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The narrow-headed vole, collared lemming and common vole were the most abundant small mammal species across the Eurasian Late Pleistocene steppe-tundra environment. Previous ancient DNA studies of the collared lemming and common vole have revealed dynamic population histories shaped by climatic fluctuations. To investigate the extent to which species with similar adaptations share common evolutionary histories, we generated a dataset comprised the mitochondrial genomes of 139 ancient and 6 modern narrow-headed voles from several sites across Europe and northwestern Asia covering approximately the last 100 thousand years (kyr). We inferred Bayesian time-aware phylogenies using 11 radiocarbon-dated samples to calibrate the molecular clock. Divergence of the main mtDNA lineages across the three species occurred during marine isotope stages (MIS) 7 and MIS 5, suggesting a common response of species adapted to open habitat during interglacials. We identified several time-structured mtDNA lineages in European narrow-headed vole, suggesting lineage turnover. The timing of some of these turnovers was synchronous across the three species, allowing us to identify the main drivers of the Late Pleistocene dynamics of steppe- and cold-adapted species.

Cynelos anubisi n. sp. (Fig. 2) Cynelos sp. nov. – Morlo et al. 2007: fig. 4G, H. cf. Ysengrinia ginsburgi – Morales et al. 2010: 48. Afrocyon burolleti – Morales et al. 2010: fig. 3. Amphicyon sp. – Morales et al. 2010: fig. 4. Cynelos sp. – Werdelin & Peigné 2010: 604. Afrocyon n. sp. – Morales et al. 2016: 143. HOLOTYPE. — CUWM 55, left mandible with alveolus of c, single root of p1, single root of p2, roots of p3, p4-m2, and alveolus of m3. PARATYPE. — DPC 14532 /2 L-7, isolated left m1 from Moghra, Egypt (Morlo et al. 2007: fig. 4G, H). REFERRED MATERIAL. — MNCN 79042 right mandible fragment with alveolus of p1, single-rooted p2, fragment of p3, and p4 from Gebel Zelten, Libya (= BIZ.2A. 15 in Morales et al. 2010: fig. 4). NHM M 82374, an isolated left m2 from Gebel Zelten, Libya (Morales et al. 2010: fig. 3). TYPE LOCALITY. — Wadi Moghra, Egypt. DISTRIBUTION. — North Africa: Egypt and Libya. AGE AND HORIZON. — Late Early Miocene, contemporaneous with European biozone MN4. ETYMOLOGY. — “ anubisi ” after “Anubis”, the Greek name for the Ancient Egyptian jackal-headed god, who watched over the dead. DIFFERENTIAL DIAGNOSIS. — Cynelos anubisi n. sp. differs from Amphicyon and Afrocyon in having a single-rooted p1 and p2, a long diastema between p2 and p3, and a short diastema between p3 and p4. The m1 metaconid of C. anubisi n. sp. is more reduced, the hypoconid is smaller, and the m1 and m2 talonids are narrower. The m2 protoconid and hy- poconid are low but massive. Cynelos anubisi n. sp. further differs from Amphicyon in having a narrower m2, as a result of an elongated talonid. All molars lack cingulids. Further differs from Afrocyon in having a single-rooted m3, and a coronoid process that rises at a shallower angle. Cynelos anubisi n. sp. is about the same size as, or is slightly smaller than C. macrodon, but C. anubisi n. sp. differs from C. euryodon and C. minor in being much larger in size, and differs from the Buluk Cynelos in being much smaller. Cynelos anubisi n. sp. differs from C. euryodon and C. lemanensis in having a single rather than a double rooted p2, and a much shorter p3-p4 diastema (see Peigné & Heizmann 2003; Morales et al. 2016); differs from C. macrodon and C. ginsburgi n. comb. in having m1 with a longer paraconid blade relative to overall tooth size; further differs from C. macrodon in having the m1 paraconid oriented slightly more anteriorly, having p4 smaller, and in possessing small p4 pre- and postprotoconulids; and further differs from C. ginsburgi n. comb. in having p4 and m2 longer relative to m1, m1 broader the m1 paraconid less angled but relatively longer, and the metaconid is located slightly more anteriorly. Cynelos anubisi n. sp. is larger than any Eurasian Cynelos species, including C. bohemicus (Schlosser, 1899) which is known from Europe and Asia during biozones MN 3 to MN 5 (Jiangzuo et al. 2018). Cynelos anubisi n. sp. also clearly differs from C. bohemicus in possessing a large diastema between p2 and p3 that is absent in this Eurasian species (see Fejfar & Heizmann 2016: fig. 9.5). Additionally, the generic assignment of material representing C. bohemicus is controversial. Schlosser (1899) originally placed the taxon in the genus Pseudocyon, although more recent contributions have discussed the material