Carnivora. Systematic Paleontology THIRTY-TWO

Transcription

1 THIRTY-TWO Carnivora LARS WERDELIN AND STEPHANE PEIGNE The order Carnivora has a shorter history in Africa than on any other continent except Australasia and South America. The definite record of the order on the African continent extends back to the Lower Miocene, though some earlier records may exist (discussed later). During this time, the order has diversified enormously, first as a result of migrations from Eurasia and later as a result of in situ speciation. Despite this, our knowledge of the history of African Carnivora still is poorer than for most continents, mainly due to the geographically biased fossil record on the African continent. For the Plio-Pleistocene, only parts of northern, eastern, and southern Africa have an adequate Carnivoran fossil record, and for the Miocene the situation is much worse, as only some time slices of this epoch have an adequate record in some parts of eastern Africa, with most of the rest of the continent simply a white spot on the map. Nevertheless, this review encompasses more than 100 genera and about twice that many species. The organization is by family (in standard order) and genus (in alphabetical order), with a series of subheadings providing the bulk of the information. These are as follows: Diagnosis We have tried to provide reasonable diagnoses of all genera. In most cases these have been taken from the original publications or from subsequent revisions. Many extant genera are diagnosed on the basis of soft-tissue characters, and for these we have tried to present provisional diagnoses based on craniodental information. These diagnoses should be treated as general indications only. African Species A list of the species of each genus that are known from the African fossil record. When "Genus sp." is listed, this means that an unnamed species is known to differ significantly from all named species, or at least cannot comfortably be included in the named species. Age The approximate first and last appearance datums for each African genus (not including extra-african occurrences). Geographic Occurrence An alphabetical list of the African countries in which each genus has been found (again, extra African occurrences are not included). Locality data are provided in the tabular material. Remarks Any comments of a mainly taxonomic nature that we have found to be relevant in our study of the various taxa. We conclude with short sections on biogeography and migration patterns, based on the data we have collected in creating the review. Systematic Paleontology Family AMPHICYONIDAE Haeckel, 1866 Table 32.1 Genus AFROCYON Arambourg, 1961 Figure 32.1 Diagnosis Revised from Arambourg (1961). Amphicyonid of large size, comparable to Amphicyol1 giganteus or A. shahbazi; p4 simple with slightly enlarged talonid and distal accessory cuspid; m1 voluminous with posteriorly located metaconid, relatively short talonid, large hypoconid in buccal position; m2 with well-developed protoconid and hypoconid, entoconid reduced; m3 longer than wide, bilobed and two-rooted; mandibular corpus very tall and narrow. African Species A. bllrolleti Arambourg, Age Ca Ma. Geographic Occurrence Libya. Remarks The single species of the genus is known from a fragmentary and poorly preserved left hemimandible with FIGURE 32.1 AfroC)'Oll burolleti, type specimen (MNHN, no number) in buccal and occlusal vievvs. 603

2 TABLE 32.1 Occurrences of Amphicyonidae species Sites....; Q) <::l :::l "0. :s.s co ;::;.l::l ::::.9< P, :::: Q) Q) :::: <::l co :S <::l co l:l '-.l "'OJ co <::l 2 ". "0 "0 "'OJ s:: e 2;. b{) s:: s:: s:: '2 '2 '-.l :::: '0' 1; :::l :::l :::l :::: 0 0 <::l ::::. P, :::: "'OJ. s::. co 'E Ci' ( s::.s :s <::l u u.:::: :::l.s... :::l.::::... '5 :E :2 :2.:::1..s..s..s..s co Ci' co co E' S 0..;::: <u <u ;:0 :::: '- :::: S :::: :::: :::: :::: :::: <::l <u <.i::.. b{)'e b{) co >-.. <=q G U G G <::l Arrisdrift Beni Mellal Bled Douarah Beglia Buluk Chamtwara Elisabethfeld Escarpment (Gona) Fejej Fiskus Fort Ternan Grillental Hamadi Das (Gona) Hiwegi R 1 Hiwegi R 3 ct. ct. Hiwegi R 5 Hondeklip Bay Jebel Zelten ct. Kalodirr Kiahera Hill Kipsaraman ct. Koru Kulu Langental Legetet Lemudong'o Lothagam Lower Nawata Lothagam Upper Nawata Maboko Malembe Mfwangano Moroto II? Napak Ngorora Member D Nyamsingula Oued Mya 1 Rusinga Samburu Hills Namurungule Songhor Toros Menalla Wadi Moghra 604 LAURASIATHERIA

3 p4-m3. Its main distinctive feature is the presence of a double-rooted m3, which is unique among Amphicyoninae. Due the fragmentary nature of the holotype, detailed comparison with other taxa must await future discoveries. Genus AGNOTHERIUM Kaup, 1833 Diagnosis Revised from Kurten (1976). Amphicyonid of medium to large size, with felinoid characters: short snout, elongated upper canines, reduced anterior premolars, small, double-rooted P3, large P4 with small parastyle and reduced protocone; large Ml-2; high-crowned, large p4, large m1 with trenchant trigonid, no metaconid, and reduced talonid with only a trenchant hypoconid crest; m2 reduced relative to m1 and lacking the paraconid; jaw deep. African Species A. d. mztiqlltl11z Kaup, 1833, A. kiptalami Morales and Pickford, 2005, A. sp. Age Ca Ma. Geographic Occurrence Kenya, Morocco, South Africa, Tunisia. Remarks This genus shows a derived condition toward hypercarnivory, including reduced premolars and m2-3, m1 lacking metaconid and with a talonid formed solely by the hypoconid, and extreme reduction of the P4 protocone. The diagnosis proposed by Kurten (1976) includes the absence of m3. However, material from Steinheim (Helbing, 1929: figure 1) and Frohnstetten (Kuss, 1962) clearly shows the alveolus for m3. In addition, the isolated m2 from Beni Mellal (Morocco) assigned to a form close to A. mztiqllll11z by Ginsburg (1977) has a small facet on its distal face that indicates the presence of an m3. There is considerable size variation in African Agnotizerillm, from the early and small species from Fort Ternan to later, larger forms such as Agnotizerizl11z kiptalami from Ngorora D or Agnotizerizll1/ sp. d. A. mztiqllllj11 from Bled Douarah. The taxonomic status of the species from Fort Ternan is not yet clear. It is smaller than other Agnotizerill1ll and the m1 has a small metaconid (Morales and Pickford, 2005a; Werdelin and Simpson, 2009, fig. 2), whereas the metaconid is completely absent in all other known specimens of Agnotizerill1ll. Genus AMPHICYON Lartet, 1836 Diagnosis Translated and revised from Kuss (1965). Medium- to large-sized amphicyonids with dental formula I 3/3, C 1/1, P4/4, M 3/3; Pl-3/pl-3 rounded, short, lacking posterior accessory cusps and with strong basal cingulum; p4 with posterior accessory cusps and sometimes a weak anterior bump; P4 somewhat shorter than m1, generally with a parastyle and a retracted and reduced protocone; m1 relatively low-crowned, paraconid somewhat truncated and with an anterior crest, metaconid reduced but always present, talonid wide with a tall hypoconid; M1 triangular or distally somewhat concave, lingual cingulum mostly present only posterior to the protocone; M2 and M3 relatively large; M2 enlarged and as wide or slightly wider than M1; m2 more or less rectangular, longer than wide, with vestigial paraconid and talonid shorter and narrower than trigonid. African Species A. gigmztells (Schinz, 1825). Age Ca Ma. Geographic Occurrence Libya, Namibia. Remarks The genus is known from complete skeletons and many dental remains from North America and Europe. It is also known from many poorly characterized Asian species, the generic assignment of which requires confirmation (see review in Peigne et ai., 2006). Although rare in Africa, the genus potentially has a Pan-African distribution, since it is present in northern (Jebel Zelten; Ginsburg and Welcomme, 2002) and southern (Arrisdrift; Morales et ai., 2003) Africa. In both cases, remains have been assigned to Ampizicyol1 gigantells or a closely related species. Amplzicyon gigantells is a large-sized generalized species based on dental remains from the middle Miocene of France. Specimens from Libya (distal humerus and astragalus) are assigned to Amplzicyol1 sp. d. A. gigmztells on the basis of their overall size only, as these remains are not diagnostic at the species level in Ampizicyoll. The presence of A. gigantells at Arrisdrift is well supported, with a sub complete mandible with p4-m2 and some metapodials (Morales et ai., 2003). Genus BONISICYON Werdelin and Simpson, 2009 Diagnosis From Werdelin and Simpson (2009). Amphicyonidae of small size; carnassial shear on m1 entirely mesiodistal; m1 hypoconid an elongated crest, separated from trigonid by a narrow postvallid notch and effectively a part of the carnassial shear; m1 metaconid in evidence only as a bulge on the lingual side of the protoconid; m1 relatively wide and bulbous at the base of the crown; m2 broad and short. African Species B. illambo Werdelin and Simpson, Age Ca Ma. Geographic Occurrence Ethiopia, Kenya. Remarks This genus and species brings together material of a small amphicyonid from a number of sites. It includes the material from the Upper Nawata Fm., Lothagam described by Werdelin (2003) as Amphicyonidae sp. B as well as the tooth described as Si111ocyon sp. from Lemudong'o by Howell and Garcia (2007). Isolated teeth from Gona, Ethiopia, bear witness to the uniqueness of this taxon. Genus CYNELOS Jourdan, 1862 Diagnosis Translated and revised from Kuss (1965) and Hunt (1998a). Small- to large-sized amphicyonids with dental formula 13/3, C 1/1, P 4/4, M 3/3; incisors tend to have accessory cusps; strong canines; long and slender premolars; p4 with posterior accessory cuspid only; M2-3 and m2-3 enlarged, with M2 only slightly more reduced than M1, and with the paracone as large or only slightly larger than the metacone; lower jaw slender. African Species C. euryodol1 (Savage, 1965), C. macrodol1 (Savage, 1965), c. minor (Morales and Pickford, 2008), C. sp. Age Ca Ma. Geographic Occurrence Egypt, Kenya, Uganda. Remarks Cynelos is the most diverse amphicyonine genus with at least six species in North America and up to nine in Europe during the late Oligocene and Miocene, although there is no consensus about the generic assignment of some European species (Hunt, 1998a; Peigne and Heizmann, 2003). In Africa, Cynelos is by far the most common Amphicyonidae with two species. C. macrodon is known only from isolated teeth but C. ellryodon, the smallest African species, is wellknown from several early Miocene localities in Uganda and Kenya. The arrival of Cynelos coincides with the first wave of migrations of Carnivora to Africa. Recently, Morales et ai. (2007) proposed resurrecting Hecllbides for the African species. In our opinion, these authors demonstrate the distinction between Cyne/os ellryodon and C. le11lmzensis only. Given the fragmentary nature of the known material, we see no strong support for a generic distinction of African Cynelos. Genus ]viyacyon Sudre and Hartenberger, 1992 THIRTY-TWO: CARNIVORA 605

4 Diagnosis Translated and modified from Sudre and Hartenberger (1992). Amphicyonid of large size characterized by its sectorial molars; m1 large with an elongated talonid, strong protoconid with tall, strong trenchant anterior crest, paraconid indistinct and separated from protoconid by a very weak notch (visible on the buccal margin of the anterior crest), metaconid reduced and situated slightly posteriorly, talonid short, with a strong, crested hypoconid and a smaller, poorly developed entoconid situated far distally and lacking crest; m2 short and oblong, with protoconid trenchant, no parac.onid, poorly developed metaconid situated at the level of the protoconid, and talonid short and narrower than the trigonid, with a strong hypoconid but no entoconid; trigonid of m1 and m2 with a strong buccal cingulum. African Species M. dojambir Sudre and Hartenberger, Age Ca Ma. Geographic Occurrence Algeria. Remarks This species is represented by a fragmentary right hemimandible with m1-m2 (m3 not yet erupted). This is a very large species that reached the size of the largest species of Amphicyon, A. ingens from North America (Hunt, 2003). As previously pointed out (Sudre and Hartenberger, 1992), Myacyon has nothing to do with any of the known Amphicyoninae. It remains a geographically and morphologically isolated species in northern Africa. Genus YSENGRINIA Ginsburg, 1965 Diagnosis Modified from Hunt (1998a). Medium- to largesized amphicyonid with dental formula I 3/3, C 1/1, P 4/4, M 3/3. Pl-3/pl-3 low, reduced and lacking accessory cuspids; p4 tall with well-developed posterior accessory cuspid; robust, massive m1 trigonid with strongly reduced metaconid, talonid dominated by a centrally to buccally placed, prominent hypoconid crest and a reduced entoconid; M2-3/m2-3 not enlarged relative to Ml/m1 as in, for example, Amphicyol1; m2 elliptical in occlusal view, short, with large trigonid comprising a vestigial paraconid, strong protoconid, reduced metaconid, and low, short, posteriorly tapering talonid with a prominent hypoconid crest but no entoconid; mandibular corpus robust and tall, especially anteriorly. African Species Y. ginsbllrgi Morales et al., 1998; Ysengrinia sp. Age Ca Ma. Geographic Occurrence Namibia, South Africa. Remarks The only African species of Ysengrinia is known through dental and postcranial remains from Arrisdrift (Morales et al., 1998, 2003). There are many morphological differences between this species and the type species Y. geml1diana (Heizmann and Kordikova, 2000; Peigne and Heizmann, 2003), notably the more reduced size of p4 relative to m1 in the African species. The species assigned to Ysengril1ia do not really form a homogeneous group (especially with the inclusion of poorly known species such as Y. depereti and Y. valel1tial1a), and a detailed analysis shows many differences between them (Peigne and Heizmann, 2003). AMPHICYONIDAE indet. Age Ca Ma. Geographic Occurrence Angola, Chad, Ethiopia, Kenya, Uganda. Remarks The earliest and the latest occurences of the Amphicyonidae in Africa are documented by indeterminate remains. An isolated incisor, particularly difficult to assign precisely, is known from Malembe (Hooijer, 1963). Though assigned to d. Amphicyon, this tooth could equally belong to Cynelos. Aside from Bonisicyon illacabo, Lothagam includes a large amphicyonid (size of A. giganteus) from the Lower Nawata Fm., known from an upper molar and fragmentary postcranial elements. Other records of Amphicyonidae are mentioned in faunal lists that require confirmation. Thus,?C,l1Jelos sp. may be present in the early middle Miocene site of Moroto II, Uganda (Pickford et al., 2003), but here we consider it an undetermined amphicyonid. One exception is the recent description of new material from the Namurungule Fm. (Samburu Hills) that includes lower teeth of a large Carnivore assigned to Amphicyonidae or Ursidae (Tsujikawa, 2005). Illustrations of the specimens show that they belong to an amphicyonid. The author suggests close relationships to Agl1otiJerill1l1, but in our opinion, the m2 is much more similar to that of Ysengril1ia spp. (especially Y. gemndial1a and Y. americana) in having an elliptical outline, a posteriorly tapering talonid with a prominent, laterally placed hypoconid, and a distinct buccal cingulum. The m2 from Samburu Hills is, however, larger in size and more elongated than in Ysengrillia, and, above all, it comes from geologically much younger strata. In addition, the p4 of the same individual differs from that of Ysengrinia in lacking accessory cuspids. The absence of the posterior accessory cuspid on p4 is a derived feature of Pseudarctos bavariclls and Ictiocyon socialis, two much smaller species with an m2 that is morphologically distinct from the Samburu Hills amphicyonid. The species from Namurungule may represent a new species, but additional remains are necessary to confirm this hypothesis. Family URSIDAE Fischer, 1817 Table 32.2 Genus AGRIOTHERIUM Wagner, 1837 Diagnosis Modified from Hunt (1998c). Large-Sized ursine with dental formula I 3/3, C 1/1, P 3-4/2-4, M 2/3; sexually dimorphic; short-snouted robust skull, somewhat brachycephalic; palate wide; premolar tootluow much shortened with anterior premolars reduced in size, single rooted, and low crowned; P4 robust, with strong parastylar cusp and protocone shelf; M2 with only rudimentary talon; ml-2 cusp pattern variable; premasseteric fossa present; symphyseal region of lower jaw ventrally produced as a "chin"; long-footed, plantigrade limbs. African Species A. africal Hendey, 1980, A. aecuatorialis Morales et al., 2005, A. sp. Age Ca Ma. Geographic Occurrence?D. R. Congo, Ethiopia, Kenya, Libya, South Africa, Uganda. Remarks This genus has a stratigraphic range in Africa from the latest Miocene to the early Pliocene. Though generally rare, it was successful and spread through the entire continent. The Muishondfontein Pelletal Phosphorite Mb. of the Varswater Fm. of Langebaanweg has yielded a large number of cranial, dental and postcranial specimens of Agriotherill111 africamlln, representing a minimum of 14 individuals. The ursid from Sahabi, previously identified as Indarctos sp. (Howell, 1987), is probably a close relative of this species (Morales et al., 2005). The possible record of the genus from the upper member of the Sinda Beds of the Democratic Republic of Congo (late Miocene to Pliocene) is speculative given the available material (Yasui et al., 1992). Additional material is also known from early Pliocene sites in Ethiopia (L.W., pers. obs.). 606 LAURASIATHERIA

5 TABLE 32.2 Occurrences of Ursidae species Sites Afalou bou Rhummel Ahl al Oughlam Ain Bahya III Ain Rouina Ali Bacha Aramis Babors Bouknadel Boulhaut Breche entre Oran et Mers-el-Kebir Djebel Thaya Doukkala I Douar Debagh El Khenzira El Ksiba Fort Bourdonneau Gratte de l'akouker Gratte d'os (Djurdjura) Gratte de l'ours (Djurdjura) Gratte des Ours (Constantine) Gratte des Ours GO (Casablanca) Gratte du Mouflon (Constantine) Hadar Denen Dora Hadar Kada Hadar Khifan bel Ghomari Koobi Fora Lonyumun Koobi Fora Okote Koobi Fora Tulu Bor L'Anou Tenechiji L'lfri en Terga Roumi La Madeleine La Pointe Pescade Langebaanweg PPM Les Falaises Menacer Nachukui Lower Lomekwi Nkondo Oulad Hamida I-rhino cave Oulad Hamida Th 1II-H. erectlls Rusinga Sahabi Sagantole Fm. Sidi Abderrahmane Sinda Takouatz Guerrissene. ] E 2 t:l t:l ;:; u u "" :t t:l t:l E E E ;:; ;:; ;:; 't 'E '6..;:: l t..;::.... "" "" "" d. d. "" 'U 0::; '.8.,.; Q) ::l CI "" e t:l P. V>.S.8 "" "" B p. Q).8 t:l V>. ro :g "" E t:l "" ;:; ;:; "" ;:; "" 0::; Vl yo yo yo 'U H :r::..::; " ;::J :.3 :.3 :.3 aff.? d. cf. THIRTY-TWO: CARNIVORA 607

6 TABLE 32.2 (CONTINUED). :9 I-.8 <::l ;:;.g i;; ':5 <, a. Sites Tamar Hat Ternifine Thomas I-H. erectus cave Thomas II Thomas III Tighenif Toulkine Tugen Mabaget <::l " EO 8 ci. "1::i <::l V) 2...: Cl) ci..g.g :::: "0 e V) <::l.8.8 l- I- i:: a Cl) ::::.8 B ':5..s:: <::l <u... co a.2 'j " :::: "0 ::l. <::l 'Vj ;:; I- "1::i ;;; b<j.,. k.:; ;:J :5 :5 :5 Genus HEMIC YON Lartet, 1851 Diagnosis Translated and modified from Ginsburg and Morales (1998). Mid- to large-sized species of Hemicyoninae with dental formula I 3/3, C 1/1, P 4/4, M 2/3; m1 talonid simple and nearly symmetrical, talonid groove shallow and roughly axial; P4 with paracone anterobuccally inflated and without anterobuccal cingulum; M1 subrectangular; M2 tends to be oval in shape; P4 protocone elongated; lower premolars simple; m1 with a strong metaconid well separated from the protoconid; talonid shallow, with low hypoconid and lingual tubercles of the talonid poorly or not developed; m2 generally broader than m1, with protoconid and metaconid very prominent; m3 small, low, with only protoconid and metaconid developed. African Species H. sp. Age Ca Ma. Geographic Occurrence Kenya. Remarks Hemicyon belongs to the Hemicyoninae, traditionally considered a subfamily of Ursidae. It is known in Africa from a single tooth, a P4 from the early Miocene of Rusinga (Schmidt-Kittler, 1987), which is only slightly younger than the earliest, much smaller, Eurasian species of the genus, H. gargan (Ginsburg and Morales, 1998). Schmidt-Kittler (1987) rightly casts doubts on the generic assignment of the Rusinga tooth, arguing that his specimen compares well with an isolated P4 from Wintershof-West (early Miocene, Germany) that is now assigned to another hemicyonine, Plloberocyon dellmi (Ginsburg and Morales, 1998). Genus INDARCTOS Pilgrim, 1913 Diagnosis Modified from Hunt (1998c). Mid- to large-sized ursid, sexually dimorphic with dental formula I 3/3, C 1/1, P 4/4, M 2/3; skull dolichocephalic and snout short; P1/ p1-p3/p3 well developed in earlier species and reduced in size but nearly always present and single rooted in advanced species; P4 robust with parastylar cusp present but usually not as developed as in Agrioti1erillm; molars low crowned; M2 with elongate talon; protoconid -metaconid-entoconid -entoconulid of m1 aligned in smooth descending curve; premasseteric fossa absent; anterior part of lower jaw tapers forward, not as squared off and blunt as in Agrioti1erill1n; "chin" present only in old individuals; plantigrade. African Species T. aff. arctoides (Deperet, 1895). Age Ca Ma. Geographic Occurrence Algeria. Remarks Apart from the Hemicyol1 sp. from RUSinga, Il1darctos is the earliest ursid in Africa. It is known through a single record from Menacer, late Miocene of Algeria (Petter and Thomas, 1986). The genus has also been identified from Sahabi (Howell, 1987) but assignment of this material is debatable, and this record is here assigned to Agrioti1erill111. According to Howell (1987), two genera, AgriotiJerillm and Tndarctos, are present at Sahabi, mainly based on the size variability of the sample. However, size alone cannot be used to distinguish the strongly sexually dimorphic species of the two genera. It is therefore most probable that only one genus (AgriotiJerilll1l) is present at the Libyan locality. Genus URSUS Linnaeus, 1758 Diagnosis Modified from Hunt (1998c). Mid- to largesized ursid; sexually dimorphic, with dental formula I 3/3, C 1/1, P 2-4/1-4, M 2/3; skull dolichocephalic; no premasseteric fossa; m2 shorter than m1 in earlier forms but m2 later increasing to exceed m1 length; great elongation of m2-3/ Ml-2; Pleistocene forms exhibit twinning of metaconid of m1-m2 and considerable widening of m2-3 relative to mi. African Species U. cf. etrusclls Cuvier, 1823, U. arctos Linnaeus, 1758, U. sp. Age Ca. 2.5 Ma-Holocene. Geographic Occurrence Algeria, Morocco, Tunisia. 608 LAURASIATHERIA

7 Remarks UrSllS originated in Eurasia and reached Africa in the mid-pliocene. It is only known from the Maghreb, where it became extinct around the mid-19th century (Servheen et ai., 1998; Nowak, 2005, p. 124). The earliest record of the genus, Urslls sp. d. U. etrusclls, is from the mid-pliocene site of Ahl al Oughlam. The presence of a form close to this typical Villafranchian Eurasian bear in northwestern Africa supports a migration event from Europe, possibly through Spain and the Gibraltar Strait, where the species is known from as early as the early Pliocene of Layna (Fraile et ai., 1997). A large number of Pleistocene to Holocene records of Ursus spp. are known from northern Africa, especially from Morocco and Algeria (Hamdine et ai., 1998). URSIDAE indet. Age Ca Ma. Geographic Occurrence Algeria, Ethiopia, Kenya. Remarks At least one species of undescribed ursine is represented by postcranial material from several members at Hadar, Koobi Fora and West Turkana (Werdelin and Lewis, 2005). Family CANIDAE Fischer, 1817 Table 32.3 Genus CANIS Linnaeus, 1758 Diagnosis Modified after Munthe (1998). Canids of large size; shorter, broader face than Vllipes; wide zygomatic arches; width of skull across widest part of arches equal to at least half the length of the skull; frontal sinuses enlarged and invading the postorbital processes; incisors with accessory cusps and 13 enlarged; P4 about the same length as M1, no parastyle; m1 talonid with crest uniting hypoconid and entoconid. African Species C. adllstlls Sundevall, 1847, C. atrox Broom, 1948, C. allrells Linnaeus, 1758, C. brevirostris Ewer, 1956, C. falconeri Forsyth Major, 1877, C. mesomelas Schreber, 1776, C. pictus (Temminck, 1820), C. sp. Age Ca. 3.5 Ma-Recent. Geographic Occurrence Algeria, Ethiopia, Kenya, Morocco, South Africa, Tanzania, Zambia. Remarks The genus Canis (here including Lymon) is widely distributed in Africa today. Modern Canis are cursorial, openhabitat adapted taxa. This is likely to have been true in the past as well and may explain their relative scarcity as fossils, since such habitats seem to be underrepresented in the carnivoran fossil record of Africa. The record of Canis sp. from South Turkwei is, at ca. 3.5 Ma, one of the oldest for the genus outside North America (Werdelin and Lewis, 2000). Genus EUCYON Tedford and Qiu, 1996 Figure 32.2A Diagnosis Modified after Tedford and Qiu (1996). Skull with frontal sinus invading the base of the postorbital process, usually removing the "vulpine-crease" on the dorsal surface of the process; paroccipital process expanded posteriorly, usually with a salient tip; mastoid process enlarged into a knob or ridgelike prominence; lacking foxlike lateral flare and eversion of the dorsal border of the orbital part of the zygoma; lacking a transverse cristid connecting the hypoconid and entoconid of the m1 talonid; second posterior cusplet present on p4. African Species E. intrepidlls Morales et ai., 2005, E. 111inimus Haile-Selassie et ai., 2009, E. wokari Garcia, 2008, E. sp. (figure 32.2A). Age Ca Ma. Geographic Occurrence Ethiopia, Morocco, Kenya, South Africa. Remarks The genus Eucyon was originally erected for some Eurasian and North American species transitional between the primitive Leptocyon spp. and derived Canis (Tedford and Qiu, 1996). Ellcyon intrepidus from the Lukeino Fm., Kenya, recently described by Morales et al. (2005), is the oldest known representative of the genus in Africa. It is based on isolated teeth, but these are diagnostic for the genus. A second possible record of this species from Lemudong'o in Kenya has recently been described by Howell and Garcia (2007). Recently, Garcia (2008) has described the new species E. wokari from the early Pliocene of Ethiopia (Aramis and Kuseralee). More complete material of Eucyon sp. is known from Langebaanweg (figure 32.2A), in the form of a skull, mandibles, and postcranial elements of more than one individual (Spassov and Rook, 2006). Ellcyon is also known from the much younger site of Ahl al Oughlam, where material described by Geraads (1997) as Canis aff. illllms at least in part is attributable to Eucyon sp. (but see Geraads, 2008). Genus NYCTEREUTES Temminck, 1838 Figllres 32.2B and 32.2C Diagnosis Modified after Ward and Wurster-Hill (1990). Skull small, greatest length <130 mm with short, narrow muzzle and low forehead; parietals with rugose surface and slight interparietal crest; mandible with a distinct rounded sub angular lobe on the posterior margin; dental formula I 3/3, C 1/1, P 4/4, M 2/3, total 42, but an extra upper molar is common. Carnassial blades reduced and molars relatively large. African Species N. abdesalami Geraads, 1997, N. terblancijei (Broom in Broom and Schepers, 1946; figure 32.2C), N.? sp. nov. Werdelin and Dehghani, in press (figure 32.2B), N. sp. Age Ca Ma. Geographic Occurrence Morocco, South Africa, Tanzania. Remarks Nycterelltes is mainly a Eurasian taxon and its lineage was one of the first can ids to evolve after the family dispersed into Eurasia near the end of the Miocene (Tedford and Qiu, 1991). Barry (1987) described a medium-sized canid from the Laetolil Beds, Upper Unit, as aff. Canis brevirostris. Tedford (pers. comm., 1999) has suggested that this taxon should be placed in Nyctereutes, and on the basis of dental characteristics we concur (Werdelin and Dehghani, in press). It should be noted, however, that the Laetoli form lacks the subangular lobe of the mandible that is characteristic of Nycterelltes. There is also a possible specimen of Nyctereutes from the latest Miocene of Lissasfa in Morocco (Geraads, 2008). Specimens definitely belonging to Nycterelltes have been recorded from Ahl al Oughlam in Morocco (Geraads, 1997) and Kromdraai and Elandsfontein in South Africa (Hendey, 1974; Ficcarelli et ai., 1984). Genus OTOCYON Muller, 1836 Diagnosis Small-sized Canidae; dentition with supernumerary molars, dental formula I 3/3, C 1/1, P 4/4, M 3-4/4-5; premolars small, with mesial and distal diastemata; upper molars similar in size, multicusped; P4 and m1 hypocarnivorous; m1 with low trigonid, long talonid; P4 triangular, with very short metastyle. African Species O. megalotis (Desmarest, 1822). THIRTY-TWO: CARNIVORA 609

