ORIGINAL PAPER

The oldest African fox (Vulpes riffautae n. sp., Canidae,Carnivora) recovered in late Miocene deposits of the Djurabdesert, Chad

Louis de Bonis & Stéphane Peigné & Andossa Likius &

Hassane Taïsso Mackaye & Patrick Vignaud &

Michel Brunet

Received: 19 October 2006 /Revised: 7 February 2007 /Accepted: 15 February 2007 /Published online: 15 March 2007# Springer-Verlag 2007

Abstract We report on the oldest fox (Canidae) ever foundin Africa. It is dated to 7 Ma based on the degree ofevolution of the whole fauna. It belongs to a new species.Its overall size and some morphological characteristicsdistinguish the Chadian specimen from all the other foxes.The presence of Vulpes and of the genus Eucyon in slightlyyounger African locality, as well as in southwestern Europein the late Miocene, may indicate that canids migrated inEurope from Africa through a trans-Mediterranean route.

Keywords Mammalia . Carnivora . Canidae . Africa .

Late Miocene

Introduction

The carnivoran mandible described in this article wasrecovered from the fossil vertebrate-bearing localities ofthe anthracotheriid unit from Toros-Menalla in the Djurabdesert, northwest of N’Djamena (Chad). During the lastdecade, more than four hundred fossiliferous sites have beendiscovered in this area. These have yielded a large number offossil vertebrates (Vignaud et al. 2002; de Bonis et al. 2005;Peigné et al. 2005a,b) and significant remains of thehominid Sahelanthropus tchadensis (Brunet et al. 2002,

2005). The fossil-bearing sediments correspond to thefilling of the Chad basin with freshwater deposits yieldinga rich vertebrate fauna that is very different from the extantone. The Toros-Menalla deposits are situated in the intra-cratonic Chad basin. The sedimentary context consists ofrepeated aeolian/lacustrine sequences indicating periodicityof dry and humid conditions (Schuster et al. 2006). Thevertebrate remains are recorded from the lacustrine depositswith several species of fishes, aquatic and terrestrial turtles,snakes, lizards, crocodiles, birds and about thirty species ofmammals in TM 266. The biochronologic data allow for aprecise dating of the localities. The suid Nyanzachoerussyrticus is associated with the anthracotheriid Libychosauruspetrochii; the latter is absent in the localities of the easternDjurab dated to the early Pliocene, such as KossomBougoudi. The third molars (M3-m3) of Nyanzachoerusare similar to the fossils of the lower member of the NawataFormation of Lothagam, which is radio-metrically dated to7.4–6.5 Ma (McDougall and Feibel 1999). In terms of thedegree of evolution, the proboscidean dentition is a little moreprimitive,soolder,thanthatwhichexistsinthecorrespondingfossilsof the Lukeino Formation radio-metrically dated to about 6Ma(Deino et al. 2002). The species of Hipparion is similarto a middle Turolian sample from the Arabian Peninsula.The other mammalian taxa are compatible with a geologicalage of late Miocene about 7 Ma. The Toros-Menalla faunacontains several carnivore taxa belonging to Felidae,Hyaenidae, Viverridae, Herpestidae, and Mustelidae but onlyone specimen of canid.

The family Canidae originated in North America andevolved in that continent from the late Eocene to the present(Wang et al. 2004). The canid from Toros-Menalla ispotentially the oldest canid in the Old World, as it may beolder than “Canis” cipio (closely related to Eucyon; Wang

Naturwissenschaften (2007) 94:575–580DOI 10.1007/s00114-007-0230-6

L. de Bonis (*) : S. Peigné : P. Vignaud :M. BrunetLaboratoire de Géobiologie, Biochronologie et Paléontologiehumaine, UMR 6046 CNRS, Université de Poitiers,86022 Poitiers cedex, Francee-mail: louis.debonis@univ-poitiers.fr

A. Likius :H. T. MackayeUniversité de N’Djamena,BP 1117 N’Djamena, Chad

et al. 2004) from late Miocene localities (middle Turolian,MN 12) of Spain dated 6.9–6.8 Ma (van der Made et al.2006). The species from Toros-Menalla is certainly the oldestcanid recovered in Africa, and it represents the onset of thedevelopment of the extant canid fauna in this continent; theChadian record is also the oldest one for foxes in the OldWorld.