as belonging to Amphicyon (Fejfar & Heizmann 2016, following Kuss 1965) or Cynelos (Jiangzuo et al. 2018, following Hunt 1998; Hunt & Stepleton 2015). DESCRIPTIONS AND COMPARISONS In the holotype mandible, CUWM 55, p4, m1, and m2 are present, while the canine, p1-p3, and m3 are represented by alveoli. Judging from the alveoli, C. anubisi n. sp. had a single-rooted p1 and p2, and a small, double rooted p3, with a long diastema between p1-p2, longer than that observed in MNCN 79042 from Gebel Zelten. An alternative possible interpretation is that p 1 in C. anubisi n. sp. is reduced, and the p2 is double-rooted, but this is not the condition observed in MNCN 79042. In this respect, C. anubisi n. sp. clearly differs from the other Moghra amphicyonid CUWM 53, the Gebel Zelten specimen NHM M 82373, Amphicyon giganteus from Arrisdrift (Morales et al. 2003), and Afrocyon burolleti (Morales et al. 2010), all of which have a double-rooted p2 and lack diastemata in the premolar series. The p4 is a typical amphicyonid tooth with the posterior part being broader than the anterior. The tooth is smaller than that of Cynelos macrodon from Kenya (Adrian et al. 2018), and is similar in size to that of MNCN 79042 from Gebel Zelten, although the p4 of C. anubisi n. sp. is slightly broader. A very low preprotoconulid is indicated by abrasion marks, a feature also present in the Gebel Zelten specimen.The protoconid is the tallest cusp, forms the most voluminous part of the tooth, and also shows strong horizontal abrasion. A strong protoconulid about half the height of the protoconid was present on the distal margin of the tooth, but the protoconulid is abraded in this specimen. A small postprotoconulid is present on the distal end of the postprotocristid, and this feature lacks abrasion marks. Such a postprotoconulid is also present, although smaller, in MNCN 79042. This is in contrast to C. ginsburgi n. comb., where the p4 is relatively shorter compared to m1, with pre- and postprotoconulids present, and C. macrodon, which lacks both a preprotoconulid and postprotoconulid. The morphology of m1 is preserved in the holotype and is also visible in DPC 14532 (Morlo et al. 2007). Both specimens are rather slender, with a low metaconid placed slightly distolingual to the protoconid. Both, the m1 of the type specimen, as well as of DPC 14532, are weathered and abraded, so the original heights of the paraconid, protoconid, and hypoconid are unknown. The lingual border of the talonid is occupied by a small hypoconulid and an even smaller entoconid. The m1 of C. anubisi n. sp. is about the same size as in C. macrodon, but C. anubisi n. sp. has a relatively longer paraconid blade, even longer than that observed for C. ginsburgi n. comb. However, C. anubisi n. sp. and C. macrodon share a more angled paraconid blade than C. ginsburgi n. comb., as well as a metaconid that is positioned slightly more anteriorly. The apex of the m1 metaconid in C. ginsburgi n. comb. is tilted slightly distally, but due to abrasion this feature cannot be determined for the m1 of C. macrodon and C. anubisi n. sp. The m 2 in C. anubisi n. sp. is a massive, elongated tooth in which the protoconid is larger than the metaconid, and the hypoconid approaches the size of the protoconid. Although the tooth is worn and abraded, the original height of the three cusps would have been low. A small entoconid is present on the lingual margin of the tooth. In contrast to Amphicyon, the talonid basin is greatly reduced. As in p4 and m1, there is no development of a cingulid. The very same features are present in an isolated m2 NHM M 82374, assigned to Afrocyon burolleti by Morales et al. (2010: fig. 3) from Gebel Zelten, although the Libyan tooth differs from CUWM 55 in being about 15% smaller. Both the Moghra and Gebel Zelten m2s differ from the holotype of Afrocyon burolleti in being proportionally broader, with a length/breadth ratio of 1.4, while this same figure is about 1.7 in Afrocyon. The m3 is represented only by its alveolus, although the depth and conical shape of the feature suggests a small but substantial tooth, which would have participated in the grinding function of the molar row. It differs from m3 of Afrocyon in being single-rooted. The mandibular symphysis in C. anubisi n. sp. extends slightly more posteriorly, past p2, as compared with MNCN 79042 from Gebel Zelten, in which the symphysis ends directly below p2. In addition, the C. anubisi n. sp. mandible has two mental foramina, one located between p1 and p2, and the other situated slightly posterior to p4. In MNCN 79042 the anterior mental