taxonID	type	description	language	source
03DC6B6BFFEDFFC9FC2AF945FC77F89D.taxon	materials_examined	Type species: Onthophagus bituberculatus Olivier, 1789. Included species: At present five Afrotropical medium-sizes species (Figs S 2, S 3 – Appendix 4, frontal carina and simple vertex carina). The granulo-punctuate pronotum is not very convex, and has evident anterior angles with divergent apices that are inferiorly prolonged to a strong prosternal carina. Head and pronotum are usually black or dark brown. The elytra are flat, narrowed backwards, with marked striae and interstriae, extremely variable in coloration, ranging from evenly black to yellow with a discal dark spot. Sexual dimorphism was shown in the protibia (carrying a tooth on the inner margin only in males) and pygidium (far more developed in males than females), as is common in Onthophagini. Supporting Information) can be included in the genus Hamonthophagus gen. nov., namely: H. acutus (d’Orbigny, 1908), H. biturberculatus (Olivier, 1789), H. depressus (Harold, 1871), H. fallax (d’Orbigny, 1913) and H. laceratus (Gerstaecker, 1871). Diagnosis: The species included in the genus Hamonthophagus are strictly allied, and share a combination of characters that distinguishes them from the other Onthophagini: the anterior margin of the clypeus mostly sinuate and bidentate, and both head carinae poorly developed (i. e. slightly marked Epipharynx: The epipharynx (Fig. 2) is characterized by a rounded anterior margin gently notched in the middle, abundant and widespread acropariae, and well-developed corypha. The pubescence of the haptomerum is thick, the chaetopariae are almost rectilinear, constituted by short and dense setae. The anterior epitorma is longitudinal and narrow, the proplegmatium well sclerotized and arched, the apotormae are present, and the crepis is small, sharp and left-turned. The dexiotorma and laeotorma are slightly asymmetrical. Male genitalia: The male is characterized by a medium-sized phallotheca (or aedaegus, Fig. 7) with symmetrical parameres, and well-developed apices, carrying ventrally a symmetrical expansion. The membranous internal sac (or endophallus, Fig. 8) carries a hook-shaped and well-sclerotized primary sclerite, and some small accessory sclerites. Female genitalia: In females, an asymmetrical, well-sclerotized funnel-shaped area is evident in the vagina (Figs 9, 10), and is perhaps the most obvious character of the genus. The membranaceous and plurisinuate infundibulum is barely visible, being basally located at a very low position, just on the oviductus. The receptaculum seminis (Figs 9, 10) is curved in the distal third, more enlarged at base and tapering to an apex, with a large desclerotized area medially. Specific diagnosis: The Hamonthophagus species can be distinguished on the basis of some external features, such as the body pubescence, the elytral striae and the punctuation of pronotum and pygidium. Clear differences in shape were underlined by the geometric morphometrics analysis of head, pronotum, elytron and mentum (see above). Marked differences can also be highlighted by analysis of the epipharynx (always, according to the geometric morphometrics approach) and genitalia. The pubescence covering the body consists of thick, ochreous and truncated setae that are short in H. depressus, H. acutus and H. fallax, and longer in H. bituberculatus, while in H. laceratus the setae are very elongate, thinner and not truncated. The pronotum has a characteristic punctuation with varyingly sized, closely spaced, double points often carrying a hook-shaped granule never covering the point. While the points are usually dense (but excluding H. laceratus), in H. bituberculatus only the larger points bear the minute and flat granules, while in H. acutus, H. fallax and H. depressus, the majority of the evident points carry well-developed and thick granules. Hamonthophagus laceratus is characterized instead by sparse and superficial points, with very minute and scattered granules. The elytral striae are constituted by a very narrow line with larger points, except H. laceratus, where there are instead large striae with small points. Rasping, dense small setigerous points are present on the interstriae, and in H. acutus and H. bituberculatus the granules are small, while in H. depressus and H. fallax they are broader and evident. Again, in H. laceratus the points are rade, and almost inapparent, with few, very small granules. Hamonthophagus fallax usually carries an evident testaceous dot on the proximal sides of the elytra, and these dots are distally narrower than those in H. depressus. In H. acutus, the pygidium is covered by