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The hyaenodontidans from the Gour Lazib area (?Early Eocene, Algeria): implications concerning thesystematics and the origin of the Hyainailourinae andTeratodontinaeFloréal Solé a b , Julie Lhuillier c , Mohammed Adaci d , Mustapha Bensalah d , M’hammedMahboubi e & Rodolphe Tabuce ca Muséum national d’histoire naturelle, Département Histoire de la Terre, CP 38; UMR 7207– CNRS: Centre de recherche sur la paléobiodiversité et les paléoenvironnements; 57 rueCuvier; , Paris , F-75005 , Franceb Institut de Génomique Fonctionnelle de Lyon , Université de Lyon, CNRS, UMR 5242, INRA,UCBL 1, Ecole Normale Supérieure de Lyon, 46 Allée d’Italie, 69364 Lyon Cedex 07 , Francec Laboratoire de Paléontologie, Institut des Sciences de l’Évolution (ISE-M, UMR – CNRS5554), C.c. 64 , Université Montpellier 2, Place Eugène Bataillon, F-34095 MontpellierCedex 05 , Franced Laboratoire de recherche n°25, Département des Sciences de la Terre , Université AbouBekr, Belkaïd, B.P. 119 , Tlemcen , 13000 , Algeriae Laboratoire de Paléontologie stratigraphique et Paléoenvironnement , Université d’Oran,B.P. 1524 El M’naouer , Oran , 31000 , AlgeriaPublished online: 16 Jul 2013.

To cite this article: Journal of Systematic Palaeontology (2013): The hyaenodontidans from the Gour Lazib area (?EarlyEocene, Algeria): implications concerning the systematics and the origin of the Hyainailourinae and Teratodontinae, Journalof Systematic Palaeontology, DOI: 10.1080/14772019.2013.795196

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Journal of Systematic Palaeontology, 2013http://dx.doi.org/10.1080/14772019.2013.795196

The hyaenodontidans from the Gour Lazib area (?Early Eocene, Algeria):implications concerning the systematics and the origin

of the Hyainailourinae and TeratodontinaeFloreal Solea,b∗, Julie Lhuillierc, Mohammed Adacid, Mustapha Bensalahd, M’hammed Mahboubie and Rodolphe Tabucec

aMuseum national d’histoire naturelle, Departement Histoire de la Terre, CP 38; UMR 7207 – CNRS: Centre de recherche sur lapaleobiodiversite et les paleoenvironnements; 57 rue Cuvier; F-75005, Paris, France; bInstitut de Genomique Fonctionnelle de Lyon,Universite de Lyon, CNRS, UMR 5242, INRA, UCBL 1, Ecole Normale Superieure de Lyon, 46 Allee d’Italie, 69364 Lyon Cedex 07,

France; cLaboratoire de Paleontologie, Institut des Sciences de l’Evolution (ISE-M, UMR – CNRS 5554), C.c. 64, Universite Montpellier2, Place Eugene Bataillon, F-34095 Montpellier Cedex 05, France; dLaboratoire de recherche n◦25, Departement des Sciences de la

Terre, Universite Abou Bekr, Belkaıd, B.P. 119, Tlemcen 13000, Algeria; eLaboratoire de Paleontologie stratigraphique etPaleoenvironnement, Universite d’Oran, B.P. 1524 El M’naouer, Oran 31000, Algeria

(Received 23 June 2011; accepted 16 November 2012)

The Algerian localities of the Gour Lazib area (Early or early Middle Eocene) have yielded an important mammalian fauna.The Hyaenodontida are well represented in this fauna: three species–two are new– are reported. The genus Glibzegdouia,which has been previously described as a possible Carnivora, is now clearly referred to the Hyaenodontida. It appearsmorphologically close to Masrasector and Dissopsalis. A new genus, Furodon, is described. It appears morphologically closeto the oldest Pterodon species. This discovery supports an African origin for the hyainailourine genus Pterodon and relatedgenera (e.g. Hyainailouros, Akhnatenavus). Two very small lower molars are referred to a new genus Parvavorodon, whichis also referred to Hyainailourinae. The localities of the Gour Lazib area therefore show important hyaenodontid diversityfor the Early or early Middle Eocene. We performed a new phylogenetic analysis to question the relationships betweenthe African, Asian, North American and European hyaenodontidans. Our study supports the endemism and originalityof the Asian ‘proviverrines’ Indohyaenodon, Paratritemnodon, Kyawdawia and Yarshea; we propose a new subfamily:Indohyaenodontinae. The African ‘proviverrines’ (e.g. Masrasector, Anasinopa, Dissopsalis and Glibzegdouia), which arenotably characterized by large premolars and the presence of a wide talonid on the molars, are close to the enigmatic AfricanTeratodon. We therefore propose to refer them to Teratodontinae. The Hyainailourinae, which include the new genus Furodon,are characterized by the presence of secant dentition related to a hypercarnivorous diet. They appear phylogenetically closeto the African Koholiinae. The genus Metapterodon is referred to Koholiinae based on the phylogenetic analysis. TheAfrican origin of the Teratodontinae and Hyainailourinae is supported by Glibzegdouia and Furodon. The origination ofseveral subfamilies in Africa supports the hypothesis of an African origin for the Hyaenodontida. The origination of theTeratodontinae in Africa contradicts the previous hypotheses of Afro-Asian ‘proviverrines’.

Keywords: Gour Lazib; Algeria; Africa; Hyaenodontida; Hyainailourinae; Teratodontinae

Introduction

The Early and Middle Eocene carnivorous placen-tal mammals from Laurasia and Africa correspondto the ‘Miacidae’, Viverravidae, Hyaenodontidae andOxyaenidae. The most characteristic dental feature of thesegroups is the presence of teeth that are devoted to ‘cut’ themeat–the carnassial teeth. The ‘Miacidae’ and Viverravidaeare referred to Carnivoramorpha, whereas the Hyaenodon-tidae and Oxyaenidae have been usually grouped amongthe ‘Creodonta’, a clade that has been demonstrated to bediphyletic (Polly 1994, 1996; Gheerbrant et al. 2006; Soleet al. 2009). Sole (2013) therefore proposed to use Oxyaen-odonta and Hyaenodontida.

In Africa, only the Hyaenodontida have been recordeduntil the Oligocene (Holroyd 1994; Gheerbrant et al.

∗Corresponding author. Email: sole@mnhn.fr

2006; Rasmussen & Guttierrez 2009) and the previ-ously described early Palaeogene Carnivora (Gheerbrant1995; Crochet et al. 2001) are now considered as hyaen-odontidans (Werdelin & Peigne 2010), so the earliestrecord of Carnivora is Late Oligocene for that conti-nent (Kenya; Rasmussen & Guttierrez 2009). Hence,Hyaenodontida represent the only specialized carnivorousmammals present in Africa until the latest part of theOligocene.

The Hyaenodontida are considered to originate eitherduring the Palaeocene in Africa (Gheerbrant 1995; Soleet al. 2009) or in Asia (Meng et al. 1998; Bowen et al.2002). Presently, two Late Palaeocene species have beenrecorded in Africa: the primitive Tinerhodon Gheerbrant,1995 (Adrar Mgorn, Morocco) and the much derivedLahimia Sole & Gheerbrant in Sole et al. (2009) (Ouled

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Table 1. Measurements (mm) of the specimens of Glibzegdouiatabelbalaensis Crochet, Peigne & Mahboubi, 2001 from GlibZegdou and Gour Lazib (Algeria).

Locus n Observed range Mean

P3 L 2 5.26–5.48 5.37W 2 3.62–3.68 3.65

M2 L 1 6.93 —W 0 — —

M1 L 1 6.68 —W 1 3.59 —

M2 L 1 7.64 —W 1 4.41 —

Abdoun Basin, Morocco). Two Early Eocene hyaenodon-tidans are known in Africa: Boualitomus Gheerbrant inGheerbrant et al. (2006) (Ouled Abdoun Basin, Morocco)and Koholia Crochet, 1988 (El Kohol, Algeria). Boualito-mus, Koholia and Lahimia belong to the Koholiinae.

Two poorly preserved Lutetian? hyaenodontidans –Pterodon sp. indet. and “Proviverrinae” genus sp. indet. –have been recently described by Pickford et al. (2008)from the Black Crow locality (Sperrgebiet, Namibia). Morerecently, Grohe et al. (2010, in press) have described newspecimens of Apterodontinae from the ?Middle Eocenelocality of Dor El Talha (Libya). Their study supports anAfrican origin and a semi-aquatic lifestyle for Apterodon(Grohe et al. in press).

In contrast to the Early and Middle Eocene povertyof specimens, the Late Eocene-Oligocene African hyaen-odontidans are numerous: seven genera (ApterodonFischer, 1880; Quasiapterodon Lavrov, 1999; Masrasec-

Table 2. Measurements (mm) of the specimens of Furodoncrocheti gen. et sp. nov. from Glib Zegdou (Algeria).

Locus n Observed range Mean

M1 L 2 4.38–4.73 4.56W 2 5.05–5.86 5.68

C L 1 6.55 —W 1 4.98 —

P1 L 1 >2.32∗ —W 1 >2.45∗ —

P2 L 1 >5.19∗ —W 1 >2.74∗ —

P3 L 1 >5.21∗ —W 1 >2.79∗ —

P4 L 1 6.27 —W 1 3.02 —

M1 L 1 5.75 —W 1 3.08 —

M2 L 1 6.6 —W 1 4.08 —

M3 L 1 8.01 —W 1 4.84 —

MD H 1 12.44 —Weight = 1.9 kg†

∗Estimated on the basis of the roots; †Weight estimated after Morlo (1999).

Table 3. Measurements (mm) of the specimens of Parvavorodongheerbranti gen. et sp. nov. from Gour Lazib (Algeria).

Locus n Observed range Mean

M1 or M2 L 2 3.82–4.03 3.93W 2 1.90–2.11 2

tor Simons & Gingerich, 1974; Metasinopa Osborn, 1909;Metapterodon Stromer, 1926; Pterodon Blainville, 1839and Akhnatenavus Holroyd, 1999) are recorded in the locali-ties of the Jebel Qatrani Formation, the Birket Qarun Forma-tion and the Qsar el Sagha Formation (Fayum, Egypt).The genera Apterodon and Quasiapterodon are referred toApterodontinae, whereas the five other genera are referredto the Hyainailourinae (Morlo et al. 2007; Lewis & Morlo2010). In addition to this, a new proviverrine hyaenodonti-dan from the earliest Late Eocene of Egypt (∼37 Ma) hasbeen recently discovered, but not yet formally describedand named (Borths et al. 2010); this taxon represents theoldest hyaenodontidan ever discovered in the Fayum.

The fossil record of Hyaenodontida in Africa showstherefore a significant sampling gap between the LatePalaeocene-Early Eocene and Late Eocene-Oligocenetimes.

Here, we describe new creodonts from several localitiesof the Gour Lazib, Tindouf Province, Algeria. The fossilscome from the Glib Zegdou (levels HGL 50, HGL 50bis andHGL 52) and from the central part of the Gour Lazib (HGL04 and HGL 10); all sites are from the middle memberof the Glib Zegdou Formation which is late Ypresian orearly Lutetian in age (Adaci et al. 2007). Recent magne-tostratigraphic and biostratigraphic studies suggest an ageranging between 49 and 45 million years ago for the GlibZegdou Formation ( = Ypresian-Lutetian boundary or mid-Lutetian) (Costeur et al. 2012). The Gour Lazib outcropscomprise a number of fossiliferous outliers, including theGlib Zegdou, having yielded a rich mammalian fauna. Onlyone carnivorous mammal was known until now; this taxonwas named Glibzegdouia and attributed to the Carnivora byCrochet et al. (2001), but Werdelin & Peigne (2010) consid-ered that its ordinal assignment is uncertain and suggestedcreodont affinities. The new material provides not only abasis for the reassessment of Glibzegdouia but also allowsus to describe two new genera. These fossils are interme-diate in age and morphology between the African earlyPalaeogene creodonts and the late Palaeogene ones, so theyare of primary importance for our understanding of theevolution of creodonts in Africa.

