius are close together, with 8���12 tubercles between the elytral base and the apical callosity. The male genitalia of the two species are distinctive with the most important differences found on the median lobes. Phoberus cyrtus have the lobes narrow and their apices sharper or angular, whereas in Phoberus fumarius the lobes are broad and their apices more rounded (as described in the text and as shown in Figs 2 and 6 versus Figs 9 and 13). Available information strongly suggests that the two species are allopatric and existent only in different, non-overlapping biomes, Phoberus cyrtus in the summer rainfall area of southern Africa and Phoberus fumarius in the winter rainfall area (see text and Fig. 21). Phoberus sulcatus (Thunberg, 1787) (Figs 17���20) is widespread and common throughout South Africa (Scholtz 1980a). Both Phoberus cyrtus and Phoberus fumarius have an external appearance similar to that species, and the three may be confused. In collections they have certainly been confused. Of the two species treated in this work, Phoberus fumarius is more likely to be confused in this way. Phoberus sulcatus differs from both the other species in bearing a distinct median pronotal depression (Fig 19) and having the pronotal and elytral carinae and tubercles more pronounced and higher (Figs 17, 19). The male genitalia of Phoberus sulcatus are indubitably different with the median lobe of this species divided into two broadly curved lobes (Fig 18). We found that Phoberus sulcatus can readily be distinguished from these other two species by the larger shadows cast under oblique lighting by its more prominent paramedian pronotal carinae; specimens should not be too soiled for this comparison to succeed. Geographic distribution. Phoberus cyrtus is known from South Africa, Lesotho and Botswana (Fig. 21). In South Africa the species has been recorded from the Eastern Cape, KwaZulu-Natal, Mpumalanga and North West Provinces. Broad habitat associations. As far as it is known, Phoberus cyrtus is restricted to the summer rainfall region of southern Africa, in contrast to its putative sister species Phoberus fumarius, which is confined to the winter rainfall region. Phoberus cyrtus is known from two biomes. In the south of its known distribution, the species lives in the Grassland Biome, where it is present in the Mesic Highveld Grassland���, Drakensberg Grassland��� and Dry Highveld Grassland Bioregions (biomes and bioregions following Mucina & Rutherford 2006; Dayaram et al. 2019). Towards the north and west of its range, the species occurs in the Savanna Biome (Central Bushveld��� and Lowveld Bioregions). Whereas the grassland habitats are cool and mesic, the savanna habitats are warm to hot and experience extended dry periods. This discordant distribution among two sets of habitats with significantly different bioregional climate regimes may indicate a eurytopic species., Published as part of Str��mpher, Werner P. & Stals, Riaan, 2021, Proper cleaning of keratin beetles reveals a cryptic species: Phoberus fumarius (Haaf, 1953) from southern Africa is reinstated as valid (Coleoptera: Trogidae), pp. 543-557 in Zootaxa 4942 (4) on pages 545-549, DOI: 10.11646/zootaxa.4942.4.3, http://zenodo.org/record/4612451, {"references":["Haaf, E. (1953) Die afrikanischen Arten der Gattung Trox F. (Col. Scarab.). 1. Beitrag zur Kenntnis der Subfam. Troginae. Entomologische Arbeiten aus dem Museum G. Frey, 4, 309 - 346.","Scholtz, C. H. (1980 a) Monograph of the genus Trox F. (Coleoptera: Trogidae) of subsaharan Africa. Cimbebasia Memoir, 4, 1 - 104.","Scholtz, C. H. (1982) Catalogue of world Trogidae (Coleoptera: Scarabaeoidea). Entomology Memoir, Department of Agriculture and Fisheries, Republic of South Africa, 54, 1 - 27.","Zidek, J. (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta Mundi, 314, 1 - 38.","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016 b) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics & Evolution, 47 (1), 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133","Zidek, J. (2017) Updated