foramen is located slightly more anteriorly below p2, and two smaller foramina are present beyond the anterior root of p4. REMARKS All African members of the genus Cynelos are united by the presence of diastemata between reduced premolars, a feature that distinguishes Cynelos from the other Early Miocene African amphicyonids, Amphicyon and Afrocyon. The occurrence of a diastema between p2 and p3, and the overall trenchant tooth morphology are features shared among Cynelos from Moghra, the type species of Cynelos (the European C. lemanensis), and African C. euryodon. Based on the mandible CUWM 55 we erect a new species of Cynelos, C. anubisi n. sp. The m1, DPC 14532 /2 (Morlo et al. 2007) described previously is designed as the paratype. Additionally, a mandibular fragment from Gebel Zelten, MNCN 79042, previously discussed as “ Amphicyon sp.” (Morales et al. 2010), is also assigned to C. anubisi n. sp., due to a shared similarity in size, a shared low and slender morphology of the mandibular ramus, and the combined presence of a single rooted p1 and p2, a long diastema between p2 and p3, a very short diastema between p3 and p4, the occurrence of pre- and postprotoconulids on p4, and lack of cingulids. In addition, an isolated m2 NHM M 82374 from Gebel Zelten is provisionally referred to C. anubisi n. sp. due to its rectangular outline and small mesiolingual enlargement, which differ both from Afrocyon, and from the m 2 in the mandible NHM M 82373 (Morales et al. 2010: fig. 2) currently attributed to cf. Amphicyon (see below). We thus assign the tooth to Cynelos anubisi n. sp., while recognizing that the Gebel Zelten specimen is slightly smaller than the m2 of the holotype. Cynelos anubisi n. sp. differs from similar sized Cynelos in having a longer paraconid blade. However, a thorough comparison between C. anubisi n. sp. and C. macrodon is hampered by the fact that the only teeth available for comparison are p4 and m1. Indeed, among species of Cynelos, C. macrodon is not very well represented. The holotype of C. macrodon is an M1 from Rusinga Island, Kenya (Savage 1965), and the hypodigm is comprised of only isolated teeth. These include m1 and M1 fragments (Schmidt-Kittler 1987), an additional m1 (Morales et al. 2016), recent material from Kalodirr, which is a p4, P4, and M1 (Adrian et al. 2018), and an M2 from Napak-I, assigned to “ Afrocyon sp.” by Morales et al. (2016: figs. 2, 6). Pickford et al. (2003) identified “? Cynelos ” as present in the fauna from Moroto, Uganda, and Leakey et al. (2011) list a few records of C. macrodon from sites in the Turkana Basin. However, except for the material discussed by Adrian et al. (2018), descriptions of C. macrodon are generally lacking. While the m1 of C. macrodon from Rusinga (Schmidt- Kittler 1987, Morales et al. 2016) is about the same size as the m1 of C. anubisi n. sp., the Rusinga specimen lacks the long paraconid blade and shorter protoconid of C. anubisi n. sp. Compared with C. macrodon from Kalodirr (Adrian et al. 2018), the p4 of C. anubisi n. sp. is smaller and possesses pre- and postprotoconulids. We consider possession of a longer m1 paraconid blade to be an important character distinguishing C. anubisi n. sp. from C. macrodon. Confidence in this feature as a diagnostic character is based on the observation that, among species of Cynelos, members of C. euryodon exhibit variation in a number of ways, including the height of the m1 hypoconid, the size of the antero-labial enlargement, cusp height, and strength of the anterior cingulid in m2, but not in the length of the m1 paraconid blade (pers. obs. MM on KNM without number, KNM RU 2986, KNM RU 4393). Finally, a lower canine and M2 from the Middle Miocene Muruyur Formation, Kenya, were attributed to C. macrodon by Morales & Pickford (2008), but were subsequently transferred to Myacyon by Morales et al. (2016). The specimen KNM-MY 89 (Fig. 3) is a left m2 from the same area. This tooth is part of the carnivoran collection described by Schmidt-Kittler (1987), but this particular specimen was not included in his monograph because it was misplaced at the time. The morphology of KNM-MY 89 is much closer to Cynelos than to Myacyon (see Morales et al. 2010: fig. 3, 6) because it is nearly identical to the m2 of CUWM 55 and to NHM M 82374 from Gebel Zelten. Due to this morphological resemblance we refer the lower canine, M2, and m2 from the Muruyur Formation to Cynelos cf. macrodon, thereby re-instating the initial view of Morales & Pickford (2008). These specimens represent the only record of a large middle Miocene Cynelos in Africa. Morales