8 TABLE 32.3 Occurrences of Canidae species..;j b 0;:: Q) "0 ;:;.;... ::l e i:: t:.s os: a a ;:; i::: '-' Q) "1::; i::: cd..l:l t: "0 'e i:: e i:: e i:: '2. o o cd Sites U CJ CJ CJ CJ CJ CJ. 0<;:;,.!;:: '-' \"i i:: : ::l "1::; F t:..l:l e.;....a o tl '-' P., os ;:; ;:: '5.. Vl i:: i:: i:: i::: i::: o a a 'e i:: G- 8- '-' ;:; ;( CJ CJ I'q &l :z; 0 " P., Vl P., Vl ;:; ;:; i::: ;u ii '-' ;( :z; ;;;, a e 0- '-' \"i P.,.s i::: a Vl t:,.!;:: :d.;... & t: i:: i::.s.,.!;:: ;:; :s ;:; i:: '-' '-' '-' '- '- i2 8-.s.s tl tl tl tl <u.s.s.s..s..s..s e e.s..s..s. 0 Q., Q., S S S S S S S tl Ahl al Oughlam Ain Bahya Ain Boucherit Ain Hanech d. Asbole d. Awash 7 (Kalb) d. Bouknadel Busidima-Telalak Cap Achakar Cooper's Dar es Soltane Diepkloof Rock Shelter Djebel Irhoud Doukkala I Doukkala II Drimolen Duinefontein 2 Equus Cave Elands Bay Cave Elandsfontein Main El Harhoura 1 Florisbad Gladysvale Gratte des felins Hadar Denen Dora Hadar Kada Hadar Hadar Sidi Hakoma Hopefield Isenya Kabwe Kifan Bel Gohmari Konso-Gardula 2 Koobi Fora KBS Koobi Fora Upper Burgi Kossom Bougoudi affo Kramdraai cf. Member A Laetolil Beds Upper Unit Lainyamok Langebaanweg Baard's Quarry Langebaanweg PPM Lemudong'o Lissasfa do d. aff.? aff. 610 LAURASIATHERIA

9 'g t::s ts " " ;; i:: 0.;:J ::: t::s '-0 E :.;;;.;.J... '-,.Si " "" :e a '12 <ll '-0 '1::,.l::l P., P., " P.,.s " " t::s a i;:; 0:; "0 ;:: '-0.r; " 12 fr t::s V> V> V>... i:: ill '12 12 :d '-0.S " " ;:: a a ;:: b i:: i:: " % g ;0,. '-0... 'g P., '-> 12 a a t::s... <l) '-> " " P.,.is... A. t::s Ci' fl :s: ;:: ;:: P., V> ;:: ;:: ;:: ;:: <EE.. 12 's.. i:: '- '- V> OJ t::s t::s,.l::l V> "0 i:: i:: i:: a.8 ".8 '-0 "l "l "l "l.;:j.;:j.;:::.;:j.;:j.;:j "l.;:j a a "" ill '2 >:: Ci' i:: i:: '12 A. i:: i:: i:: i:: Ci'.8.8 " " " ",.s., " " OJ t::s t::s t::s t::s t:i t:i t:i t::s " " " " " Sites U V V V V V V V V &l &l &l 0 P..; P..; $ $ $ $ $ $ $ Lothagam Kaiyumung Makapansgat d. Member 3 Makapansgat Member 4 Melkbos Melkbosstrand Mursi Fm. Nachukui Kalochoro Nachukui Natoo Olduvai Bed I d. Olduvai Bed II Olduvai Bed IV? Oulad Hamida aff. aff. I-rhino cave Oulad Hamida Th III-H. erectlls aff. Plovers Lake Sagantole Fm. Saldanha Lime Quarry Saldanha Sea Harvest Sibudu HP Slangkop South Turkwel Sterkfontein d. Member 4 Sterkfontein d. Member 5 Sterkfontein Jacovec Cave Swartklip 1 Swartkrans d. Member 2 Swartkrans d. Member 3 Taung Dart deposits Thomas I-H. aff. d. erectlls cave Tighenif d. Toros Menalla Tugen Lukeino Ysterfontein THIRTY-TWO: CARN1VORA 611

10 100 mm c FIGURE 32.2 African Canidae. A) Eucyoll sp. PQ-L from the Muishondfontein Pelletal Phosphorite Mb., Varswater Fm., Langebaanweg, cranium in left lateral view and right hemimandible in lingual view. B)?Nyctere!ltes sp. nov. (part) LAET (specimen incorrectly labeled) from the Upper Laetolil Beds, Laetoli, right hemimandible in lingual view. C) Nyctereutes terblallcijei type specimen KA 1290 from Kromdraai Mb. A skull in right lateral view. Age Ca. 0.3 Ma-Recent. Geographic Occurrence Kenya. Remarks The extant species O. mega/otis is in the fossil state only known from Lainyamok in Kenya at ca 300 ka (Potts and Deino, 1995). Earlier records of bat-eared foxes are referred to ProtOtOCYOl1 (see next section). Genus PROTOTOCYON Pohle, 1928 Diagnosis Translated and modified from Pohle (1928). Small, foxlike genus of dogs, close to OtOC)!Ol1; frontal sinuses and sagittal crest absent; the weak lambdoid crests form a central invagination; dental formula I 3/3, C 1/1, P 4/4, M 2/3; premolars small; carnassial with a weak parastyle, width more than 60% of length; molars relatively large and very similar to each other, resembling those of OtOCYOl1; lower jaw with a processus subangularis. African Species P. recki Pohle, 1928, P. sp. Age Ca Ma (possibly as early as 3.7 Ma). Geographic Occurrence Tanzania. Remarks This genus, known through material of several individuals from Olduvai, Bed I (Petter, 1973), is only doubtfully distinct from the modern genus, OtOCYOl1. The main differences lie in the somewhat more primitive dentition of P. recki. It is a typical bat-eared fox and probably had the same ecological habits as its modern counterpart. Remains of a canid in the Otocyoll lineage have been recorded from Laetoli (Werdelin and Dehghani, in press), but whether they can truly be referred to the genus Prototocyon remains to be seen. Genus VULPES Frisch, 1775 Diagnosis Modified after Munthe (1998). Skull with long, sharp muzzle; postorbital process thin and concave dorsally; frontal bones less convex than in Canis; simple incisors; canine teeth slender, relatively longer than in Callis; cheek teeth slender with sharp, well-defined cusps and crests. African Species V. chama (Smith, 1833), V. pattisol1i Broom, 1948, V. pllicher Broom, 1939, V. riffalltae Bonis et al., 2007, V. rueppellii (Schinz, 1825), V. vlilpes (Linnaeus, 1758). 612 LAURASIATHERIA

11 Age Ca. 7 Ma-Recent. Geographic Occurrence Algeria, Chad, Kenya, Morocco, South Africa, Tanzania. Remarks Most finds of foxes in Africa are from the southern part of the continent. However, some of the oldest finds thus far (e.g., an unidentified?vulpini from the Mursi Fm., Ethiopia) have come from eastern Africa, suggesting that much of the group's history on the continent remains to be discovered. Recent discoveries have demonstrated that a small-sized fox is present in Toros Menalla, late Miocene of Chad, which implies a much earlier occurrence for the genus in Africa, and for the whole tribe Vulpini in the Old World (Bonis et al., 2007b). CANIDAE indet. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, South Africa. Remarks As noted earlier, a probable vulpine has been recovered from the Mursi Fm. Since it may not belong to any of the listed genera, we prefer to list it here. Indeterminate remains of canids are also known from Kossom Bougoudi, Chad (Bonis et al., 2008). Family MUSTELIDAE Fischer, 1817 Table 32.4 Genus AONY Lesson, 1827 Diagnosis After Willemsen (1992). Dentition more robust than in Lutra; teeth broad; P4 with large talon, protocone elongated, extending distally nearly to the metacone, no hypocone; m1 with talonid broader than trigonid, hypoconid present, entoconid absent or smaller than hypoconid, buccal cingulum of talonid strong. African Species A. capensis (Schinz, 1821). Age Ca. 4.3 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, South Africa, Tanzania. Remarks Compared to other Lutrinae, members of the Aonyxini are rare in the African fossil record. The oldest record is from the Laetolil Beds, Lower Unit. Genus DJOURABUS Peigne et al., 2008 Diagnosis After Peigne et al. (2008a). Large-sized species; tall and thick mandibular corpus; m 1 bunodont, short, and very broad with a trigonid broader than the talonid, transversely oriented paraconid shelf, bases of the metaconid and paraconid connate, short; m1 talonid molarised and short, with squared-off distal margin and individualized and low entoconid and hypoconid, and cingulid very reduced and restricted to the mesial margin of the paraconid. African Species D. dabba Peigne et al., Age Ca. 7 Ma. Geographic Occurrence Chad. Remarks This species is based on a fragmentary mandible on which the canine and the m1 are the only preserved teeth. Despite the fragmentary nature of the material, the morphology of the mandible and carnassial of this bunodont lutrine is easily distinguished from that of the other bunodont genera (Ellhydriodol1, Enhydl'ithel'ill1ll, Pailidoilitra, and Sivaonyx). As it is one of the earliest-known bunodont lutrines, the origin of Djollrablls remains unclear. Genus EKORUS Werdelin, 2003 Diagnosis After Werdelin (2003). Mustelidae of gigantic size; dentition highly modified for slicing, with narrow canines and slender premolars; m1 lacking metaconid, talonid reduced to a single, tall cusp placed directly posterior to the trigonid blade; m2 small and peg shaped; M1 very reduced and much broader than long; appendicular skeleton modified, with relatively long limbs but short, broad, semiplantigrade feet; humerus lacking entepicondylar foramen; vertebral column slender; tail long. African Species E. ekakeran Werdelin, Age Ca Ma. Geographic Occurrence Kenya. Remarks This form was described on the basis of a nearly complete skeleton from the Lower Nawata Mb. at Lothagam (Werdelin, 2003). It is by far the most hypercarnivorous and cursorial mustelid known. Its origin is unknown but may lie with the Eurasian Miocene IschYl'ictis group, which also includes large, hypercarnivorous forms. If so, the Ekol'lls lineage has a long, independent history in Africa. Genus EOMELLIVORA Zdansky, 1924 Diagnosis Modified after Wolsan and Semenov (1996). Musteline mustelids of very large size; P3 with one or two posterior accessory cusps; P4 with a subconical protocone, and with paracone-protocone and paracone-parastyle crests; M1 with a vestigial metacone, an arched, ridge-shaped protocone continuing into the anterior protocone crest, and a talon about equally expanded anteriorly and posteriorly; p4 with a posterior accessory cusp; m1 metaconid absent and replaced by a distinct crest, talonid with lingual talonid crest enclosing a vestigial basin, with single but strong, nearly centrally positioned hypoconid; m2 elongated anteroposteriorly, with a low crown surrounded by a Cingulum and a very small, almost centrally placed protoconid, linked with the cingulum by crests anteriorly, posteriorly, and lingually. African Species E. tllgenensis Morales and Pickford, Age Ca Ma. Geographic Occurrence Kenya. Remarks Morales and Pickford (2005a) report material from the Ngorora Fm. that they refer to a new species of Eomellivora. This material is morphologically similar to the Eurasian E. wil1lal1i but much smaller, being approximately the size of the extant Mellivora capensis. Genus EROKOMELLIVORA Werdelin, 2003 Diagnosis After Werdelin (2003). Small-sized Mellivorinae with slender mandibular horizontal ramus; premolars long and slender; m1 low and long with small metaconid; m2 present and single rooted. African Species E. lothagamemis Werdelin, Age Ca Ma. Geographic Occurrence Kenya. Remarks This taxon resembles Mellivora dentally but lacks several of the derived features of the extant genus, such as the loss of m2. It is possible that Mellivora is a direct descendant of Erokomellivora. Genus HOWELLICTIS Bonis et al., 2009 Diagnosis After Bonis et al. (2009). Small-sized Mellivorinae without PI/pI differing from Mellivora in its smaller size, less THIRTY-TWO: CARNIVORA 613

12 imbricated and narrower premolars, lack of a sagittal crest, and presence of a reduced m2. African Species H. valentini Bonis et ai., Age Ca. 7 Ma. Geographic Occurrence Chad. Remarks The species is represented by a fairly good sample from Toros-Menalla, Chad. It is morphologically different from Mellivora and could belong the ancestral stock of this genus, along with Erokol1lellivora. Genus ICTONY Kaup, 1835 Diagnosis Small-sized Mustelidaei dental formula I 3/3, C 1/1, P 3/3, M 1/2i premolars small and slenderi m1 with a low trigonid and broad and flat talonid about as long as the trigonidi P4 slenderi M1 short with reduced metastyle wing. African Species 1. libyca (Hemprich and Ehrenberg, 1833), 1. striatlls (Perry, 1810). Age Ca. 2.5 Ma-Recent. Geographic Occurrence Angola?, Morocco, South Africa. Remarks Ictonyx is only known as a fossil from two records of extant species, since 1. boiti has been transferred to the genus Prepoecilogale (Petter and Howell, 1985). Genus KENYALUTRA Schmidt-Kittler, 1987 Diagnosis After Schmidt-Kittler (1987). Lutrine of the size of, or somewhat smaller than, the Recent clawless otteri trigonid of the lower carnassial as low as the talonid, metaconid and protoconid equal in size, talonid markedly broader that the trigonid, and hypoconid situated far backward. African Species K. songhorensis Schmidt-Kittler, Age Ca Ma. Geographic Occurrence Kenya. Remarks Kenyalutra was erected by Schmidt-Kittler (1987) on the basis of two lower carnassials, one of them fragmentary, from Songhor, Kenya. He assigned them to the Lutrinae mainly on the basis of two features: trigonid and talonid of nearly equal height and talonid very broad, considerably broader than the trigonid. However, these are characters of questionable Significance given the age difference between Kenyailltra and other African Lutrinae, the very small size of Kenyalutra compared to comparable Lutrinae, and the highly derived nature of some features, such as the breadth of the talonid and the posteriorly placed hypoconid. Morales et ai. (2000) suggested that the Kenyailltra specimens might be the unknown lower dentition of Kelba qlladeemae, a mammal of uncertain affini s first described by Savage (1965) on the basis of isolated uppe'': molars. They further suggested that this combination might be referred to the hemigaline viverrids. However, recently described material clearly shows that Kelba is not a carnivoran (Cote et ai., 2007), whereas the Kenyailltra specimens almost certainly are. Kenyalutra might still be a viverrid, but more material is needed to make any sort of assessment of its phylogenetic relationships. Thus, for the time being we retain it in the Mustelidae. Genus LUOGALE Schmidt-Kittler, 1987 Diagnosis After Schmidt-Kittler (1987). Mandibular ramus very strong: m1 shorter and taller than in the Laphictis Ischyrictis group, with small but clearly developed metaconid and short talonid with a single blunt CUSPi P4 with strong carnassial blade and well-separated rounded protocone, small parastyle present. African Species L. rusingensis Schmidt-Kittler, Age Ca Ma. Geographic Occurrence Kenya. Remarks Schmidt-Kittler (1987) described this new genus and species on the basis of some tooth and mandible fragments from Rusinga, Kenya. It shows similarities with less derived forms of the Isci1yrictis group in the structure of mi. Schmidt-Kittler (1987) suggests that it may have originated from the European Paragale-Plesiogale group. Genus LUTRA Brisson, 1762 Diagnosis After Willemsen (1992). Lutrine with wide, flat headi fingers and toes extensively webbed and with welldeveloped clawsi tail tapering evenly, somewhat flattened. Skull long and depressed, facial part rather long, intertemporal region not swolleni dental formula I 3/3, C 1/1, P 4/3, M 1/2i teeth with sharp cutting edges, never blunt and not very robusti P4 with a sharp cutting blade on the buccal side, formed by paracone and metacone, talon not covering entire lingual side of trigonim1 with sharp cuspids, hypoconid forming a sharp edge, external cingulum of talonid not strongly developed. African Species L. fatimazoi1rae Geraads, 1997, L. libyca Stromer, Age Ca. 7.1 Ma-Recent. Geographic Occurrence Egypt, Morocco. Remarks The species listed here are all quite similar to each other and to Torolutra (discussed later). The number of valid species and their relationship to each other and to other species of Lutra (e.g., L. simplicidens Thenius, 1948) remains obscure at present. Genus MARTES Pinel, 1792 Diagnosis After Anderson (1970). Skull ranging in basilar length from 60 to 115 mmi facial angle slighti auditory bullae elongated, thin walled, moderately inflated, not in close contact with paroccipital processesi auditory meatus short but distincti palate extends beyond last upper molari dental formula I 3/3, C 1/1, P 4/4, M 1/2i P4 trenchant with small, well-developed protocone; ml with small metaconid, trigonid longer than talonid, talonid semibasined. African Species M. khelifensis Ginsburg, Age Ca Ma. Geographic Occurrence Morocco. Remarks Pre-Pleistocene Martes is in need of revision. This genus, like Mllstela, has become a wastebasket nomen for various small mustelids of uncertain relationship to each other. Martes khelifensis is one of these. It is known from a single Ml germ that Ginsburg (1977) compares with the homologous element of the Asian M. anderssoni, another small Martes-like mustelid of uncertain affinity. Nevertheless, M. khelifensis is the only such taxon in the African fossil record. Genus MELLALICTIS Ginsburg, 1977 Diagnosis Translated from Ginsburg (1977). Mellivorine with elongated facei p4 tall, lacking anterior accessory CUSPi ml short with well-developed metaconid and short talonid where the hypoconid is the largest CUSPi M1 with strongly developed para style, metacone reduced to a narrow crest posterior to the paracone, paraconule isolated, protocone elongated surrounded by an extensive cingulum. 614 LAURASIATHERIA

13 African Species M. mel/aiensis Ginsburg, Age Ca Ma. Geographic Occurrence Morocco. Remarks Mel/a/ictis is known from a number of isolated teeth and postcranial fragments from Beni Mellal, Morocco. It has similarities with both ischyrictines and mellivorines, but Ginsburg (1977) suggests that its relationships probably lie with the former, particularly with the genera Ischyrictis and Hadrictis. If this is correct, it may also be related to the much later and larger Ekorus from Kenya (see earlier discussion). Genus MELLIVORA Storr, 1780 Diagnosis Large-sized Mustelidae; molar dentition reduced, dental formula I 3/3, C 1/1, P 3/3, M 1/1; premolars slender, m1 with short, trenchant talonid, metaconid low; M1 anteroposteriorly shortened, paracone and metacone small. African Species M. benfieidi Hendey, 1978, M. capensis (Schreber, 1776), M. caroiae Michel, Age Ca. 5.8 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks The oldest published record of Mellivora is from Langebaanweg, based on material referred to M. benfieidi (Hendey, 1974). However, Petter (1987) suggests that this species may simply be the ancestral starting point of the M. capel1sis lineage, in which case the two would be conspecific. Only better material can elucidate the intraspecific variation of these species. M. caroiae from the Pleistocene of Morocco (Michel, 1988) does not appear to differ from the living species, though this is difficult to evaluate due to the absence of illustrations of the relevant specimens. Genus MUSTELA Linnaeus, 1758 Diagnosis Modified from Bryant et al. (1993). Smallsized Mustelidae; skull with anterior opening of palatine canal in maxilla, inflated bulla, and no paroccipital process; dental formula I 3/3, C 1/1, P 3/3, M 2/2; no accessory cuspid on p4; M1 transversely elongated and with buccal cusps reduced; no metaconid on m1, m1 talonid not basined, with a strong central hypoconid, an entoconid, and a buccal cingulum. African Species M. plltorills Linnaeus, Age Ca Ma. Geographic Occurrence Morocco. Remarks The only fossil record of the genus in Africa is Mllsteia plltorills from the Upper Pleistocene of EI Harhoura 1 cave, Morocco (Aouraghe, 2000). The species is currently absent from the continent. Genus NAlvIIBICTIS Morales et ai., 1998 Diagnosis After Morales et al. (1998). Mustelinae with hypercarnivorous dentition; lower canine with very high crown; lower premolars and m1 mediolaterally compressed; p4 high crowned; m1 with a vertical paraconid, a residual metaconid, a small talonid comprised of a bevelled hypoconid. African Species N. sel111ti Morales et ai., Age Ca Ma. Geographic Occurrence Namibia. Remarks Namibictis is a medium-sized mustelid with hypercarnivorous dentition known from Arrisdrift, Namibia. It is similar in carnassial morphology to ischyrictines, especially Mel/a/ictis, and is probably related to the latter taxon, as suggested by Morales et al. (1998). Genus PLESIOGULO Zdansky, 1924 Diagnosis After Kurten (1970). Wolverine with relatively large anterior premolars, carnassials relatively shorter than in Gllio; upper molar larger and more expanded anteroposteriorly than in Guio, with inner lobe produced posterad; lower carnassial with a long, weakly basined talonid, and with or without a metaconid; m2 less reduced than in Guio. African Species P. botori Haile-Selassie et ai., 2004, P. monspessllim1lls Viret, 1939, P. praecocidens Kurten, Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, South Africa. Remarks P. m011spessllianlls, from South Africa (Hendey, 1978) and P. botori from Kenya and Ethiopia (Haile-Selassie et ai., 2004a) are both large species of the genus, while P. praecocidens from Kenya is a smaller species. Whether P. monspessllimllls and P. praecocidens are correctly assigned to species is at present a moot point, but it is clear that Piesiogllio had a short-lived presence in Africa at the end of the Miocene-beginning of the Pliocene. Genus PREPOECILOGALE Petter and Howell, 1985 Diagnosis After Cooke (1985). Musteline slightly smaller than IctOI1Y striatus, resembling Poeciiogaie aibil111c/1a in size, skull proportions, and in the extent of the lambdoid crest; P2 present as in IctOI1Y; P4 intermediate between IctOI1Y and Poecilogaie in protocone development; tympanic bullae less inflated than in IctOI1Y striatus. African Species P. balti (Cooke, 1985). Age Ca Ma. Geographic Occurrence Morocco, South Africa, Tanzania. Remarks P. bolti is known from Bolt's Farm in South Africa, where it was originally described as Ictol1Yx bolti (Cooke, 1985), from Ahl al Oughlam, and from Laetoli, indicating a wide distribution over a relatively short time span. Genus SIVAONY Pilgrim, 1931 Figure 32.3 Diagnosis After Morales et al. (2005). Lutrinae with P4 with continuous cutting blade, with no incision separating the paracone and metastyle, protocone moderate, hypocone moderate, well separated from the protocone and the paraconemeta style; M1 somewhat wider than long, with peripheral cusplets and a wide, flat central valley; m1 with relatively long trigonid, and unmolarised talonid with a wide basin. African Species S. africmjlls (Stromer, 1931), S. beyi Peigne et ai., 2008, S. hel1de)!i Morales et ai., 2005, S. ekecamal1 (Werdelin, 2003)(figure 32.3D), S. kmnuij(ll1girei Morales and Pickford, 2005, S. soriae Morales and Pickford, 2005, S. sp. Age Ca Ma. Geographic Occurrence Chad, Egypt, Ethiopia, Kenya, South Africa, Uganda. Remarks The majority of the taxa listed above have in the past been referred to Enhydriodon, but Morales and Pickford (2005b) and Morales et al. (2005) have transferred them to Sivaonyx. Neither taxon is particularly well characterized. The lineage includes an impressive array of African species, the geologically younger of which were among the most massive carnivorans of all time (figures 32.3A-C, E). THIRTY-TWO: CARNIVORA 615

