taxonID	type	description	language	source
03FB2F7EF40EFFDBFCF2FB045F72F8E6.taxon	description	A g e. Early Miocene according to Reynoso (2014) (originally correlated to the Late Oligocene). D i a g n o s i s. See Rasmussen and Gutiérrez (2009). D i s c u s s i o n. Mlanyama sugu differs from Anasinopa species by the broadening of the metastyle in the M 1 / and the reduction and narrowing of the isthmus linking the protocone to the buccal cusps. The m / 3 of M. sugu is very sectorial, with strong reduction of the talonid and absent metaconid, a cusp which is present but reduced in the other molars. Rasmussen and Gutiérrez (2009) related it closely to Dissopsalis pyroclasticus in particular because of the transverse orientation of the anterior premolars.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40EFFDAFC47F8CB5943FE09.taxon	description	D i a g n o s i s. Hyaenodontidae with elongated upper molars (M 1 – M 2), with strong protocone separated from the buccal cusps by a narrow isthmus, long in the type genus Dissopsalis, shorter in the other genera of the tribe, metacone better developed than the paracone, the two cusps separated from each other by a well-marked vertical groove. Lower dentition with very sectorial m 3 without a metaconid and a reduced talonid, m 2 with greatly reduced, even residual, metaconid, wide talonid with hypoconid and entoconid, m 1 with strong metaconid and wide accented talonid. Robust anterior premolars (P 4 and p 4). Anterior premolars not broadened. I n c l u d e d g e n e r a. Dissopsalis PILGRIM, 1912,	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40FFFDAFE9BFD975AFEFCDB.taxon	description	D i a g n o s i s. See Barry (1988). O t h e r s p e c i e s. Dissopsalis pyroclasticus SAVAGE, 1965.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40FFFDAFF7DFCA45F03FCCB.taxon	description	T y p e l o c a l i t y. Maboko, Kenya (erroneously given as Kaboor (error for Kabua = alternative name for Maboko )), Northern Frontier District, Kenya by Savage (1965). A g e. Basal Middle Miocene, ca. 15 Ma. D i a g n o s i s. See Savage (1965). D i s c u s s i o n. Barry (1988) cited several quite obvious differences between the lower dentitions of Dissopsalis carnifex PILGRIM, 1910 and Dissopsalis pyroclasticus. Among these were the more pronounced metaconid in the molars of D. pyroclasticus, the differences in length of the p 4 and m 3, and the taller crown and the diminutive talonid of the m 3 of D. pyroclasticus. To these differences could be added the greater size of the m 2 with respect to the m 3 in D. pyroclasticus. These differences, although important, do not exclude the possibility of the two species being classified in the same genus. Borths et al. (2016: supplementary tab. 2) included, but did not describe, additional upper and lower dentitions in this species, the study of which would, without doubt, help to clarify the relationship between the two species.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF408FFDDFCD0FA6F59DBF8A3.taxon	description	D i a g n o s i s. See Morlo et al. (2007). A g e. Basal Middle Miocene O t h e r s p e c i e s. Buhakia hyaenoides (MORALES et al., 2003), Arrisdrift, Namibia. Buhakia sp., Karungu, Kenya (Savage 1965), Buhakia sp., Grillental VI, Sperrgebiet, Namibia.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF408FFDDFF7AFF075F4DFA01.taxon	description	H o l o t y p e. Left maxillary fragment with M 1 – M 2 (NHMUK M 19083) (Text-fig. 1). T y p e l o c a l i t y. Rusinga, Kenya. A g e. Early Miocene. D i a g n o s i s. See Lewis and Morlo (2010). D i s c u s s i o n. As for Dissopsalis pyroclasticus, Borths et al. (2016: supplementary tab. 2) cited the existence of additional unpublished teeth, comprising both M 1 / M 2 occlusal shape: quadrangular (0), isosceles 1 triangle (1), rectangle (2), linear (3) 2 M 1 / M 2 protocone size: large (0), medium (1), reduced (2) M 1 / M 2 protocone isthmus: wide (0), narrow (1), 3 reduced (2) M 2 L / W ratio: <0.85 (0), between 0.86 – 1.15 (1), 4 between 1.16 – 1.35 (2),> 1.35 (3) M 1 L / W ratio: <0.85 (0), between 0.86 – 1.15 (1), 5 between 1.16 – 1.35 (2),> 1.35 (3) M 1 / M 2 protocone position: in front of parastyle / 6 paracone (0), anterior position to parastyle / paracone (1) M 2 metacone / paracone: partially separated (0), fused with 7 vertical separation groove (1), fused without separation groove (2) M 1 metacone / paracone: partially separated (0), fused with 8 vertical separation groove (1), fused without separation groove (2) 9 M 2 metastyle: small (0), short (1), long (2), very long (3) M 1 / M 2 paracone / metacone morphology: coniform (0), 10 sectorial (1) 11 M 1 / M 2 parastyle: present (0), reduced or absent (1) M 1 / M 2 buccal cingulum: present (0), reduced or 12 absent (1) 13 M 1 / M 2 buccal stylar area: present (0), reduced or absent (1) M 1 / M 2 paracone / metacone relative size: paracone 14 larger than metacone (0), paracone somewhat larger than metacone (1), metacone larger