et al. (2010) referred the Moghra specimens described by Morlo et al. (2007) as “ Cynelos sp. nov.” to the species “ Ysengrinia ” ginsburgi from Arrisdrift (Morales et al. 1998: fig. 4; Morales et al. 2003; Werdelin & Peigné 2010). Due to the presence of a large M2, Morales et al. (2016) excluded the taxon from Ysengrinia and transferred it to Afrocyon. However, the Arrisdrift species differs from Afrocyon burolleti and resembles C. anubisi n. sp. in possessing a much more trenchant dentition with narrower m1 and m2 talonids, and in having a diastema between p2 and p3 (Morales et al. 1998, 2003). We thus corroborate the exclusion of the species from Ysengrinia, but transfer it from Afrocyon to Cynelos as C. ginsburgi n. comb. Cynelos anubisi n. sp. and C. macrodon differ from C. ginsburgi n. comb. in having the m1 metaconid located slightly more anteriorly, and C. anubisi n. sp. further differs from C. ginsburgi n. comb. in having p4 and m2 larger relative to m1. Cynelos ginsburgi n. comb. differs from C. macrodon in having the P4 protocone less reduced, M1 less triangular, and pre- and postprotoconulids present on p4 (see Morales et al. 1998). Cynelos ginsburgi n. comb. also has a shortened M1 talon (see Adrian et al. 2018 for C. macrodon and Morales et al. 2016 for C. ginsburgi n. comb.) relative to C. macrodon, but this abbreviated talon occurs to a much lesser degree in C. ginsburgi n. comb. than it does in Amphicyon giganteus from Moghra. Genus Amphicyon Lartet in Michelin, 1836 (see Peigné 2012)

Genus Cynelos Jourdan, 1862 TYPE SPECIES. ��� C. lemanensis (Pomel, 1846), subsequent designation by Jourdan (1862). DIAGNOSIS. ��� Emended after Peign�� & Heizmann (2003), Werdelin & Peign�� (2010): small to large sized amphicyonids with low, slender mandibles; diastemata between anterior premolars; premolars widest distally; p4 with strong postprotocuspid; the p4 is larger in relation to m1 and to m2 than in Amphicyon, the tip of the main cusp of p4 does not project posteriorly, and the p4 talonid is wider; m1 with low metaconid and tall hypoconid crest, entoconid crest distinct but low, talonid wider than trigonid; m2 mesiodistal length about two thirds the length of m1, m2 lacking the paraconid, with a long and wide talonid, protoconid lacking a distal crest; m3 single-rooted and not double rooted as in Afrocyon, P4 with small protocone, M1 rectangular, M2 slightly more reduced than M1, with paracone slightly larger than metacone, and v-shaped hypocone crests in African specimens. REFERRED AFRICAN SPECIES. ��� Cynelos anubisi n. sp., from Moghra (includes ��� Amphicyon sp.��� in Morales et al. 2010: fig. 4), Cynelos euryodon (Savage, 1965), Cynelos ginsburgi (Morales, Pickford, Soria & Fraile, 1998) n. comb., Cynelos macrodon (Savage, 1965), Cynelos minor (Morales & Pickford, 2008), Cynelos sp. from Buluk, Kenya (KM WS 12663, Anemone et al. 2005). REMARKS Cynelos is a widespread genus of amphicyonid carnivoran known from the late Oligocene through Early Miocene of Africa (Hooijer 1963; Werdelin & Peign�� 2010) and Europe (Peign�� & Heizmann 2003), Early to Middle Miocene of North America (Hunt 1998; Hunt & Stepleton 2015), and the Middle Miocene of Asia (Jiangzuo et al. 2018). North American and European members are well known (e.g., Viranta 1996; Hunt 1998; Peign�� & Heizmann 2003; Hunt & Stepleton 2015), and the Asian material was recently reviewed (Jiangzuo et al. 2018). Individual species representing the African record of the genus have been discussed by several authors (e.g., Morales & Pickford 2005, 2008; Morlo et al. 2007; Morales et al. 1998, 2003, 2008, 2010; Adrian et al. 2018), with overviews given in Werdelin & Peign�� (2010) and Morales et al. (2016). urn:lsid:zoobank.org:act: EED887B0-9F1C-4ADD-B4CF-7E8BBEB7514D Savage (1965) described two amphicyonids, a small species, Hecubides euryodon Savage, 1965 from Napak-I in Uganda, and a large one, Hecubides macrodon Savage, 1965, from Rusinga Island, Kenya. Hecubides Savage, 1965 was later included in Cynelos (Ginsburg 1980), a decision followed by nearly all researchers (e.g., Schmidt-Kittler 1987; Morales et al. 1998; Pickford et al. 2003; Peign�� & Heizmann 2003; Peign�� et al. 2006a, b; Morlo et al. 2007; Werdelin & Simpson 2009; Werdelin & Peign�� 2010; Leakey et al. 2011; Hunt & Stepleton 2015; Jiangzuo et al. 2018; Adrian et al. 2018). One exception to this is the work of Morales and colleagues (Morales & Pickford 2005; Morales et al. 2007, 2008, 2010, 2016), who emphasized differences between H. euryodon and the type species of Cynelos, C. lemanensis (Morales et al. 2016). However, as Jiangzuo et al. (2018) discussed, the features cited in favor of retaining the name Hecubides (Morales et al. 2016) can be regarded as intrageneric if other species of Cynelos are considered. Here we follow the majority of authors in recognizing Hecubides as a junior synonym of Cynelos. Despite acknowledging the close morphological relationship between ��� Hecubides ���, Cynelos macrodon, and ��� Cynelos sp. nov.