14 FIGURE 32.3 SivaollYx Spp. A-C) SivaollYx n. sp. A, KNM-ER 3110 left ramus from the Tulu Bor Mb., Koobi Fora Fm. in buccal (A), lingual (B), and occlusal (C) views. D) S. ekecamall type specimen KNM-KP (part), right ml from Kanapoi in occlusal view. E) SivaollYx n. sp. B, L56-11eft from Mb. C, Shungura Fm in occlusal view. Genus TOROLUTRA Petter et ai., 1991 Diagnosis Translated from Petter et al. (1991). Mandible with relatively deep masseteric fossa, separated from the ventral margin of the ramus and with anterior border behind alveolus for m2; p4 robust, oval in occlusal view, surrounded by a thick, continuous cingulum; m1 with tall and sharp trigonid cusps, the protoconid being the tallest trigonid cusp, talonid narrow, with flat and obliquely oriented lingual face of the hypoconid. African Species T. ollgandensis Petter et ai., 1991, T. sp. Age Ca Ma. Geographic Occurrence Egypt, Ethiopia, Kenya, Uganda. Remarks The taxonomy of small and medium-sized lutrines is in a state of confusion at present (see Llltm), and where TOro/lltm might fit into this picture is currently not determinable. Genus VISHNUONY Pilgrim, 1932 Diagnosis Modified after Pilgrim (1932). Lutrinae of rather small size; P4 triangular, buccal anteroposterior diameter greater than lingual, and also much exceeding transverse diameter, with high pointed paracone, metacone lower but elongated, parastyle weak, protocone and hypocone both much lower than paracone, protocone situated rather far forward; internal cingulum slight; M1 rather small; mandible with deep ramus; p4 elongate, only slightly broader posteriorly, with strong posterior accessory cusp, not situated so much to the outside of the main cusp as in Sivaonyx, without anterior accessory cusp, with broad cingulum; m1 with talonid broader and shorter than trigonid, surrounded by a crenulated rim, entoconid as strong as hypoconid; m2 elongate oval, rather longer than in Sivaonl'x. African Species V. cijinjiensis Pilgrim, 1932, V. angolo/ensis Werdelin, Age Ca S Ma. Geographic Occurrence Kenya. Remarks The V. cijinjiensis specimen from Ngorora matches that described from India by Pilgrim (1932) in size and morphology, while V. angolo/ensis is rather larger. Morales and Pickford (200Sa) suggest that V. ang%lensis might belong to Toroilltm and this merits further consideration, as none of these taxa is particularly well characterized. LUTRINAE indet. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Tanzania. Remarks Lutrinae that do not fit into the aforementioned genera or that are indeterminate are found at. a number of localities, but none of these are outside the stratigraphie or geographic range of the more complete material. In addition to Djollmblls dabba and Sivaon)!x beyi, the late Miocene locality of Toros-Menalla, Chad, has also yielded at least two additional species known from fragmentary material. One species displays piscivorous dental adaptations and is related to Toro/lltm ollgaljdensis and the other species is represented by a complete femur very similar to that of Aonyx capensis (Peigne et al., 2008a). A partial juvenile skeleton with maxilla fragment from the lower unit of the Laetolil beds is allied to Aonyx. MUSTELIDAE indet. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Tanzania, South Africa, Uganda. Remarks Mustelidae of uncertain affinity are known from a number of localities (table 32.4). Most of these are known from fragmentary material, and several represent small species that may belong to known genera but have not been sufficiently studied. In a faunal list, Pickford et al. (1990) record? Poeci/ictis from Cangalungue Cave in Angola. The generic attribution of this material needs verification, though clearly a small mustelid is indicated. Family NANDINIIDAE Pocock, 1929 Table 32.5 Genus NANDINIA Gray, LAURASIATHERIA

15 ::r: :r: p. t::1 p. "' 0 >-1 >-; ::Y. t::1 ;J (1) ;J ::l (]) n ro ::l >n ttl ttl ttl ttl """""' Vl;..O""""",....?,g;:;::r:;;:: o-e;e:. 0: n::r;j""" p. (3 0 0 o 8. :::. ';j ttl t::1 n - 0: n 6' 0.. ro <'l...,... ro a '-< (]. ;J (1) N too too too 00(1) :::: :=t 0... ro >- :: ",::r. ' >- > 0- r:: e; 2- ::r 5',... (]) o n >- >- >- >- >- VJ Joooi!-i"""""' ::r 0-. S ;? ;; '? p.. too ' - >- e:... '7j s p Aon)'x capellsis Aonyxini indet. Djollrablls dabba Ekorlls ekakeran Eomellivora tllgenensis Erokomellivora lotiwgamellsis Howellictis valentini I Icton/,x lib)lca Icton/,x striatlls Icton)'x sp. Ken)'ailitra song/jorel1sis Lllogale rusingensis I Llltra fatimazo/jrae Llltra lib)'ca I Lutrinae indet. Mattes klzeiifensis Mellalictis mellalensis?:: I Mellivora bel1fieldi Po I Mellil'ora capensis Mellivora carolae Mllstela plltorills Mustelidae indet. o n 0: ::; (1) ;J n o ' >- ;;::r "" 0: m w g v o:+ (1) {l (1) o. I Namibictis Sf1Juti I Plesiogulo botori Plesiogulo monspesslllal111s Plesiogulo praecocidens n Prepoecilogale bolti Po I Sivaon)'x africalw SivaolJ)'x be)li Sivaon/,x ekecaman Sivaon)'x ljeljde)'i Sivaon)'x kamuhangirei Sivaoll)'x soriae Sivaon)'x sp. n Toroilitra ollgandensis Toroilitra sp. Vishlluoll)'x angololensis VishmlOlJ)'x c/jinjiensis

16 r',ob C/)O«::;::8- O'q :::: t"" "'0 "'OQ'q ::;::8-0«* r' r' r',ot;j:j cro r'ro.o[ a8" '-< VJ-' Pol...-j to I-i to n C (1) C ro ::: (ti _. Vl _. Vl Q'q r+ r+ q, A A A 000 8,g:&c'& -l '0. C. o (tl >-r:i "7j I-i f"i 0 ti:! 0 g'pss: 03. n n A A CS r' g g 0 g ::;::. g9ga; g 2: g: g P' r+ N q, n A A ::c 0 Q;l t:l.'o::r:," l:j E:; O(DOC/J. 0: S p:. V') AOIl)'x capensis Aonyxini indet. Djoumblls dabba Ekorus ekakemll Eomellivom tllgel1ensis Erokomellivom lotijagamensis Howellictis valentini Ictol1)! lib)'ca IctOIl)' striatus Ictol1),x sp. Kel1),alutm sol1ghorel1sis Lllogale ntsingensis Llltm fatimazohme Lutm lib)'ca Lutrinae indet. Martes khelifel1sis Mellalictis mellalensis Mellivom benfieldi Mellivom capellsis Mellivom carolae Mllstela putorills Mustelidae indet. Namibictis sel111ti Plesiogulo botori Plesiogulo mol1spesslllanlis Piesiogllio pmecocidells Prepoecilogale bolti SivaolJ)'x africaija Sivaol1)' be)'i Sivaon)'x ekecamal1 Sivaon)'x hende)'i Sivaoll)'x kamuhallgirei Sivaoll)'x so1'iae Sivaoll)'x sp. Toroilltm ougandel1sis Toroilltm sp. Vis1117U0I1)' allgololel1sis Visimuon)'x chilljiensis n>-l o ;, Z OJ >-I r Zw C [oj ",. o-l'

17 Lemudong'o Lothagam Lower Nawata Lothagam Upper Nawata Manonga 1 Nachukui Lower cf. Lomekwi Nachukui Upper Lomekwi Nakali Nakoret Napak Ngorora Member D Nkondo Omo Shungura B Omo Shungura C Omo Shungura E+F Omo Shungura H Omo Us no d. Plovers Lake Rusinga Sagantole Fm. aff. Saldanha Sea Harvest Sibudu HP d. Songhor South Turkwel Swartklip 1 Toros Menalla aff. Tugen Lukeino aff. Tugen Mabaget Tygerfontein Wadi Natrun cf. Warwire

18 TABLE 32,5 Occurrences of Nandiniidae and Viverridae species ';:: <u,:3 'E ::l '1::l 1:::,;::,;:: 0') 0') '1::l e 0') E '<;; 0') 2 1::: '0 '1::l 1::: 1::: 0 <u 1::: 1::: ' "" <..J <u -S : 'B' 2!..; <u 1::: QJ '1;,; <u ".8 '2.. <u.s 2 -<..s:: ;s: " 1:::. ' ' "CI <..J <u..s:: E ;0. ".S,;:: a 1::: : <,S " 0') 0 ;s: <u 't; 0') 0..s:: 0') 1::: 0 :S 1:::, ;s: z Q :d E,;:: v, '1::l 0.. "" <u <..J QJ 0') 0 " :E :E,;,; v, ;0. E "" ',S :d 0 P2 " '1::l '1::l. m <..J 'S::,;,;,;,; :;::; ' '1::l :;::; ';:0 E..s::,. :g 5 'i2 1::: 1::: <..J <..J E :.8.8 ' ' <..J..g <..J B 2! g, 's :;::; 0,;,; ' 'i2 2! 2! H " " 1::: Z.;::: <..J <..J <u <u " '- " h 1::: > 1::: 1::: 1:::. 1:::. i:o 2! 1U..s:: 1:::..s:: <u QJ ;0. 0') -< <t:... ;0..;:,: :> Sites Z "<: "<: "<: U U C) C) C) ("0., 0 V) V) Adu-Asa Fm, >= 0 t:: s: s: s: :> Ahl al Oughlam Allia Bay Aramis Arrisdrift As Duma Awash 10 (Kalb) Awash 7 (Kalb) Beni Mellal Chamtwara ChOl'ora Duinefontein 2 Elands Bay Cave Elandsfontein Main Elisabethfeld Fort Ternan Hadar Denen Dora Hadar Pinnacle Hadar Sidi Hakoma Kanapoi Kipsaraman Klein Zee Konso-Gardula 1 Konso-Gardula 2 Koobi Fora Chari Koobi Fora KBS Koobi Fora Lokochot Koobi Fora Okote Koobi Fora Tulu Bor Koobi Fora Upper Burgi Koru Laetolil Beds Upper Unit Langebaanweg PPM Langebaanweg QSM Legetet Lemudong'o Lothagam Lower Nawata Lothagam Upper Nawata Manonga 2 Melka Kunture Garba IV cf, 620 LAURASIATHERIA

19 Sites Z :z; Nachukui Kalochoro Nachukui Lower Lomekwi Nachukui Upper Lomekwi Napak Ngorora Member A Ngorora Member B Olduvai Bed I Olduvai Bed II Omo Shungura B.,... Q; 0 '1::i 1:: <u 'E ' 1 tl.;;:; 1::: 03.l:::.s tl.l::: E.,... ;0. " 0 <u.s 'c.l::: p -c '';:: tl.:::1 -c C2.:::.,... '';:: :5 'E" 'E" 1:: :;:: '1::i tl tl tl.,....,....,....,... Q; Q; Q; Q; 1:: <u <u > ;0. ;0. 1:: 1:: 1:: :E.. :E...2: 'U Q; Q; U C) C) C) -< <u 8 -< Z 23 " it.:::.s ; ; Z.::: -< tl " " ;;: ij ij " " " Omo Shungura C d. Omo Shungura D d. Omo Shungura E+F Omo Shungura G Omo Shungura L Plovers Lake Rusinga Sagantole Fm. Sahabi Saldanha Sea Harvest d. Sibudu HP d. Songhor South Turkwel d. Sterkfontein Jacovec Cave Swartkrans Member 3 Toros Menalla Tugen Lukeino Tugen Mabaget Wadi Moglua.:::1 e 'E.:;:; :::? ',.... tl.::: <2.. E 1:: tl 1:: 1:: 1:: 0 <u <u 1:: tl " " 'U.l::: tl.,.; <u tl ;:: Q) '5.. Q; <2.. tl.'it. E 2 1:: :E.. ".. "0 tl. a 1:: 0 tl tl 0.:::1.s tl 03 <u E.S 0 'U '1::i.,....,... Q) Q; <u..2l Q; 0 " 13 " tl co '1::i ;0. 'c 1:: " :;::.:;:;.l:::.,.....2l :;:: 'c '';:: "0 0 <u. :;:: " 2 " " " 's.. 'E 2 2 c Q; Q; Q; f2:.l::: '1::i 1::... '-' ",., ",., <u <u ".I:l 6<' ii:l 1::.l::: Q; Q) <u 2.l::: 2.;0 ;0. ;0. ;> tl.::l :::s Q:; s: <'-. '< '< 0 V) V) t::: s: s: ;> S1 d. aff. Diagnosis Partly modified from Veron (1995) and Nowak (2005). Small-sized carnivoran with dental formula I 3/3, C 1/1, P 4/4, M 1-2/1-2; skull with triangular palate; primitive auditory bulla (no inflation during ontogeny, single rather than double chamber, no septum bullae, cartilaginous caudal entotympanic); paroccipital apophysis relatively thick; lacrymal and jugal bones not in contact; anterior rim of orbit formed by frontal, maxillary and jugal; posterior carotid foramen located between the basisphenoid and the caudal entotympanic; upper incisors forming a transverse line; lingual cusp present on 1'3; M2/m2 very reduced and rounded, sometime absent; m2 with only a small buccal cuspid. African Species N. sp. Age Ca. 6.1 Ma-Recent. Geographic Occurrence Kenya. Remarks Nmldinia is an arboreal forest taxon and as such less likely to be represented in the fossil record than terrestrial taxa. However, it is possible that postcranial remains, especially femora, of Nandinia are mistaken for monkey postcrania (B. Senut, pers. comm. to L.W., 2006) and that the family has a more extensive record than a single tooth from the Lukeino Fm. suggests. Family VIVERRIDAE Gray, 1821 Table 32.5 Genlls AFRICANICTIS Morales et al.,1998 Diagnosis After Morales et al. (1998). Feloid carnivoran of the size of the European Miocene hyaenid Protictitherilllll THIRTY-TWO: CARNIVORA 621

20 crassll711; pi reduced; premolars tall and narrow; m1 with tall trigonid and short talonid with a small hypoconid; P4 narrow and elongate, with strong parastyle and conical protocone; M2 small; M1 with well-developed parastylar area, shortened in the type species. African Species A. hyael10ides Morales et ai., 2003, A. meini Morales et ai., 1998, A. Sch711idtkittleri Morales et a!., Age Ca Ma. Geographic Occurrence Kenya, Namibia. Remarks This genus of small carnivorans, at present known only from Arrisdrift, Namibia, and Chamtwara, Kenya (Schmidt-Kittler, 1987; Morales et ai., 1998, 2003), is a morphological link between Stel10plesictis and the percrocutids. Its dental morphology is similar to primitive hyaenids such as Protictitherillll1 (Morales et ai., 1998), and Trmgllrictis (Colbert, 1939; Werdelin and Solounias, 1991). Of the three species, the older A. schmidtkittleri is the smallest and most primitive, while A. meini and A. hyae/1oides differ in the structure of the ml. The latter is similar in many respects to primitive Percrocuta spp. Genus CIVETTICTIS Pocock, 1915 Diagnosis Large-sized Viverridae; dental formula I 3/3, C 1/1, P 4/4, M 2/2; P4 protocone enlarged, metastyle short; m1 with short trigonid and broad, three-cusped talonid; premolars robust but slender; m1 and Ml-2 broad and with crushing adaptations. African Species C. civetta (Schreber, 1776), C. howelli Morales, Pickford and Soria, 2005, C. sp. Age Ca. 5.1 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, South Africa. Remarks Civettictis spp. are rare members of eastern African Pliocene faunas. Most fossil finds are fragmentary. Genus GENETTA Cuvier, 1817 Diagnosis From Gaubert et ai. (2004). Small-sized viverrid with dental formula I 3/3, C 1/1, P 4/4, M 2/2; plesiomorphic dental anatomy; M2 present; m1 talonid developed; m2 well developed. African Species G. genetta (Linnaeus, 1758), G. tigrina (Schreber, 1776), G. sp. Age Ca. 14 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks Fossils of Genetta spp. are not uncommon in the African Upper Miocene-Pleistocene, but referral to specific species is hampered by the fragmentary nature of most remains, especially the earliest records (Beni Mellal, Lothagam, Lemudong'o). The first definite record of the genus is from Kanapoi. The oldest material that is indistinguishable from modern G. genetta on diagnostic features is from the Upper Burgi Mb. at Koobi Fora. Genus HERPESTIDES Beaumont, 1967 Diagnosis Small-sized viverrid with dental formula I 3/3, C 1/1, P 4/4, M 2/2; skull with strongly developed sagittal crest; ossified auditory bulla with inflated ectotympanic, bilaminar septum, and so forth (see Hunt, 1991); P3 with third root more or less well individualized and supporting a small cuspid or a prominent lingual cingulum; P4 with protocone enlarged and parastyle developed and trenchant; M1 with crescent-shaped protocone much larger than buccal cusps; hypoconid prominent on m1; Single-rooted m2 lacking the paraconid. African Species H. aegypticus Morlo et ai., 2007, H. aeqllatorialis Schmidt-Kittler, 1987,?H. afarensis Geraads et ai., Age Ca. 18-?1O Ma. Geographic Occurrence Egypt,?Ethiopia, Kenya. Remarks The European H. antiqlllls is known through abundant material from the Lower Miocene of St-Gerand-Ie-Puy, France (Beaumont, 1967). Hunt (1991) has shown this taxon to be a true viverrid. In Africa, the genus is known only from dental remains, of which those of H. afarensis from Chorora (Geraads et ai., 2002) cannot be considered diagnostic at the generic level. This species may belong to Kmlllites (see next section). Genus KANUITES Dehghani and Werdelin, 2008 Diagnosis From Dehghani and Werdelin, Viverridae with long and narrow skull and moderately hypercarnivorous dentition; dental formula (upper dentition only): I?3, C, P 4, M 2; PI simple, peglike; P2-3 tall, slender; P4 robust, protocone slightly mesial to paracone, metastyle robust; M1 mesiodistally reduced, triangular, buccal cingulum strongly developed; M2 small, triangular. African Species K. lewisae Dehghani and Werdelin, Age Ca Ma. Geographic Occurrence Kenya. Remarks Known from the middle Miocene of Fort Ternan, Kenya, Kmlllites is the oldest known viverrid from Africa with intact basicranium. Its dentition and skull morphology are quite generalized, but dentally it shows resemblances to Helpestides and may be close to that taxon. Genus KETKETICTIS Morlo et ai., 2007 Diagnosis Modified after Morlo et ai. (2007). Differs from other African Miocene Viverridae s. I. except Helpestides and Kichechia in having a very broad and plesiomorphic m1; differs from Helpestides and Kichechia in having the trigonid cingulid surrounding the anterior tip of the tooth, the metaconid placed more posteriorly, and the low talonid equipped with extremely low cuspids and low crista obliqua; differs from Helpestides in being broader and having a less trenchant trigonid; differs from Kichechia in being much larger, having a smaller entoconid, and the cristid obliqua oriented more buccally. African Species K. solida Morlo et ai., Age Ca Ma. Geographic Occurrence Egypt. Remarks Ketketictis is known from a single, isolated m1 from Wadi Moghara initially identified as Helpestides sp. indet. A (Miller, 1999). Due to the fragmentary nature of this species, and its resemblance to Kicheci1ia, an assignment to Herpestidae cannot be ruled out. It should be noted that the familial allocation of poorly known early Miocene African taxa to either Viverridae s. I. or Herpestidae is strongly subjective and generally not based on apomorphies. Genus MIOPRIONODON Schmidt-Kittler, 1987 Diagnosis Revised after Schmidt-Kittler (1987). Comparable in general features of the dentition to Stel10plesictis but smaller and differing from that genus in: m1 trigonid lower and metaconid less reduced; pi two rooted; P4 less elongate; M1 relatively larger. 622 LAURASIATHERIA

21 African Species M. pickfordi Schmidt-Kittler, Age Ca Ma. Geographic Occurrence Kenya. Remarks Mioprionodon is a genus of small, relatively hypercarnivorous viverrids known from a number of mandible fragments with teeth from Songhor, Kenya. It resembles Stenoplesictis, Semigenetta, and Palaeoprionodon in the general morphology of its dentition, but it is considerably less derived in its hypercarnivorous features. Mioprionodon is distinguished from the aforementioned taxa on the basis of plesiomorphic characters and probably is a basal member of that clade. Genus MOGHRADICTIS Morlo et ai., 2007 Diagnosis After Morlo et ai. (2007). Large, hypercarnivorous "stenoplesictid," differing from all others in having an ml with small metaconid combined with long talonid, which is unbasined due to the presence of a tall hypoconid and no entoconid; differs from European "stenoplesictids" in having a P4 paracone that protrudes backwards; differs from "Stenoplesictis" l1l11ijoronii, in being much larger and having the ml postparacristid pointing more anteriorly. African Species M. nedjema Morlo et ai., Age Ca Ma. Geographic Occurrence Egypt. Remarks This recently described carnivoran from Wadi Moghara is represented only by a few dental remains. The species has been previously listed as Herpestides sp. indet. B (Miller, 1999). Morlo et ai. (2007) consider assignment of "Stenoplesictis" JOronii from Kenya to this genus possible. Genus ORANGICTIS Morales et al., 2001 Diagnosis After Morales et ai. (2001a). Primitive viverrine intermediate in size between Viverricliia indica and Viverra zibetlw; dentition robust; p4 with greatly reduced anterior cusplet; m1 short with high and closed trigonid, in which the metaconid is important and the paraconid is in a very lingual position, talonid small with very well-developed entoconid, attaining the height of the hypoconid; m2 relatively large, open trigonid with small paraconid and metaconid slightly taller than protoconid, talonid deeply excavated like that of m1, but with hypoconulid higher than entoconid and separated from it and the hypoconid. African Species O. gariepensis Morales et ai., Age Ca Ma. Geographic Occurrence Namibia. Remarks This genus is based on two mandibular specimens from Arrisdrift, Namibia (Morales et al., 2003). It shows some dental affinities with Helpestides and, like it, may be close to the basal stock of the Viverridae and/or Viverrinae. It also bears resemblances to Tllgenictis (discussed later), though it is much smaller than that taxon. Morales and Pickford (2005a) suggest that they may belong to the same lineage. Genus PSEUDOCIVETTA Petter, 1967 Diagnosis Modified and translated from Petter (1973). Civet of large size with bunodont dentition; M1 and M2 rectangular, with low, rounded cusps; p4 with large, rounded main cusp followed by a talonid with bunodont cuspids. African Species P. ingens Petter, Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, Tanzania. Remarks Pselldocivetta is by far the most aberrant viverrid, though it bears some resemblances to Tllgenictis and Orangictis (see these genera). Its cheek dentition is very bunodont, with extremely low relief and with a p4 that has a low, rounded main cusp and a posterior area that supports a number of low, bunodont cusps. Pselldocivetta was a short-lived but successful taxon that was present over a relatively large geographic area for about 1 Ma near the Plio-Pleistocene boundary. Genus SAHELICTIS Peigne et al., 2008 Diagnosis Modified from Peigne et al. (2008b). Viverrid of large size. Compared to Viverra leakeyi, V. howelli, V. pepratxi, V. peii, m1 with reduced metaconid; compared to these species and to Megaviverra cmpati1orllll1, ml talonid narrower, with a reduced hypoconid, no entoconid, no or greatly reduced hypoconulid, and a very low mesiolingual rim; compared to Viverra leake)'i, V. howelli, V. pepratxi, V. peii, V. (?) chinjie11sis, and Vishmtictis salmontanlls, m1 talonid shorter; compared to V. leakeyi and V. bakerii, M1 more reduced relative to P4 (or less shortened P4 relative to M1); compared to V. leakeyi, Ml transversely less elongated; compared to Visll1111ictis dllra11di, long rostrum, skull lacking expanded frontal regions and marked postorbital constriction; compared to Viverra howelli, V. pepratxi, V. (7) chi11jiensis, and Visllllllictis salmo11tanlls, much larger size. African Species S. korei Peigne et ai., Age Ca. 7 Ma. Geographic Occurrence Chad. Remarks Sahelictis korei is thus far known only from Toros Menalla, Chad. However, the large viverrid described from Langebaanweg and previously assigned to V. leakeyi, could be closely related to the Chadian species and belong to the same genus. With V. /eakeyi and P. ingens, S. korei is one of the rare fossil viverrids from Africa to reach a size larger than the extant C. civetta. Genus STENOPLESICTIS Filhol, 1880 Diagnosis Modified from Peigne and de Bonis (1999) and Peigne (2000). Small feloid with dental formula I 3/3, C 1/1, P 3-4/3-4, M 2/2; snout narrow; auditory region with inflated ectotympanic that widely contacts the basicranium; caudal entotympanic not ossified; septum bullae bilaminar for half of its height; lateral rims of basicranium ventrally depressed; P3 with small but distinct posterior accessory cusp; M2 reduced but double rooted in earlier species; p3-4 with distinct posterior accessory cuspid; posterior cingulum of lower premolars not trenchant; m1 metaconid reduced but relatively larger and less retracted than in Palaeoprionodon and Hap/ogale; m1 talonid basined, with a tall buccal crest on which the hypoconid may be prominent; m2 reduced and single rooted, with trigonid including a dominant protoconid. African Species "5." l1111horonii Schmidt-Kittler, Age Ca Ma. Geographic Occurrence Kenya. Remarks Like Africanictis and Mogi1radictis (discussed earlier), Stenoplesictis is included in the family Stenoplesictidae in recent contributions (e.g., Morales et ai., 1998; Morlo et al., 2007). However, this name used as a family-group taxon is paraphyletic, as pointed out as early as 1931 by Pilgrim, and is supported only by dental convergences. Therefore, it should not be used as valid (see also Hunt, 1989, 1998b). It THIRTY-TWO: CARNIVORA 623

