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Churchill, Morgan, Boessenecker, Robert W., Clementz, Mark T. (2014): Colonization of the Southern Hemisphere by fur seals and sea lions (Carnivora: Otariidae) revealed by combined evidence phylogenetic and Bayesian biogeographical analysis. Zoological Journal of the Linnean Society 172 (1): 200-225, DOI: 10.1111/zoj.12163, URL: http://dx.doi.org/10.1111/zoj.12163

Type.-CGM 30798, mandible with both dentaries including left i3, m2-3 and right i3, p4-m3 (Fig. 2A) (also figured in Fourtau, 1920; 46, 50), M 29372 is a cast. Diagnosis.-Revised: very large species of Brachyodus, mlm 3 length about 143 mm, C-Pl diastema about 95 mm, i1-i2 absent, i3 tusk-like, at least in males, pl occasionally absent. Occurrence.-Early Miocene, Wadi Moghra, Egypt; Gebel Zelten, Libya; Sperrgebiet, Namibia. Material.-cGm 7210, right M; CGM 30762 right dentary, roots of p3, p4-m3 (Andrews, 1899, pl. XXIII, see below); CGM 30764, left dentary, p3-m2; CGM, 30797, left dentary p2-m2; CGM 30800, right dentary p2-m3; CGM 30801, left maxilla P1, P4; CGM 32984, symphysis; cGm 32985, symphysis; CGM 73660, left dentary p1; CGM unnumbered, right dentary alveolus p1, crown p2 (Andrews, 1899, pl. XXIII); CGM unnumbered, right dentary m2-3; CGM unnumbered, left maxilla P3-M1; CGM unnumbered, right dentary m1-2; CGM unnumbered, right dentary m2-3 (Fourtau, 1920, figs. 32, 33); CGM unnumbered, left dentary, worn m1-3. CUWM 10, right M3; CUWM 14, right maxilla P2-3; CUWM 82, left m3, CUWM 69, right maxilla M2- 3; DPC 2518, left dentary m1, m2 erupting; DPC 4064, right M2; DPC 4065, right maxillary fragment with dP3-M1; DPC 4068, left M3; DPC 4107, left M3 fragment; DPC 4421, left dentary dp4-m2, DPC 4426, right dentary p4-m2; DPC 4438, left M1; DPC 4465, left dentary worn m3; DPC 4586, right dentary p4- m3; DPC 4593; left and right dentaries worn right and left m1-3; DPC 6238, left M2; DPC 6440, left dentary m2-3; DPC 6467, left M2; DPC 6620, right dentary, m1 broken, m2-3 worn; DPC 7373, right upper dP4; DPC 7385, left maxilla fragment; DPC 7501, left dentary m2-3; DPC 7746, left dentary p2-m2; DPC 8930, isolated i3; DPC 8965, right dentary broken m2, m3 erupting; DPC 9028, left dentary m1-3; DPC 9046, isolated i3; DPC 12553, left dentary, p4-m3; DPC 12930, left dentary dp3-4; DPC 14540, molar fragment; DPC 14563, right M1; DPC 14580, left P4, M3; DPC 17679, left maxilla P3-M3; DPC 17681, right dentary m2-3; DPC 17683, right M3; DPC 17684, left maxilla P2-4; DPC 17689, left M3; DPC 17697, left maxilla, P4-M3; M 11067, left P4-M3 (Tables 1, 2). Remarks.-Brachyodus depereti has been described previously (Dineur, 1982; Fourtau, 1920; Holroyd et al., 2010; Pickford, 1991). Premolars and molars are morphologically similar to the type species, B. onoideus from Europe, but the teeth of B. depereti are larger. Upper molars are five-cusped with pinched rather than loop-like styles. The dental enamel is finely wrinkled and there is a prominent lingual cingulum. No skull preserving the premaxilla of B. depereti is known, but it is likely that the central upper incisors were tusk-like and pointed anterioinferiorly, as both B. onoideus and B. aequatorialis show a similar condition (Pickford, 1991). The i1-2 are absent in B. depereti and, at least in males, i3 is a robust, tusk-like tooth. Upper dP4 is molariform. Black (1978) suggested that the canine is possibly absent in females but no material is available from Moghra that would either support or refute this suggestion (after Pickford, 1991). Two specimens are identified here as B. depereti, CGM 30762, a right dentary with p4-m3, and an unnumbered dentary fragment with the alveolus for p1 and the crown of p2. These specimens were originally described and figured by Andrews (1899, pl. XXIII), and subsequently CGM 30762 was also figured by Pickford (1991, pl. 3; Figs.1, 2). Both of these authors cited the dentary with p4-m3 as CGM 2849, but the specimen now has the number CGM 30762. How two different catalog numbers came to be associated with this specimen is unknown, but we identify the specimen using the number it has now. In addition, both of these specimens have a complicated past. Cairo Geological Museum 30762 was originally published by Andrews 1899 as the holotype of Brachyodus africanus. Fourtau (1918, 1920) and Forster-Cooper (1924) subsequently assigned a small number of specimens