��� from Moghra (Morlo et al. 2007), Morales et al. (2016) moved these large Cynelos species from the genus Hecubides/ Cynelos to Afrocyon, a monotypic taxon erected on the basis of a partial mandible from the Early Miocene of Gebel Zelten in Libya (Arambourg 1961). This assignment effectively promoted the view that Cynelos was a taxon restricted to northern continents, and that all Early Miocene African amphicyonids belong to either the small Hecubides or the large Afrocyon, with the exception of Amphicyon giganteus from Arrisdrift, Namibia. Recently, Adrian et al. (2018) described new Early Miocene material from Kalodirr, Kenya, and attributed it to C. macrodon, although without mentioning the previous allocation of Cynelos to the North African taxon Afrocyon. Here we follow Adrian et al. (2018) and discuss clear differences between Cynelos and Afrocyon, which unite C. macrodon with C. lemanensis, C. euryodon and other African specimens. Consequently, we not only re-establish the occurrence of Cynelos in Africa, but also recognize six African species in the genus: small C. euryodon and C. minor, large C. macrodon, C. ginsburgi n. comb., Cynelos anubisi n. sp. from Moghra, and a very large species from Buluk, Kenya., Published as part of Morlo, Michael, Miller, Ellen R., Bastl, Katharina, Abdelgawad, Mohamed K., Hamdan, Mohammed, El-Barkooky, Ahmed N. & Nagel, Doris, 2019, New Amphicyonids (Mammalia, Carnivora) from Moghra, Early Miocene, Egypt, pp. 731-745 in Geodiversitas 41 (21) on page 734, DOI: 10.5252/geodiversitas2019v41a21, http://zenodo.org/record/3695843, {"references":["JOURDAN C. 1862. - Des terrains siderolitiques. Revue des Societes savantes. Sciences mathematiques, physiques et naturelles, Paris 1: 130 - 133. https: // gallica. bnf. fr / ark: / 12148 / bpt 6 k 202286 k / f 135. item","POMEL A. 1846. - Memoire pour servir a la geologie paleontologique des terrains tertiaires du departement de l'Allier. Bulletin de la Societe geologique de France, serie 2, 3: 353 - 373. https: // biodiversitylibrary. org / page / 54306487","PEIGNE S. & HEIZMANN E. P. J. 2003. - The Amphicyonidae (Mammalia: Carnivora) from Ulm-Westtangente (MN 2, Early Miocene), Baden Wurttemberg, Germany - systematics and ecomorphology. Stuttgarter Beitrage zur Naturkunde, Serie B (Geologie und Palaontologie) 343: 1 - 133.","WERDELIN L. & PEIGNE S. 2010. - Carnivora in WERDELIN L. & SANDERS W. J. (eds), Cenozoic Mammals of Africa. University of California Press: 603 - 657.","MORALES J., BREWER P. & PICKFORD M. 2010. - Carnivores (Creodonta and Carnivora) from the basal middle Miocene of Gebel Zelten, Libya, with a note on a large amphicyonid from the middle Miocene of Ngorora, Kenya. Bulletin of the Tethys Geological Society 5: 43 - 54.","SAVAGE R. J. G. 1965. - Fossil mammals of Africa: 19. The Miocene Carnivora of East Africa. Bulletin of the British Museum of Natural History (Geology) 10 (8): 239 - 316. http: // www. biodiversitylibrary. org / page / 36359760","MORALES J., PICKFORD M., SORIA D. & FRAILE S. 1998. - New carnivores from the basal Middle Miocene of Arrisdrift, Namibia. Eclogae Geologicae Helvetiae 91 (1): 27 - 40.","ANEMONE R., GROSSMAN A., MILLER E. R. & WATKINS R. 2005. - Biochronology and paleoecology of the Buluk fauna, Early Miocene of northern Kenya. Journal of Vertebrate Paleontology 25 (3, Supplement): 32 A. https: // doi. org / 10.1080 / 02724634.2005.10009942","HOOIJER D. A. 1963. - Miocene Mammalia of Congo. Annales, Musee royal de l'Afrique Centrale, Ser. in 8 °, 46: 1 - 77.","HUNT R. M. JR 1998. - Amphicyonidae in JANIS C. M., SCOTT K. M. & JACOBS L. L. (eds), Evolution of Tertiary Mammals of North America Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Volume 1. Cambridge University Press, Cambridge: 196 - 227.","HUNT R. M. JR & STEPLETON E. 2015. - A Skull of the Immigrant Eurasian Beardog Cynelos (Carnivora, Amphicyonidae) from the Early Miocene of Southern California. Journal of Vertebrate Paleontology 35 (1) e 891229: 1 - 19. https: // doi. org / 10.1080 / 027246 34.2014. 