identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03BB87868B07533CFEFCC5FCFB9B299E.text	03BB87868B07533CFEFCC5FCFB9B299E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Enhydriodon Falconer 1868	<div><p>Genus Enhydriodon Falconer, 1868</p><p>TYPE SPECIES. — Enhydriodon sivalensis Falconer, 1868 by original designation.</p><p>OTHER INCLUDED SPECIES. — Enhydriodon africanus Stromer, 1931, E. falconeri Pilgrim, 1931, E. ekecaman Werdelin, 2003, E. hendeyi Morales, Pickford &amp; Soria, 2005, E. kamuhangirei Morales &amp; Pickford, 2005,? E. soriae (Morales &amp; Pickford, 2005), E. dikikae Geraads, Alemseged, Bobe &amp; Reed, 2011, E. afman Werdelin &amp; Lewis, 2013 .</p><p>GEOGRAPHICAL AND STRATIGRAPHIC RANGE OF THE GENUS IN AFRICA (Fig. 1). – Late Miocene to early Pleistocene based on the earliest occurrences in the Tugen Hills, Kenya (5.88-5.72 Ma, Lukeino Formation) and in the Middle Awash, Ethiopia (5.77-5.54 Ma, Adu-Asa Formation) and on the latest occurrences in the Turkana Basin, Kenya (c. 2-1.87 Ma, Koobi Fora Formation and c. 2 Ma at Nakoret) and in Pinnacle, Ethiopia (early Pleistocene). Countries: Egypt, Ethiopia, Kenya, Uganda, Chad, South Africa (see Appendix 2 for details).</p><p>REMARKS</p><p>The attribution of fossil specimens to either Enhydriodon or Sivaonyx Pilgrim, 1931, its close relative, is problematic and has often been debated (e.g. Pickford 2007; Peigné et al. 2008; Werdelin &amp; Peigné 2010; Geraads et al. 2011; Valenciano &amp; Govender 2020). The morphological differences between Sivaonyx and Enhydriodon are difficult to assess because the main differences are carried by the P4, unknown in some species, and because the specimens are fragmentary, preventing from observing the intraspecific variation. We consider here that Sivaonyx beyi Peigné, Bonis, Likius, Mackaye, Vignaud &amp; Brunet, 2008, the oldest species of Sivaonyx in Africa, definitely belongs to this genus, because its dental morphology is very similar to that of the type species Sivaonyx bathygnathus (Lydekker, 1884) (e.g. M 1 with a narrow trigonid relative to the talonid, a protoconid higher and larger than the metaconid, a lingual rim of the talonid lower than the labial rim, M1 with parallel mesial and distal borders, distantly located paracone and metacone, and a trenchant parastyle). However, we consider that all other species of African Sivaonyx or Enhydriodon (except maybe “ Sivaonyx ” soriae Morales &amp; Pickford, 2005) may belong to Enhydriodon, based on some dental traits: metaconid higher than the protoconid on M 1, presence of a carnassial notch and one or more cusps between the protocone and the hypocone on P 4, and/or distolingual expansion on M1.</p><p>Enhydriodon and/or Sivaonyx are recorded in Africa (Stromer 1920; Hendey 1978; Werdelin 2003; Morales &amp; Pickford 2005, 2007; Morales et al. 2005; Haile-Selassie 2008; Lewis 2008; Bonis et al. 2008; Peigné et al. 2008; Geraads et al. 2011; Werdelin &amp; Manthi 2012; Werdelin et al. 2014; Werdelin &amp; Lewis 2020), in southern Asia (Falconer 1868; Lydekker 1884; Pilgrim 1931; Chow 1961; Qi 1985, 2006; Verma &amp; Gupta 1992; Grohé et al. 2013) and in southeastern Europe (Lydekker 1890; Peigné 2016; Koufos et al. 2018). Sivaonyx originated from southern Asia during the middle Miocene (Grohé et al. 2013) and then probably migrated to Europe and Africa during the late Miocene. Other dispersal waves of Sivaonyx and/or Enhydriodon between Africa and southern Asia could have happened during the Pliocene or Pleistocene, although their number and their polarity are difficult to assess due to the lack of cladistic analysis of these otters. Alternately, it is also possible that the African Plio-Pleistocene Enhydriodon evolved from an African Sivaonyx, independently from the Asian species of Enhydriodon .</p><p>From the Miocene to the Pleistocene, two main trends characterized these otters: 1) their teeth became more and more bunodont, a trait observed in other fossil otters (Wang et al. 2017); and 2) their body size increased, with some Plio-Pleistocene forms becoming gigantic, reaching the size of a large male lion (e.g. Enhydriodon dikikae in Geraads et al. 2011; Enhydriodon dikikae and Omo Enhydriodon in Valenciano et al. 2017).</p></div>	https://treatment.plazi.org/id/03BB87868B07533CFEFCC5FCFB9B299E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grohé, Camille;Uno, Kevin;Boisserie, Jean-Renaud	Grohé, Camille, Uno, Kevin, Boisserie, Jean-Renaud (2022): Lutrinae Bonaparte, 1838 (Carnivora, Mustelidae) from the Plio-Pleistocene of the Lower Omo Valley, southwestern Ethiopia: systematics and new insights into the paleoecology and paleobiogeography of the Turkana otters. Comptes Rendus Palevol 21 (30): 681-705, DOI: 10.5852/cr-palevol2022v21a30, URL: https://doi.org/10.5852/cr-palevol2022v21a30