from Moghra to this taxon. Pickford (1991) felt that the holotype jaw of B. africanus did not belong in Brachyodus, although the rest of the hypodigm did. To remedy this, Pickford reassigned CGM 30762 to the genus Afromeryx, designated it as the type of a new species, A. africanus, and identified one additional sub-adult mandibular specimen (M 15021) as a member of the A. africanus hypodigm (see below). Pickford then created a new species, Brachyodus mogharensis (see below), to house the ������genuine������ Brachyodus material formerly recognized as B. africanus. Based on size and overall morphology, we believe that CGM 30762 can comfortably be included as a specimen of Brachyodus depereti (see discussion below under Afromeryx for more details) and support that assignment here. Brachyodus depereti is well represented at Moghra but is a much rarer element among the Gebel Zelten (Libya) and Sperrgebiet (Namibia) faunas. In Libya, the taxon is documented by the presence of a very large talus in the size range for B. depereti (Pickford, 2003), and in Namibia the presence of B. depereti has been identified on the basis of a distal lateral metacarpal (AM 1' 97), a femur lacking the head and neck (AM 02), and possibly a large talus (unnumbered) (Pickford, 2003). An occurrence of B. depereti has also been reported from Siwa Oasis, Egypt (M 11967) although this is likely an error. Hamilton (1973) published a description of six early Miocene fossil specimens housed in the Natural History Museum (London). A note in the collection indicates that these fossils were purchased by the Museum in 1920 from Lady Moon, who found them ������ near Siwa Oasis������. However, there appears to be no continental Miocene deposits in or around Siwa. Instead, Siwa has long been known as a crossroad for caravans traversing northern Africa, and the fact that B. depereti is a common element in the Moghra fauna suggests that Moghra may be the original source of the material and that people may have carried material from Moghra to somewhere ������ near Siwa������., Published as part of Miller, Ellen R., Gunnell, Gregg F., Gawad, Mohammad Abdel, Hamdan, Mohamad, El-Barkooky, Ahmed N., Clementz, Mark T. & Hassan, Safiya M., 1914, Anthracotheres from Wadi Moghra, early Miocene, Egypt, pp. 967-981 in Journal of Paleontology 88 (5) on pages 967-968, DOI: 10.1666/13-122

Jaggermeryx naida new species Type.-DPC 2499, left dentary, alveoli i3-c, and partial crowns of p2-4 (Fig. 6A-6C). Diagnosis.-Differs from all other anthracotheres in having multiple (six to ten) mental foramina present on both sides of the symphysis, inferior boarder of symphysis inflected inferiorly. Further differs from Brachyodus in having dental formula 3.1.3.3, and lacking specializations of the anterior dentition such as suppression of i1-2, or development of a tusk-like i3. Differs from Afromeryx in having a deeper mandibular symphysis and in lacking p1. Further differs from Sivameryx in being of larger size, and in having a bunodont rather than selenodont dentition. Description.-The most characteristic feature of this taxon is the presence of multiple mental foramina. These foramina vary in size, but extend posteriorly from below i1/i2 to p4, most typically with four in a rhomboid configuration under i1-3, two superiorly and two inferiorly, and with the medial and inferior foramen much larger than the other three. Three additional foramina are usually present below c-p2, and at least one is positioned below p4. The symphysis is unfused, rugose, heavily constructed and broad across its base but tapers superioanteriorly. interestingly, although the symphysis remains unfused, one specimen (CUWM 115) shows an anomaly whereby plastic deformation during fossilization pressed the right and left mandibular halves together in such a way that, at first glance the specimen appears to have four left incisors and two right ones. The ventral boarder of the symphysis in J. naida curves slightly ventrally so that the symphysis is angled ventrally at about 45-48��. The symphysis (see Fig. 6D) can be twice as deep in males (~65-80 mm) as in females (~42 mm). Incisor alveoli are of approximately equal size, although in some cases the i3 alveolus is slightly larger than for i1-2, and is about the same size as the canine alveolus. Small diastemata separate alveoli of i1-3 (~3 mm), and i3-c (9 mm). The species is sexually dimorphic