891229","JIANGZUO Q., CHUNXIAO L., ZHANG X., WANG S., YE L. & LI Y. 2018. - Diversity of Amphicyonidae (Carnivora, Mammalia) in the Middle Miocene Halamagai formation in Ulungur River area, Xinjiang, Northwestern China. Historical Biology, https: // doi. org / 10.1080 / 08912963.2018.1477142","VIRANTA S. 1996. - European Miocene Amphicyonidae: taxonomy, systematics, and ecology. Acta Zoologica Fennica 204: 1 - 61.","MORLO M., MILLER E. R. & EL- BARKOOKY A. N. 2007. - Creodonta and Carnivora from Wadi Moghra, Egypt. Journal of Vertebrate Paleontology 27 (1): 145 - 159. https: // doi. org / 10.1671 / 0272 - 4634 (2007) 27 [145: CACFWM] 2.0. CO; 2","MORALES J., PICKFORD M., FRAILE S., SALESA M. J. & SORIA D. 2003. - Creodonta and Carnivora from Arrisdrift, early Miocene of Southern Namibia. Memoirs of the Geological Survey of Namibia 19: 177 - 194.","ADRIAN B., WERDELIN L. & GROSSMAN A. 2018. - New Miocene Carnivora (Mammalia) from Moruorot and Kalodirr, Kenya. Palaeontologia Electronica 21.1.10 a 1 - 19. https: // doi. org / 10.26879 / 778.","MORALES J., PICKFORD M. & VALENCIANO A. 2016. - Systematics of African Amphicyonidae, with descriptions of new material from Napak (Uganda) and Grillental (Namibia). Journal of Iberian Geology 42 (2): 131 - 150. http: // hdl. handle. net / 10261 / 137042","GINSBURG L. 1980. - Hyainailouros sulzeri, mammifere creodonte du Miocene d'Europe. Annales de Paleontologie, Vertebres, 66 (1): 19 - 73.","PICKFORD M., SENUT B., GOMMERY D. & MUSIIME E. 2003. - New Catarrhine fossils from Moroto II, early middle Miocene (c. 17.5 Ma) Uganda. Comptes rendus Palevol 2: 649 - 662. https: // doi. org / 10.1016 / j. crpv. 2003.10.001","PEIGNE S., CHAIMANEE Y., YAMEE C., TIAN P. & JAEGER J. - J. 2006 a. - A new amphicyonid (Mammalia, Carnivora, Amphicyonidae) from the late middle Miocene of northern Thailand and a review of the amphicyonine record in Asia. Journal of Asian Earth Sciences 26 (5): 519 - 532. https: // doi. org / 10.1016 / j. jseaes. 2004.11.003","WERDELIN L. & SIMPSON S. W. 2009. - The last amphicyonid (Mammalia, Carnivora) in Africa. Geodiversitas 31 (4): 775 - 787. https: // doi. org / 10.5252 / g 2009 n 4 a 775","LEAKEY M., GROSSMANN A., GUTIERREZ M. & FLEAGLE J. G. 2011. - Faunal Change in the Turkana Basin during the Late Oligocene and Miocene. Evolutionary Anthropology 20: 238 - 253. https: // doi. org / 10.1002 / evan. 20338","MORALES J., PICKFORD M. & SORIA D. 2007. - New carnivores (Creodonta and Carnivora) from the Early Miocene of Napak, Uganda. Paleontological Research 11: 71 - 84. https: // doi. org / 10.2517 / 1342 - 8144 (2007) 11 [71: NCMCCA] 2.0. CO; 2","ARAMBOURG C. 1961. - Note preliminaire sur quelques Vertebres nouveaux du Burdigalien de Libye. Compte rendu de la Societe Geologique de France 4: 107, 108."]}

Amphicyon giganteus (Schinz, 1825) (Fig. 4) In the synonym list, only first description and citations of the African record is given. Canis giganteus Schinz, 1825: 342. Canis d���une taille gigantesque ��� Cuvier 1824: pl. 193, fig. 20. Amphicyon giganteus ��� Kuss 1965: 66. ��� Morales et al. 1998: fig. 7; 2003: 180. ��� Jiangzuo et al. 2019: 6. Amphicyonidae gen. et sp. indet. ��� Hendey 1978: fig. 4. Megamphicyon giganteus ��� Morales et al. 2016: 147. HOLOTYPE. ��� M7753, Mus��um d���Orl��ans, France. TYPE LOCALITY. ��� Avaray, France. AGE AND HORIZON IN AFRICA. ��� Late Early Miocene, contemporary to European biozone MN4. AFRICAN RECORD. ��� Left P4 (DPC 14532 /1), left M1 (DPC 5426), Left M2 (DPC 8981) from Egypt (Morlo et al. 2007), PQAD 1520, right mandible fragment from Arrisdrift, Namibia (Morales et al. 1998, 2003, 2016). NEW SPECIMEN. ��� CUWM 53, left mandible fragment with crowns of m1-2, and alveoli for a single-rooted p1, double-rooted p2-p4, and a single-rooted m3. DESCRIPTION AND COMPARISONS The left mandible fragment CUWM 53 preserves the crown of m1, part of m2, and alveoli for a single-rooted p1, doublerooted p2-p4, and a single-rooted m3. No diastemata are present. As the inferior part of the mandible is broken, it is not possible to assess its original height. Both m1 and m2 are robust teeth, including their roots. The m1 paraconid-protoconid blade appears to show strong wear, but this feature may be due in part to abrasion of this specimen. The protoconid is the tallest cusp of the trigonid, with the metaconid and paraconid subequal in height. The metaconid is tilted slightly posteriorly. The talonid makes up more than half of the tooth and a large hypoconid, which approaches the metaconid and paraconid in height, is present. The talonid also has a small entoconid, and a cingulid is present. The m2 is broken mesiolingually but a tall protoconid and lower metaconid are preserved. A