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† Myzopoda africana, new species Figures 2, 3 Myzopoda sp., Butler, 1978: 65; Gunnell, 2010: 586. HOLOTYPE: NMT.010/Bat, complete left humerus (see table 1 for measurements). ETYMOLOGY: The species name is given for the African continent, since the new species represents the only known occurrence of the genus Myzopoda on continental Africa; extant species are restricted to Madagascar. TYPE LOCALITY: Tanzania: Arusha Province, Olduvai Gorge, Bed I, FLK NI, Layer 123. DIAGNOSIS: Differs from living Myzopoda schliemanni and M. aurita in having a larger humerus (mean dimensions 21% larger) that is more robust and has a relatively longer deltopectoral crest, a more rounded humeral head, a more robust lesser tubercle, a more distinct and elongate lateral capitular tail, a more distinct and laterally compressed capitulum, and a relatively broader epitrochlea with a more distally extended epitrochlear process. REFERRED SPECIMENS: NMT.008/Bat, left proximal humerus from Olduvai Bed I, FLK NI, layer 123; NMT.009/Bat, right distal humerus, Olduvai Bed 1, FLK NI, layer 4 (collected 1960). DESCRIPTION: Three humeri from Olduvai can be assigned to Myzopoda based on the presence of the following combination of characters: round, bulbous capitulum, extended capitular tail with flaring lip, a broad epicondyle with two distinct processes, and a humeral head placed distal to the trochiter (greater tubercle). The proximal humerus (fig. 2) of M. africana has a semi-rounded, distolaterally slightly flattened head. The proximal extent of the head does not extend as far proximally as the trochiter and is even slightly below the proximal extent of the lesser tubercle. The deltopectoral crest is elevated anteriorly, is relatively long and curving, and has a sharply defined anterior margin with a slight overhanging lip developed medially. Distally (figs. 2, 3), the humerus of M. africana has a rounded and slightly laterally compressed capitulum that is robust and not offset from the long axis of the humeral shaft. The lateral capitular tail is as broad as the trochlear surface. The trochlear groove is distinct but not deeply invaginated and with a sharply defined trochlear lip. The medial epicondyle is robust with a relatively elongate process that extends distally beyond the trochlear ridge and is developed as two rounded surfaces aligned anteroposteriorly. There is a small but distinct groove on the lateral surface of the epicondyle. Other fossil material of Myzopoda is unknown from both Africa and Madagascar, although more ancient myzopodids referred to another genus have been described from Egypt (Gunnell et al., 2014)., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 3-7, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press.","Gunnell, G. F., N. B. Simmons, and E. R. Seiffert. 2014. New Myzopodidae (Chiroptera) from the late Paleogene of Egypt: emended family diagnosis and biogeographic origins of Noctilionoidea. PLoS ONE 9 (2): e 86712. [doi: 10.1371 / journal. pone. 0086712]"]}

Cf. Eptesicus isabellinus Figure 8 Eptesicus cf. hottentotus, Butler, 1978: 65. REFERRED SPECIMEN: NMT.032/Bat, right distal humerus, FLK NI, layer 6, 1964 (see table 1 for measurements). DESCRIPTION AND DISCUSSION: Olduvai specimen NMT.032/Bat represents a vespertilionid humerus very similar in size and morphology to that of extant Eptesicus isabellinus. NMT.032/Bat (fig. 8, bottom row) has a narrow but robust capitulum that is not angled or offset from shaft. The lateral capitular tail is much less broad than the trochlear surface, and is separated from the capitulum by a shallow groove. A trochlear groove is absent and the trochlear surface is relatively wide and robust with a sharply defined trochlear lip. The medial epicondylar process is low and robust and does not extend distally beyond the trochlear ridge. Groove on the lateral surface of epicondyle is weak to absent. These are all features typical of Eptesicus. The taxonomy of Eptesicus isabellinus is unresolved at the moment, but it appears to be the North African representative of the Eptesicus serotinus radiation sensu lato (Simmons, 2005). However, it is not clear whether E. isabellinus can be maintained as a species separate from E. serotinus (Simmons, 2005). We have chosen to retain it as distinct subject to revision of circumMediterranean Eptesicus species. NMT.032/Bat very closely resembles E. isabellinus (NHMUK 19.7.7.1160, fig. 8, top row) in nearly every detail of size and morphology. The fossil specimen has a slightly less distally extended epicondylar process and the distal end is slightly deeper anteroposteriorly. NMT.032 differs from Eptesicus furinalis (AMNH 278332) only in being relatively larger with a slightly broader capitulum and lacking any groove between the capitulum and trochlea. The lateral ridge is slightly broader than in the extant taxon but otherwise similar. The only records of Eptesicus fossil material from Africa all come from southern Africa (Hendey, 1981; Pocock, 1987; Avery, 1998), mostly from the early to late Pliocene deposits, and they are based on teeth and jaws that are not directly comparable to the specimen described here. Eptesicus is a common element of other Plio-Pleistocene faunal communities and is widespread across the northern continents during this time period (Simmons and Gunnell, in prep.)., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 13-15, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press.","Hendey, Q. B. 1981. Paleoecology of the late Tertiary fossil occurrences in \" E \" Quarry, Langebaanweg, South Africa, and a reinterpretation of their geological context. Annals of the South African Museum 84: 1 - 104.","Pocock, T. N. 1987. Plio-Pleistocene fossil mammalian microfauna of southern Africa - a preliminary report including description of two new fossil Muroid genera (Mammalia: Rodentia). Palaeontologia Africana 26: 69 - 91.","Avery, D. M. 1998. An assessment of the Lower Pleistocene micromammalian fauna from Swartkrans members 1 - 3, Gauteng, South Africa. Geobios 31: 393 - 414."]}

† Cardioderma leakeyi, new species Figures 4, 5 Megadermidae Butler and Greenwood, 1965: 14. Cardioderma sp., Butler, 1978: 65; Gunnell, 2010: 586. HOLOTYPE: NMT.003/Bat, left maxilla with P4–M3 (fig. 4), from the 1960 Olduvai Collection (see table 2 for measurements). REFERRED SPECIMEN: NMT.002/Bat, right dentary with m1–3, FLK NI, Layer 3. ETYMOLOGY: Named in honor of L.S.B. Leakey who was instrumental in initiating and leading the search for vertebrate fossils, especially fossil humans, in East Africa. TYPE LOCALITY: Tanzania: Olduvai Gorge, Bed I, FLK NI, Layer 2. DIAGNOSIS: Differs from extant Cardioderma cor in averaging 18%–20% larger in tooth dimensions; P4 with relatively larger parastyle and metastyle and better developed labial cingulum; M1–2 with relatively more robust mesostyle, deeper parafossa and metafossa, and deeper trigon basin; less reduced M3/m3; m1 and m2 with relatively broader talonid basin and more robust metaconid; m3 with broader trigonid and less reduced talonid. DESCRIPTION AND COMPARISONS: The specimens referred to Cardioderma leakeyi can be recognized as megadermatids based on the robust nature of the cusps and crests on upper and lower cheek teeth, the large hypocone shelves and the narrow and labiolingually restricted protofossae on M1–2, the broadly open trigonid on m1 and the high and short lower molar talonids. The holotype maxilla (NMT.003/Bat, fig. 4) has a maxillary foramen that opens over the anterior root of P4 as in the extant species of Cardioderma, C. cor. There is a secondary and smaller foramen that opens more ventrally on the maxilla over the posterior root of P4 (this foramen is slightly more posteriorly placed in the extant species). In lateral view, the anterior labial root of M1 is exposed through the bony surface of the maxilla, as is often the case in C. cor. The root of the zygomatic arch is dorsal to M2 as in the living form, but it is much more robust in C. leakeyi, as is the anterior orbital process. The optical foramen is the same size and in the same position in both species of Cardioderma. The P4 of Cardioderma leakeyi has a robust paracone, relatively large parastyle and metastyle, and a relatively heavy labial cingulum. The lingual cingulum is relatively broad anteroposteriorly and extends lingually farther than is seen in C. cor. M1–2 each have a prominent hypocone shelf, an anteroposteriorly narrow but labiolingually extended and deep protofossa, a robust mesostyle, and parastylar and metastylar foveae that are nearly equivalent in size (whereas in C. cor the metastylar fovea is typically larger). M3 has a relatively robust parastyle and a labiolingually short lingual shelf and is less anteroposteriorly compressed than seen in C. cor. The horizontal ramus of the dentary (NMT.002/Bat) is relatively deep in Cardioderma leakeyi (3.0 mm beneath m1) compared to C. cor (2.0 mm below m1). The lower molars of C. leakeyi (fig. 5) are all quite similar to those of extant C. cor. The m1 is broken anteriorly, making it impossible to tell whether C. leakeyi had a small, low, and centered paraconid as seen in C. cor. All three lower molars of C. leakeyi have a prominent protoconid and a somewhat lower, but distinct, metaconid. Lower m2 and m3 have distinct paraconids that are placed somewhat lower than the metaconids. The hypoflexid is deep and the cristid obliqua is angled and joins the postvallid of the trigonid well lingual of center on all molar teeth. The hypoconid on m1–2 is distinct, but it is less so on m3; the entoconid is distinct only on m1. Molar talonids are broader than those seen in C. cor, but the talonid is not as broad as the trigonid on any tooth. All molars have moderate and complete labial cingulids. The only other published record of a fossil Cardioderma species (Louchart et al., 2009) is from the early Pliocene in Ethiopia. However, the material upon which that assignment is based has never been described or figured making comparisons impossible at this time.