Material and methods

The terminology of the molar dental cusps and crestsfollows Van Valen (1966). The measurements (in mm)follow those used by Gingerich & Deutsch (1989) for the

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Hyaenodontidans from the Gour Lazib area 3

hyaenodontidans. The material here presented is housed inthe collections of the Universite Sciences et Techniques duLanguedoc, Montpellier 2.

Institutional abbreviationsHGL, Hammada Gour Lazib, Algeria; GZC, Glib Zegdou,Oran University collections, Oran-Es Senia, Algeria.

Systematic palaeontology

Order Hyaenodontida Leidy, 1869Family Hyaenodontidae Leidy, 1869

Subfamily Teratodontinae Savage, 1965.

Emended diagnosis. Teratodontinae differ fromHyainailourinae by the combination of derived andprimitive features. Their primitive features are: the pres-ence of a wide-basined talonid with developed entoconidon molars, and the absence of protocone on P3. The derivedfeatures are: the robustness of the premolars; the widepostfossid on P4; the postfossid lingually closed and wideron molars than in primitive hyaenodontidans; the talonid aswide as the trigonid; the developed ectocingulid on molars;the bulbous and anteriorly directed protocone on P4; the

absence of the parastyle on P4; the separated paraconeand metacone on molars. As numerous hyaenodontidangroups, the evolution of the Teratodontinae is characterizedby the development of a secant dentition (e.g. loss ofthe metaconid). The entoconid is reduced in derivedgenera (e.g. Dissopsalis), but the talonid remains wide andlingually closed on M1 and M2 in all Teratodontinae.

Included genera. Dissopsalis Pilgrim, 1910; TeratodonSavage, 1965; Anasinopa Savage, 1965; MasrasectorSimons & Gingerich, 1974; Glibzegdouia Crochet, Peigne& Mahboubi 2001; Buhakia Morlo, Miller & El-Barkooky,2007; Mlanyama Rasmussen & Guttierrez, 2009.

Geographical and stratigraphical range. Possibly EarlyEocene to late Middle Miocene; Africa and Asia.

Glibzegdouia Crochet, Peigne & Mahboubi, 2001

Diagnosis. Same as for the type and only known species.

Glibzegdouia tabelbalaensis Crochet, Peigne &Mahboubi, 2001

(Fig. 1)

Diagnosis. Small hyaenodontidan characterized by thepresence on molars of an ectocingulid, enlarged talonid,closed postfossid, and separated paracone and metacone. It

Figure 1. Glibzegdouia tabelbalaensis Crochet, Peigne & Mahboubi, 2001. Scanning electron micrographs. HGL 10-15, LM2: A, occlusalview (reversed). HGL 50-411, LM1: B, occlusal view; E, labial view; F, lingual view. GZC 35, LM2 (Holotype): C, occlusal view; D,lingual view; G, labial view. Scale: 1.5 mm.

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differs from its closest genus, Masrasector, notably by lessreduced metaconid, less mesially located paraconid, andnarrower talonid. It differs from other teratodontines by amore developed entoconid on molars.

Holotype. GZC 35, LM2.

Referred specimens. HGL 50-411, LM1; HGL 50-406,RP3; HGL 50-408, RP3; HGL 10-15, LM1.

Age and locality. HGL 10 and HGL 50, Gour Lazib, Alge-ria, late Ypresian or middle Lutetian.

Description – Upper dentition. Premolars. P3: Theenamel is slightly crenulated. The tooth is elongatedmesiodistally. There is a small cingulum surrounding thetooth. A swelling is present lingually, but no protocone ispresent.

Molars. M1: The only available specimen is unfortunatelyincomplete. The protocone area is missing, but it was proba-bly large. The paracone and metacone are almost fully sepa-rated. They are almost the same height. The parastylar areais reduced, but present. The stylar shelf is very narrow. Thepostmetacrista is not significantly shifted distally. More-over, the postmetacrista is short compared with the observa-tion in primitive hyaenodontidans (e.g. Prototomus, Arfia).A carnassial notch is present along the postmetacrista.

Description – Lower dentition. Molars. M1: The trigonidon HGL 50-411 is longer than the talonid. The trigonid andtalonid have the same width. The protoconid is the highestcusp. The paraconid is slightly lower than the metaconid.The metaconid is clearly the less developed cusp ofthe trigonid. The metaconid is transversally aligned withthe protoconid, while the paraconid is mesially located. Thetalonid is wide and bears three distinct cusps (entoconid,hypoconulid and hypoconid). The entoconid is the highestcusp, the hypoconid the lowest. The hypoconulid andentoconid are close and more distal than the hypoconid.The cristid obliqua is slightly oblique (distally and labiallyshifted). It reaches the trigonid on the protoconid distalfacet. The talonid is closed lingually by an entoconulid.The ectocingulid is well developed. The anterior keel (=precingulid) is short and is not linked to the ectocingulid.

M2: GZC 35 was the first described specimen of Glibzeg-douia (Crochet et al. 2001). It was identified as an M1. HGL50-411 is morphologically similar to GZC 35 but differs bya clearly smaller size and a lower paraconid. On GZC 35,the paraconid is only slightly smaller than the protoconidand the talonid is slightly narrower than the trigonid. Thecristid obliqua is more oblique on GZC 35 than on HGL50-411. The cristid obliqua reaches the trigonid just belowthe intraprotocristid notch. The differences between GZC35 and HGL 50-411 correspond to those usually observedbetween M1 and M2 among Hyaenodontida–the M2 beinglarger and more secant. We consider now that GZC 35 is

an M2. The crown bases of the M1 and M2 are convex inlateral views.

Comparison. The reference of HGL 50-411 to Glibzeg-douia is supported by the commonly shared morphologywith GZC 35: talonid wide and basined with developedentoconid, metaconid transversally aligned with the proto-conid, enlarged hypoconid compared with primitive hyaen-odontidans, and close entoconid and hypoconulid. We referHGL 10-15 as an M1 because the parastylar area is toofeebly developed to correspond to an M2. The importantfeatures of HGL 10-15 are the important separation of theparacone and metacone, both cusps being almost subequalin height. The wear facets 3 (lingual wall of the paracone)and 4 (lingual wall of the metacone) are well marked onHGL 10-15. These facets are similar to those observed onthe labial side of the hypoconid on the M2 (GZC 35, holo-type of Glibzegdouia). The wear facet 3 on lower molarsis caused by the occlusion with the paracone of the M2,while the wear facet 4 is caused by the occlusion with themetacone.

Thanks to the discovery of new specimens, the hyaen-odontidan status of Glibzegdouia is now clearly established:the presence of a secant M2, which is larger than the M1, andthe presence of a secant M2, clearly supports the reference ofGlibzegdouia to the Hyaenodontida rather than Carnivora.

The Hyaenodontida are poorly known in the EarlyEocene of Africa. The two previously described species,Lahimia and Koholia, have been referred to the Koholi-inae (Crochet 1988; Sole et al. 2009). These species arecharacterized by the absence of P1 and important devel-opment of the prevallum–postvallid shearing (Sole et al.2009). The absence of P1 cannot be checked based on theavailable specimens from the Gour Lazib area. However,we notice that the prevallum–postvallid shearing is not asdeveloped as in Lahimia and Koholia. Moreover, Glibzeg-douia differs from koholiines by a wider talonid, devel-oped ectocingulid, and separated paracone and metacone(derived features), and less reduced entoconid (primitivefeature). Hence Glibzegdouia clearly differs from koholi-ine genera. The only known African ?Middle Eocene hyaen-odontidan is the Apterodontinae from Dor El Talha, Libya(Grohe et al. 2010, 2012). The age of the fauna from Dor ElTalha is in fact problematic and a mid-Priabonian (middleLate Eocene) age could be more likely according to Sallamet al. (2011). Jaeger et al. (2010a, b) proposed a late MiddleEocene age for Dor El Talha. However, Glibzegdouia differsfrom Apterodon by the presence of a wide and deep post-fossid and developed trigonid; this contrasts with impor-tant reduction of the paraconid, metaconid, entoconid andhypoconulid observed in Apterodon (Szalay 1967).

When compared with the Late Eocene-Middle MioceneAfrican hyaenodontidans, Glibzegdouia appears morpho-logically close to the genera Masrasector, Anasinopa andDissopsalis. Glibzegdouia shares with these genera the

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Hyaenodontidans from the Gour Lazib area 5

presence of an enlarged and basined talonid, the developedentoconid, the convex morphology of the base of the lowermolars in lateral view, the lingual closure of the talonid,the metaconid transversally aligned with the protoconid,the paraconid higher than the metaconid, and the pres-ence of a developed ectocingulid. Glibzegdouia also shareswith Anasinopa, Masrasector and Dissopsalis the pres-ence of separated paracone and metacone on upper molars.Glibzegdouia is morphologically closer to the oldest genus,Masrasector, than to Anasinopa and Dissopsalis. Glibzeg-douia and Masrasector differ from the two latter generaby a more developed metaconid, the presence of a distinctentoconid on molars, a talonid more elongated mesiodis-tally on molars, and by a more developed ectocingulid.However, these features are primitive among Hyaenodon-tida, except the developed ectocingulid; we think that theectocingulid is reduced because of the enlargement of thetalonid in the younger Anasinopa and Dissopsalis. Glibzeg-douia and Masrasector also differ from Anasinopa andDissopsalis by a narrower postfossid and talonid, and amore oblique cristid obliqua. These are primitive features.Finally, Glibzegdouia tabelbalaensis is clearly smaller thanthe species of Anasinopa and Dissopsalis.

Glibzegdouia differs from Masrasector by the presenceof a more developed metaconid, narrower talonid, moreoblique cristid obliqua, and less mesially located para-conid. As indicated above, all these features are primitiveamong Hyaenodontida. This is not surprising when consid-ering the oldest age of Glibzegdouia compared with generaAnasinopa, Masrasector and Dissopsalis.

Finally, the morphology of the lower and upper molarsof Glibzegdouia supports a close relationship betweenGlibzegdouia and the Late Eocene-Early Oligocene genusMasrasector. Moreover, our phylogenetic study (see below)shows a close relationship between Glibzegdouia, Masra-sector, and the younger genera Anasinopa and Dissopsalis.Because the genera Glibzegdouia is morphologically closeto Masrasector, Dissopsalis and Anasinopa, we propose torefer Glibzegdouia to Teratodontinae.

As indicated above, Pickford et al. (2008) described thetaxon ‘Proviverrinae’ genus and species indet. from the?Lutetian locality of Black Crow (Namibia). The sole speci-men that is presently available is a DP4. The specimen shareswith the M1 of Glibzegdouia the presence of separated para-cone and metacone and short postmetacrista. Moreover, theNamibian specimen shares with the upper molars of Masra-sector the presence of wide and long protocone. Thus theNamibian specimen should be referred to Teratodontinaetogether with Glibzegdouia.

Subfamily Hyainailourinae Pilgrim, 1932

Emended diagnosis. Hyaenodontids with metaconidsmall to absent, with distinct but unbasined talonid onlower molars, connate metacone and paracone present onM1–M2, a weak to absent P3 lingual cingulum, P4 lackingcontinuous lingual cingulum, relatively large anterior keels

on lower molars, M3 talonid reduced relative to that ofM1–M2, lower molar protoconids and paraconids subequalin length, circular subarcuate fossa present on petrosal, andnuchal crest not extending laterally to mastoid processes(after Lewis & Morlo 2010).

Included genus. Pterodon Blainville, 1839; Hyainailouros]Biedermann, 1863; Paroxyaena Martin, 1906; MetasinopaOsborn, 1909; Isohyaenodon Savage, 1965; LeakitheriumSavage, 1965; Megistotherium Savage, 1973; ParapterodonLange-Badre, 1979; Akhnatenavus Holroyd, 1999; Furodongen. nov.; Parvavorodon gen. nov.