and taller than paracone (2) 15 M 1 / M 2 trigon valley: wide (0), narrow (1) 16 P 4 L / W ratio: <1.1 (0),> 1.1 (1) 17 P 3 W / L ratio:> 0.55 (0), <0.55 (1) 18 m 3 metaconid: present (0), absent (1) 19 m 3 W / L ratio:> 0.60 (0), <0.60 (1) 20 m 2 metaconid: present or residual (0), absent (1) m 2 talonid morphology: basined with hypoconid, 21 hypoconulid and entoconid (0), narrow with hypoconid in central position (1) 22 m 2 talonid size: well-developed (0), reduced (1) p 4 talonid morphology: basined (0), narrow, hypoconid in 23 central position (1) 24 p 3 W / L ratio: <0.65 (0),> 0.65 (1) 25 p 4 W / L ratio: <0.57 (0),> 0.57 (1) p 2 / p 3 orientation with respect to axis of mandible: 26 linear (0), transversal (1) upper and lower dentitions. The lingual wall of the M 2 in the holotype maxilla of the species is heavily worn, which partly obscures proper assessement of its characters, just as in a second individual, attributed to this species but which shows important differences from the holotype, rendering its identification doubtful. Therefore, it is not strange to find that its systematic position is erratic. It is possibly related to Isohyaenodon andrewsi (van Valen 1967, Dashzeveg 1985) or to Hyainailouros napakensis (Morales et al. 1998 a) or to the more hypercarnivorous species of the subfamily Hyainailourinae (Solé et al. 2014, Borths et al. 2016). Another possible correlation for Leakitherium hiwegi, not only morphological, but also dimensional, could apply to Dissopsalis pyroclasticus, which has a robust lower dentition. The two forms also share the presence of wrinkled enamel and in addition have posterior molars (M 2 – M 1 and m 3 – m 2) which are similar in length. This is highly divergent from the interdental proportions in Dissopsalis carnifex, in which the M 2 and the m 3 are noticeably larger than the M 1 and m 2. In fact, Morlo et al. (2007) indirectly suggested the possible generic separation of these taxa in their interpretation of Buhakia from Wadi Moghara, a new genus proposed by the authors, which they considered to be closer to D. carnifex than to D. pyroclasticus, but the separation was not formally proposed (Lewis and Morlo 2010). In their supplementary table 2, Borths et al. (2016) pointed out that the protocones in the holotype of Leakitherium hiwegi “ suggest larger talonid basins are expected on the lower dentition …. ”. This is possible as indicated above. Even though the holotype of Leakitherium hiwegi has a heavily worn lingual wall of the M 2, the impression is that the protocone would have been at least as well-developed as in the M 1. Another important feature is the strong groove between the paracone and metacone visible in the buccal side of the two cusps. The metacone is much better developed than the paracone in both molars, a condition also found in other taxa such as Dissopsalis carnifex. An analogous morphology, even though the teeth are more robust, occurs in the maxilla from Karungu described by Savage (1965) as Metapterodon kaiseri, which we here interpret as an indeterminate species of Buhakia, and therefore relatively close to Dissopsalis.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF408FFDCFC15F80C582CF846.taxon	description	Text-fig. 2 2003 Africanictis hyaenoides; Morales et al., p. 183. H o l o t y p e. Left m 2 (GSN AD 241 ’ 99) (Text-fig. 2). T y p e l o c a l i t y. Arrisdrift, Namibia. A g e. Basal Middle Miocene. D i s c u s s i o n. Morales et al. (2003) defined Africanictis hyaenoides at the locality of Arrisdrift (Namibia) on the basis of a suspected m 1 with specialised features (loss of the metaconid, narrow, cutting trigonid and reduced talonid, although it possesses a hypoconid and entoconid) thereby differing from the type species of the genus, Africanictis meini (Morales et al. 1998 b). Among the carnivores described by Morlo et al. (2007) from Wadi Moghra, Egypt, there was a new creodont, Buhakia moghraensis, based on a subadult mandible of which the dentition was interpreted to be dp 4, m 1 and m 2. In particular, the last molar in this specimen is morphologically quite close to the Arrisdrift species, which is only slightly larger (Text-fig. 3). The Arrisdrift molar is unworn, and has open roots, indicting its subadult status, similar in age to the individual of B. moghraensis. The Arrisdrift molar has a basal cingulum on the buccal side of the paraconid which sweeps upwards along the anterior margin and swells such that occlusion with the talonid of the preceding molar (m 1) is via this cingulum and the anteriormost extremity of the paraconid, a conformation visible in Buhakia moghraensis and frequent in hyaenodonts in which the molars are imbricated. Other characters common to the two species are the loss of the metaconid and the presence of cutting talonid and hypoconid.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF409FFDCFEE4F82A5F05FA79.taxon	description	2010 Metapterodon kaiseri Stromer; Lewis and Morlo, p. 551. L o c a l i t y. Karungu, Kenya. A g e. Early Miocene. M a t e r i a l. Maxilla with P 3 – M 3 (Savage 1965: textfig. 