Materials and methods

The material consists of two mandibular corpi of a singleindividual (TM 32-04-78), the right one with the alveolusof the canine and the p1 and p2-m3, the left one with thecanine root, broken p1, broken p2, p3, whose distal half ismissing, and p4-m3. Size and morphology indicate a canidclose to the foxes. First, the specimen will be describedand then compared to extant or fossil foxes of similar size.The measurements were taken with a vernier calliper to1/100 mm and given to 1/10 mm.

Systematics

Carnivora Bowdich, 1821Canoidea Fischer von Waldheim, 1817Canidae Fischer von Waldheim, 1817Caninae Fischer von Waldheim, 1817Vulpini Hemprich and Ehrenberg, 1832Genus Vulpes Frisch, 1775Vulpes riffautae nov. sp.Origin of the name: dedicated to our colleague SabineRiffaut for her fine drawingsHolotype: two mandibular corpi TM 32-04-78 housed inthe Centre National d’Appui à la Recherche (CNAR),N’Djamena, ChadType locality: Toros-Menalla 32, Djurab desert, ChadLevel and geological age: Anthracotheriid unit, lateMioceneDiagnosis: Vulpes with size between that of V. zerda andV. rueppellii; lower premolars p3 and p4 with tiny posterioraccessory cuspids (PACs) and separated by large diastemas;m1 with metaconid a little less reduced than in extantspecies, no crest between hypoconid and entoconid; noentoconulids in m1 or m2; m2 with occlusal outline morerectangular than trapezoidal, rather long talonid comparedto trigonid, weak mesio-buccal cingulum; m3 with rela-tively large mesial and distal foveae.

Description

The mandible is shallow and slender (Table 1). Thesymphysis ends just before the posterior root of p2. There

is a rather deep masseteric fossa. The anterior mentalforamen lies between the root of p1 and the anterior root ofp2 and the posterior smaller mental foramen below theposterior root of p3 (Fig. 1a,b). The incisors are missing, asare the crowns of the canines. Only the root of the left canineis present. The p1s, whose crowns are not preserved, aresingle-rooted. The p2, preserved in the right side, isasymmetric, the anterior part being smaller than the posteriorone while its mesial face slopes more steeply than the distalone. The base of the crown bulges a little buccally butwithout a distinct cingulum. There is a blunt mesial ridge anda slightly sharper distal one. There is no trace of an accessorycuspid and no talonid. The p3 is larger than p2 and slightlyless asymmetric with the same ridges but with a tiny PACand a very small talonid. In p4, there is a small PAC at mid-height of the disto-buccal side of the main cuspid. Distally, asmall and shallow talonid basin is limited by a low crest and

Table 1 Measurements of teeth and mandible of TM 32-04-78

TM 32-04-78 sin TM 32-04-78 dex

c-m3 48.4p1-m3 41.5 41.0p1-p4 23.7 23.2m1-m3 18.8i3LI3wclcwp1Lp1wp2L 4.9p2w 2.0p3L 5.5 5.5p3w 2.0p4L 6.7 6.6p4w 2.4 2.5m1L 10.3m1w 4.0m1trL 7.0m1talw 3.7m2L 5.5 5.5m2trw 3.6m2talw 3.0m3L 2.7m3w 2.3 2.1Ldias c_p1 2.9Ldias p1_p2 0.9Ldias p2_p3 1.1Ldias p3_p4 1.5 1.2H md p2-p3 6.8 7.1H md p3-p4 7.3 7.5H md p4-m1 8.2 8.2H md m1-m2 8.1 8.9

L Length; w breadth; tr trigonid; tal talonid; dias diastema; hmd heightof the mandible