Miniopterus cf. M. schreibersi (Kuhl, 1817) Figures 14, 15 Miniopterus cf. schreibersi, Butler, 1978: 65; Gunnell, 2010: 588. REFERRED SPECIMENS: NMT.007/Bat, right dentary p3–4 (p4 now lost but shown in drawings of MG), from FLK NI, layer 5, Tr. IV; NMT.011/Bat, left and right distal humeri, from FLK NI, layer 3; NMT.017/Bat, right distal humerus, from FLK NI, layer 2; NMT.019/Bat, right proximal humerus, FLK NI, layer 2 (maybe not Miniopterus, poorly preserved); NMT.022/Bat, left proximal humerus, right distal humeri (2), humeral shaft, from FLK, Main Dig Z level; NMT.038/Bat, right distal humerus, from FLK NI, layer 3, Tr. IV (see tables 1 and 2 for measurements). DESCRIPTION AND DISCUSSION: One dental specimen representing Miniopterus cf. M. schreibersi is known from the Olduvai sample, a right dentary (NMT.007/Bat) that originally had p3–4 and the alveoli for all other teeth preserved. The p3 is double rooted with a single, lingually placed cusp that has a sloping, convex buccal surface and straight, flat lingual surface. A continuous, weakly developed cingulid is present that is developed into a tiny posterolingual extension. In detailed morphology this p3 is quite similar to that of extant M. schreibersi (fig. 14). The p4 was originally in place in this specimen and based on the available drawings (see fig. 14) it was very similar to p4 of extant M. schreibersi. It shared the presence of a tall, pointed protoconid and a broad but short talonid shelf formed by the distal cingulid with the extant taxon. Both extinct and extant specimens lack any indication of a para- or metaconid and both have well-developed labial and lingual cingulids that join anteriorly with the labial cingulid sloping ventrally from anterior to posterior. The dentary has a large mental foramen beneath the c1–p2 roots. Based on alveoli, p2 was single rooted as in living M. schreibersi. The i3 alveolus is slightly separated from the conjoined i1–2 alveoli and is larger than either of the other incisor alveoli as in extant Miniopterus species, and all incisor alveoli are aligned mesiodistally. There is a smaller mental foramen below the i1 alveolus on anterior surface of dentary. The humeral fragments of the Olduvai Miniopterus specimens are very similar in size and morphology to the humerus of extant Miniopterus schreibersi. The proximal end of the humerus in Olduvai Miniopterus specimens has a moderately high (anteriorly) deltopectoral crest that is relatively long and sharply defined. It has a distinct, overhanging medial lip. The crest curves to the base of the trochiter and anteriorly encloses a relatively deep intertrochanteric fossa. The head of the humerus is ovate and angled. The proximal margin of the head does not extend as far proximally as the trochiter, instead extending proximally about the same distance as the lesser trochanter. The distal end of the Miniopterus humerus has a narrow, rounded capitulum that is slightly angled and is not offset from shaft. The lateral capitular tail is somewhat broader than the trochlear surface and is separated from the capitulum by a deep and distinct groove. In contrast, the trochlear groove is faint and the trochlear surface is relatively narrow with a sharply defined trochlear lip. The medial epicondylar process is broken in all Olduvai specimens, but, judging by what remains, it would have been distinct and would have extended well beyond the trochlear ridge as it does in extant Miniopterus (fig. 15). The groove on the lateral surface of the epicondyle is absent, but there is a distinct fossa that occurs on the lateral surface below the root of the epicondylar process. Like all other Miniopterus species, the specimens from Olduvai have well developed and deep olecranon fossae (see fig. 15B, top and bottom), a feature not commonly found in bats. These Olduvai specimens are assigned to Miniopterus cf. M. schreibersi because they are very similar in morphological detail to the humerus of the living taxon. However, Miniopterus schreibersi is not the only similar-sized Miniopterus species to occupy sub-Saharan Africa today. Therefore it is possible that the Olduvai species might be more closely related to one of the other species, especially M. africanus or M. inflatus, given that both of these species have been confused with M. schreibersi in the past (Simmons, 2005). We have chosen to follow Butler’s (1978) original assignment of these specimens to Miniopterus cf. M. schreibersi but acknowledge that more complete specimens are needed to assess the precise relationship of the Olduvai Miniopterus species to living members of that genus. The African fossil record of Miniopterus is limited to the record from Olduvai, two Pliocene records from South Africa (Pocock, 1987) and a late Pliocene record from Morocco (Gunnell et al., 2011). The South African Pliocene occurrences are recorded in faunal lists, but the specimens upon which the assignments are based have never been described or figured., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 23-28, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Kuhl, H. 1817. Die deutschen Fledermause. Hanau 22: 1 - 67.","Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press.","Vaughan, T. A. 1976. Nocturnal behavior of the African false vampire bat (Cardioderma cor). Journal of Mammalogy 51: 227 - 248.","Csada, R. 1996. Cardioderma cor. Mammalian Species 519: 1 - 4.","Rosevear, D. R. 1965. The bats of West Africa. London: British Museum (Natural History).","Smithers, R. H. N. 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria.","Johnston, D. S. 2006. Nycticeinops schlieffeni. Mammalian Species 798: 104.","Kingdon, J. 1974. East African mammals: an atlas of evolution in Africa. II (A). Insectivores and bats. London: Academic Press.","Hill, J E. 1974. A review of Scotoecus Thomas, 1901 (Chiroptera: Vespertilionidae). Bulletin of the British Museum (Natural History), Zoology 27: 169 - 188.","Jacobs, D. 2008. Scotoecus albofuscus. The IUCN red list of threatened species. Version 2015.2.","Van der Merwe, M. 1975. Preliminary study on the annual movements of the Natal clinging bats. South African Journal of Science 71: 237 - 241.","Nowak R. M. 1994. Walker's bats of the world. Baltimore: Johns Hopkins University Press.","Taylor, P. 1998. The smaller mammals of KwaZulu-Natal. Pietermaritzburg, South Africa: University of Natal Press.","Mickleburgh, S., A. M. Hutson, W. Bergmans, and J. Fahr. 2008 b. Nycteris thebaica. The IUCN red list of threatened species. Version 2015.2.","Happold, D. C. D. 1987. The mammals of Nigeria. London: Oxford University Press.","Hayman, R. W. 1954. Notes on some African Bats, mainly from the Belgian Congo. Revue de Zoologie et de Botanique Africaines 50: 277 - 295.","Goodman, S. M., et al. 2005. The distribution and conservation of bats in the dry regions of Madagascar. Animal Conservation 8: 153 - 165.","MacKinnon, J., C. E. Hawkins, and P. A. Racey. 2003. Pteropodidae. In S. M. Goodman and J. P. Benstead (editors), The natural history of Madagascar: 1299 - 1302. Chicago: University of Chicago Press.","Mickleburgh, S., A. M. Hutson, W. Bergmans, J. Fahr, and P. A. Racey. 2008 c. Eidolon helvum. The IUCN red list of threatened species. Version 2015.2.","Mickleburgh, S., A. M. Hutson, W. Bergmans, F. W. Cotterill, J. Fahr, and D. Jacobs. 2008 a. Tadarida aegyptiaca. The IUCN red list of threatened species. Version 2015.2.","Olsson, A., D. Emmett, D. Henson, and E. Fanning, E. 2006. Abundance and activity patterns of microchiropteran bats at a cave roost in southwest Madagascar. African Journal of Ecology 44: 401 - 403.","Pocock, T. N. 1987. Plio-Pleistocene fossil mammalian microfauna of southern Africa - a preliminary report including description of two new fossil Muroid genera (Mammalia: Rodentia). Palaeontologia Africana 26: 69 - 91.","Gunnell, G. F., T. P. Eiting, and D. Geraads. 2011. New late Pliocene bats (Chiroptera) from Ahl al Oughlam, Morocco. Neues Jahrbuch fur Geologie und Palaontologie 260: 55 - 71."]}