with females having a shallower symphysis and a shorter c-p2 diastema (ranging from ~ 16-21 mm, e.g., CUWM 1578b), and males having a deep symphysis and a c-p2 diastema up to twice the length of that seen in females (~45 mm, e.g., CUWM 115). As represented on the type specimen, some but not all individuals attributed to this taxon have an unusual morphology associated with the anterior dentition, whereby the lower incisors and canine seem to be ������perched������ on the lateral edge of the dentary rather than situated perpendicular to the cheek tooth row (Fig. 6C). Also, where known, lower i1-2 alveoli are slightly procumbent anteriorly, and i3 is procumbent anterolaterally. One explanation for the ������splay������ of the lower anterior teeth is that, in addition to the normal cropping function of artiodactyl incisors, the anterior dentition may have had a scoop-like or sieve-like function. Etymology.-" Naida������ derived from the Greek ���Naias,��� waternymph, in recognition of the species��� semi-aquatic habitat. Material.-CGM 30775, edentulous left dentary symphysis, alveoli for i2-3,c, p2-4; CGM 30885, right symphysis fragment; CGM 83752, left dentary alveoli i3-c, roots of p2-3, partial roots of p4; CUWM 115 edentulous dentary complete symphysis, alveoli left i1-p2, alveoli right i1-p2; CUWM 157a, b right dentary p3-4, m3; DPC 17702, right symphysis fragment; DPC 17738, right dentary alveoli i1-3,c, p2-4 (Table 1). Remarks.-The type specimen of J. naida preserves only part of the premolar series and is otherwise edentulous as is most of the hypodigm, which is less than desirable. However, all of the specimens assigned to J. naida show a number of distinctive morphologies that individually-and certainly collectively-are unknown among other anthracothere taxa. Interestingly, one of the J. naida specimens has been known for a long time because it was figured in Fourtau (1920, Fig. 31) as an unnumbered dentary of Brachyodus africanus. Pickford (1991) subsequently included this specimen as part of the hypodigm of B. mogharensis, but we assign this specimen here (now carrying the number CGM 30775) to J. naida, owing to the presence of multiple mental foramina arranged in the characteristic pattern, as well as the unusual canted morphology of the symphyseal region. When only one unnumbered specimen collected almost a century ago was known, it was possible to consider the unusual morphology as representing idiosyncratic variation. However, sustained collecting efforts at Moghra over the last decade have recovered additional specimens with the same unusual morphology, making clear that these specimens represent a new anthracothere taxon. The occurrence of a large number of mental foramina present bilaterally in this species is unique among the Moghra anthracotheres. Because the mental foramen transmits the mental nerve, which provides sensory innervation to the chin and lower lip, we interpret the morphology of J. naida as indicating that the animal must have had either a very sensitive lower lip or an acutely sensitive tactile snout, perhaps combined with perceptive vibrissae. Also, the spacing of the lower incisor alveoli, and the fact that the symphyseal region shows slight splay anteriorly, suggests that the anterior lower dentition may have functioned as a kind of scoop-like or sieve-like structure. This morphological information, together with sedimentological data showing that the Moghra deposits formed under wet conditions, and preliminary results from stable isotope analyses indicating that many of the Moghra anthracotheres, including J. naida, were semiaquatic, leads to the conclusion that J. naida may have foraged along the river banks, using its sensitive tactile snout, and employing the lower anterior teeth in a cropping and scoop-like fashion., Published as part of Miller, Ellen R., Gunnell, Gregg F., Gawad, Mohammad Abdel, Hamdan, Mohamad, El-Barkooky, Ahmed N., Clementz, Mark T. & Hassan, Safiya M., 1914, Anthracotheres from Wadi Moghra, early Miocene, Egypt, pp. 967-981 in Journal of Paleontology 88 (5) on pages 977-978, DOI: 10.1666/13-122

Jaggermeryx new genus Type species.- Jaggermeryx naida new species by monotypy. Diagnosis.-As for species. Etymology.- ‘‘Jagger’’ for Sir Mick Jagger, in recognition of his famous lips, and meryx’’ Gr. ‘‘ruminant,’’ a common suffix for artiodactyls. Occurrence.-Early Miocene, Wadi Moghra, Egypt.