576 Naturwissenschaften (2007) 94:575–580

a very tiny distal cuspid. There is a moderate buccalcingulum on m1. The protoconid is robust and taller thanthe paraconid. The upper border of the latter stretches almosthorizontally at a 45° angle relative to the long axis of thetooth row. A conical metaconid, rather large for a canid, lieson the disto-lingual face of the protoconid. Both protoconidand metaconid are clearly separated from the cuspids of thetalonid. The conical hypoconid and entoconid are of equalsize. The talonid basin is narrow and it opens distally;insofar, there is neither a hypoconulid nor any distal crest. Anoticeable cingulum runs along the base of the mesial face ofm2. The trigonid of this tooth is shorter than the talonid; theprotoconid and metaconid, both conical, are of equal sizeand are fused together at mid-height. A small and shallowanterior fovea is limited mesially by a low blunt crest butwithout any paraconid. In the talonid, the hypoconid is largerthan the entoconid; the former is in contact with theprotoconid and the latter is linked to the metaconid by alingual crest. The wide talonid basin tapers off posteriorlyand it is closed distally by a low crest without any trace of ahypoconulid. The m3 is oval in occlusal outline. It is reducedbut the trigonid and talonid are clearly indicated. The

metaconid is larger than the protoconid, the two beinglinked for most of their height, with an anterior crest joiningtogether to close a mesial fovea as wide as the talonid basinand a posterior crest that distally encloses the talonid basin.There is a small buccal depression between the protoconidand the posterior crest, and a minute buccal ridge runsvertically along the protoconid.

Comparisons

Among the modern Canidae, the Vulpini (Vulpes, Otocyonand, Urocyon) is distinguished essentially by cranialcharacters (Berta 1987; Tedford et al. 1995). Nevertheless,it also shows other characteristics. The size is small, themandibular corpus is shallow and slender, and the heightdifference between paraconid and protoconid of m1 issmaller than in the Canini (Tedford et al. 1995, Fig. 7) likein TM 32-04-78. Vulpes and the TM canid differ fromOtocyon by the simple p4, the smaller m3, and the lack ofm4 and from Urocyon by the lack of protostylid on thebuccal cingulum of the lower molars and from both generaby the more compressed lower dentition.

Despite the large number of extant fox species, none iscertainly known from the late Miocene of the Old Worldand only a few from North America, where canidsoriginated. We will begin our comparisons with extantAfrican species and then continue with fossil ones. Severalspecies may, today, share the same geographic range andhave different ecological niches, as is the case in MiddleEast with the red, Blandford’s, Rueppell’s and the fennecfoxes (Dayan et al. 1989, 1992). Overall, size must play asignificant role in the definition of these niches.

Vulpes vulpes, the red fox, is the most common species ofVulpes, with a range extending from North America to Africa.It is considerably larger than TM 32-04-78, the third andfourth premolars have more developed PACs, the m1 has arelatively more reduced metaconid and a shorter talonid withlower cuspids relative to the height of the protoconid. The m2is relatively shorter with a smaller mesio-buccal fovea, themesial surface of the protoconid is less steeply inclined andthe talonid is narrower, with a more reduced entoconid. Them3 is more reduced and has smaller foveae.

The scrawny or Rueppell’s fox, V. rueppellii, which issmaller than V. vulpes, is nevertheless larger than the TMcanid. The p3 and p4 have a more developed PAC with asmall up-curling talonid. Hypoconid and entoconid are welldeveloped on m1 and m2, but the talonid narrowsposteriorly on the latter. There is a minute accessory cuspidin m1 between the metaconid and entoconid and, in one ofthe individuals used in the comparisons, a hypoconulid.

The Corsac or steppe fox,Vulpes corsac, is also larger thanthe TM fox. The premolars are relatively higher with more

Fig. 1 Vulpes riffautae n. sp., TM 32-04-78, Toros-Menalla, Chad,late Miocene. a Left corpus and b right corpus of the mandible.1 Lingual view, 2 buccal view, 3 occlusal view. Scale bar=1 cm

Naturwissenschaften (2007) 94:575–580 577

developed PAC; there is no diastema between p3 and p4.The m2 trigonid is broader and has a huge metaconid and anarrower talonid.

The fennec, Vulpes zerda is smaller than TM 32-04-78. Italso differs in its shorter, taller and sharper premolarsseparated by shorter diastemas. The p3 and p4 lack talonidand accessory cuspids. The carnassial has a more transverse-ly oriented paraconid, a smaller metaconid and a shortertalonid with more distinct cuspids. The m2 is shorter relativeto m1, its trigonid is higher and its talonid narrower.

