First occurrence of the 'hunting hyena' Chasmaporthetes in the Late Miocene fossil bearing localities of Toros Menalla, Chad (Africa)

First occurrence of the ‘hunting hyena’ Chasmaporthetes in the Late Miocenefossil bearing localities of Toros Menalla, Chad (Africa)

LOUIS de BONIS1, STEPHANE PEIGNE1,3, ANDOSSA LIKIUS2, HASSANE T. MAKAYE2, MICHEL BRUNET1

and PATRICK VIGNAUD1

Key words. – Late Miocene, Africa, Chad, Mammalia, Carnivora, Hyaenidae.

Abstract. – Late Miocene localities of Toros Menalla (Chad) have yielded many bones of fossil vertebrates with a lot ofmammalian remains. Among the mammals, there are several Carnivora taxa, especially hyenids. The family Hyaenidaeis very well developed during this period with classical bone crusher species but also with flesh eater taxa which are cal-led hunting hyenas. The genus Chasmaporthetes is one of these taxa. It was described from North America, Asia, Euro-pa and South Africa but it is recorded for the first time in central Africa. The Chadian specimens are close to the SouthAfrican species C. australis (HENDEY, 1974) but differs through some morphological and metrical details. C. australisis a huge hunting hyena, a little bigger than the extant species Crocuta crocuta, the spotted hyena. An isolated premolarrecorded in the locality Sahabi (Libya) belongs probably to the same group. The spreading of this large hunting speciesis probably correlated with the abundance of large ungulates in the local faunas.

Découverte de la hyène “chasseresse” Chasmaporthetes dans les gisements fossilifères duMiocène supérieur de Toros Menalla (Tchad)

Mots-clés. – Miocène supérieur, Afrique, Tchad, Mammalia, Carnivora, Hyaenidae

Résumé. – Les gisements de vertébrés du Miocène supérieur de Toros-Menalla (Tchad) ont livré un grand nombre derestes fossiles de mammifères parmi lesquels les carnivores de la famille des Hyaenidae sont fort bien représentés.Celle-ci est particulièrement diversifiée à cette époque. En effet, à coté des formes classiques de hyènes nécrophagesdont la denture extrêmement robuste permet de broyer les os, existaient d’autres formes, appelées parfois hyènes “ chas-seresses “, spécialisées pour un mode de vie plus prédateur. C’est le cas du genre Chasmaporthetes. Décrit pour la pre-mière fois en Amérique du Nord, il a été retrouvé en Asie, Europe et Afrique. Les spécimens du Tchad sont proches dela plus grande espèce du genre, C. australis du Pliocène inférieur de Langebaanweg (Afrique du Sud) dont la taille dé-passe légèrement celle de l’actuelle hyène tachetée, Crocuta crocuta. Ils diffèrent cependant de l’espèce sud-africainepar un certain nombre de traits qui pourraient justifier la création d’une espèce distincte mais, dans l’attente d’une meil-leure connaissance de l’étendue des variations sur le matériel de Langebaanweg, il nous paraît préférable de considérerle carnivore tchadien comme Chasmaporthetes cf. australis. Une prémolaire isolée récoltée à Sahabi (Liby) pourraitaussi être rattachée au même groupe. L’extension de cette espèce géante sur l’ensemble du continent africain à la fin duMiocène et au début du Pliocène pourrait être liée à l’abondance à cette époque de gros gibier comme les anthracothèresou les hippopotames.

INTRODUCTION

Located in the Djurab desert, 500 km north-east ofN’Djamena (Chad), the fossiliferous localities of Toros Me-nalla, well known by the discovery of the hominine Sahe-lanthropus tchadensis [Brunet et al., 2002], have yieldednumerous Late Miocene (c.a. 7 Ma) vertebrate remains re-corded by surface collection and excavation as well. Thechronologic assessment of these faunas is mainly based onthe biochronology of mammalian taxa. More than 300 fossilbearing sites have been located by the MPFT (Mission Pa-léoanthropologique Franco-Tchadienne). The carnivores arewell represented and, among them, the family Hyaenidae isquite common through several species of different sizes [

Vignaud et al., 2002; Bonis et al., 2005]. The study of oneof them, remarkable by its large dimensions, constitutes theaim of this article. The remains of this species come fromthree localities (TM 60; TM 112; TM 289), which are situa-ted in the “Anthracotheriid level” and associated to remainsof, among other taxa, Lybicosaurus petrochii and the suidNyanzachoerus syrticus [Vignaud et al., 2002].