checklist and bibliography of family Trogidae (Coleoptera: Scarabaeoidea). Folia Heyrovskyana, Series A, 25 (1), 93 - 127.","Mucina, L. & Rutherford, M. C. (Eds.) (2006) The vegetation of South Africa, Lesotho and Swaziland. South African National Biodiversity Institute, Pretoria, Strelitzia, 19, i - viii + 1 - 807.","Dayaram, A., Harris, L. R., Grobler, B. A., Van der Merwe, S., Rebelo, A. G., Powrie, L. W., Vlok, J. H. J., Desmet, P. G., Qabaqaba, M., Hlahane, K. M. & Skowno, A. L. (2019) Vegetation map of South Africa, Lesotho and Swaziland 2018: a description of changes since 2006. Bothalia, 49 (1), a 2452, 1 - 11. https: // doi. org / 10.4102 / abc. v 49 i 1.2452"]}

Phoberus bulirschi n. sp, a new species from KwaZulu-Natal, South Africa, is described and compared to Phoberus ngomensis (Van der Merwe & Scholtz, 2005) and P. sternbergi (Van de Merwe & Scholtz, 2005), its most morphologically related species. The main diagnostic characteristics distinguishing these taxa are indicated and illustrated. The distribution map of the Phoberus species belonging to the "natalensis" group is updated., Phoberus bulirschi n. sp., nouvelle espèce du KwaZulu-Natal, Afrique du Sud, est décrite et comparée à Phoberus ngomensis (Van der Merwe & Scholtz, 2005) et P. sternbergi (Van de Merwe & Scholtz, 2005), espèces qui lui sont morphologiquement les plus proches. Les principaux caractères diagnostiques distinguant ces taxons sont indiqués et illustrés. La carte de distribution des Phoberus du groupe « natalensis » est mise à jour.

Phoberus nyassicus (Haaf) new status (Fig. 5A ���D) Trox nigrociliatus nyassicus Haaf, 1953: 330 Trox (Trox) nigrociliatus nyassicus: Scholtz 1980: 93; Scholtz 1982: 5 Trox (Phoberus) nigrociliatus nyassicus: Scholtz 1986: 361; Zidek 2013: 13 Phoberus nigrociliatus nyassicus: Str��mpher et al. 2016: 78 Diagnosis. Phoberus nyassicus is very similar to P. nyikanus in appearance, but can be distinguished from the latter by distinct, evenly spaced tubercles on the elytra; P. nyikanus has the tubercles partially fused, forming semicontinuous ridges., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2017, A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group, pp. 292-298 in Zootaxa 4216 (3) on pages 296-297, DOI: 10.5281/zenodo.231847, {"references":["Haaf, E. (1953) Die afrikanischen Arten der Gattung Trox F. (Col. Scarab.). Entomologische Arbeiten aus dem Museum Georg Frey, 4, 309 - 346.","Scholtz, C. H. (1980) Monograph of the genus Trox F. (Coleoptera: Trogidae) of Subsaharan Africa. Cimbebasia Memoir, 4, 1 - 104.","Scholtz, C. H. (1982) Catalogue of world Trogidae (Coleoptera: Scarabaeoidea). Entomology Memoir, Department of Agriculture and Fisheries, Republic of South Africa, 54, 1 - 27.","Scholtz, C. H. (1986) Phylogeny and systematics of the Trogidae (Coleoptera: Scarabaeoidea). Systematic Entomology, 11, 355 - 363.","Zidek, J. (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta Mundi, 314, 1 - 38.","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics and Evolution, 47, 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133"]}

Phoberus nyikanus Str��mpher, new species (Figs. 1A ���D, 2A) Type material. Holotype: (1♂: NCSA): MALAWI / SE1033BD / Nyika National Park / 2.iii.1987 / 2607 m. / J. & A. Londt / Nganda summit���grassveld (typeset on white rectangle). The aedeagus of the holotype is placed separately in a transparent genital vial. Dissected metathoracic wing set separately on white cardboard rectangle. Condition of Holotype: Good, except that tarsi on protibia are missing. 2 Paratypes: (1♂, 1♀: TMSA): Chapolo for[est] (handwritten) / Nyika National Park / Malawi / 2.i.1981 (handwritten) / Coll: C.O. Dudley (typeset on white rectangle). 1 Paratype: (♀: UPSA): Chilenda (handwritten) / Nyika Nat [ional] Park / Malawi / 31.xii.1980 (handwritten) / Coll: C.O. Dudley (typeset on white rectangle) // Grassland (typeset on white rectangle). 