"Character support: Onitini, Oniticellini, and the Onthophagini monophyly (unweighted data)" "Uncontroverted synapomorphies:" "1) pygidium median groove absent (59-2);" "2) propygidium transversely with an even width (61-0);" "3) propygidium posterior/ventral border medially without either an angulate emargination anteriorly or rounded shape posteriorly (62-2);" "4) elytral striae composed of a single line (94-0);" "5) cervical lateral sclerite apex lacking a lateral pocket or cavity (104-1);" "6) meso- and metatibia broad and greatly expanded apically (125-1);" "7) metatibia not elongate and parallel sided at middle ½ (130-1)." "Controverted synapomorphies:" "8) cephalic projection consisting of a central horn positioned posteriorly (46-1);" "9) eye canthus completely dividing eye (49-1);" "10) ventral lacinial articulation sclerite with a short “tail” extending only part of length (76-1);" "11) four teeth on the protibia; (80-0);" "12) pronotal posterior margin with a slight but distinct posteriorly directed point or angulate edge (89-0)." "Character support: Oniticellini + Onthophagini monophyly (unweighted data)" "Uncontroverted synapomorphies:" "1) labial palps with the third palpomere reduced in size and appearing as having only two palpomeres (21-0);" "2) mentum paraglossal strut laterally at distal apex expanded but not bifurcate (29-1);" "3) angle of parameres to basal piece where attached approximately 45 degrees (54-1);" "4) length of parameres to basal piece, excluding narrow projections approximately ½ the length of basal piece (55-2);" "5) dorsal plate of the genital capsule oriented transversely (56-1)." "Controverted synapomorphies:" "1) mentum glossal lobe narrowly rounded (31-1);" "2) the antennae with vessicles present and easily visible (36-1);" "3) blunt paramere apex (53-1);" "4) mesonotum with the prescutum anteriorly transverse plate approximately ventrally directed and angularly emarginate (107-1);" "5) mesocutum lateral projection extended with a pointed apex (114-2);" "6) mesoscutum lateral edge lacking a pocket (115-1);" "7) metascutellum posterior edge with a slight margin or remnant edge (119-1)." "Character support: Oniticellini + Onthophagini monophyly (weighted data, k = 10)" "Uncontroverted synapomorphies:" "1) labial palps with the third palpomere reduced in size and appearing as having only two palpomeres (21-0);" "2) angle of parameres to basal piece where attached approximately 45 degrees (54-1);" "3) length of parameres to basal piece, excluding narrow projections approximately ½ the length of basal piece (55-2);" "4) dorsal plate of the genital capsule oriented transversely (56-1);" "5) wing AA vein posteriorly forming a cell partially closed with extension toward costal margin (8-2)." "Controverted synapomorphies:" "1) antennae with vessicles present and easily visible (36-1);" "2) blunt paramere apex (53-1);" "3) mesonotum with the prescutum anteriorly transverse plate approximately ventrally directed and angularly emarginate (107-1);" "4) paraglossal strut laterally at distal apex expanded, but not bifurcate (29-1);" "posterior margin of the gula on the head truncate and slightly projecting (51-1)." "Character support: Oniticellini monophyly (weighted data, k = 10)" "Uncontroverted synapomorphies:" "1) pygidium lacks a transverse ridge defining or separating the propygidium from the pygidium (58-0)." "Character support: Onthophagini monophyly (unweighted data)" "Uncontroverted synapomorphies:" "1) scutellum (metanotum) posterior margin with a short apical projection and with a truncate apex and laterally with rapidly converging sides (119-5)." "Controverted synapomorphies:" "1) dorso-lateral edge of propygidium angulate and not rounded; (63-1);" "2) mesonotum scutellum apex triangular shaped (106-1);" "3) prescutum-scutum junction when viewed in a horizontal position broadly rounded (109-2);" "4) mediophragma