28, pl. 4, fig. 2, Lewis and Morlo 2010: fig. 26.5). D i s c u s s i o n. The species Metapterodon kaiseri was recorded by Savage (1965) at the locality of Karungu. The maxilla from this site differs from that of M. kaiseri (holotype from Elisabethfeld) and M. stromeri from Langental (Morales et al. 1998 a) by the scalene triangle occlusal outline of the M 2, in which the protocone-parastyle is quite a bit shorter than the length of the parastyle-metastyle. This is related to the lesser development of the isthmus which unites the protocone to the base of the paracone. The M 1 and P 4 of the holotype of M. kaiseri also have more developed and individualised protocones than the Karungu form, and in addition, the latter form possesses a stronger, very robust P 3, which contrasts with the extremely gracile P 3 of M. kaiseri (holotype; Pl. 2, Fig. 4). Like Morales et al. (1998 a) we conclude that none of the forms attributed by Savage (1965) to Metapterodon, belong to this genus. An alternative interpretation of the Karungu maxilla is that it could correspond to a form akin to Dissopsalis, a genus which has a peculiar admixture of characters, among which we emphasize the strong development of the protocone in the upper molars (M 1 – M 2), which agrees with the relatively strong development of the talonid of m 2. As we saw above, it is one of the few characters clearly observed in the genus Buhakia, which were described by Morlo et al. (2007) and Lewis and Morlo (2010). The dimensions of the Karungu maxilla suggest that it could correspond to the available lower dentition of Buhakia. Additionally, the Karungu maxilla falls outside the known range of variation of Dissopsalis carnifex, with which it shares the robustness of the premolars (P 3 – P 4), and the linear disposition of the buccal cuspids of the upper molars (M 1 – M 2), such that the metacone is more developed than the paracone.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF409FFDEFBABFA4759AFFDC5.taxon	description	L o c a l i t y. Grillental VI, Sperrgebiet (Namibia). A g e. Early Miocene. M a t e r i a l. Right mandible with the p 2 – m 1 and the alveolus of the m 2 and the anteriormost part of the alveolus of m 3 (GSN GT VI 22 ’ 17). Measurements p 3 (L = 9 mm, W = 4.5 mm), p 4 (L = 9.15 mm, W = 5 mm), m 1 (L = ca. 8.3 mm, W = ca. 4.2 mm), alveolus m 2 (L = 10 mm, W = 4.8 mm) (Pl. 1). D e s c r i p t i o n. GT VI 22 ’ 17 is a right mandible with the p 2 – m 1 and the alveolus of the m 2 and the anteriormost part of the alveolus of m 3. The front of the p 2 and the lingual side of the m 1 are broken. The talonid of the m 1 is relatively short but it would have been broad, with a strong, tall hypoconid. The p 4 is quite a bit bigger than the m 1 with a well-developed, tall main cusp. The anterior and posterior cristids are almost vertical and are well-marked, especially the anterior one. There is no anterior cuspid. The talonid is short but relatively broad, and the hypoconid is strong and occupies the buccal half of the talonid, and a low crest forms the lingual border. A moderate cingulum is irregularly developed around the premolar, more marked on the anterior part of the tooth. The p 3 is almost as large as the p 4 but is lower with less vertical cristids. Like the p 4 it has no anterior cuspid and the talonid is short, almost reduced to a posterior cingulum. The p 2 has lost the anterior extremity but its basic morphology seems to be like that of the p 3, and it is appreciably smaller. The two anterior premolars overlap and are oriented obliquely with respect to the p 4. The mandibular ramus is deep and the symphysis very rugose, with its posterior margin at the level of the posterior root of the p 3. On the buccal side of the jaw there are two foramina visible at half the height of the jaw beneath the gap between the p 3 and p 4. Of these the anterior one is strong and oval, the posterior one small and circular. D i s c u s s i o n. The premolars of the Grillental VI jaw are robust and big, the two anterior premolars (p 2 and p 3) being only slightly smaller than the p 4. The p 4 is quite peculiar because of its tall main cusp with almost vertical, concave anterior cristid, and the absence of a paraconid. In effect, it is quite close in morphology to the p 4 of Dissopsalis carnifex (Barry 1988: fig. 2) and Dissopsalis pyroclasticus (Savage 1965: pl. 3), even though the tooth is smaller than in these two species. In contrast if we attribute the p 2 or p 3 to D. carnifex (GSP 16036; Barry 1988: fig. 3) the differences from the Grillental specimen would be important. Unfortunately, in D. pyroclasticus the anterior premolars are not preserved although the alveoli suggest that they would have been large relative to the p 4. It is possible that the p 3 attributed to Dissopsalis carnifex mentioned above could be a P 3 as it is morphologically close to the