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TOKTOKKUS HERERO GEARNER lsid urn:lsid:zoobank.org:act: DE95E0D8-39B9-4555- AF0C-77AEDD599E20 Type material: Holotype (deposited at USNM), male: ‘ Abachaus. / Otjiwarongo / S.W. A/ G. Hobohm / 24-2- 1945 ’, ‘ Psammodes / tuberculipennis/ Haag / det. Penrith’. Diagnosis: This species is the only known species in this genus with macroscopic punctation on the pronotum and setae on the elytra (Fig. 3D). This species can be further differentiated from T. mashunus, T. mulleri, T. schultzei, T. vialis and T. waclawae by the presence of microtubercles between tuberculate rows on the elytra. This species, while round, is much more elongate than T. tschinkeli and lacks the lip-like margin of the prosternum characteristic of T. tschinkeli. The tubercles in this species are smaller and less prominent than those of T. tuberculipennis and T. makuya. Description: Length 25.0 mm, width of pronotum 8.0 mm and elytra 12.0 mm. Head: Hypognathous. Frons coarsely punctate (1–2 diameters apart); frontoclypeal suture course, with deep groove in middle; apical clypeal margin broadly shallowly emarginate; clypeus projected anterioventrally; apical margin of labrum emarginate medially, sparsely punctate in apical half, margin of labrum densely covered with yellowish, acuminate setae. Eye comma-shaped, with reduced ventral part, strongly emarginate around epistomal base; with deep groove on temporal side. Mentum trapezoidal, with straight base, not fully filling buccal cavity; anterior margin not emarginate; covered with fine setae. Submentum semicircular, concave basally. Antenna slender, moderately covered in recumbent acuminate goldish setae; antennomere 2 short, ~0.1– 0.2 of antennomere 3 length; antennomere 4 less than half of antennomere 3 length. Prothorax: Pronotal lateral margin rounded, ‘flared’ at anterior third, well visible. Pronotum widest above middle. Disc dull, with dense punctation; anterior fully marginate, posterior margin absent at middle, anterior apices strongly produced. Hypomeron convex, without submarginal groove, rugulose and occasionally punctate. Prosternal process rounded in lateral view with small projection by coxa, longitudinally depressed in middle (ventral view) between procoxae. Anterior margin of prosternum straight, with gold setae. Pterothorax: Scutellum densely covered with microtubercles. Elytra widest in basal third, slightly rounded, covered in scattered gold setae; disc impunctate, with scattered microtubercles; lateral half at and below humerus and declivous portion (apical third) with alternating rows of tubercles and microtubercles (larger than those on the disc); elytral margin not visible dorsally except in apical quarter, tuberculate rows extend to lateral margin. Elytral slope relatively steep, elytral apex flattened. Epipleura impunctate, not tuberculate, clearly distinguishable from neighbouring portion of elytra, widely enfolding fifth ventrite. Mesoventrite with deep median groove and elevated sides. Metaventrite impunctate, densely setose. Lateral regions of metaventrite (between coxae) extremely short. Metaepisternal suture abbreviated posteriorly. Legs: Covered with dense gold setae. Procoxa exposed basally. Apex of protibia with prominent denticle on outer margin, lateral carina terminating in basal third; median spur reduced, reaching 0.5 of outer lateral spur length. Spurs on meso- and metatibiae of equal length. Tarsi narrowed laterally. Abdomen: Ventrites 1–3 medially densely covered with goldish setae moderately punctate and weakly rugulose; ventrite 5 densely punctate, each puncture with single gold seta (~0.5 diameters apart); ventrite 5 with submarginal sulcus only at base. Terminalia: Due to scarcity of available specimens, terminalia were not dissected. Etymology: Named in honour of the Herero people of Namibia, where this species occurs.

Kamiński, Marcin J., Gearner, Olivia M., Kanda, Kojun, Swichtenberg, Kali, Purchart, Luboš, Smith, Aaron D. (2021): First insights into the phylogeny of tok-tokkie beetles (Tenebrionidae: Molurina, Phanerotomeina) and examination of the status of the Psammodes vialis species-group. Zoological Journal of the Linnean Society 191: 883-901, DOI: 10.1093/zoolinnean/zlaa052

TOKTOKKUS VIALIS SELLATUS (HAAG- RUTENBERG) = Psammodes kuisup P��ringuey, 1908, synon. nov. (as P. ��� kuisip ��� in Gebien, 1937 and Kaminski et al., 2019a). = Psammodes sellatus uriai Koch, synon. nov. Justification: When describing Psammmodes kuisup (in erratum, referred to in the description as Psammodes tuberculipennis), P��ringuey (1908) was apparently unaware of Psammodes sellatus Haag-Rutenberg and, therefore, did not provide diagnosable characters to differentiate the two species. Examination of specimens, including the P.sellatus holotype and a specimen collected at the type locality and determined by Koch as P. kuisup, and comparison of the species descriptions, reveal no easily diagnosable characters between these two species. Koch (1953b) described the subspecies P. sellatus uriai as differing from P. sellatus sellatus by the following characters: a more elongate body, a slenderer pronotum with less prominent punctation and tubercles less dense and organized in rows (as opposed to denser and more irregular in sellatus). However, when comparing specimens, these differences appear minor and not sufficient to justify classifying these as separate subspecies. NOTES ON PHANEROTOMEINA: GENUS CHILIARCHUM KOCH (SEPIDIINI: PHANEROTOMEINA) l s i d u r n:l s i d:z o o b a n k.o r g:a c t: C B7 1 2 3 7 1-B6 0 4- 4D1B-93A2-1258C6920572 Type species: Moluris (Phanerotoma) bertolonii Gu��rin-M��neville (by original designation). Diagnosis: From Koch (1954): submarginal area of the sides of the pronotum, and often that of the elytra, with either a broad stripe of white to yellow subtomentose hairs along the lateral carina or with densely conglomerated micropunctation or granulation with associated scattered short, fine, squamulated bristles; pronotum with two prebasilar impressions at or near the posterior angles, filled with pale subtomentose hairs or densely micropunctured with scattered short, fine bristles; males with a reddish brown brush of hairs on the outer edge of the upper surface of the mesotibiae, sometimes extending to the tibial base. Species and subspecies included (8): Chiliarchum arnoldi arnoldi (Koch, 1952) comb. nov., C. arnoldi sabianus (Koch, 1952) comb. nov., C. bertolonii (Gu��rin-M��neville, 1844) comb. nov., C. freyi (Koch, 1952) comb. nov., C. guerini guerini (Haag- Rutenberg, 1871) comb. nov., C. guerini lawrencii (Koch, 1954) comb. nov., C. guerini mancus (Koch, 1954) comb. nov., C. junodi (P��ringuey, 1899) comb. nov., Published as part of Kami��ski, Marcin J., Gearner, Olivia M., Kanda, Kojun, Swichtenberg, Kali, Purchart, Lubo�� & Smith, Aaron D., 2021, First insights into the phylogeny of tok-tokkie beetles (Tenebrionidae: Molurina, Phanerotomeina) and examination of the status of the Psammodes vialis species-group, pp. 883-901 in Zoological Journal of the Linnean Society 191 on page 899, DOI: 10.1093/zoolinnean/zlaa052, http://zenodo.org/record/5724192, {"references":["Peringuey LA. 1908. Tenebrionidae und Curculionidae. Denkschriften der Medicinisch- Naturwissenschaftlichen Gesellschaft zu Jena 13: 393 - 424.","Gebien H. 1937. Katalog der Tenebrioniden (Col. Heteromera). Teil I. Pubblicazioni del Museo Entomologico \" Pietro Rossi \" 2: 505 - 883.","Kaminski MJ, Kanda K, Lumen R, Ulmer JM, Wirth CC, Bouchard P, Aalbu R, Mal N, Smith AD. 2019 a. A catalogue of the tribe Sepidiini Eschscholtz, 1829 (Tenebrionidae, Pimeliinae) of the world. ZooKeys 844: 1 - 121.","Koch C. 1953 b. The Tenebrionidae of southern Africa XVII. Contributions to the fauna of Angola. Publicacoes Culturais da Companhia de Diamantes de Angola 16: 61 - 96.","Koch C. 1954. The Tenebrionidae of southern Africa XXVI. New Port. East African species collected by Dr A. J. Barbosa. Revista da Faculdade de Ciencias, Universidade de Lisboa. Serie C. Ciencias Naturais 3: 239 - 310.","Koch C. 1952. Die Tenebrioniden des sudlichen Afrikas XIII. Vorstudien zu einer Monographie der Molurini, 3. (Col. Tenebrionidae). Entomologische Arbeiten 3: 214 - 349."]}