† Nycticeinops serengetiensis, new species Figures 9, 10 Cf. Nycticeius (Scoteinus) schlieffeni, Butler and Greenwood, 1965: 15; Butler, 1978: 65; Gunnell, 2010: 588. HOLOTYPE: NMT.005/Bat, right dentary with p4–m1 and alveoli for all other teeth (fig. 9F–H; see table 2 for measurements). REFERRED SPECIMENS: NMT.001/Bat, left dentary with m2–3 and alveoli for all anterior teeth, from FLK NI, layer 123; NMT.006/Bat, left edentulous dentary preserving all tooth alveoli, from FLK NI, layer 1; NMT.013/Bat, left distal humerus, from FLK NI, layer 2; NMT.015/Bat, right distal humeri (3), from FLK NI, layer 123; NMT.016/Bat, left and right distal humeri, from FLK NI, layer 3; NMT.023/Bat, right distal humerus (and most of shaft), from Main Dig, FLK 1, Z level; NMT.025/Bat, left distal humerus, from FLK1, Z level; NMT.034/Bat, left proximal humerus, from FLK NI, layer 1, Tr. IV, 1964; NMT.035/Bat, right distal humeri (2), from FLK NI, layer 5, Tr. IV, 1964; NMT.036/Bat, left distal humerus, from FLK NI, layer 2, Tr. V; NMT.037/Bat, left and right distal humeri, from FLK NI, layer 1, Tr. III (see table 1 for measurements). ETYMOLOGY: Named for the famed Serengeti region in northern Tanzania where Olduvai Gorge is located. TYPE LOCALITY: Tanzania: Olduvai Gorge, Bed I, FLK NI, layer 5, Tr. IV. DIAGNOSIS: Differs from extant Nycticeinops schlieffeni (the only other recognized species of the genus) in having teeth on average 20% larger; p4 relatively taller and labiolingually narrower with a relatively longer talonid shelf; m1 with more curved paracristid and more angled entocristid, hypoconulid relatively lower and talonid basin more deeply excavated; m2 with relatively lower hypoconulid; and m3 with heavier anterolabial cingulid. DESCRIPTION AND DISCUSSION: All three of the known fossil jaws have slightly different arrangements of their incisor alveoli. The holotype of N. serengetiensis (NMT.005/Bat) has a small i3 alveolus directly in front of the canine alveolus while a slightly larger i2 alveolus is anteromedial to that of i3 and directly posterior to an equally large i1 alveolus. NMT.001/Bat has a similar arrangement except that the i1 alveolus is noticeably larger than either the i2 or i3 alveoli and is placed somewhat more laterally, producing a triangular arrangement of alveoli. The edentulous dentary of N. serengetiensis (NMT.006/Bat) is more similar to the holotype except that the alveoli of i2 and i3 are nearly mediolaterally opposite one another and the i2 alveolus is smaller. The arrangement of incisors in extant Nycticeinops schlieffeni (e.g., AMNH 257406) is somewhat different in that the i2 alveolus is not as far medial but instead forms part of a non-arcuate row of incisors running from the canine to the midline of the jaw with i1 most anterior and i3 most posterior. The offset arrangement found in N. serengetiensis may, in part, be due to the larger teeth of the fossil form and subsequent crowding of teeth in the jaw (although certain other extant vespertilionids also have a similar arrangement of incisor alveoli to those seen in N. serengetiensis; N.J. Czaplewski, personal commun.). As further evidence of this crowding, the p4 of NMT.005/Bat is very closely appressed to the m1 with the m1 overlapping the p4 talonid shelf. The dentaries of N. serengetiensis are all broken posteriorly but preserve enough of the ascending ramus to show that it rose dorsally at a shallower angle than that in N. schlieffeni, in which the ramus rises more steeply and then turns anteriorly (fig. 9A). Like in the extant taxon, there is typically a single mental foramen located beneath the area of contact between c1–p3, but it is relatively larger in N. serengetiensis, which occasionally has a smaller foramen located more anteriorly as well (fig. 9D). The dentaries of N. serengetiensis also differ from the modern species by having their ventral borders gently sloping dorsally from posterior to anterior and then ending anteriorly by turning dorsally. In N. schlieffeni the dorsal border of the dentary angles dorsally from posterior to anterior and forms a continuous surface with no angulation producing a jaw that is deeper beneath p4 than it is beneath m3. The Olduvai collection includes 13 distal humeri and one proximal humerus that we refer to N. serengetiensis based on their similarity to those of extant N. schlieffeni. NMT.034/Bat is a left proximal humerus that has a semi-rounded and slightly angled head that does not extend quite as far proximally as the trochiter but does extend slightly past the lesser trochanter. The deltopectoral crest is high anteriorly and fairly long, and has a sharply defined edge with a slightly overhanging lip developed medially. The crest curves toward the base of the trochiter and anteriorly encloses a moderately deep intertrochanteric fossa. The distal humerus of N. serengetiensis is characterized by a relatively narrow capitulum that is neither angled nor offset from the shaft. The lateral capitular tail is much less broad than the trochlear surface and is separated from the capitulum by a very shallow groove. The trochlear groove is weak and the trochlear lip is sharply defined. The medial epicondyle has a small process that extends distally only slightly beyond trochlear ridge. The groove on lateral surface of epicondyle is moderately developed. Other than the specimens described here no other fossils of Nycticeinops are known (Simmons and Gunnell, in prep.)., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 17-19, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M., and M. Greenwood 1965. Chiroptera. In L. S. B. Leakey (editor), Olduvai Gorge 1951 - 1961, vol. 1. A preliminary report on the geology and fauna: 14 - 15. Cambridge: Cambridge University Press.","Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press."]}

Cf. Myotis sp. Myotis sp., Butler, 1978: 65; Gunnell, 2010: 588. REFERRED SPECIMEN: NMT.012/Bat, left damaged distal humerus, FLK NI, layer 3 (see table 1 for measurements). DESCRIPTION AND DISCUSSION: NMT.012/Bat is the only specimen in the Olduvai sample that may represent Myotis. The distal end of the humerus of NMT.012/Bat has typical Myotis features including having a relatively narrow capitulum that is not offset from shaft, a lateral capitular tail that is less broad than trochlear surface and proximolaterally extended into a distinct lip, a trochlear groove that is essentially absent, and a trochlear surface that is relatively broad with a trochlear lip that is not sharply defined. Additionally, the medial epicondyle has a small process that does not extend distally beyond the trochlear ridge and the groove on the posterolateral surface of the epicondyle is deep, narrow, and distinct. Of the 12 Myotis species living in Africa today (Simmons, 2005), only M. dieteri (Democratic Republic of Congo), M. morrisi (Ethiopia, Nigeria), M. scotti (Ethiopia), M. tricolor (subSaharan Africa), M. bocagii (western and southern Africa), and M. welwitschii (sub-Saharan Africa) are found south of the Sahara, and of these only M. welwitschii lives in Tanzania today. NMT.012/Bat represents a relatively large species of Myotis, such as M. welwitschii, and may well be related to, or even represent, that species. Myotis is poorly represented in the fossil record of Africa in the Plio-Pleistocene and is restricted to southern Africa (Broom, 1948; Pocock, 1987) outside of the record from Tanzania and an occurrence in Morocco (Gunnell et al., 2011). Myotis is a common element of other Plio-Pleistocene faunal communities and is widespread across the northern continents during this time period (Simmons and Gunnell, in prep.)., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on page 15, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press.","Broom, R. 1948. Some South African Pliocene and Pleistocene mammals. Annals of the Transvaal Museum 21: 1 - 38.","Pocock, T. N. 1987. Plio-Pleistocene fossil mammalian microfauna of southern Africa - a preliminary report including description of two new fossil Muroid genera (Mammalia: Rodentia). Palaeontologia Africana 26: 69 - 91.","Gunnell, G. F., T. P. Eiting, and D. Geraads. 2011. New late Pliocene bats (Chiroptera) from Ahl al Oughlam, Morocco. Neues Jahrbuch fur Geologie und Palaontologie 260: 55 - 71."]}