Blanford’s fox, Vulpes cana, whose size is similar to that ofV. zerda (Geffen 1994), inhabits western Asia and the MiddleEast (Geffen 1994; Geffen et al. 2004) and may be present inEthiopia (Geffen in litteris). It differs from the TM specimen

in its dental morphology. The premolars are less separated andsharper; the m1 has a more transversely oriented paraconidlike in V. zerda. The diet of both recent small foxes includes aconsiderable amount of invertebrates, principally insects, andtheir dentitions are a little insectivore-like.

Few Eurasian fossil species can usefully be compared toTM 32-04-78. The genus Vulpes has been cited from thelocality of Venta del Moro, Spain (Ginsburg 1999; Moraleset al. 1999) with a magnetostratigraphic age of 5.8 Ma(Opdyke 1990), but this material is now assigned to Eucyonmonticinensis (van der Made et al. 2006); it is known froman upper carnassial whose size and morphology would besimilar to those of “Canis” adoxus Martin 1973 from thePliocene locality of Saint-Estève (Eastern Pyrénées). The

Fig. 2 a Size comparisons onthe basis of m1 length betweenTM 32-04-78 and some recentand fossil foxes. b Comparisonsof indices (m2 length/m1 length)! 100 of TM 32-04-78 and somerecent and fossil foxes. Datafrom Hendey (1974), Koufos(1993), Kurtén and Crusafont(1977), Odinsov (1967), Poplin(1972), Qiu and Tedford (1990)and our own measurements

578 Naturwissenschaften (2007) 94:575–580

latter is generally assigned to the genus Vulpes (Pons Moyaand Crusafont 1978a,b), but its dental dimensions exceedthose of any species of that genus (i.e. m1=18.1!7.6; m2=0.4!6.3 mm; cf. Fig. 2) and are closer to the dimensions ofEucyon. Vulpes alopecoides Major, 1875 was first describedfrom late Pliocene deposits of the Val d’Arno, Italy butrecovered later in several localities in France, Spain andGreece at the same geological level. It is larger than the TMspecimen (Kurtén and Crusafont 1977; Koufos 1993); thePACs of p3 and p4 are more developed, the metaconid of m1is posteriorly situated, m1 is more robust and m2 is relativelyshorter. Vulpes praecorsac Kormos, 1932 was recoveredfrom Pliocene localities in Hungary and southern Ukraine. Itis larger than the Chadian fossil fox and close to V. corsac(Fig. 2a). Vulpes beihaiensis Qiu and Tedford, 1990 is alsolarger and has a deeper mandible. The premolars are lessasymmetrical and separated by shorter diastemas. The sizedifferences between the TM specimen and the otherEurasiatic foxes are shown on the Fig. 2a. TM 32-04-78 isclearly smaller than the other species except V. zerda. Theratio m2 length/m1 length (Fig. 2b) is a little lower in theTM specimen than in V. cana and similar to that of V. chama.We note that the smallest fox, V. zerda, has the longest m2relative to the length of m1. This characteristic could belinked to the diet. Although V. zerda can sometimes killrabbit-sized animals, it generally feeds on small mammalsand insects like V. cana, another very small fox, hence theirsharp and insectivore-like dentition.

Insofar as canids originated in North America, it isnecessary to take a look at the oldest representative of thegenus Vulpes on that continent. Vulpes stenognathus Savage1941 is larger than TM specimen and “most living speciesof Vulpes”, the premolars are spaced but less than in TMspecimen, there is no diastema between p3 and p4 and themorphology of “m1 and m2 show same cusp arrangementas V. fulvus” (Savage 1941), i.e. the characters of V. vulpes,the red fox. Vulpes sp. was described from the earlyHemphillian (Stevens and Stevens 2003) on the basis ofsome jaw remains with quite worn upper and lowerpremolars. The lower premolars compare quite closely withthose of the TM specimen in the overall morphology of thecrowns, but the wear has obliterated the details, and theyare far larger and more compressed than the TM premolars.