superficial points and evident, roundish and small granules, while in H. depressus and H. fallax the dense, large setigerous points are without granules on the disc, sometimes carrying rough points only on the sides. Besides, the latter species both have an evident and cerebroid microsculpture on the surface, which in H. acutus is less marked. The pygidium of H. bituberculatus has an opaque, smooth surface with few, scattered, shallow points (sometimes with minute granules), but an evident, very thick microsculpture. Also in H. laceratus, the pygidium is almost smooth, with an evident microsculpture, with only few and sparse points lacking granules. The fore margin of the epipharynx (Fig. 2) is only weakly notched in the middle in H. acutus, while in H. bituberculatus, H. depressus and H. fallax the notch is V-shaped, more marked and large. In H. laceratus the fore margin is slightly more squared than in the other species. The apotormae are less developed in H. bituberculatus and H. laceratus than in the other species. The crepis is more reduced in H. acutus and H. fallax. The medial triangular sclerotized area of the proplegmatium is far shorter in H. fallax than in the other species. In H. laceratus, the rear sclerotized part between the proplegmatium and crepis is much longer than in any other species. In males, the apices of parameres are elongate, large and only slightly hooked in H. acutus, H. bituberculatus and H. fallax, and more slender in H. depressus. In H. laceratus, the parameres of the aedeagus are narrower than in the other Hamonthophagus, rounded at the apex and slightly downcurved. The small, rounded ventral expansion is well developed mainly in H. laceratus (see Fig. 7 for a comparison among the species). The primary lamella of the endophallus is elongate with a large hook at the base in H. acutus, H. bituberculatus, H. depressus and H. fallax, with small differences in the longitudinal development among these species. In H. laceratus the primary lamella is more peculiar, being tougher and half as long as in the other species, but always hook-shaped (see Fig. 8 for a comparison among the species). These species can also be easily identified by the shape of the peculiar asymmetrical, funnel-shaped sclerotization of the vagina that shows a characteristic and differentiated development in the five species (see Fig. 10 for a comparison among the species). General remarks: No preimaginal stages have been described so far. Distribution: The genus Hamonthophagus is distributed in arid and savannah Afrotropical regions (Fig. 14). Etymology: The new genus was named after the Latin word hamo (= hook), with reference to the characteristic shape of the primary lamella of the internal sac.	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFEEFFC8FF37FF02FB50FD13.taxon	description	(FIGS 2, 7, 8, 10) Type material: NAMIBIA [Sud-Ouest africain allemand]: Okahandja [MNHN]. Paralectotypes: BOTSWANA: lake Ngami [MNHN]. DEMOCRATIC REPUBLIC OF CONGO: [Tanganyika,] region de Mpala [MNHN]. MALAWI: Malawi Lake [= Nyassa] [not located]. NAMIBIA: Salem [not located]. SOUTH AFRICA: Eastern Cape province [= Cafrerie] [MNHN]. For the morphological account, please refer to the original description. Geographical distribution (Fig. 14): The species distribution comprises Namibia, south-west Botswana and north-west South Africa (see Appendix 3, Supporting Information for a detailed list of the localities). Besides, in the type series d’Orbigny (1908) also included material from the Tanganika area (Democratic Republic of Congo) and Nyassa (i. e. Malawi). The former specimen was reported in the Collection Oberthur, MNHN, and the latter was reported in ‘ coll. du British Museum’ (now BMNH) where, however, it has not been traced (M. Barclay, pers. comm.). Neither specimen could be examined by us. As no other collection data from these areas were found within the studied material, these records were here regarded as uncertain until further confirmation. Also, a specimen from the MNHN labelled as ‘ Senegal provenance tres douteuse’ was not included in the present analysis. HAMONTHOPHAGUS BITUBERCULATUS (OLIVIER, 1789: 131) (FIGS 2, 7, 8, 10) Synonymy: Onthophagus discoideus (Olivier, 1789: 171) teste Harold 1880. Type material: At present, the typical material of H. bituberculatus could not be found. Although various materials of the Olivier collection were traced in several museum collections over the years (Bragg, 1996; Staines & Whittington, 2003; G ultekin € & Korotyaev, 2011), most specimens are still missing. The type material of this species was collected from ‘ Senegal’ by Geoffroy de Villeneuve, as well as its synonym O. discoideus (which was recorded also from Goree Island). As the type material of this species could not be located at present, no lectotype could be designed here. For the morphological account, please refer to the original description. Geographical distribution (Fig. 14): The species is widely distributed in the whole sub-Saharan area [Benin, Burkina Faso, Eritrea, Ethiopia, Gambia, Ghana, Guinea, Guinea Bissau, Ivory Coast, Mauritania, Niger, Nigeria, Senegal (the type locality), Sudan and Togo], and in Central and Eastern Africa (Central African Republic, Chad, Democratic Republic of Congo, Gabon, Kenya, Republic of Congo and Uganda], extending eastwards and southwards toward Tanzania and Malawi (see Appendix 3, Supporting Information for more details). The species is also recorded from Cairo (Egypt, Schatzmayr, 1946; Baraud, 1985) and Arabia (Paulian, 1980), but these data need to be confirmed. Accidental introduction is reported in Antilles (Martinique), where an anthropogenic cause was hypothesized to explain these findings (Matthews, 1966; Chalumeau, 1983).	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFEEFFD7FC0DFCCEFEE0FE20.taxon	description	(FIGS 2, 7, 8, 10) Synonymy: Onthophagus laceratus Peringuey 1901 nec Harold. Onthophagus carteri Blackburn, 1904: 147 teste Cartwright, 1938 depressus var. marmoreus d’Orbigny 1904: 309 Type material: Lectotype here designated: (male) SOUTH AFRICA: [Caffraria =] Eastern Cape province [ZMHB]. For the morphological account, please refer to the original description. Geographical distribution (Fig. 14): The species was originally described from South Africa, Caffraria (now Eastern Cape Province), but shows a wide distribution (the full list of the localities can be found in Appendix 3, Supporting Information) extending in a large part of the Afrotropical region (Angola, Botswana, Burundi, Democratic Republic of Congo, Kenya, Malawi, Mozambique, Namibia, South Africa, Tanzania, Zambia and Zimbabwe). Accidental introduction has been reported in Madagascar, Mauritius, USA (Florida, Georgia and South Carolina) and Australia (New South Wales and Queensland). Howden & Cartwright (1963) reported that specimens were collected at light in Georgia by Fattig. In the USA, H. depressus has been recorded in Georgia, south-west South Carolina and Florida (Hunter & Fincher, 1996; Hoebeke & Beucke, 1997; Evans, 2014), with a scattered distribution, since 1937 (Cartwright, 1938). The species was unintentionally introduced in Australia probably before 1900 (Matthews, 1972; Woodruff, 1973), when Blackburn (1904) described H. depressus specimens as a new species naming it O. carteri. This species was later properly identified as O. depressus by Arrow (see Cartwright, 1938). It is likely that the first introduction in Australia could be localized to the area near Sydney, from where it began to expand its range starting from 1941 (Matthews, 1972; Woodruff, 1973).	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF1FFD7FF07FDDDFEB8FB60.taxon	description	(FIGS 2, 7, 8, 10) Type material: Lectotype here designated: (male) MALAWI: [Nyassa Zomba haut Chire =] Zomba, Shire river upper course, Malawi Lake [MNHN]. Paralectotype: (female) TANZANIA [= Afrique Or Alem]: Dar-es-Salaam [MNHN]. For the morphological account, please refer to the original description. Geographical distribution (Fig. 14): The species has been described from Malawi and Tanzania, and at present is recorded from Botswana, Burundi, Democratic Republic of Congo, Kenya, Malawi, Namibia, Tanzania and Zambia. The record from Graaf-Reinet (Eastern Cape province, South Africa) is very interesting, but needs to be confirmed by further records. A detailed list of the localities is given in the Appendix 3 (Supporting Information).	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF1FFD7FE83FB12FDF1F89D.taxon	description	(FIGS 2, 7, 8, 10) Synonymy: Onthophagus laceratus subsp. benadirensis M uller € 1942: 82. Type material: Lectotype here designated: (male) TANZANIA: Zanzibar [ZMHB]. Paralectotype: (female) same locality [ZMHB]. The subspecies benadirensis from the Mogadishu area (Somalia) was examined and no marked differences were evident from the nominal species. For the morphological account, please refer to the original descriptions. Geographical distribution (Fig. 14): The species was described from Zanzibar, and shows a wide distribution extending to Burundi, Democratic Republic of Congo, Ethiopia, Kenya, Somalia, Sudan and Tanzania (see Appendix 3, Supporting Information for the full list of localities).	