TOKTOKKUS MAKUYA GEARNER lsid urn:lsid:zoobank.org:act: 7265FD25-E2AF- 49BD-947A-A4C6D80E0C97 Type material: Holotype (USNM), male: ‘South Africa: Limpopo: Makuya / N. R., Mutale Falls Safari Camp, / 300m, 22.42°S 31.05°E, / 2.ii.2018. KK18_064. K. Kanda, / R. A. Gomez, J. M. Pflug’. Paratypes: Four males and three females (Kojun Kanda): same data as holotype; male (PERC): same data but with: ‘Tenebrionid Base / Aaron D Smith / Catalog # 22651’; female (CASC): ‘ MOZAMBIQUE / Lourenco Marques’, ‘ ii 1957 / NLHKrauss’; male (CASC): ‘ 28°E 18°S, Rhodesia / Hostes Nicolle Inst. / Wildlife Res. Jan. / 1974 M. B. Fenton’; male and three females (T. Keith Philips): ‘SO. AFRICA: Northern Prov. just NW of Sukses, Madikela / Game Res., 12–14.III.1999 / Philips, Gerofsky, and Kryger / S24°05’, E28°18 ’’; male (CASC): ‘S. AFRICA / Bechuanaland / Tsuagara / Jan. 7, 1965 / John W. Neal’; male (CASC): ‘Naawpoort; / Pbg 21-11- 21 / G. V. Son’, ‘ PSAMMODES / vialis / Burchell / det. Dr. C. Koch’; three females (LuboŠ Purchart): ‘ AFRICA, MOZAMBIQUE / S 22°04.963’; E 33°55.577’ / camp site / 3–4. iii. 2011 ’, ‘ R. Blažek lgt.’; male (BMNH): ‘Naauwpoort, / Petersburg / distr. / 25/x.1928 / G. V.Son’; male (BMNH): ‘Naauwpoort, / Petersburg Distr. / 25/ xii. 27. / G. van Son’; female (BMNH): ‘Lake / Nigami’; male (BMNH): ‘ P. vialis Burchell / = pierreti Amyot / det. K.G.Blair.’, ‘ Psammodes tuberculipennis’, ‘L. M. / 20-1-09’, ‘Pres. by / Imp. Bur. Ent. / Brit. Mus. 1925–93.’, ‘Lourenco Marques: / 20.1.1909 / Howard Coll.’; female (BMNH): ‘ ZAMBIA 1147m / Lukwakwa, West Lunga N.P. / S12°39’40”, E24°26’13” / 28–29.iv.14. Light Trap / leg. Smith, R., Takano, H., / Chmurova, L, & Smith, L.’, ‘ Psammodes / vialis / E. Ruzzier det. 2015’; male (BMNH): ‘Damara Land’, ‘F. Bates / 81–19’; male and two females (Ted MacRae): ‘ R.S. Africa Northern Prov. / waterberg, Goelhoutbosh / 24°22’34’’ S, 27°33’ 64’’ E’, ‘ 29.xi.1999, T.C. MacRae / Nocturnally in sandy / ground in open woodland’; male (MIZ PAS): same data; three females (PERC): same data; female (XXX): ‘ SOUTH AFRICA 1965 / Bechuanaland 5 IV / Ngamiland Nokaneng’; two males and a female (CASC): ‘SW Africa / 19° 14’S / 20° 14’E’, ‘CO Handley Jr / XI. 28, 1952 ’. Diagnosis: The presence of microtubercles between the tuberculate rows on the elytra of this species distinguishes it from the following species: T. mashunus, T. mulleri, T. schultzei, T. vialis and T. waclawae. This species can be differentiated from T.tschinkeli by the lack of a prominent lip-like structure on the margin of the prosternum, and the elytra, while round, are more elongate than T. tschinkeli. The tubercles in this species are taller and less dense than those of T. tuberculipennis and T. herero, with those of T. tuberculipennis being nearly confluent at times. Description: Length 28.0–33.0 mm, width of pronotum 10.0–13.0 mm and elytra 17.0–22.0 mm. Head: Hypognathous. Frons finely punctate (2–4 diameters apart); frontoclypeal suture course, with deep groove in middle; apical clypeal margin broadly shallowly emarginate to non-emarginate; clypeus projected toward front of body; apical margin of labrum shallowly to sharply emarginate medially, densely punctate (although punctures often fine) in apical half, margin of labrum densely covered with yellowish, acuminate setae. Eye comma-shaped, with reduced ventral part, strongly emarginate around epistomal base; with deep groove on temporal side. Mentum trapezoidal, with straight base, not fully filling buccal cavity; anterior margin not emarginate. Submentum semicircular, concave basally. Antenna slender, moderately covered in recumbent acuminate goldish setae; antennomere 2 short, equal to 0.1–0.2 of antennomere 3 length; antennomere 4 about half of antennomere 3 length; length of antenna slightly longer than pronotal length. Prothorax: Pronotal lateral margin rounded, well visible. Pronotum widest at middle. Disc dull, impunctate; anterior fully marginate, posterior margin occasionally absent at middle, anterior apices strongly produced. Hypomeron convex, without submarginal groove, impunctate but weakly rugulose in places. Prosternal process rounded in lateral view with small projection by coxa, longitudinally depressed in middle (ventral view). Anterior margin of prosternum straight to slightly projecting ventrally (lateral view). Pterothorax: Scutellum densely covered with microtubercles. Elytra widest at middle, slightly rounded; disc impunctate, not covered by tubercles; lateral part (below humerus) covered with tubercles (organized in ~7–8 more or less regular rows on each elytron), with microtubercles scattered between rows; declivous portion on each elytron with additional 3–4 tuberculate rows with microtubercles in between; elytral margin not visible dorsally except in apical quarter, tuberculate rows extend more or less to lateral margin. Elytral slope relatively steep, elytral apex flattened. Epipleura, impunctate, not tuberculate, clearly distinguishable from neighbouring portion of elytra, widely enfolding fifth ventrite. Mesoventrite with deep median groove and elevated sides. Metaventrite impunctate, often densely setose (in males). Lateral regions of metaventrite (between coxae) short. Metaepisternal suture abbreviated posteriorly. Legs: Covered with dense gold setae. Procoxa exposed basally. Apex of protibia with prominent denticle on outer margin, lateral carina terminating at middle to basal third; median spur reduced, reaching 0.5 of outer lateral spur length. Spurs on meso- and metatibiae of equal length. Tarsi narrowed laterally. Abdomen: Ventrites 1–3 medially densely covered with goldish setae (males), moderately punctate and weakly rugulose; ventrite 5 densely punctate; ventrite 5 without submarginal sulcus. Terminalia: Aedeagus as in Figure 1E. Ovipositor similar to others in the genus (Fig. 6A–H). Etymology: Named after the locality from which the holotype was collected and is also the name of a tribe in the Limpopo province of South Africa.