Another American genus, Leptocyon, is sometimes con-sidered close to Vulpes or an ancestor of Vulpes, and someAmerican specimens have been moved from one genus tothe other. Nowadays, Leptocyon is considered the sistergroup of all the living canids (Tedford et al. 1995). Thespecies L. vafer (Leidy 1869) has been recovered from thelate Miocene (Hemphillian) and the early Pliocene. It was a“foxlike canid with slender jaw” (Munthe 1998). It is largerthan the TM fox and the corpus is deeper, the anteriorpremolars are narrow and uncrowded, but without large

diastemas like those of TM fox, with small PACs. Thetrigonid of m1 is open, and the talonid has a much morereduced entoconid relative to the hypoconid. The presence ofa “paraconid of m2 low, vestigial, shelf like” (Munthe 1998)can be considered as plesiomorphic character in comparisonto the Chadian material. The metaconid of m2 is smaller thanthe protoconid, and the mesial cingulum is weak. Accordingto Munthe (1998), the m3 would be very small.

Conclusion

The specimen TM 32-04-78 is the oldest canid thus far foundin Africa and extends the time range of the family in thiscontinent; until now, the oldest African canid was a Canini,Eucyon intrepidus Morales et al. 2005, from the LukeinoFormation, Kenya, dated to 6.1–5.7 Ma. V. riffautae n. sp. isalso potentially the oldest canid of the Old World, insofar, asCanini are known only from the late Miocene of Europe andVulpini from the Pliocene of China and Europe. TheChadian carnivore shows derived features of the Canidae(hypoconulid shelf on m1, m2 with inflated metaconid andlarge mesial cingulum). Its characteristics fit those of theVulpini as a whole but do not match any known species. Itssize, when compared to those of recent or fossil foxes,indicates a fox a little larger than V. zerda but smaller thanmost of the others (Fig. 2a). It does not share any apomorphieswith one of the recent species. V. zerda and V. cana are quiteclose in size to the TM fox, but their dental morphologies,which are a little “insectivore-like”, are very different. LateMiocene canids are unknown in Asia, although the continentis necessarily on the route of their migration from NorthAmerica. The rare late Miocene European canid localities aresituated in south–west Europe (Spain and southern Italy).Therefore, it is possible that canids spread first in Africabefore invading Europe at the end of the Miocene through asouth-to-north trans-Mediterranean route.

Acknowledgements Many thanks are due to the authorities from Chadand France for supporting the field expeditions in the Djurab. This workwas supported by a grant from the RHOI/NSF program. For access tocomparative material, we thank D. Robineau and Alexis Martin,Museumnational d’Histoire naturelle, Paris. Many thanks are also due to SabineRiffaut who skillfully prepared the drawings and Xiaoming Wang(Natural History Museum, Los Angeles) for critics on an earlier draft ofthe paper. We are especially grateful to Lars Werdelin (Swedish Museumof Natural History, Stockholm) for his insightful comments and forimproving the language. We also thank two anonymous referees for theircomments and all the MPFT members who helped to collect the fossils.

References

Berta A (1987) Origin, diversification, and zoogeography of the SouthAmerican Canidae. Fieldiana Geol 39:455–471

Naturwissenschaften (2007) 94:575–580 579

de Bonis L, Peigné S, Likius A, Mackaye H T, Vignaud P, Brunet M(2005) Hyaenictitherium minimum, a new ictithere (Mammalia,Carnivora, Hyaenidae) from the Late Miocene of Toros-Menalla,Chad. Comptes Rendus Palevol 4:671–679

Bowdich TE (1821) An analysis of the natural classification ofMammaliafor the use of students and travellers. J. Smith, Paris, pp 1–115

Brunet M, Guy F, Pilbeam D, Mackaye H T, Likius A, Ahounta D,Beauvilain A, Blondel C, Bocherens H, Boisserie J R, Bonis Lde, Coppens Y, Dejax J, Denys C, Duringer P, Eisenmann V,Fanone G, Fronty P, Geraads D, Lehmann T, Lihoreau F,Louchart A, Mahamat A, Merceron G, Mouchelin G, Otero O,Campomanes PP, De Leon MP, Rage JC, Sapanet M, Schuster M,Sudre J, Tassy P, Valentin X, Vignaud P, Viriot L, Zazzo A,Zollikofer C (2002) A new hominid from the Upper Miocene ofChad, Central Africa. Nature 418:145–151