P 3 in the maxilla from Karungu, here identified as Buhakia sp., sharing the presence of a weakly developed posterior cuspid, and above all a characteristic postero-buccal cingulum or basal swelling. Some information can be obtained from Anasinopa leakeyi (Savage 1965) in which the p 4 possesses a structure similar to that described in Dissopsalis, although it differs from it by the lower height of the main cusp and the less vertical anterior cristid, as well as the greater development of the talonid. The p 2 and p 3 of D. pyroclasticus have a more usual morphology similar to that in the Grillental VI specimen, even though, at least in the p 3 the main cusp is lower than in the Namibian form. In conclusion, the mandible from Grillental VI has several features more specialised than in Anasinopa leakeyi, a characteristic of the tribe Dissopsalini, as we point out in this paper. The dimensions of the Grillental VI specimen are smaller than Dissopsalis or Leakitherium hiwegi, and could correspond closely to Buhakia (Morales et al. 2003, Morlo et al. 2007). The alveoli of the m 2 in GT VI 22 ’ 17 measure about 11 mm, which suggests that the tooth would have been close in dimensions to the m 2 of B. moghraensis or B. hyaenoides. However, in this genus the lower premolars are unknown, and as mentioned previously, they are poorly represented in the larger genus Dissopsalis. But, we can have a reasonable idea about the characters of the premolars in this group that we call Dissopsalini which are remarkably divergent from those of the very bunodont genus Teratodon, approaching the hyaenid pattern, tall and robust premolars. This morphological pattern is clearly recognisable in the mandible from Grillental VI, from which we deduce that its inclusion in Dissopsalini is the most plausible hypothesis. In contrast, it differs in size from the species of Dissopsalis and Leakitherium, and is closer to species of Buhakia, which is for the moment, the only clear representative of this size in this tribe. For these reasons, and in the hope of finding new material in the Early Miocene sites of the Sperrgebiet, Namibia, we prefer to classify this fossil as Buhakia sp., following the same reasoning that we used for the controversial maxilla from Karungu. Nevertheless, some doubt remains concerning the specific identification of the two fossils.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40BFFDEFC5CFDAB5F6DFC75.taxon	description	D i a g n o s i s. Hyainailourinae with sectorial upper molars (M 2 – M 1) with the paracone and metacone subequal in size and fused together, nevertheless with a moderate vertical groove between the two cusps. It has a strong protocone united to the rest of the tooth by a long, narrow isthmus. Lower molars with metaconid, with unicuspidate talonid which is very reduced in the m 3. Premolars gracile. I n c l u d e d g e n e r a. Type genus only.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40BFFD1FC2EFC5B5E36FE78.taxon	description	1985 Pterodon Blainville, 1839; Dashzeveg, p. 234. T y p e s p e c i e s. Metapterodon kaiseri STROMER, 1926 (with type locality: Elisabethfeld, Sperrgebiet, Namibia). Holotype is left maxillary fragment with M 2 – P 3 (BSPG Nr. 1926 X 1). A g e. Early Miocene. D i a g n o s i s. Metapterodon, being the only known genus in the tribe, has the same diagnosis as the tribe. O t h e r s p e c i e s. Metapterodon stromeri MORALES, PICKFORD et SORIA, 1998 from the Early Miocene of Langental, Sperrgebiet, Namibia.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF40BFFD1FC2EFC5B5E36FE78.taxon	description	A g e. Early Miocene. M a t e r i a l. Right M 2 (L = 12.8 mm, W = 12.2 mm) (NAP V 121 ’ 08). D e s c r i p t i o n. The occlusal outline is an obtuse isosceles triangle, in which the two equal sides are formed of the posterior surface (protocone-parastyle) and the buccal surface (parastyle-metastyle), while the unequal side is longer and extends from the paracone to the metastyle. Outstanding is the strong development of the protocone, in the shape of a pointed clog, which extends well forwards. The parastyle is strong and basally is contiguous with the moderately well-developed buccal cingulum. The paracone and metacone are largely fused together, but are slightly separated at their apices by a shallow incision on the buccal surface and are transversely compressed. The metastyle is similar in size to the paracone-metacone. The lingual cingulum is weak. D i s c u s s i o n. The occlusal morphology and the great development of the protocone (Pl. 2, Figs 1 – 2) approach this molar to that of Metapterodon kaiseri STROMER, 1926 and Metapterodon stromeri MORALES et al., 1998 a. It differs from the former by its greater dimensions (ca. 10 %) and by the greater development of the parastyle. It differs from M. stromeri by its smaller dimensions (ca. 15 %; Text-fig. 4). Holroyd (1999) included three species from the Eocene-Oligocene of the Fayum, Egypt, in Metapterodon, but she raised