TOKTOKKUS WACLAWAE KAMIŃSKI lsid urn:lsid:zoobank.org:act: CCA6EFFB-401D-40AF- 9341-AB40107E2698 Type material: Holotype (MIZ PAS), male: ‘N Rhodesia / 8 I 1942 / Dr W. Eichler’, ‘42’, ‘ Psammodes / pierreti Amyot’. Paratypes (MIZ PAS): male and female: ‘N Rhodesia / 10 I 1942 / Dr W. Eichler’; male: ‘KafueR / Rhodesia / UnivFilmEx’, ‘HCRaven / Dec`19 col’; male: ‘ Victoria Falls / Zimbabwe /??? 11–89’; female: ‘ Psammodes Pierreti / Solier / Zamberer’, ‘ Sammlung / Schroeder’; female: ‘ ZIMBABWE Victoria / Falls, 17°56’S 25°50’E / 19–22 Dec 1995 / W.J. Pulawski collector’; male and five females (LuboŠ Purchart): ‘ AFRICA, S ZAMBIA / Victoria falls env. (Livingstone) / 26.–30.xii.1993 / leg. J. Moravec’. Diagnosis: Similar to T. vialis due to the lack of microtubercles between the lateral tubercles, and presence of longitudinal ridges on elytral sides and slope. Both species can be distinguished by different morphology (larger and often confluent in T. waclawae) and arrangement (rows extend over humeri to scutellum in T. waclawae) of tubercles; and elytral disc (almost flat in lateral view in T. waclawae) (Fig. 3F). Description: Length 29.0–30.0 mm, width of pronotum 9.0–10.0 mm and elytra 19.0–20.0 mm. Head: Hypognathous. Frons finely punctate (3–4 diameters apart); frontoclypeal suture course, with deep groove in middle; apical clypeal margin broadly shallowly emarginate; clypeus projected toward front of body; apical margin of labrum sharply emarginate medially, densely punctate (although punctures fine) in apical half, apical side of labrum densely covered with yellowish, acuminate setae. Eye comma-shaped, with reduced ventral part, strongly emarginate around epistomal base; with deep groove on temporal side. Mentum trapezoidal, with straight base, not fully filling buccal cavity; anterior margin not emarginate; covered with fine setae. Submentum semicircular, concave basally. Antenna slender, moderately covered in recumbent acuminate goldish setae; antennomere 2 short, equal to 0.2 of antennomere 3 length; antennomere 4 about half of antennomere 3 length; length of antenna equal to 0.85 of pronotal length. Prothorax: Pronotal lateral margin rounded, well visible. Pronotum widest above middle. Disc dull, impunctate; anterior and basal margins, anterior apices strongly produced. Hypomeron convex, without submarginal groove, impunctate. Prosternal process rounded in lateral view, longitudinally depressed in middle (ventral view). Anterior margin of prosternum labiate, strongly projecting ventrally (lateral view). Pterothorax: Scutellum densely covered with microtubercles. Elytra widest in basal third, rounded laterally; disc impunctate, not covered by tubercles; lateral part (below humerus) covered with tubercles (organized in more or less regular rows) and microtubercles (2–4 diameters apart); remaining lateral part of elytra visible only ventrally, impunctate, without tubercles and microtubercles. Elytral slope steep, densely covered with microtubercles (1–4 diameters apart), with sparsely distributed tubercles, elytral apex flattened. Epipleura, impunctate, not tuberculate, clearly distinguishable from neighbouring portion of elytra, widely enfolding fifth ventrite. Mesoventrite with deep median groove and elevated sides. Metaventrite impunctate, densely setose. Lateral regions of metaventrite (between coxae) extremely short. Metaepisternal suture abbreviated posteriorly. Legs: Covered with dense, goldish setae. Procoxa exposed basally. Apex of protibia with prominent denticle on outer margin, lateral carina terminating in basal third; median spur reduced, reaching 0.5 of outer lateral spur length. Spurs on meso- and metatibiae of equal length. Tarsi narrowed laterally. Abdomen: Ventrites 1–4 medially densely covered with goldish setae moderately punctate and weakly rugulose; ventrite 5 densely punctate and setose; ventrite 5 without submarginal sulcus, densely punctured (~0.5 diameters apart), each puncture with single goldish setae. Terminalia: Aedeagus as in Figure 1E. Ovipositor as in Figure 6C–H. Etymology: This new species is dedicated to the memory of the first author’s grandmother, Wacława Kamińska (born on 5 November 1927 in Bartniki, Poland, died on 29 September 2010, in Warsaw, Poland). SYNONYMY NOTES

KEY TO THE SPECIES OF THE GENUS TOKTOKKUS 1. Apex of elytra not sloped (dorsal view) and depressed with clear margin (Fig. 3B)............... T. congolensis- Apex of elytra sloped (dorsal view) and flat to slightly convex (e.g. Fig. 3A)...............................................2 2. Margin of prosternal collar expanded and folded out into a large lip (Fig. 3C); elytra round, almost heart- shaped (Fig. 3C); declivous portion of elytra with little to no tubercles..................................... T. tschinkeli- Margin of prosternal collar not expanded, only occasionally folded out; elytra round or elongate; declivous portion of elytra with tuberculate rows.........................................................................................................3 3. Disc of pronotum with prominent punctures (Fig. 3D); gold setae present on elytra (Fig. 3D)..... T. herero- Disc of pronotum only with micropunctures; elytra not covered with setae................................................44. Tubercles on elytral sides round/globular, vertically direct (Fig. 2B, C)......................................................5- Tubercles on elytral sides pointed, directed posteriorly (e.g. Fig. 3F)..........................................................6 5. Elytral tubercles dense (1.0–1.5 diameters apart), laterally reaching humerus; elytra round (Fig. 2B)........................................................................................................................................................... T. mulleri - Elytral tubercles sparse (2–3 diameters apart), laterally terminating prior to humerus; elytra elongate to round (Fig. 2C)................................................................................................................................ T. schultzei6. Elytral tubercles sparse (4–6 lateral rows), deeply angled...........................................................................7- Elytral tubercles dense (more than 6 lateral rows), slightly angled (Fig. 2B–F).........................................8 7. Elytral tubercles distinct, not confluent, nearly reaching humeri (Fig. 3A)................................ T. barclayi - Elytral tubercles small and short, confluent into rows, terminating well before humeri (Fig. 2A)...................................................................................................................................................... T. mashunus8. Microtubercles present between tuberculate rows (e.g. Fig. 3E), tuberculate rows rarely elevated on ridges...............................................................................................................................................................9- Elytral tubercles all relatively the same size, no microtubercles present between rows, tuberculate rows often elevated on ridges................................................................................................................................10 9. Body size fairly small (18.0–23.0 mm); elytral tubercles relatively dense, almost confluent; tubercles relatively short.................................................................................................................... T. tuberculipennis - Body size medium to large (28.0–32.0 mm); elytral tubercles less dense, only occasionally confluent; tubercles taller................................................................................................................................ T. makuya 10. Elytral tubercles large, often confluent (Fig. 3F); tuberculate rows extend over humeri to scutellum; humeri prominent; disk of elytra where smooth often flat (Fig. 3F)......................................... T. waclawae- Tubercles medium sized (Fig. 2E, F), not confluent; tuberculate rows end at humeri; humeri not prominent; disc of elytra where smooth generally convex.................................................................................... T. vialis

