First insights into the phylogeny of tok-tokkie beetles (Tenebrionidae: Molurina, Phanerotomeina) and examination of the status of the Psammodes vialis species-group Author Kamiński, Marcin J. Author Gearner, Olivia M. Author Kanda, Kojun Author Swichtenberg, Kali Author Purchart, Luboš Author Smith, Aaron D. text Zoological Journal of the Linnean Society 2021 2020-06-16 191 883 901 journal article 3320 10.1093/zoolinnean/zlaa052 fd4b2931-7bad-4644-b485-f365040ae01e 0024-4082 5724192 0810A91E-7D8D-4F56-B468-9D5B3A6115E9 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 .