03BB87868B075337FC2CC39EFA052FE1.text	03BB87868B075337FC2CC39EFA052FE1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Enhydriodon omoensis Grohé & Uno & Boisserie 2022	<div><p>Enhydriodon omoensis n. sp.</p><p>(Figs 1-4; Tables 1; 2)</p><p>urn:lsid:zoobank.org:act: B1424D23-7B3B-4639-8D1C-E0611DF8D6D8</p><p>HOLOTYPE. — P 791-18, right P4 (unit C-8, Shungura Formation) .</p><p>REFERRED MATERIAL. — B 426, left I 3 (Usno Formation, correlated with unit B-2 of Shungura Formation); W 8-4S, left P 3 (Usno Formation, equivalent to unit B-2 of Sh. Fm); B 376, right P 4 (Usno Formation, equivalent to unit B-2 of Sh. Fm); L 2-10012, left hemimandible with P 4 and crown base of M 1 (unit B-10, Sh. Fm); L 56-1, left M 1 (unit C-6, Sh. Fm); OMO 18-1972-99, right broken M 1 (unit C-8, Sh. Fm); L 2-148, left M 2 (unit B-10, Sh. Fm); OMO 3/0-10084, left hemimandible with alveolus of M 2, broken roots of M 1 and posterior root of P 4 (unit B-12, Sh. Fm); L 23-10021, right M1 (unit C-6, Sh. Fm); W 717, labial fragment of left M1 ( Usno Fm, correlated with unit B-2 of Sh. Fm); L 183- 14, right femur (unit C-5, Sh. Fm) .</p><p>ETYMOLOGY. — Species name is derived from the Lower Omo Valley, where it has been recovered.</p><p>DIAGNOSIS. — Enhydriodon with at least three lower premolars; P 3 two-rooted, M 1 with a high talonid border, a high hypoconid, a smaller hypoconulid relative to the hypoconid and in distolingual position relative to the latter, an entoconulid higher than the entoconid, and two cuspids on the distal border between the hypoconulid and the entoconulid; P4 with a labial border concave at the level of the region between the parastyle and paracone and between the paracone and metastyle, with two post-protocone cusps aligned distolingually with the protocone, the distalmost post-protocone cusp being much smaller than the mesialmost post-protocone cusp, with a short V-shaped notch between the protocone region and the hypocone; M1 with a large parastyle, closely located paracone and metacone, and an elongated distolingual hypocone; femur with a neck oriented at 40° relative to the longitudinal axis of the diaphysis, a head much higher than the greater trochanter, a strong and very medially oriented lesser trochanter, a diaphysis compressed dorsoventrally with a flat ventral surface, a slightly curved pattelar groove, and a medial condyle that does not project much more posteriorly than the lateral condyle.</p><p>DIFFERENTIAL DIAGNOSIS. — Differs from E. soriae by larger and taller cuspids and cusps, a wider M 1 trigonid, P4 with two post-protocone cusps, fusion of the post-protocone cusp with the protocone, and a narrow separation between the protocone region and the hypocone, M1 with closely located paracone and metacone, and a more distal hypocone; differs from E. ekecaman by a higher crown on M1, a more squared P4 with the distalmost post-protocone cusp more lingually located relative to the hypocone, M1 with a smaller parastyle, a larger metaconule, and a less reduced metacone; differs from E. hendeyi by less bulbous cuspids and cusps, P 4 with a smaller distal accessory cuspid, M 1 with a smaller paraconid, an entoconulid higher than the entoconid, and presence of an hypoconulid, M 2 with a more distally placed metaconid, P4 with post-protocone cusps and lacking a mesiolingual cingulum, longer femur with a head that projects further above the level of the greater trochanter, a more medial lesser trochanter, a concave pattelar groove, and a medial condyle that projects much less ventrally relative to the lateral condyle; differs from E. africanus by a double-rooted P 3, P 4 with a smaller distal accessory cuspid, M 1 with a shorter and higher metaconid and a higher-crowned talonid, M1 with a higher protocone relative to the paraconule and a larger and higher hypocone; differs from E. kamuhangirei by a larger size; differs from E. afman by a double-rooted P 3, M 1 with an entoconulid higher than the entoconid and the presence of an hypoconulid and cuspids on the talonid distal rim; differs from E. dikikae by the presence of P 3, a more squared P 4 with the mesialmost post-protocone cusp being more individualized and the distalmost protocone cusp being less individualized, and a femur with a head projecting further above the level of the greater trochanter; differs from Djourabus dabba Peigné, Bonis, Likius, Mackaye, Vignaud &amp; Brunet, 2008 by a longer M 1 relative to its width and a wider and longer M 1 talonid relative to its trigonid.</p><p>REMARKS</p><p>We attribute the specimens described below (two fragmentary hemimandibles, several isolated lower and upper teeth, and a femur) to a single species, as the range of size and morphology is similar with the intraspecific variation ranges observed in extant otters.</p><p>DESCRIPTION</p><p>Mandible and lower dentition (Fig. 2 A-I, K-N; Table 1)</p><p>On OMO 3/0-10084 (Fig. 2A), a mental foramen is present below the middle part of the P 4, closer to the ventral border of the hemimandible than to the dorsal one. The anteriormost part of the masseter fossa is preserved. The masseter fossa is deep and extends towards the M 2 position. The second hemimandible fragment (L 2-10012; Fig. 2F, G) belongs to a different individual. It only preserves the dorsalmost part beneath the P 4 and does not allow a more detailed description of the mandibular morphology.</p><p>The I 3 is triangular in shape in occlusal view (Fig. 2B). It has a small cuspid. The lingual cingulid is asymmetrical. It extends from the mesial base of the tooth distoventrally then distodorsally, describing a V shape. The mesial cingulid is in a more dorsal position compared to the distal cingulid (Fig. 2B, C).</p><p>The P 3 is almost complete and only lacks its mesialmost and labialmost parts, just at the base of the main cuspid (Fig. 2D, E). It is longer than wide (Table 1). It has two roots. The main cuspid is mesially placed relative to the tooth length, with the mesial face being more vertical than the distal one. Two crests extend from its tip to the mesial and distolabial corners of the tooth. The cingulid is well-marked around the tooth, where the base is preserved.