22 should be noted, however, that integration of these three genera within the Viverridae s. 1., which is also paraphyletic, is not satisfying, either. A second alternative would be to include these genera in the Percrocutidae. The sale African species of Stenoplesictis is known from a maxillary fragment from Songhor and a mandibular fragment from Rusinga (Schmidt-Kittler, 1987). As previously pointed out (Peigne and de Bonis, 1999), the generic assignment of the species is debatable. It may be closer to Semigenetta (Hunt, 1996). Genus TUGENICTIS Morales and Pickford, 2005 Diagnosis After Morales and Pickford (2005a). Viverridae the size of Civettictis civetta; m1 robust, with high, bunodont cusps, paraconid located anteriorly, metaconid smaller that other trigonid cusps, talonid relatively short, with hypoconid and entoconid as tall as trigonid, hypoconid pyramidal, with an anterolingual crest running obliquely to the axis of the tooth as far as the contact between protoconid and metaconid, entoconid high and crescentiform; m2 with reduced trigonid, formed of a voluminous protoconid, small residual paraconid, and metaconid somewhat narrower than protoconid, and talonid dominated by a strong, rounded hypoconid well separated from the protoconid and a hypoconid joined to the entoconid, which is relatively small and well separated from the metaconid. African Species T. ngororaensis Morales and Pickford, Age Ca Ma. Geographic Occurrence Kenya. Remarks T. ngororaensis is a large, bunodont viverrid known from a single lower carnassial from Member A of the Ngorora Formation (Morales and Pickford, 2005a). This tooth is reminiscent of the lower carnassial of Orangictis (Morales et ai., 2001a) but is much larger. It also bears similarities to the lower carnassial of Pselldocivetta (Petter, 1967) but is overall far less derived. Nevertheless, Morales and Pickford (2005a) suggest that these three genera might be related as part of a lineage of bunodont viverrids culminating with the highly derived Pselldocivetta. Genus VISHNUICTIS Pilgrim, 1932 Diagnosis Modified after Pilgrim (1932). Medium- to large-sized viverrids with elongate, high, and narrow skull; contracting gradually to a considerable distance behind the postorbital processes, also contracting in front of the postorbital processes; slender muzzle; sides of face steeply descending from a nasal maxillary ridge; teeth not compressed laterally and with rather blunt cusps; braincase exceptionally narrow; upper molars rather large, relatively broad lingually; P3 without a lingual cusp; premolar series rather spaced, premolars simple; mandible rather stout but shallow; ml with relatively long trigonid, relatively short talonid; m2 rather large, oblong. African Species V. africana lvforales and Pickford, Age Ca Ma. Geographic Occurrence Kenya. Remarks The only record of this Asian genus in Africa is a fragmentary mandible with p4-m1 from the middle Miocene of Kipsaraman, Tugen Hills, Kenya (Morales and Pickford, 2008). According to these authors, this species is very similar to Viverra (7) cllinjiensis, compared to which it is slightly smaller and differs in the more gracile dentition and a larger distal accessory cuspid in p4. They provisionally assign V. (7) cizinjiensis and its African relative to the genus Visi1l1!lictis. Genus VIVERRA Linnaeus, 1758 Figures 32.4A and 32.4B Diagnosis Modified after Veron (1995). Medium-sized viverrids with dental formula I 3/3, C 1/1, P 4/4, M 2/2; skull with posterior rim of nasals across anterior rim of orbits, posterior opening of the carotid canal located in the middle of the medial rim of the bulla; anteromedial ventral process of the promontorium of the petrosal flattened and located level with the basisphenoid-basioccipital suture; paroccipital process very thin in the lower part, thicker in the upper part. African Species V. howelli Rook and Martinez-Navarro, 2004, V. leake)ji Petter, 1963 (figures 32.4A and 32.4B). Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, Libya, Morocco, South Africa, Tanzania. Remarks Small Viverra species are difficult to positively identify. Such records are hidden among the Viverridae indet. For reasons stated in Werdelin (2003), we are inclined to place V. leakeyi in the extant genus rather than in Megaviverra as in Morales et a1. (2005)(but see also Saizelictis, earlier). VIVERRIDAE indet. Figure 32.4C Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Morocco, Namibia, South Africa, Tanzania. Remarks The numerous mentions of Viverridae indet. are, except for those from the Ngorora Fm., Fort Ternan, and Lothagam, and Toros-Menalla, Pliocene to Pleistocene in age. Most can be assigned to extant genera, though others, including one from the late Pliocene of Koobi Fora (figure 32.4C), represent extinct taxa, in some cases much larger than any living viverrid. The specimen from Member B of the Ngorora Formation (Morales and Pickford, 2005a) represents a small, Genetta-like species. A tooth from Beni Mellal, identified by Ginsburg (1977) as Felis sp. is more likely to belong to a viverrid. Family HERPESTIDAE Bonaparte, 1845 Table 32.6 Genus ATILA F. Cuvier in Geoffroy and Cuvier, 1826 Figllres 32.SG-32.SJ Diagnosis Skull broad, supraoccipital crest large, sagittal crest well developed; mandible strong, ventral surface curved; coronoid and angular processes large; dental formula I 3/3, C 1/1, P 3-4/3-4, M 2/2; upper canine straight, lower curved; premolars robust; m1 with robust, long trigonid, short talonid; m2 with trigonid and talonid of similar length; P4 robust with large protocone; Ml with large metastyle wing; Ml and M2 bunodont. African Species A. l7lesotes (Ewer, 1956) (see figures 32.5G ), A. paludinoslls (Cuvier, 1829), A. sp. Age Ca. 1.7 Ma-Recent. Geographic Occurrence South Africa, Tanzania. Remarks A. mesotes, known only from Kromdraai Mb. A, was described as a species of Helpestes, but following suggestions by Ewer (1956) that this species is on the lineage to the 624 LAURASIATHERIA

23 B FIGURE 32.4 African Viverridae. A-B) Viverra ieakeyi, PQ-L 51590, unknown member, Varswater Fm., Langebaanweg, skull in ventral view and left hemimandible in lingual view; C) Viverridae n. gen. and sp., KNM-ER 5339, KBS Mb., Koobi Fora Fm., in left lateral view. extant marsh mongoose, we here transfer it to Atilax. In addition, the genus is known from Olduvai, Bed II. Genus CROSSARCHUS F. Cuvier in Geoffroy and Cuvier, 1825 Diagnosis Modified after Goldman (1987). Small-sized herpestids; dental formula I 3/3, C 1/1, P 3/3, M 2/2; skull with elongated snout, triangular palate, ectotympanic developed and greatly inflated, but much less than caudal entotympanic; PI/pI absent; P4 wide, with short metastylar shearing surface, well-developed para style, and prominent, slender protocone; M1 wide, paracone and metacone sub equal, buccally projecting parastyle and meta style; M2 similar to M1; p3 with posterior accessory cusp projecting buccally; m1 with sharp trigonid, paraconid and metaconid well defined, talonid widened; m2 well developed, trigonid low, paraconid not visible, talonid with basin and hypoconulid rim. African Species C. transvaalensis Broom, Age Ca. 1.6 Ma-Recent. Geographic Occurrence South Africa. Remarks Fossil Crossarciws is only known from Bolt's Farm and possibly from Kromdraai Mb. A. The relationships of C. tmnsvaalensis to extant cusimanses, none of which are known from southern Africa, are obscure. Genus CYNICTIS Ogilby, 1833 Diagnosis Modified after Veron (1995) and Taylor and Meester (1993). Small-sized herpestids with dental formula I 3/3, C 1/1, P 4/4, M 2/2; skull relatively tall; orbit completely surrounded by bony ring; anterior part of bulla as large as posterior; anterior chamber of bulla more inflated than posterior; upper incisors disposed on a transverse line; parastyle of P4 enlarged, nearly the size of the protocone; carnassial notch of m1 deep. African Species C. penicillata Cuvier, Age Ca. 3.3 Ma-Recent. Geographic Occurrence South Africa. Remarks CYlJictis was tentatively reported from Laetoli by Petter (1987), but this record cannot be considered valid (Werdelin and Dehghani, in press). The only fossil records of the genus are from South Africa. Genus GALERELLA Gray, 1865 Diagnosis Small-sized Herpestidae; skull smaller than Herpestes; dental formula I 3/3, C 1/1, P 4/3, M 2/2; p2 relatively smaller than in Helpestes. African Species G. debilis Petter, 1973, G. paiaeoserengetensis (Dietrich, 1942), G. plllve1'lllenta (Wagner, 1839), G. sangllinea (Ruppell, 1835), G. sp. Age Ca. 7.4 Ma-Recent. Geographic Occurrence Chad, South Africa, Tanzania. Remarks For convenience, we retain Galerella as a separate genus despite evidence that it does not constitute a monophyletic group (Veron et al., 2004). One of us (LW) is inclined to doubt the specific identification of G. sangllinea from the Miocene of Chad, though this material clearly does not belong to either of the previously named fossil species of Galerella. Genus HELOGALE Gray, 1861 Figures 32.SA-32.SC Diagnosis Small-sized herpestid with dental formula I 3/3, C 1/1, P 3/3, M 2/2; skull with no postorbital bar; premolars tall and sharp; P4 protocone very large and tall, much larger than the parastyle; no accessory cusps on P2-3/ p2-3, a small distal one on p4; carnassial notches shallow; m1 trigonid tall with paraconid and protoconid of roughly equal height, and metaconid tall but more reduced than the paraconid. THIRTY-TWO: CARNIVORA 625

24 FIGURE 32.5 African Herpestidae. A-C) HeZogaZe pazaeogmcilis, LAET from the Upper Laetolil Beds, Laetoli: A) cranium in ventral view, B-C) right ramus in lingual and buccal view; D) Helpestes sp., PQ-L from the Langeberg Quartzose Sand Mb., Varswater Fm., Langebaanweg, snout in ventral view; E-F) Helpestes sp., PQ-L from the Langeberg Quartzose Sand Mb., Varswater Fm., Langebaanweg, right mandible in buccal and lingual view; G-I) Atilax mesotes type specimen KA 86 from Kromdraai Mb. A, skull in ventral view and left mandible in lingual and buccal view. African Species H. hirtuia Thomas, 1904, H. kitafe Wesselman, 1984, H. paiaeogmcilis (Dietrich, 1942) (figures 32.5A-32.5C), He/oga/e sp. Age Ca. 6 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Tanzania. Remarks He/ogaie spp. are present at a number of Pliocene localities in eastern Africa, with the youngest record being from the Shungura Fm., Member G. No Heiogaie has been reported from the Pleistocene of Africa. Whether this represents a bias due to collecting methods or a bias in the sampled environments is not clear. Genus HERPESTES Illiger, 1811 Figures 32.5D-32.5F Diagnosis Modified after Veron (1995). Small- to mediumsized herpestids with dental formula I 3/3, C 1/1, P 4/4, M 2/2; skull elongated with palate triangular, lacrymal and jugal not 626 LAURASIATHERIA

25 ::r:::r:::r: ::r: ::r:o:::lrnrnrn rnrnuu unnti:lti:l>->->->- > " _. PI:l f-i 0..0 t:::: -- "... I-j -'::r C 0 :::s VI f-i::l 0.. r-;. : : : ::r: 0. up..:::: "' C Ai ::r: b b. S. c+ g 8 C , 03, 2; Q e; g g- e; e:. & 0. &. ; ro. :::. 8 ;o;o;og"'u8'b..n;::;.g"8'tno ro;;g:...,.8 0.>" 0",0.> Wf-lf-1p.J po.... :::I ro "'C <rt'_ ""''''' "<:1"";-.-nl-jro8 o.>::r:o.> (]Q;::"'S C7''Tl ro 0:. n, S e; e:. 2: ::l p.j "' () r-t" Pol 0' ;;1N 7;" a ""' A III AWax mesotes A Wax pazlidil10sus AWax sp. CrossarcJws tral1svaazel1sis CYl1ictis penicillata GaZerella debilis GaZerella DaZaeoserenuetel1sis GaZerella puzveruzenta GaZerella sanguil1ea GaZerella sp. HeZogaZe hirtula p, HeZogaZe kitafe HeZogaZe pazaeogracilis HeZogaZe sp. Helpestes abdezalii Helpestes azayzaii P, Helpestes icijneumon Helpestes sp. Herpestidae indet. IcJ171ellmia azbicallda n C It> n It> " co -l o >...,. ::r:r It> >ow D; ;:::r. 0\ 0. It>.ij It> n. Ichnellmia l1ims p, IcJ171ellmia sp. Kichechia zamal1ae KicJlechia sp. Legegetia Ilandii Leptoplesictis mbitensis LeptopZesictis namibiellsis LeptopZesictis rangwai LeptopZesictis SeJllltae MZlIlgos dietriciji MZl11gos minlltlls MZl1lgos 11lungo MZl11gos sp.."" Suricata major."" Sliricata suricatta Sllricata sp. Ugal1dictis l1apakensis

26 >-5: 0.0. C ro >-> ::s ::T r; r; a;? a ::J 8. (ti 0 rl- OJ::J Oq o 0' t-' OJ e (l) Oq t-' OJ ;j::j ;joq OJ ::J ::s (l) Oq t-' OJ C:: '0 0 '0 (l),... b:j c:: (l) ::J 0. ::;,.' Vl t""" A 1'\ A Cl "'i 0 0 bsifs2g. E a :;:; -. co c::: g; 8. ro 1"'1,... PJ g >- A b:j en 2S A. t;;' 0. N C ::g S-. A A A 1-",... PJ 1;1 2 e; 0; PJ PJ PJ aq a ::0 8- c c ::J " -e P1 f? o cr' ::s 0. ::s (l) 1-" ro (fq ::i _. ::0 S2 Atilax mesotes Atilax paludinosus Atilax sp. Crossarclllls tmnsvaalensis C},nictis penicillata Galerella debilis Galerella oalaeoseren'letensis Galerella PUlVetlilenta Galerella sanguinea Galerella sp. Helogale hirtlila Helogale kita(e () Helogale palaeogmcilis Helogale sp. Herpestes abdelalii He1pestes ala)llaii Helpestes ichneumon Helpestes sp. n -l o P -l Z r tn Z w c:: tv tn I:) Herpestidae indet. Ic/mewnia albicauda Iclmeu11lia nims Icil11ellmia sp. I Kichechia zalnanae I Kichechia sp. Legegetia nandii Leptoplesictis mbitensis Leptoplesictis namibiemis Leptoplesictis mngwai Leptoplesictis senlltae Mungos dietriclli Mungos minutus Mungos l1111ngo Mzmgos sp. Suricata major Suricata sllricatta Suricata sp. Ugandictis napake71sis

27 MoruorotHill Nachukui Middle Lomekwi Nachukui Upper Lomekwi Napak Ndolanya Beds Upper Unit Ngorora Member A Olduvai Bed I aff. aff. Olduvai Bed II Omo Shungura B Omo Shungura C Omo Shungura E+F Omo Shungura G Plovers Lake d. d. Rusinga Saldanha Sea Harvest Sibudu HP Songhor Sterkfontein Member 2 Sterkfontein Member 4 Sterkfontein Member 5 Sterkfontein Jacovec Cave Swartklip 1 Thomas I Toros-Menalla Tugen Lukeino d. Tugen Mabaget

28 in contact; anterior rim of orbits formed by the frontal, maxillary, and jugal; auditory meatus somewhat triangular in shape; anterior chamber of the bulla less inflated than the posterior one; dentition trenchant; upper incisors disposed on a straight line; three-rooted P3; carnassial notch on ml deep. African Species H. abdeia/ii Geraads, 1997, H. aiayiaii Haile Selassie and Howell, 2009, H. icll17elll1zoi1 (Linnaeus, 1758), H. sp. (figures 32.5D-32.5F) Age Ca Ma-Recent. Geographic Occurrence Chad, Ethiopia, Kenya, Morocco, South Africa, Tanzania, Zambia. Remarks Helpestes sp. indet. was recently described from Lemudong'o in Kenya by Howell and Garcia (2007). H. abdeia/ii may be close to Gaiel-ella plliveruienta (and thus not a species of Helpestes), but this requires further analysis. Toros Menalla, Chad, has yielded the earliest definite African record of the genus, though older material has been attributed to it (Morales and Pickford, 2008). Genus ICHNEUMIA 1. Geoffroy St.-Hilaire, 1837 Diagnosis Modified after Taylor (1972). Frontal region of skull expanded and more elevated than parietal region; anterior orbital margin above P4; postorbital processes well developed, forming complete postorbital bridge in fully adult skulls; sagittal and lambdoidal crests well developed; posterior chamber of bulla strongly inflated; dentition heavy, especially P4, with inner lobe of tooth occupying considerably more than half the anteroposterior diameter of the crown and metastyle reduced; Ml fairly symmetrical, parastyle slightly projecting; anterior root of zygomatic arch far behind P4; ml with connate paraconid and metaconid; m2 elongate with six cusps, lower canines recurved, upper canines only slightly recurved; dental formula I 3/3, C 1/1, P 4/4, M 2/2. African Species I. albicauda (Cuvier, 1829), I. nims Geraads, 1997, I. sp. Age Ca. 2.5 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Morocco, Tanzania. Remarks White-tailed mongooses are known only from a pair of records that probably pertain to the extant species (Petter, 1973; Geraads et ai., 2004) and the single tooth of I. nims from Ahl al Oughlam (Geraads, 1997). Howell and Garcia (2007) describe as Icl1I1ellmia aff. aibicallda a right mandibular corpus with p4 from Lemudong'o in Kenya. In view of the much greater age of this specimen than other known Ichnellll1ia, this record requires additional taxonomic scrutiny. Genus KICHECHIA Savage, 1965 Diagnosis Modified after Savage (1965). Herpestid with upper dental formula I 3/?3, C 1/1, P 4/4, M 2/2; teeth not compressed; canine long and slender; parastyle present only on P4; upper molars without conules and without hypocone, protocone crescentic and without anterior and posterior wings. African Species K. zall1c1lwe Savage, 1965, K. sp. Age Ca Ma. Geographic Occurrence Kenya, Uganda. Remarks K. ZCIlnal1ae is the most common small carnivoran of the eastern African Lower Miocene and is known from a considerable number of localities. Genus LEGETETIA Schmidt-Kittler, 1987 Diagnosis After Schmidt-Kittler (1987). Equal in size to Kichechia or somewhat smaller. Differs from Kichechia in the following characters: all cones of the dentition more acute; m2 two-rooted and very elongate bearing a complete trigonid and strong hypoconulid; p4 with strongly developed anterior accessory cusp; pi two rooted; Ml and M2 with broader shelf buccally to the external cusps; M2 with well-developed lingual cingulum; P4 more elongate. African Species L. l1al1dii Schmidt-Kittler, Age Ca Ma. Geographic Occurrence Kenya. Remarks Legetetia is known from a large number of isolated teeth and mandible fragments from several west Kenyan Lower Miocene localities. It is notable for its hypocarnivorous dentition and particularly the elongated, two-rooted m2. Schmidt-Kittler (1987) notes similarities with the European SivCllwslla-ElIboictis group and suggests that they and Legetetia may be derived from a common stock. Genus LEPTOPLESICTIS Forsyth Major, 1903 Diagnosis After Roth (1988). Small-sized carnivoran; dental formula (lower dentition only) I 3, C 1, P 4, M 2; premolars with tall cusps; p4 posterior accessory cusp very large; ml postvallid notch less deep than in Herpestes; m2 trigonid and talonid distinct. African Species L. mbitel1sis Schmidt-Kittler, 1987, L. l1al1libiensis Morales et ai., 2008, L. l'cingwai Schmidt Kittler, 1987, L. Sel111tae Morales et ai., Age Ca Ma. Geographic Occurrence Kenya, Namibia. Remarks Leptoplesictis is poorly defined, as the characters cited by Roth (1988) to distinguish it from Helpestes are likely to be plesiomorphic. It is generally considered to be a herpestid because of its similarity to Herpestes, though definitive proof from basicrania is still lacking. It is an occasional member of the European carnivoran guild in the early and middle Miocene (Roth, 1988). In Africa the genus is known from the early Miocene of Kenya and Namibia. Genus MUNGOS E. Geoffroy St.-Hilaire and Cuvier, 1795 Diagnosis Modified after Petter (1973) and Veron (1995). Small-sized herpestids with dental formula I 3/3, C 1/1, P 3/3, M 2/2; skull with orbits closed or nearly closed by a postorbital bar; posterior rim of nasals across the anterior rim of orbits; anterior rim of orbits formed by the frontal, maxillary, jugal, and lacrymal; auditory meatus wider than tall; PlIpl absent (pi present in M. dietrichi); P3 with three roots; P4 molariform, broader than long; talonid of ml present but simple and with no cuspids. African Species M. dietrichi Petter, 1963, l'vi. mil111tus Petter, 1973, Mlll1gos sp. Age Ca. 3.7 Ma-Recent. Geographic Occurrence Kenya, South Africa, Tanzania. Remarks All fossil records of lvlzll1gos are of extinct species. MZll1gos dietrichi accounts for most of these records with five occurrences. The record of this species from the Laetolil Beds, Upper Unit, is the earliest appearance of MZll1gos in Africa. In Olduvai, Bed I it is sympatric with another Mzmgos, M. minutlls, which differs from it in the loss of pi (as in the extant species) and in being markedly smaller, although Petter later (1987) synonymized the two species. 630 LAURASIATHERIA

29 Genus SURICATA Desmarest, 1804 Diagnosis Partly modified after van Staaden (1994). Smallsized Herpestidae; dental formula I 3/3, C 1/1, P 3/3, M 2/2; skull high and rounded, with large closed orbits and interorbital space greater than or equal to postorbital constriction; incisor rows slightly curved; all cheek teeth have tall cusps adapted for puncture-crushing of small prey; PI/pI absent; protocones of P4-M2Iarge, metastyles very short; m1 trigonid short, talonid long. African Species S. suricatta (Schreber, 1776), S. major Hendey, 1974, S. sp. Age Ca. 1.8 Ma-Recent. Geographic Occurrence South Africa. Remarks All fossil meerkat records are from southern Africa and all except one are referable to the extant species S. sllricatta. S. major was described by Hendey (1974) on the basis of material from Elandsfontein. It differs from S. suricatta mainly in size. Genus UGANDICTIS Morales et a!., 2007 Diagnosis Adapted from Morales et a!. (2007). Small size; m2 medium sized; m1 with v-shaped trigonid, metaconid lower than paraconid, which is located anterolingually, talonid smaller than trigonid with strong hypoconid, subdivided hypoconulid and entoconid; p4 narrow with anterior, central and posterior cuspids and a basal platform in a posterolingual position; p3 and p2 relatively short with posterior cuspids; P4 with short metastyle, paracone similar in size to the metastyle, conical in form, prominent parastyle, protocone strong, located anterior to parastyle, and posterior to the basal cingulum of the metastyle; P3 and P4 elongated, with weak basal cingula; upper molars with three roots. African Species U. napakemis Morales et ai., Age Ma. Geographic Occurrence Kenya, Uganda. Remarks Some of the material recently assigned to this new taxon by Morales et a!. (2007) was previously referred to Leptoplesictis mngwai and Kichechia zamanae. HERPESTIDAE indet. Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, Namibia, South Africa. Remarks Indeterminate Herpestidae are known from a variety of sites, the oldest of which is Ngorora Mb. A (Morales and Pickford, 2005a). Family PERCROCUTIDAE Werdelin and Solounias, 1991 Table 32.7 Genus DINOCROCUTA Schmidt-Kittler, 1976 Diagnosis Modified from Howell and Petter (1985). Percrocutids of large to very large size; P3 without internal root; p4 long relative to p3; p2 hypertrophied with a high robusticity index. African Species D. algeriensis (Arambourg, 1959), D. senyllreki (Ozansoy, 1957). Age Ca Ma. Geographic Occurrence Algeria, Libya. Remarks DinocroCllta is mainly distributed in Eurasia, where it was prominent until some time before the Miocene Pliocene boundary (Werdelin, 1996). In Africa, the genus TABLE 32.7 Occurrences of Percrocutidae species.:::j :;;, :::: "" 'r::: ;( :::: "" "" 0- Vl {l {l c: c:... ;:J ;:: ;:: '-' a '-' a '-' ;:: :::: Sites Ci Q., "" Bled Douarah Beglia Bou Hanifia Fort Ternan Menacer Nakali Ngorora Member B Ngorora Member D Ngorora Member E Sahabi Samburu Hills Namurungule.;( :::: "" :.Ei "" ;:J '-' a ;:: ;:: 2; 2; is only known from North Africa (Bou Hanifia, Menacer, Sahabi). It may be found to be present elsewhere, but the absence of DinocroCllta from Ngorora suggests that they cannot have been common members of the late middle-early late Miocene fauna of sub-saharan Africa. Genus PERCROCUTA Kretzoi, 1938 Diagnosis Modified from Howell and Petter (1985). Percrocutids of relatively small size. Last molars (M2/m2) lost; m1 metaconid absent or vestigial; tendency toward shortening of the talonid and elongation of the trigonid, accentuated in more evolved representatives; P3 with or without an internal root; P4 with a reduced protocone, situated more or less posterior to the anterior margin of the parastyle; anterior premolars hypertrophied; p2 and p3 short (relative to p4 and m1) and broad with high robusticity indices. African Species P. tobieni Crusafont Pair6 and Aguirre, 1971, P. leakeyi (Howell and Petter, 1985), P. sp. Age Ca Ma. Geographic Occurrence Kenya, Tunisia. Remarks Percrocllta tobieni is a fairly generalized, mediumsized Percrocllta. It is only known with certainty from eastern Africa, but the specimen described as Capsatl1erillm lllciae from Bled Douarah, Tunisia (Kurten, 1976), likely also belongs to this species. More than one species may be involved, as material from the Namurungule Formation, Kenya (Nakaya et ai., 1984), differs from the type specimen of P. tobieni, but the necessary comparisons have yet to be made. Aguirre and Leakey (1974) mention the presence of "Hiperl1yaena leakeyi" from Nakali, but as pointed out by Morales and Pickford (2006), this is a nomen nudllm and the name dates from Howell and Petter (1985). Morales and Pickford (2006) transfer the species from DinocroCllta, in which it was placed by Howell and Petter (1985), to Percrowta, and this is probably correct. THIRTY-TWO: CARNIVORA 631

30 Family HYAENIDAE Gray, 1821 Table 32.S Genus ADCROCUTA (Kretzoi, 1938) Diagnosis Emended after Kretzoi (1938). Hyaenidae of large size; teeth massive; premolars powerful with weak accessory cusps; m1 talonid composed of entoconid and hypoconid; metaconid poorly developed or absent; P4 protocone very reduced. African Species A. eximia (Roth and Wagner, 1854). Age Ca Ma. Geographic Occurrence Libya. Re'marks AdcroCllta is nearly ubiquitous in Eurasian faunas from MN 10-13, but in Africa is known only from Sahabi. No derived, bone-cracking hyaenid (type 6 of Werdelin and Solounias, 1996) is known from sub-saharan Africa. Genus CHASMAPORTHETES Hay, 1921 Figure 32.6D Diagnosis Emended after Kurten and Werdelin (1988). Medium- to large-sized hyaenids with long, slender limbs; rostrum broad; frontal profile stepped; cheek teeth slender, trenchant; dental formula I 3/3, C 1/1, P 3-4/3-4, M 1/1; PI large, with a tendency to be shed early in life; anterior premolars with strong posterior accessory cusps, anterior cusps variable but usually strong; P4 with large protocone, short paracone, and elongated parastyle; p4 elongate, with large accessory cusps; m1 without metaconid and with single, laterally compressed, blade-like talonid cusp. African Species C. australis (Hendey, 1974), C. darelbeidae Geraads, 1997, C. l1itidula (Ewer, 1954) (figure 32.6D), C. sp. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Libya, Morocco, South Africa, Tanzania. Remarks Chasmaporthetes is a common member of the carnivoran guild in Pliocene Old World faunas and is the only hyaenid to migrate to North America (Kurten and Werdelin, 1988). Its earliest African appearance is at Toros Menalla (Bonis et ai., 2007a), but it is best known from South Africa, where the species C. l1itidula is a characteristic member of the Transvaal cave assemblages (Ewer, 1954b; Werdelin and Turner, 1996). We here consider C. darelbeidae from Ahl al Oughlam (Geraads, 1997) to be a valid species. Genus CROCUTA Kaup, 1828 Figures 32.6A-32.6C Diagnosis Emended after Kretzoi (1938). Hyaenidae of large size; premolars massive, P4 paracone weak, metastyle blade greatly elongated; m1 trigonid elongated, metaconid vestigial or lost, talonid greatly reduced. African Species C. crocuta (Erxleben, 1777), C. dietrichi Petter and Howell, 1989 (figures 32.6A-32.6C), C. eturo110 Werdelin and Lewis, 2008, C. ultra Ewer, 1954, C. sp. Age Ca. 3.8 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks We consider C. dbaa (Geraads, 1997) to be a synonym of C. dietrichi. The history of CroCllta in Africa is complex and has only been presented in outline (Lewis and Werdelin, 1997, 2000). Werdelin and Lewis (2008) have recently described the new and aberrant C. etllrono from the Pliocene of the Turkana Basin. Still more recently, Gilbert et ai. (2008) have described a new subspecies, C. crocllta yangllia from the Daka Mb. of the Bouri Fm., Middle Awash, Ethiopia. Genus HYAENA Brisson, 1762 Figllre 32.6E Diagnosis Large-sized Hyaenidae with moderate bonecracking adaptations; limbs slender with elongated metapodials; skull robust: dental formula I 3/3, C 1/1, P 4/3, M 1/1; premolars with some bone-cracking adaptations; P4 short, with metastyle shorter than paracone; m1 with short but well-formed talonid, metaconid usually present. 100 mm FIGURE 32.6 African Hyaenidae. A-C) CroCllta dietricl1i type specimen LAET from the Upper Laetolil Beds, Laetoli, left ramus in lingual, buccal, and occlusal view; 0) Clwsmaportl1etes i1itidula SK from Member 1, Swartkrans, right and left rami in right lateral view; E) Hyaena sp. KNM-ER 1548, Upper Burgi Mb., Koobi Fora Fm., cranium in left lateral view. 632 LAURASIATHERIA

31 0 0 E v e. e. e. e. ci (l). ::c ::. -. g. 0 g. v ::0: ::l ::l ::l C (l) c ::l v - 8 = ::c = 0 = ' v",ngo'o'ciro8gg(l)"","", ::l8go", (l) (l) C ::l ::l ::l (l) u c ::r:: ro i? rv p>.., - Q1 -- e:. e:. P"" 2:. tl.j::l:::-:.-+ ::l p>::r 0'0"... ::;: 825' N 8 8;5. ce. f-'. [.I'J j:ll "' '-+... PJ CJ I"! A n tl.j aq 8 Vo Co Adcrocuta eximia Chasmaporthetes allstralis Chasmaporthetes nitidula Chasmaportlzetes darelbeidae p, CiJasmaporthetes sp. () CroClita crocuta Crocuta dietrichi Crocuta etllrono CroCllta ultra Crocuta sp. p, Hyaena ijyaena Hyaena makapani Hyaena sp. Hyaenictis graeca Hyaenictis hendeyi Hyaenictis wehaietll Hyaenictis sp. HyaenictitiJeriwl1 minimul1l HyaenictitlJeriwl1 namaqllensis o () () S (l) ::l () (l).., > o OJ ""0-< ::Crn ci3 V.J (l) N. (l) -. co (l) (). HyaenictitlJerillm parvlll1l H)laenictitlJeriwl1 sp. Hyaenidae indet. IctitiJerilll1l ebll P, IkeloiJ)laella abronia Lycyaena C7"llsa(onti Lycyaenops silberbergi PaciJycroclita brevirostris PaciJycroCllta sp. J ParaiJyaena bnmnea Parai1yaena 110welli () () () () PlioC7"ocuta perrieri Proteles amplidentus Proteles sp. () ProtictitiJeriul1l CraSS!l111 ProtictitiJeri!l111 plmicll11j ProtictitiJeriwll sp.