† Scotoecus olduvensis, new species Figures 6, 7 Cf. Pipistrellus (Scotozous) rueppelli, Butler and Greenwood, 1965: 15; Butler, 1978: 65; Gunnell, 2010: 588. HOLOTYPE: NMT.004/Bat, left dentary with c1–m3 (fig. 6B; see table 2 for measurements). REFERRED SPECIMEN: NMT.024/Bat, left distal humerus (also includes right proximal humeral fragment), FLK Main Dig 1, Z level (see table 1 for measurements). ETYMOLOGY: Named for Olduvai Gorge, Tanzania. TYPE LOCALITY: Tanzania: Olduvai Gorge, Bed I, FLK NI, Layer 3. DIAGNOSIS: Similar to extant Scotoecus albofuscus (fig. 6A) and S. hindei (fig. 6C) but differs from both in being, on average, 12% larger in tooth dimensions; S. olduvensis further differs from S. hindei in having: c1 lacking a buccal cingulid and anterolingual and posterolingual basal cuspules; a relatively longer posterior shelf that is not notched; p2 relatively larger compared to p4; p4 more similar in size to p2 (not larger as in extant taxa); p4 with a buccal cingulid lacking or very weak, a more robust cusp, and a lingual cingulid lacking an anterolingual extension. S. olduvensis further differs from S. albofuscus in having: c1 more robust, lacking a heavy lingual cingulid, with a more posteriorly extended posterior shelf that is not notched; p4 more similar in size to p2 (not larger as in extant taxon), lacking a lingual crest off the posterior part of the cusp that terminates in a posterior lingual extension of the lingual cingulid, which lacks an anterolingual extension, but has a better developed posterior shelf; S. olduvensis further differs from both extant taxa in having: p2 lacking a distinctive anterior extension of the anterolingual cingulid that forms a protrusion that fits into the notch of the canine posterior shelf; m3 with a somewhat lower hypoconid and entoconid and lacking a hypoconulid. DESCRIPTION: The lower canine of S. olduvensis is robust and relatively broad with a convex anterior surface, a flat posterior surface, and a lingual cingulid continuous with a low, moderate posterior shelf. The posterior shelf is overlapped by p2, which is single rooted and slightly wider than long but essentially circular in occlusal view. Anteriorly p2 overlaps the posterior shelf of the canine and has a continuous cingulid that is heaviest buccally. Lower p4 is similar to p2 but is slightly larger and relatively wider than long. It has a developed lingual cingulid, but the buccal cingulid is very weak. Like p2 there is only a single cusp developed on p4 that is slightly more robust than that found in p2. The lower molars are very similar to those of S. hindei and S. albofuscus: in having very tall and prominent talonid cusps and high entocristids, in being nyctalodont with well-developed crests, and in having relatively strong buccal cingulids. The lower molar trigonids are nearly as wide as talonids, in contrast to the living species in which the trigonid is noticeably more restricted buccolingually. The bases of the hypoconid and entoconid are inflated producing a relatively narrow and restricted talonid basin compared to the extant taxa. The distal end of NMT.024/Bat, the left distal humerus that we refer to S. olduvensis (fig. 7, bottom row), has a narrow but robust capitulum that is slightly angled and not offset from the shaft. The lateral capitular tail, even though somewhat broken, appears to have been much less broad than trochlear surface and is essentially continuous with the capitulum (shallow to absent groove). The trochlear groove is weak and the trochlear surface is relatively wide and robust with a sharply defined trochlear lip. The medial epicondylar process is broken and the groove on lateral surface of epicondyle is weak to absent. Also included with NMT.024/Bat is a potentially associated right proximal humerus. The head of this specimen is ovate and somewhat angled. The proximal margin of the head does not extend beyond the proximal extent of trochiter and is about at the same height as the proximal extent of the lesser trochanter. There is a high (anteriorly) deltopectoral crest. It is relatively long and sharply defined with a slight overhanging lip developed medially. The deltopectoral crest curves toward base of trochiter and anteriorly encloses a moderately deep fossa. No other fossils of Scotoecus have been described or figured (Simmons and Gunnell, in prep.).

Cf. Pipistrellus sp. REFERRED SPECIMENS: NMT.030/Bat, right distal humerus, FLK NNI, layer 2 or 3; NMT.040/ Bat, left distal humerus, FLK NI, layer 3, Tr. IV, 1964 (see table 1 for measurements). DESCRIPTION: Two fragmentary distal humeri from Olduvai represent pipistrelles, similar in size and morphology to Pipistrellus nanulus and P. rueppelli, the former known today from equatorial Africa and the latter from much of the African continent (Simmons, 2005). In anterior view the distal end of NMT.030/Bat has a narrow capitulum that is slightly angled and not offset from shaft. The lateral capitular tail is much less broad than the trochlear surface and is not separated from capitulum by a groove. The trochlear groove is moderately deep, the trochlear surface is relatively wide and robust, and the trochlear lip not sharply defined. The medial epicondylar process is small and extends distally only slightly beyond trochlear ridge. In posterior view, the groove on the lateral surface of the epicondyle is weak to absent. NMT.040/Bat differs slightly from NMT.030/Bat in lacking the slight angulation of the capitulum and in having a more sharply defined trochlear lip. Otherwise the two humeri are nearly identical. Pipistrelles are unknown from the fossil record in Africa outside Olduvai (Butler, 1978; Simmons and Gunnell, in prep.)., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 15-16, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press.","Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press."]}

Mops cf. M. thersites Figures 12, 13 Molossidae indet. (in part), Butler, 1978: 66; Gunnell, 2010: 587. REFERRED SPECIMENS: NMT.014/Bat, right distal humerus, FLK N1, layer 3, no. 4; NMT.028/Bat, right distal humerus, FLK NNI, layer 2; NMT.029/Bat, left complete humerus, FLK NNI, layer 2; NMT.033/Bat, left distal humerus, FLK NI, layer 3, Tr. III (see table 1 for measurements). DESCRIPTION AND DISCUSSION: The Olduvai specimens referred above are very similar in size and morphology to the humeri of extant Mops thersites, which is today known from western and central equatorial Africa south and east to Mozambique and Zanzibar (Simmons, 2005). The proximal end of NMT.029/Bat (fig. 12E) is similar in shape and proportions to all extant molossids except that head on the Olduvai specimen is relatively larger, especially proximodistally. The head is ovate with the long axis oriented slightly proximomedially to distolaterally. The proximal margin of the head is situated well below the proximal extent of the trochiter, but is even with the proximal extent of lesser tubercle. The deltopectoral crest is high anteriorly and relatively long. It is a very sharply defined crest with a well-developed overhanging lip developed medially, a crest that curves laterally toward the trochiter at its proximal end and forms the anterior border of a deep intertubercular fossa. The distal end of the humerus is preserved in four specimens of Mops cf. M. thersites (figs. 12, 13). The capitulum is narrow, angled, and not offset from shaft. The lateral capitular tail is slightly less broad than the trochlear surface, and is separated from capitulum by a moderate groove. The trochlear groove is weak, the trochlear surface is relatively broad, and the trochlear lip is not sharply defined. The medial epicondylar process in NMT.029/Bat extends a short distance past the trochlear ridge and only slightly less far distally than is seen in Mops thersites. The groove on lateral surface of epicondyle is either absent or only weakly developed. In general, molossids are rare in the African fossil record. There is an early Miocene Tadarida recorded from Kenya (Arroyo-Cabrales et al., 2002) and a late Pleistocene or Holocene record of Mormopterus (Sabatier and Legendre, 1985) from Madagascar and nothing else outside of the occurrences from Olduvai., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on pages 22-23, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press.","Arroyo-Cabrales, J., R. Gregorin, D. A. Schlitter, and A. Walker. 2002. The oldest African molossid bat cranium (Chiroptera: Molossidae). Journal of Vertebrate Paleontology 22: 380 - 387.","Sabatier, M., and S. Legendre. 1985. Une faune a rongeurs et chiropteres Plio-Pleistocene de Madagascar. 110 Congres national des Societes savantes, Montpellier 4: 21 - 28."]}