</p><p>The P 4 (Fig. 2G, H) has an ovoid shape: the tooth is wider distally than mesially, it is longer than wide (Table 1), it has a weak concave outline at the middle of the lingual border and a straight and slightly distolabially oriented distal border. It bears a small mesiolingual accessory cuspid, a prominent main cuspid located mesially relative to the tooth length, and a high and large distal accessory cuspid, just distolabial to the main cuspid (Fig. 2 F-H). The tip of the distal accessory cuspid reaches more than half of the height of the main cuspid. Crests are visible on the mesiolingual and distolabial faces of the main and distal accessory cuspids. The cingulid is thick on the mesiolingual side of the tooth, until it reaches the level of the main cuspid tip, and on its distal side, where it encloses a small distolingual unbasined shelf. The small mesial accessory cuspid and the distal border of P 4 are more visible on L 2-10012 (Fig. 2F, G) than on B 376 (Fig. 2H). The distal and mesiolingual borders of P 4 show wrinkles.</p><p>The M 1 has a low trigonid, as wide as the talonid, and slightly longer than the latter (Fig. 2I, L-N; Table 1). The paraconid is the lowest trigonid cuspid. It is subdivided into two cuspids in L 56-1 (Fig. 2L, M). The metaconid is placed distally relative to the protoconid and it is higher and shorter than the protoconid. A short and deep U-shaped lingual valley separates the paraconid from the metaconid. The carnassial notch, between the paraconid and protoconid, is shallow. The metaconid bears a mesial and distal bulge at the cuspid midheight (Fig. 2M). A small protoconulid rises from the lower half of the distal face of the protoconid. The protoconulid is separated from the talonid by a sharp notch, which is positioned approximately at the same level as the metaconid base. The M 1 talonid has a shallow basin. On the labial side, it has a strong hypoconid, whose tip aligns with that of the protoconid, and a smaller hypoconulid, located directly distolingual to the hypoconid. On the lingual side of the talonid, the entoconulid is located sligtly mesially with respect to the hypoconulid, but distally with respect to the hypoconid. It is as high as the hypoconulid. The entoconid is the most mesial talonid cuspid. It is closer to the lingual border compared to the entoconulid and it is lower than the latter. Two additional small and very low cuspids are present between the hypoconulid and entoconulid, on the distal border of the talonid. The hypoconid is as high as the paraconid. The cingulid is particularly well developed around the paraconid.</p><p>The M 2 (Fig. 2K) is wider than long. An oval-shaped mesial contact facet with M 1 is visible. The paraconid is practically indistinct, but a carnassial notch is well visible, even if it is very small and very mesially located. The M 2 protoconid is placed on the distal half of the tooth. The paracristid is very short between the paraconid and the carnassial notch, but longer between the carnassial notch and the protoconid. The paracristid forms a straight line in front of the carnassial notch and follows the curved labial border behind the carnassial notch. The metaconid is the highest cuspid and points mesially relative to the protoconid. The very shallow protocristid notch is located at the middle of the labiolingual axis of the tooth. The talonid is short, representing less than half of the M 2 length. A hypoconid is present at the distal border of the tooth, lingually placed with respect to protoconid. The cingulid is thick on the labial side.</p><p>Upper dentition (Fig. 2 O-U; Table 2)</p><p>The P4 (Holotype; Fig. 2 O-Q) has a large and heavily worn paracone, occupying about one third of the labial side, and located slightly mesially relative to the tooth length (Fig. 2O). It is just distal to a smaller and much lower parastyle. The distolabial corner of the tooth is broken, but the mesial part of the metastyle is preserved, and the outline of the distal border of P4 indicates that the metastyle occupied more than half of the tooth length. Two crests extend mesially from the paracone tip: one extends to the parastyle base, the other to the protocone base. Three cusps constitute the protocone region. They are aligned distolingually on the mesiolingual portion of the P4. The protocone is as large as the paracone, but lower. Its tip is worn (but less than the paracone one) and is displaced mesially with respect to the paracone and distally relative to the parastyle. A second, high cusp, is present on the distolingual face of the protocone (Fig. 2 P-Q). This cusp is higher than the worn hypocone. A third, very small cusp (second and distalmost post-protocone cusp) is present at the base of the distolingual face of the second cusp (mesialmost post-protocone cusp) (Fig. 2 P-Q). A notch separates the protocone region from the large hypocone. The hypocone is lower than the paracone. It is also heavily worn so that its relative height compared to the protocone cannot be assessed. It is positioned distally relative to the paracone and is more lingual relative to the protocone. The base of the protocone region extends more lingually than the base of the hypocone. The broad lingual shelf of P 4 extends until the distal end of the metastyle. The labial outline of P4 describes three convex curves: one around the parastyle, another one around the paracone, and one around the metastyle. We can only observe the mesial part of the latter. The cingulum is well-marked labially and distal to the hypocone.