32 t:-'t-'t-'t-' t-' r t-'aaa AAAAAAr::r::::C g S b (:J 0. 0:; g 8 g g g ;; r::: r::: 22.::l ;:iggggg::; n;'::; ':9 0. uu;:;u'+;::;;:;:;::;ef 8 <1aq'Tj'Tj'Tj'Tj'TjN"'t;;:JO (l) PJOPJ"'iPJ PJ... - ooooo(d(ti- -- a g. ci (D. C; CL,s- 0' :::::: ::: C ;? 0 r A ::r a C bsg :g2"3 8 ;'. e::g g' :g c:t o:;rtx g 0; 0; Q g 5- _. >- Adcrocuta exi111ia Chasmaporthetes australis Chasmaporthetes nitidula Clwsmaporthetes darelbeidae Chasmaporthetes sp. CroCllta crocllta Crowta dietrichi CroCllta eturono Crocuta ultra Crocuta sp. H)!aena h)!aena Hyaena 111akapani p., Hyaena sp. p., H)'aenictis graeca H)!aenictis hendeyi Hyaenictis wehaietu H)!aenictis sp. Hyaenictitheriul11 minimum Hyaenictitheriu111 namaquensis Hyaenictitheriul11 parvum Hyaenictitherium sp. Hyaenidae indet. n >-l 0» z >-l r M Zw c:: t:i 2. (oj Co Ictitherill111 ebu p., p., Ikeloh)'aena abronia Lycyaena C1'usafonti Lycyaenops silberbergi p., Paci1YCl'ocuta brevirostris p., Pac/J)lcrocuta sp. Parahyaena bnl1111ea Parai1yaena lwwelli Pliocrowta perrier! Proteles amplidentus Proteles sp. Protictitherium crass1l111 Protictitheri1l111 pllnicul11 Protictitheriu111 sp. ----'-

33 Lothagam Lower d. d. Nawata Lothagam Upper d. el cf. Nawata Makapansgat Member 3 Melkbos Middle Awash Adu Asa Naehukui Kataboi Naehukui Lower Lomekwi Naehukui Middle Lomekwi Nakoret Nkondo Olduvai Bed I d. Olduvai Bed II Olduvai Bed IV Omo Shungura K Plovers Lake d. Sagan tole Fm. Sahabi Saldanha Sea Harvest d. Samburu Hills Namurungule South Turkwel d. cf. Sterkfontein Member 2 Sterkfontein Member 4 Sterkfontein Member 5 Sterkfontein Jacovee Cave Swartklip 1 Swartkrans Member 2 Swartkrans Member 3 Toros-Menalla Tugen Lukeino Tugen Mabaget xl

34 African Species H. h)'aena (Linnaeus, 1758), H. makapani Toerien, 1952, H. sp. (figure 32.6E). Age Ca. 3.6 Ma-Recent. Geographic Occurrence Chad, Ethiopia, Kenya, South Africa, Tanzania, Tunisia. Remarks The evolutionary history of H)'aena in Africa after the possible divergence of the genus from Ike/oh)'aena or some closely related form is fairly straightforward. H. makapani is present until ca. 2 Ma, while the extant species H. h),aena appears somewhat earlier than this, suggesting a moderate temporal overlap. The record of H. h)'aena precllrsor from A'in Brimba, Tunisia requires further analysis. Genus HYAENICTIS Gaudry, 1861 Diagnosis Emended after Werdelin et al. (1994). Mediumto large-sized Hyaenidae; mandible long and slender; dental formula I 3/3, C 1/1, P 4/3-4, M 1/1-2; p2-3 slender, with weakly developed anterior accessory cusps; P4 with metastyle slightly longer than paracone; m1 with short, two- or threecusped talonid; m2 generally present. African Species H. graeca Gaudry, 1861, H. hende)'i Werdelin et ai., 1994, H. welwietll Haile-Selassie and Howell, 2009, H. sp. Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa. Remarks H),aenictis is present at a number of African localities, of which Langebaanweg has by far the best sample. Ginsburg (1977) referred a H),aenictis from Beni Mellal to the Eurasian H. graeca, but the specific referral must be considered doubtful. Genus HYAENICTITHERIUM Zdansky, 1924 Diagnosis Modified after Kretzoi (1938). Medium- to largesized hyaenids; dentition more crushing than in Ictitherillln, with shorter muzzle, stronger mandibular ramus, shorter and more massive canines; m1 talonid with two cusps; m2 and M2 present but reduced. African Species H. minimllm Bonis, Peigne, Likius, Mackaye, Vignaud and Brunet, 2005, H. na1naqllensis (Stromer, 1931), H. d. parvliill (Khomenko, 1914). Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Libya, South Africa. Remarks Most records of H),aenictitherill11l in Africa are referable to H. na1naqllensis, but some are smaller. It may turn out that the material referred to H. d. parvlll11 by Werdelin (2003) should be placed in the approximately coeval species from Chad, H. minimllm. Genus ICTITHERIUM Roth and Wagner, 1854 Diagnosis Modified after Kretzoi (1938). Medium-sized forms with generalized doglike hyaena morphology; m1 talonid large, low; P4 with metastyle shorter than paracone and protocone reduced; m2 and M2 present and unreduced; premolars with incipient crushing morphology. African Species I. ebll Werdelin, Age Ca Ma. Geographic Occurrence Kenya. Remarks Ictitherill1n is a common member of the later Miocene carnivoran guild of Eurasia but is rare in Africa, with the only record being the aberrant, long-limbed I. ebll from Lothagam (Werdelin, 2003), though Semenov (2008) has questioned the generic attribution of this species. Genus IKELOHYAENA Werdelin and Solounias, 1991 Diagnosis Modified from Werdelin and Solounias (1991) Slightly smaller than extant H. h)'aena in size; maxillary contribution to zygomatic arch large; premolars, especially P3 and p3, enlarged, but not strongly conical, and anterior edge only slightly convex; anterior accessory cusps not appressed to main cusp of premolars; infraorbital foramen positioned above midline of P3; P4 metastyle short; M2 and m2 generally present. African Species 1. abrollia (Hendey, 1974). Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks Ike/oh)'aena abronia, known from both southern and eastern Africa, is very similar to the extant striped hyaena except for the retention of the second molars. It is not clear whether 1. abronia should be placed before or after the split between H)'aella and Parah),aella, and we therefore retain the genus Ike/oh)'aena here. Genus LYCYAENA Hensel, 1863 Diagnosis Modified after Kretzoi (1938). Large-Sized forms with slender premolars with moderately strong accessory cusps; m1 talonid short; m2 very reduced; M2 lost. African Species L. C1'llsafonti Kurten, Age Ca Ma. Geographic Occurrence Tunisia. Remarks The Eurasian hyaenid genus L)'c)'aella, a primitive member of the hunting hyaena lineage (Werdelin and Solounias, 1991, 1996), is only known from a single African record; a lower jaw fragment from Bled Douarah, Tunisia, and apparently never reached sub-saharan Africa. Genus LYCYAENOPS Kretzoi, 1938 Diagnosis Emended after Kretzoi (1938). Closely related to L)'c)'aena but with lower and longer m1; significantly more massive premolars with well-developed accessory cusps; p2-p3 posteriorly expanded, with posterior end squared off. African Species L. si/berbergi (Broom in Broom and Schepers, 1946). Age Ca Ma. Geographic Occurrence South Africa. Remarks This species is commonly placed in Chasmaporthetes (see Werdelin and Turner, 1996), but Werdelin (1999b) noted similarities between it and the European L. rhomboideae and suggested that they might be congeneric. Genus PACHYCROCUTA Kretzoi, 1938 Diagnosis Modified after Kretzoi (1938). Very large, robust Hyaenidae; premolars very massive; p2-3 lacking anterior accessory cusps; P4 with massive paracone and protocone; m1 short and broad lacking metaconid; talonid consisting only of the hypoconid. African Species P. brevirostris (Aymard, 1846), P. sp. Age Ca Ma. Geographic Occurrence Kenya, South Africa, Tanzania. 636 LAURASIATHERIA

35 Remarks This highly distinctive species has long been known from South Africa as P. bel/ax (Ewer, 1954a). More recently, it has been identified from eastern Africa as well (Werdelin, 1999a). Genus PARAHYAENA Hendey, 1974 Diagnosis Large-sized Hyaenidae with moderate to strong bone-cracking adaptations; limbs slender with elongated metapodials; premolars robust; P3 and p3 with strong bonecracking ability; dental formula I 3/3, C 111, P 4/3, M 1/1; P4 with large protocone, metastyle longer than paracone; m1 short with short but well-formed talonid, metaconid commonly lost. African Species P. brzl11lzea (Thunberg, 1820), P. howelli Werdelin, Age Ca. 4.3 Ma-Recent. Geographic Occurrence Kenya, South Africa, Tanzania, Zambia. Remarks Living brown hyenas are restricted to southern Africa, but P. brllljljea has been recorded as a fossil in eastern Africa (Werdelin and Barthelme, 1997). The presence of P. Iwwelli at several eastern African localities suggests that the origin of the species may lie there rather than in southern Africa. Genus PLIOCROCUTA Kretzoi, 1938 Diagnosis Modified after Kretzoi (1938). Hyaenidae of large size; with unreduced P4 protocone; short, massive m1 lacking metaconid but with a wide talonid bearing ento- and hypoconid. African Species P. perrieri (Croizet and Jobert, 1828). Age Ca Ma. Geographic Occurrence Ethiopia, Morocco. Remarks The only certain African record of this species (P. perrieri latidens Geraads, 1997) is from Ahl al Oughlam. The taxon has been reported from other North African sites, such as Aln Boucherit, but the only eastern African record that may be valid is from the Sidi Hakoma and Denen Dora Mbs at Hadar, though this material also requires renewed evaluation. Genus PROTELES 1. Geoffroy St.-Hilaire, 1824 Diagnosis Hyaenidae with cheek teeth reduced to pegs; dental formula I 3/3, C 1/1, P+M 3-4/2-4, sometimes less; canines not reduced; skull slender; palate long and rectangular. African Species P. a17lplidentus vverdelin and Solounias, 1991, P. cristatus (Sparrman, 1783), P. sp. Age Ca. 2.5 Ma-Recent (possibly as old as 4 Ma). Geographic Occurrence South Africa. Remarks Although Prateles is easy to recognize from craniodental remains, it is only known from scarce records in South Africa. P. amplidentus differs from the extant species in its slightly less reduced cheek teeth. Postcranial remains of a very small hyaenid from the Laetolil Beds, Lower Unit may possibly belong to Prateles or a related form (Werdelin and Dehghani, in press). Genus PROTICTITHERIUM Kretzoi, 1938 Diagnosis Modified after Kretzoi (1938). Smale slender Hyaenidae; well-developed P4 protocone; m1 with very tall talonid cusps, tall, well-developed metaconid, talonid relatively short; upper molars relatively reduced. African Species P. CmSSll111 (Deperet, 1892), P. PZlI1iCllll1 (Kurten, 1976), P. sp. Age Ca Ma. Geographic Occurrence Kenya, Morocco, Tunisia. Remarks The African representation of this primitive hyaenid genus is limited. The only certain records are from Bled Douarah (see Werdelin and Solounias [1991) for a discussion of this material) and Beni Mellal, described as IctitiJerilllll d. ammbourgi, a synonym of P. cmssllll1, d. Schmidt-Kittler (1976). A single sub-saharan record also exists, described as Ictitizeriwl1 sp. by Nakaya et al. (1984). HYAENIDAE indet. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Libya, South Africa, Uganda. Remarks Unidentified and indeterminate Hyaenidae are known from a number of localities of late Miocene to early Pleistocene age. With the exception of "hyaenid sp. E" from Langebaanweg (Hendey, 1974) and a specimen from Chad showing similarities to it, none appear to represent taxa other than those already listed. Family BARBOUROFELIDAE Schultz et al., 1970 Table 32.9 Genus AFROSMILUS Kretzoi, 1929 Figures 32.7C-32.7E Diagnosis Modified after Morlo et al. (2004). Mediumsized barbourofelids with upper canines very compressed; P2 absent; P3 with anterior accessory cusp present; P4 protocone about the size of the parastyle; M1 reduced with P4 lengthlml width ratio ; M1 with protocone very reduced but still distinct and metacone completely reduced and not projected posteriorly; m1 with talonid vestigial or absent; mandibular flange poorly developed. African Species A. africmllls (Andrews, 1914), A. turkanae Schmidt-Kittler, 1987 (figures 32.7C-32.7E), A. sp. Age Ca Ma Ma. Geographic Occurrence Kenya, Namibia. Remarks The genus is the best known of the family in Africa, although our knowledge remains rather limited. This is apparently the only Lower Miocene African genus that migrates out of the continent, with A. hispaniclis known from the Lower Miocene of Spain (Morales et al., 2001b); this constitutes evidence of a first migration of the family to Europe. Genus GINSBURGSMILUS Morales et al., 2001 Figures 32.7F-32.7K Diagnosis Barbourofelids with upper canine transversely compressed, with crenulated anterior and posterior crests, and vertical grooves on both faces; P2 present but very reduced; p3 with anterior accessory cuspid present. African Species G. napakel1sis Morales et al., 2001 (figures 32. 7F-32. 7K). Age Ca Ma. Geographic Occurrence Kenya, Uganda. Remarks The single species assigned to the genus is the oldest and most primitive of the family, the origin of which remains unknown. The available material of Ginsblirgsmillis does not display any derived character of the upper dentition; consequently, those mentioned in the revised diagnosis THIRTY-TWO: CARNIVORA 637

36 F B c E K J o 50mm FIGURE 32.7 African Barbourofelidae. A-B) Syrtos1IJilus syrtensis type specimen MNHN right ramus from Jebel Zelten in buccal and occlusal view; C-E) Afrosmillis tllrkolloe type specimen KNM-MO left ramus from Moruorot in buccal, lingual, and occlusal views; F-K) GillSblirgsmillis Ilopokellsis type specimen UM-P67-13 right C and P3-P4, from Napak I: F-H) right upper canine in lateral, medial, and anterior views, I-K) right P3-P4 in buccal, lingual, and occlusal views. proposed here could apply to any basal barbourofelid (see Morlo et al., 2004). Morales et al. (200lb) also mentioned diagnostic characters of the lower dentition, based on a specimen initially described by Schmidt-Kittler (1987, p. 118) as AfroS11lilliS turkanae. The only significant character may be the presence of a small anterior accessory cuspid on p3 (not "p2" as specified in the original diagnosis [Morales et al., 200lb: 98]). If correctly assigned, the calcaneum of this species recently described from Napak I (Morales et al., 2007) indicates plantigrade hindlimb posture. Genus SYRTOSMILUS Ginsburg, 1978 Figures 32.7A and 32.7B Diagnosis Medium-sized barbourofelid, smaller than Sa7Js(l11OsmilliS pa/11lidens; pi, p2, and m2 absent; mandibular flange little developed and less pronounced than in Sansanosmillls; mandibular corpus strongly curved; diastema between c and p3 longer; p3 less reduced than in Sa11Sanosmilus. African Species S. syrtensis Ginsburg, 1978 (figures 32.7A and 32.7B). Age Ca Ma. Geographic Occurrence Libya. Remarks The single and holotype specimen from Jebel Zelten is a fragmentary right hemimandible with p4 and ml crown base. The absence of p2, the length of the c-p3 diastema and the strong curvature (in dorsal view) of the dentary indicate that this is an advanced species of the family, more derived than Ginsburgs11li/liS and AfrosmilliS. Genus VAMPYRICTIS Kurten, 1976 Diagnosis Derived from Morlo et al. (2004). Large-sized barbourofelid with crenulated deciduous upper canine; lower carnassial elongated, extremely compressed, with strongly curved blade, no talonid and vestigial metaconid. African Species V. vipem Kurten, Age Ca Ma. Geographic Occurrence Tunisia. Remarks This species was the youngest member of the family in Africa until the recent discovery of cranial material from Kenya (discussed later). Although poorly known (fragment of upper canine and ml), the familial assignment is well supported. The ml of Vanzpyrictis is much more derived than the contemporary machairodontine felid MachairodZls aphal1istlls and has a barbourofelid dental character: the reduction of the talonid (absent in Vampyrictis) before that of the metaconid (vestigial). The species has reached the same evolutionary grade as the latest European Sal1sanosmilus in terms of sabertooth adaptation. From what we know of the 638 LAURASIATHERIA

37 TABLE 32.9 Occurrences of Barbourofelidae species :::t i:! <::l u :t...i 2 p Sites "" Bled Douarah Beglia Fiskus Grillental Jebel Zelten Karungu (Nira & Kachuka) Langental Locherangan Moruorot Hill Napak Rusinga Samburu Hills Namurungule Songhor <::l <u.;:; <::l ;:: ;:: [:' <::l aj <u OJ t 0., :g 2!...;::.:;,. QJ. i '+< Yl.- : ] : :C 0!-;..i ;::i E ;:: E 0...;..0 aj -;:; <::l p p 3!-;'lj ;:: E' OJ G "" "" <'0. ;:: dental eruption sequence in barbourofelids and given the young age of the holotype individual, the fragment of upper canine is probably deciduous (Bryant, 1988; pers. obs.). BARBOUROFELIDAE indet. Age Ca Ma. Geographic Occurrence Kenya. Remarks Tsujikawa (2005) recently described a nearly complete skull with damaged/worn dentition from the Namurungule Formation and assigned it to the felid subfamily Machairodontinae. However, this specimen undoubtedly belongs to a derived barbourofelid species. From the available illustrations (Tsujikawa, 2005: figure 2), the skull is ca. 20 cm long from the posterior margin of the OCCipital condyle to the anteriormost border of the premaxilla. In addition, the general morphology of the skull (short, wide snout, strong maxillary constriction just in front of P3, short palate, strongly lowered postglenoid process, reduced interbullar space, dorsally located occipital condyles, etc.), and the upper dentition (anteroposteriorly short but strongly compressed upper canine, absence of P2, short postcanine diastema, reduced P3 and much more elongated P4) distinguishes this specimen from contemporary machairodontines and indicates that it probably belongs to Smlsanosmillis (size of S. va/lesiensis) or to a new, closely related genus. The published information is too limited to allow for a precise taxonomic allocation of the Namurungule specimen. Family FELIDAE Fischer, 1817 Table Genus ACINONY Brookes, 1828 Diagnosis Large-sized Felidae; dental formula I 3/3, C 1/1, P 3/2, M 1/1; skull rounded with high vault, splanchnocranium nearly vertical; limbs slender, claws semiretractile, claw sheaths reduced; cheek dentition sectorial; premolars with tall accessory cusps. African Species A. aic/w Geraads, 1997, A. jllbatus (Schreber, 1776), A. sp. Age Ca. 4 Ma-Recent. Geographic Occurrence Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks The fossil history of the cheetah in Africa is poorly known. The earliest Acinonyx are recorded from Sterkfontein Mb. 2 and the Laetolil Beds, Upper Unit, but these and most later records cannot be definitely assigned to the living species. The modern species is, however, present at Swartkrans, Mb. 3, in the latest Pliocene or early Pleistocene. Genus AMPHIMACHAIRODUS Kretzoi, 1929 Diagnosis Modified from Sardella (1994). Felids of large size; skull massive and elongated; symphyseal region of mandible robust; ascending ramus of mandible low; teeth crenulated; incisors robust; lower incisors elevated relative to cheek teeth; upper canine compressed and elongated; lower canine reduced in height; P2 present but very reduced; P3, p3, and p4 reduced relative to P4 and m1 but with well-developed cusps; upper carnassial with small preparastyle and moderately reduced protocone; m1 with very small talonid. African Species A. kabir Peigne et ai., 2005, A. sp. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Libya, South Africa, Tanzania. Remarks AmphimaciJairodlls spp. are not uncommon elements in late Upper Miocene faunas in Africa, but the remains are nearly always very limited and the only named species is A. kabir from Chad (Peigne et ai., 2005). The African record of Amphimaclwirodlls was recently reviewed by Sardella and Werdelin (2007). THIRTY-TWO: CARNIVORA 639

38 u u u u u u n n > > > > > > > > > > > > > a g g g g g g & a " ; s s..s..::j i"<'a"d" CLAvi Vl Vl en en Vl S PJ tobj;:l}:l> s - ro c U ro _ - crtdoroe:.gpjopjgtdoo'lulf-" ::J,"""", Op.j :::1PJ::JPJPJ"""'OVl (DgPJO---'-"''-''o PJrl- ej-pjcfl1 r+"<(ti --?:l' >--'::J>-iAAA'-" PJ 3.8 (!l n PJ ro PJ >-l PJ PJ PJ A,...J 5' n. t; ::J 0 0= cr rr e:.. PJ" (: N c:cn : '-''-'--- a PJ A ce. C ' Acil1ol1)' jllbatlls Acil1oll),x aiclw Acinoll),x sp. AI1lphi11lachairodlls kabir A11lphimaclwirodllS sp. Carneal earncal Carneal sp. Dimnanto(elis (erox Dino(elis aronoki Dino(elis barlowi Dino(elis darti Dillo(elis diastemata Dino(elis petteri Dino(elis pivetemti Dino(elis sp. Felis margarita Po Felis silvestris Po Felis sp. Felidae indet. Po Homotheri1l111 lwdarellsis HOl71otherill17J problematicll171 H011lotheriu17J a(rieaml171 HOlllotherilll1l sp. LeptailllntS serval LeptailllnlS sp. Po Lokotllnjailunts el1lageritus Lynx sp. Lynx tho17jasi Machairodus aphanistus Machairodlls robimol1i Machairodlls sp. Megantereon ekidoit Megantereon ekidoit/whitei l\;[egantereol1 whitei Metailunts ObSCllntS lvietailllrus sp. Nmna(elis minor Po Po PantiJern leo.-..) Po PantiJern pm'dus Panthern sp o n 8 :; ro ej» ro r Q.,l-l '-'0 ro s: 0 ro.cj ro n.

39 E1 Harhoura Equus Cave Eshoa Kakurangori I I I I I I I I I I I I I I Fish Hoek Gladysvale I Gratte des felins xix Hadar Denen Dora Hadar Kada Hadar Hadar Lip Hadar Pinnacle Hadar Sidi Hakoma Bopefield I Inolelo 1 Isenya d. Jebel Zelten Kabwe I I II Kanam East Kanapoi Kifan Bel Ghomari Kipsaraman I Klasies River Mouth Konso-Gardula 1 Konso-Gardula 2 Koobi Fora KBS Koobi Fora Lonyumun Koobi Fora Okote I I I I I Koobi Fora Tulu Bar I I I I Koobi Fora Upper Burgi I I I I d. I Kora Tara Kosia Kassam Bougoudi I Kramdraai Member A I I I I II Laetolil Beds Lower Unit Laetolil Beds Upper I I I I I I I I I I I d. I Unit Langebaanweg PPM I I Langebaanweg QSM Leba I I I I I I I I I I I I I I I lx

40 OOOZ A" ZZ o..::>j CO"," CCC::l::l-O o..rl: ;:r;... f-" 0 >...::::: r+ tjj tjj tjj rt> rt> rt> < :::: tjj rt> 0. V> C '0 '0 Z Z r n o r n ::r' o ::r' S C -C rt> ". e '0 '0 S C rt> :>;". 0: 0. ;;; 6:;;:: ;;:: S&8-&8: CJW SCCC;;;vS ;g.222;p.8 vp. ;; ;; 8. 9e:.; - n; a.;p. to (D foot goo... C (;:' ;;:: ;;:: " v ""d 0 0 ::l V> oq ;';. ;;:: rt> g. r o z" ::r' :2 8' S C '0 '0 r r r r o 0 0 rt> :2 0. Pol PJ PJ PJ 0 8'SSS b 0-' :2. rp 2 g ::l aq c.,., &' Acinonyx jllbatlls Acinonyx aieha Aeinonyx sp. Amphimaehairodlls kabir AmphimaclwirodllS sp. Caraeal earaeal Caraeal sp. Diamantofelis ferox Dinofelis aronoki Dinofelis barlowi Dinofelis darti Dinofelis diastemata Dinofelis petted Dinofelis pivetealli Dinofelis sp. Felis margarita Felis silvestris Felis sp. Felidae indet. n o-j o > z o-j r ZW c:: to r.j H o 0 H017lotheri1l11l hadare/jsis H011lotiJerillm problematicum Homot/Jerill171 african lim HOl1lotherillm sp. Leptailll1'llS serml p., I Leptailll1'llS sp. I Lokotll11jailll1'llS emageritlls Lynx sp. Lynx tilomasi Maehairodlls apiwnistlls Maehairodlls robinsoni Maehairodlls sp. Megantereon ekidoit Megantereon ekidoitlwhitei Megantereon whitei Metailll7'llS ObSCllntS n n I Metailll1'llS sp. Namafelis minor Panthera leo PantiJera pardlls Pa1Jthera sp.