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF1FFD6FC18FF03FED2FB81.taxon	materials_examined	Type species: Onthophagus pallens d’Orbigny, 1908 Included species: M. pallens (d’Orbigny, 1908), and M. utete sp. nov. Diagnosis: Species of the genus Morettius (Fig. S 3 – Appendix 4, Supporting Information) are characterized by the mostly rounded and only slightly notched anterior margin of the clypeus, and the pronotum covered by granules, sometimes mixed to points. The pygidium is always smooth, with some rade points. The species show a moderate sexual dimorphism in the fore tibiae, and the pygidium is larger in males than in females. Epipharynx: The epipharynx (Fig. 2) fore margin is arched, with a largely V-shaped notch in the middle. The corypha is reduced, the chaetopariae are arched with short, thick and almost equal-length setae. The pubescence of the haptomerum is dense. The proplegmatium is subequal along the whole length, with the posterior triangular sclerotization reaching at least as much or more than half the length of the anterior epitorma, which is rectilinear, well sclerotized and thin. The base of the triangular sclerotization reaches the small, thick and upwardturned apophyses. Laeotorma and dexiotorma are symmetrical, short and stout. Pternotormae are well sclerotized, and the crepis is short but evident, with a sharp apex. The plegmatic area is visible. Male genitalia: Only the male genitalia of M. pallens could be examined. The phallobase (Fig. 7) is short, only slightly arched, slender and of equal size along the whole length. The parameres are symmetrical, and squared, with a small tip at the apex, and a small, rounded protrusion ventrally. The internal sac (Fig. 8) is membranous, with various well-sclerotized parts differing greatly from Hamonthophagus species. Female genitalia: The female genitalia are very peculiar (Fig. 11), as the vagina of both species is entirely membranous, and no sclerotization is present at all. Furthermore, the infundibular tube is lowered as in Hamonthophagus, but here an expanded portion is identifiable in the central part of the vagina, which is differently shaped in the two species. The receptaculum seminis is sickle-shaped, namely slim, arched and apically sharp (Fig. 9). Specific diagnosis: These species can be easily distinguished on the basis of external morphology, epipharynx and female genitalia. The pronotum in M. pallens is covered by distinct rasping points mixed with smaller, yellow granules; the granules of the rasping points instead are large, darker than the background surface, and carry long, thick and light yellow setae. The pronotum of M. utete is covered by only a few rasping setigerous points with thin, yellow setae and many small granules, which are very thick, evenly coloured as the base and without points. Elytral striae of M. pallens are as large as the points, being instead larger than the points in M. utete. The smooth pygidium carries in M. pallens few, small, rade and deep setigerous points that are not granulated, and in M. utete only some large and superficial vanishing points without setae. The epipharynx (see Fig. 2 for a comparison among species) has the characteristic shape of Morettius species, but can be distinguished from M. utete by the more developed apotormae, and the more slender laeotorma and dexiotorma. In both species, the vagina is wholly desclerotized, but carries two globose symmetrical expansions that encircle the desclerotized and lowered infundibulum in M. pallens, while in M. utete there is a single, large expansion (see Fig. 11 for a comparison among species). As the male of M. utete is unknown, no comparison can be made between species. General remarks: No preimaginal stages have been described so far.	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF0FFD6FC19FF03FD16FA96.taxon	description	(FIGS 2, 11, 17) Etymology: The species was named after the collection locality. Type material: Holotype: Female, TANZANIA: Utete-Rufijikindwjivi [MHNL]. Paratypes: 2 females, same locality [MHNL] [EBCT]. Description Male: Unknown. Female: Length: 6.1 – 6.6 mm. Head bronze, transverse (length / width ratio: 0.72), with maximum width just anteriorly to the eyes. Clypeus sinuate, with clypeal edge reddish. Clypeo-genal junction not sinuate. Frontal carina fine, weakly curved, placed at the mid-length of the head, short and low, occupying half of the interocular space. Surface markedly reticulate. Clypeus covered by flat and transverse granules, more or less merged. Genal granules large and round. Vertex unarmed, weakly concave, with round and fine granules. Antennal scape normally shaped, not dentate or serrulate. Antennal club yellow. Pronotum bronze, with hind angles bearing a bronze callus surrounded by a wide yellowish area Distribution: The genus Morettius is characterized by a disjoint distribution, being found in central west Africa, and south-eastern Africa (Tanzania). Etymology: The genus is named after our colleague, the French entomologist Philippe Moretto, who works extensively on African Scarabaeoidea.	