TOKTOKKUS CONGOLENSIS KAMIŃSKI & GEARNER (FIG. 3B) lsid urn:lsid:zoobank.org:act: 86883CCF-78A5-4DD3- A2C6-0AFA496EB42A Type material: Holotype (BMNH), male: ‘ Congo / 80-32’. Diagnosis: This species is most similar to T. waclawae due to its large, dense, confluent tubercles, but can be distinguished by the apex of the elytra, which is straight and depressed with raised margins (Fig. 3B), while in T. waclawae this structure is sloped and lacking a depression. Description: Length 30.0 mm, width of pronotum 9.0 mm and elytra 12.0 mm. Head: Hypognathous. Frons finely punctate (2–4 diameters apart) except at middle; frontoclypeal suture course, with deep groove in middle; apical clypeal margin broadly shallowly emarginate to non-emarginate; clypeus projected toward front of body; apical margin of labrum shallowly emarginate medially, scattered punctation in apical half, margin of labrum densely covered with yellowish, acuminate setae. Eye comma-shaped, with reduced ventral part, strongly emarginate around epistomal base; with deep groove on temporal side. Mentum trapezoidal, with straight base, not fully filling buccal cavity; anterior margin not emarginate. Submentum semicircular, concave basally. Antenna slender, moderately covered in recumbent acuminate goldish setae; antennomere 2 short, equal to 0.2 of antennomere 3 length; antennomere 4 about twothirds of antennomere 3 length; length of antenna equal to or slightly longer than pronotal length. Prothorax: Pronotal lateral margin rounded, well visible. Pronotum widest above middle. Disc dull, impunctate; anterior margin complete, basal margin absent at middle, anterior apices strongly produced. Hypomeron convex, without submarginal groove, impunctate. Prosternal process rounded in lateral view, longitudinally depressed in middle (ventral view). Anterior margin of prosternum straight. Pterothorax: Scutellum densely covered with microtubercles. Elytra widest in basal third, slightly rounded; disc impunctate, without tubercles in middle; lateral part covered with nearly confluent tubercles organized in rows; declivous portion on each elytron with additional 3–4 tuberculate rows. Elytral slope relatively steep, elytral apex depressed with raised margin. Epipleura, impunctate, not tuberculate, clearly distinguishable from neighbouring portion of elytra, widely enfolding fifth ventrite. Mesoventrite with deep median groove and elevated sides. Metaventrite impunctate, setose. Lateral regions of metaventrite (between coxae) short. Metaepisternal suture abbreviated posteriorly. Legs: Covered with goldish setae. Procoxa exposed basally. Apex of protibia with prominent denticle on outer margin, lateral carina terminating at middle to basal third; median spur reduced, about half of outer lateral spur length. Spurs on meso- and metatibiae of equal length. Tarsi narrowed laterally. Abdomen: Ventrites 1–4 medially densely covered with goldish setae, weakly punctate and weakly rugulose; ventrite 5 without submarginal sulcus, densely punctured (~0.5 diameters apart), each puncture with single goldish setae. Terminalia: Due to scarcity of available specimens, terminalia were not dissected. Etymology: Specific epithet is derived from type locality, Congo, an area in Central Africa surrounding the Congo River.

TOKTOKKUS TSCHINKELI KAMIŃSKI & GEARNER (FIG. 3C) lsid urn:lsid:zoobank.org:act: 61AAB5AF-1529-4D7D- A25C-90FD186FE037 Type material: Holotype (deposited at USNM), male: ‘ Chiqubo, Mozam. / No. 11–20, II 1964 / Coll. A.L. Moore. ’. Paratypes (TMNH): male: ‘LimpopoRiv. / Mozambique Territory. / Feb. 1924. / F.Streeter’, ‘ Psammodes / sp? near / tuberculipennis / det. AJH.’; female: same data but lacking the identification label. Diagnosis: This species can be distinguished from its congeners by having margin of prosternal collar expanded and folded out into a large lip, extremely rounded elytra – almost heart-shaped, and declivous portion of elytra with little to no tubercles (Fig. 3C). Description: Length 29.0–32.0 mm, width of pronotum 9.0–10.0 mm and elytra 19.0–23.0 mm. Head: Hypognathous. Frons finely punctate (3–4 diameters apart); frontoclypeal suture course, with deep groove in middle; apical clypeal margin broadly shallowly emarginate; clypeus projected toward front of body; apical margin of labrum sharply emarginate medially, densely punctate (although punctures fine) in apical half, apical side of labrum densely covered with yellowish, acuminate setae. Eye comma-shaped, with reduced ventral part, strongly emarginate around epistomal base; with deep groove on temporal side. Mentum trapezoidal, with straight base, not fully filling buccal cavity; anterior margin not emarginate; covered with fine setae. Submentum semicircular, concave basally. Antenna slender, moderately covered in recumbent acuminate goldish setae; antennomere 2 short, equal to 0.2 of antennomere 3 length; antennomere 4 about half of antennomere 3 length; length of antenna equal to 0.85 of pronotal length. Prothorax: Pronotal lateral margin rounded, well visible. Pronotum widest above middle. Disc dull, impunctate; anterior and basal margins, anterior apices strongly produced. Hypomeron convex, without submarginal groove, impunctate. Prosternal process rounded in lateral view, longitudinally depressed in middle (ventral view). Anterior margin of prosternum labiate, strongly projecting ventrally (lateral view). Pterothorax: Scutellum densely covered with microtubercles. Elytra widest in basal third, rounded laterally; disc impunctate, not covered by tubercles; lateral part (below humerus) covered with tubercles (organized in more or less regular rows) and microtubercles (2–4 diameters apart); remaining lateral part of elytra visible only ventrally, impunctate, without tubercles and microtubercles. Elytral slope steep, densely covered with microtubercles (1–4 diameters apart), with sparsely distributed tubercles, elytral apex flattened. Epipleura, impunctate, not tuberculate, clearly distinguishable from neighbouring portion of elytra, widely enfolding fifth ventrite. Mesoventrite with deep, median groove and elevated sides. Metaventrite impunctate, densely setose. Lateral regions of metaventrite (between coxae) extremely short. Metaepisternal suture abbreviated posteriorly. Legs: Covered with dense, goldish setae. Procoxa exposed basally. Apex of protibia with prominent denticle on outer margin, lateral carina terminating in basal third; median spur reduced, reaching 0.5 of outer lateral spur length. Spurs on meso- and metatibiae of equal length. Tarsi narrowed laterally. Abdomen: Ventrites 1–4 medially densely covered with goldish setae moderately punctate and weakly rugulose; ventrite 5 densely punctate and setose; ventrite 5 without submarginal sulcus, densely punctured (~0.5 diameters apart), each puncture with single goldish setae. Terminalia: Due to scarcity of available specimens, terminalia were not dissected. Etymology: Named in honour of Dr Walter R.Tschinkel (Florida State University), an internationally renowned entomologist, for his outstanding contributions to darkling beetle systematics.

