Trox cyrtus Haaf, 1953: 333 Trox ( Trox ) cyrtus : Scholtz 1980a: 66 Scholtz 1982: 3 Trox ( Phoberus ) cyrtus : Zidek 2013: 9 Phoberus cyrtus : Strümpher et al. 2016b: 78 Zídek 2017: 99 Proper cleaning of keratin beetles reveals a cryptic species: Phoberus fumarius (Haaf, 1953) from southern Africa is reinstated as valid (Coleoptera: Trogidae) Strümpher, Werner P. Stals, Riaan Zootaxa 2021 2021-03-16 4942 4 543 557 4GJWC (Haaf, 1953) Haaf 1953 [151,509,930,957] Insecta Trogidae Phoberus Animalia Coleoptera 2 545 Arthropoda species cyrtus  ( 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: Strümpher et al.2016b: 78(new combination);  Zídek 2017: 99(checklist).     Typelocality. Vallée du Haute Orange[Valley of high Orange River] (border of Mohale’s Hoekand Quthing Districts, Lesotho)  Type material examined.All type specimens were examined through detailed photographs.   HOLOTYPEƋ ( MNHN, aedeagus extracted) ( Figs 1–3): Rectangle, faded blue cardstock: “MUSEUM PARIS | Basoutoland, Lessouto [ Lesotho] | Vallée du HtOrange [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” || Rectangularcuratorial label, white: “ MNHN| EC10025”.   PARATYPES2ƋƋ ( MNHN): Bluelabel identical || Rectangle, red cardstock: “ PARATYPE” || Foxywhite label identical || Whitecuratorial labels: “ MNHN| EC10027” and “ MNHN| EC10029”.   PARATYPES 3♀♀( MNHN): Labelsidentical 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”.   PARATYPE1Ƌ ( MNHB): Bluelabel 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♁ Nova667 [farm], near Ladybrand, SE2927 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♀ idembut  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, Museumstaff ( 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 Endrödy-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: idembut  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 cyrtusand  Phoberus fumariusare 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 cyrtushas these tubercles widely separated, with 5–8 tubercles between the elytral base and the apical callosity, whereas these tubercles in  Phoberus fumariusare 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 cyrtushave the lobes narrow and their apices sharper or angular, whereas in  Phoberus fumariusthe lobes are broad and their apices more rounded (as described in the text and as shown in Figs 2and 6versus Figs 9and 13). Available information strongly suggests that the two species are allopatric and existent only in different, non-overlapping biomes,  Phoberus cyrtusin the summer rainfall area of southern Africa and  Phoberus fumariusin 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 cyrtusand  Phoberus fumariushave 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 fumariusis more likely to be confused in this way.  Phoberus sulcatusdiffers 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 sulcatusare indubitably different with the median lobe of this species divided into two broadly curved lobes ( Fig 18). We found that  Phoberus sulcatuscan 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 cyrtusis known from South Africa, Lesothoand Botswana( Fig. 21). In South Africathe species has been recorded from the Eastern Cape, KwaZulu-Natal, Mpumalangaand North WestProvinces.  Broad habitat associations.As far as it is known,  Phoberus cyrtusis 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 cyrtusis 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. 3062300334 Vallee du Haute Orange Lesotho Districts 2 545 1 Mohale's Hoek holotype 3062300332 MNHN Rectangle & Rectangular Lesotho Dr. E. 1500 -30.283333 Rectangle 1222 27.766666 Rectangle 2 545 EC10025 2 holotype 3062300309 MNHN Foxy & White Lesotho Blue Rectangle 2 545 EC10027, EC10029 2 paratype 3062300301 MNHN Labels Lesotho 2 545 3 3 paratype 3062300336 MNHN Lesotho 2 545 EC10028, EC10026 1 1 paratype 3062300335 MNHB Lesotho Dr. E. Blue Rectangle 2 545 2 paratype 3062300318 1971-02-15 1971-02-25 1971-02-15 TMSA Snyman & Jones South Africa -30.7 Aliwal North 1220 26.7 2 545 1 Eastern Cape 3062300310 1976-01-19 1976-01-22 1976-01-19 BMSA GW Ferreira & S van Ee South Africa -29.116667 Ab 1228 27.366667 Nova 2 545 1 Free State 3062300348 1977-02-01 1977-02-12 1977-02-01 BMSA Ferreira & Strydom South Africa -29.116667 Ab 1228 27.366667 Nova 2 545 3 3 Free State 3062300305 1976-11-08 1976-11-18 1976-11-08 BMSA Museum South Africa Kerkenberg -28.50389 Ac 20 29.108334 Harrismith 2 545 4 4 Free State 3062300314 1981-11-13 1981-11-14 1981-11-13 SANC Tonder & Kok South Africa -29.116667 Mdedelelo Forestry Area 1228 29.433332 2 545 1 KwaZulu-Natal 3062300316 [151,1433,151,179] 1994-04-24 TMSA Harrison & R Stals 2300 -29.25 Castle Game Reserve 1228 29.433332 Bannerman Hut 3 546 1 3062300320 1994-01-28 TMSA S Endrody-Younga -24.983334 Skukuza Rest Camp 1248 31.6 Kruger National Park 3 546 1 Mpumalanga Province 3062300338 1970-10 TMSA L Schulze -26.5 Ermelo district 1241 29.116667 Winkelhaak 3 546 1 Mpumalanga Province 3062300340 1956-10-28 1961-08-13 1956-10-28 NHMB L Schulze Botswana -25.65 Rustenburg 1245 27.233334 Winkelhaak 3 546 1 North West 3062300342 1963-09-02 TMSA JA Nagle Botswana -25.033333 Otse 925 25.73889 3 546 1 South East