crest extends from the metaconid distolingually to join the cingulid around the talonid. Specimen CUWM 53 resembles Amphicyon giganteus and is assigned to that species rather than Afrocyon (Arambourg 1961) on the combined basis of having more slender m1 and m2, presence of a massive m1 talonid with the hypoconid as the most prominent cusp, an m2 which is subequal in height to the m1 talonid, lack of a talonid basin on m1, and the presence of an m3 alveolus indicating a single-rooted m3. The Moghra mandible also shares a suite of features with Amphicyon giganteus from Arrisdrift (Hendey 1978; Morales et al. 1998, 2003). These include comparable size, a double-rooted p2, lack of premolar diastemata, m1 with a low metaconid and well-developed hypoconid, the presence of a paraconid on m2, and presence of a single-rooted m3. Morales et al. (2016) referred Amphicyon giganteus from Arrisdrift to the genus ��� Megamphicyon ���, a reassignment of the species suggested by the work of Kuss (1965), but a move that has been rejected by many authors (e.g., Ginsburg & Telles-Antunes 1968; Viranta 1996; Morales et al. 1998, 2003; Peign�� et al. 2006a, 2008; Jiangzuo et al. 2018, 2019; Bastl et al. 2018). Here we join the majority and so recognize the species A. giganteus for the Arrisdrift and Moghra material. European A. giganteus differs from A. major (Middle Miocene of Europe) in being larger on average, lacking diastemata between its premolars, having p4 larger (see Morales et al. 2003), m1 with an entoconid that is narrower and tapers posteriorly, and m2 with a higher trigonid (Jiangzuo et al. 2018). All of these characters also separate the African record of A. giganteus from A. major. ADDITIONAL SPECIMENS Three isolated upper teeth (P4: DPC 14532 /1, M1: DPC 5426, M2: DPC 8981) from Moghra previously described as belonging to ��� Cynelos sp. nov.��� (Morlo et al. 2007) are here provisionally attributed to A. giganteus. These same specimens have been discussed by other authors as perhaps belonging to Ysengrinia ginsburgi (Morales et al. 2010: 48), ��� Afrocyon n. sp.��� (Morales et al. 2016), or Cynelos macrodon (Adrian et al. 2018). The P4 resembles that of A. giganteus from La Barranca and Arroyo de Val (Peign�� et al. 2006b: pl. 2, fig. 1-2), but has the protocone placed slightly more anterior. It differs from C. macrodon and C. ginsburgi n. comb. in having a much larger protocone, a feature that is nearly absent in C. macrodon and is greatly reduced in C. ginsburgi n. comb. The M1 resembles A. giganteus from Farinheira (Ginsburg & Antunes 1995: pl. 1 fig. 2) and La Barranca (Peign�� et al. 2008: pl. 2, fig. 5), but the talon of the Moghra specimen is shorter and narrower. In the former respect the Moghra M1 is reminiscent of the M1 of A. eppelsheimensis from the Late Miocene of Anjou (Gagnaison et al. 2017: fig. 2b). However, the two species are clearly different in that the M1 of A. eppelsheimensis displays a triangular occlusal outline (Kuss 1965, Morlo pers. observ.). The M1 from Moghra differs from members of Cynelos in having a mesiodistally very short talon relative to the trigon, with the protocone and metacone higher, more pronounced cingulae, and a small metaconule present. In these characters the Moghra specimen is unlike the holotype of C. macrodon from Rusinga (see Morales et al. 2016: fig. 9, 2A), and is even more distant from the C. macrodon material from Kalodirr described by Adrian et al. (2018: fig. 5.2-4). The short talon also distinguishes DPC 5426 from C. ginsburgi n. comb. (Morales et al. 2016: fig. 9, 5), C. euryodon, and C. lemanensis. As with P4 and M1, the isolated M2 DPC 8981 from Moghra resembles European specimens of A. giganteus, in this case from B��zian (Ginsburg & Bulot 1982: pl. 1, fig. 2), Olival de Suzana (Ginsburg & Antunes 1995: pl. 1, fig. 2), and La Barranca (Peign�� et al. 2008: pl. 2, fig. 6). The Moghra specimen differs from the M 2 in C. euryodon (Morales et al. 2016: fig. 2, 4), C. lemanensis (Peign�� & Heizmann 2003), C. macrodon from Napak-I, and C. cf. macrodon from the Muruyur Formation, in having a more rectangular shape, possessing stronger labial and lingual cingulae, and with u-shaped, rather than v-shaped hypocone crests. Another African specimen possibly belonging to Amphicyon is the left mandible NHM M 82373 from Gebel Zelten, Libya, which has been discussed elsewhere as representing Afrocyon burolleti (Morales et al. 2010: fig. 2). However, the Gebel Zelten mandible differs from the poorly preserved holotype of Afrocyon burolleti (Arambourg 1961; Morales et al. 2010: fig. 1; Werdelin & Peign�� 