</p><p>The M 1 is wider than long and shows a distolingual expansion (Fig. 2 S-U). The tooth has bulbous cusps. It has a large parastyle, mesiolabially placed relative to the paracone and about the same height as the latter. The metacone is slightly more distal and higher than the paracone. There is no metastylar region labial to the metacone. A paraconule-protocone blade, close to the mesial border of the tooth, extends lingually and slightly distally. The protocone is higher than the paraconule and a shallow notch separates these two cusps. Two crests extend labially from the paraconule: one extends distally until it reaches a lingual bulge at the base of the paracone, the other one extends mesially to follow the mesial border of M1 until the base of the parastyle (Fig. 2S). The distolingual border of the tooth is inflated and shows a hypocone, as high as the metacone. A large metaconule is placed in distolingual position relative to the metacone and reaches about half of the metacone height. The metaconule is more labial compared to the paraconule. The cingulum is thick on the mesial and distolingual sides of the tooth.</p><p>Postcranium (Fig. 3 A-F)</p><p>The femur L 183-14 was already described and measured in detail by Lewis (2008: 611, table 4 for measurements, 614 for description). The neck is short and the head is oriented more proximally than medially, as the neck is shifted at around 40° relative to the longitudinal axis of the diaphysis (Fig. 3A). Because of this feature, the lateromedial width of the proximal epiphysis is narrow (Fig. 3E). There is a wellexcavated proximal surface along the neck and the medial border of the greater trochanter (Fig. 3A, C). The head is round. The articular surface for the acetabulum slightly extends lateroventrally on the neck (Fig. 3A, E). The fovea capitis is large and is located on the ventromedial side of the head (Fig. 3B). The greater trochanter is bending dorsally (Fig. 3D) and it is lower than the femoral head (Fig. 3A, C). The femur has a short (proximodistally) and deep trochanteric fossa. There is a strong lesser trochanter, which is directed more medially than ventrally, so that it is visible in dorsal view (Fig. 3C). The diaphysis is more compressed anteroposteriorly than lateromedially and its ventral surface is flat. The patellar groove is weakly concave and is slightly wider mediolaterally than anteroposteriorly (Fig. 3F). The medial condyle is larger than the lateral condyle (Fig. 3A), but it does not extend much ventrally compared to the lateral condyle (Fig. 3F). The intercondylar fossa is rectangular and particularly wide lateromedially. Two pits are present above the lateral border of the lateral condyle and above the medial border of the medial condyle and could correspond to the insertion sites for the lateral and medial heads of the gastrocnemius (Fig. 3A, B, D). One large pit is present on the medial epicondyle for the attachment of the medial collateral ligament (Fig. 3B). Several pits are present on the lateral epicondyle: one rather centrally located, where the lateral collateral ligament attached, one beneath, closer to the distal border of the epicondyle, where the extensor digitorum longus inserted, and two along the ventral margin of the epicondyle, where at least the popliteus was present (Fig. 3D).</p><p>COMPARISONS</p><p>Asian Enhydriodon</p><p>The first fossils of Enhydriodon were found in Asia. Two species, E. sivalensis Falconer, 1868 and E. falconeri Pilgrim, 1931 were initially recorded from unknown localities of the Siwalik Hills of the Indian Subcontinent, their age being therefore debated. Only one specimen, an isolated m1, which belongs either to E. falconeri or E. sivalensis (see Peigné et al. 2008), was reported from deposits of the “Dhok Pathan Stage” in Hasnot, Punjab province of northern Pakistan (Pilgrim 1931). The “Dhok Pathan Stage” of the Punjab province is equivalent to the late Miocene-early Pliocene Dhok Pathan Formation, dated from 9.8 Ma to c. 3.5 Ma (Barry et al. 2013). More recently, another specimen from Himachel Pradesh, in India, attributed to E. sivalensis, could suggest a late Pliocene age for the species (Verma &amp; Gupta 1992). Here we compare the upper teeth of the Asian Enhydriodon to our material as the holotype of E. falconeri is an isolated P4 and most specimens of the larger E. sivalensis, including its lectotype, are partial crania.</p><p>The Omo P4 differs from that of E. falconeri (holotype BMNH-M 4847, left P4; Pilgrim 1931: pl. 2, fig. 3-3a; Pickford 2007: 97, fig. 9D) by having a longer metastyle and lower post-protocone cusps, which are oriented on a more distal axis. Because of this last feature, the distalmost post-protocone cusp is far more lingually placed in E. falconeri compared to the same cusp in the Omo P 4. This cusp is also much lower than the mesialmost post-protocone cusp and well-separated from the latter, unlike in the Omo P4.