Tribe Platynotini Mulsant & Rey, 1853 Subtribe Eurynotina Mulsant & Rey, 1854 Genus Menederopsis Koch, 1954b: 16 Type species. Menederopsis constrictus Koch, 1954b (by original designation). = Archinamaqua Schawaller, 2012: 80 syn. nov. Type species. Archinamaqua lyleae Schawaller, 2012 (by original designation), Published as part of Kamiński, Marcin J. & Schawaller, Wolfgang, 2019, The taxonomic identity of some enigmatic darkling beetle genera: Archinamaqua Schawaller, 2012 and Menederopsis Koch, 1954 (Coleoptera: Tenebrionidae) from Namaqualand, South Africa, pp. 291-294 in Zootaxa 4543 (2) on page 293, DOI: 10.11646/zootaxa.4543.2.8, http://zenodo.org/record/2617837, {"references":["Koch, C. (1954 b) Die Tenebrioniden des sudlichen Afrikas XV. Revisioin der Oncotini nov. trib. Opatrinae (Psectropini Kaszab p. p.). Arkiv Zoologi Stockholm, 7, 1 - 96.","Schawaller, W. (2012) A new genus and species of Tentyriini (Coleoptera: Tenebrionidae) from sand dunes in Namaqualand, South Africa. Zootaxa, 3514, 79 - 83"]}

Menederopsis constrictus Koch, 1954b: 16 Type data: Holotype, female (Ditsong Museum): “Buffels River near Grootmist / 21 Nov. 1948 /leg. Univ. Calif. Transv. Mus. Exp. ” = Archinamaqua lyleae Schawaller, 2012: 80 syn. nov. Type data: Holotype (Ditsong Museum) and 35 paratypes (Ditsong Museum and Stuttgart State Museum of Natural History). For label information see Schawaller (2012)., Published as part of Kamiński, Marcin J. & Schawaller, Wolfgang, 2019, The taxonomic identity of some enigmatic darkling beetle genera: Archinamaqua Schawaller, 2012 and Menederopsis Koch, 1954 (Coleoptera: Tenebrionidae) from Namaqualand, South Africa, pp. 291-294 in Zootaxa 4543 (2) on page 293, DOI: 10.11646/zootaxa.4543.2.8, http://zenodo.org/record/2617837, {"references":["Koch, C. (1954 b) Die Tenebrioniden des sudlichen Afrikas XV. Revisioin der Oncotini nov. trib. Opatrinae (Psectropini Kaszab p. p.). Arkiv Zoologi Stockholm, 7, 1 - 96.","Schawaller, W. (2012) A new genus and species of Tentyriini (Coleoptera: Tenebrionidae) from sand dunes in Namaqualand, South Africa. Zootaxa, 3514, 79 - 83"]}

Anaxius pseudoloensus sp. nov. http://zoobank.org/urn:lsid:zoobank.org:act:13054B1E-F5D9-43A0-BAE0-BAA29297C843 (Figs. 1G, 3G, 5) Type data. Holotype, male (TMSA): “Espungabera / 80.K.Jutha 1954”. Etymology. The name highlights the morphological resemblance of this species to representatives of the genus Loensus Lucas, 1920 (Pedinini: Pedinina). Diagnosis. This species is similar to A. campbellae by having, inter alia, protibia with shallow median cavity, widened protarsus, with subsequent tarsomeres slightly narrower, and not apically depressed intercoxal process of prostenum (lateral view). Anaxius pseudoloensus can be easily distinguished by its smaller body size (4.5 mm in pseudoloensus; 6.5–7.0 in campbellae), more slender elytra (Figs 1B vs 1G), and simple mesofemora in males (with preapical denticle in campbellae). Description. Body—length ca. 4.5 mm. Habitus as in Fig. 1G. Dorsal side of head covered with fine punctures (0.5–1.0 diameters apart). Frontoclypeal suture not visible. Canthus rounded at base; narrower than head at level of eyes. Clypeal emargination shallow (clypeal emargination width: depth ratio ca. 8.0). Labrum wide (width: length ratio ca. 1.9); emarginate medially; with basal membrane exposed. Ligula exposed, not covered by mentum. Mentum widest in apical part; with well-developed medial keel; with exposed lateral lobes; not covering cardo or basistipes; apically with median indentation. Submentum pentagonal; short. Apical segment of maxillary palpus trapezoidal; with sensory field occupying whole proximal edge. Second segment of maxillary palpus elongate (length: width ratio ca. 2.4). Palpifer located laterally on basistipes. Anterior tentorial pit circular. Eye not dorsally shielded by any keel. Margin between maxillary fossa and antenna insertion well marked. Antenna not available for study (broken off holotype). Pronotum slightly narrower than elytra (elytra: pronotum width ratio ca. 1.1); relatively narrow (length: width ratio of pronotal disc ca. 0.8); disc shiny, covered with fine punctures (0.5–1.0 diameters apart). Lateral sides of pronotal disc straight at basal half. Disc without apophyseal depressions. Basal, anterior, and lateral emargination of pronotal disc present. Base of pronotum straight. Posterior angles not protruding beyond base. Pronotal hypomeron dull; covered with very fine longitudinal rugosities; without submarginal indentation. Intercoxal process of prosternum not depressed apically (lateral view); apically with oval depression on ventral surface; slightly protruding towards first abdominal segment. Procoxae narrowly separated (procoxa: intercoxal process width ratio ca. 2.0). Five elytral intervals finely marked; remaining ones indistinguishable. First interval elevated in disc. Elytron covered by fine punctures (0.1 diameters apart; sometimes confluent). Elytral base straight; not emarginate. Elytral humerus rounded. Epipleuron slightly narrowing apically, then of constant width; basally covering all elytral intervals (ventral view). Scutellum relatively large; triangular. Metathoracic wings absent. Metaventrite extremely short (metacoxal cavity: metaventrite (between insertions of meso- and metacoxae) length ratio ca. 8.0). Process of first abdominal ventrite wide (distances between mesocoxae: metacoxae ratio ca. 0.9). Fifth abdominal ventrite without submarginal sulcus; covered with fine punctures (4–5 diameters apart). Male protarsus widened, with subsequent tarsomeres slightly narrower. Protibia with shallow longitudinal cavity. Metafemora with fringe of setae distributed on whole length. Other leg parts simple. Meso- and metatibiae not avaliable for study (broken out of holotype). Aedeagal tegmen widest in middle; unipartite; without basal gap. Apical part divided (up to 0.10 of whole tegmen length). Clava straight. Penis of even width along most of its length; with apical part not covered by tegmen, exposed dorsally. Basal apophyses of penis relatively short (ca. 0.4 of whole tegmen length). Distribution. This species has been collected in the following ecoregion of Mozambique (Fig. 5): Southern Miombo woodlands., Published as part of Kamiński, Marcin J. & Schoeman, Colin S., 2018, Taxonomic revision of a darkling beetles genus Anaxius (Tenebrionidae: Pedinini: Helopinina), pp. 471-485 in Zootaxa 4455 (3) on pages 481-482, DOI: 10.11646/zootaxa.4455.3.4, http://zenodo.org/record/1457467