MATERIAL AND METHODS

List of the material: TM 112-00-98; right maxilla (crushedcanine alveolus, P1 alveolus, P2-M1); left maxilla (canine al-veolus, P1 alveolus, partially broken P2, broken P3, broken

Bull. Soc. géol. Fr., 2007, t. 178, no 4, pp. 317-326

Bull. Soc. géol. Fr., 2007, no 4

1. Laboratoire de Géobiologie, Biochronologie et Paléontologie humaine, UMR 6046 CNRS, Univ.Poitiers, Fac. Sciences, 40 avenue du recteur Pineau,86022 Poitiers cedex, Email: [email protected]. Université de N’Djamena, BP 1117, N’Djamena, Chad.3. MNHN, Paléontologie, 8 rue Buffon, 75005 Paris.Manuscrit déposé le 3 juillet 2006 ; accepté après révision le 18 janvier 2007.

anterior part of P4). TM 60-02-04: left maxilla (part ofI1-I2 alveoli, I3 alveolus, canine alveolus, P1 alveolus, ba-dly broken P2 and P3, damaged P4). TM 289-02-02: rightupper third incisor. Fragment of upper left P4; left mandible(p3-m1); left isolated p2, right isolated p2 on a piece ofmandible; right isolated p3 on a piece of mandible; rightisolated p4; right isolated m1; very damaged right lower ca-nine; all of which are situated at the same place and they be-long to the same individual on the basis of vicinity andstage of wear.

The specimens are described then compared to othertaxa from a morphological point of view and metrically.The use of bivariate and multivariate statistical analyses al-lows an extended comparison of size and proportions aswell. Measurements are given in millimetres and our ownmeasurements were taken with a vernier calliper to1/10 mm.

SYSTEMATICS

Order: Carnivora Bowdich, 1821Family: Hyaenidae Gray, 1869

Genus: Chasmaporthetes Hay, 1921Species: Chasmaporthetes cf. australis (HENDEY, 1974)Localities: Toros Menalla (TM 60; TM 112; TM 289),

Sahabi?Geological level: Late Miocene

Description

Upper jaw

The upper third incisor TM 289-02-02(fig. 3 F, 3G) resem-bles that of the extant Hyaena, although slightly less tall, ofthat of Chasmaporthetes lunensis from Saint Vallier [Viret,1954] but far smaller than that of Adcrocuta. The maxillaTM 112-00-98 is so damaged that we cannot see any detail;in TM 60-02-04 (fig. 1), the infra-orbital foramen, which isquite small, opens between the posterior root of P2 and the

anterior root of P3, just behind a bulging caused by the ca-nine root. The upper dentition is well represented on themaxilla TM 112-00-98 (fig. 2). The first incisor was cer-tainly small, the second one medium sized and the third onevery big. The canine alveolus is large but not very deep. Sothe canine root was robust but rather short. This charactercould correspond to a lesser ability for loading strong forceto the top of the canine and linked to lesser bone crushing inthe diet. Present on both side of TM 112-00-98, the P1 al-veolus is rounded, quite large and situated between the ca-nine and P2 but in contact with the distal border of thecanine alveolus; P1 was probably developed as in speci-mens of Chasmaporthetes from China [Galiano and Frailey,1977]. In TM 60-02-04 it is smaller and situated in contactwith the disto-buccal border of the canine alveolus. All thepremolars have a horizontal wear facet on the top of thecusps. P2 and P3 are elongated and quite slender so diffe-rent from the common hyaenid conditions. P2 is two rooted;the anterior part of the crown is broken but we can measurethe length on the right side; there is a small but well definedposterior accessory cusp (PAC); the great axis of the crownis oblique relative to the tooth row. P3 is larger than P2 andimbricated with it; there is no anterior accessory cusp(AAC) but a well-developed PAC followed by a cingulum;the base of the crown is bulging buccally and a little distally(and is supported by a third root). P4 is elongated with alarge lingually and anteriorly directed protocone, which isvisible in buccal view. M1 is reduced and transversely di-rected; the parastyle and the paracone seem to be fused andbulge out buccally; the metacone is a well defined smallcusp; both, paracone and metacone are linked by a crest tothe crescent shaped protocone. There is no M2.

Lower jaw

The lower part of the piece of a left mandible is broken butit was probably rather deep (fig. 3). The p1 is probably ab-sent as in other species of Chasmaporthetes. The secondpremolar is asymmetrical, the mesial part being shorter than


318 BONIS L. de et al.

FIG. 1. – Chasmaporthetes cf. australis. TM 60-02-04. Left maxilla: A – inlateral view; B – in lingual view; C – in occlusal view (scale bar = 1 cm).FIG. 1. – Chasmaporthetes cf. australis. TM 60-02-04. Maxillaire gauche :A – en vue latérale ; B – en vue linguale ; C – en vue occlusale (échelle =1 cm).

