Cynelos Jourdan 1862

Genus Cynelos Jourdan, 1862

TYPE SPECIES. — C. lemanensis (Pomel, 1846), subsequent designation by Jourdan (1862).

DIAGNOSIS. — Emended after Peigné & Heizmann (2003), Werdelin & Peigné (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 & Peigné 2010) and Europe (Peigné & 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; Peigné & 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 & Peigné (2010) and Morales et al. (2016).

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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; Peigné & Heizmann 2003; Peigné et al. 2006a, b; Morlo et al. 2007; Werdelin & Simpson 2009; Werdelin & Peigné 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.