Mops cf. M. condylurus Figure 11 Molossidae indet. (in part), Butler, 1978: 66; Gunnell, 2010: 587. REFERRED SPECIMEN: NMT.031/Bat, right distal humerus from FLK NI, layer 3, Tr. III. DESCRIPTION AND COMPARISONS: The humeral specimens described below can all be recognized as molossids based on the presence of a relatively small and rounded humeral head that is positioned well below the trochiter, a bilaterally narrow and angled (proximomedially to distolaterally) capitulum, a bilaterally narrow trochlea separated from the capitulum by a moderate to distinct trochlear groove, an extended and robust capitular tail that is separated from the capitulum by a deep groove, and a well-developed and distally extended epitrochlear process. The specimen referred here to Mops cf. M. condylurus (as well as those referred to Mops cf. M. thersites below) resembles those of extant species of Mops in having an angled capitulum that is separated from the trochlea by a shallow yet distinct groove and from the lateral capitular tail by a deeper and well-formed groove. The epicondylar process is distinct but does not extend very far beyond the distal edge of the trochlea. Molossus differs from the Olduvai molossids in having: a less distinct groove between the capitulum and trochlea, a shallower groove between the capitulum and capitular tail, and a much more distally extended epitrochlear process (see Simmons and Geisler, 1998: fig. 33). The Olduvai molossids resemble Eumops in possessing a shallow groove between the capitulum and the trochlea but differ in having a narrower (proximodistally) lateral capitular tail that is separated from the capitulum by a shallower groove. The Olduvai molossids also are similar to Tadarida, Otomops, Myopterus, and Chaerephon in sharing a relatively broader and angled capitulum but differ in having a relatively more elongate trochlea (proximodistally). Additionally, the Olduvai molossids have a more robust and more distinctly separated capitular tail than either Myopterus or Otomops do. Olduvai specimen NMT.031/Bat represents a molossid very similar in size (see table 1 for comparative measurements) and morphology to extant Mops condylurus, which is today known from most of Africa south of the Sahara (Simmons, 2005). In anterior view, the distal end of NMT.031/Bat shows a narrow, medially angled capitulum that is not offset from the shaft. The lateral capitular tail is not as broad as the trochlea but is nearly as deep and is separated from the capitulum by a relatively deep groove. The trochlear surface is relatively broad and well defined, is separated from the capitulum by a weak groove, and has weakly defined medial trochlear lip. The medial epicondyle is not wide but is robust. The epicondylar process on NMT.031/Bat is broken so it is not possible to determine its extent. Posteriorly, the groove on the lateral surface of the epicondyle is very weak to absent. NMT.013/Bat differs from M. condylurus in having a relatively less deep trochlea and a somewhat wider capitulum, especially at its distal extent., Published as part of Gunnell, Gregg F., Butler, Percy M., Greenwood, Marjorie & Simmons, Nancy B., 2015, Bats (Chiroptera) from Olduvai Gorge, Early Pleistocene, Bed I (Tanzania), pp. 1-35 in American Museum Novitates 2015 (3846) on page 20, DOI: 10.1206/3846.1, http://zenodo.org/record/4598312, {"references":["Butler, P. M. 1978. Insectivora and Chiroptera. In V. J. Maglio and H. B. S. Cooke (editors), Evolution of African mammals: 56 - 68. Cambridge, MA: Harvard University Press.","Gunnell, G. F. 2010. Chiroptera. In L. Werdelin and W. J. Sanders (editors), Cenozoic Mammals of Africa: 581 - 597. Berkeley: University of California Press.","Simmons, N. B., and J. H. Geisler. 1998. Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments on the evolution of echolocation and foraging strategies in Microchiroptera. Bulletin of the American Museum of Natural History 235: 1 - 182.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, a taxonomic and geographic reference, vol. 1: 312 - 529. Baltimore: Johns Hopkins University Press."]}

Pteronotus quadridens (Gundlach, 1840) Figures 3, 5 MATERIAL EXAMINED: Cueva de Lily: 1 complete skull, 2 skull fragments, 1 dentary, 2 radii. Oleg’s Bat Cave: 1 complete skull, 1 dentary. EXTANT DISTRIBUTION: Cuba, Hispaniola, Jamaica, and Puerto Rico (Dávalos and Turvey, 2012; Simmons, 2005). FOSSIL RECORD: Pteronotus quadridens has previously been recovered from a Quaternary cave deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1). Additionally, P. quadridens has been found in Pleistocene or Holocene cave deposits in Cuba (Silva Taboada, 1974, 1979; Woloszyn and Silva Taboada, 1977) and the Bahamas (Andros, Great Abaco, and New Providence; Morgan, 2001). REMARKS: No consistent differences in cranial or postcranial morphology or size were found between our sample and the comparative material (appendix)., Published as part of Velazco, Paúl M., O'Neill, Hannah, Gunnell, Gregg F., Cooke, Siobhán B., Rimoli, Renato, Rosenberger, Alfred L. & Simmons, Nancy B., 2013, Quaternary Bat Diversity in the Dominican Republic, pp. 1-20 in American Museum Novitates 2013 (3779) on page 11, DOI: 10.1206/3779.2, http://zenodo.org/record/5364403, {"references":["Davalos, L. M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B. D. Patterson and L. P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical mammals: 157 - 202. Chicago: University of Chicago Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world: a taxonomic and geographic reference. 3 rd ed., 1: 312 - 529. Baltimore, MD: Johns Hopkins University Press.","Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with description of two new species (genera Pteronotus and Mormoops) and the first West Indian record of Mormoops megalophylla. Acta Zoologica Cracoviensia 19: 33 - 73.","Silva Taboada, G. 1979. Los murcielagos de Cuba. La Habana: Editorial Academia, Academia de Ciencias.","Woloszyn, B. W., and G. Silva Taboada. 1977. Nueva especie fosil de Artibeus (Mammalia: Chiroptera) de Cuba, y tipificacion preliminar de los depositos fosiliferos cubanos contentivos de mamiferos terrestres. Poeyana 161: 1 - 17.","Morgan, G. S. 2001. Patterns of extinction in West Indian bats. In C. A. Woods and F. E. Sergile (editors), Biogeography of the West Indies: patterns and perspectives: 369 - 406. Boca Raton: CRC Press."]}

Pteronotus macleayii (Gray, 1839) Figures 3, 5, 6 MATERIAL EXAMINED: Cueva de Lily: 4 radii. Oleg’s Bat Cave: 6 complete skulls, 2 skull fragments, 2 dentaries, 1 humerus, 3 femora. EXTANT DISTRIBUTION: Cuba and Jamaica (Dávalos and Turvey, 2012). FOSSIL RECORD: Fossil and subfossil remains are known from Pleistocene and Holocene deposits in Cuba and the Bahamas (Silva Taboada, 1974; Morgan, 1989). REMARKS: No consistent differences in cranial or postcranial morphology or size were found between our sample and the comparative material (appendix). This is the first record of P. macleayii for Hispaniola. Absence of this species from the modern fauna despite years of extensive collecting suggests that it has been extirpated from the island., Published as part of Velazco, Paúl M., O'Neill, Hannah, Gunnell, Gregg F., Cooke, Siobhán B., Rimoli, Renato, Rosenberger, Alfred L. & Simmons, Nancy B., 2013, Quaternary Bat Diversity in the Dominican Republic, pp. 1-20 in American Museum Novitates 2013 (3779) on page 10, DOI: 10.1206/3779.2, http://zenodo.org/record/5364403, {"references":["Davalos, L. M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B. D. Patterson and L. P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical mammals: 157 - 202. Chicago: University of Chicago Press.","Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with description of two new species (genera Pteronotus and Mormoops) and the first West Indian record of Mormoops megalophylla. Acta Zoologica Cracoviensia 19: 33 - 73.","Morgan, G. S. 1989. Fossil Chiroptera and Rodentia from the Bahamas, and the historical biogeography of the Bahamian mammal fauna. In C. A. Wood (editor), Biogeography of the West Indies: past, present and future: 685 - 740. Gainesville: Sandhill Crane Press."]}

Pteronotus parnellii (Gray, 1843) Figures 3, 5 MATERIAL EXAMINED: Cueva de Lily: 4 complete skulls, 1 skull fragment. Oleg’s Bat Cave: 19 complete skulls, 6 dentaries. EXTANT DISTRIBUTION: Cuba, Jamaica, Puerto Rico, Hispaniola, Saint Vincent, and possibly Trinidad and Tobago (Dávalos and Turvey, 2012; Clare et al., 2013). FOSSIL RECORD: In Hispaniola Pteronotus parnellii has been recovered from fossilized owl pellets (Diquini and Gonâve Island) in Haiti and from a Quaternary deposit (Cerro de San Francisco) in the Dominican Republic (fig. 2; table 1). Additionally, P. parnellii sensu stricto (see below) has been found in Pleistocene or Holocene cave deposits in Antigua, the Bahamas (New Providence), Cuba, Grand Cayman, Jamaica, Puerto Rico, and Tobago (Martin, 1972; Morgan, 1989, 2001). REMARKS: No consistent differences in cranial morphology or size were found between our sample and comparative material from the modern fauna of Hispaniola (appendix). The taxonomy and biogeography of bats of the Pteronotus parnellii complex is currently in a state of flux. Although traditionally recognized as a single species ranging through the Greater Antilles and from Mexico south to Peru and Brazil (e.g., Simmons, 2005), recent authors have found multiple diagnosable species within what was once called Pteronotus parnellii. Morphological and molecular studies have demonstrated that this complex includes at least five species and perhaps more, only some of which seem to correspond to previously delimited subspecies (Gutiérrez and Molinari, 2008; Clare et al., 2013). Because the holotype P. parnellii is from Jamaica and Antillean populations are typically much smaller than mainland forms, it seems likely that the name P. parnellii properly applies to all these bats including those from Hispaniola., Published as part of Velazco, Paúl M., O'Neill, Hannah, Gunnell, Gregg F., Cooke, Siobhán B., Rimoli, Renato, Rosenberger, Alfred L. & Simmons, Nancy B., 2013, Quaternary Bat Diversity in the Dominican Republic, pp. 1-20 in American Museum Novitates 2013 (3779) on page 10, DOI: 10.1206/3779.2, http://zenodo.org/record/5364403, {"references":["Davalos, L. M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B. D. Patterson and L. P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical mammals: 157 - 202. Chicago: University of Chicago Press.","Clare, E. L., et al. 2013. Diversification and reproductive isolation: cryptic species in the only New World high-duty cycle bat, Pteronotus parnellii. BMC Evolutionary Biology 13: 26.","Martin, R. A. 1972. Synopsis of late Pliocene and Pleistocene bats of North America and the Antilles. American Midland Naturalist 87: 326 - 335.","Morgan, G. S. 1989. Fossil Chiroptera and Rodentia from the Bahamas, and the historical biogeography of the Bahamian mammal fauna. In C. A. Wood (editor), Biogeography of the West Indies: past, present and future: 685 - 740. Gainesville: Sandhill Crane Press.","Morgan, G. S. 2001. Patterns of extinction in West Indian bats. In C. A. Woods and F. E. Sergile (editors), Biogeography of the West Indies: patterns and perspectives: 369 - 406. Boca Raton: CRC Press.","Simmons, N. B. 2005. Order Chiroptera. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world: a taxonomic and geographic reference. 3 rd ed., 1: 312 - 529. Baltimore, MD: Johns Hopkins University Press.","Gutierrez, E. E., and J. Molinari. 2008. Morphometrics and taxonomy of bats of the genus Pteronotus (subgenus Phyllodia) in Venezuela. Journal of Mammalogy 89: 292 - 305."]}