41 Omo Shungura A d. Omo Shungura B d. Omo Shungura C d. d. Omo Shungura D Omo Shungura E+F Omo Shungura G d. d. Omo Shungura L Omo Usno d. d. d. Oulad Hamida I-rhino cave Oulad Hamida III-H. erectlls Plovers Lake d. d. Sagantole Fm. Sahabi aff. Saldanha Sea Harvest Samburu Hills Namurungule Shoshamagai Sibudu HP Sidi Abderrahman Songhor South Turkwel Sterkfontein Member 2 Sterkfontein Member 4 Sterkfontein Member 5 Sterkfontein Jacovec Cave Swartklip 1 Swartkrans Member 2 Swartkrans Member 3 Taung Hrdli<;:ka deposits Thomas I-H. erectlls cave Tighenif? Toros-Menalla Tugen Lukeino d.

42 Genus CARACAL Gray, 1843 Diagnosis Derived from Salles (1992). Medium-sized felidsj skull with deep pterygoid fossa with marked lateral pterygoid processj subarcuate fossa of the petrosal residualj internal auditory meatus with distinctive, salient borderj marked angle formed by the nasal and frontal parts of the profile (45 0 or higher)j rostral constriction limited to the posterior frontonasal regionj anterolateral projection of the jugal over the infraorbital foramen present. African Species C. cameal (Schreber, 1776), C. sp. Age Ca. 4-Recent. Geographic Occurrence Ethiopia, Morocco, South Africa, Tanzania. Remarks Throughout the Plio-Pleistocene, there are a number of finds of medium-sized felids. These come in two size groups. One of these matches C. eameal in size and the other matches 1. serval. On the assumption that these size groups at least represent the modern genera, if not species, the oldest Cameal are from Sterkfontein, Mb. 2 and from the Us no Fm, Ethiopia. Genus DIAMANTOFELIS Morales et al., 1998 Diagnosis Emended after Morales et al. (1998). Mediumsized Felidae with rounded mandibular symphysisj p2 absentj short postcanine diastemaj premolars narrow and highj m1 with talonid reduced and lacking a metaconid. African Species D. ferox Morales et al., Age Ma. Geographic Occurrence Namibia. Remarks This early felid, known from a partial mandibular corpus and some tentatively referred postcrania from Arrisdrift, bears similarities in the cheek teeth to Afrosmiitls (Morales et al., 1998) but is clearly not of the sabertoothed type. Its affinities are obscure, except for an apparent close relationship to Nmnafelis minor (discussed later). D Genus DINOFELIS Zdansky, 1924 Figure 32.8B Diagnosis After Werdelin and Lewis (2001). Machairodontinae of moderate to large sizej cranium rounded, with large to very large sagittal crestj lower jaw stoutj coronoid process large compared to other Machairodontinaej no anterior flange present but anterior margin of ramus strong and flatj upper canine short and only very slightly mediolaterally compressedj no longitudinal groovesj posterior crest well developed, but no serrations presentj P2 lost except in D. cristataj P3 not or slightly reduced and slender, but less so than in other Machairodontinaej P4 with somewhat reduced protoconej lower canine reduced but not incisiformj p2-p4 slenderj m1 elongated, but less so than in other Machairodontinae. African Species D. aronoki Werdelin and Lewis, 2001, D. barlowi (Broom, 1937), D. darti (Toerien, 1955), D. diastemata (Astre, 1929), D. petted Werdelin and Lewis, 2001, D. pivetealli (Ewer, 1955) (see also figure 32.8B), D. sp. Age Ca Ma. Geographic Occurrence Chad, Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks Dinofelis is perhaps the most common fossil felid taxon in the African Neogene. There was an extensive endemic radiation of the genus in Africa that is still only 100 mm FIGURE 32.8 African Felidae. A) HOl11otlleriwl1 sp. KNM-ER 931 right ramus from the KBS Mb., Koobi Fora Fm., in buccal view; B) Dillofelis pivetealli KNM-ER left ramus (reversed) from the Okote Mb., Koobi Fora Fm., in buccal view; C) Palltilem pardlls KNM-ER 3848 right hemimandible from the Upper Burgi Mb., Koobi Fora Fm., in buccal view; D) Felidae indet. PQ-L left hemimandible (reversed), unknown member, Varswater Fm., Langebaanweg in buccal view. partly explored (Werdelin and Lewis, 2001). The referral of material from Lukeino to the European species D. diastemata (Morales et al., 2005) is in our opinion based on material that is not diagnostic at the species level, and in any case the referral of material from Langebaanweg to that species cannot be maintained (Hendey, 1974j Werdelin and Lewis, 2001). Genus FELIS Linnaeus, 1758 Diagnosis Modified after Salles (1992). Small-sized Felidaej dental formula I 3/3, C 1/1, P 3/2, M l/1j premolar cusps tall, trenchant; m1 metaconid and talonid absent or vestigial; 644 LAURASIATHERIA

43 except F. ChCllIS, skull with deep pterygoid fossa with marked lateral pterygoid process, marked angle formed by the nasal and frontal parts of the profile (45 0 or higher), and modified flat, enlarged, and medially inflected lower rim surface of the orbit; great lateral expansion of the frontal bone, frontal breadth measuring more than half the condylobasal length; anterolateral projection of the jugal over the infraorbital foramen present. African Species F. margarita (Loche, 1858), F. sill'estris Schreber, 1775, F. sp. Age Ca. 7 Ma-Recent. Geographic Occurrence Chad, Egypt, Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks Felid remains of the size of Felis spp. are known from many localities in Africa. The oldest of these is Toros Menalla (Peigne et ai., 2008b). The fragmentary nature of most of the material makes referral to a species difficult. The oldest material referred to a specific species of Felis is F. sill'estris lybica from Ahl al Oughlam. The tooth from Beni Mellal identified by Ginsburg (1977) as Felis sp. is more likely to belong to a viverrid. Genus HOMOTHERIUM Fabrini, 1890 Figure 32.8A Diagnosis Emended after Sardella (1994). Machairodont of large size; dentition serrated and crenulated; incisors robust, 13 caniniform; incisors placed in an arch; C flattened and elongated, crenulations large; c small, subequal to i3; p3 reduced, sometimes lost in derived forms; P4 with completely reduced protocone, preparastyle present; p4 reduced, main cusp large; ml elongated, metaconid and talonid absent; skull elongated, especially the splanchnocranium; sagittal and lambdoid crests large and robust. African Species H. afriea Petter and Howell, 1987, H. hadarel1sis Petter and Howell, 1988, H. problematiclli1j (Collings, 1972), H. sp. (figure 32.8A) Age Ca Ma. Geographic Occurrence Algeria, Ethiopia, Kenya, Morocco, South Africa, Tanzania. Remarks H0111otheriz1711 is one of the most common carnivorans in African Plio-Pleistocene sites, though it is rarely present in large numbers. Because of the fragmentary nature of much of the material, the taxonomy and evolution of H0111otherill171 in Africa is poorly understood. Genus LEPTAILURUS Severtzow, 1858 Diagnosis Derived from Salles (1992). Medium-sized felid; skull with marked lateral expansion of the frontal bone; marked angle formed by the nasal and frontal parts of the profile (45 0 or higher); rostral constriction limited to the posterior frontonasal region; anterolateral projection of the jugal over the infraorbital foramen present; protocone of P4 markedly reduced. African Species 1. serl'al (Schreber, 1776),1. sp. Age Ca. 4 Ma-Recent (possibly as old as 6 Ma). Geographic Occurrence Ethiopia, Kenya, South Africa, Tanzania, Zambia. Remarks If the comments provided under Carneal apply, the earliest LeptaihtnlS is from Makapansgat Mb. 3 and from the Laetolil Beds, Upper Unit, unless the record from Lemudong'o (Howell and Garcia, 2007) can be verified as coming from this genus. Genus LOKOTUN/AILURUS Werdelin, 2003 Diagnosis After Werdelin (2003). Felidae of large size; mandibular horizontal ramus slender, ascending ramus relatively tall; upper canine strongly laterally compressed, with serrations present on both anterior and posterior edges; P2 present but small and peglike; upper carnassial long and slender with completely reduced protocone; p3 small and single rooted, though not peglike; lower carnassial long, slender, and low; metaconid-talonid complex absent; appendicular skeleton relatively slender, lacking extreme machairodont features; first digit of manus very robust, with extremely large claw, more than twice the size of the claws on the other digits. African Species 1. emageritlls Werdelin, Age Ca Ma. Geographic Occurrence Kenya. Remarks This form, known from a partial skeleton and other remains from Lothagam, shows a mixture of derived and primitive features (Werdelin, 2003). Its relationship to other African machairodont cats is unknown at present. It is most similar to a medium-sized machairodont from Langebaanweg, but that taxon is considerably less dentally derived (Hendey, 1974; Sardella and Werdelin, 2007). A second possible record of the species was recently reported from Lemudong'o by Howell and Garcia (2007). Genus LYN Kerr, 1792 Diagnosis Small- to medium-sized felids with short face; P2 absent; three grooves on upper canine; limbs long, especially posterior limb; tail short. African Species 1. thomasi Geraads, 1980, 1. sp. Age Ca Ma. Geographic Occurrence Morocco. Remarks Fossils of Lyl1x spp. are rare in Africa and have thus far only been identified in North Africa. Lyl1x thomasi from Oulad Hamida is a clearly distinct species, but its relationships to other members of the genus remain obscure. Genus MACHAIRODUS Kaup, 1833 Diagnosis After Anton et al. (2004). Lion-sized felid; skull narrow across the zygoma, with moderately convex dorsal profile, well-developed sagittal crest, zygomatic arch low and gently curved in side view, temporal fossa elongated, paroccipital process well developed and projecting inferiorly beyond the relatively small mastoid process, nasofrontal suture intermediate between pantherine (pointed) and evolved machairodontine (straight) condition, postorbital processes large but low; small lower incisors arranged in a straight row; large lower canines with flattened roots and an oval cross section to the crown; small diastema between c and p3; large lower premolars with a complete set of accessory cusps and p3 large relative to p4; well-developed metaconid-talonid complex on ml; mandibular horizontal ramus thick and high, with an undeveloped mandibular flange, coronoid process high and posteriorly inclined; relatively large upper incisors set in a shallow arc; high-crowned and very flattened upper canines; P2 variably present; P3 with posterior expansion; upper carnassial with a distinct protocone and preparastyle. African Species?M. aphcl11istus Kaup, 1833, M. robinsoni Kurten, Age Ca Ma. Geographic Occurrence?Ethiopia, Tunisia. THIRTY-TWO: CARNIVORA 645

44 Remarks Anton et al. (2004) restrict Maclzairodus to M. aphmzistus only, but we feel that M. robinsoni, though poorly known, is so similar to M. aplwnistus in its known features that it is difficult to distinguish them at the specific, let alone generic level. The record of M. aphanistus from Chorora (Geraads et ai., 2002) must at present be considered non-diagnostic and pertains to a machairodont of unknown affinities. Genus MEGANTEREON Croizet and Jobert, 1828 Diagnosis Medium-sized sabertooth felids; powerful forequarters, relatively short limbs; dental formula I 3/3, C 1/1, P 2/2, M 1/1; C very long, lacking serrations; m1 not elongated, metaconid and talonid lost. African Species M. ekidoit Werdelin and Lewis, 2000, M. whitei (Broom, 1937). Age Ca Ma. Geographic Occurrence Ethiopia, Kenya, South Africa. Remarks The specific identity of African Megantereon has been intensively debated over the past 20 years (Turner, 1987; Martinez Navarro and Palmqvist, 1995, 1996; Palmqvist, 2002; Werdelin and Lewis, 2002) but now appears to be resolved, at least so far as M. wijitei is concerned (Palmqvist et ai., 2007). Genus METAILURUS Zdansky, 1924 Diagnosis Medium-sized machairodont felids with rounded skull; dental formula I 3/3, C 1/1, P 2/2, M 1/1; C straight, uncompressed, lacking longitudinal grooves, lacking serrations; cheek teeth sectorial; P4 protocone reduced but present; M1 very small; m1 with vestigial metaconid. African Species M. major Zdansk»), 1924, M. ObSCZLrlIS (Hendey, 1974), M. sp. Age Ca Ma. Geographic Occurrence Kenya, South Africa, Tanzania. Remarks Hendey (1974) suggested relationships of his F. obscllm to Sivapanthem Kretzoi, 1929, while Turner et al. (1999) refer Langebaanweg material (presumably F. obscllm) to the North American genus Adelphailurus. Recently, Morales et al. (2005) placed the species in Megantereon, but this seems in part to be based on a mistaken belief in the presence of Megantereon in the Upper Miocene of Baode, China (Zdansky, 1924). In fact, the Chinese specimen can be referred to Pammaclwerodlls. Instead, the Lukeino material shows no appreciable differences from MetaihLrlIS spp. Petter (1973) tentatively attributes a specimen from Olduvai, Bed II to MetailllntS, but, though compelling, this requires further material for confirmation. Most recently, Howell and Garda (2007) report M. major from Lemudong'o, Kenya on the basis of a mandible fragment with p3 and p4. Though referral to Metailzmls seems indicated, we doubt that this material is diagnostic at the species level. Genus NAMAFELIS Morales et ai., 2003 Diagnosis After Morales et al. (2003). Felidae the size of Camcal camenl; lower dental series p2-m1; symphysis rounded; m1 trigonid lacking a metaconid, talonid formed of a welldefined cusp surrounded posteriorly by a low cingulum. African Species N. minor Morales, Pickford, Fraile, Salesa and Soria, Age Ca Ma. Geographic Occurrence Namibia. Remarks Like Oiamantofelis (see above), this is a felid of pseudaelurine grade with no clear affinities to any particular Eurasian Pseudaelzmls. Genus PANTHERA Oken, 1816 Figllre 32.8C Diagnosis Large-sized Felidae; dental formula I 3/3, C 1/1, P 3/2, M 1/1; hyoid ossified; premolar accessory cusps, especially the posterior, generally large; small talonid present on m1; canines short and robust. African Species P. leo (Linnaeus, 1758), P. pm dus (Linnaeus, 1758) (see also figure 32.8C), P. sp. Age Ca. 4 Ma-Recent. Geographic Occurrence Algeria, Ethiopia, Kenya, Morocco, South Africa, Tanzania, Zambia. Remarks The early history of Panthem in Africa is very fragmentary and although some commentators (e.g., Turner, 1990) have suggested that the extant species P. leo and P. pm dus are present in the Laetolil Beds, Upper Unit (ca. 3.7 Ma), no Panthem material older than ca. 2 Ma is diagnostic at the species level. From about this time onwards the extant species are the only species of Panthem in Africa. FELIDAE indet. Figure Age Ca. 18 Ma-Recent Geographic Occurrence Angola, Chad, Ethiopia, Kenya, Libya, Morocco, South Africa, Tunisia, Uganda. Remarks Records of undetermined felids in Africa are numerous (figure 32.8D). Most of these records are based on fragmentary material that does not allow for a better assignment. Other records, from faunal lists, need to be confirmed. This is the case for early and middle Miocene records such as Maboko (Pickford, 1986) and Jebel Zelten (Savage and Hamilton, 1973; Ginsburg, 1979). The earliest undoubted record of Felidae in Africa is an undescribed small species from Songhor (Kenya), represented by a partial hemimandible. Some records are based on recent finds, like the rich samples from Toros Menalla (Chad), Koobi Fora (Kenya), and the Middle Awash (Ethiopia). A first glance at the Chadian material indicates that the Felidae is the most diverse family there, with a minimum of seven species, from the size of a domestic cat (Felis sp., discussed earlier) to a size larger than a tiger (Al1lphi11laciJairodus kabil). The diversity and richness of fossil Felidae in Africa could therefore substantially increase in the next few years. Family OTARIIDAE Gray, 1825 Table Genus ARCTOCEPHALUS Geoffroy and Cuvier, 1826 Diagnosis Modified from Berta and Wyss (1994); Nowak (2003). Otariid characterized by facial angle of the skull always greater than 125, P3 and Ml single rooted, and calcaneal secondary shelf present. Differs from CalloriJinlls in having tibia and fibula fused proximally; differs from Otariinae (i.e., all other extant genera except Callorhinus) in retaining an 13 incisiform and oval in cross-section. African Species A. pllsillus (Schreber, 1775). Age yrs BP-Recent; Geographic Occurrence South Africa. Remarks The Cape fur seal is known from large samples from several coastal archeological sites in South Africa. 646 LAURASIATHERIA

45 TABLE 32.II Occurrences of pinniped species '2;; :::: 0 EO -< 0 S Z co.a 'U u :::: Sites Q., co ;:: -< t;..!:: B u.8 i::: 0 B t.i. ::::.:;; co t.i J:l ::l...;..!::..!:: Q) :::: 1:: t.i u t.i "0 :::: t.i u.s.s.;:: :::: co 0 S g Q) u i::: 0 P., oj t.i t.i S t.i -< u u u.s B ::l..!:: is..!:: 8..c:..!:: tj.s.s :::: u 0 u 't.i ;:: t.i..!:: oj 0 0 co S :2 :::: c :r; 0 0 >-1 " z:: Ahl al Oughlam Die Kelders 1 Duinefontein 2 Klasies River Mouth Main Site Langebaanweg Baard's Quarry Langepaanweg PPM Langebaanweg indet. Member Raz-el-Aln Ryskop Sahabi Sea Harvest Site Thomas Quarry 1 level G Wadi Natrun? aff. Family ODOBENIDAE Allen, 1880 Table Genus ONTO CETUS Leidy, 1859 Diagnosis Modified from Kohno and Ray (2008). Odobenine odobenid distinguished from other genera by having tusklike canine with thin cementum and orthodentine layers and trefoiled incisive foramina. Differs from AivllkliS and Protodobenlls by having tusklike upper canine with welldefined core of globular osteodentine. Differs from Valenictlls and Odobenlls by retention of the tusk-like upper canine with strong curvature, taper, lateral compression, and longitudinal surface fluting; two upper incisors not in line with the cheek teeth; and two well-developed lower incisors. Functional dental formula 12-3/2, C 1/1, P 4/4, M 1-0/1-0. African Species O. emmonsi Leidy, Age Ca. 2.5 Ma. Geographic Occurrence Morocco. Remarks The African material of this species was described by Geraads (1997) as Alachtherilll1J africa11l1111. It was recently synonymized with the Plio-Pleistocene O. emmollsi by Kohno and Ray (2008). The Ahl al Oughlam material represents the southernmost record of a walrus in the Old World. Family PHOCIDAE Gray, 1821 Table Genus HOlvIIPHOCA Muizon and Hendey, 1980 Diagnosis Modified after Muizon and Hendey (1980). A monachine phocid. Dental formula: I 2/2, C 1/1, P 4/4, M 1/1. Differs from Monac/llls in having a relatively large rostrum, wide posteriorly and narrow anteriorly. The premaxillae terminate against the nasals and have prominent anterior tuberosities. The coronoid process is relatively high as in Lobodontini and not strongly medially recurved as in Monac/llls. Premolars similar to Monac/llls but differ in being lower crowned, relatively narrower, and having a pronounced posterolingual expansion of the cingulum. Small but distinct premolar accessory cusps. M1 distinct in having strongly recurved, sharp principal cusp. M1 largest of cheek teeth with posteriorly slanted principal cusp. Interorbital region broad, tapering posteriorly as in Lobodon but unlike all other monachines. Tympanic bulla covers petrosal; mastoid forms a lip overlapping the posterior border of the bulla. Humerus lacking entepicondylar foramen; tibia and fibula anteriorly fused. African Species H. capensis (Hendey and Repenning, 1972). Age Ca Ma. Geographic Occurrence South Africa. Remarks Muizon and Hendey (1980) consider this species to be related to LobodolJ, taking a morphologically intermediate position between that genus and MOl1ac/llIs. The Langebaanweg material, which is extensive, is the only sub-saharan record of a pre-middle Pleistocene pinniped. Genus LOBODON Gray, 1844 Diagnosis Modified after Bininda-Emonds and Russell (1996). Characterized by the following features: anterior nasal bones trident shaped, with lateral prongs shorter than the median prong; size of pre orbital process of maxilla small; medium or great degree of invagination of frontal (posterodorsal) edge of widened maxillofrontal suture; tendency to reduce the orbitosphenoid; least interorbital width located distinctly posterior to middle of interorbital region; THIRTY-TWO: CARNIVORA 647

46 greatest zygomatic width at level with glenoid fossa (i.e., at squamosal); weak degree of interlock between jugal and dorsal process of squamosal process of zygomatic arch; outline of palatine bones "butterfly-shaped" in ventral view; shape of posterior edge of palatine roughly triangular; medial portion of caudal entotympanic inflated; posterior opening of carotid canal directed roughly 45 medially; mastoid lip in region of external cochlear foramen present; dorsal region of petrosal expanded; intermediate size and shape of petrosal apex; incisors caniniform; outermost lower incisor about equal in size to remaining incisors; postcanine teeth multicuspate; size of accessory cusps in postcanines large, distinct from major cusp; first postcanines smaller than rest, which are subequal; slanted (relative to vertical) implantation of postcanines; lingual face of mandible concave at middle postcanines; distinct medially directed flange present along ventral edge of jaw located posterior to mandibular symphysis and ventral to posterior postcanines; gluteal fossa on ilium shallow; no curvature of pelvis around long axis (Le., pelvis straight); ischiatic spine located in posterior post acetabular region; medium depth of trochanteric fossa on femur. African Species L. carcinopijaga (Hombron and Jacquinot, 1842). Age < yrs BP-Recent (occasional). Geographic Occurrence South Africa. Remarks A single specimen of crabeater seal has been reported from the late Pleistocene Sea Harvest Site, South Africa. Genus MESSIPHOCA Muizon, 1981 Diagnosis Translated and modified from Muizon (1981). Monachinae with greatly developed ulna olecranon and radial facet anteriorly oriented. Differs from Monaclllls and PliopllOca in the curvature of the ulna diaphysis and in the shape of the articular facet with the pyramidal. The radius and ulna are greatly flattened. The humerus lacks an entepicondylar foramen. African Species M. mauretal1ica Muizon, Age Ca. 7-6 Ma. Geographic Occurrence Algeria. Remarks This poorly known taxon from Raz-el-Aln, Algeria, is of particular interest for the light it may shed on the state of the Mediterranean during the Messinian salinity crisis (Muizon, 1981). Genus MIROUNGA Gray, 1827 Diagnosis Modified after Bininda-Emonds and Russell (1996). Basal monachine phocid characterized by: nasal processes of maxilla extending along maxilla only part way of nasals; nasal bones not trident shaped; presence of a deep fossa on the ventrolateral side of premaxilla; preorbital process of maxilla large; incisive foramina absent; pterygoid hamuli directed medially; basioccipital-basisphenoid region concave; hypomastoid fossa absent; median lacerate foramen in the auditory bulla absent; stylomastoid and auricular foramina confluent; parietal contributing to the bony falx; one lower incisor per hemimandible; upper incisors rounded in cross section; postcanine teeth peglike (unicuspate); first and fifth lower postcanine teeth noticeably smaller and first and fifth upper postcanine teeth noticeably larger than the other postcanines, which are subequal; tendency to single-rooting of lower postcanines; reversely arched upper postcanine tooth row; lingual face of mandible at middle postcanines concave; scapular spine of medium size; gluteal fossa on ilium shallow; no distinct trochanteric fossa on femur; claw semicircular in cross section. African Species M. leonina (Linnaeus, 1758), M. sp. Age Ca. 0.1 Ma-Recent (possibly present as early as 18 Ma). Geographic Occurrence South Africa. Remarks Pickford and Senut (2000) refer remains from Ryskop in Namibia to?mirozmga sp. These remains are by far the oldest pinniped remains in Africa. Remains of the extant southern elephant seal have been found at the middle Pleistocene archeological sites of Die Kelders and Klasies River Mouth in South Africa. Genus MONACHUS Fleming, 1822 Diagnosis Modified after Bininda-Emonds and Russell (1996). Derived monachine phocid characterized by: Broad contact between nasal processes of premaxilla and nasals; fossa on ventrolateral side of premaxilla shallow; interorbital septum anterior to optic foramina present; incisive foramina of medium size; foramen ovale entirely in the squamosal; medial portion of caudal entotympanic not inflated; petrosal not covered by the auditory bulla; hypomastoid fossa deep; external cochlear foramen open; petrosal apex unexpanded and pointed; incisors not procumbent; no tendency to form or lose additional cusps in triconodont postcanine teeth; accessory cusps in postcanine teeth small; postcanine teeth touching or overlapping; anterior/posterior end of postcanine teeth directed laterally; upper postcanine tooth row kinked between postcanine 1 and 2, otherwise straight; gluteal fossa on the wing of ilium absent; no ridges in anterior portion of obturator foralnen. African Species M. l710nacijus (Hermann, 1779). Age Ca yrs BP-Recent? Geographic Occurrence Morocco. Remarks There is a single record of Mediterranean monk seal from the Thomas Quarry Level G, Morocco. Genus PLIOPHOCA Tavani, 1941 Diagnosis Modified from Koretski and Ray (2008). Medium-sized phocid with relatively broad and robust teeth; P1 and pi single rooted, the other postcanine teeth double rooted; in the maxilla, anterior alveoli smaller than posterior alveoli, and slightly oblique to the labial side, while the posterior alveoli are slightly oblique to the lingual side; the opposite is true in the dentary; retromolar space elongated, mandibular body not high, ramus very strong and high; p2-m1 five cusped, with basal cingula very well developed; alveoli of m1, M1 shorter than those of p4, P4; diastemata present in maxilla, absent in the mandible; proximal part of deltoid crest of humerus located higher than the head, but lower than the minor tubercle; deltoid crest strongly developed, terminates slightly lower than the middle of diaphysis; greater breadth of deltoid crest located on its middle part; greater trochanter of femur rectangular, placed at the same level as head; trochanteric fossa shallow and caudally elongated; intertrochanteric crest not developed; least width of femur diaphysis is at the middle of the bone; femoral condyles placed wide apart and flattened, with greatest distance between then being 0.15 of the bone length; distal end of femur wider than proximal end by African Species P. etrusca Tavani, Age Ca. 6-4 Ma. 648 LAURASIATHERIA