Geographical and stratigraphical range. Possibly EarlyEocene to late Middle Miocene; Africa, Europe, Asia andpossibly North America (Hemipsalodon Cope, 1885).

Furodon gen. nov.

Diagnosis. Same as for the type and only known species.

Etymology. Combination of fur (Latin, thief) and odon(Latin, tooth).

Furodon crocheti sp. nov.(Fig. 2)

Diagnosis. Furodon differs from koholiine genera(Lahimia, Boualitomus) by the presence of a P1. It differsfrom Glibzegdouia and related genera by the reduction ofthe metaconid, entoconid and talonid in general. Furodonis close to Late Eocene species of Pterodon and Akhnate-navus, with which it shares the reduction of the metaconidand talonid, and the fusion of the paracone and meta-cone. However, Furodon is clearly less advanced towardshypercarnivory than Pterodon and Akhnatenavus becauseit still possesses a distinct metaconid and entoconid; theprotocone on the molars is less reduced than in Pterodonand Akhnatenavus. The presence in Furodon of closelyappressed premolars, P2 and P3 almost equal in length,and wider talonid than in Akhnatenavus and related taxasupport a close relationship between this genus and earliestPterodon species. Furodon differs from earliest Pterodonspecies by the presence of a developed P1.

Etymology. Dedicated to Jean-Yves Crochet, who hasdescribed numerous African Palaeogene mammals,included several hyaenodontidans.

Holotype. HGL 50bis-56, left mandible with canine, P4-M3 and roots of P1-P3.

Referred specimens. HGL 50-410 LM1; HGL 50-404,RM1; HGL 50-405, LM1; HGL 50-407, LM1.

Age and locality. HGL 50 and HGL 50 bis, Gour Lazib,Algeria, late Ypresian or middle Lutetian.

Description.

Mandible. Two large foramina are present, respectivelybelow the mesial root of P2 and between P3 and P4. The

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Figure 2. Furodon crocheti gen. et sp. nov. HGL 50bis-56 (holotype), left mandible with roots of canine, P1-P4 and M1-M3: A, Computedtomography scan view; B, C, general views of the holotype in B, labial view; C, lingual view. Scale: 10 mm. Detailed scanning electronmicroscope views of the dentition; HGL 50-404, RM1: D, occlusal view (reversed); HGL 50bis-56 (holotype), LP4-LM3: E, occlusal view;F, lingual view; G, labial view. Scale: 4 mm.

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mandible is slightly curved ventrally in the anterior part,which is only shallower than the distal part. The symph-ysis is high and extends below the P3. The coronoid crestis clearly vertical. The condyle is partially broken. Themasseteric fossa is deep. There is no diastema between thepremolars.

Lower dentition – Canine. C1: The canine is oval inocclusal outline. The canine is broken at mid-height, butwas surely very high. Its root extends below the P3. Thecanine is projected mesially and labially.

Premolars. P1, P2, P3: According to the preserved alve-oli, P1 is single-rooted (Fig. 2A) and P2 and P3, which seemto have almost the same length, are double-rooted. P2 andP3 are longer than P1, but distinctly shorter than P4.

P4: The P4 is only slightly asymmetric in lateral view. Thetooth is elongated mesiodistally, but it is slightly enlargedtransversally compared with other primitive hyaenodonti-dans. The paraconid is poorly defined and developed. Itspresence is marked by the occurrence of precingulids onmesiolabial and mesiolingual parts of the crown. The proto-conid is high and pointed. The talonid and postfossid arenarrow. Only a hypoconid is labially present. This cuspappears high and secant in lateral view. A vestigial ento-conid is lingually present.

Molars. M1: The M1 is clearly the shortest and narrow-est molar. The trigonid is importantly worn horizontally,so the difference in height between the cuspids cannotbe evaluated. The paraconid is projected mesially, butless than on M2 and M3. The metaconid is alignedtransversally with the protoconid. The talonid is narrowerthan the trigonid, but the postfossid remains wide. Thehypoconulid is not significantly projected distally. Theentoconid is small but present and is lingually located.The hypoconid is the largest cuspid of the talonid.The cristid obliqua is oblique and reaches the trigonidbelow the intraprotocristid notch. An anterior keel (=precingulid) is present, but is short. A small ectocingulid ispresent, but it does not reach the precingulid.

M2: The trigonid is more developed than on M1. Theprotoconid is the highest cusp. The paraconid is higher thanthe metaconid. The trigonid is twice as high as the talonid.The paracristid is long and the paraconid is clearly mesiallylocated. However, the paraconid remains lingually located.The talonid and postfossid are narrower than on M1. Thehypoconulid is slightly more distally located than on M1.The entoconid is more reduced, crestiform and fused withthe hypoconulid than on M1. It is also less lingually located.The anterior keel (= precingulid) is present, but it is short.As observed on M1, the ectocingulid is developed and thereis no postcingulum.

M3: The trigonid is higher than on previous molars. Themetaconid is more reduced than on M1 and M2. The para-conid is higher than on M1 and M2, and is prominentlymesially located. As usually observed on the M3 of hyaen-

odontidans, the talonid is narrow and short. On the worntalonid, the hypoconulid is large and pointed, while theentoconid is crestiform. A short precingulid is present.

Upper dentition. Molars. M1: The specimens HGL 50-404, HGL 50-405 and HGL 50-407 are considered as M1.The stylar shelf and parastylar area are well reduced. Thepostmetacrista is very elongated and secant but it is notsignificantly distally projected. The paracone is higher thanthe metacone but its base is less developed. The paraconeand metacone are connate, but their apices remain sepa-rated. The protocone is mesiodistally short and transver-sally developed. The paraconule and metaconule are small,notably the paraconule. The paraconule is linked to theparastylar area through the paracingulum. No lingual cingu-lum is present.

Comparison. Furodon is only slightly smaller thanGlibzegdouia, but differs by the important reductions of themetaconid and talonid and the fusion of paracone and meta-cone. These features also distinguish the new genus fromthe African genera Dissopsalis, Anasinopa and Masrasec-tor. Furodon also distinguishes from the latter genera andGlibzegdouia by the lingual opening of the talonid (primi-tive feature).

Furodon is morphologically similar to the KoholiinaeBoualitomus and Lahimia in the simplification of themolars. The three genera shared the reduction of the meta-conid, talonid and entoconid. They also share a simi-lar morphology of the paraconid, which is higher andmore mesially located than in primitive hyaenodontidans.However, Furodon, which is younger than the two koholi-ine genera, differs by the presence of a single-rooted P1,whereas the P1 is clearly absent in Koholiinae (Gheerbrantet al. 2006; Sole et al. 2009). Furodon also differs fromKoholia, which is the only known Koholiinae documentedby upper molars (Crochet 1988), by more slightly sepa-rated paracone and metacone and a less developed parasty-lar area. The development of the parastylar area is a peculiarfeature of the Koholiinae. It is related to the developmentof the prevallum–postvallid shearing, which is peculiar toKoholiinae (Sole et al. 2009). The lesser fusion of the para-cone and metacone of Furodon, compared with that ofKoholia, is a primitive feature. Finally, Furodon appearsmore primitive than the Koholiinae, even if, surprisingly, itis younger than the latter genera.

Furodon is morphologically similar to the hyainailourinegenera Pterodon and Akhnatenavus, which are first recordedin the Late Eocene of the Jebel Qatrani Formation (local-ity L-41, Fayum, Egypt). Furodon, Pterodon and Akhnate-navus differ from Koholiinae by the presence of P1.Furodon also shares with Pterodon and Akhnatenavus thereduction of the metaconid, talonid and entoconid. Theyalso share the development of the paraconid, which ismore mesially located than in primitive hyaenodontidans.The reduction of the crushing structures (e.g. talonid) anddevelopment of the secant structures (e.g. paracristid and

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postmetacrista) are adaptations towards a hypercarnivo-rous diet. However, this adaptation is highly convergentamong Hyaenodontida (Polly 1996). Furodon, Pterodonand Akhnatenavus share a reduced M3 talonid relativeto that of M1 and M2. This feature distinguishes thesetaxa from Orienspterodon (Middle Eocene, Myanmar),which was considered by Egi et al. (2007) as the oldesthyainailourine, and is characteristic of the Hyainailouri-nae (Lewis & Morlo 2010). They also share the importantelongation of the postmetacrista on molars.

Furodon differs from Pterodon and Akhnatenavus by thepresence of a slightly basined talonid, which still bearsan entoconid on M1 and M2. The talonid and postfos-sid in Furodon are wider than in Pterodon and Akhnate-navus. Furodon differs significantly from Pterodon andAkhnatenavus by the presence of a distinct metaconid. Theanterior keel is also less developed than in Pterodon andAkhnatenavus. The protocone is more developed and lessmesially located than in the two derived hyainailourines.The paracone and metacone are less fused than in the otherhyainailourine genera. Furodon also differs by the develop-ment of the paraconule and metaconule on upper molars.The postmetacrista is shorter and less distally located thanin Pterodon and Akhnatenavus.

The new genus is closer to Pterodon than to Akhnate-navus in the closely appressed premolars. It also shareswith Pterodon the presence of a transversally enlarged P4.This feature distinguishes also Furodon (and Pterodon)from Metapterodon. Pterodon and Furodon also differ fromAkhnatenavus by a larger talonid on the molars (primi-tive feature), and by P2 and P3 subequal in length (derivedfeature). Furodon differs from earliest Pterodon species:P. phiomensis by a double-rooted P2 (single-rooted in P.phiomensis) and P. africanus by a less reduced P1 (P1 absentor represented only by a shallow alveolus in P. africanus).Finally, the peculiar features of Furodon support the erec-tion of a distinct genus.

Pickford et al. (2008) described Pterodon sp. indet. basedon a maxillary fragment, which bears P4 and M2, and rootsof P3 and M1. The morphology of the P4 supports the refer-ence of the specimen to Hyainailourinae; indeed, converselyto Koholiinae, the postmetacrista is short and low on P4

(see below for a discussion concerning the morphology ofthe P4 among the Koholiinae). Moreover, the upper molarspresent on the Namibian specimen share with Furodon andPterodon the reduced and mesially shifted protocone, andthe distally elongated postmetacrista on the molars. Thespecimen cannot be directly compared with those fromGlib Zegdou. However, as with Furodon, the Namibianspecimen differs from the oldest species of Pterodon andAkhnatenavus from the Jebel Qatrani Formation (Egypt,Fayum), by less reduced protocone area (primitive feature).Pterodon sp. indet., together with Furodon, must be referredto Hyainailourinae. Moreover, the two taxa represent prim-itive hyainailourines.

Parvavorodon gen. nov.

Diagnosis. Same as for the type and only known species.

Etymology. Combination of parva (Latin; small), voro(Latin; to devour) and odon (Latin; tooth).

Parvavorodon gheerbranti sp. nov.(Fig. 3)

Figure 3. Parvavorodon gheerbranti gen. et sp. nov. Scanningelectron micrographs. HGL 04-17, RM1 or RM2: A, occlusal view;B, lingual view; C, labial view. Scale: 1.5 mm.

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Diagnosis. Parvavorodon is a very small hyaenodonti-dan characterized by secant lower molars. Compared withFurodon, the metaconid and entoconid are more reduced,the paraconid is more developed and the talonid is narrower.Parvavorodon is clearly smaller than Furodon.

Holotype. HGL 04-17, RM1 or RM2

Referred specimen. HGL 04-16, LM1 or LM2

Age and locality. HGL 04, Gour Lazib, Algeria, late Ypre-sian or middle Lutetian.

Etymology. Dedicated to Emmanuel Gheerbrant, whogreatly increased our knowledge of the earliest Africanmammals, notably the hyaenodontidans.

Description.