reasonable doubts concerning the generic attribution. In particular, in the two species represented by upper dentitions (and thus comparable to the holotype of Metapterodon kaiseri) there are important differences, noted in the descriptions by Holroyd (1999: 12, 14). According to this author, the molars of M. schlosseri and Metapterodon markgrafi are strongly modified for cutting, with a small protocone in the first species and none in the second, and in addition, the two species possess long metastyles. These differences prevent the use of Metapterodon as a possible genus for these Palaeogene forms. In contrast, as we will see later, the species could be related to Isohyaenodon, an idea indirectly supported by Savage (1965) who included in I. andrewsi one of the mandibles later identified as M. schlosseri by Holroyd (1999).	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF404FFD0FC35FE48586BFE7A.taxon	description	D i a g n o s i s. Hyainailourinae with sectorial upper molars (M 2 – M 1) with the paracone somewhat larger than the metacone, generally fused together, some genera retain a moderate vertical sulcus between the two cusps, but which can be completely lost. Protocone is in general reduced, in some genera it is relictual or even absent. Lower molars without metaconid and with talonid very reduced or absent. I n c l u d e d g e n e r a. Miocene: Hyainailouros BIEDERMANN, 1863, Isohyaenodon SAVAGE, 1965, Sectisodon gen. nov. and Exiguodon gen. nov. Eocene (after Solé et al. 2015): Akhnatenavus HOLROYD, 1999, Kerberos SOLÉ et al., 2015, Parapterodon LANGE- BADRÉ, 1979, Pterodon BLAINVILLE, 1839, Falcatodon gen. nov.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF405FFD0FF70FE465A85FD0C.taxon	description	1863 (with type locality Veltheim, Switzerland). D i a g n o s i s. See Ginsburg (1980). A f r i c a n s p e c i e s. Hyainailouros napakensis GINSBURG, 1980 and Hyainailouros osteothlaste s (SAVAGE, 1973).	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF405FFD0FF47FC9358D2FA31.taxon	description	Text-fig. 5 1965 Pterodon africanus Andrews; Savage, p. 272. 1965 Pterodon nyanzae sp. nov.; Savage, p. 274. 1999 Pterodon nyanzae Savage; Holroyd, p. 7. 2009 Hyainailouros sp.; Rasmussen and Gutiérrez, p. 35. 2010 Hyainailouros napakensis Ginsburg; Lewis and Morlo, p. 547. 2010 Hyainailouros nyanzae Savage; Lewis and Morlo, p. 547. H o l o t y p e. Left maxilla with P 4 – M 2 (NHMUK M 19090) described and figured by Savage (1965). T y p e l o c a l i t y. Napak I, Uganda. A g e. Early Miocene. D i a g n o s i s. See Ginsburg (1980). O t h e r l o c a l i t i e s. Ombo, Rusinga, Kenya (Savage 1965), Losodok, Meswa Bridge, Kenya (Rasmussen and Gutiérrez 2009). With reserve, given the incomplete nature of the material, we may include the localities of Grillental and Elisabethfeld, Namibia (Morales et al. 2007).	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF405FFD5FF70F99E5846FA83.taxon	description	Text-fig. 6 1965 Pterodon nyanzae sp. nov.; Savage, p. 274. L o c a l i t y. Arrisdrift, Namibia. Morales et al. (1998 b) and Morales et al. (2003) referred an M 1 (Text-fig. 5 c), an upper canine and a mandible with m 1 and unerupted molar (Text-fig. 6) to this species. A g e. Basal Middle Miocene.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF405FFD5FF70F99E5846FA83.taxon	description	1947 Hyaenaelurus fourtaui; Koenigswald, p. 292. 1973 Megistotherium osteothlastes; Savage, pp. 485 – 486. 1989 Megistotherium osteothlastes Savage; Rasmussen et al., p. 443. 2007 Hyainailouros bugtiensis Pilgrim; Morlo et al., p. 147. 2010 Megistotherium osteothlastes Savage; Lewis and Morlo, p. 550. H o l o t y p e. Skull (NHMUK M 26173). T y p e l o c a l i t y. Gebel Zelten, Libya. D i a g n o s i s. See Savage (1973). A g e. Basal Middle Miocene. O t h e r l o c a l i t i e s. Wadi Moghara, Egypt (Fourtau 1920, Koenigswald 1947, Rasmussen et al. 1989, Morlo et al. 2007); Cheparawa, Muruyur Formation (ca. 14.5 Ma), Tugen Hills, Kenya (Morales and Pickford 2008); Fort Ternan, Kenya (ca. 13.7 Ma) (Savage 1973); Bartule, Ngorora Formation, Member A (ca. 13 – 12.5 Ma) and Kabarsero, locality 2 / 10, Ngorora Formation, Member D (ca. 12 Ma), both Kenya (Morales and Pickford 2005). D i s c u s s i o n. Hyainailouros has a long and complicated history of study, influenced by two opposing aspects; on the one hand, it is quite rare in the fossil record, and on the other, its gigantic size makes it easily recognisable even with little material. The sample of the type species described from the Swiss locality at Veltheim by Biedermann (1863) was revised by Helbing (1925), to which he added other fossils from Europe and Asia. Later, Beaumont (1970) revised the fossils from the type locality, and fossils from Chevilly Aérotrain, France, were added to the hypodigm (Ginsburg 1980) which are the most complete dental remains associated with elements of the postcranial skeleton. Two additional Miocene genera were attributed to