Monophyllus redmani Leach, 1821 Figures 4–6 MATERIAL EXAMINED: Cueva de Lily: 14 complete skulls, 14 skull fragments, 2 dentaries, 7 humeri, 9 radii. Oleg’s Bat Cave: 20 complete skulls, 9 dentaries, 8 humeri, 2 femora. EXTANT DISTRIBUTION: The Bahamas (Acklins, Crooked Island), Cuba, Hispaniola, Jamaica, Puerto Rico, Turks and Caicos Islands (Middle Caicos, North Caicos, and Providenciales) (Dávalos and Turvey, 2012). FOSSIL RECORD: Monophyllus redmani has previously been recovered from fossilized owl pellets (Diquini and Gonâve Island) from Haiti as well as from Quaternary cave deposits (Cerro de San Francisco) from the Dominican Republic (fig. 2; table 1). Additionally, M. redmani has been found in Pleistocene or Holocene cave deposits in the Bahamas, Cayman Islands, Cuba, Jamaica, Puerto Rico, and Middle Caicos in the Turks and Caicos Islands (Anthony, 1925; Koopman and Willliams, 1951; Williams, 1952; Koopman, 1955; Koopman and Ruibal, 1955; Choate and Birney, 1968; Silva Taboada 1974; Morgan, 2001). REMARKS: No consistent differences in cranial or postcranial morphology or size were found between our sample and the comparative material (appendix)., Published as part of Velazco, Paúl M., O'Neill, Hannah, Gunnell, Gregg F., Cooke, Siobhán B., Rimoli, Renato, Rosenberger, Alfred L. & Simmons, Nancy B., 2013, Quaternary Bat Diversity in the Dominican Republic, pp. 1-20 in American Museum Novitates 2013 (3779) on page 12, DOI: 10.1206/3779.2, http://zenodo.org/record/5364403, {"references":["Davalos, L. M., and S. Turvey. 2012. West Indian mammals: the old, the new, and the recent. In B. D. Patterson and L. P. Costa (editors), Bones, clones, and biomes. The history of recent Neotropical mammals: 157 - 202. Chicago: University of Chicago Press.","Anthony, H. E. 1925. Mammals of Porto Rico, living and extinct. In Scientific survey of Porto Rico and the Virgin Islands 9: 1 - 241. New York: New York Academy of Sciences.","Williams, E. E. 1952. Additional notes of fossil and subfossil bats from Jamaica. Journal of Mammalogy 33: 171 - 179.","Koopman, K. F., and R. Ruibal. 1955. Cave-fossil vertebrates from Camaguey, Cuba. Breviora 46: 1 - 8.","Choate, J. R., and E. C. Birney. 1968. Sub-recent Insectivora and Chiroptera from Puerto Rico, with the description of a new bat of the Genus Stenoderma. Journal of Mammalogy 49: 400 - 412.","Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with description of two new species (genera Pteronotus and Mormoops) and the first West Indian record of Mormoops megalophylla. Acta Zoologica Cracoviensia 19: 33 - 73.","Morgan, G. S. 2001. Patterns of extinction in West Indian bats. In C. A. Woods and F. E. Sergile (editors), Biogeography of the West Indies: patterns and perspectives: 369 - 406. Boca Raton: CRC Press."]}

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.

Type.-M 15021, sub-adult left dentary, edentulous symphysis, crowns of p2-m3, m3 in crypt (Table 1, Fig. 5A). Diagnosis.-Differs from A. zelteni in being larger (m2 area>60% larger), having a much longer molar series length (m1-3 ~ 108 mm in A. grex versus ~58 mm in A. zelteni), p4 with a well-developed talonid; differs from A. palustris (n. sp. below) in having smaller premolars (~25-30%), lower premolars with pustulate anterior and posterior crests, lowers premolars with distinct and elevated cingulids, p2 lacks development of an anterior crest. Etymology.-"Grex," Greek for flock or herd, in recognition of the herd behavior of many artiodactyls. Occurrence.-Early Miocene, Wadi Moghra, Egypt. Description.-Description of holotype specimen and occlusal details of teeth are available in Pickford (1991). Remarks.- Afromeryx grex resembles the better known A. zelteni from Gebel Zelten, in having an unfused symphysis, dental formula 3.1.4.3; no specializations of the anterior dentition, i1-3 are small spatulate teeth; no i3-c diastema, c-p1 diastema present, p1 single rooted, and mental foramina below i1, p2, p4. in all known comparable parts, A. grex seems to be a larger version of A. zelteni. As Pickford (1991) noted, the presence of pustulate crests on the premolars in species of Afromeryx is reminiscent of the condition seen in Gonotelma shabazi from the Bugti Beds, early Miocene, Pakistan, although what this resemblance means regarding the extent of a possible relationship between the two taxa has not been explored. The taxonomic history of Afromeryx is complex. Pickford (1991) named Afromeryx africanus on the basis of a right dentary with roots of p3, p4-m3 (CGM 30762), and attributed one additional sub-adult mandibular specimen (M 15021) to this species. The type specimen of A. africanus had previously been recognized as representing Brachyodus africanus (Andrews, 1899), but Pickford (1991) stated that, ������ The material does not closely resemble Brachyodus, the molars and premolars being bunodont, the jaw being short with an unspecialized symphyseal region������ (1991, p. 1503). In terms of size and degree of bunodonty, we find that CGM 30762 cannot be distinguished from members of Brachyodus depereti and so we have assigned the specimen to that taxon. In addition, it is unknown how the assessment was made that CGM 30762 represented a taxon with a short jaw and an unspecialized symphyseal region. The dentary is broken in front of p3 and so is missing the anterior portion of the jaw. This is critical because many of the features that distinguish anthracothere species are located in the anterior dentition. For example, species of Afromeryx have an unspecialized symphyseal region (i1-3 small, spatulate teeth, no i3-c diastema, p2 single-rooted), whereas species of Brachyodus show a number of extreme specializations (suppression of i1-2, i3 tusk-like, peglike canine, long diastema). The fact that the holotype of A. africanus designated by Pickford (1991) is a dentary with only roots of p3 and crowns p4-m3 means that most of the key diagnostic criteria of Afromeryx are not visible on this specimen. Therefore, we feel that CGM 30762 cannot serve as the holotype of a species of Afromeryx. To rectify this situation we have reassigned CGM 30762 to Brachyodus depereti where it fits comfortably based on size and known morphology and we have assigned M15021 as the holotype and currently only know specimen of a new species of Afromeryx, A. grex., 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 page 974, DOI: 10.1666/13-122

Anthracotheriidae species indeterminate The following specimens from Wadi Moghra are placed in an indeterminate group because either the fragmentary nature of the material, extreme wear, serious abrasion, or a combination of all three prevents assigning the specimen to a known taxon. DPC 2504, left dentary fragment; DPC 3629, left maxilla M2-3; DPC 4245, edentulous partial palate left P2-3, right P2-4; DPC 4421, left dentary dp4-m2, p4, p4 in crypt; DPC 4434 right dentary m3 fragment; DPC 4437, left dentary p2, dp3-4; DPC 4585, left dentary fragment with tooth fragments; DPC 6426, right m3; DPC 8932, right edentulous dentary fragment; DPC 8933, right MX; DPC 8952, right edentulous dentary fragment; DPC 12593, left dentary symphysis, c, dp4-m1, m2 in crypt; DPC 12930, left dentary dp3-4; DPC 14576, left dentary dp3-4, m1 in crypt; DPC 14577, right m3 fragment; DPC 17705, left edentulous dentary fragment; DPC 17734, right dentary with m3., 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 page 978, DOI: 10.1666/13-122