1 Paratype: (♀: UPSA): M9724 / Nyika Nat [ional] Park / Malawi / iv.1976, V. Taylor / 8000 [feet] / (handwritten on white rectangle). Specimens bear new type labels: HOLOTYPE [or PARATYPE] / Phoberus nyikanus Str��mpher, 2016 (printed on red rectangle). Diagnosis. This new species is morphologically similar to other members in the group, Phoberus nigrociliatus (Kolbe) new status (Fig. 3A ���C), P. nyansanus (Haaf) new status (Fig. 4A ���C), and P. nyassicus (Haaf) new status (Fig. 5A ���C). However, it can be reliably distinguished from other members by the presence of low, semicontinuous ridges (partially fused tubercles) on the even-numbered costae (Fig. 1A), whereas in the other three species the tubercles are distinct, evenly spaced, and not fused (Figs. 3A, 4A, 5A). Furthermore, P. nyikanus lacks the distinct longitudinal ridges marking intercostal margins, which are prominent in the other three species (Figs. 1A, 3A, 4A, 5A, 5 C). Holotype description. Colour: Black. Size: length 15.1 mm, width (across humeri) 6.9 mm. Head: Clypeus triangular, apical portion deflexed; lateral margins with pointed setae; surface of clypeus and frons punctate; frons with raised, rounded setose ridges; antennal scape black, elongate, with long, black setae; pedicel attached to apex of scape; club black, velutinous (Fig. 1A ���B). Pronotum: attenuate anteriorly, sides broad, lateral margins irregular, with fringes of dark brown setae; surface coarsely punctate; median depression distinct and except for median obstruction stretches over entire length of pronotum; anterior half of median depression shallow, posterior half deeper and broader; discal area raised; discal ridges rounded, broad anteriorly, attenuate anteriorly to short narrow ridge; median basal tubercles and lateral basal tubercles present, lateral basal tubercle partially fused to anterolateral tubercle; posterior margin with dense fringe of flat setae; pronotal ridges and tubercles with setae (Fig. 1A ���B). Scutellum: Oval, as wide as long, sunken basally (Fig. 1A). Elytra: humeral calli present, reduced; lateral margins smooth to slightly irregular; sutural margin raised, with small, round to slightly elongate tubercles with tufts of black setae; even-numbered costae distinct, tubercles partially fused to form low, continuous ridges with regular tufts of setae; odd-numbered costae with small tubercles; indistinct/vestigial longitudinal ridges mark intercostal margin, and intercostal punctures round to evenly undulate with transverse ridges between them; elytral profile convex, attaining maximum height behind middle (Fig. 1A ���B). Metathoracic wing: wing length approximately equal to length of elytron; apical region distal to apical hinge reduced in size, RP branches incomplete and lacking sclerotization (Fig. 2 A). Legs: protibia bifid, outer margin with 3 small, median teeth, inner margin with long, dense setae, tibial spur as long as, or longer than, third tarsomere; mesotibia and metatibia with long, dense, black setae, 2 apical spurs on inner margin, spurs as long as, or slightly longer, than first tarsomere; tarsomeres ventrally with sparse setae, claws simple (Fig. 1A). Male genitalia: See Fig 1 C. Variation (1 male and 3 female paratypes). Size: length 10.9���14.8 mm, width (across humeri) 5.6���6.9 mm. Legs: outer margin with 2���3 small median teeth. The external morphology of males and females is identical. Distribution. Nyika Plateau in northern Malawi (Fig. 6). Etymology. This species is named for the Nyika Plateau, where the type series was collected. Comments. Examination of the metathoracic wings of Phoberus nyikanus (Fig. 2 A) and P. nyassicus (Haaf) (Fig. 5 D) clearly shows these two species to be in the early stages of flight loss (stage 1 and 2; See Browne et al. 1993). In both species the region distal to the apical hinge is reduced in size, the wing membrane is thin, and the RP branches are incomplete and lack the typical dense sclerotization; also the anal region of the metathoracic wing shows some degree of reduction. The metathoracic wings of beetles are used for propulsion during flight (posteromotorism) and, although