FIG. 2. – Chasmaporthetes cf. australis. TM 112-00-98. Right maxilla: A –in lateral view; B – in lingual view; C – in occlusal view (scale bar = 1 cm).FIG. 2. – Chasmaporthetes cf. australis.TM 112-00-98. Maxillaire : A – envue latérale ; B – en vue linguale ; C – en vue occlusale (échelle = 1 cm).

the distal one and more steeply inclined; a small crest runsalong the mesial face from the top to the base where it turnsa little lingually without AAC; there is a well developedPAC. The p3 is larger than p2 and less asymmetrical; thereare an AAC and a PAC. The p4 is quite symmetrical as is afelid premolar with two well-developed accessory cuspids.The lower carnassial m1 is long and slender; the protoconidis slightly taller than the paraconid; there is no trace of ametaconid; a nodulous cingulum runs along the lingual baseof the paraconid; the talonid is quite small with a trenchanthypoconid linked by a distal crest to a tiny entoconid. Thereis no m2. The right isolated premolars and m1 are similar tothe left ones.

COMPARISONS

Genus level

The Hyaenidae with tall and elongated premolars (trenchantfunction) belong to a group often called “hunting hyaenas”which is composed of three genera: Lycyaena, Hyaenictisand Chasmaporthetes.

For Pilgrim [1931, p. 103] Lycyaena is a “Hyaenidaewith narrow, elongate skull; occipital of medium height, notmarkedly triangular, narrow below; sagittal crest rather low;lambdoidal crest moderately prominent; orbit rather large;post-orbital processes not very prominent; premaxillae se-parated by an interval from the frontals; dentition I 3/3 CI/1 P 3/3 or 4, M 1/1(2); third incisor somewhat enlarged;canine large; anterior premolars in a straight line, generallywell spaced (slightly overlapping in L. lunensis), long, withlarge posterior and practically no anterior cusp; P4 withprominent protocone; M1 rather large, transverse to P4 butslightly behind it; mandible with deep masseteric fossa, lo-wer premolars with well marked anterior and posteriorcusps; ml with metaconid and small talonid; m2 lost in allexcept the most primitive species.” Although Pilgrim inclu-ded in his diagnosis the genus Chasmaporthetes (Chasma-porthetes lunensis), the shape of M1 [Pilgrim 1931,Fig. 29], less triangular, the spaced premolars, the lowercarnassial with a metaconid and a better developed taloniddo not fit TM specimen characters.

For Pilgrim [1931, p.101] Hyaenictis, whose type spe-cies is H. graeca [Gaudry, 1861], is a “Hyaenidae with


FIRST OCCURRENCE OF THE ‘HUNTING HYENA’ CHASMAPORTHETES, CHAD (AFRICA) 319

FIG. 3. – Chasmaporthetes cf. australis. TM289-02-02. Left mandible and isolated p2: A – in buc-cal view; B – in lingual view; C – in occlusal view;TM 289-02-02. Isolated right p2-p3-p4-m1: D – inbuccal view; E – in lingual view; TM 289-02-02.Right upper I3: F – buccal view; G – mesial view(scale bars = 1cm).FIG. 3. – Chasmaporthetes cf. australis. TM289-02-02. Mandibule et p2 isolée gauches : A – envue buccale ; B – en vue linguale ; C – en vue occlu-sale. TM 289-02-02. P2-p3-p4-m1 droites isolées : D– en vue buccale ; E – en vue linguale ; TM289-02-02. Troisième incisive supérieure droite : F –en vue buccale ; G – en vue mésiale (échelles = 1 cm).

moderately long, slender, rather shallow mandible and (byinference) with somewhat elongated facial region”; Ml large,triangular, almost at right angles to P4; M2 absent; P4 withlong posterior lobe and large protocone; ml short, hardly lon-ger than P4, without metaconid; m2 small; pl present butwith a tendency to be deciduous; p2 with large posterior andsmall anterior cusps; p3 with large posterior and small ante-rior cusps; p4 with large anterior and posterior cusps. In a re-vised diagnosis [Werdelin et al., 1994], Hyaenictis is a“medium to large sized Hyaenidae; mandible long and slen-der; dental formula I3/3 C1/1 P4/3-4 M1/1-2; p2 and p3 slen-der with weakly developed anterior accessory cusps; P4 withmetastyle slightly longer than paracone; m1 with short, twoor three cusped talonid, m1 metaconid lost; m2 present”. Inm1, a very small bulge is in the place of the metaconid andthe talonid is made up of two well separated cuspids. TheTM lower premolars are not spaced, there is no m2 and themandible is deeper. TM differs from the upper teeth allocatedto this species by a less slender P4 (shorter metastyle).