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF0FFD5FF1DFA48FE91F989.taxon	description	(FIGS 2, 7, 8, 11) Type material: Lectotype here designated: (male) CHAD: Kiao-Kata, Moyen-Chari, south to Chad lake [= moyen Chari rives, Kiao-Kata] [MNHN]. Paralectotype: (female) same locality [MNHN]. For the morphological account, please refer to the original description. Geographical distribution (Fig. 14): The species was reported from Cameroon, Chad, Nigeria (southern border), Republic of Congo and Sudan (collection localities are listed in Appendix 3, Supporting Information). covering more than half the pronotal length and prolonged narrowly on the sides to reach the anterior angles. Pronotal pubescence black and very short, only evident on the lateral edges. Pronotum unarmed, very weakly transverse (length / width ratio: 0.55). Base evenly curved, markedly bordered. Posterior angles not sinuate. Anterior angles strongly sinuate, sharply projected outwards. Surface reticulate. Area surrounding the callus not granulate; remaining pronotal surface entirely covered by small granules. Elytra yellow with black symmetrical spots, one basal on the fifth interstriae, one on the first third of the sixth and seventh interstriae, another four connected on the middle of the second to fifth interstriae forming a zig-zig pattern. Juxtasutural interstriae yellow – orange, darkened anteriorly. Basal carina of interstriae bronze. Pubescence yellow, very short and scattered, only evident posteriorly. Elytral ground reticulate. Interstriae bearing small yellow granules, arranged on the juxtasutural interstriae in a regular row. Interstriae weakly convex, basally carinate. Striae narrow, well marked, yellow. Punctures of striae never wider than the striae. The seventh stria sinuate basally. Pygidium yellow almost smooth, finely microreticulate, with small, rare, hardly perceptible punctures. Base carinate. Epipleura yellow. Sternal thoracic surface bronze, except for the base of propleurae bronze. Abdominal sternites yellow. Metasternal pubescence scattered, short and yellow. Coxae yellow. Trochanters bronze. Femura yellow, apically bronze. Tibiae bronze, meso- and metatibiae apically yellowish. Tarsomeres weakly bronze. Pubescence yellow. Fore tibiae three-toothed. Tibial spur elongate, bent inward, apically rounded. Tarsi normally shaped. Individual variation: Paratype: the wide posterior spot of the right elytra is extended on the sixth interstria. Spot of the scond interstria obviously longer than in the hotolypus. Seventh elytral stria only weakly sinuate. Epipharynx (Fig. 2): See the above generic diagnosis.	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
03DC6B6BFFF3FFD5FC99FD42FA22F937.taxon	description	3. Pronotum black or dark brown with yellowish spots at the hind angles, and covered by sparse points, with very minute and rade granules .. ... ... ..... laceratus (Gerstaecker) 3 0. Pronotum entirely black or dark brown, and covered by varyingly sized, closely spaced, dense double points often carrying hook-shaped granules .. .... ... ... ... .... ... ... ... ... .... ..... 4 4. Pronotum with simple, small points and larger points with minute and flat granules. Setae of the dorsal surface obviously longer than wide .. ... .... .. bituberculatus (Olivier) 4 0. Pronotum covered by granulate points. Setae of the dorsal surface largely as long as wide ...................... 5 5. Pronotum evenly covered by granulate points. Pygidium covered by scattered but evident granules .......... acutus (d’Orbigny) 5 0. Pronotum covered by large granulate points and few, very smal simple points. Pygidium covered by rather large, more or less dense ocellate points ........................ 6 6. In males, apices of parameres elongate, large and only slightly hooked. In females, the sclerotized area asymmetrically developed with the apex on the right. Elytra black, with one or several testaceous, symmetrical, small patches .... .... ..... .... ... ... ... ... ... ... ... ... ..... fallax (d’Orbigny) 6 0. In males, apices of parameres more slender and pointed. In females, the sclerotized area well developed and triangular. Elytra usually entirely black or sometimes dark brown ........................... .. depressus (Harold)	en	Roggero, Angela, Dierkens, Michael, Barbero, Enrico, Palestrini, Claudia (2017): Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society 180: 298-320