the flight capability of the two species is not known, we can assume that it is greatly compromised by the degree of wing reduction that likely renders the two species flightless., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2017, A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group, pp. 292-298 in Zootaxa 4216 (3) on pages 293-295, DOI: 10.5281/zenodo.231847, {"references":["Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics and Evolution, 47, 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133","Browne, D. J., Scholtz, C. H. & Kukalova-Peck, J. (1993) Phylogenetic significance of wing characters in the Trogidae (Coleoptera: Scarabaeoidea). African Entomology, 1, 195 - 206."]}

Phoberus nigrociliatus (Kolbe) new status (Fig. 3A ���C) Trox nigrociliatus Kolbe, 1904: 292; Arrow 1927: 465 Trox nigrociliatus nigrociliatus: Haaf 1953: 328 Trox (Trox) nigrociliatus nigrociliatus: Scholtz 1980: 92; Scholtz 1982: 5 Trox (Phoberus) nigrociliatus nigrociliatus: Scholtz 1986: 361; Zidek 2013: 13 Phoberus nigrociliatus nigrociliatus (Kolbe): Str��mpher et al. 2016: 78 Synonym. Trox asmaraensis Balthasar, 1939: 32; synonymy by Haaf 1953: 328; Scholtz 1980: 92, 1982: 5; Zidek 2013: 7 Diagnosis. Phoberus nigrociliatus can be easily separated from other members in the group on the basis of the pronotum. It has the pronotum smoothly rounded, whereas the other members have the pronotum ridged with large tubercles. P. nigrociliatus has only been recorded from the Ethiopian highlands., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2017, A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group, pp. 292-298 in Zootaxa 4216 (3) on page 295, DOI: 10.5281/zenodo.231847, {"references":["Kolbe, H. J. (1904) Uber einige interessante Lamellicornier und Tenebrioniden Afrikas. Berliner Entomologische Zeitschrift, 49, 282 - 302.","Arrow, G. J. (1927) A note on the coleopterous genus Trox, with description of a few new Asiatic species. Annals and Magazine of Natural History, series 9, volume 19, 465 - 468.","Haaf, E. (1953) Die afrikanischen Arten der Gattung Trox F. (Col. Scarab.). Entomologische Arbeiten aus dem Museum Georg Frey, 4, 309 - 346.","Scholtz, C. H. (1980) Monograph of the genus Trox F. (Coleoptera: Trogidae) of Subsaharan Africa. Cimbebasia Memoir, 4, 1 - 104.","Scholtz, C. H. (1982) Catalogue of world Trogidae (Coleoptera: Scarabaeoidea). Entomology Memoir, Department of Agriculture and Fisheries, Republic of South Africa, 54, 1 - 27.","Scholtz, C. H. (1986) Phylogeny and systematics of the Trogidae (Coleoptera: Scarabaeoidea). Systematic Entomology, 11, 355 - 363.","Zidek, J. (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta Mundi, 314, 1 - 38.","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics and Evolution, 47, 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133","Balthasar, V. (1939) Neue Arten der coprophagen Scarabaeiden aus dem Museo Zoologico della Universita di Firenze. Redia, 25, 1 - 36."]}

Phoberus nigrociliatus group The four species are grouped together on the basis of the general shape and structure of the male genitalia. All species have distinctive external morphology and reliable characters for distinguishing species are the shape and sculpture of the pronotum and elytra. The male genitalia are not sufficiently diagnostic for species identification without reference to other characters. We stress that species in the P. nigrociliatus group are grouped together intuitively, on the basis of overall similarity; groupings therefore do not necessarily reflect phylogenetic relationship. Diagnosis. Size: Length 9���15 mm, width 5���13 mm; Colour: Black or reddish brown. Head: Frons with paired, rounded, setose ridges. Pronotum: Variable from smoothly rounded to prominently ridged with large tubercles, median depression deep or absent; attenuate anteriorly; lateral margins irregular; fringes and tufts of yellow or black setae. Elytra: Lateral margins serrate; humeral calli prominent; sutural margin with small tubercles; costae separated from