</p><p>The Omo upper teeth differ from the lectotype of E. sivalensis (BMNH-M 37153, a cranium with P4-M1 partially preserved and broken P3; Pickford 2007: 104, fig. 12C) by a less squared appearance of P4 and a more developed labial shelf of the paracone and more closely located paracone and metacone on M1. The P4 protocone is lacking on the lectotype, but one cusp is present at the position of the mesialmost post-protocone cusp in the Omo P 4. In that case, the Omo upper carnassial further differs from that of E. sivalensis by having two post-protocone cusps instead of one. Interestingly, two other crania attributed to E. sivalensis by Pilgrim (1932) (BMNH-M 37154 and BMNH-M 37155; Pickford 2007: 104, fig. 12B, D) have a P4 morphology different from that of the lectotype. A wide U-shaped valley filled by several small cusps occurs between the protocone and hypocone, differing greatly from our specimen. Other crania attributed to E. sivalensis (Lydekker 1884: pl. XXVI, fig. 5; GSI NRV2/468, described by Verma &amp; Gupta 1992, refigured by Pickford 2007: 104, fig. 12A) have not been observed and cannot be compared here.</p><p>African Enhydriodon</p><p>One of the oldest possible Enhydriodon species in Africa is E. soriae (Morales &amp; Pickford, 2005), originally described as Sivaonyx soriae based on remains from latest Miocene deposits of the Lukeino Formation, in Kenya (Morales &amp; Pickford 2005; Fig. 1). It is known by lower and upper dentition, the latter originally assigned to S. senutae Morales &amp; Pickford, 2005 in the same paper and re-attributed to S. soriae by Peigné et al. (2008). It is a much smaller otter than the species from Omo (Tables 1, 2; Fig. 4), which shows very different morphological features too. The Omo M1s differ from the M 1 of E. soriae by having larger, taller, and more bulbous cusps, closer paracone and metacone, with the distal crest of the paracone and the mesial crest of the protocone of similar lengths (contrary to a distal crest of the paracone much longer relative to the mesial crest of the metacone in E. soriae), and an hypocone that is more projected linguodistally. The Omo P4 has also larger and taller cusps, clear concavities on its labial wall, a post-protocone cusp fusing with the distolingual face of the protocone (instead of well-separated cusps in E. soriae), two post-protocone cusps (instead of one in E. soriae), and it lacks a wide U-shaped valley between the protocone region and the hypocone. The Omo M 1 has a wider trigonid, which approximates the width of the talonid, while the M 1 of E. soriae has an oval outline because the trigonid is narrower than the talonid.</p><p>Enhydriodon ekecaman Werdelin, 2003 was first described from the early Pliocene fluvio-lacustrine Kanapoi Formation, northern Kenya (Werdelin 2003; Werdelin &amp; Manthi 2012; Werdelin &amp; Lewis 2020), then found in the Mabaget Formation, in the Baringo district of western Kenya (Morales &amp; Pickford 2005; Fig. 1). The Kanapoi material includes complete M 1, M 1 and a partial P 4, while, in Baringo, specimens comprise a complete P 4, P 3, worn and broken M 1 s and a fragmented hemimandible (seeTables 1, 2 and Fig. 4 for measurements). A complete P4 from Kossom Bougoudi, Chad, dated to the Mio-Pliocene boundary, assigned to E. aff. ekecaman (Bonis et al. 2008), is also compared here. Unlike the Omo P4, none of the P4s from Kanapoi, Baringo, and Kossom Bougoudi show any concave border labially to the parastyle-paracone region. Additionally, the upper carnassial from Baringo is narrower relative to its width (Table 2; Fig. 4), and the distalmost post-protocone cusp is more labially placed compared to that of the Omo carnassial. In E. aff. ekecaman from Chad, three cusps are present between the protocone and the hypocone (instead of two in the Ethiopian species); they are smaller and more separated from each other and from the protocone than in the Omo P4. Compared to the M1 from Kanapoi, the Omo M1 has larger cusps in general (especially the metaconule), a smaller parastyle, a paracone located closer to the metacone and linguodistally to the parastyle (contrary to directly lingual to the parastyle in E. ekecaman), a less reduced metacone, and a mesiolingual cingulum that does not extend mesially relative to the protocone position. Finally, the Omo M 1 s have a higher crown and distinctive hypoconid and hypoconulid compared to the lower carnassial from Kanapoi.</p><p>1 1</p><p>Enhydriodon hendeyi from river channel deposits of the Mio-Pliocene Langeberg Quartz Sand (Valenciano &amp; Govender 2020) and Muishond Fontein Pelletal Phosphate members of the Varswater Formation, “E ” quarry of Langebaanweg, South Africa, was originally described as E. africanus by Hendey (1978). The holotype includes a hemimandible bearing alveolus for P 2, alveoli for P 3 with a remaining distal root, and P 4 -M 2 (SAM-PQL 50000A) and an isolated P4 (SAM-PQL 50000B), both possibly belonging to the same individual. The hemimandible shows four mental foramina under the P 3 and P 4, unlike in our Ethiopian form. The cuspids are more bulbous than in the material of Omo Enhydriodon . Contrary to our taxon, in E. hendeyi, the P 4 has a larger distal accessory cuspid; the M 1 has a wider paraconid, poorly differentiated entoconid and entoconulid, with the entoconulid being lower than the entoconid, and no hypoconulid; the M 2 has a more mesially placed metaconid. The P4 from South Africa differs also greatly from the Omo P 4 by having lower cusps, a thick mesiolingual cingulum, and by lacking post-protocone cusps, showing instead a long and shallow V-shaped valley between the protocone and the hypocone. Additionaly, a complete left femur (SAM-PQL 41523), initially attributed to Enhydriodon africanus by Hendey (1978), has been reassigned to E. hendeyi (Morales et al. 2005) and can be compared here with the femur from the Lower Omo Valley. SAM-PQL 41523 is about half the length of L 183-14 (total length of 164.5 mm for SAM-PQL 41523 and 326.3 mm for L 183-14; measurements from Lewis 2008: table 4), the femoral head does not project as far proximally (Lewis 2008: fig. 1A; Valenciano &amp; Govender 2020: fig. 6C), the proximal surface along the neck and the medial border of the greater trochanter is poorly excavated, the proximal epiphysis is wider lateromedially, the lesser trochanter does not project as medially (Lewis 2008: fig. 1A; Valenciano &amp; Govender 2020: fig. 6C, D), the medial condyle projects more ventrally, and the patellar groove is not concave.