Lower dentition. Molars. M1 or M2: The fossils HGL04-16 and HGL 04-17 are morphologically similar andalmost the same size. The enamel of HGL 04-17 has beenworn out, but the morphology of the tooth is clearly distin-guishable. Because of the short and wide talonid, HGL04-17 could not correspond to a M3. The paraconid is wellprojected mesially. The paraconid is clearly higher thanthe metaconid but smaller than the protoconid, which isthe highest cusp of the trigonid. The talonid is reducedcompared with the trigonid. The talonid and postfossidare narrow. The entoconid is crestiform and fused withthe hypoconulid. The postfossid is opened lingually. Theentocristid is low. The entocristid and cristid obliqua areboth oblique (distally shifted labially). The hypoflexid isnarrow. The precingulid is present, together with a smallectocingulid.

Comparison. HGL 04-16 and HGL 04-17 are puzzlingbecause they are very small and possess very secantmorphology. This morphology, which is characterized bythe important paraconid and the reduction of the talonid,recalls that of the koholiines Lahimia and Boualitomus.However, the presence of a developed ectocingulid (clearlyvisible on HGL 04-16) and wider talonid permit us to distin-guish the fossils of Parvavorodon from those of Koholiinae.

HGL 04-16 and HGL 04-17 differ from the other taxaof the Gour Lazib by a distinctly smaller size. They furtherdiffer from those of Glibzegdouia by a narrower talonidand more reduced metaconid and entoconid. Parvavorodonand Furodon share the reduction of the metaconid (whichremains transversally aligned with the protoconid), themesial localization of the paraconid, and the narrow talonid.These derived features clearly distinguish Parvavorodonfrom the very small Late Palaeocene Tinerhodon fromMorocco.

In addition to its smaller size, Parvavorodon differ fromFurodon by a more mesially located paraconid, a narrowertalonid, and a more reduced entoconid. These features arederived among Hyaenodontida.

As Furodon, Parvavorodon probably lies close to theLate Eocene-Early Oligocene hyainailourine Pterodon andAkhnatenavus rather than to any other hyaenodontidans.All these four genera present adaptations towards hyper-carnivory. Because of the presence of a narrow talonid onParvavorodon compared with Pterodon and Furodon, webelieve that Parvavorodon could represent a morphologi-cal ancestor to Akhnatenavus. Unfortunately, the availablematerial does not allow further and formal evaluations ofthe relationships of Parvavorodon within hyainailourines.

Phylogenetic analysis of the oldest Africanhyaenodontidans

Aim of the phylogenetic analysisThe phylogenetic positions of the African ‘proviverrines’(e.g. Anasinopa, Masrasector and Dissopsalis) amongHyaenodontida have been the subject of debate for a longtime. Holroyd (1994) and Egi et al. (2005) have proposeda close relationship between Anasinopa, Masrasector,Dissopsalis, the European proviverrines (e.g. Eurotherium,Prodissopsalis, Cynohyaenodon and Paracynohyaenodon)and the Asian proviverrines (e.g. Paratritemnodon, Kyaw-dawia and Yarshea). These studies led to the hypothesisof faunal exchanges between Europe, Africa and Asia. Egiet al. (2005) proposed that the ‘Afro-Asian’ hyaenodonti-dans, which had roots in European forms, dispersed towardsthe south and southeast.

However, the phylogenetic tree obtained by Egi et al.(2005) has been criticized by Peigne et al. (2007),who notably underlined that some of the charactersthat defined the ‘Afro-Asian’ proviverrines, such as thesmall M2 metaconid, are homoplastic and convergent.Finally, Peigne et al. (2007) proposed to remove Dissop-salis and Anasinopa from the ‘Afro-Asian proviver-rines’ of Egi et al. (2005) and referred them toHyainailourinae. This left Masrasector as the only possi-ble African ‘proviverrine’ related to the Southeast Asiantaxa; however, Peigne et al. (2007) considered that Masra-sector could be close to Prototomus-like Proviverrinae.Lewis & Morlo (2010) doubtfully included this genusin Hyainailourinae. Paratritemnodon, Kyawdawia andYarshea are grouped under ‘Asian’ Proviverrinae in Peigneet al. (2007). The two groups–Hyainailourinae and ‘Asian’Proviverrinae–are considered to have roots in Arfia. TheEuropean taxa–Proviverra-like Proviverrinae–are consid-ered to be monophyletic (Peigne et al. 2007).

The most recent study focused on the systematics of theearliest hyaenodontidans by Sole (2013) leads to a clarifi-cation of the systematics of earliest Laurasiatic ‘Proviver-rinae’. He proposed separating the ‘Proviverrinae’ intothree groups: Proviverrinae [= Proviverra-like Proviver-rinae of Peigne et al. (2007)], Sinopaninae [= Prototomus-like Proviverrinae of Peigne et al. (2007)] and Arfianinae

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(which only contains the peculiar genus Arfia). However,Sole (2013) did not address questions on relationships of‘Afro-Asian’ hyaenodontidans.

Finally, Grohe et al. (2012) recently performed a phylo-genetic analysis to ascertain the phylogenetic relationshipsof the Apterodontinae. This latter subfamily appears to beclosely related to hyainailourine Metasinopa.

To test the hypotheses provided by the previous authors,we performed a phylogenetic analysis of the oldest Africanhyaenodontidans. We therefore included 29 hyaenodonti-dan taxa, which represent the Hyainailourinae, Proviver-rinae, Sinopaninae, Arfianinae and ‘Afro-Asian’ hyaen-odontidans. We also included Apterodontinae, which areconsidered to appear during the ?Middle Eocene in Africa(Grohe et al. 2012). To limit the study, we did not include allthe representatives of each group. Indeed, the resolution ofthe relationships between all the African taxa is beyond thesubject of this article and only the earliest representativesof each group are included. Finally, we did not include thesubfamily Limnocyoninae, which is clearly monophyleticand apart because of the reduction and loss of the last molars(Gunnell 1998; Morlo & Gunnell 2003, 2005; Sole 2013).

The list of the examined taxa is in Online SupplementalMaterial Appendix 1.

Material and methodsThis analysis is focused on dental characters because of thelack of cranial and postcranial specimens for most of thetaxa studied. The character matrix is derived from those ofPolly (1996), Egi et al. (2005) and Sole (2013) with newcharacter additions and definitions. The data matrix consistsof 60 dental characters (50 binary characters and 10 multi-state characters) (Online Supplemental Material, Appendix2). The maximum number of characters was analysed basedon the earliest species, and the matrix was completed ifnecessary with younger species.

The polarization of the characters was based on outgroupcomparison criteria. As most of the multistate charactersconcern features that are convergent and represent adap-tations towards secant dentition [e.g. length of the post-metacrista (character 47), position of the paraconid (char-acter 23)] or are convergent among Hyaenodontida [e.g.relationship between the paracone and metacone (character49)], they were treated as unordered.

Because ‘Creodonta’ are surely diphyletic, the oxyaen-odontans are not the sister group of the hyaenodonti-dans (Polly 1994, 1996). Therefore, Oxyaenodonta are notused as an outgroup. Instead three cimolestid taxa, Cimo-lestes magnus, Procerberus formicarum and Maelestesgobiensis, were used as outgroups. Following Lillegraven(1969), Cimolestes magnus is considered as the best plau-sible ancestral morphotype of Hyaenodontida. A eutherianmorphotype based on study of Eomaia, Acristatherium andProkennalestes was also included in this analysis.

The data matrix (Online Supplemental Material,Appendix 3) was assembled with WINCLADA (Nixon 1999)and the parsimony analysis was performed using TNT(Goloboff et al. 2008) using a traditional search.

Results

The parsimony analysis yielded three equally parsimonioustrees, with a tree length of 161 steps, consistency index(CI) of 0.42 and retention index (RI) of 0.71. The RI istypical for a matrix mostly comprising dental characters.The CI is weak, probably because of homoplasies. Thestrict consensus tree is 162 steps long (CI 0.42 and RI 0.71)with two collapsed nodes (Fig. 4 depicts the consensus treewith stratigraphical and geographical information).

Bremer supports were calculated for 10 supplementarysteps (Fig. 4). Fifteen nodes have Bremer support >3. Thesubfamilies are well supported as their Bremer supportsare ≥3. It is worth noting that several clades correspond torecently established accepted clades: Proviverrinae (node22), Sinopaninae (node 14), Koholiinae (node 7) and Arfi-aninae (node 25) (Sole 2013). Only the Sinopaninae arepoorly supported. However, this group was well supportedin a previous phylogeny (Sole 2013). Finally, the relation-ships between the subfamilies are poorly supported; theBremer supports are equal to 1.

The African taxa referred to by Lewis & Morlo (2010)to Hyainailourinae are separated in two distinct clades,here named Hyainailourinae (node 4) and Teratodonti-nae (node 18). The Hyainailourinae retains the generaincluded in Hyainailourinae in Holroyd (1994, 1999)except Metapterodon, which is referred to Koholiinae(node 7). The Teratodontinae group the generally African‘Proviverrinae’ (e.g. Holroyd 1994) and the particulargenus Teratodon. The Asian ‘proviverrines’ (e.g. Para-tritemnodon, Kyawdawia and Yarshea) are grouped in anew subfamily: Indohyaenodontinae.

Relationships and morphological evolutionEight distinct subfamilies are distinguished in the presentphylogenetic analysis; among them, seven correspondto already known subfamilies–Proviverrinae, Sinopani-nae, Arfianinae, Koholiinae, Apterodontinae, Teratodonti-nae and Hyainailourinae–whereas one clade correspondsto a new subfamily: Indohyaenodontinae. However, thedefinitions of Teratodontinae and Hyainailourinae impor-tantly differ from the previous studies. Because Sole (2013)is interested in Laurasiatic arfianines, sinopanines andproviverrines, we will not discuss the evolution of thesegroups. This section will focus on African Teratodonti-nae, Koholiinae and Hyainailourinae, and on Asian Indo-hyaenodontinae. We will describe their internal nodes andmorphological evolution. The abbreviations FO and SO

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Hyaenodontidans from the Gour Lazib area 11

Figure 4. Phylogeny of the earliest Hyaenodontida with stratigraphical and geographical information. Tinerhodon is not referred to asubfamily. Teratodon is known in Miocene; Dissopsalis is from Middle Miocene; Anasinopa is from Early to ?Middle Miocene (Lewis& Morlo 2010); Brychotherium is from Late Eocene to Early Oligocene (Holroyd 1994). Abbreviations: Megisto., Megistotherium;Hyainailou., Hyainailourinae; Indohyaeno., Indohyaenodontinae; Ap., Apterodontinae; Ar., Arfianinae. Above line (italic) = Bremersupport; below line = node number.

correspond respectively to Fast Optimization and SlowOptimization.

First, it is worth noting that the genus Tinerhodon, whichis considered as the most primitive hyaenodontidan (Gheer-

brant 1995; Gheerbrant et al. 2006; Sole et al. 2009; Sole2013) is at the base of the consensus tree. This is in agree-ment with its features, which are reminiscent of ‘cimolestid’taxa (Gheerbrant 1995; Gheerbrant et al. 2006).

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Node 2 corresponds to a separation between two distinctclades: (1) Koholiinae + Hyainailourinae, (2) Proviver-rinae + Sinopaninae + Arfianinae + Teratodontinae +Indohyaenodontinae + Apterodontinae. The first groupis notably supported by a shallow postfossid [29(1)], ashort metacingulum [52(1)], and reduced metacone on M3

[58(1)]. These features are all related to the development ofa secant dentition. The second clade is notably supportedby an enlargement of the talonid [30(1)], and develop-ment of the cingulid on molars [34(1)], and separatedparacone and metacone on molars [49(2)]. These featuresare related to the development of an omnivorous diet. Thedistinction between the two groups is therefore correlatedto dietary adaptations. However, as indicated above, theBremer supports for the two clades are weak. Becauseof this, we chose not to create a family based on theserelationships.