Chasmaporthetes, whose type species is C. ossifragusHAY, 1921, had its diagnosis revised [Galiano and Frayley,1977, p. 2] as follows “P1 larger than in other hyaenid gene-ra (absent in Euryboas). Anterior accessory cusps of p2/2and p3/3 more prominent than in other hyaenids; upper andlower tooth rows curved with premolars slightly imbricated,as opposed to a straight tooth row as in Euryboas”. The dia-gnosis did not include the genus Euryboas which was latter

synonymised with Chasmaporthetes. In another revision[Kurten and Werdelin, 1988, p. 48], the two genera were in-cluded in Chasmaporthetes as “medium sized hyaenids withlong, slender limbs; rostrum broad; frontal profile stepped;cheek teeth slender, trenchant; dental formula 13/3, C 1/1,P3-4/3-4, M1/1; P1 large, with a tendency to be shed earlyin life; anterior premolars with strong posterior accessorycusps, anterior cusps variable; P4 with large protocone,short paracone, and elongated metastyle; M1 transverse; p4elongate, with large accessory cusps; m1 without metaconidand with talonid bearing a single, laterally compressed,blade-like cusp with a rounded lateral profile”. These cha-racters fit those of the TM specimens despite the talonid ofm1 has a tiny entoconid. The genus Euryboas SCHAUB,1941, established for E. lunensis from the Early Pleistoceneof Roccaneyra, Massif Central, France, has been synonymi-sed with Chasmaporthetes.

Species level

Morphological comparisons

Several species have been included in the genus Chasma-porthetes from North America, Asia, Europa, and Africa.All are dated from Late Miocene to Early Pleistocene.

The type species, C. ossifragus HAY, 1921, comes fromthe North American Early Pleistocene locality CoconinoForest region, Arizona, USA. It was established on a badlypreserved piece of mandible with the diagnosis: “fourth pre-molar and first molar of nearly the same length; the molarapparently thicker. Jaw deeper and thicker than in existingspecies. Referred second molar two-rooted and its socketnearly two third as long as that of the molar. Symphysis ex-tending back to at least the middle of the second premolar”[Hay, 1921, p. 636]. It is clear that the referred second mo-lar does not belong to the same genus. A beautiful right lo-wer jaw coming from North Cita Canyon, bed 4, RandallCounty Texas and dated to Late Pliocene has been describedas Ailuraena johnstoni STIRTON & CHRISTIAN, 1940, and la-ter synonymized with the former allows to complete thedescription and to show that the second molar is absent.Other specimens of Chasmaporthetes have been recorded inlatest Pliocene-Early Pleistocene (Late Blancan) layers ofNorth America [Berta, 1981]. In the maxilla [Berta, 1981,fig. 2], C. ossifragus differs from the TM material by thenasal aperture plan more steeply inclined, the infra-orbitalforamen more backwardly located (between the hinder rootof P3 and the front root of P4), the P1 alveolus clearly sepa-rated from the canine alveolus, the P4 protocone more back-wardly situated (not visible on a buccal view of P4). In themandible [Hay, 1921; Stirton and Christian, 1940, Fig. 1;Berta, 1981, Fig. 3], its differs by diastema between p2-p3and p3-p4, relatively shorter p2 and p3 and shorter talonidof m1.

The geologically oldest species allocated to Chasma-porthetes are C. bonisi KOUFOS, 1987 from the Turolian ofnorthern Greece and C. exitelus KURTEN & WERDELIN,1988 from the Turolian of China. The former differs fromthe Chad specimens by the relatively larger M1 especiallythe bucco-lingual diameter and the shape of the upper pre-molars whose maximum width is more central. The latterdiffers from TM 112 by the same last character.



FIG. 4. – Metrical comparisons of the species of Chasmaporthetes. A –Scatterplot: P4 length x P4 width; B – Scatterplot: P4 length x P3 length.FIG. 4. – Comparaisons métriques des espèces de Chasmaporthetes. A –Diagramme de dispersion : longueur de P4 x largeur de P4 ; B – Dia-gramme de dispersion : longueur de P4 x longueur de P3.

The Pleistocene species C. melei ROOK, FERRETI, AREA& TUVERI, 2004 from Sardinia is very small when compa-red to TM specimens. The cheek teeth look thinner.