47 Geographic Occurrence Egypt. Remarks The single African record of this species is based on a fragmentary mandible with one postcanine tooth and an indeterminate deciduous tooth from Wadi Natrun, Egypt (Stromer, 1907, 1913). The species was first assigned to Pristiphoca sp. aff. P. occital1a by Stromer (1913). According Koretski and Ray (2008), however, this species is based on a canine tooth of an indeterminate carnivore, which is presumably lost. Pending rediscovery of the holotype and availability of associated material, these authors regard the species as a nomen nudum. They also consider that the fragmentary right hemimandible with two postcanines referred to as the same species by Gervais (1853) cannot be confidently assigned to the same species and assign it to Pliophoca etmsca. Following Howell and Garcia (2007), we then assign the species from Wadi Natrun to PliopllOca etmsca. MONACHINAE indet. Age Ca. 7-6 Ma. Geographic Occurrence Algeria, Libya. Remarks Muizon (1981) and Howell (1987) record additional monachines from Raz-el-Aln and Sahabi, respectively, although the material is too fragmentary to allow for a more precise diagnosis. Biogeography Two uncertain occurrences may attest to a pre-miocene presence of the order Carnivora in Africa. A Carnivora indet. is represented by several isolated premolars from the Ouarzazate Basin (Paleocene of Morocco; Gheerbrant, 1995), while the second Paleogene record, Glibzegdollia tabelbalael1sis, is based on an isolated ml from Glib Zegdou (Eocene of Algeria; Crochet et al, 2001). In both cases, the ordinal assignment is uncertain. Lower molars of recently described proviverrine creodonts from the early late Paleocene and early Eocene of Morocco (Gheerbrant et al, 2006) show some resemblance to those of G. tabelbalaensis. More precise comparisons of these teeth may therefore indicate that these putative carnivorans belong to the Creodonta. Like many nonafrotherian mammalian orders, the presence of carnivorans in Africa is correlated with the collision between the Afroarabian and Eurasian plates near the Oligocene-Miocene boundary. Unambigous remains of Carnivora in Africa are from early Miocene strata in Kenya, Uganda, Congo, and Namibia (but see Note added in proof). Soon after their arrival, the Carnivora successfully and rapidly expanded across the entire continent. The quality of the fossil record is very heterogeneous in Africa, which renders the distribution of Neogene carnivorans difficult to interpret. The Neogene fossil record of carnivorans is at best extremely poor in western Africa (all countries south of Mauritania, Mali, Niger), central and westcentral Africa (Central African Republic, Sudan, Cameroon, Gabon, CongO-Brazzaville, Democratic Republic of Congo), and some southern countries (Angola, Botswana, Mozambique, Zambia). Some regions in Africa have, in contrast, yielded extremely rich faunas for certain times and/or have a good and continuous fossil record for a long time. Eastern Africa is the best "hot spot" for Neogene paleomammalogy in Africa, and in particular for the Plio-Pleistocene (Werdelin and Lewis, 2005); southern Africa also has rich and diversified Plio-Pleistocene faunas but not as many Miocene mammal sites as eastern Africa. This does not imply that carnivorans were absent from southernmost Africa, since they are diverse and numerous, from, for example, the early-middle Miocene of Namibia. With the increasing amount and intensity of field research in Africa, some regions or countries are showing their scientific potential, demonstrating that eastern African dominance may not only be due to its unique geographic and tectonic position, which created exceptional conditions of fossilization, but also to the near absence of a fossil record in most other African regions. One of these emerging regions is northern Chad, the fossiliferous sites of which to date have yielded thousands of Mio-Pliocene fossils. Over 30 years ago, Savage (1978) concluded his contribution to Maglio and Cooke's Evolution of African Mammals about creodonts and carnivorans thus: "This account of origins and migrations is unhappily more conjecture than fact. Only many more facts will substantially improve it. Pliny wrote nearly 2,000 years ago, 'Ex Africa semper aliquid novi': it is still true today." Although Pliny's statement will be true in paleontology for many years to come, 30 years after this confession, the late R. J. G. Savage would certainly acknowledge that the paleontology of carnivores in Africa has been much improved, especially in the last decade, thanks to a much better, and still improving, fossil record. A comparison between the state of the art of Savage (1978) and the present account shows this (see table 32.12). In particular, the total number of extinct genera and occurrences is much greater. All the families are involved, but the Ursidae (Savage did not really provide a synthesis of the numerous Quaternary records), Hyaenidae, and Mustelidae (in particular the Lutrinae) display a tremendous improvement in their fossil record (number of species with fossil record and occurrences). Hunt (1996) provided a detailed analysis of the biogeography of each family incorporating the fossil record, while Werdelin and Lewis (2005) discussed the fossil record of Plio Pleistocene carnivorans in Africa. Therefore, we here summarize the main biogeographic trends of each family, especially for the Miocene. Many recent papers have described new or unpublished material from this epoch that has revealed a much greater richness and diversity than previously known. Significant examples include, for example, Arrisdrift (eight species; Morales et al., 1998, 2003), Ngorora Fm. (seven species; Morales and Pickford, 2005a), Lukeino and Mabaget Fms (19 species; Morales et al, 2005), Nawata Fm. (18 species; Werdelin, 2003), Adu-Asa and lower Sagantole Fms (at least 14 species; Haile-Selassie and Howell, 2009), and Lemudong'o (12 species; Howell and Garcia, 2007). In Africa, there are no fossils assigned to the Miacidae, Viverravidae, Nimravidae, Procyonidae, or Ailuridae (with the Simocyoll sp. of Howell and Garcia [2007] reassigned to Amphicyonidae). Despite the lack of a fossil record in many African countries, these families of carnivores, the three former being typical of the Paleogene, apparently never reached Africa. More surprising is the apparent absence of a fossil record for the Eupleridae (including all the Malagasy carnivorans), although they had an African ancestry with a divergence time extending back to the early Miocene (24-18 Ma; Yoder et al, 2003). The extant carnivore fauna of Africa is composed of five feliform (Felidae, Hyaenidae, Viverridae, Nandiniidae, Herpestidae) and two caniform (Canidae, Mustelidae) families, including about 80 species (Sunquist and Sunquist, 2002; Gaubert et al., 2004; Sillero-Zubiri et al., 2004; Nowak, 2005). In addition, Amphicyonidae, THIRTY-TWO: CARNIVORA 649

48 TABLE Extant and fossil records as reported by Savage and this study The table only reports the extant and fossil records for Africa. Savage listed the known total number of extinct and living genera for all Carnivora, which is not provided here. Extinct Genera Genera with Fossil Record Living Genera" Total of Genera Species with Fossil Record Occurrences Family Savage This Study Savage This Study Savage This Study Savage This Study Savage This Study Savage This Study Amphicyonidae Canidae Ursidae Mustelidae Nandiniidae Viverridae I-Ierpestidae Percrocutidae I-Iyaenidae Barbourofelidae Felidae Otariidae Odobenidae Phocidae Total ,009 "Living genera include the genera currently present in Africa, with or without a fossil record. Differences observed between our study and Savage's in the living genera are due to the classification used. See appendix for further explication.

49 Barbourofelidae, and Ursidae, the former two wholly extinct, have fossil records in Africa. Except for Nandiniidae, the fossil record in Africa for each extant family is continuous or nearly continuous since its first occurrence. The Barbourofelidae and Amphicyonidae are common but not abundant members of predator communities of the early and middle Miocene of Africa. Amphicyonids in particular are diverse in eastern Africa, where the fossil record of the family is more or less continuous from the early to the late Miocene. They are also found in Mediterranean countries (Morocco, Tunisia, Algeria, Libya, Egypt) and southern Africa (Namibia and South Africa). The extinction of the Amphicyonidae (ca. 5.5 Ma) is slightly later than the arrival of large ursines (Indarctos and Agriotl1erium) and canids. No known locality documents the sympatry of these three families, however. Some sites have either Ursidae and Canidae (e.g., Lukeino) or Amphicyonidae and Canidae (e.g., Toros-Menalla, Lemudong'o), but there is no known late Miocene locality documenting a potential sympatry of Amphicyonidae and Ursidae in Africa. Evidence supporting competitive exclusion is lacking, but competition for large prey was possible between large amphicyonids (Agl1oti1erill11l) and large ursids (Agriotiz e rill 111). Early African canids (late Miocene-early Pliocene) were much smaller species and not able to compete with any of the contemporary amphicyonids or ursids (though competition with the small amphicyonid BOl1isicYOI1 may have occurred). The last Barbourofelidae are probably Vampyrictis vipera from Bled Douarah and the Barbourofelidae indet. from Namurungule. Both have reached the evolutionary grade of Sal1sm1Osmillls. Felids, and in particular machairodontines, were their main competitors. The extinction of this already rare family in the late Miocene of Africa may be a result of the arrival of Machairodontinae on the continent. Remains of Ursidae are rare but they had a pan-african distribution (from Libya to South Africa). A single tooth from Rusinga (early Miocene, Kenya) indicates an early expansion of this originally Holarctic family southwards. However, this record is separated from the next oldest record of the family (Irzdarctos from Menacer, Algeria) by a lo-million-year hiatus. Urslls, the only extant genus with a fossil record in Africa was restricted to northwestern Africa but is now extinct there (Servheen et a!., 1998). Mustelids are among the first carnivoran immigrants to Africa, with otterlike forms from the early Miocene of East Africa. The radiation of otters during the late middle and late Miocene is one of the most significant traits of the family in Africa. Lutrines are by far the most common mustelids, with about two-thirds of the total occurrences of the family. Also worth noting is the great size range of the species, spanning from the size of the living river otter, Llltra llltra, to a much greater size than any living otter (»100 kg). These giant otters were distributed from Egypt to South Africa, including a great diversity in eastern Africa. The Canidae are exclusive to North America until the late Miocene, after which EllCYOrz- and Vlllpes-like taxa appear in western Europe, Asia, and Africa. Our knowledge of the Canidae in Africa is still limited though rapidly improving. Recently, Morales et a!. (2005) described from Lukeino (ca. 6 Ma) what was then the oldest canid of Africa, EllCYOI1 il1trepidlls. Even more recently, however, a foxlike canid has been discovered from older sediments in Chad (Toros-Menalla, ca. 7 Ma; Bonis et a!., 2007b). The family reaches southern Africa in the early Pliocene and northwestern Africa in the mid-pliocene. The final radiation of the Canidae (Caninae) in the Pleistocene is seen mainly in eastern Africa, but also in South Africa and Algeria. The Viverridae and the Herpestidae are among the first immigrants to Africa during the early Miocene and are first known from the same eastern African localities (e.g., Koru, Napak, Rusinga, Songhor). Viverrids are found in northern (Egypt) and southern (Namibia) parts of the continent soon after, but remain rare or absent during the middle Miocene, with a few records in Kenya (Fort Ternan, Ngorora), Morocco (Beni Mellal), and Namibia (Arrisdrift), and are only slightly more abundant during the late Miocene in Ethiopia (Chorora, Adu-Asa), Kenya (Lothagam, Lukeino), and Chad (Toros-Menalla). After the early Miocene records, herpestids are nearly absent in Africa until the end of the Miocene. Solitary mongooses (Galerella and Iclmellll1ia) are first found in the late Miocene of Chad (only record for central Africa) and Kenya. Social mongooses (Helogale) appear in Africa in the latest Miocene of Ethiopia (Adu Asa Fm.). Plio-Pleistocene taxa show greater diversity than today and are known mainly from eastern (Ethiopia, Kenya, Tanzania) and South Africa. In addition to these areas, modern mongooses are known only from a single northern African site (mid-pliocene, Morocco). The Hyaenidae has a particularly rich and diverse fossil record in Africa, which strongly contrasts with the present low diversity (four extant species). In contrast to many families, the oldest hyaenids in Africa are from northern Africa, with Hyael1ictis?gmeca, Protictitl1eriwl1 plll1icum, P. CraSSlll1l, and Lycyaena crllsafol1ti, all from roughly contemporaneous, late middle-early late Miocene localities of Morocco (Beni Mellal) and Tunisia (Bled Douarah). Protictitl1erillln expands its range southwards to Kenya (Namurungule). A hyaenid fauna composed of Chasmaporti1etes, Ikeloizyael1a, Hyael1ictis, Hyael1ictitl1eri1l111, Ietitizeriwll, and AdcTOCllta, of which the latter never reaches sub-saharan Africa, characterizes the late Miocene-earliest Pliocene. Except for Clwsmaporthetes and Ikelol1yaena, all these genera become extinct ca. 5 my a ago, while bone-cracking species of Croellta, P/ioeroCllta, PacJlyero Cllta, Pamizyaena, and H)Jael1a appear soon after this date. The Percrocutidae is represented in Africa by two genera, DinocroCllta and Percroellta. Many records are late middle Miocene in age, with the oldest being from Kenya (PercroClita sp. from Fort Ternan and P. tobiel1i from Ngorora). Slightly younger records of PereroCllta are from Tunisia (Bled Douarah) and Kenya (Nakali, Namurungule). Dil1oeroClita is slightly younger and is found at early late Miocene localities of Algeria and Kenya. The last records of the family are from the latest Miocene. The Felidae was a very diverse family, with at least 15 genera and 35 taxa. Including undetermined material would certainly raise the total number of felid species to over 40. The Felidae first appears in Africa during the early Miocene. The family therefore has a long history on this continent though the distribution is, as for other families, very heterogeneous. About 90% of the occurences are in eastern or southern Africa. The subfamily Machairodontinae, which includes the sabertooth taxa, first appears in the late middle Miocene and becomes extinct during the middle Pleistocene. For more than 10 million years, sabertooth cats were diverse (they account for about half of the total number of African species of Felidae) and significant members of the large predator guild. They had their greatest THIRTY-TWO: CARNIVORA 651

50 diversity during the late Miocene-early Pliocene with about 10 species, most of which were large-sized taxa. Extant genera and species have a fossil record restricted to the Pliocene (after ca. 4.3 Ma). Migration Patterns As noted, the Carnivora arrives in Africa in the Lower Miocene or slightly earlier, which is quite late in the history of the order. Clearly, their arrival in Africa was due to a migration event, from either Europe or Asia, or both. Throughout the history of Carnivora in Africa, there has been a series of such events, both into and out of Africa. Although the fossil record biases the data, a number of these events are distinct and broadly associated with tectonic and climatic trends of the Neogene The first such event occurred some 20 Ma ago. It involved members of the families Amphicyonidae, Ursidae, Felidae, Musteiidae, Herpestidae, Viverridae, and Barbourofelidae. The latter two families may have migrated from Asia via Africa to Europe, while the other five most likely are immigrants from Europe. There is no evidence for size sorting during this event, as it involves both very small (e.g., Herpestidae) and very large (e.g., Amphicyonidae) taxa. This event is broadly correlative with the first land passage between Africa and Eurasia, the so-called G0l11photheriu111 land bridge (Ragl, 1998), a result of the isolation of the Indo-Pacific from the Paratethys during the later part of sea level cycle TB2.1 (Haq et ai., 1987, 1988). The next event occurs prior to 14 Ma and involves immigration of the family Percrocutidae, the subfamily Mustelinae (earlier mustelids are of uncertain subfamily, but possibly Lutrinae), the amphicyonid genus Agnotherium, and the lutrine VishIllLOI1Y, the latter of which almost certainly arrived from the Indian subcontinent. The paleogeography of this time is very similar to the time of the Gomphotherium land bridge, with the Indo-Pacific and Paratethys once again isolated from one another during sea level cycle TB2.4, which is dated ca Ma (Berggren et ai., 1995). Again, no clear size sorting is discernible during this event. The third event occurred prior to 10 Ma and involved the family Hyaenidae and the genus Machairodlls, with the latter appearing nearly simultaneously in Eurasia and Africa. This event seems to have involved the immigration to Africa of mainly medium- to large-sized species and is associated with sea-level cycles TB2.6 (ca Ma) and TB3.1 (ca Ma). It should be noted that as presently reconstructed, there is no direct land connection between Africa and Europe at this time, and exchange between these continents may have occurred via an easterly route around the Paratethys. The next event occurred prior to ca 7.5 Ma and involves a series of genus-level taxa: the ursids 1I1darctos and Agriotherilll11, the mustelids Piesioglilo and Sivaol1)'x, the hyaenids AdcroCllta, Chasmaporthetes, and Hyaenictis, and the felid Metailurus. All these are relatively large taxa, indicating significant size sorting. Agrioti1eri1l111 and Sivaonyx appear more or less simultaneously in Africa and Eurasia. This event may also, if recent data are accurate, involve the family Canidae as the last carnivoran family to migrate into Africa. It is broadly correlative with sea-level cycle TB3.3, ca Ma. The Pliocene also saw a series of carnivoran migration events, but this time mainly out of Africa. The first such event occurred ca Ma and involved migration into Africa of the canid genus Nycterelltes and migration out of Africa of the felid genera Megmztereon and H0111oti1eriWll. All these are medium- to large-sized taxa. The event may be correlated with sea level cycle TB3.4, ca Ma, but it is more likely that at this point migrations were climatically rather than tectonically mediated, as the paleogeography changes very little after the middle Upper Miocene. An additional set of genera of medium- to large sized carnivorans migrates out of Africa ca. 3.5 Ma: the canid Canis, the hyaenid PachycroCllta, and the felid Acinonyx. A final event occurs before ca. 2 Ma and involves migration out of Africa of the hyaenid genera Hyaena and CroCZlta and the felid genus Panti1era. At this time the mustelid genus Aonyx may have migrated to Africa, but this is very uncertain. Except Aonyx these are all large-sized taxa. In summary, carnivoran migrations in the Miocene appear to be mainly into Africa and to be tectonically controlled. The earliest migrations show no size sorting, but gradually migrations appear to involve mostly medium- to large-sized taxa. In the Pliocene, migrations are mainly out of Africa and are climatically controlled. They involve mostly medium- to large-sized taxa. Conclusions Despite their late appearance, the Carnivora have diversified greatly on the African continent, as shown in the present compilation. At the same time, the geographic and temporal biases in the material show just how much remains to be discovered. The Plio-Pleistocene, up to about one million years ago, is reasonably well-known for some African regions, but the origin of the extant fauna is obscure (Werdelin and Lewis, 2005). The latest Miocene has seen a tremendous upswing in interest and discoveries in the last few years (Werdelin, 2003; Haile-Selassie et ai., 2004b; Morales et ai., 2005; Howell and Garcia, 2007), and we hope that in the next decade a similar upswing will occur for the middle Miocene, which is very poorly known. One aspect of carnivoran evolution in Africa that is becoming evident as new discoveries accumulate is the importance of Africa to the evolution of carnivorans elsewhere. Though we would perhaps not go quite as far as Pickford (2004), it is becoming clear that important Plio Pleistocene carnivoran taxa such as Canis, Homotherizll1z, Acinonyx, Panthera, and others may have had an African origin, or at least that Africa was important to their gobal diversification. This makes the factors allowing intermittent migration of mammals out from Africa of critical importance for understanding the origin and composition of modern carnivoran guilds in the Old World and in some cases North America. Carnivorans are also important in the evolutionary history of our own genus (Turner, 1984), and hence understanding carnivoran evolution will help in understanding human evolution as well. ACKNOWLEDGMENTS We would like to thank all those many people who collected, curated, and initially described the specimens on which the taxa incorporated in this chapter are based. vve would also like to thank many colleagues for fruitful discussions over the years about carnivores and African paleontology. L.W. would like to thank Susanne Cote, Meave Leakey, Margaret Lewis, Jorge Morales, and Martin Pickford for help that significantly improved the chapter, and the Swedish Research Council for a series of grants that made the work possible. S.P. would like to thank the Swedish Museum of Natural History 652 LAURASIATHERIA

51 and financial support through the Synthesys project (SE-TAF 1380), which was made available by the European Community's Research Infrastructure Action under the FP6 "Structuring the European Research Area" Program. APPENDI This appendix contains details of the analysis presented in table For Canidae, we include Fenneells in Vllipes and Lymon in Canis (see Werdelin and Lewis, 2005); for Ursidae, we consider Urslls as extinct in Africa; for Mustelidae, we consider, following current use, Paraonyx as used by Savage a junior synonym of Aon)!x and Poecilietis to be a junior synonym of Icton)!x; Viverridae in Savage (1978) are now separated into four families (Viverridae, Herpestidae, Nandiniidae, and Eupleridae); Stenoplesictidae is not regarded as valid here. The differences from Savage concern the Herpestidae, in which we recognize Galerella, Paraeynietis, Liberiictis, and Dologale as valid genera and enogale as a junior synonym of Helpestes, and the African Viverridae, in which we follow current usage by considering Osbomietis a junior synonym of Genetta. Differences among the Hyaenidae are due to the recognition of Parahyaena as distinct from Hyaena and the recognition of the Percrocutidae as a distinct family. For Felidae, we recognize Barbourofelidae as a distinct family. For extant genera, we follow recent systematists and recognize Caraeal, Leptailll1'llS, and Protelis as valid genera distinct from Felis; species number may be difficult to establish because of the many indeterminate records. Like Savage (1978), we attempt here to include some of them. Species numbers are therefore a lowest estimate that will increase when better material is found to document indeterminate or poorly known taxa. We report evidence that supports the recognition of new taxa in addition to the named species. For Amphicyonidae, we have 10 species and consider that Amphicyonidae indet. (including sp. A) includes at least one additional species. For Ursidae, in addition to the five named species, there are at least two additional species: (1) a new ursine to be described from East Africa; (2) the hemicyonine species from Rusinga. For Canidae, in addition to the 20 named species, we have added two species, considering (1) that Callis sp. includes at least one new species (from Turkwel and possibly Laetoli); other, more recent mentions may be referred to either of the previously described species; (2) that Canidae indet. from the Mursi Formation probably represents a species new for the continent. For Mustelidae, in addition to the 32 named species, we consider that Mustelidae indet. includes at least one additional species. For Viverridae, some undetermined mentions certainly represent new taxa (especially from Koobi Fora). The exact number is not possible to establish, but we consider that Viverridae indet. comprises at least two additional species. For Herpestidae, we have included only the named species. For Percrocutidae, we did not consider PercroCllta from Namurungule Fm. as a distinct species. For Hyaenidae, in addition to the 27 named species, we have considered as distinct species (1) in Hyaenidae indet., Hyaenidae indet. E from Langebaanweg; and (2) that at least one record of Proteles sp. may correspond to the extant species Proteles cristatlls. For Barbourofelidae, we did not recognize Barbourofelidae indet. from Namurungule as a distinct species because of a possible identity with Vmnp)'rietis of the same age. For Felidae, in addition to the 27 named species (iviegalltereon ekidoit/whitei does not correspond to a species but represents uncertainty of assignment), two species have been added, considering that (1) Felis sp. certainly includes at least one species distinct from Felis silvestris I)'bim and (2) that Felidae indet., especially the record from Songhor, certainly includes another one. For pinnipeds, "living genera" include those that are frequently observed along the African coasts that is, 1vIonaclllls monac7111s, Aretoeephaills pllsilllls, and A. tropimlis (Nowak, 2003). In addition to the six named species, we have considered that?miroll1jga sp. from Ryskop certainly represents a species distinct from M. leonina. NOTE ADDED IN PROOF Rasmussen and Gutierrez (2009) recently described the carnivoran Mioprionodon llodopells from the late Oligocene of Nakwai, northwestern Kenya. This represents the first record of the Carnivora in Africa and suggests dispersal opportunities from Eurasia earlier than previously thought (see also Morlo et al. (2007»). Literature Cited Adam, P. J Lobodon carcil1opilaga. Mammalian Species 772:1-14. Aguirre, E., and P. Leakey Nakali: Nueva fauna del Hipparion de Rift valley de Kenya. Estlldios Geol6gicos 30: Anderson, E Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta Zoologica Fenl1ica l30:1-l32. Ant6n, M., M. J. Salesa, ]. Morales, and A. Turner First known complete skulls of the scimitar-toothed cat Maclwirodlls aplwnistlls (Felidae, Carnivora) from the Spanish Late Miocene site of Batallones: 1.1ollnJai of Vertebrate Paleontology 24: Aouraghe, H Les carnivores fossiles d'el Harhoura 1, Temara, Maroc. L'Antilropologie 104: Arambourg, C Note preiiminaire sur quelques Vertebres nouveaux du Burdigalien de Libye. Compte Rendll s011lmaire des seances de la Societe Geologiqlle de France 1961(4): Barry,]. C Large carnivores (Canidae, Hyaenidae, Felidae) from Laetoli; pp in M. D. Leakey and J. M. Harris (eds.), Laetoli: A Pliocene Site in Northern Tanzania. Clarendon Press, Oxford. Beaumont, G. de Observations sur les Herpestinae (Viverridae, Carnivora) de l'oligocene superieur avec quelques remarques sur des Hyaenidae du Neogene. Archives des Sciences, Geneve 20: Berggren, W. A., D. V. Kent, C. C. Swisher, III, and M.-P. Aubry A revised Cenozoic geochronology and chronostratigraphy; pp in W. A. Berggren, D. V. Kent, M.-P. Aubrey, and J. Hardenbol (eds.), Geochrollology, Ti11le Scales alld Global Stratigraphic Correlation. SEPM Special Publication 54. Berta, A., and A. R. Wyss Pinniped phylogeny. Proceedings of tile San Diego Society ofnatllral Histor)' 29: Bininda-Emonds, O. R. P., and A. P. Russell A morphological perspective on the phylogenetic relationships of the extant phocid seals (Mammalia: Carnivora: Phocidae). Bonner Zoologische Monograplzien 41: Bonis, L. de, S. Peigne, A. Likius, H. T. Mackaye, M. Brunet, and P. Vignaud. 2007a. First occurence of the "hunting hyaena" Clwsl1laportlzetes in the Late Miocene fossil bearing localities of Toros-Menalla, Chad (Africa). Blllletin de la Societe Geologique de Frallce 178: Bonis, L. de, S. Peigne, F. Guy, A. Likius, H. T. Mackaye, P. Vignaud, and M. Brunet A new mellivorine (Carnivora, l'viustelidae) from the late Miocene of Toros Menalla, Chad. Nelles Tahrbllc/l fiir Geologie wld Paliiolltologie 252: Bonis L. de, S. Peigne, A. Likius, H. T. Mackaye, P. Vignaud, and M. Brunet. 2007b. The oldest African fox (\!zilpes riffalltae n. sp., Canidae, Carnivora) recovered in late Miocene deposits of the Ojurab desert, Chad. Natllnvissensc/zaften 94: Bonis, L. de, S. Peigne, H. T. Mackaye, A. Likius, P. Vignaud, and M. Brunet The fossil vertebrate locality Kossom Bougoudi, Ojurab desert Chad: A window in the distribution of the carnivoran faunas at the Mio-Pliocene boundary in Africa. COl1lptes Rendlls PalevoI7: Bryant, H. N Delayed eruption of the deciduous upper canine in the sabertoothed carnivore Barbollrofelis lovei (Carnivora, Nimravidae). Tournai of Vertebrate Paleontolog), 8: THIRTY-TWO: CARNIVORA 653