</p><p>Enhydriodon africanus is known by two specimens from the early Pliocene of Klein Zee, Namaqualand of northwestern South Africa: an M1 and a hemimandible with alveolus for P 2, bases of canine and P 3 crowns, P 4 -M 1, and alveolus for M 2 (Stromer 1931: pl. I, figs 3a, b, 4a, c; refigured in Morales &amp; Pickford 2005: 238, 239, fig. 2I, P from MNHN casts, and in Valenciano &amp; Govender 2020: 16, fig. 4D-F from the original holotype). It is similar in size to E. ekecaman and E. hendeyi, so it is much smaller than the Omo Enhydriodon (Fig. 4; Tables 1, 2). In our Ethiopian species, the M1 has a higher protocone relative to the paraconule and the hypocone is clearly larger and higher than in the South African species. The mental foramen is located under the P 4 in OMO 3/0-10084 while it is present under the P 3 in the holotype of E. africanus . The Omo Enhydriodon has a two-rooted P 3 (instead of a single-rooted P 3 in E. africanus), a smaller distal accessory cuspid on P 4, and an M 1 with a shorter and higher metaconid and higher-crowned talonid, especially lingually, where the entoconid and the entoconulid are separated by a deeper notch. Stromer (1931) also assigned another M 1 from the Pliocene of Wadi Natrun, in Egypt, to E. africanus . He first described that specimen as Lutra aff. capensis (Stromer 1920), but we agree with him (Stromer 1931) and other authors (Peigné et al. 2008) that it belongs to either Sivaonyx or Enhydriodon . We cannot make relevant comparisons here from the figures of the specimen we have seen (Stromer 1920: fig. 11). Also, because part of the talonid is absent, we cannot evaluate what was its width relative to the trigonid width or the height of its lingual rim relative to the labial rim. Yet those two characters can give indications to determine whether the Egyptian specimen belongs to Sivaonyx or Enhydriodon . However, this form is smaller than the Omo taxon; it is closer in size to E. africanus, E. ekecaman and E. hendeyi, and larger than E. soriae .</p><p>Enhydriodon kamuhangirei is represented by very worn M 1 s from the early Pliocene of Kazinga and Warwire, Uganda (Fig. 1). The holotype resembles the Omo M 1 in the number and general position of the talonid cuspids, although it is difficult to assess whether there were two (as in the Ethiopian specimens) or just one cuspid between the entoconulid and the hypoconulid, due to the wear. This species is much larger than E. ekecaman, E. hendeyi and E. africanus, whose sizes are similar. It is still smaller than the Omo otter (Tables 1, 2; Fig. 4). It is also the only Enhydriodon recorded on the western branch of the East African Rift.</p><p>Enhydriodon afman is recorded from the late Pliocene of the Lokochot Member of the Koobi Fora Formation, northern Kenya (Fig. 1). Although similar in age to some of the Omo otter specimens, and located in the eastern part of the Turkana Basin, which is close to the Lower Omo Valley, it is a smaller species (Table 1; Fig. 4) with a distinct dental morphology. E. afman is represented by a hemimandible with alveolus for P 3 and M 2, alveoli for P 4, and M 1, and possibly a few other isolated teeth and bones. The holotype differs from the Omo material by having three mental foramina under P 3 -P 4 (although this feature can be variable within species), a single-rooted P 3, an M 1 talonid without hypoconulid and without cuspids on the distal rim, with the entoconid and entoconulid subequal in size, and with a shallower notch separating the entoconid from the metaconid. The description of the M 2 of E. cf. afman, from the Tulu Bor Member of Koobi Fora (Werdelin &amp; Lewis 2013), matches the morphological features of the Omo Enhydriodon, but probably of other species of the genus too (e.g. E. hendeyi). This M 2 is larger than that of E. omoensis n. sp. (Table 1).</p><p>Enhydriodon dikikae is represented by a cranium and an associated mandible, as well as several isolated lower teeth and postcranial remains. It comes from the early Pliocene of Dikika, in the Lower Awash Valley of Ethiopia (Fig. 1). The Lower Awash otter has smaller tooth dimensions compared to the Omo otter (Tables 1, 2; Fig. 4). However, the M1 of E. dikikae is more elongated than that of E. omoensis n. sp. Morphological differences with the Omo Enhydriodon are mostly visible on the P4 and M 1. The P4 of the holotype of E. dikikae has a different outline of the labial rim, the latter being only concave at the level of the paracone-metastyle region. The tooth is less square in occlusal view, with different proportions and configuration of the lingual cusps between the protocone and the hypocone compared to the Omo P4: the mesialmost