Chasmaporthetes borissiaki (KHOMENKO, 1932) comesfrom a Moldavian locality of Early Pliocene age. It wouldbe characterised by relatively small size; a muzzle long andnarrow; P4 with very strong, trenchant protocone; P2-4 andm1 narrow; pI at least occasionally present; forelimb relati-vely short [Kurten and Werdelin, 1988]. We do not knowlimb bones of this TM hyaenid. We cannot compare the TMadult maxillae with the holotype of C. borissiaki, which be-longs to a young individual with an erupting canine, but themuzzle of the Moldavian species is closer to that of C. ossi-fragus than to that of TM species. P4 protocone is less pro-jecting anteriorly and not visible in buccal view, P3 is moreasymmetrical with the mesial surface more steeply inclined.As for the maxilla, in the mandible the premolars p2 and p3of C. borissiaki are more asymmetrical and the talonid ofm1 seems to be a little shorter.

Another European species, C. lunensis (DEL CAMPANA,1914) from the Early Pleistocene of Olivola, Val di Magra,Italy, and to which material from many other European lo-calities is referred [Kurten and Werdelin, 1988], differsfrom C. borissiaki by its larger size although smaller andless robust than C. ossifragus, with distal forelimb marke-dly lengthened; P2-4 and m1 broader; anterior cusps onP2-3/p2-3 smaller than in C. nitidula (see below). It hasbeen subdivided into two sub-species. The nominate onehas a P4, and perhaps the other premolars, longer than thatin the other one C. lunensis honanensis (ZDANSKY, 1924).Some authors believe the latter to be a different species, C.kani GALIANO & FRAILEY, 1977 from the Middle to LateVillafranchian (Late Pliocene and Early Pleistocene) ofChina but honanensis has priority on kani at the species le-vel. For Qiu [1987] another sub-species does exist called C.kani progressus. The type specimen of C. lunensis differsfrom TM 112-00-98 by the more distal protocone of P4, P3with a slightly more steeply inclined mesial surface and aless posterior lingual bulging; and M1 less reduced. The up-per teeth of the Chinese sub-species or species [Zdansky,1924; Galiano and Frailey, 1977] differ from TM 112-00-98by the same characters. In the mandible of the Chinese spe-cimens, p2 and p3 have a more steeply inclined mesial sur-face and the three premolars have a more marked buccalcingulid. The snout is also more steeply inclined than that

of TM 60-02-04 and the infra-orbital foramen is more pos-teriorly located.

Chasmaporthetes silberbergi (BROOM in BROOM &SCHEPERS, 1946) is a quite poorly known species. The typespecimen, a piece of snout and mandible, comes from a lo-wer infilling of the Pliocene hominid bearing locality Sterk-fontein (South Africa). Other specimens were describedfrom Swartkrans, South Africa [Ewer, 1955] and other le-vels of Sterkfontein [Turner, 1987]. In the mandible, thepremolars are more symmetrical than that of TM283-02-02, the distal part of the crown being less developedrelative to the mesial one. There are two mental foraminaunder the anterior and posterior roots of p2 respectively. Apiece of maxilla with P3-P4 from Sterkfontein (STS 127)allocated to this species, or to C. nitidula (EWER, 1955) dif-fers from TM 112-00-98 by the shorter main cusp, the wea-ker lingual bulging at the base of the crown, the lingualposition of the AAC and the more steeply inclined anterioredge of P3. The P4 seems to have a less developed proto-cone.

Chasmaporthetes nitidula (EWER, 1955), whose sizematches that of C. silberbergi, was described from the EarlyPleistocene of Swartkrans, South Africa. A revised diagno-sis [Kurten and Werdelin, 1988, p. 50] defines the species as“resembles lunensis, but anterior cusps of P2-3/3 larger.”For Turner [1987, p. 323], “the lower premolars of C. niti-dula are more oval in occlusal outline whereas those of C.silberbergi tend to be rectangular and give more massiveappearance”. They have “a rather ’fleur de lys’ profile onboth the anterior and posterior edges of the main cusps,whereas both edges of C. silberbergi tend to be straight”.The AAC of the lower p2-p3 of the type specimen of thisspecies are relatively larger than those of TM but it is morepuzzling concerning the upper premolars. On both speci-mens SK 309 [Ewer, 1955, Fig.6] the anterior accessorycusps are very well developed and visible in buccal view buton the fragment of maxilla SK 305 with P2-P4 [Ewer, 1955,Fig. 5] the P2 and P3 posterior accessory cusps are smalland lingually situated; on the same maxilla P4 looks verymuch like that of TM 112-00-98; the infra-orbital foramenis posteriorly situated, between the posterior root of P3 andthe anterior one of P4, relative to that of TM maxillae. Asub-species, C. n. darelbediae GERAADS, 1997 from theLate Pliocene of Ahl al Oughlam (Morocco), was establis-hed on a piece of mandible with p3-p4 and a couple of iso-lated teeth. It would differ from the nominate sub-species



TABLE. I. – Dental measurements (mm) of Chasmaporthetes cf. australis from Toros Menalla, Late Miocene, Chad. L: total length; w: width; Lp: linguallength; Lm: length of the metastyle; wa: anterior width; wm: width of the metastyle; wp: posterior width; Ltr: length of the trigonid; taw: width of the talo-nid.TABL. I. – Mesures dentaires (mm) de Chasmaporthetes sp. cf. C. australis de Toros Menalla, Miocène supérieur, Tchad. L : longueur totale ; w : largeur ;Lp : longueur linguale ; Lm : longueur du métastyle ; wa : largeur antérieure ; wm : largeur du metastyle ; wp : largeur postérieure ; Ltr : longueur dutrigonide ; taw : largeur du talonide.

by a smaller size and a lesser length difference between p3and p4.