The Koholiinae (node 7) are supported by numer-ous derived features: loss of P1 [2(1)], reduced ento-conid [27(2)], reduced exodaenodonty [38(2)], and elon-gated postmetacrista on P4 [45(1)]. Except for the lastone, these features have already been discussed by Soleet al. (2009) and Sole (2013). The position of the Africangenus Metapterodon is interesting, because it groups withthe koholiines rather than with the hyainailourine taxa.Metapterodon differs from other koholiines by the absenceof metaconid on molars [27(2)], and fused paraconeand metacone [49(1)]. Metapterodon thus appears morederived towards a hypercarnivorous diet than the otherkoholiines. The remaining koholiines are supported by theprefossa/postfossid shearing [59(1)] (see Sole et al. 2009).

The hyainailourine clade (node 4) contains the generaAkhnatenavus, Pterodon, Megistotherium, and the newgenus Furodon. The Hyainailourinae are here restricted tothe definition provided by Holroyd (1994, 1999) except forMetapterodon (see above). Node 4 is notably supported bya high hypoconid on P4 [16(1)], protocone developed onP3 [39(2)FO], and paraconid mesially located on molars[23(2)SO]. As for the Koholiinae, the Hyainailourinaeare characterized by several trends towards reductionand simplification of the dentition (e.g. reduction ofthe entoconid) and development of secant features (e.g.development of the paraconid on molars and hypoconidon premolars). The youngest hyainailourines (node 5)are characterized by the loss of the metaconid on molars[26(1)], a three-rooted P3 [40(1)], the fusion of theparacone and metacone on molars [49(1)], and the loss ofmetaconule [53(1)]. These features, which are the resultof adaptations to a hypercarnivorous diet, are convergentwith those of koholiines (e.g. Metapterodon).

It is worth noting that the P1 is lost among Hyainailouri-nae (e.g. Pterodon). This loss is convergent with thatobserved in Koholiinae, which happens during the LatePalaeocene (Sole et al. 2009)–the loss of the P1 occurredafter Middle Eocene in the hyainailourine clade. As noted

by Sole (2013), the reduction of the P1 contributes to short-ening of the dentary and reinforces the power of the canines,which are closer to the condyle in this condition.

The Asian ‘proviverrines’ are grouped with theApterodontinae and Sinopaninae (node 10) based on thepresence of cingulid on P4 [18(1)], talonid not oblique[33(1)], and the presence of a postcingulid on molars[35(1)]. However, these features are convergent in Hyaen-odontida (Sole 2013). The grouping of these subfamiliesmay therefore be due to a lack of knowledge concerningthe oldest representatives of these groups, notably for theApterodontinae and Indohyaenodontinae.

The Asian ‘proviverrines’ (Indohyaenodon, Kyawdawiaand Paratritemnodon) are grouped together (node 11).The features that support the node are notably the fusedlabial cingulids [36(1)], the paraconid well mesially located[23(2) FO/SO], the short metacingulum [52(1)FO], andthe high protocone on molars [57(2)FO]. The dentitionof indohyaenodontines is: (1) reinforced by developmentof the cingulids on molars and premolars: (2) secant dueto the development of paraconid on molars: and (3) punc-turing thanks to the pointed morphology of the protoconeon molars. The combination of these features underlinesthe originality of the dentition of the South and South-eastAsian hyaenodontidans. Moreover, the Indohyaenodontinaeare also characterized by primitive features; these featuresmostly concern the morphology of the P4, which doesnot have a developed protocone and postmetacrista, butdisplays a short but high paracone. We propose to create anew subfamily for this group: Indohyaenodontinae; Indo-hyaenodon is the oldest representative of this group. TheIndohyaenodontinae include genera that are only recordedin South (India, Pakistan) and Southeast (Myanmar)Asia.

The former African ‘proviverrines’ are grouped withthe Arfianinae and Proviverrinae (node 17). This group issupported by a wide postfossid [28(1)], protocone shiftedmesially [42(1)], and paraconid mesially located [23(1)SO].The first two features are the expression of adaptationsto an omnivorous diet, so are possibly convergent (Sole2013).

Node 18 groups the former African ‘proviverrines’ (e.g.Dissopsalis, Anasinopa). It is worth noting that the genusGlibzegdouia, for which we here describe new specimens, isclose to the former African ‘proviverrines’, as discussed inthe Systematics section. The genus Masrasector, for whichthe phylogenetic position has been long debated, is alsoclose to the typical African ‘proviverrines’ in the presentanalysis. The most intriguing taxon present in this group isTeratodon, which is a durophagous adapted genus (Savage1965; Morlo et al. 2007; Lewis & Morlo 2010). Because ofthe presence of Teratodon in this African group, we proposeto use the subfamily name ‘Teratodontinae’ for the entiregroup; this subfamily name has been previously proposedby Savage (1965) for Teratodon only.

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The Teratodontinae (node 18) are notably supportedby a deep postfossid [17(1)], and low trigonid on molars[32(1)]. The Teratodontinae are therefore characterized bythe presence of robust molars. Two distinct groups are foundamong Teratodontinae: (1) Masrasector + Teratodon, and(2) Dissopsalis, Brychotherium and Anasinopa; the genusGlibzegdouia, which is here discussed, does not group witheither clade. This is probably because of the lack of informa-tion concerning this latter genus; indeed, only two lower (M1

and M2) and two upper (P3 and M1) positions are presentlyknown. The first clade (node 19) is supported by P2 as longas P3 [6(1)], P3 as long as P4 [10(1)] and robust premolars[21(1)]. This close relationship is not surprising because arelationship between Teratodon and Masrasector has beenalready envisaged (Savage 1965; Lewis & Morlo 2010).The two genera appear to have developed a durophagousdiet. This adaptation is clearly supported by the unusualmorphology of Teratodon, which is reminiscent of the Euro-pean proviverrine Quercytherium. The second clade (node20) is supported by a secant hypoconid [16(1)], talonidnot oblique [33(1)], loss of the cingulae [54(0)], absenceof metacone on M3 [58(1)], and presence of a cuspid ‘e’[22(1)FO]. These features, which convey adaptations for amore carnivorous diet, are known in other hyaenodontidansubfamilies (Sole 2013). The Miocene genus Dissopsalis,which is characterized by the loss of the metaconid [26(1)]and distinctly mesially located paraconid [23(2)], shows themost derived morphology.

As discussed above, the Teratodontinae are supported bynumerous adaptations (e.g. talonid enlargement, paracone-metacone separation), which clearly distinguishes themfrom Hyainailourinae. Among Hyaenodontida, such adap-tations are known in the Laurasiatic Arfia, American Sinopaand Proviverroides, and European Allopterodon. Theseadaptations (e.g. enlargement of the postfossid) clearlycontrast with the reduction of the metaconid, and elongationof the paracristid and postmetacrista observed in Dissop-salis. These latter features are related to the development ofa hypercarnivorous diet. The combination of two differenttendencies (enlargement of the talonid versus developmentof the paracristid) indicates the peculiar morphology ofDissopsalis. All these features appear to be related to thedevelopment of a hypercarnivorous/durophagous diet simi-lar to that of several carnivoran Borophaginae.

It is worth noting that Arfianinae (node 25) are not closelyrelated to ‘Asian’ (= Indohyaenodontinae) and ‘African’(= Teratodontinae) proviverrines, but to the Proviverrinae(node 21), which are only recorded in Europe (Sole 2013).They therefore do not appear to be at the root of the twolatter subfamilies (contra Peigne et al. 2007). This is notsurprising because Arfianinae are original among Hyaen-odontida as underlined by Sole (2013). Indeed, Arfiani-nae differ from all the other hyaenodontidans in having anarcuate postmetacrista without carnassial notch [45(1)] anddeveloped conules, which display internal crests (postpara-

conule and premetaconule cristae) (Smith & Smith 2001).The Indohyaenodontinae differ notably from Arfianinae bya primitive P4, narrower talonid and reduced entoconid. TheTeratodontinae are notably distinguished from Arfianinaeby the reduced metaconid on molars, the enlarged premo-lars and presence of a carnassial notch on upper molars.Because of the phylogeny and because numerous differ-ences can be identified, the three subfamilies cannot begrouped presently in the same family.

Discussion concerning the systematicsLewis & Morlo (2010) listed five hyaenodontidan subfam-ilies in Africa: Apterodontinae (two genera), Hyainailouri-nae (12 genera), Koholiinae (three genera), Proviverrinae,which only contain Tinerhodon, and Teratodontinae, whichonly includes the enigmatic and durophagous Teratodon.The recent phylogeny of Sole (2013) showed that Tiner-hodon is not a proviverrine taxon.

Metapterodon, which was generally referred toHyainailourinae, appeared at the same time as thehyainailourines Pterodon and Akhnatenavus–its earli-est occurrence is Late Eocene in age (Jebel Quatrani,Egypt; and possibly Qasr El-Sagha, Egypt). The oldestMetapterodon species is smaller than those of Pterodon, butlarger than those of Akhnatenavus and importantly differfrom Pterodon and Akhnatenavus species by numerousderived features such as absence of P1, more fused para-cone and metacone, smaller protocone on upper molars, P4

postparacrista developed into a shearing crest, blade-likeP4 postmetacrista, and double-rooted P3 (Holroyd 1994,1999). The morphology of the P4 therefore importantlydistinguishes Metapterodon from Pterodon and Akhnate-navus: the postmetacrista on P4 of Metapterodon is longand high, whereas the postmetacrista is always short andlow in Hyainailourinae. Among the African hyaenodonti-dans, this peculiar morphology of the P4 is solely known inthe koholiine Koholia. The comparison between Koholiaand Metapterodon indicates that they also share derivedfeatures: trending to a fusion of the paracone and metaconeand reduction of the protocone. Metapterodon shares withother koholiines (Lahimia and Boualitomus) the adaptationstowards a hypercarnivorous dentition such as the reductionof the metaconid and talonid. It is worth mentioning that theearliest koholiines are extremely derived compared with theearliest hyainailourines described in this article–Furodonand Parvavorodon–as is Metapterodon when comparedwith Pterodon and Akhnatenavus. Moreover, Metapterodonshares with the Late Palaeocene Lahimia and Early EoceneBoualitomus the loss of P1, whereas its contemporaneousgenera Pterodon and Akhnatenavus generally still possess aP1. The phylogenetic analysis supports a close relationshipbetween Metapterodon and Koholiinae. Finally, based onthese features and as a result of the cladistic analysis herepresented, we propose to refer Metapterodon to Koholiinae.This new placement importantly extends the stratigraphic

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range of this subfamily from the Late Palaeocene to theEarly Miocene. Koholiinae and Hyainailourinae sharesimilar features such as the reduction of the first premolarsand simplification of the molars, so they were probablycompetitors; however, despite this competition, they wereboth successful in Africa. Finally, a close relationshipbetween the two subfamilies needs to be tested.

The Hyainailourinae were originally defined by hyper-carnivorous genera such as Hyainailouros and Pterodon.Morlo et al. (2007) proposed to refer the genera Buhakia tothe Hyainailourinae because of a similar pattern in tootheruption sequence; this reference implied the referenceof Buhakia’s closest genera (e.g. Dissopsalis, Anasinopa),which were formerly named African ‘proviverrines’ toHyainailourinae (Morlo et al. 2007; Peigne et al. 2007;Lewis & Morlo 2010) Finally, Peigne et al. (2007) andLewis & Morlo (2010) considered only the relationshipsof Masrasector as unresolved, and its systematic positionas uncertain; however, Lewis & Morlo (2010) referred thegenus to Hyainailourinae.

Our phylogenetic study shows that Dissopsalis, Masra-sector, Anasinopa and Glibzegdouia are not closely relatedto the oldest hyainailourine genera (e.g. Pterodon, Akhnate-navus). The group of Dissopsalis and related genera differsdistinctly from the hyainailourine genera closely related toPterodon by numerous features (see above).