intercostae by distinct elevated bands; elytral tubercles on even-numbered costae large, shiny, and evenly spaced, or partially fused forming a distinct ridges with tufts of yellow or black setae; odd-numbered costae with small tubercles; intercostae smooth or with distinct longitudinal ridges, with undulate depressions separated by transverse ridges; elytral profile slightly convex to convex, attaining maximum height at or behind midlength. Metathoracic wing: Complete or reduced. Male genitalia: See Figs, 1C; 3B���C; 4B���C; 5B���C., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2017, A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group, pp. 292-298 in Zootaxa 4216 (3) on page 293, DOI: 10.5281/zenodo.231847

Phoberus nyansanus (Haaf) new status (Fig. 4A ���C) Trox nigrociliatus nyansanus Haaf, 1953: 330 Trox (Trox) nigrociliatus nyansanus: Scholtz 1980: 93; Scholtz 1982: 5 Trox (Phoberus) nigrociliatus nyansanus: Scholtz 1986: 361; Zidek 2013: 13 Phoberus nigrociliatus nyansanus: Str��mpher et al. 2016: 78 Diagnosis. Phoberus nyansanus can be easily separated from other members of the group by the yellow fringes and tufts of setae on the pronotum and elytra; the other members have black setae. P. nyansanus is restricted to the lake areas of east-central Africa., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2017, A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group, pp. 292-298 in Zootaxa 4216 (3) on page 295, DOI: 10.5281/zenodo.231847, {"references":["Haaf, E. (1953) Die afrikanischen Arten der Gattung Trox F. (Col. Scarab.). Entomologische Arbeiten aus dem Museum Georg Frey, 4, 309 - 346.","Scholtz, C. H. (1980) Monograph of the genus Trox F. (Coleoptera: Trogidae) of Subsaharan Africa. Cimbebasia Memoir, 4, 1 - 104.","Scholtz, C. H. (1982) Catalogue of world Trogidae (Coleoptera: Scarabaeoidea). Entomology Memoir, Department of Agriculture and Fisheries, Republic of South Africa, 54, 1 - 27.","Scholtz, C. H. (1986) Phylogeny and systematics of the Trogidae (Coleoptera: Scarabaeoidea). Systematic Entomology, 11, 355 - 363.","Zidek, J. (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta Mundi, 314, 1 - 38.","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics and Evolution, 47, 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133"]}

Phoberus MacLeay, 1819 Phoberus MacLeay, 1819: 137���138; Str��mpher et al. 2014: 557���558; Str��mpher et al. 2016: 15; Zidek 2013: 6; Z��dek 2017: 96. Type species. Phoberus horridus MacLeay, 1819, by monotypy., Published as part of Str��mpher, Werner P. & Scholtz, Clarke H., 2019, A new Phoberus MacLeay species (Coleoptera: Scarabaeoidea: Trogidae) from the Maloti-Drakensberg Mountains of southern Africa and the Phoberus caffer species group reviewed, pp. 551-563 in Zootaxa 4674 (5) on page 552, DOI: 10.11646/zootaxa.4674.5.4, http://zenodo.org/record/3465084, {"references":["MacLeay, W. S. (1819) Horae entomologicae; or essays on the annulose animals. Containing general observations on the geography, manners, and natural affinities of the Insects which compose the genus Scarabaeus of Linnaeus; to which are added a few incidental remarks on the genera Lucanus and Hister of the same author. With an appendix and plates. Volume 1, Part 1. S. Bagster, London, United Kingdom, 160 pp.","Strumpher, W. P., Sole, C. L., Villet, M. H. & Scholtz, C. H. (2014) Phylogeny of the family Trogidae (Coleoptera: Scarabaeoidea) inferred from mitochondrial and nuclear ribosomal DNA sequence data. Systematic Entomology, 39 (3), 548 - 562. https: // doi. org / 10.1111 / syen. 12074","Strumpher, W. P., Villet, M. H., Sole, C. L. & Scholtz, C. H. (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea). Insect Systematics & Evolution, 47 (1), 53 - 82. https: // doi. org / 10.1163 / 1876312 X- 46052133","Zidek, J. (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta Mundi, 314, 1 - 38.","Zidek, J. (2017) Updated checklist and bibliography of family Trogidae (Coleoptera: Scarabaeoidea). Folia Heyrovskyana, Series A, 25 (1), 93 - 127."]}