Brunet M, Guy F, Lieberman DE, Likius A, Mackaye HT, Ponce deLeón MS, Zollikofer CPE, Vignaud P (2005) New material of theearliest hominid from the Upper Miocene of Chad. Nature434:752–755

Dayan M, Tchernov E, Yom-Tov Y, Simberloff D (1989) Ecologicalcharacter displacement in Saharo-Arabian Vulpes: outfoxingBergmann’s rule. Oikos 55:263–272

Dayan M, Simberloff D, Tchernov E, Yom-Tov Y (1992) Caninecarnassials: character displacement in the wolves, jackals, andfoxes of Israel. Biol J Linn Soc 45:315–331

Deino AL, Tauxe L, Monaghan M, Hill A (2002) 40Ar/39Argeochronology and paleomagnetic stratigraphy of the Lukeinoand lower Chemeron Formations at Tabarin and Kapcheberek,Tugen Hills, Kenya. J Hum Evol 42:117–140

Fischer von Waldheim G (1817) Adversaria Zoologica. Mem Soc ImpNat Mosc 5:368–428

Frisch J L (1775) Das Natur-System der vierfüßigen Thiere inTabellen, darinnen alle Ordnungen, Geschlechte und Arten, nichtnur mit bestimmenden Benennungen, sondern beygesetztenunterscheidenden Kennzeichen angezeigt werden zum Nutzender erwachsenen Schuljugend. Günther, Glogau, pp [1–6],1–30, [1–4]

Geffen E (1994) Vulpes cana. Mamm Species 462:1–4Geffen E, Hefner R, Wright P (2004) Blanford’s fox (Vulpes cana).

In: Sillero-Zubiri C, Hoffmann M, Macdonald DW (eds) Canids:foxes, wolves, jackals and dogs. Status survey and conservation:action plan. IUCN, Gland Switzerland, pp 194–198

Ginsburg L (1999) Order Carnivora. In: Rössner GE, Heissig K (eds)The Miocene land mammals of Europe. Verlag Dr Friedlich Pfeil,Munich, pp 109–148

Hemprich FG, Ehrenberg CG (1828–1834) Symbolae physicae, seuicones et descriptiones corporum naturalium novorum aut minuscognitorum quae ex itineribus per Libyam,Aegyptium, Nubiam,Dongalam, Syriam, Arabiam et Habessiniam, pars zoologica II,anima, pp. Plates. Officina Academica, Berlin

Hendey QB (1974) The late Cenozoic Carnivora of the south-westernCape Province. Ann S Afr Mus 63:1–369

Kormos T (1932) Die Fuchse des ungarischen Oberpliozän. Foliazoologica et hydrobiologica 4(2):167–188

Koufos GD (1993) Late Pliocene carnivores from western Macedonia(Greece). Paläontol Z 67:357–376

Kurtén B, Crusafont M (1977) Villafranchian carnivores (Mammalia)from La Puebla de Valverde (Teruel, Spain). Commentat Biol85:1–139

Leidy J (1869) The extinct mammalian fauna of Dakota and Nebraska,including an account of some allied forms from other localities,together with a synopsis of the mammalian remains of NorthAmerica. Journ Acad Nat Sci Phila 7:1–472

Major F (1875) Considerazioni sulla fauna dei mammiferi plioceni epost plioceni della Toscana. Atti Soc Toscana Sci Nat Mem I7–40:223–245

Martin R (1973) Trois nouvelles espèces de Caninae (Canidae,Carnivora) des gisements plio-villafranchiens d’Europe. Noteset Mémoires. Doc Lab Geol Fac Sci Lyon 57:87–95

McDougall I, Feibel CS (1999) Numerical age control for theMiocene-Pliocene succession at Lothagam, a hominoid-bearingsequence in the northern Kenya Rift. J Geol Soc 156:731–745