Chasmaporthetes australis (HENDEY, 1974) is betterknown from more complete material recorded at the localityLangebaanweg, South Africa and considered of Early Plio-cene age [Hendey, 1974]. The type specimen (SAM PQL14199) is a crushed skull with part of the mandible inclu-ding the following teeth: part of a right P3, left and right P4,left i1 to i3, /c and p4-m1; right i1 I1, c, and p2 to m1. ForHendey [1974, p.] it is a species “similar in size to P. eximiabut differs in lacking p1, P2/2 and P3 without anterior acces-sory cusps; premolars longer; m1 metaconid very small orabsent. Mc I not as reduced as in modern hyaenids”. ForWerdelin and Solounias [1991, p.] it “differs from C. nitidu-la, particularly in the relative length of p3-p4. The latter to-oth is much shorter in C. australis”. The specimens recordedfrom Langebaanweg after Hendey [1974] show that the p2may have an anterior accessory cuspid [Hendey, 1978,Fig. 6B] which indicates an intraspecific variation. TM spe-cimens differ from C. australis from Langebaanweg by a P3more developed relative to P4, especially at the mesial half ofthe crown whose mesial edge is straight and not slightlyconcave as TM P3, and, a P4 protocone more developed. Itdiffers also from the mandible of the type specimen mandibleby the developed AAC of the lower premolars; but, other spe-cimens from Langebaanweg (SAM PQ L22004, for instance)have also more developed AAC.

Metrical comparisons

The comparisons were made using our own measurements(tabl. I) and those of different authors [Argant, 2004; Ber-ta, 1981; Del Campana, 1914; Ewer, 1955; Ficarelli andTorre, 1967; Galiano and Frailey, 1977; Geraads, 1997;Hendey, 1974; Hendey, 1978; Khomenko, 1932; Kurtenand Werdelin, 1988; Qiu, 1987; Rook et al., 2004; Werde-lin et al., 1994]. The acronyms of the used specimens arein table II.

Upper teeth

The size of P4 (buccal length x anterior width) can distin-guish clearly three groups (fig. 4 A). The first one, with onlyC. melei, is smaller with a relative smaller width. Anotherone with C. australis and TM specimens is also clearly sepa-rated by its large size. The third one is composed by all theother taxa, which are grouped with the specimens allocatedto C. ossifragus. We note nevertheless a specimen from Chi-na whose width is a little bigger. A close result is reachedwhen using P2 (length and breadth) or P3 (length andbreadth) as well as P3 length and P4 length (fig. 4 B). In thatlatter case, a specimen of C. ossifragus and another one fromSterkfontein (C. silberbergi or nitidula) are distinguishedfrom the bulk of middle-sized specimens by a little biggersize. In principal component analysis (PCA) on the measure-ments of the three premolars P2-P3-P4 (fig. 5 A), TM and C.australis plot together on the first factor, which explain71.09% of the total variance and corresponds to the overallsize. A correspondence factor analysis (fig. 5 B) allows exa-mining the differences of proportions between the studiedspecimens [Bonis and Lebeau, 1974]. On the plan of the twofirst factors (57.79% and 25.14% of the total variance respec-tively), the South African species is separated from the Cha-dian specimen on the second axis by the relative size of thepremolars. P2 is clearly larger in the former and P3 is



TABLE. II. – Explanation of the acronyms used in the bivariate and multiva-riate analysis. Abbreviations are explained in the text.TABL. II. – Signification des acronymes utilisés dans les analyses bi- et mul-tivariées. Les abréviations sont expliquées dans le texte.

relatively larger in the latter. This result is clearly demonstra-ted if a PCA is focused only on P2 and P3 dimensions (fig. 6A). The Chasmaporthetes specimens from Chad are clearlyseparated from C. australis specimens from Langebaanwegon the second axis in function of the P2 and P3 length. Theformer are situated close the mean value of the other Chas-maporthetes species whereas the latter are close to C. ossi-fragus and, in a lesser extent, to C. borissiaki and opposed toC. nitidula or Chinese specimens.