Interestingly, the African ‘proviverrine’ genera are closein our cladistic analysis to the enigmatic Teratodon whichis only recorded in the Miocene of Africa. The phyloge-netic position of Teratodon, which displays durophagousadaptations characterized by an important enlargementof the premolars (Savage 1965) was poorly under-stood. Savage (1965) placed this genus at the familylevel–Teratodontindae–but it was later reduced to a subfam-ily of Hyaenodontida by Van Valen (1967). Lewis & Morlo(2010) noted that Teratodon shares with Masrasector andthe European proviverrine Quercytherium the presenceof blunt and large durophagous premolars. Our phylo-genetic analysis provides arguments for a close relation-ship between Teratodon and Masrasector, which are bothfrom Africa. First, the close relationship between Teratodonand African ‘proviverrines’ therefore gives important cluesconcerning the origination of Teratodon, which representsan endemic and particular offshoot of African ‘proviver-rines’. Second, the present topology allows us to referthe African ‘proviverrines’ to the Teratodontinae subfam-ily. The origination of the Teratodontinae can be tracedto the ?Early Eocene because of the presence of Glibzeg-douia. The Teratodontinae now range from ?Early Eoceneto Middle Miocene, and are mostly present in Africa (seebelow).

The global enlargement of the premolars in teratodon-tines probably represents adaptations to durophagy. Severalteratodontines (e.g. Dissopsalis, Buhakia, Mlyanama)developed a hypercarnivorous dentition, which is charac-

terized by the reduction and loss of the metaconid onmolars. The morphology of these teratodontines clearlyrecalls the carnivoran Borophaginae (e.g. ProtomarctusWang et al., 1999, Phlaocyon Matthew, 1899). The ecologyof several teratodontines, as Borophaginae, was probablysimilar to the extant raccoon. Their diets were omnivo-rous/durophagous (insects, plant foods and vertebrates).

In contrast to the Teratodontinae, the Hyainailourinaeonly developed a hypercarnivorous dentition. Indeed, theiradaptations (e.g. important reduction of the metaconid andtalonid, reduction of first premolars) are related to the devel-opment of a strict meat diet. The Hyainailourinae were–withthe Koholiinae–among the major mammalian predators inNorth Africa during the Palaeogene. The origination of thisgroup of predators can now be traced to the Gour Lazibfauna with the presence of the newly described Furodonand Parvavorodon.

The Asian ‘proviverrines’ Paratritemnodon, Kyawdawia,and the recently described Indohyaenodon represent amonophyletic clade in our analysis. This latter clade ischaracterized by the combination of developed cingulids onpremolars, complete labial cingulid on molars, crestiformentoconid on molars, a short and pointed P4 without devel-oped protocone, and occurrence of developed precingu-lum and postcingulum, separated paracone and metacone,reduced protofossa, plus high protocone on upper molars.This group is only recorded in South (India, Pakistan) andSoutheast Asia (Myanmar) and range from Early Eocene toMiddle Eocene (Ranga Rao 1973; Kumar 1992; Egi et al.2004, 2005; Bajpai et al. 2009). Because of their monophylyand restricted distribution, which was probably limited inthe north by the Himalayas, we propose to create a newsubfamily and to name this group as Indohyaenodontinae.

Concerning the Apterodontinae, it is worth noting thatthey are not closely related to Hyainailourinae, in contrastto the recent phylogenetic study of Grohe et al. (2012). Thisdifference could be a result of differences in the includedtaxa.

As a conclusion, it is worth noting that the phylogenyhere proposed and the subsequent systematics are supportedboth geographically and temporally (Figs 4, 5).

Amended classificationThe subfamilies Proviverrinae, Sinopaninae, Arfianinaeand Apterodontinae have already been presented anddescribed elsewhere (Lewis & Morlo 2010; Grohe 2012;Sole 2013).

Order Hyaenodontida Leidy, 1869Family Hyaenodontidae Leidy, 1869

Subfamily Indohyaenodontinae subfam. nov.

Diagnosis. South and Southeast Asian group characterizedby derived features such as the development of cingulids onpremolars and molars, and cingulae on molars, reductionof the entoconid on molars, separation of the paracone and

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Figure 5. Palaeogeographical map of the Early Eocene/Middle Eocene transition (modified from Ron Blakey, Department of Geology,Northern Arizona University, Flagstaff, AZ, http://cpgeosystems.com/050Marect.jpg) showing the distribution of hyaenodontidans records.

metacone and development of a high protocone on molars.Their primitive features mostly concern the P4, which isshort but high, without secant postmetacrista and devel-oped protocone. The talonid of the molars is not reduced orenlarged.

Included genera. Paratritemnodon Ranga Rao, 1973;Yarshea Egi, Holroyd, Tsubamoto, Shigehara, Takai, Aung,Soe & Tun, 2004; Kyawdawia Egi, Holroyd, Tsubamoto,Soe, Takai & Ciochon, 2005; Indohyaenodon Bajpai, Kapur& Thewissen, 2009.

Geographical and stratigraphical range. Early toMiddle Eocene; South (India, Pakistan) and Southeast(Myanmar) of Asia.

Etymology. Derived from oldest genus Indohyaenodon,which has been named in reference to the country of itsorigin and to its familial relations.

Subfamily Teratodontinae Savage, 1965.

Emended diagnosis. Teratodontinae differ fromHyainailourinae by the combination of derived andprimitive features. Their primitive features are: the pres-ence of a wide basined talonid with developed entoconidon molars, and the absence of protocone on P3. The derivedfeatures are: the robustness of the premolars, the widepostfossid on P4; the postfossid lingually closed and wideron molars than in primitive hyaenodontidans; the talonid aswide as the trigonid; the developed ectocingulid on molars;the bulbous and anteriorly directed protocone on P4; theabsence of the parastyle on P4; and the separated paraconeand metacone on molars. As numerous hyaenodontidan

groups, the evolution of the Teratodontinae is characterizedby the development of a secant dentition (e.g. loss ofthe metaconid). The entoconid is reduced in derivedgenera (e.g. Dissopsalis), but the talonid remains wide andlingually closed on M1 and M2 in all Teratodontinae.

Included genera. Dissopsalis Pilgrim, 1910; TeratodonSavage, 1965; Anasinopa Savage, 1965; MasrasectorSimons & Gingerich, 1974; Glibzegdouia Crochet, Peigne& Mahboubi, 2001; Buhakia Morlo, Miller & El-Barkooky.2007; Mlanyama Rasmussen & Guttierrez, 2009.

Geographical and stratigraphical range. Possibly EarlyEocene to late Middle Miocene; Africa and Asia.

Subfamily Koholiinae Crochet, 1988

Emended diagnosis. Koholiinae are clearly distinct fromother hyaenodontidan subfamilies, especially in the short-ening of the anterior lower dentition (e.g. loss of P1

and reduction of P2), the development of the preval-lum/postvallid shearing in addition to the classical special-ized postvallum/prevallid shearing, the early reduction andsimplification of the talonid and elongation of the post-metacrista on P4 (after Sole et al. 2009).

Included genera. Metapterodon, Stromer, 1926; Koho-lia Crochet, 1988; Boualitomus Gheerbrant in Gheerbrantet al., 2006; Lahimia Sole & Gheerbrant in Sole et al. 2009.

Geographical and stratigraphical range. LatePalaeocene to Early Miocene, Africa.

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Discussion

Implications of the hyaenodontidansfrom Glib Zegdou concerning the originand radiation of the HyaenodontidaBased on the present fossil record, the Hyaenodontida areconsidered to have originated either in Africa (Gheerbrant1995; Sole et al. 2009; Sole 2013) or in Asia (Meng et al.1998; Bowen et al. 2002) during the Palaeocene. Therecent discovery of koholiine Lahimia showed that the LatePalaeocene African hyaenodontidans were already morediversified than the hyaenodontidans recorded in Laurasia(Sole et al. 2009). Sole (2013) recently argued for anAfrican origin based notably on the Palaeogene diversityof African hyaenodontidans. The presence of MoroccanLate Palaeocene Tinerhodon at the base of our cladisticstree supports an African origin for the Hyaenodontida.

The data provided by the hyaenodontidans from GlibZegdou (late Ypresian or middle Lutetian) are impor-tant because they give clues concerning the radiation anddiversification of the Hyaenodontida in Africa. Indeed, thestudy of Glibzegdouia, Furodon and Parvavorodon, andthe phylogenetic analysis clearly demonstrate that bothTeratodontinae and Hyainailourinae originated in Africaduring the beginning of the Eocene (Fig. 4). This agreeswith the hypothesis of Peigne et al. (2007) and disagreeswith that of Egi et al. (2007); it is worth mentioningthat we follow Lewis & Morlo (2010) in considering thatOrienspterodon belongs to the Hyaenodontinae rather thanto the Hyainailourinae. Moreover, the Teratodontinae andHyainailourinae, which range from the ?Early Eocene tothe Middle Miocene, mostly radiated in Africa.

At least three subfamily rank groups were presentin Africa during the Early/Middle Eocene transi-tion: Teratodontinae, Hyainailourinae and Koholiinae(Fig. 5). The diversity of the subfamilies (Teratodonti-nae, Hyainailourinae) in the Gour Lazib area in earli-est part of Eocene implies that these groups separated atleast during the earliest part of the Early Eocene. TheHyaenodontida appear to have diversified ecologically inAfrica at this time: the Teratodontinae represent an omniv-orous to durophagous/hypercarnivorous group, Koholiinaeand Hyainailourinae both correspond to hypercarnivoroushyaenodontidans. Our study therefore provides evidence foran unexpected diversity of the Hyaenodontida in Africa atleast since the Middle Eocene.

It is worth noting that the evolution of the Apterodon-tinae is now better known (Grohe et al. 2012). The oldestApterodontinae (?Middle Eocene) already possess impor-tantly derived dentition and postcranium. Moreover, thenew fossils suggest a semi-aquatic lifestyle and a dietbased on marine invertebrates for Apterodon (Grohe et al.2012). The peculiar adaptations of Apterodontinae there-fore imply an Early or at least Middle Eocene origin for thegroup.

Following the argument of Sole (2013), we think thatthe important diversity of Hyaenodontida in Africa duringthe first half of the Eocene could only be explained by anAfrican origin for the group.

Moreover, the presence of Teratodontinae andHyainailourinae in both Black Crow locality (Sperrgebiet,Namibia) and Gour Lazib area indicates that they werealready widespread in Africa at least during the MiddleEocene.

Finally, the description of the diversity of Hyainailouri-nae in Glib Zegdou supports an important–but presentlyunderestimated–African radiation of the Hyaenodontida asproposed previously (Gheerbrant 1995; Sole 2013).

Palaeobiogeographic implicationsThe phylogenetic tree shows that the Hyaenodontida prob-ably experienced a Palaeocene/Early Eocene phase of radi-ation. This phase has been followed by endemic evolution,which often results in convergences. These convergencescan significantly perturb the phylogenetic reconstructionsand determination of the geographic origin of Hyaenodon-tida. However, as indicated above, numerous argumentssupport an African origin for this group.

Sole (2013) recently discussed the dispersions ofSinopaninae, Proviverrinae and Limnocyoninae. We there-fore only discuss here the cases of Indohyaenodontinae,Apterodontinae, Hyainailourinae and Teratodontinae.

As indicated above, the indohyaenodontines Para-tritemnodon, Kyawdawia, Yarshea and Indohyaenodon,which represent a monophyletic clade in our analysis, areonly recorded in South and Southeast Asia. The recentdiscovery of the genera Indohyaenodon in the Indian local-ity of Vastan, which dates from the beginning of the EarlyEocene (Bajpai et al. 2009; Clementz et al. 2010) supportsa separation of the indohyaenodontines from the Africanhyaenodontidans at least during the latest part of the LatePalaeocene. There is however little other evidence for sucha mammalian dispersal event between Africa and India. Theonly putative direct evidence could concern the adapisori-culid ‘insectivores’, which may have dispersed from Indiato Africa at the beginning of the Tertiary and before theLate Palaeocene (Prasad et al. 2010). The subsequent EarlyEocene partial isolation of India could have favoured theendemic evolution of the Indohyaenodontinae in India.