Hyainailouros on the basis of their large dimensions and their resemblance to this genus; the first was Megistotherium SAVAGE, 1973 and the second was Sivapterodon erected by Ginsburg (1980) for the species Hyainailouros lahirii PILGRIM, 1932. The latter species is known only by a fragment of mandible with m 2 – m 3 close in size to Hyainailouros sulzeri. According to its creator, the genus is based on the greater reduction of the paraconid in the molars (m 2 – m 3). However, the two molars are poorly preserved “ both of them considerably worn and damaged in their anterior halves ” (Pilgrim 1932: 170) which prevents us from taking this character seriously, and the same applies to the apparent robusticity of the m 3, which could have been overestimated due to the damage to the anterior part of the crown. The great reduction of the talonid of the m 3, which is almost obsolete, is comparable to that in the fossils of H. sulzeri from Chevilly Aérotrain. In the current state of our knowledge, it is difficult to sustain the validity of Sivapterodon.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF405FFD5FF70F99E5846FA83.taxon	description	The opposite extreme of the size spectrum is occupied by the huge species Megistotherium osteothlastes. The upper dentition is known only from the holotype skull, which, as mentioned previously, is incomplete and damaged, measurements being obtained from the roots or alveoli. However, the specimen indicates that the teeth were large, much longer than other specimens of Hyainailouros (Text-fig. 7). Lower teeth attributed to this species were published by Rasmussen et al. (1989) and Morales and Pickford (2005) from Wadi Moghara, Egypt, and Ngorora, Kenya, respectively. The differences in dimensions from the maximal values of H. sulzeri are not great, but they fall outside the range of variation of this species. It is possible that the metric differences could correspond to extreme intra-specific variability, within which the specimens of H. osteothlastes would be the largest (perhaps males with hypertrophic canines), but it is also possible that, without denying the existence of dimorphism or great variability, there could be two or three species with overlapping dimensions. Morphological comparisons do not resolve the problems because of the meagre quality of the teeth of H. osteothlastes, impossible to observe in the upper dentition, and very limited for the lower teeth. The mandible from Wadi Moghara described by Rasmussen et al. (1989) has an m 3 missing the posterior part, and an m 2 with a reduced talonid, similar to the m 2 of H. sulzeri and to an m 2 from Ngorora (Morales and Pickford 2005).	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF400FFD5FE9CFAED58E8F873.taxon	description	1999. D e r i v a t i o n o m i n i s. From “ falcate ”, the cutting sword used by the Iberians. D i a g n o s i s. Medium sized Hyainailourinae, differing from Metapterodon by the reduction of the protocone in the upper molars (M 1 – M 2), which is located in a very anterior position. It differs from Hyainailouros by the more sectorial morphology of the upper molars, with more advanced fusion of the paracone-metacone with, nevertheless, a groove separating the cusps visible in the M 1. Lower molars sectorial with reduced talonid and without a metaconid. It differs from Isohyaenodon, Sectisodon and Exiguodon, by the lesser reduction of the protocone in the upper molars which, above all, retain a stretched out subtriangular occlusal outline.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF400FFD5FF41F85C5EFBFD68.taxon	description	p 2 – m 3 (DPC 4877). T y p e l o c a l i t y. Quarry V, upper sequence of Jebel Qatrani Formation, Fayum Province, Egypt. A g e. Rupelian (Seiffert 2010). D i a g n o s i s. The same as for the genus. D i s c u s s i o n. Holroyd (1999) recognized that “ although the Eocene-Oligocene species referred here to Metapterodon may ultimately prove to be a seperate genus from M. kaiseri, I have here united them in an attempt to demonstrate their distinctiveness from both Hyaenodon and Pterodon, and their probable closer relationship to one another than to contemporaneous species of Pterodon ”. The weaker development of the isthmus between the protocone and paracone and the more distal position of the protocone in the upper molars of Falcatodon schlosseri are closer to Isohyaenodon zadoki than to Metapterodon spp. but in any case, are different from both genera.