Lower teeth

The scatterplot of the lower carnassial measurements(length x width) is less clear than that of the upper one(fig. 6 B). The TM m1 is the largest, but specimens of C. os-sifragus plot within the C. australis plotting. In a PCA onmeasurements of p2, p3, p4, and m1 (fig. 7), TM and onespecimen of C. australis (AU5 = L20988) plot close toge-ther on the two axis. The first axis (71.35% of the variance)corresponds to the overall size differences and the secondone (11.97% of the variance) opposes the length to thewidth of the teeth, these specimens being more linked to thelength. But we can see that another specimen (AU12 =L22204) of C. australis plots with C. ossifragus close to the



FIG. 5. – Metrical comparisons of the species of Chasmaporthetes. A –Principal component analysis on measurements of P2-P3-P4; B – Corres-pondence factor analysis on measurements of P2-P3-P4.FIG. 5. – Comparaisons métriques des espèces de Chasmaporthetes. A –Analyse en composantes principales sur les mensurations de P2-P3-P4 ; B– Analyse des correspondances sur les mensurations de P2-P3-P4.

FIG. 6. – Metrical comparisons of the species of Chasmaporthetes.A – Principal component analysis on measurements of P2-P3; B – Scatter-plot: m1 length x m1 width.FIG. 6. – Comparaisons métriques des espèces de Chasmaporthetes.A – Analyse en composantes principales sur les mensurations de P2-P3 ;B – Diagramme de dispersion : longueur de m1 x largeur de m1.

larger specimens on the first axis but in a median place forthe proportions between length and width. We can note alsoa large dispersion of the Shansi specimens concerning boththe overall size and the proportions. In a CFA computed onthe same data (fig. 8 A), the first axis (35.21% of the totalvariance) separates the specimens in function of the lengthof m1 which is opposed to the length of the premolars; To-ros Menalla mandible is in the first group and L20988 in thesecond one but L22204 plots in the first one. The secondaxis, as significant as the first one (32.60% of the total va-riance), is essentially defined by the relative width of thedentition, especially p2, with grouping of specimens fromRocca Neyra, Dytiko, Shansi (C. lunensis) and a specimenof C. ossifragus in the upper part of the graph but other spe-cimens from Shansi or of C. ossifragus are in the lower partof the graph. In a scatterplot of p3 and p4 lengths (fig. 8 B),specimens from Langebaanweg, except AU 12, and Chadgroup together whereas C. melei is on the opposed place ofthe graph. C. nitidula darelbediae plots between the latterand specimens from China and Kuruksai; but the propor-tions between the two premolars are similar to that of other

species. So only a size difference separates this putativesub-species from C. lunensis or C. nitidula.

CONCLUSIONS

The materiel from Toros Menalla indicates for the first timethe presence of the large hunting hyaenid Chasmaporthetesin central Africa in, at least, three different localities. Thisgenus spread over a large territory during the Late Mioceneand even the Plio-Pleistocene, having been found in NorthAmerica, Far East, western Asia, Europe and Africa. In thislarge realm, several species were described which differfrom each other by slight characters although they can beseparated from other hyaenids. Thus, in an unpublishedmultivariate analysis, the specimens belonging to Chasma-porthetes, including Late Miocene C. exitelus and C. bonisi,are clearly distinct from Adcrocuta eximia, another LateMiocene hyaenid whose size is quite similar. By its size andmorphology the Chadian hyaenid differs from most of theother species but it is close to the South African C. australis



FIG. 7. – Metrical comparisons of the species of Chasmaporthetes. Princi-pal component analysis on measurements of p2-p3-p4-m1.FIG. 7. – Comparaisons métriques des espèces de Chasmaporthetes. Ana-lyse en composantes principales sur les mensurations de p2-p3-p4-m1.

FIG. 8. – Metrical comparisons of the species of Chasmaporthetes. A –Correspondence factor analysis on measurements of p2-p3-p4-m1; B –Scatterplot: length of p4 x length of p3.FIG. 8. – Comparaisons métriques des espèces de Chasmaporthetes. A –Analyse des correspondances sur les mensurations de p2-p3-p4-m1 ; B –Diagramme de dispersion : longueur de p4 x longueur de p3.

from its Early Pliocene type locality, Langebaanweg. Thesize of the specimens of the two samples are similar andthey differ only by features which could be of minor signifi-cance. In Langebaanweg specimens, the P4 protocone isless developed, the shape of P3 is different, P2 and p2 arelarger and the m1 is smaller. Nevertheless considering thelarge variation which does exist in other species (C. lunen-sis for example), it seems better to keep the Toros-Menallasample as Chasmaporthetes cf. australis despite these diffe-rences. So the territory of this species may have extendedfrom southern to central Africa and may be northern Africa.An isolated germ of premolar from Sahabi, Libya, was figu-red by Howell [1987, Fig. 5B] as a p3 of Euryboas sp. Thisgerm could be a p2 as well and in this case it would matchvery well C. australis. The presence of the large hunting

hyenas together with large machairodontid felids [Peigné etal., 2005] in Late Miocene and Early Pliocene layers couldbe linked to the abundance of big game, large ungulates likeanthracotherids or hippos.