Anaxius meletsensis sp. nov. http://zoobank.org/urn:lsid:zoobank.org:act:79A5A46A-8614-410A-A322-AC3CF9933242 (Figs. 1D, 2G, I, 3D, 5) Type data. Holotype, male (TMSA): “ S. Afr.; Limpopo Prov. / Meletse Reserve 1003m / 24.36 S – 27.39 E ”, “ 27.11.2014; E-Y:3954 / on ground, bushveld / leg. Ruth Müller ”. Paratypes, 2 males and female (TMSA) and male (MIZ PAN): same data as holotype. Other material, female (TMSA): “ S. Afr.: Waterberg / Geelhoutbush farm / 24.22S – 27.33 E ”, “ 3.10.1955; E-Y:3143 / singled / Endrödy & Bellamy ”. Etymology. The name refers to the type locality, Meletse Reserve (Limpopo, South Africa). Diagnosis. This species is similar to A. obesus by having, inter alia, protibia with deep metian cavity (visible laterally) (Fig. 2G), widened protarsus, with subsequent tarsomeres slightly narrower, and not apically depressed intercoxal process of prostenum (lateral view). Anaxius meletsensis can be easily distinguished by having simple mesofemora (with preapical denticle in obesus) and metafemora (with fringe of bristles on underside in obesus). Moreover, some differences might be observed in the structure of the metatibae of both species, which are curved in the middle in meletsensis (Fig. 1D), and more basally in obesus (Fig. 1E). Description. Body—length = 7.0– 7.5 mm. Habitus as in Fig. 1D. Dorsal side of head covered with confluent punctures. Frontoclypeal suture smooth. Canthus rounded at base; narrower than head at level of eyes. Clypeal emargination shallow (clypeal emargination width: depth ratio ca. 9.0). Labrum wide (width: length ratio ca. 1.9); slightly emarginate medially; with basal membrane exposed. Ligula exposed, not covered by mentum. Mentum widest in apical part; with well-developed medial keel; with exposed lateral lobes; not covering cardo or basistipes; apically with median indentation. Submentum pentagonal; short. Apical segment of maxillary palpus trapezoidal; with sensory field occupying whole proximal edge. Second segment of maxillary palpus elongate (length: width ratio ca. 2.6). Palpifer located laterally on basistipes. Anterior tentorial pit circular. Eye not dorsally shielded by any keel. Margin between maxillary fossa and antenna insertion well marked. Antenna relatively long (antenna: pronotum length ratio ca. 1.7); third antennomere short (3rd: 2nd antennomere length ratio ca. 3.0); fifth antennomere elongated (length: width ratio ca. 2.0). Pronotum slightly narrower than elytra (elytra: pronotum width ratio ca. 1.2); relatively narrow (length: width ratio of pronotal disc ca. 0.9); disc shiny, covered with fine punctures (0.5–1.0 diameters apart). Lateral sides of pronotal disc straight at basal half. Disc without apophyseal depressions. Basal, anterior, and lateral emargination of pronotal disc present. Base of pronotum straight. Posterior angles not protruding beyond base. Pronotal hypomeron dull; covered with very fine longitudinal rugosities; without submarginal indentation. Intercoxal process of prosternum not depressed apically (lateral view); apically with oval depression on ventral surface; slightly protruding towards first abdominal segment. Procoxae narrowly separated (procoxa: intercoxal process width ratio ca. 2.0). Five elytral intervals finely marked; remaining ones indistinguishable. Elytron covered by fine punctures (0.1 diameters apart; sometimes confluent). Elytral base straight; not emarginate. Elytral humerus rounded. Epipleuron slightly narrowing apically, then of constant width; basally covering all elytral intervals (ventral view). Scutellum relatively large; triangular. Metathoracic wings absent. Metaventrite extremely short (metacoxal cavity: metaventrite (between insertions of meso- and metacoxae) length ratio ca. 10.0); with longitudinal process at middle in males (Fig. 2I). Process of first abdominal ventrite wide (distances between mesocoxae: metacoxae ratio ca. 0.9). Fifth abdominal ventrite without submarginal sulcus; covered with fine punctures (4–5 diameters apart). Male protarsus widened, with subsequent tarsomeres slightly narrower. Protibia with deep longitudinal cavity. Mesotibiae, profemora, mesofemora, and metafemora simple. Metafemora similar to that of A. obesus. Female legs simple. First tarsomere of metatarsi elongated (1.7× longer than 4th one). Length of metatarsus equal to 0.75 of metatibial length. Aedeagal tegmen widest in middle; unipartite; with wide basal gap. Apical part slightly divided (up to 0.10 of whole tegmen length). Clava straight. Penis of even width along most of its length; with apical part not covered by tegmen, exposed dorsally. Basal apophyses of penis relatively short (ca. 0.4 of whole tegmen length). Ovipositor relatively short (body: ovipositor length ratio ca. 3.0). Paraproct much longer than coxites (paraproct: coxites length ratio ca. 1.8); triangular, not shielding valvifer or any other lobes of coxites. Valvifer and second lobe slightly transverse, while third one triangular. Fourth lobe rounded, situated dorsally. Gonostylus located on dorsal side of fourth lobe. Vagina and bursa copulatrix without sclerites. Spermatheca sac-like. Proctiger covering nearly whole ventral side of ovipositor. Distribution. This species has been collected in the following ecoregion of South Africa (Fig. 5): Southern Africa bushveld.