Afromeryx palustris new species Type.-CGM 83751, right dentary with symphysis, alveoli i1- p1, crowns p2-4 (Table 1, Fig. 5B-5D). Diagnosis.-Differs from A. zelteni and A. grex in being much larger (m2 area>70% larger than A. zelteni, m2 area>25% larger than A. grex); further differs from A. grex in having premolars with weaker pustulate crest development, premolars lacking distinct and elevated premolar cingulids, taller and more sharply tapering p2-3, p2 with better developed anterior crest. All premolar crowns have very finely wrinkled enamel. Etymology.-"Palustris,'" marshy or swampy, from the Latin ‘‘palus,’’ marsh, in recognition of the anthracothere habitat at Moghra. Occurrence.-Early Miocene, Wadi Moghra, Egypt. Material.-DPC 2542, right dentary with symphysis, p3-m1; DPC 4244 right dentary p2-3; DPC 4604, left dentary p4-m3; DPC 12593, right dentary dp4-m1, m2 in crypt. Description.-The dentary, as in other anthracotheres, has an unfused symphysis. It angles inferioposteriorly and is relatively short, not extending beyond p1. The dentary is deepest just posterior to m3 and has a rounded angle. The dentary tapers gently to the level of m1-p4 and then tapers more dramatically anteriorly. Incisors and canine: alveoli for three lower incisors are present, and judging from their size these would have been small teeth. The incisors are followed by an alveolus for a relatively small canine, which is separated from the single-rooted and relatively small p1 (see Fig. 5D) by a modest diastema (ranging from 20-30 mm in length). p2: relatively robust with a tall and tapering protoconid, a convex labial surface, and concave lingual surface. The labial aspect is covered with crenulated enamel that extends lingually around the anterior and posterior surfaces. There is a welldeveloped anterior crest extending from the tip of the protoconid along the anterolingual margin-there are three well-developed accessory cuspules arrayed down the lingual surface of the crest. The posterior portion of the tooth is broken but it is clear that there was no talonid development. A weak and low lingual cingulid is present but it extends only about half the length of the tooth, from the anterior crest posteriorly. Weak antero- and posterolabial cingulids are present. p3: is generally similar to p2 but differs in several important respects. Both the anterior and posterior surfaces are more extensive with the posterior surface having a strong ridge extending down the crown to reach a relatively well developed cingulid. In addition, the cingulid wraps around the base of the tooth both labially and lingually, but it is not continuous with a similarly developed but less extensive anterior cingulid. As on p2, there is no talonid development but the posterior cingulid on p3 is better developed and forms a weak shelf. A heavy anterior crest extends to the base of the tooth-this crest is weakly cuspate, showing development of bulbous but indistinct cuspules formed along its length. There is no distinct paracristid extending to the protoconid. A slightly weaker and less robust posterior crest extends to the tooth margin. This posterior crest is not cuspate and, while robust, is not as heavy as the anterior crest. A narrow, deep, dorsoventrally elongate and mesiodistally constricted valley extends between the anterior and posterior crests. A weak, low and short lingual cingulid closes the base of this valley. The tooth crown is covered by complex, rugose enamel. p4: is a more elongate version of p3, differing principally by having a distinct and broad talonid region that extends posteriorly as a flat shelf. The talonid shelf is weakly divided by a vague crest extending down the posterior surface and bifurcating with one branch extending to the talonid and the other terminating at the metaconid. The anterior and posterior lingual crests are slightly less robust than on p3 and are more angled anteriorly and posteriorly, respectively, producing a less deep, but equally extensive, lingual valley (CGM 83751, DPC 2542). However, this development appears variable as other specimens (e.g., DPC 4604) show more closely juxtaposed anterior and posterior crests that turn toward each other dorsally, producing a very narrow and restricted lingual valley. In DPC 2542 there are no cuspules developed on the anterior and posterior crests, while in DPC 4604 and CGM 83751, bulbous cuspules are present. The lingual cingulid is better developed on p4 than the other premolars, and produces a sloping lingual shelf that closes the base of the lingual valley. p4 has no crest connecting the paraconid and protoconid, but there is a weak crest connecting the metaconid to the apex of the protoconid. A much better developed anterior cingulid that forms a low but distinct shelf is present on p4. Molars: the molars of Afromeryx palustris are poorly represented but, where known, differ little from a common bunoselenodont anthracothere pattern. Lower first and second molars have trigonids and talonids of equal size and disposition. The cristid obliqua nearly reaches the lingual border producing a deep hypoconid notch that is closed ventrally by a hypocristid. Lingual cusps are aligned along the lingual margin, except for the small hypoconulid that is positioned lingual of center. There are weak pre- and postcristids but these are not labially continuous with the hypocristid. No lingual cingulids are present. The third lower molar appears to be a more robust version of m2, but with a large hypoconulid extension that forms a weakly oblique, single loop that is separated from the talonid by a deep labial valley. Like the premolars, all molars are covered by finely wrinkled enamel. Remarks.-Material assigned to A. palustris resembles other species of Afromeryx in having an unspecialized symphyseal region, a single-rooted p1, and in placement of the mental foramina. At present there are few comparable parts across all species, but the three species of Afromeryx (A. zelteni, A. grex,A. palustris) differ greatly in size, premolar proportions, and elaboration of the premolar series. To investigate more fully the nature of the relationship between the larger (A. palustris) and smaller (A. grex) specimens from Moghra, we looked at m2 dimensions. Specifically we examined the square root of m2 area in a range of extant male and female artiodactyl species, to test hypotheses about whether the size difference between the m2 of A. palustris and that of A. grex was on a par with what might be expected for males and females within a species, two species in a genus, or perhaps between individuals in two different genera. In general, artiodactyls are not particularly sexually dimorphic in the size of their postcanine dentition. Instead, sexual dimorphism in artiodactyls is primarily manifest in body size differences, elaboration of horns or antlers, and sometimes also in canine size and pelage characteristics (e.g., presence of a dewlap). However, we chose to use m2 dimensions partially because m2 size was one of the few attributes available in both A. palustris and A. grex, but also because the postcanine dentition in artiodactyls is morphologically conservative with regard to sexual dimorphism, so that a large degree of difference between males and females in this feature would be a clear result. Table 3 presents summary statistics involving m2 proportions in 14 artiodactyl species compared with the same proportions in A. grex and A. palustris. The raw data from which these calculations are derived are available as Appendix 1. Table 3 shows the average m2 area square root for males divided by that of females. This ratio is used to approximate the degree of m2 sexual dimorphism present among the extant sample. The values for the extant sample are then compared with the m2 square root ratio of A. palustris (DPC 4604) versus A. grex (M 15021). Results show that the ratio of m2 size in A. palustris to A. grex is outside what might be expected for males and females in a single artiodactyl species. The ratio for A. palustris to A. grex is about 1.17 while members of the extant sample fall within a fairly restricted range of 0.92 to 1.08. The ratio of A. palustris to A. grex also exceeds what might be expected for two species in the same genus, as the values for Kobus ellipsiprymnus versus K. kob were 0.92 to 1.04, and Hippotragus equinus versus H. niger ranged from 0.99 to 1.03. In fact, based on the data presented here, the ratio of A. palustris m2 area to that of A. grex is even greater than might be expected for members of two different artiodactyl genera. However, we recognize A. palustris and A. grex as two species of Afromeryx on the basis of anterior tooth morphology, and also because the ratio of m2 size in A. palustris versus A. grex does not exceed that which might be expected between a large male of one species and a small female of a second species within the same genus. For example, Appendix 1 shows that the m2 ratio of a large male Hippotragus equinus (NMNH 164790) compared with that of a small female of H. niger (NMNH 545318) is about 1.2, and a large male Kobus ellipsiprymnus (NMNH 61731) compared with a small female of K. kob (NMNH 164784) is about 1.54, ratios that are either on a par with or exceed the ratio of 1.17 observed for A. palustris to A. grex.