of the metanotum with a distinct longitudinal ridge (123-0)." "Character support: Onthophagini monophyly (weighted data, k = 10)" "Controverted synapomorphies:" "1) dorso-lateral edge of propygidium angulate, and not rounded (63-1);" "2) the scutellum (mesonotum) apex triangular shaped (106-1);" "3) prephragma of the prescutum (mesonotum) in ventral view recurved in shape (116-1);" "4) mediophragma (metanotum) with a distinct longitudinal ridge (123-0)." "Onthophagini clade 1 support: Haroldius + Cassolus through to Sukulus (unweighted data)" "Uncontroverted synapomorphy:" "1) apical extension of pigmented mesal comb compared to sclerotized (darkened) area near opposite lateral edge approximately the same length or slightly shorter; (92-2)." "Controverted synapomorphies:" "1) mentum length approximately equal to width (24-1);" "2) inner strut of lacinia (nearest to palpifer), distal tip, notch absent and tapered tip (75-2);" "3) scutum distinctly transverse (113-0);" "4) metatarsi: length of first metatarsis compared to second: first nearly 2X or more the length of second (126-0)." "Onthophagini clade 2 support: Megaponerophilus through to Sukulus (unweighted data)" "Uncontroverted synapomorphies:" "1) proximal antennomere of the apical club in lateral view (~4 to 5 times as long as wide) (37-1);" "2) mandibular cuticle on the lateral edge and part of ventral surface bright chestnut red color (93-0)." "Controverted synapomorphies:" "1) prothoracic apodemes with a complete oblique suture (40-0);" "2) apex of the paramere blunt and not pointed (53-1)." "Oniticellini clade 1 support: Helictopleurina (unweighted data)" "Controverted synapomorphies:" "1) prothoracic apodeme with a large flattened single flange and with one lateral edge slightly expanded perpendicularly (39-0) [only seen elsewhere in Paroniticellus];" "2) pronotal surface glabrous (88-0);" "3) elytral seventh stria strongly curved (100-0);" "4) anterior margin of scutum with an pale colored, transverse region of cuticle (111-0)." "Oniticellini clade 2 support: Drepanocerina: Cyptochirus + Afrodrepanus (unweighted data and weighted data, k = 10)" "Controverted synapomorphies:" "1) anal region of wing with a posterior notch (4-0);" "2) apodeme of cervical sclerite lacking a carina (40-2);" "3) a single carinae anterior of eyes (47-1);" "4) a transverse ridge defining or separating propygidium from the pygidium present (58-1);" "5) Elytral umeral angle in dorsal view with a lateral bulge (97-0);" "6) Mesonotum prescutum: prephragma in ventral view with recurve present (116-1)." "Oniticellini clade 3 support: Oniticellina and Drepanocerina (unweighted data)" "Controverted synapomorphies:" "1) RA vein extension along apical margin relatively great (7-1) [only seen elsewhere in Phalops and Onitis];" "2) internal apodeme (V or Y shaped usually with one arm relatively short) adjacent to paraglossal apodeme in dorsal view, from anterior to posterior obliquely angled outwards (33-0) [only seen elsewhere in Helictopleurus quadripunctatus]." "Oniticellini clade 4 support: Attavicinus + Paroniticellus (unweighted data)" "Controverted synapomorphies:" "1) internal accessory sclerite within the maxilla that is curved near the proximal end (74-1);" "2) the protibia on proximal side with a line of setae associated with a carina (81-0);" "3) males with two postero-lateral ridges on the pronotum (85-0);" "4) scutum distinctly transverse (113-0)." "Oniticellini clade 5 support: Liatongus californicus to Euoniticellus (unweighted data)" "Uncontroverted synapomorphies:" "1) metacoxal separation at middle relatively large (129, 0), uncontroverted." "Controverted synapomorphies:" "2) labium internal apodeme adjacent to paraglossal apodeme in dorsal view, from anterior to posterior, appearing V shaped (32, 1);" "3) visible portion of the first ventrite projecting between metacoxae distinctly truncate at apex (67, 0)."