post-protocone cusp, at the midheight of the protocone, is less individualized, and the distalmost post-protocone cusp is well-individualized and closer to the hypocone. The M 1 of the holotype of E. dikikae has only one cuspid between the entoconulid and hypoconulid (instead of two cuspids in the Omo M 1 s), and this cuspid is closer to the entoconulid, and as high as the hypoconulid. The entoconulid is also higher than the hypoconulid. These features differ from the position and proportion of the talonid cuspids observed in the Omo Enhydriodon . Another M 1 attributed to E. dikikae (DIK 24-1) shows a different talonid morphology compared to that of the Dikika holotype: the distal rim of the talonid is lower than the labial or lingual cuspids, there are two cuspids distal to the entoconulid, and the entoconid and entoconulid are not well separated. Again, those features are different from what can be seen in the Omo M 1 s. On the other hand, the bicuspid M 1 paraconid of E. dikikae (at least present in DIK 24-1; it is impossible to check its presence in DIK 56-9), mentioned in the species diagnosis, is also present in the Omo Enhydriodon . However, it is less pronounced in E. omoensi s n. sp. and it is a variable feature within the same individual. In fact, a mandible from Hadar (AL 254-1, under study by L. Werdelin and M. Lewis, paper in preparation) shows a left M 1 with a clear bicuspid paraconid, while the right one has a single-cusped paraconid. Finally, E. dikikae does not have a P 3, the first premolar on the dental row of the mandible being a P 4, unlike in the Omo species. Nevertheless, it is morphologically the closest species to the Omo Enhydriodon . In addition to the dental material, Geraads et al. (2011) described three epiphyses of femora of E. dikikae . They are smaller than the epiphyses of the Omo femur (lateromedial width of the proximal epiphysis is 78.3 mm for DIK 44-1 and 88.5 mm for L 183-14, lateromedial width of the distal epiphysis is 65.5 mm for DIK 44-1 and 75.2 mm for L 183-14; measurements from Lewis 2008: table 4). E. dikikae and E. omoensis n. sp. share a similar shape of the trochanteric fossa and a very medially oriented lesser trochanter. However, Geraads et al. (2011) described a less medially oriented lesser trochanter for DIK 4-1, implying that the orientation of the lesser trochanter is variable among individuals of E. dikikae . The proximal femur DIK 44-1 also has a rather deeply excavated proximal surface along the neck and the medial border of the greater trochanter, although not as deep as in the Omo femur. The head only reaches the same level as the greater trochanter and the latter has deeper imprints for the insertion of the hip extensors compared to the Omo femur. Werdelin &amp; Manthi (2012) also described a partial cranium and isolated upper teeth of E. cf. dikikae from Kanapoi. Therefore, E. dikikae could have been present in both the Afar region and the Turkana Basin during the Pliocene.</p><p>Other postcranial and dental specimens of several Enhydriodon species are reported from the late Pliocene Sidi Hakoma and Denen Dora members of the Hadar Formation, north of the Dikika research area (Howell &amp; Petter 1979; Lewis 2008; Werdelin &amp; Lewis 2013). The only specimen described so far is the femur AL 166-1 (Lewis 2008). Smaller than the Omo femur (total length of 259.9 mm for AL 166-1 and 326.3 mm for L 183-14; measurements from Lewis 2008: table 4), the lesser trochanter is very medially oriented, as in E. omoensis n. sp. As for the Dikika femora, the head only reaches the same level as the greater trochanter and the proximal surface along the neck and the medial border of the greater trochanter is deeply excavated, but not as much as in the Omo femur.</p><p>Another Ethiopian occurrence of Enhydriodon is reported from the early Pliocene of Woranso-Mille, central Afar region (Werdelin et al. 2014; Fig. 1), where a few isolated teeth and postcranial bones were discovered. Unfortunately, we could not compare our material to the P 4 and M 1 s of this locality, and the illustration of one M 1 does not permit us to observe detailed features.We can only note that the Omo M 1s are larger than those from Woranso-Mille (Werdelin et al. 2014: fig. 5).</p><p>Finally, other remains of Enhydriodon found in several plio-pleistocene localities of Ethiopia and Kenya await description (e.g. Werdelin &amp; Peigné 2010; Werdelin &amp; Lewis 2013; Appendix 2). It is notably the case for specimens of Enhydriodon sp. from the late Pliocene of the lower Lomekwi Member of the Nachukui Formation, western Turkana Basin of Kenya, where only the femur has been described by Lewis (2008). Lewis (2008) argued that this femur was very similar to that of E. hendeyi, although larger. It is much smaller than the Omo femur (total length of 225.4 mm for KNM-WT 37400 and 326.3 mm for L 183-14; measurements from Lewis 2008: table 4). The femoral head is lower than the greater trochanter. Compared to the Omo femur, the proximal surface along the neck and the medial border of the greater trochanter is poorly excavated, the lesser trochanter does not project as medially, the medial condyle projects more ventrally, and the patellar groove is not concave.</p></div>	https://treatment.plazi.org/id/03BB87868B075337FC2CC39EFA052FE1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grohé, Camille;Uno, Kevin;Boisserie, Jean-Renaud	Grohé, Camille, Uno, Kevin, Boisserie, Jean-Renaud (2022): Lutrinae Bonaparte, 1838 (Carnivora, Mustelidae) from the Plio-Pleistocene of the Lower Omo Valley, southwestern Ethiopia: systematics and new insights into the paleoecology and paleobiogeography of the Turkana otters. Comptes Rendus Palevol 21 (30): 681-705, DOI: 10.5852/cr-palevol2022v21a30, URL: https://doi.org/10.5852/cr-palevol2022v21a30