52 Bryant, H. N., A. P. Russell, and W. D. Fitch Phylogenetic relationships within the extant Mustelidae (Carnivora): Appraisal of the cladistic status of the Simpsonian subfamilies. Zoological Touroal of the Linman Society 108: Colbert, E. H Carnivora of the Tung Gur Formation of Mongolia. Bulletin of the American IvIlIsellin of Nat lim I History 76: Cooke, H. B. S Ictonyx bolti, a new mustelid from cave breccias at BoIt's Farm, Sterkfontein area, South Africa. SOlltil African TOllmal ofsci llce 81: Cote, S., L. WerdeIin, E. R. Seiffert, and J. C. Barry Additional material of the enigmatic early Miocene mammal Kelba and its relationship to the order Ptolemaiida. Proceedings of tlze National Academy of Sciences, USA 104: Crochet, ].-Y., S. Peigne, and!vi. Mahboubi Anciennete des Carnivora (Mammalia) en Afrique; pp in C. Denys, L. Grangeon, and A. Poulet (eds.), Proceedings of the Stil Intemational Symposium on African Small Mammals, Paris, 4-9 TlIly IRD Editions, Collection Colloques et Seminaires, Orleans. Dehghani, R., and L. Werdelin A new small carnivoran from the Middle Miocene of Fort Ternan, Kenya. Nelles Tallrbllc1l fiir Geologie liild Pa/dontologie 248: Ewer, R. F. 1954a. The fossil carnivores of the Transvaal caves: The Hyaenidae of Kromdraai. Proceedings of tile Zoological Society of London 124: b. The fossil carnivores of the Transvaal caves: The lycyaenas of Sterkfontein and Swartkrans, together with some general considerations of the Transvaal fossil hyaenids. Proceedings of tile Zoological Society of London 124: The fossil carnivores of the Transvaal caves: Two new viverrids, together with some general considerations. Proceedings of tile Zoological Society of London 126: FiccarelIi, G., D. Torre, and A. Turner First evidence for a species of racoon dog, Nyctereutes Temminck, 1838, in South African Plio Pleistocene deposits. Bollettino della Societii Paleontologica Italiana 23: Fraile, S., B. Perez, l. de Miguel, and]. Morales Revisi6n de los carnivoros presentes en los yacimientos del Ne6geno espanol; pp in J. P. Calvo and J. Morales (eds.), Avances en el Conocimiento del Terciario Iberico. III Congreso del Grupo Espanol del Terciario, Cuenca. Garcia, N New Eucyon remains from the Pliocene Aramis Member (Sagantole Formation), Middle Awash Valley (Ethiopia). Comptes Rendus Palevol 7: Gaubert, P., C. A. Fernandes, M. W. Bruford, and G. Veron Genets (Carnivora, Viverridae) in Africa: an evolutionary synthesis based on cytochrome b sequences and morphological characters. Biological Toumal oftlze Linnean Society 81: Geraads, D Carnivores du Pliocene terminal de Ahl al Oughlam (Casablanca, Maroc). Geobios 30: Plio-Pleistocene Carnivora of northwestern Africa: a short review. Comptes Rendus Palevol 7: Geraads, D., Z. Alemseged, and H. Bellon The Late Miocene mammalian fauna of Chorora, Awash Basin, Ethiopia: Systematics, biochronology and the.ok.4 0 Ar ages of the associated volcanics. Tertimy Researc11 21: Geraads, D., Z. Alemseged, D. H. Reed, and J. G. Wynn The Pleistocene fauna (other than Primates) from Asbole, lower Awash Valley, Ethiopia, and its environmental and biochronological implications. Geobios 37: Gervais, P Description de quelques ossements de phoques et de cetaces. Memoires de la section des Sciences, Acadelllie des sciences et lettres de Montpellier 2: Gheerbrant, E Les mammiferes paleocenes du Bassin d'ouarzazate (Maroc): III. Adapisoriculidae et autres mammiferes (Carnivora,?Creodonta, Condylarthra,?Ungulata et incertae sedis). Palaeontograpilica, Abt. A 237: Gheerbrant, E., M. larochene, M. Amaghzaz, and B. Bouya Early African hyaenodontid mammals and their bearing on the origin of the Creodonta. Geological Magazine 143: Gilbert, W. H., N. Garcia, and F. C. Howell Carnivora; pp in W. H. Gilbert and B. Asfaw (eds.), Homo erectus: Pleistocene Evidence frolll the Middle Awasll, Ethiopia. University of California Press, Berkeley. Ginsburg, L Les carnivores du Miocene de Beni Mellal (Maroc). Geologie Mediterraneenne 4: Les migrations de mammiferes carnassiers (Creodontes + Carnivores) et Ie probleme des relations intercontinentales entre l'europe et l'afrique au Miocene inferieur. Annales Geologiqlles des Pal's Hellbliqlles H.S. 1979: Ginsburg, L., and J. Morales Les Hemicyoninae (Ursidae, Carnivora, Mammalia) et les formes apparentees du Miocene inferieur et moyen d'europe occidentale. Annales de Paleontologie 84: Ginsburg, L., and J.-L. Welcomme Nouveaux restes de Creodontes et de Carnivores des Bugti (Pakistan). S)'mbioses (n.s.) 7: Goldman, C. A Crossarclllls obscl/l'lls. lviammalian Species 290:1-5. Haile-Selassie, Y., and F. C. Howell Carnivora; pp in Haile-Selassie, Y. and WoldeGabriel, G. (eds.), Ardipithecus kadabba: Late Miocene Evidence ftom the Middle Awasll, Etiliopia. University of California Press, Berkeley. Haile-Selassie, Y., L.]. HIusko, and F. C. Howell. 2004a. A new species of Piesiogllio (Mustelidae: Carnivora) from the Late Miocene of Africa. Palaeontologia Africana 40: Haile-Selassie, Y., G. WoldeGabriel, T. D. White, R. L. Bernor, D. Degusta, P. R. Renne, W. K. Hart, E. Vrba, S. Ambrose, and F. C. Howell. 2004b. Mio-Pliocene mammals from the Middle Awash, Ethiopia. Geobios 37: Hamdine, W., M. Thevenot, and]. Michaux Histoire recente de l'ours brun au Maghreb. Comptes Rendus de lacademie des Sciences, Paris, ScieJJces de la Vie 321: Haq, B. U., J. Hardenbol, and P. R. Vail Chronology of fluctuating sea levels since the Triassic (250 million years ago to Present). Science 235: Mesozoic and Cenozoic chronostratigraphy and cycles of sea level change; pp in C. K. Wilgus, B. J. Hastings, H. Posamentier, J. C. van Wagoner, C. A. Ross, and C. G. S. C. Kendall (eds.), Sea-Level Change: An Integrated Approach. SEPM Special Publication 42. Heizmann, E. P. J., and E. G. Kordikova Zur systematischen Stellung von 'Ampllicyon" intermedius H. v. Meyer, 1849 (Carnivora, Amphicyonidae). Carolinea 58: Helbing, H PselldoC)'on saljsaniensis Lartet von Steinheim am Albuch. Eclogae Geologicae Helvetiae 22: Hendey, Q. B The late CenozoiC Carnivora of the south-western Cape Province. Annals of the SOlltil African Musellm 63: Late Tertiary Mustelidae (Mammalia, Carnivora) from Langebaanweg, South Africa. Annals of the Soutil African Musellll1 76: Hooijer, D.A Miocene Mammalia of Congo. IvIusee Royal de lafriqlle Centrale, TervllrflJ, Belgiqlle, Amwles, Serie in 8, Sciences Geologiqlles 46:1-77. Howell, F. C Preliminary observations on Carnivora from the Sahabi Formation (Libya); pp in N. T. Boaz, A. EI-Arnauti, A. W. Gaziry, J. de Heinzelin, and D. D. Boaz (eds.), Neogene Paleontolog)' and Geolog)' ofsalwbi. Liss, New York. Howell, F. C., and N. Garcia Carnivora (Mammalia) from Lemudong'o (Late Miocene, Narok District, Kenya). Kirtlandia 56: Howell, F. c., and G. Petter Comparative observations on some Middle and Upper Miocene hyaenids. Genera: PercroCllta Kretzoi, Alloilyaena Kretzoi, AdcroCllta Kretzoi (Mammalia, Carnivora, Hyaenidae). Geobios 18: Hunt, R. M. Jr Evolution of the aeluroid Carnivora: significance of the ventral promontorial process of the petrosal, and the origin of basicranial patterns in the living families. American Mliselllll Novitates 2930: Evolution of the aeluroid Carnivora: Viverrid affinities of the Miocene carnivoran Herpestides. American lvfllsellm Novitates 3023: Biogeography of the order Carnivora; pp in ]. L. Gittleman (ed.), Camivore BelwviOl; Ecolog)" and Evoilition, vol. 2. Cornell University Press, Ithaca, N.Y a. Amphicyonidae; pp in C. M. Janis, K. M. Scott, and L. L. Jacobs (eds.), Evoilition of Tertiary Mammals of Nor til America: Valli me 1. Terrestrial Camivores, Ungulates, and Unglliatelike Mammals. Cambridge University Press, Cambridge b. Evolution of the aeluroid Carnivora: diversity of the earliest aeluroids from Eurasia (Quercy, Hsanda-Gol) and the origin of felids. American l\;ilisellm Novitates 3252: LAURASIATHERIA

53 c. Ursidae; pp in C. M. Janis, K. M. Scott, and L. L. Jacobs (eds.), Evoilltioll of Tertiary Mamlllais of North America: Voilime 1. Terrestrial Carnivores, Ullgulates, alld Ullgulatelike Mmllmals. Cambridge University Press, Cambridge Intercontinental migration of large mammalian carnivores: Earliest occurrence of the Old World beardog Amphicyoll (Carnivora, Amphicyonidae) in North America. Bulletill of tile AmericaJl Museum ofnatuml History 279: Kohno, N. and C. E. Ray Pliocene walruses from the Yorktown Formation of Virginia and North Carolina, and a systematic revision of the North Atlantic Pliocene walruses. Virgillia Museum of Natural History Special PlIblicatioll 14: Koretski, 1. A., and C. E. Ray Phocidae of the Pliocene of Eastern USA. ViIgillia Museum of Nat lira I History Special PublicatioJl 14: Kretzoi, M Die Raubtiere von Gombasziig nebst einer Ubersicht der Gesamtfauna. Al11zales Historico-Naturales Mllsei NatiolJalis HlIl1garici 31: Kurten, B The Neogene wolverine Plesiogulo and the origin of Gulo (Carnivora, Mammalia). Acta Zoologica Fel11zica 131: Fossil Carnivora from the Late Tertiary of Bled Douarah and Cherichira, Tunisia. Notes dll Service Geologique de Tzmisie 42: Kurten, B., and L. Werdelin A review of the genus Clzasmaportlzetes Hay, 1921 (Carnivora, Hyaenidae). Joumal of Vertebrate Paleolltology 8: Kuss, S. E Problematische Caniden des europaischen Tertiars. Bericilte del' Natllrforscilellden Gesellsciwft Zll Freibll1g im Brisgall 52: Revision der europaischen Amphicyoninae (Canidae, Carnivora, Mamm.) ausschliesslich der voroberstampischen Formen. Sitzzmgsbericilte der Heidelbe/gerAkademie del' WissenscizafteJl, Mati1e1llatiscil-Natllrwissel1scizaftlige Klasse 1965: Lewis, M. E., and L. Werdelin Trends in the evolution and ecology of the genus CroCllta. JOllmal of Vertebrate Paleolltology 17 (suppl. to number 3):60A The evolution of spotted hyaenas (CroCl/ta). Hyaella Specialist GrollP Newsletter 7: Martinez Navarro, B., and P. Palmqvist Presence of the African machairodont Megmztereol1 wilitei (Broom, 1937)(Felidae, Carnivora, Mammalia) in the lower Pleistocene site of Venta Micena (Orce, Granada, Spain), with some considerations on the origin, evolution and dispersal of the genus. Joumal of Arciweological Science 22: Presence of the African saber-toothed felid Megalltereon whitei (Broom, 1937)(Mammalia, Carnivora, Machairodontinae) in Apollonia-l (Mygdonia Basin, Macedonia, Greece). Journal of Arclzaeological Science 23: Michel, P Un nouveau Mellivorinae (Carnivora, Mustelidae) du Pleistocene de Buknadel (region de Rabat-Maroc) = Me/livora carolae n. sp. COl1zptes ReJldllS de l'academie des Sciellces, Paris, Serie II, 306: Miller, E. R Faunal correlation of Wadi Moghara, Egypt: Implications for the age of Prohylobates tandyi. JOllrnal of Hllman Evolutioll 36: Morales,]., and M. Pickford. 2005a. Carnivores from the Middle Miocene Ngorora Formation (13-12 Ma), Kenya. Estudios Geologicos 61: b. Giant bunodont Lutrinae from the Mio-Pliocene of Kenya and Uganda. EStlldios Geologicos 61: A large percrocutid carnivore from the Late Miocene (ca Ma) of Nakali, Kenya. Al11wles de PaleolJtologie 92: Creodonts and carnivores from the Middle Miocene Muruyur Formation at Kipsaraman and Cheparawa, Baringo District, Kenya. Comptes ReIJdus Palevol 7: Morales, ]., M. Pickford, S. Fraile, M. Salesa, and D. Soria Creodonta and Carnivora from Arrisdrift, early Middle Miocene of southern Namibia. lvlemoirs of tile Geological Survey of Namibia 19: Morales, J., Pickford, M., and Salesa, M. J Creodonta and Carnivora from the early Miocene of the northern Sperrgebiet, Namibia. J'vlemoirs oftlze Geological Survey of Namibia 20: Morales,]., M. Pickford, M. Salesa, and D. Soria The systematic status of Kelba, Savage, 1965, Kezzyalutra, Schmidt-Kittler, 1987 and Ndamatlwia Jacobs et ai, 1987 (Viverridae, Mammalia) and a review of early Miocene mongoose-like carnivores of Africa. A/males de Paleontologie 86: Morales, J., M. Pickford, and D. Soria Carnivores from the Late Miocene and basal Pliocene of the Tugen Hills, Kenya. Revista de la Sociedad Geologica de Espana 18: New carnivoran material (Creodonta, Carnivora and Incertae sedis) from the early Miocene of Napak, Uganda. Paleontological Researclz 11: Morales,]., M. Pickford, D. Soria, and S. Fraile New carnivores from the basal Middle Miocene of Arrisdrift, Namibia. Ec/ogae Geologicae Helvetiae 91: a. New Viverrinae (Carnivora: Mammalia) from the basal Middle Miocene of Arrisdrift, Namibia. Palaeontologia Africmw 37: Morales, ]., M. ]. Salesa, M. Pickford, and D. Soria. 2001b. A new tribe, new genus and two new species of Barbourofelinae (Felidae, Carnivora, Mammalia) from the early Miocene of East Africa and Spain. Transactions of the Royal Society of EdinblI1glz, Earth Sciences 92: Morlo, M., E. R. Miller, and A. N. EI-Barkooky Creodonta and Carnivora from Wadi Moghra, Egypt. Journal of Vertebrate Faleontolog)' 27: Morlo, M., S. Peigne, and D. Nagel A new species of Prosmlsanosmilus: Implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia). Zoological Joumal oftlze Li/mean Society 140: Muizon, C. de Premier signalement de Monachinae (Phocidae, Mammalia) dans Ie Sahelien (Miocene superieur) d'oran (Algerie). Palaeovertebrata 11: Muizon, C. de, and Q. B. Hendey Late Tertiary seals of the South Atlantic Ocean. Annals of the South African Museum 82: Munthe, K Canidae; pp in C. M.Janis, K. M. Scott, and L. L. Jacobs (eds.), Evolution of Tertiar), Mammals of Nortlz America: Volume 1. Terrestrial Carnivores, Ungulates, and Unglllatelike Mamlllais. Cambridge University Press, Cambridge. Nakaya, H., M. Pickford, Y. Nakano, and H. Ishida The Late Miocene large mammal fauna from the Namurungule Formation, Samburu Hills, Northern Kenya. African Study Monographs, Supplementary Issue 2: Nowak, R. M Walker's Marine Ma17l11zals of the World. Johns Hopkins University Press, Baltimore, 264 pp Walker's Ca1'1livores of tile World. Johns Hopkins UniverSity Press, Baltimore, 328 pp. Palmqvist, P On the presence of Megantereon whitei at the South Turkwel hominid site, northern Kenya. Joumal of Paleontolog)' 76: Palmqvist, P., V. Torregrosa, J. A. Perez-Claros, B. Martinez-Navarro, and A. Turner A re-evaluation of the diversity of lvlegantereon (Mammalia, Carnivora, Machairodontinae) and the problem of species identification in extinct carnivores. Joumal of Vertebrate Paleontology 27: Peigne, S Systematique et evolution des Feliformia (Mammalia, Carnivora) du Paleogene d'eurasie. Unpublished PhD dissertation, Universite de Poitiers, Poitiers, France, 396 pp. Peigne, S., Y. Chaimanee, C. Yamee, P. Tian, and J.-J. Jaeger A new amphicyonid (Mammalia, Carnivora, Amphicyonidae) from the late Middle Miocene of northern Thailand and a review of the amphicyonid record in Asia. JOllmal of Asian Earth Sciences 26: Peigne, S., and L. de Bonis The genus Stenoplesictis Filhol (Mammalia, Carnivora) from the Oligocene deposits of the Phosphorites of Quercy, France. JOllmal of Vertebrate Paleontolog)' 19: Peigne, S., L. de Bonis, A. Likius, H. T. Mackaye, P. Vignaud, and M. Brunet A new machairodontine (Carnivora, Felidae) from the Late Miocene hominid locality of TM266, Toros-Menalla, Chad. Comptes Rendlls PalevoI4: a. Late Miocene Carnivora from Chad: Lutrinae (Mustelidae). Zoological Journal Of tile Lilllzmn Society 152: Peigne, S., L. de Bonis, H. T. Mackaye, A. Likius, P. Vignaud and M. Brunet. 2008b. Late Miocene Carnivora from Chad: Herpestidae, Viverridae, and small-sized Felidae. C017lptes Rendus Palevol 7: Peigne, S., and E. P. ]. Heizmann The Amphicyonidae (Mammalia: Carnivora) from Ulm-Westtangente (MN 2, early Miocene), Baden-Wtirttemberg, Germany: Systematics and ecomorphology. Stuttgarter Beitriige zur Natll1kwzde, Serie B (Geologie wzd Paliiontologie) 343: THIRTY-TWO: CARNIVORA 655

54 Petter, G Petits carnivores villafranchiens du Bed I d'oldoway (Tanzanie); pp in Problel11es actuels de Paleontologie (Evolution des Vertebres). Colloques Internationaux du CNRS, Paris Carnivores pleistocenes du ravin d'olduvai (Tanzanie); pp in L. S. B. Leakey, R.]. G. Savage, and S. C. Coryndon (eds.), Fossil Vertebrates of Africa, vol. 3. Academic Press, London Small carnivores (Viverridae, Mustelidae, Canidae) from Laetoli; pp in M. D. Leakey and]. M. Harris (eds.), Laetoli: A Pliocene Site in Northern Tanzania. Clarendon Press, Oxford. Petter, G. and F. C. Howell Diversite des Carnivores (Mammalia, Carnivora) dans les faunes du Pliocene moyen et superieur d'afrique orientale. Indications paleoecologiques; pp in Fondation Sinl'er-Polignac (ed.), L'environnement des HOl11inides au Plio-Pleistocene. Masson, Paris. Petter, G., M. Pickford, and F. C. Howell La loutre piscivore du Pliocene de Nyaburogo et de Nkondo (Ouganda, Afrique occidentale): Torolutra ougandemis n. g., n. sp. (Mammalia, Carnivora). COil1ptes Rendus de l'academie des Sciences, Paris 312: Petter, G., and H. Thomas Les Agriotherinae (Mammalia, Carnivora) neogenes de I'ancien monde. Presence du genre Indarctos dans la faune de Menacer (ex-marceau), Algerie. Geobios 19: Pickford, M Cainozoic Palaeontological Sites of Western Kenya. Miincilller GeolVissensciwftlicile Abhmidiungen A 8: Southern Africa: A cradle of evolution. Soutiz African Journal of Science 100: Pickford, M., T. Fernandes, and S. Ao Nouvelles decouvertes de remplissages de fissures it primates dans Ie "Planalto da Humpata," Hulla, Sud de Angola. Comptes Rendus de l'academie des Sciences 310: Pickford, M., and B. Senu! Geology and paleobiology of the Namib Desert, southwestern Africa. Memoirs oftize Geological Survey of Namibia 18: Pickford, lv!., B. Senut, D. Gommery, and E. Musiime New Catarrhine fossils from Moroto II, early Middle Miocene (ca Ma) Uganda. Comptes Rendus Palevol 2: Pilgrim, G. E Catalogue oftize POlltian Carnivora of Europe ill the Department of Geolog)'. British Museum (Natural History), London, 174 pp The fossil Carnivora of India. PalaeolJtologia Indica 18: Pohle, H Die Raubtiere von Oldoway. WisseJ1sciwftliclle Eigebilisse der Oldoway-Expedition 1913 (N.F.) 3: Potts, R., and A. Deino Mid-Pleistocene change in large mammal faunas of eastern Africa. Quaternary Researciz 43: Rasmussen, D. T. and!v!. Gutierrez A mammalian fauna from the Late Oligocene of Northwestern Kenya. Palaeontograpizica Abt. A 288:1-52. Ragl, F Palaeogeographic considerations for Mediterranean and Paratethys seaways (Oligocene to Miocene). Annalen des Naturizistoriscizes Museums in Wien 99: Roth, C Leptoplesictis Major 1903 (Mammalia, Carnivora, Viverridae) aus dem Orleanium und Astaracium/Miozan von Frankreich und Deutschland. Paliiontoiogiscile Zeitscizrift 62: Salles, L. O Felid phylogenetics: Extant taxa and skull morphology (Felidae, Aeluroidea). Americall Museum Novitates 3047:1-67. Sardella, R Sistematica e distribuzione stratigrafica dei Macairodontini dal Miocene Superiore al Pleistocene. Unpublished PhD dissertation, Department of Geology, University of Florens, Florens, Italy, 137 pp. Sardella, R., and L. Werdelin Aniphimacizairodus (Felidae, Mammalia) from Sahabi (latest Miocene-earliest Pliocene, Libya), with a review of African Miocene Machairodontinae. Rivista Italiana di Paleontologia e Stratigrafia 113: Savage, R.]. G Fossil mammals of Africa: 19. The Miocene Carnivora of East Africa. Bulletin oft/ie British lvfuseum (Natural HistOlY): Geolog)' 10: Carnivora; pp in V. ]. Maglio and H. B. S. Cooke (eds.), Evolution of African Mammals. Harvard University Press, Cambridge. Savage, R. J. G., and W. R. Hamilton Introduction to the Miocene mammal faunas of Gebel Zelten, Libya. Bulletin of the Britisil Museum (Natural History), Geolog)' 22: Schmidt-Kittler, N Raubtiere aus dem Jungtertiar Kleinasiens. Paleontographica, Abt. A 155: The Carnivora (Fissipedia) from the Lower Miocene of East Africa. Palaeontographica, Abt. A 197: Servheen, c., S. Herrero, and B. Peyton eds Bears. Status Survey and Conservation Action Plan. IUCN/SSC Bear and Polar Bear SpeCialist Groups. IUCN, Gland, Switzerland. Sillero-Zubiri, c., M. Hoffmann, and D. W. Macdonald CmIids: Foxes, Wolves, Jackals and Dogs. Status Survey and Conservation Action Plan. IUCN, Gland, Switzerland. Spassov, N., and L. Rook EuC)'on marinae sp. nov. (Mammalia, Carnivora), a new canid species from the Pliocene of Mongolia, with a review of forms referable to the genus. Rivista Italiana di Paleontologia e Stratigmfia 112: Staaden, lv!. J. van Suricata suricatta. Mammalian SpeCies 483:1-8. Stromer, E Fossile Wirbeltier-Reste aus dem Uadi Faregh und Uadi Natnln in Agypten. Abhandlrlllgen Hemusgegeben von der Senckenbelgisc1len Naturforsclienden Gesellsc1zaft 29: Mitteilungen liber Wirbeltierreste aus dem Mittelpliocan des Natrontales (Agypten). Zeitschrift der Deutscilen Geologischen Gesellsc1wft, Ablwndhlllgen A 65: Sudre, ]., andj.-l. Hartenberger Oued Mya 1, nouveau gisement de mammiferes du Miocene superieur dans Ie sud Algerien. Geobios 25: Sunquist, M., and F. Sunquist Wild Cats of the World. University of Chicago Press, Chicago, 452 pp. Taylor, M. E Ici11leumia albicallda. Mammalian Species 12:1-4. Taylor, P. J., and J. A.]. Meester C)'nictis penicillata. Mammalian Species 432:1-7. Tedford, R. H., and Z. Qiu Pliocene Nyctereutes (Carnivora: Canidae) from Yushe, Shanxi, with comments on Chinese fossil raccoon-dogs. 11ertebrata PalAsiatica 29: A new canid genus from the Pliocene of Yushe, Shanxi Province. 11ertebmta PalAsiatica 34: Tedford, R. H.,. Wang, and B. E. Taylor Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae). Bulletin of the A11lericmzlvfuseum ofnatuml History 325: Tsujikawa, H The updated Late Miocene large mammal fauna from Samburu Hills, northern Kenya. African Stud)' A onogmphs 32:1-50. Turner, A Hominids and fellow travellers: Human migration into high latitudes as part of a large mammal community; pp in R. Foley (ed.), Hominid Evolution alld Commzl11ity Ecology Academic Press, London lvfegantereon CllltridellS (Cuvier)(Mammalia, Felidae, Machairodontinae) from Plio-Pleistocene deposits in Africa and Eurasia, with comments on dispersal and the possibility of a New World origin. JOll1'1wl of Paleontology 61: The evolution of the guild of larger terrestrial carnivores during the Plio-Pleistocene in Africa. Geobios 23: Turner, A., L. Bishop, C. Denys, and]. K. McKee Appendix: A locality-based listing of African Plio-Pleistocene mammals; pp in T. G. Bromage and F. Schrenk (eds.), African BiogeograpllY, Climate Change, and HllInan Evolution. Oxford University Press, Oxford. Veron, G La position systematique de Cryptoprocta ferox (Carnivora) : Analyse cladistique des caracteres morphologiques de carnivores Aeluroidea actuels et fossiles. AIa11l11laiia 59: Veron, G., M. Colyn, A. E. Dunham, P. Taylor, and P. Gaubert Molecular systematics and origin of sociality in mongooses (Herpestidae, Carnivora). lvfolecular Phylogenetics and Evolution 30: Ward, O. G., and D. H. Wurster-Hill N),ctereutes procyonoides. Mammalian Species 358:1-5. Werdelin, L Carnivores, exclusive of Hyaenidae, from the later Miocene of Europe and Western Asia; pp in R. L. Bernor, V. Fahlbusch, and H.-W. Mitlmann (eds.), The Evolution of West em Eurasian Neogene lvfa11l11lal Fml11as. Columbia University Press, New York. Werdelin, L. 1999a. PaciIycroCllta (hyaenids) from the Pliocene of east Africa. Palaontoiogiscile Zeitsc1lrift 73: b. Studies of fossil hyaenas: Affinities of LyC)'aenops rl boideae Kretzoi from PestlOrinc, Hungary. Zoological Joumal of the Li1l1Iemi Society 126: Mio-Pliocene Carnivora from Lothagam, Kenya; pp in M. G. Leakey and ]. D. Harris (eds.), Lot/wgal1z: Dawn of Hwnal1it)' in Eastem Africa. Columbia University Press, New York. Werdelin, L., and]. Barthelme Brown hyena (Para hyaena brw11lea) from the Pleistocene of Kenya. Journal of 11ertebmte Paleontology 17: LAURASIATHERIA