Morales J, Nieto M, Khöler M, Moyà-Solà S (1999) Large mammalsfrom the Vallesian of Spain. In: Agusti J., Rook L., Andrews P(eds) Hominoid evolution and climatic change in Europe, vol. 1.The evolution of Neogene terrestrial ecosystems in Europe.Cambridge University Press, Cambridge, pp 113–126

Morales J, Soria D, Pickford D (2005) Carnivores from the lateMiocene and basal Pliocene of the Tugen Hills, Kenya. Rev SocGeol Esp 18:39–61

Munthe K (1998) Canidae. In: Janis CM, Scott K, Jacobs LL (eds)Evolution of tertiary mammals of North America. CambridgeUniversity Press, Cambridge, pp 124–143

Odinsov IA (1967) New type of Pliocene beast Vulpes odessana fromkarst caves in Odessa. Paleontologiceskij sbornik 4(1):130–137

Opdyke ND (1990) Magnetic stratigraphy of Cenozoic terrestrialsediments and mammalian dispersal. J Geol 98:621–637

Peigné S, Bonis L de, Likius A, Mackaye H T, Vignaud P, Brunet M(2005a) A new machairodontine (Carnivora, Felidae) from theLate Miocene hominid locality of TM 266, Toros-Menalla, Chad.Comptes Rendus Palevol 4:243–253

Peigné S, Bonis L de, Likius A, Makaye H T, Vignaud P, Brunet M(2005b) The oldest modern mongoose (Carnivora, Herpestidae)from Africa: Galerella sanguinea from the late Miocenehominid-bearing locality TM 266, Toros-Menalla, Chad. Natur-wissenschaften 92:287–292

Pons Moya J, Crusafont Pairo M (1978a) Sobre la identidad del«Canis» adoxus Martin (1973) y su implicación en el conoci-miento del genero Vulpes. Acta Geol Hisp 13:129–132

Pons Moya J, Crusafont Pairo M (1978b) El Canis cipio Crusafont(1950), comparación con los canidos del Plioceno y Pleistocenoeuropeo. Acta Geol Hisp 13:133–136

Poplin F (1972) Contribution à la morphologie et à la biométried’Alopex lagopus (Linné) et de Vulpes vulpes (Linné). Lesrenards d’Arcy-sur-Cure. Thesis, Univ Paris-6, pp 1–149

Qiu Z, Tedford RH (1990) A Pliocene species of Vulpes from Yushe,Shanxi. Vertebr Palasiatica 28:245–258

Savage D (1941) Two new middle Pliocene carnivores from Oklahomawith notes on the Optima fauna. Am Midl Nat 2:692–710

Schuster M, Duringer P, Ghienne JF, Vignaud P, Makaye HT, Likius A,Brunet M (2006) The age of the Sahara desert. Science 311:821

Stevens MS, Stevens JB (2003) Carnivora (Mammalia, Felidae,Canidae, and Mustelidae) from the earliest Hemphillian screwbean local fauna, Big Bend National Park, Brewster County,Texas. Bull Am Mus Nat Hist 279:177–211

Tedford RH, Taylor BE, Wang X (1995) Phylogeny of the Caninae(Carnivora: Canidae): the living taxa. Am Mus Novit 3146:1–37

van der Made J, Morales J, Montoya P (2006) Late Miocene turnoverin the Spanish mammal record in relation to palaeoclimate andthe Messinian Salinity Crisis. Palaeogeogr Palaeoclimatol Palae-oecol 238:228–246

Vignaud P, Duringer P, Mackaye HT, Likius A, Blondel C, BoisserieJR, de Bonis L, Eisenmann V, Etienne ME, Geraads D, Guy F,Lehmann T, Lihoreau F, López-Martínez N, Mourer-Chauviré C,Otero O, Rage JC, Schuster M, Viriot L, Zazzo A, Brunet M(2002) Geology and paleontology of the Upper Miocene Toros-Ménalla hominid locality, Chad. Nature 418:152–155

Wang X, Tedford RH, Van Valkenburgh B, Wayne RK (2004)Ancestry: evolutionary history, molecular systematics, andevolutionary ecology of Canidae. In: Macdonald DW, Sillero-Zubiri C (eds) The biology and conservation of wild canids.Oxford University Press, Oxford, pp 39–54

580 Naturwissenschaften (2007) 94:575–580