03BB87868B0C5337FBEDC2FEFBA22AFA.text	03BB87868B0C5337FBEDC2FEFBA22AFA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Torolutra Petter, Pickford & Howell 1991	<div><p>Genus Torolutra Petter, Pickford &amp; Howell, 1991</p><p>TYPE AND ONLY SPECIES. — Torolutra ougandensis Petter, Pickford &amp; Howell, 1991 by original designation.</p><p>DIAGNOSIS (translated from Petter et al. 1991: 952). — “Mandible with a relatively deep masseteric fossa separated from the ventral margin of the ramus and with the anterior border located behind the alveolus for M 2; P 4 robust, with an oval outline, surrounded by a thick and continuous cingulid; M 1 with tall and sharp trigonid cuspids, the protoconid being the tallest trigonid cuspid, a narrow talonid, with a flat and obliquely oriented lingual face of the hypoconid.”</p><p>GEOGRAPHICAL AND STRATIGRAPHIC RANGE. — Late Miocene to early Pleistocene based on the earliest occurrences in the Tugen Hills, Kenya (c. 5.7 Ma, Lukeino Formation) and in the Middle Awash, Ethiopia (5.77-5.54 Ma, Adu-Asa Formation) and on the latest occurrence in the Turkana Basin, Kenya (1.56-1.38 Ma, Koobi Fora Formation). Countries: Egypt, Ethiopia, Kenya, Uganda (see Appendix 2 for details).</p></div>	https://treatment.plazi.org/id/03BB87868B0C5337FBEDC2FEFBA22AFA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grohé, Camille;Uno, Kevin;Boisserie, Jean-Renaud	Grohé, Camille, Uno, Kevin, Boisserie, Jean-Renaud (2022): Lutrinae Bonaparte, 1838 (Carnivora, Mustelidae) from the Plio-Pleistocene of the Lower Omo Valley, southwestern Ethiopia: systematics and new insights into the paleoecology and paleobiogeography of the Turkana otters. Comptes Rendus Palevol 21 (30): 681-705, DOI: 10.5852/cr-palevol2022v21a30, URL: https://doi.org/10.5852/cr-palevol2022v21a30
03BB87868B0D5329FBE2C45CFECB29BE.text	03BB87868B0D5329FBE2C45CFECB29BE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lutrinae Bonaparte 1838	<div><p>Lutrinae indet.</p><p>(Fig. 3 G-K; Table 3)</p><p>MATERIAL. — F 508-26, right humerus lacking the distal epiphysis (lower part of Member H, Shungura Formation).</p><p>DESCRIPTION</p><p>The humerus (Fig. 3 G-K) is broken distally, at the level of the entepicondylar foramen and the olecranon fossa. It has an S-shape in lateral view and is laterally compressed. The head is laterally compressed too and the neck greatly bows ventrally. The lesser tuberosity is lower than the head and is pressed against the dorsomedial surface of the latter. The greater tuberosity reaches the same height as the lesser tuberosity. The greater tuberosity is positioned dorsolaterally to the head and greatly extends dorsally. A narrow and shallow bicipital groove separates it from the lesser tuberosity. The deltopectoral and lateral supinator crests are strong. The proximal border of the entepicondylar foramen is visible. The remaining proximalmost part of the olecranon fossa does not show any supratrochlear foramen.</p><p>COMPARISONS AND DISCUSSION</p><p>The S-shape laterally compressed diaphysis, the presence of an entepicondylar foramen, the strong supinator crest together with low tuberosities on the proximal epiphysis suggest this specimen belongs to an otter. The size is much smaller than the postcranial material of Enhydriodon and is rather reminiscent of a medium-sized otter. Compared to an humerus of the extant African species Aonyx capensis (Schinz, 1821) (SAM-ZM 1474), the tuberosities are lower and the lesser tuberosity is more extended dorsomedially. Moreover, contemporaneous humeri of the medium-sized otters Torolutra cf. ougandensis and cf. Torolutra sp. are known from the Upper Burgi and KBS members of Koobi Fora (Werdelin &amp; Lewis 2013). Two of the humeri from Koobi Fora are larger, and two others are smaller than the Omo humerus. However, it is likely that all these remains correspond to species of similar sizes. Morphologically, the proximal epiphysis on the Koobi Fora fossils (Werdelin &amp; Lewis 2013: fig. 4.7a-g) resembles that of the Omo specimen. Nevertheless, the distal epiphysis does not have any entepicondylar foramen (Werdelin &amp; Lewis 2013: figs 4.5e, f; 4.7b, c, h-j), unlike the Omo specimen, and the shaft is thicker distally, where the supinator crest emerges (Werdelin &amp; Lewis 2013: fig. 4.7b, c). These morphological differences and the fact that the humerus from the Shungura Formation has not been found in association with the above-described teeth of Torolutra render a finer taxonomic attribution premature.</p></div>	https://treatment.plazi.org/id/03BB87868B0D5329FBE2C45CFECB29BE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Grohé, Camille;Uno, Kevin;Boisserie, Jean-Renaud	Grohé, Camille, Uno, Kevin, Boisserie, Jean-Renaud (2022): Lutrinae Bonaparte, 1838 (Carnivora, Mustelidae) from the Plio-Pleistocene of the Lower Omo Valley, southwestern Ethiopia: systematics and new insights into the paleoecology and paleobiogeography of the Turkana otters. Comptes Rendus Palevol 21 (30): 681-705, DOI: 10.5852/cr-palevol2022v21a30, URL: https://doi.org/10.5852/cr-palevol2022v21a30