Gheerbrant & Rage (2006) noted that the history of trans-Tethyan Palaeogene mammal dispersals is characterizedby at least four dispersal phases: during the Early Thane-tian, and by the Thanetian/Ypresian, Bartonian/Priabonianand Priabonian/Rupelian transitions. Two other dispersalsbetween Africa and Laurasia (by the Ypresian/Lutetianand by the Lutetian/Bartonian transitions) are presentlyless supported. The dispersals principally occurred betweenAfrica and Europe via the Iberian route or Apulian route.

The presence in Europe of the genera Apterodon andPterodon during the Late Eocene and Early Oligocene

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respectively indicates that these hyaenodontidans dispersedfrom Africa to Europe. The oldest European recordof Pterodon (Pterodon dasyuroides) is Late Eocene inage (MP18, La Debruge) (Lange-Badre 1979). BecausePterodon is unknown in Asia, we propose that Pterodonprobably dispersed from Africa to Europe via the Iberianroute or Apulian route during the Bartonian/Priaboniantransitions. However, as noted by Gheerbrant & Rage(2006), the Bartonian/Priabonian Dispersal Phase, which ischaracterized by the immigration of the anthracothere artio-dactyls, and the ‘baluchimyine’ rodent Protophiomys intoAfrica, is only southward. Hence, Pterodon could representthe sole case of northward dispersal.

Apterodon appeared in Europe probably during the EarlyOligocene. Lange-Badre & Bohme (2005) proposed threemigration routes for explaining the presence of Apterodonin Europe: Apulian route, Iberian route and East–Westdispersal route. The last migration way is based on ahypothetical Asian origin of Apterodon. However, the newdiscoveries from Libya reinforce the hypothesis of anAfrican origin for Apterodontinae (Grohe et al. 2012).Because Apterodon is unknown in Asia and Arabia, wepropose that Apterodon probably dispersed via the Iberianroute or Apulian route as did Pterodon. However, the disper-sal of Apterodon occurred during the Priabonian/Rupeliantransition. Gheerbrant & Rage (2006) noted that the disper-sal of Apterodon into Europe is the sole case of northwarddispersal during the Priabonian/Rupelian Dispersal Phase.The recent work of Grohe et al. (2012) supports this disper-sal event.

The new classification of the African hyaenodontidansimplies that the species Dissopsalis carniflex is the solerepresentative of the Teratodontinae in Asia; the genus isrecorded in Pakistan (Barry 1988). The African origin ofTeratodontinae implies that Dissopsalis migrated at leastfrom Africa to Asia during the Middle Miocene or before.As the genus is unknown in Europe, it probably dispersedvia the Iranian route, which comprises the Iranian block andArabian Peninsula. Gheerbrant & Rage (2006) noted thatthis route was possibly active since the Middle Eocene.

Biostratigraphic implications of thehyaenodontidans from the Gour LazibThe hyaenodontidan fauna from the Gour Lazib is signifi-cantly diversified: the three genera (Glibzegdouia, Furodonand Parvavorodon) represent two subfamilies (Teratodon-tinae and Hyainailourinae). This diversity clearly exceedsthat from the Palaeocene locality of Adrar Mgorn (Tiner-hodon), the Palaeocene-Eocene localities of Ouled AbdounBasin (Lahimia is reported from the Late Palaeocene;Boualitomus is reported from the Early Eocene), the EarlyEocene locality of El Kohol (Koholia) and the ?MiddleEocene locality of Dor El Talha (Apterodon). However, it isworth mentioning that the ?Lutetian locality of Black Crow

(Sperrgebiet, Namibia) yielded taxa that can be referred toTeratodontinae and Hyainailourinae (see above). However,the Algerian locality is slightly more diversified than theNamibian locality (three taxa versus two taxa).

The specific and high rank diversities of Glib Zegdouare clearly less than that of the Fayum (Jebel QatraniFormation, Qsar el Sagha Formation and Birket QarunFormation, Late Eocene to Oligocene, Egypt). In the latterlocalities, seven genera are recorded (plus one proviver-rine not yet formally described; see Borths et al. 2010).The genera are referred to Teratodontinae (Masrasector),Hyainailourinae (Metasinopa, Pterodon, Akhnatenavus),Apterodontinae (Apterodon, Quasiapterodon) and Koholi-inae (Metapterodon). The Gour Lazib and Fayum sites shareonly the presence of Teratodontinae and Hyainailourinae(Fig. 4).

The three genera recorded in the Gour Lazib appearclearly smaller and more primitive than those of the Fayum.This agrees with the antiquity of the Algerian sites (lateYpresian or middle Lutetian). Moreover, the presence ofclosely related genera in the Gour Lazib and Fayum under-lines an apparent continuity of the fauna in the NorthernAfrican area during the Eocene. The faunal continuity inNorth Africa during Palaeogene is also supported by thereference of Metapterodon to Koholiinae proposed in thisarticle (Fig. 4).

Surprisingly, the fauna from the Gour Lazib appearscloser to the Black Crow locality (Namibia) than to thoseof El Kohol and Ouled Abdoun Basin (Late Palaeoceneto Early Eocene). Pickford et al. (2008) already noted thestriking similarities of the faunas from North Africa andNamibia. These similarities suggest biogeographic rela-tionships between the two areas. Finally, the fauna fromthe Gour Lazib appears also close to that of the Fayum.However, Glib Zegdou is distinguished by the absenceof Koholiinae and Apterodontinae. This absence in GlibZegdou could be explained by environmental differences(e.g. prey availability).

Conclusions

The Algerian localities of the Gour Lazib area (Early orearly Middle Eocene) have yielded three hyaenodontidans:Glibzegdouia, is now clearly referred to the Hyaenodon-tida, while Parvavorodon and Furodon represent twonew taxa. Their discovery increases our knowledge ofthe earliest hyaenodontidans and shows an unexpecteddiversity for this group in Africa. Indeed, Glibzegdouiaappears to be the first representative of the Teratodontinae,which now include several African genera such as Masra-sector and Anasinopa. On the other side, Parvavorodonand Furodon represent the earliest hyainailourines everdiscovered. Moreover, the study of the fossils from GourLazib supports an African origin of Hyaenodontida. The

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diversity of African hyaenodontidans from the earliestpart of Eocene clearly supports a Late Palaeocene-EarlyEocene radiation of this group in Africa.

Our study significantly fills the gap concerning the radia-tion of the African hyaenodontidans. However, the relation-ships between the different subfamilies of Hyaenodontidaare still poorly resolved. The establishment of these rela-tionships will allow us to better reconstruct the radiationand dispersions of the Hyaenodontida.

Acknowledgements

We thank the vice-chancellor of Tlemcen University, andthe authorities from Bechar and Tindouf districts, whoassisted with fieldwork in the Gour Lazib area. We thankL. Marivaux, L. Hautier, R. Lebrun, A. Mahboubi and A.-L. Charruault for their help. We are grateful to C. Caze-vieille for access to the scanning electron microscope ofthe Centre de Ressources en Imagerie Cellulaire in Mont-pellier. This research was supported by the French ANR-PALASIAFRICA Program (ANR-08-JCJC-0017).

Supplemental Material

Supplemental material is available online DOI:10.1080/14772019.2013.795196

References

Adaci, M., Tabuce, R., Mebrouk, F., Bensalah, M., Fabre,P.-H., Hautier, L., Jaeger, J.-J., Lazzari, V., Mahboubi,M., Marivaux, L., Otero, O., Peigne, S. & Tong, H. 2007.Nouveaux sites a vertebres paleogenes dans la region des GourLazib (Sahara nord-occidental, Algerie). Comptes Rendus dePalevol, 6, 535–544.

Bajpai, S., Kapur, V. V. & Thewissen, J. G. M. 2009. Creodontand condylarth from the Cambay Shale (Early Eocene), Vastanlignite mine, Gujarat, Western India. Journal of the Palaeon-tological Society of India, 54, 103–109.

Barry, J. C. 1988. Dissopsalis, a Middle and Late Mioceneproviverrine creodont (Mammalia) from Pakistan and Kenya.Journal of Vertebrate Paleontology, 8, 25–45.

Biedermann, W. G. A. 1863. Petrefacten aus der Umgegendvon Winterthur: II. Die Braunkohlen von Elgg. Anhang:Hyainailouros sulzeri. Bleuler-Hausheer, Wintherthur, 23 pp.

Blainville, H. M. D. de 1839. Sur les insectivores fossilesd’Auvergne. Annales Francaises et Etrangeres d’Anatomieet de Physiologie, 3, 60.

Borths, M., Seiffert, E., Goodenberger, K. & Simons, E. 2010.The oldest Fayum creodont: dental and humeral morphol-ogy of a new proviverrine hyaenodontid from the earliestLate Eocene of Egypt. Journal of Vertebrate Paleontology, 29(supplement), 63A.

Bowen, G. J., Clyde, W. C., Koch, P. L., Ting, S., Alroy, J.,Tsubamoto, T., Wang, Y. & Wang, Y. 2002. Mammaliandispersal at the Paleocene/Eocene boundary. Science, 295,2062–2065.

Clementz, M., Bajpai, S., Ravikant, V., Thewissen, J. G. M.,Saravanan, N., Singh, I. B. & Prasad, V. 2010. Early Eocenewarming events and the timing of terrestrial faunal exchangebetween India and Asia. Geology, 39, 15–18

Cope, E. D. 1885. The White River beds of Swift Current River,Northwest Territory. American Naturalist, 19, 163.

Costeur, P., Benammi, M., Mahboubi, M., Tabuce, R., Adaci,M., Marivaux, L., Bensalah, M., Mahboubi, S., Mahboubi,A., Mebrouk, F., Maameri, C. & Jaeger, J.-J. 2012.Chronology of the Eocene continental deposits of Africa:Magnetostratigraphy and biostratigraphy of the El Koholand Glib Zegdou Formations, Algeria. Geological Society ofAmerica Bulletin, doi: 10.1130/B30565.1.

Crochet, J.-Y. 1988. Le plus ancien creodonte africain: Koholiaatlasense nov. gen., nov. sp. (Eocene inferieur d’El Kohol,Atlas saharien, Algerie). Comptes Rendus de l’Academie desSciences, Serie II, 307, 1795–1798.

Crochet, J.-Y., Peigne, S. & Mahboubi, M. 2001. Anciennete desCarnivora (Mammalia) en Afrique. Pp. 91–100 in C. Denys,L. Granjon & A. Poulet (eds) African Small Mammals. Petitsmammiferes africains. IRD Publishers, Paris.

Egi, N., Holroyd, P. A., Tsubamoto, T., Shigehara, N., Takai,M., Tun, S. T., Aye Ko Aung A. K. & Soe A. N. 2004. Anew genus and species of hyaenodontid creodont from thePondaung Formation (Eocene, Myanmar). Journal of Verte-brate Paleontology, 24, 502–506.

Egi, N., Holroyd, P. A., Tsubamoto, T., Soe, A. N., Takai,M. & Ciochon, R. L. 2005. Proviverrine hyaenodontids(Creodonta, Mammalia) from the Eocene of Myanmar anda phylogenetic analysis of the proviverrines from the para-Tethys area. Journal of Systematic Paleontology, 34, 337–358.

Egi, N., Tsubamoto, T. & Takai, M. 2007. Systematic status ofAsian “Pterodon” and early evolution of hyaenaelurine hyaen-odontid creodonts. Journal of Paleontology, 81, 770–778.