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF400FFD7FC22FD785838FEA5.taxon	description	1999 Metapterodon Stromer; Holroyd, p. 11. T y p e s p e c i e s. Isohyaenodon andrewsi SAVAGE, 1965 (holotype: right mandible with m 1 – m 3 (M- 15048); type locality: Ombo, Kenya). D i a g n o s i s. See Morales et al. (1998 a). O t h e r s p e c i e s. Isohyaenodon zadoki SAVAGE, 1965 (= Isohyaenodon matthewi SAVAGE, 1965) and Isohyaenodon sp. (Morales et al. 2008). N. B. Part of Savage’s (1965) hypodigm of Isohyaenodon matthewi (CMF 4060 NHMUK M 2947), left m 2 in a mandible fragment, fits perfectly onto a specimen attributed by the same author to Isohyaenodon andrewsi (CMF 4023 NHMUK M 2948) a left m 3 in a mandible fragment. Both fragments came from site R 3, Rusinga Island and represent a single individual. D i s c u s s i o n. Soon after its creation, this genus, as was mentioned in the introduction, was the subject of discussion by van Valen (1967) and subsequently by Morales et al. (1998 a, 2007) and Lewis and Morlo (2010). The holotype mandible is similar in dimensions (at least the m 3) to Metapterodon stromeri from the locality of Langental (Morales et al. 1998 a). However, there are morphological reasons for separating the two species, I. andrewsi possesses a slightly more sectorial m 3, with a moderate tendency to enlarge the protoconid. The two species share the absence of the metaconid and the greatly reduced talonid in the m 3. But in M. stromeri the m 2 retains a more developed talonid. Reasonably, the mandible of I. andrewsi, which shows a greater sectorial tendency, could correspond to the upper dentitions more sectorial than the Metapterodon species. Borths et al. (2016: supplementary table 2) support the speculation that Isohyaenodon andrewsi and Metapterodon represent the lower and upper dentition of the same taxon based on the size of the occluding carnassial portion of the upper and lower dentitions. However, very sectorial lower teeth are known in Africa from the Late Eocene in the Fayum succession, the case with the species previously identified as Metapterodon schlosseri by Holroyd (1999). This was realised by Savage (1965), when he included in Isohyaenodon one of the Fayum mandibles described by Andrews (1906). The upper teeth of this species are known from a single maxilla (Holroyd 1999: fig. 8 B) which differs from species of Metapterodon by the reduction of the protocone and the extension of the metastyle in the M 1 – M 2 (Text-fig. 8 k). This morphological pattern can be seen in the form described by Savage (1965: text-fig. 29, pl. 4, fig. 2) as Metapterodon zadoki from the site of Rusinga, Kenya, which could correspond to the lower dentition of the species Isohyaenodon matthewi from the locality of Songhor, Kenya (Savage 1965: text-figs 41 – 43). Apparently, M. zadoki seems to have progressed further in the reduction of the protocone of the molars, as was noted by Savage (1965), which are also more robust and without a parastyle (Pl. 3, Fig. 1). To a certain extent, these characters approach Isohyaenodon to the new genus Sectisodon gen. nov., but the difference in robustness of the M 1 is more than noticeable. It is evident that the solution is not easy, as there exists a whole range of morphologies in the upper teeth which do not correspond directly to the sectorial lower teeth. Thus, we propose to restrict the species content of Isohyaenodon to I. andrewsi and Isohyaenodon zadoki (= I. matthewi) from Kenya, as well as Isohyaenodon sp. (Pl. 3, Fig. 2) from Elisabethfeld, Namibia (Morales et al. 2008) in the hope that new fossil finds may clarify the characteristics of this genus.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF402FFD7FC4BFAAC5F4EF874.taxon	description	SAVAGE, 1965. D e r i v a t i o n o m i n i s. From the Latin “ exiguus ” meaning exiguous. D i a g n o s i s. Hyainailourinae of diminutive dimensions, lower molars (m 3 – m 2) with greatly reduced talonid; protoconid and paraconid similar in size. Paraconid of the molars lingually oriented. M 2 and M 1 close in size and morphology. Occlusal outline sub-triangular, with greatly reduced protocone, which appears like an antero-lingual cingulum which extends anteriorly and buccally. Presence of a strong parastyle in an antero-buccal position, united to the apex of the paracone by a well defined crista. The buccal cingulum borders a wide buccal platform, particularly large in the M 2. Paracone tall and narrow, elongated blade-like metastyle. P 4 broadened, with reduced protocone and presence of a notch between the main conical cusp and the blade-like posterior cusp.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF402FFD7FC0CFD395EA2FAC5.taxon	description	1965 Metasinopa (?); Savage, p. 264. 1999 Metapterodon markgrafi; Holroyd, p. 11. H o l o t y p e. Left maxilla containing alveoli for C – Pl, roots of P 2 – P 3, partial P 4, Ml and partial M 2 (AMNH 14452). T y p e l o c a l i t y. Fayum, Egypt. D i a g n o s i s. See Holroyd (1999). D i s c u s s i o n. As defined by Holroyd (1999) Sectisodon markgrafi possesses highly sectorial molars with complete fusion of the paracone-metacone and strong reduction of the protocone, which is nevertheless visible in occlusal view. It is quite a bit larger than the Napak species, and the greater development of the protocone in the upper molars differentiates it from S. occultus.