Type.-CGM 30780, left dentary with p3-m3, (Fig. 3A, 3B) (also figured by Fourtau, 1920, fig. 1). Diagnosis.-A small species of Sivameryx, length of M1-3 59 mm (S. africanus comparable measurement 80 mm), differs from Asian Sivameryx species in having lower molar paracristid extending to lingual margin and in possessing a lingual postprotocrista on upper molars that extends distally to the transverse valley. Occurrence.-Early Miocene, Wadi Moghra, Egypt. Material.-CGM 30781, left dentary p3-m3 (m3 broken); CGM 39779, left dentary m3; CGM 73692, left dentary m2; CGM 73679, edentulous left dentary; CGM 84434, edentulous right dentary with symphysis; CUWM 30, left dentary, m2-3; CUWM 31, left maxilla M2-3; CUWM 64a, right maxilla P2-M3, CUWM 64b left M3; CUWM 70, right maxilla M1-2; CUWM 100a, left dentary alveoli pl-ml, crowns m2-3; CUWM 100b, left dentary m2-3; CUWM 172, skull; DPC 3421, right dentary m1-2 fragments; DPC 3932, right dentary eroded m2-3; DPC 4066, right maxilla M2-3; DPC 4067, right dentary p3-m1 (abraded); DPC 4425, right dentary m1-3: DPC 4434, right dentary with broken m3; DPC 4452, left dentary with broken m1-3; DPC 4535, right maxilla P3, P4; DPC 5968, right m3 fragment; DPC 5973, left dentary p1-2; DPC 5979, left M2; DPC 6234, left M1; DPC 6243, right maxilla P3-M2; DPC 6289, left maxilla M1-3, M1-2 broken; DPC 7281, right M2; DPC 7659, left maxilla M3: DPC 8931, right maxilla fragment; DPC 8972, right dentary p3- m3; DPC 9039, left dentary p1-dp4 roots; DPC 12538, left dentary alveolus for p1, broken crowns p2-4; DPC 12546, right dentary m2-3; DPC 12547, right maxilla P4-M1 (abraded); DPC 12605, left dentary p4-m3, m1-2 broken; DPC 12608, left dentary m1-3, m1 broken; DPC 12609, right dentary p4, m2; DPC 12610, left dentary p3-m3; DPC 12632, left M3 fragment; DPC 12940, right dentary p1-2, p4-m1, m3 abraded; DPC 14547, right dentary m2-3; DPC 14559, right dentary p1-3; DPC 17680, left dentary p4-m2; DPC 17682, left dentary m1-2; DPC 17685, left maxilla M2-3; DPC 17687, left dentary fragments; DPC 17701 right m3 fragment; DPC 17710, dentary fragments; DPC 17711, right dentary roots of p4; DPC 17714, left maxilla P4-M3, M1 and M3 broken; DPC 17742, right dentary p4-m3; DPC 21506, right maxilla M3 (Tables 1, 2). Description.- Sivameryx has been described and discussed previously (Black, 1978; Fourtau, 1918, 1920), but most recently by Pickford (1991) who noted important distinguishing features of the genus such as: quasi-penticuspidate upper molars featuring loop-like parastyles and mesostyles, protoconule is almost fused with the protocone, canines are markedly sexual dimorphic, p2 is double-rooted, the dentition is selenodont, and the m3 talonid is loop-like and strongly obliquely oriented. Pickford (1991, p. 1511) noted that S. moneyi was mostly represented by lower jaws and the species appeared to be ������ in all its known parts, merely a smaller version of S. africanus " from Gebel Zelten. Recovery of CUWM 172, a nearly complete adult skull of S. moneyi, greatly expands our understanding of the taxon. Because the focus of this paper is on the diversity of anthracotheres found at Wadi Moghra, we provide an initial description of this Sivameryx skull here. Detailed study of the skull and its implications for larger issues involving the functional morphology, systematics, and paleobiogeography of anthracotheres in general is beyond the scope of this work. The skull is well preserved (Fig. 4A-4D) and cranial sutures are distinct throughout allowing for clear identification of key anatomical features. Nasal bones: the nasals are broken anteriorly but originate at anterior margin of orbit and are relatively broad posteriorly but tapering anteriorly as far as they are preserved. Frontal bones: the frontal bones are relatively short and wide, forming the roof of the skull and comprise about two-thirds of the medial wall of the orbit. Orbital processes are broken but are presumed to have been relatively robust judging by the remaining portion of their bases. The frontals extend rostrally as bilateral narrow splints separating the caudal part of the nasals from the lacrimals. The frontal has considerable contact with the lacrimal, palatine, and orbitosphenoid but only a very small area of contact with the maxillary bone. Dorsally the frontal forms distinct ridges that sweep caudally from the orbital processes to converge towards the sagittal crest. Orbit: the orbital mosaic is comprised of the lacrimal, maxillary, palatine, frontal, and orbitosphenoid bones, but is dominated by contributions from the frontal and palatine bones with a lesser contribution from the maxilla. The lacrimal foramen is small and located well within the orbit and the facial wing of the lacrimal extends relatively far anteriorly. The optic foramen is relatively large and enters the orbit through the orbitosphenoid. There are two smaller foramina anterior to the canal, a very small one in the frontal and a larger one formed on the maxillopalatine suture. Parietal bones: a pronounced sagittal crest is present superiorly on the parietals, and the crest extends caudally onto the occiput where it helps form the most caudal part of the skull. The parietals are rounded laterally and demarcate a relatively small braincase that tapers anteriorly towards the parieto-frontal contact as well as caudally toward the parieto-occipital suture. Ventrally and caudally, the parietals converge to help form part of the nuchal crest, and large gutters for the temporalis muscle are found caudal and lateral to the braincase, extending along the lateral margins of the parietal and temporal bones. Temporal bones: the temporal forms the basal and lateral portion of the braincase. The temporal extends and contacts the parietal dorsally, the orbital wing of the sphenoid rostrally, and the basisphenoid ventrally. At the juncture of the basisphenoid and temporal is the internal auditory meatus and slit-like formen lacerum medium that is continuous with that for the internal auditory meatus. The bony auditory bulla is large and bulbous, and has a distinct and elongate external auditory tube that extends caudally and dorsally, as in some suids, peccaries and hippopotami. Just rostral to the bulla is a distinct foramen ovale formed in the caudal-most aspect of the sphenoid. Caudal to the bulla is a small posterior lacerate foramen. The temporal contacts the lateral wing of the basioccipital in a relatively long dorsoventrally extend suture. The glenoid surface is flat, and almost as wide as it is long. There is a distinct postglenoid process but both an anterior process and a postglenoid foramen are absent. Dorsocaudally the temporal bone forms a robust portion of the lateral aspect of the nuchal crest. Occipital bone: the inferior portion of the occipital is occupied by large, robust occipital condyles. Anterior and lateral to the condyles, two bony wings sweep forward to form dorsoventrally flattened surfaces. The rostral portion of these surfaces form a wall caudal and lateral to the auditory structures. These structures also extend inferiorly to form small anteroposteriorly compressed mastoids, although no stylomastoid foramina are evident. Hypoglossal foramina are present rostral to the condyles, and small but distinct jugal foramina are located inferiorly to the nuchal crest and lateral on the basioccipital. Sphenoid bone: ventrally, the sphenoid has a distinct central ridge that continues rostrally to meet a sharply defined and high vomer. Caudally the basiosphenoid is bifurcate with bilaterally developed low processes extending caudally to the internal auditory meatus. Maxillary bone: the maxilla is robust and forms a rostrum that is dorsoventrally relatively deep. However, the maxilla is transversely compressed just rostral to the zygoma, so that the rostrum appears relatively deep but mediolaterally fairly narrow. The rostrum is concave and narrowest superior to P4 and then broadens modestly rostrally. The concave portion of the rostrum is occupied by a small infraorbital foramen positioned superior to the posterior root of P3. From the infraorbital foramen, the maxilla extends a short distance caudally to a point superior to M3, where the bone terminates in a pointed process that is notched medially and ventrally, near where the maxilla meets the palatine. The maxilla forms the floor of the orbit and encloses the infraorbital foramen. The portion of the palate formed by the maxilla is relatively long and narrow, but not as elongate as in Afromeryx or Brachyodus. The incisive foramina are broken rostrally so their extent is hard to estimate. The caudal margin of these foramina does not extend past I2. There is a paired set of palatal foramina present at the rostral margin of P3 and a second very small pair of palatal foramina present at the rostral margin of M3. Zygomatic bones: the rostral roots of right and left zygoma are present but most of the zygoma are absent. Palatine bones: the palatine extends rostrally as a broad plate with a narrow central ridge until it reaches the rostral margin of M3, where it tapers to a point that extends to the rostral margin of M2. The palatine extends caudally well past the tooth row to the caudal rim of the orbit, and a distinct post-palatine process is present. Dentition: the dentition is represented by right P4-M3, left P3 (broken), and P4-M3. All teeth except right P4 and right M3 are heavily worn. The anterior premolars are represented only by alveoli and it is difficult to be certain of homologies because supernumerary teeth were present, not only a common occurrence in Libycosaurus (Lihoreau et al., 2006; Pickford, 2006) but also in anthracotheres in general (personal observ.). The right dentition anterior to P4 has 9 alveoli. We interpret these to represent P? (double-rooted), P1 (double-rooted), P2 (double-rooted), and P3 (triple-rooted). The left side of the palate has six alveoli anterior to P3 which we interpret to represent the homologs of the right side. Recovery of CUWM 172 confirms a number of features previously cited as characteristic of S. moneyi and also reveals some new details. The new skull confirms that P1 and P2 are double-rooted (Pickford, 1991). The occurrence of supernumerary premolars in S. moneyi has not been documented previously but extra premolars seem to be a fairly common occurrence among anthracothere species in general (personal observ.)., 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 970-974, DOI: 10.1666/13-122