Gheerbrant, E. 1995. Les mammiferes paleocenes du Bassind’Ouarzazate (Maroc) III. Adapisoriculidae et autresmammiferes (Carnivora, ?Creodonta, Condylarthra, ?Ungu-lata et incertae sedis). Palaeontographica, Abteilung A, 237,39–132.

Gheerbrant, E. & Rage, J.-C. 2006. Paleobiogeography ofAfrica: How distinct from Gondwana and Laurasia? Palaeo-geography, Palaeoclimatology, Palaeoecology, 241, 224–246.

Gheerbrant, E., Iarochene, M., Amaghzaz, M. & Bouya, B.2006. Early African hyaenodontid mammals and their bearingon the origin of the Creodonta. Geological Magazine, 134,475–489.

Gingerich, P. D. & Deutsch, H. A. 1989. Systematics andevolution of Early Eocene Hyaenodontidae (Mammalia,Creodonta) in the Clarks Fork Basin, Wyoming. Contribu-tions from the Museum of Paleontology, 27, 327–391.

Goloboff, P. A., Farris, J. S. & Nixon, K. C. 2008. TNT, a freeprogram for phylogenetic analysis. Cladistics, 24, 774–786.

Grohe, C., Morlo, M., Salem, M. & Jaeger, J.-J.. 2010. Theoldest Apterodontinae (Hyaenodontidae, “Creodonta”) fromthe Middle Eocene of Dor El Talha (Libya). Journal of Verte-brate Paleontology, 29 (supplement), 99A.

Grohe, C., Morlo, M., Chaimanee, Y., Blondel, C., Coster, P.,Valentin, X., Salem, M., Bilal, A. A., Jaeger, J.-J. & Brunet,M. New Apterodontinae (Hyaenodontida) from the EoceneLocality of Dur At-Talah (Libya): Systematic, Paleoecologi-cal and Phylogenetical Implications. PLoS ONE, forthcoming.

Grohe, C., Morlo, M., Chaimanee, Y., Blondel, C., Coster,P., Valentin, X., Salem, M., Bilal, A. A., Jaeger, J.-J.,& Brunet, M. 2012. New Apterodontinae (Hyaenodon-tida) from the Eocene Locality of Dur At-Talah (Libya):Systematic, Paleoecological and Phylogenetical Implications.

Dow

nloa

ded

by [

Nor

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akot

a St

ate

Uni

vers

ity]

at 0

2:53

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Aug

ust 2

013

Hyaenodontidans from the Gour Lazib area 19

PLoS ONE 7, 1–19. e49054. doi:10.1371/journal.pone.0049054

Gunnell, G. F. 1998. Creodonta. Pp. 91–109 in C. M. Janis,K. M. Scott & L. L. Jacobs (eds) Evolution of the TertiaryMammals of North America. Volume 1: Terrestrial carnivores,ungulates, and ungulatelike mammals. Cambrige UniversityPress, Cambridge.

Holroyd, P. A. 1994. An examination of dispersal origins forFayum Mammalia. PhD thesis, Duke University, 328 pp.

Holroyd, P. A. 1999. New Pterodontinae (Creodonta: Hyaenodon-tidae) from the late Eocene-early Oligocene Jebel QatraniFormation, Fayum province, Egypt. PaleoBios, 19, 1–18.

Jaeger, J.-J., Beard, K. C., Chaimanee, Y., Salem, M.,Benammi, M., Hlal, O., Coster, P., Awad, A., Bilal,Duringer, P., Schuster, M., Valentin, X., Marandat, B.,Marivaux, L., Metais, E., Hammuda, O. & Brunet, M.2010a. Late middle Eocene epoch of Libya yields earli-est known radiation of African anthropoids. Nature, 467,1095–1098.

Jaeger, J.-J., Marivaux, L., Salem, M., Abolhassan Bilal, A.,Benammi, M., Chaimanee, Y., Duringer, P., Marandat, B.,Metais, E., Schuster, M., Valentin, X. & Brunet, M. 2010b.New rodent assemblages from the Eocene Dur At-Talahescarpment (Sahara of central Libya): systematic, biochrono-logical, and palaeobiogeographical implications. ZoologicalJournal of the Linnean Society, 160, 195–213.

Kumar, K. 1992. Paratritemnodon indicus (creodonta:Mammalia) from the early Middle Eocene Subathu Forma-tion, NW Himalaya, India, and the Kalakot mammaliancommunity structure. Palaontologische Zeitschrift, 66,387–403.

Lange-Badre, B. 1979. Les creodontes (Mammalia) d’Europeoccidentale de l’Eocene superieur a l’Oligocene superieur.Memoires du Museum National d’Histoire Naturelle, 42,1–249.

Lange-Badre, B. & Bohme, M. 2005. Apterodon intermedius, sp.nov., a new European creodont mammal from MP22 of Espen-hain (Germany). Annales de Paleontologie, 91, 311–328.

Lewis, M. E. & Morlo, M. 2010. Chapter 26 – ‘Creodonta’.Pp. 543–560 in L. Werdelin & W. Sanders (eds) Cenozoicmammals of Africa. University of California Press, Berkeley.

Lillegraven, J. 1969. Latest Cretaceous mammals of upper partof Edmonton Formation of Alberta, Canada, and reviewof marsupial-placental dichotomy in mammalian Edmon-ton. University of Kansas Paleontological Contributions, 50,1–122.

Martin, R. 1906. Revision der obereocaenen und unteroligo-caenen Creodonten Europas. Revue Suisse de Zoologie, 14,405–600.

Matthew, W. D. 1899. A provisional classification of the fresh-water Tertiary of the West. Bulletin of the American Museumof Natural History, 12, 19–75.

Meng, J., Zhai, R. & Wyss, A. R. 1998. The Late PaleoceneBayan Ulan fauna of Inner Mongolia, China. Pp. 148–185in K.C. Beard & M.E. Dawson (ed.) Dawn of the Age ofMammals in Asia. Carnegie Museum of Pittsburgh, Pitts-burgh.

Morlo, M. 1999. Niche structure and evolution in creodont(Mammalia) faunas of the European and North AmericanEocene. Geobios, 32, 397–305.

Morlo, M. & Gunnell, G. F. 2003. Small limnocyonines (Hyaen-odontidae, Mammalia) from the Bridgerian Middle Eoceneof Wyoming: Thinocyon, Prolimnocyon, and Iridodon, newgenus. Contributions from the Museum of Paleontology, 31,43–78.

Morlo, M. & Gunnell, G. F. 2005. New species of Limno-cyon (Mammalia, Creodonta) from the Bridgerian (middleEocene). Journal of Vertebrate Paleontology, 25, 247–251.

Morlo, M., Miller, E. R. & El-Barkooky, A. N. 2007. Creodontaand Carnivora from Wadi Moghra, Egypt. Journal of Verte-brate Paleontology, 27, 145–159.

Nixon, K. C. 1999.Winclada (BETA) ver. 0.9.9. Published by theauthor, Ithaca, NY.

Osborn, H. F. 1909. New carnivorous mammals from the FayumOligocene, Egypt. Bulletin of the American Museum of Natu-ral History, 26, 415–424.

Pilgrim, G. E. 1910. Preliminary note on a revised classificationof the Tertiary freshwater deposits of India. Records of theGeological Survey of India, 40, 185–205.

Peigne S., Morlo, M., Chaimanee, Y., Ducrocq, S., Tun, S.T. & Jaeger, J.-J. 2007. New discoveries of hyaenodon-tids (Creodonta, Mammalia) from the Pondaung Formation,Middle Eocene, Myanmar – paleobiogeographic implications.Geodiversitas, 29, 441–458.

Pickford, M., Senut, B., Morales, J., Mein, P. & Sanchez, I. M.2008. Mammalia from the Lutetian of Namibia. Memoir ofthe Geological Survey of Namibia, 20, 465–514.

Polly, P. D. 1994. What if anything is a creodont? Journal ofVertebrate Paleontology, 14 (supplement), 42A.

Polly, P. D. 1996. The skeleton of Gazinocyon vulpeculus gen. etcomb. nov. and the cladistic relationships of Hyaenodontidae(Eutheria, Mammalia). Journal of Vertebrate Paleontology,16, 303–319.

Prasad, G. V. R., Verma, O., Gheerbrant, E., Goswami, A.,Khosla, A., Parmar, V. & Sahni, A. 2010. First mammalevidence from the Late Cretaceous of India for biotic disper-sal between India and Africa at the KT transition. ComptesRendus de Palevol, 9, 63–71.

Ranga Rao, A. 1973. Notices of two new mammals from theUpper Eocene, Kalakot beds, India. ONGC special paper, 2,1–6.

Rasmussen, D. T. & Guttierrez, M. 2009. A mammalian faunafrom the Late Oligocene of Northwestern Kenya. Palaeonto-graphica Abteilung A, 288, 1–52.

Sallam, H. M., Seiffert, E. R. & Simons, E. L. 2011. Cran-iodental Morphology and Systematics of a New Familyof Hystricognathous Rodents (Gaudeamuridae) from theLate Eocene and Early Oligocene of Egypt. PLoS ONE, 6,e16525.

Savage, R. J. G. 1965. Fossil mammals of Africa: 19. The MioceneCarnivora of East Africa. Bulletin of the British Museum ofNatural History, 10, 239–316.

Savage, R. J. G. 1973. Megistotherium, gigantic hyaenodontfrom Miocene of Gebel Zelten, Libya. Bulletin of the BritishMuseum (Natural History), Geology, 22, 485–511.

Simons, E. L. & Gingerich, P. D. 1974. New carnivorousmammals from the Oligocene of Egypt. Annals of the Geolog-ical Survey of Egypt, 4, 157–166.

Smith, T. & Smith, R. 2001. The creodonts (Mammalia, Ferae)from the Paleocene-Eocene transition in Belgium (TienenFormation, MP7). Belgian Journal of Zoology, 131, 117–135.

Sole, F. New proviverrine genus from the Early Eocene of Europeand the first phylogeny of Late Palaeocene–Middle Eocenehyaenodontidans (Mammalia). forthcoming.

Sole, F. 2013. New proviverrine genus from the Early Eocene ofEurope and the first phylogeny of Late Palaeocene–MiddleEocene hyaenodontidans (Mammalia). Journal of SystematicPalaeontology, 11, 375–398.

Sole, F., Gheerbrant, E., Mbarek, A. & Baadi, B. 2009.Further evidence of the African antiquity of hyaenodontid

Dow

nloa

ded

by [

Nor

th D

akot

a St

ate

Uni

vers

ity]

at 0

2:53

17

Aug

ust 2

013

20 F. Sole et al.

(“Creodonta”, Mammalia) evolution. Zoological Journal ofthe Linnean Society, 156, 827–846.

Stromer, E. 1926. Reste land-und susswasser-bewohnenderWirbeltiere aus dem Diamentenfelden Deutsch-Sudwest-afrikas;. Pp. 107–153 in E. Kaiser (ed.) Die DiamentenwusteSudwestafrikas, vol. 2. Ditrich Reimer, Berlin.

Szalay, F. S. 1967. The affinities of Apterodon (Deltatheridia,Hyaenodontdae). American Museum Novitates, 2293, 1–17.

Van Valen, L. 1966. Deltatheridia, a new order of mammals.Bulletin of the American Museum of Natural History, 132,1–126.

Van Valen, L. 1967. New Paleocene insectivores and insectivoreclassification. Bulletin of the American Museum of NaturalHistory, 135, 217–284.

Wang, X., Tedford, R. H., & Taylor, B. E. 1999. Phyloge-netic systematics of the Borophaginae (Carnivora: Canidae).Bulletin of the American Museum of Natural History, 243,1–391.

Werdelin, L. & Peigne, S. 2010. Carnivora. Pp. 603–657in L. Werdelin & W. Sanders (eds) Cenozoicmammals of Africa. University of California Press,Berkeley.

Dow

nloa

ded

by [

Nor

th D

akot

a St

ate

Uni

vers

ity]

at 0

2:53

17

Aug

ust 2

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