Anaxius bloubergensis sp. nov. http://zoobank.org/urn:lsid:zoobank.org:act:81D10A96-9238-4501-AF69-048DFF06453F (Figs. 1A, 2E, F, 3A, 5) Type data. Holotype, male (TMSA): “ S. Afr.; Limpopo Prov. / Blouberg Nat. Res. / 22.59 S – 29.08 E ”, “25– 27.11.2016 E-Y:3987 / day, sandy bushveld / leg. Ruth Müller ”. Paratypes, 3 males (TMSA): same data as holotype; male (TMSA) and male (MIZ PAN): “ S. Afr.; Limpopo Prov. Blouberg NR North / dd22 98S, dd 29.12E ”, “ 6.12.2012; 866m, BLN1 / Roodeberg Bushveld / leg. Colin Schoeman ”; male (MIZ PAN): “ Blouberg NR / Rodeberg Bushveld / -22.98; 29.12”, “leg. Colin Schoeman / BLN7c / I”. Etymology. The name refers to the type locality, Blouberg Nature Reserve (Limpopo, South Africa). Diagnosis. This species is distinguished from all other congeners by a unique structure of pro- (first tarsomere extremely widened, following tarsomeres narrow; Fig. 1A), mesotarsi (first tarsomere widened; Fig. 2F), mesotibiae (flattened laterally; protruding outwards apically; Fig. 2F), and aedeagal tegmen (without basal gap; Fig. 3A). Description. Body—length = 8.0-9.0 mm. Habitus as in Fig. 1A. Dorsal side of head covered with fine confluent punctures (0.5–1.0 diameters apart). Frontoclypeal suture rough. Canthus rounded at base; narrower than head at level of eyes. Clypeal emargination shallow (clypeal emargination width: depth ratio ca. 8.0). Labrum narrow (width: length ratio ca. 1.1); strongly emarginate medially; with basal membrane exposed. Ligula exposed, not covered by mentum. Mentum widest in apical part; with well-developed medial keel; with exposed lateral lobes; not covering cardo or basistipes; apically with median indentation. Submentum pentagonal; short. Apical segment of maxillary palpus trapezoidal; with sensory field occupying whole proximal edge. Second segment of maxillary palpus elongate (length: width ratio ca. 2.4). Palpifer located laterally on basistipes. Anterior tentorial pit circular. Eye not dorsally shielded by any keel. Margin between maxillary fossa and antenna insertion well marked. Antenna relatively long (antenna: pronotum length ratio ca. 1.4); third antennomere short (3rd: 2nd antennomere length ratio ca. 3.0); fifth antennomere elongated (length: width ratio ca. 2.0). Pronotum narrower than elytra (elytra: pronotum width ratio ca. 1.3); relatively narrow (length: width ratio of pronotal disc ca. 0.7); disc dull, covered with fine punctures (3–4 diameters apart). Lateral sides of pronotal disc straight. Disc widest in basal half; without apophyseal depressions. Basal and lateral emargination of pronotal disc present; anterior interrupted in middle. Base of pronotum straight. Posterior angles not protruding beyond base. Pronotal hypomeron dull; without punctures; covered with very fine longitudinal rugosities; without submarginal indentation. Intercoxal process of prosternum depressed apically (lateral view); apically with oval depression on ventral surface; slightly protruding towards first abdominal segment. Procoxae narrowly separated (procoxa: intercoxal process width ratio ca. 2.0). Elytral striae and intervals not clearly visible. Elytron covered by shiny tubercles (0.1 diameters apart) displayed on dull surface. 1st, 3rd, 5th and 7th intervals protruding on disc; transforming into sparsely (1–2 diameters apart) distributed, large, tubercles on elytral apex. Elytral base straight; not emarginate. Elytral humerus rounded. Epipleuron slightly narrowing apically, then of constant width; basally covering all elytral intervals (ventral view). Scutellum relatively large; triangular. Metathoracic wings absent. Metaventrite extremely short (metacoxal cavity: metaventrite (between insertions of meso- and metacoxae) length ratio ca. 10.0). Process of first abdominal ventrite wide (distances between mesocoxae: metacoxae ratio ca. 0.9). Fifth abdominal ventrite without submarginal sulcus; covered with fine punctures (4–5 diameters apart). Male pro- and mesotarsi with basal tarsomere widened. Protibia with deep longitudinal cavity. Profemora simple. Mesotibiae flattened laterally; protruding outwards apically. Mesofemora with internal extension in apical part. Metatibia and metafemora with fringe of setae distributed on whole length. Metatibia curved inwards. First tarsomere of metatarsi elongated (1.5× longer than 4th one). Length of metatarsus equal to half of metatibial length. Aedeagal tegmen widest in middle; unipartite; without basal gap. Apical part divided (up to 0.15 of whole tegmen length). Clava straight. Penis of even width along most of its length; with apical part not covered by tegmen, exposed dorsally. Basal apophyses of penis relatively short (ca. 0.4 of whole tegmen length). Distribution. This species has been collected in the following ecoregions of South Africa (Fig. 5): Drakensberg montane grasslands, woodlands and forests; and Southern African bushveld.

Anaxius obesus F��hraeus, 1870 (Figs. 1E, 5) Anaxius obesus F��hraeus, 1870: 307.- Koch, 1958: 211. Notes. F��hraeus (1870) did not specify the number of studied specimens, however from the context it seems to be clear that he was dealing with at least two specimens. Therefore, a lectotype designation is needed to fix the taxonomic status of the genus and the species. Material studied. Lectotype, designated here, male (TMSA): ���Caffra- / ria���. Paralectotype, male (TMSA): same data as holotype. Other material, male (TMSA): " Lichtenburg / Transvaal / Dr. Brauns ", male (TMSA): " Plansberg / XI.1950 / P.L. Breutz "; 2 males (TMSA): " S. Afr.; Limpopo Prov. / Mabote farm / 24.07 S ��� 28.39E ", " 12.12.2009 / in the Waterberg / leg. Ruth M��ller "; male (TMSA): ��� S. Afr.: Transvaal / Pienaars River 8km / 25.17S ��� 28.17E, 28.12.1994; E-Y 3234, on ground / leg. CL Bellamy���. Distribution. This species has been collected in the following ecoregions of South Africa (Fig. 5): Highveld grasslands; and Southern Africa bushveld., Published as part of Kami��ski, Marcin J. & Schoeman, Colin S., 2018, Taxonomic revision of a darkling beetles genus Anaxius (Tenebrionidae: Pedinini: Helopinina), pp. 471-485 in Zootaxa 4455 (3) on pages 479-481, DOI: 10.11646/zootaxa.4455.3.4, http://zenodo.org/record/1457467, {"references":["Fahraeus, O. I. (1870) Coleoptera Caffrariae a J. A. Walhberg collecta. - Ofversigt af Kongl. Akademiens Ftirhandlingar 4: 243 - 358.","Koch, C. (1958) Tenebrionidae of Angola. Publicacoes Culturais da Companhia de Diamantes de Angola, 39, 1 - 231."]}

Genus Anaxius F��hraeus, 1870 Anaxius F��hraeus, 1870: 307.- Koch 1958: 208. Type species. Anaxius obesus F��hraeus, 1870; by monotypy. Diagnosis. According to Koch���s (1958) hypothesis, this genus is close to Aptila F��hraeus, 1870 and Asidodema Koch, 1958 based on pronotal shape (subparallel sided; flat disc) and male terminalia (relatively short tegmen; clavae as large as penis). All three genera were previously classified within a single subtribe Aptilina. From Aptila and Asidodema, Anaxius can be distinguished by having widened protarsi and the apical segment of the maxillary palpus enlarged in males. Moreover, it differs from Aptila by the lack of scale-like vestiture on the upper surface of the body, while the relatively reduced prosternum differentiates it from Asidodema (see Koch 1958). Species included (7) . bloubergensis sp. nov., campbellae Koch, 1958, limpopoensis sp. nov., meletsensis sp. nov., obesus F��hraeus, 1870, prozeskyi Koch, 1958, pseudoloensus sp. nov. Distribution. Representatives of this genus have been collected in the following ecoregions of Namibia, Mozambique, South Africa and Zimbabwe (Fig. 5): Drakensberg montane grasslands, woodlands, and forests; Highveld grasslands; Kalahari xeric savanna; Southern Africa bushveld; and Southern Miombo woodlands. Anaxius was recorded from localities distributed between 874 and 1544 meters in elevation (Fig. 6). Note. During the present investigation 49 Anaxius specimens were studied. Among this number a total of three females was identified. Because of this the following key includes only the male morphology., Published as part of Kami��ski, Marcin J. & Schoeman, Colin S., 2018, Taxonomic revision of a darkling beetles genus Anaxius (Tenebrionidae: Pedinini: Helopinina), pp. 471-485 in Zootaxa 4455 (3) on page 472, DOI: 10.11646/zootaxa.4455.3.4, http://zenodo.org/record/1457467, {"references":["Fahraeus, O. I. (1870) Coleoptera Caffrariae a J. A. Walhberg collecta. - Ofversigt af Kongl. Akademiens Ftirhandlingar 4: 243 - 358.","Koch, C. (1958) Tenebrionidae of Angola. Publicacoes Culturais da Companhia de Diamantes de Angola, 39, 1 - 231."]}