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF402FFD7FE80FE0C5B17FBCF.taxon	description	D e r i v a t i o n o m i n i s. From the Latin “ sectis ” meaning cutting. D i a g n o s i s. Hyainailourinae of small dimensions, M 1 and M 2 of similar length. M 2 with paracone and metacone fused together, tall and pointed. Protocone very reduced, flattened and extending basally beyond the base of the paracone. Anterior cingulum with a well-defined parastyle with a moderate buccal cingulum. M 2 and M 1 with the metastyle quite a bit longer than the paracone-metacone. P 3 short, with tall main cusp, posterior cusplet moderate, and much reduced protocone. Basal lingual cingulum strong, weaker on the buccal side. Lower molars with protoconid bigger than the metaconid, m 2 with talonid present, much reduced in the m 3. O t h e r s p e c i e s. Sectisodon markgrafi (HOLROYD, 1999), Fayum, Egypt.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF402FFD7FE9EFBD15EA2FD56.taxon	description	T y p e l o c a l i t y. Napak V, Uganda. A g e. Early Miocene. O t h e r l o c a l i t i e s. Napak I and Napak IV, Uganda. D e s c r i p t i o n. NAPIV 72 ’ 04, right P 3 (L = 7.75 mm, W = 5.46 mm, H paracone = 7.01 mm). This is a narrow premolar with the main cusp tall and sharp. There is a small posterior cusplet and a well-marked basal cingulum completely encircling the tooth. The protocone is reduced to a lingual bulge in the middle of the tooth joined to the cingulum and quite a bit deeper than the base of the main cusp. The morphology is analogous to that seen in the protocones of the molars. NAP IV 01 ’ 09, right M 1 (L = 8.1 mm, W = 5.58 mm, H. paracone = 6.19 mm, mestastyle length = 3.63 mm). Short molar with tall cusps. Paracone and metastyle of similar size. The protocone is greatly reduced, flattened and extending low down beyond the base of the paracone. Reduced parastyle included in the anterior cingulum. Basal cingula almost imperceptible. The molar has a vertical wear facet on the lingual surface of the paracone and the metastyle (Pl. 4, Fig. 2 a). NAP V 178 ’ 08, right M 2 (L = 8.57 mm, W = 4.32 mm, H paracone = 5.52 mm, Mestastyle length = 3.85 mm). This specimen is morphologically similar to the preceding one, similar in length, but narrower and with lower cusps. The parastyle and protocone seem to be quite big and the metastyle elongated. There is a small interstitial facet at the posterior part of the base of the metastyle (Pl. 4, Fig. 3 a) probably produced by contact with a small M 3. D i s c u s s i o n. Sectisodon represents a more advanced sectorial adaptation than other hyaenodonts from the Miocene of Africa, with the exception of Exiguodon. The upper molars (M 1 – M 2) almost lack protocones, which differentiates them from those of Metapterodon, in which the M 2 has a hyperdeveloped protocone. With respect to Exiguodon described in this work, apart from the reduction of the protocone in the upper molars, the morphological differences are important, in particular the strong development of the cingulum and the buccal platform that occurs in this genus.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
03FB2F7EF402FFD6FC22F85C5838F8B4.taxon	description	1965 Hyaenodon (Isohyaenodon) pilgrimi; Savage, p. 284 1998 Isohyaenodon pilgrimi Savage; Morales et al., p. 636. 2007 Isohyaenodon pilgrimi Savage; Morales et al., p. 72. 2010 Isohyaenodon pilgrimi Savage; Lewis and Morlo, p. 548. H o l o t y p e. Mandible with both rami and part of the neurocranium attached to the seven cervical vertebrae (NHMUK M 19100 a-c; Pl. 5). T y p e l o c a l i t y. Site R 114, (Whitworth’s Pothole), Rusinga Island, Kenya. A g e. Early Miocene. D i a g n o s i s. The same as for the genus. O t h e r l o c a l i t i e s. Songhor and Koru, Kenya and Napak IV, Uganda. D e s c r i p t i o n o f K o r u f o s s i l s. KO 38 ’ 04 (Pl. 6, Fig. 1) is a right mandible with m 3 (L = 4.7 mm, W = 2.3 mm) and m 2 (L = 4.4 mm, W = 2 mm). In the m 3 the paraconid and protoconid are robust, sub-equal in size and morphology. The paraconid is located in quite a lingual position, such that the molar appears to be inclined with respect to the antero-posterior axis of the dental series. There is no sign of a metaconid, and the talonid is reduced to a small tubercle at the base of the crown in a posterior-lingual position. The notch that separates the paraconid from the protoconid is deep. In the paraconid a basal vertical tubercle develops on the antero-buccal surface that serves to lodge the talonid of the m 2. The buccal surface shows a vertical wear facet, while lingually a deep valley is developed at the base of the paraconid-protoconid. The m 2 is somewhat worn anteriorly, and is smaller than the m 3, although its morphology is similar. The talonid is quite well-developed and possesses a small sharp hypoconid.	en	Morales, Jorge, Pickford, Martin (2017): New Hyaenodonts (Ferae, Mammalia) From The Early Miocene Of Napak (Uganda), Koru (Kenya) And Grillental (Namibia). Fossil Imprint 73 (3 - 4): 332-359, DOI: 10.2478/if-2017-0019, URL: http://dx.doi.org/10.2478/if-2017-0019