2010: fig. 32.1) in possessing a single-rooted m3, and in having a shallower mandible and less acute coronoid angle. The mandible also differs from Cynelos in lacking premolar diastemata, having p4 shorter relative to m1 and m2, and a broader talonid in m1 and the m2 with a wider mesiolingual enlargement. In all these features, NHM M 82373 resembles Amphicyon giganteus, although given the Gebel Zelten specimen���s much smaller size, we refer the mandible to ���cf. Amphicyon ���. REMARKS Fossil material of Amphicyon giganteus has been known since the time of Cuvier (1824), but the range of morphological variation within the species remains an open discussion. Until now, A. giganteus was represented in Africa only by the single mandible reported from Arrisdrift, South Africa (Morales et al. 1998) and some postcranial remains from Gebel Zelten, attributed to the species by Ginsburg (1980) and Ginsburg & Welcomme (2002). With the addition of CUWM 53 from Moghra, the species is again interpreted as having a pan-African distribution (Werdelin & Peign�� 2010). The three isolated upper teeth from Moghra here attributed to A. giganteus are the first upper teeth described from A. giganteus in Africa. Even though there is little appropriate material available for comparison, the three specimens show features that are similar to those seen in A. giganteus from Europe (Ginsburg & Bulot 1982; Ginsburg & Antunes 1995; Peign�� et al. 2008) and which have not been documented in Cynelos, namely: P4 has a much stronger protocone, M1 has an extremely short talon, and M2 has a peculiar morphology with u-shaped hypocone crests and an enlarged lingual cingulum. Re-evaluation of the mandible NHM M 82373 from Gebel Zelten suggests that a second, smaller species of Amphicyon, cf. Amphicyon, is also present in Africa. The Gebel Zelten mandible resembles the Moghra material of A. giganteus much more than it does to specimens of Afrocyon burolleti or Cynelos. However, NHM M 82373 is much smaller than A. giganteus and its m2 shows a slight mesiolingual enlargement not observed among other published A. giganteus specimens. Amphicyon has already been reported from Gebel Zelten based on postcranial material (Ginsburg 1980; Ginsburg & Welcomme 2002) that, however, is too large to belong to the same species as NHM M 82373. Large Amphicyon survived in Africa longer than on any other continent as is evidenced by ��� Amphicyonidae species A��� from Lothagam (Werdelin 2003; Werdelin & Peign�� 2010)., Published as part of Morlo, Michael, Miller, Ellen R., Bastl, Katharina, Abdelgawad, Mohamed K., Hamdan, Mohammed, El-Barkooky, Ahmed N. & Nagel, Doris, 2019, New Amphicyonids (Mammalia, Carnivora) from Moghra, Early Miocene, Egypt, pp. 731-745 in Geodiversitas 41 (21) on pages 738-740, DOI: 10.5252/geodiversitas2019v41a21, http://zenodo.org/record/3695843, {"references":["SCHINZ H. R. 1825. - Das Thierreich eingetheilt nach dem Bau der Thiere als Grundlage ihrer Naturgeschiechte und der vergleichenden Anatomie von dem Herrn Ritter von Cuvier. Volume 4. Stuttgart and Tubingen, 792 p. https: // doi. org / 10.5962 / bhl. title. 120160","CUVIER G. 1824. - Recherches sur les Ossemens fossils ou l'on retablit les caracteres de plusieurs animaux dont les revolutions du globe ont detruit les especes. 2 nd edition. G. 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We present the first record of the mustelid Trochictis from the Late Miocene (MN 9/10) of Germany, a partial mandible with p4, m1, and m2 from Eppelsheim. Trochictis peignei sp. nov. is characterised by the combination p4 with a distal accessory cuspid and a basal lingual enlargement, m1 with a length/width index larger than 2.5, a rounded lingual wall of the paraconid, an entoconulid present, the metaconid as high as the paraconid, and cuspules present on the posterior talonid edge, and m2 with a very reduced talonid. Comparison to similar sized Middle and Late Miocene mustelids and a stratocladistic analysis place T. peignei sp. nov. closest to T. narcisoi from MN 9 of Can Llobateras, T. depereti from several MN 6 to MN 7/8 European localities, and cf. Trochictis sp. from MN 9 of Rudabánya. We also suggest synonymy of T. carbonaria and T. artenensis and verify that m2 of Trochictis occasionally is double-rooted. The analysis does not corroborate a close relationship of Trochictis to Taxodon or the subfamily Ictonychinae, but can also not discard confidently such an assignment. With a body mass of about 3 kg, Trochictis peignei sp. nov. represents the hitherto smallest described carnivoran of the Eppelsheim Formation.