Anaxius prozeskyi Koch, 1958 (Figs. 1F, 2C���D, 5) Anaxius prozeskyi Koch, 1958: 209. Material studied. Holotype, male (TMSA): ��� Anaxius / magnificus / Koch / C. Koch det. 195���, ���Blouberg, Tvl. / N. side, Glenferness / 16���21.I.1955 / Transva.Mus.Exp.���, ���HOLOTYPE / Anaxius / PROZESKYI���. Paratype, male (TMSA): ���Helopininae / anaxius /?���, ���186���, ������3811���, ���3427���, ���Makgaberg. / - 2.03. / 2457���, ���PARATYPE / Anaxius / PROZESKYI��� Other material, 5 males (TMSA) and male (MIZ PAN): ���S. Afr.: Limpopo Prov. / Blouberg Nat. Res. / 22.59S���29.08E���, ���25��� 27.11.2016 E-Y:3987 / day, sandy bushveld / leg. Ruth M��ller���; 3 males (TMSA): ���Namibia, Okahandja Dis/ 45 kmSE���Okakarara / 20.56S���17.42E���, ��� 4.3.2006; E-Y:3713 / single, sandveld, 1345m, Ruth M��ller���. Distribution. This species has been collected in the following ecoregions of Namibia and South Africa (Fig. 5): Kalahari xeric savanna; and Southern Africa bushveld., Published as part of Kami��ski, Marcin J. & Schoeman, Colin S., 2018, Taxonomic revision of a darkling beetles genus Anaxius (Tenebrionidae: Pedinini: Helopinina), pp. 471-485 in Zootaxa 4455 (3) on page 481, DOI: 10.11646/zootaxa.4455.3.4, http://zenodo.org/record/1457467, {"references":["Koch, C. (1958) Tenebrionidae of Angola. Publicacoes Culturais da Companhia de Diamantes de Angola, 39, 1 - 231."]}

Anaxius limpopoensis sp. nov. http://zoobank.org/urn:lsid:zoobank.org:act:0719746F-D175-4BF0-9B46-561914F390E1 (Figs. 1C, 3C, 5) Type data. Holotype, male (TMSA): “ S. Afr.; Limpopo Prov. / Lindani Nat. Res 1336m / 24.02 S – 28.23 E ”, “ 8.12.2005; E-Y:3687 / single, bushveld / leg. Gusmann, Müller ”. Paratypes, male (TMSA) and male (MIZ PAN): same data as holotype; 2 males (TMSA): “S. Afr. Limpopo Prov. / Mabote farm / 24.07S 28.39 E ”, “ 14.11.2009 / Leg. Ruth Müller ”; male (TMSA) “ S. Afr.; Limpopo Prov. / 25 km N Mookgophong / 24.25 S – 28.41 E ”, “12 – 15.1.2014 E-Y:3951 / mixed moodland 1185m / leg. Ruth Müller ”; male (TMSA): “ S. Afr.; Limpopo Prov. / Waterberg Game Res. / 24.11 S – 28.20 E ”, “4 – 7.11.2002 / leg. B. Dombrowsky”. Etymology. The name refers to the type locality, Limpopo Province (South Africa). Diagnosis. This species is similar to A. prozeskyi by having relatively narrow protarsi in males (like in Fig. 1F), strongly depressed apical part of the intercoxal process of prosternum (lateral view), and dorsal body surface covered with soft and dense bristles, with additional stiff and black setae on elytra (Fig. 1C, F). Anaxius limpopoensis can be easily distinguished by having simple mesotibia (curved in limpopoensis; flattened apically, protruding in middle in prozeskyi) profemora (with fringe of bristles on underside in prozeskyi), and mesofemora (with prominent denticle in prozeskyi). Description. Body—length = 5.0–7.0 mm. Habitus as in Fig. 1C. Dorsal side of head covered with fine confluent punctures (0.5–1.0 diameters apart) and fine yellow bristles. Frontoclypeal suture smooth. Canthus rounded at base; narrower than head at level of eyes. Clypeal emargination shallow (clypeal emargination width: depth ratio ca. 9.0). Labrum wide (width: length ratio ca. 1.5); shallowly emarginate medially; with basal membrane exposed. Ligula exposed, not covered by mentum. Mentum widest in apical part; with well-developed medial keel; with exposed lateral lobes; not covering cardo or basistipes; apically with median indentation. Submentum pentagonal; short. Apical segment of maxillary palpus trapezoidal; with sensory field occupying whole proximal edge. Second segment of maxillary palpus elongate (length: width ratio ca. 2.3). Palpifer located laterally on basistipes. Anterior tentorial pit circular. Eye not dorsally shielded by any keel. Margin between maxillary fossa and antenna insertion well marked. Antenna relatively long (antenna: pronotum length ratio ca. 1.6); third antennomere short (3rd: 2nd antennomere length ratio ca. 3.0); fifth antennomere relatively short (length: width ratio ca. 2.0). Pronotum narrower than elytra (elytra: pronotum width ratio ca. 1.3); relatively narrow (length: width ratio of pronotal disc ca. 0.7); disc dull, covered with confluent punctures (space between punctures shiny) and yellow bristles, which are becoming denser in posterior anges. Lateral sides of pronotal disc rounded. Disc widest in middle; without apophyseal depressions; with basal depressions. Basal, anterior and lateral emargination of pronotal disc present. Base of pronotum slightly bisinuate. Posterior angles not protruding beyond base. Pronotal hypomeron dull; with fine punctures; covered yellow bristles; without submarginal indentation. Intercoxal process of prosternum depressed apically (lateral view); not protruding towards first abdominal segment. Procoxae narrowly separated (procoxa: intercoxal process width ratio ca. 2.0). Elytral striae and intervals not visibly distinguishable. Elytron covered by confluent punctures, yellow hair, which become more dense closer to elytral suture and epipleuron, and additional stiff and black setae (Fig. 1C). Elytral base slightly bisinuate; not emarginate. Elytral humerus rounded. Epipleuron slightly narrowing apically, then of constant width; basally covering all elytral intervals (ventral view). Scutellum relatively large; triangular. Metathoracic wings absent. Metaventrite extremely short (metacoxal cavity: metaventrite (between insertions of meso- and metacoxae) length ratio ca. 10.0). Process of first abdominal ventrite wide (distances between mesocoxae: metacoxae ratio ca. 0.9). Fifth abdominal ventrite without submarginal sulcus; covered with fine punctures (4–5 diameters apart). Male protarsi relatively narrow. Protibia with shallow median cavity on inner side. Pro- and mesofemora simple. Meso- and meta curved; with fringe of setae distributed on whole length. Metafemora with fringe of setae distributed along whole length; with three spines. First tarsomere of metatarsi elongated (1.5× longer than 4th one). Metatarsus nearly as long as metatibia. Aedeagal tegmen widest in middle; bipartite. Apical part divided (up to 0.15 of whole tegmen length). Clava straight. Penis of even width along most of its length; with apical part not covered by tegmen, exposed dorsally. Basal apophyses of penis relatively short (ca. 0.4 of whole tegmen length). Distribution. This species has been collected in the following ecoregion of South Africa (Fig. 5): Southern Africa bushveld.