Acknowledgements. – We are grateful for financial support to the NSF pro-gram Revealing Hominid Origins Initiative. Many thanks are due to autho-rities from Chad (Ministère de l’Education Nationale, de l’EnseignementSupérieur et de la Recherche, Université de N’Djamena, CNAR, DRGM)and France (Ministère de l’Enseignement Supérieur, de la Recherche et dela Technologie, Ministère des Affaires Etrangères, CNRS, Université dePoitiers) for supporting the field expeditions in the Djourab and work inthe lab. We also thank all the MPFT participants and people who gave usaccess to comparative materials. The fine arts are from Sabine Riffaut. Wethank very much our colleague, the late Pr. F. Clark Howell for his scienti-fic comments and editing remarks, which greatly improved a first versionthe manuscript as well as the two referees, our colleagues M. Morlo and L.Rook.

Reference

ARGANT A. (2004). – Les carnivores du gisement Pliocène final deSaint-Vallier (Drôme, France). – Geobios, 37, 133-182.

BERTA A. (1981). – The Plio-Pleistocene Hyaena Chasmaporthetes ossifra-gus from Florida. – J. Vert. Paleont., 3-4, 341-356.

BOWDICH T.E. (1821). – An analysis of the natural classification of Mam-malia for the use of students and travellers. – J. Smith, Paris,1-115.

BONIS L. de & LEBEAU M.O. (1974). – A propos d’un crâne de Canidé dé-couvert dans le Quaternaire du Cantal. Etude multidimension-nelle des caractères discriminants entre chiens et loups. –Mammalia, 38 (4), 717-728.

BONIS L. de, PEIGNÉ S., LIKIUS A., MACKAYE H.T., VIGNAUD P. & BRUNET

M. (2005). – Hyaenictitherium minimum, a new ictithere (Mam-malia, Carnivora, Hyaenidae) from the late Miocene of To-ros-Menalla, Chad. – C.R. Palevol (Acad. des Sciences) 4, (8),671-679.

BROOM R. & SCHEPERS G.W.H. (1946). – The South African fossilape-men: the Australopithecinae. – Transvaal Museum Mem., 2,1-272.

BRUNET M., GUY F., PILBEAM D., MACKAYE H. T., LIKIUS A., AHOUNTA D.,BEAUVILAIN A., BLONDEL C., BOCHERENS H., BOISSERIE J. R.,BONIS L. DE, COPPENS Y., DEJAX J., DENYS C., DURINGER P., EI-

SENMANN V., FANONE G., FRONTY P., GERAADS D., LEHMANN T.,LIHOREAU F., LOUCHART A., MAHAMAT A., MERCERON G., MOU-

CHELIN G., OTERO O.,CAMPOMANES P. P., DE LEON M. P., RAGE

J.-C., 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 of Chad, Central Afri-ca. – Nature, 418, 145-151.

DEL CAMPANA D. (1914). – La Lycyaena lunensis n.sp. dell’ossario plioce-nico di Olivola (Val di Magra). – Paleontographia Italica, 20,87-103.

EWER R. F. (1955). – The fossil carnivores of the Transvaal Caves. The Ly-cyaenas of Sterkfontein and Swartkrans together with some ge-neral considerations of the Transvaal fossil Hyaenids. – Proc.Zool. Soc. London, 124, (4), 839-857.

FICCARELLI G. & TORRE D. (1967). – Una mandibola di Euryboas lunensis(DEL CAMPANA) nel giacimento villafranchiano di Olivola (Valdi Magra). – Atti della Società Toscana di Scienze Naturali Resi-dente in Pisa, Memorie, 74, 193-198.

GALIANO H. & FRAILEY D. (1977). – Chasmaporthetes kani, new speciesfrom China, with remarks on phylogenetic relationships of gene-ra within the Hyaenidae (Mammalia, Carnivora). – Am. Mus.Novitates, 2632, 1-16.

GAUDRY A. (1861). – Note sur les carnassiers fossiles de Pikermi (Grèce).– Bull. Soc. géol. Fr., 2, XVIII, 527-538.

GERAADS D. (1997). – Carnivores du Pliocène terminal de Ahl al Oughlam(Casablanca, Maroc). – Geobios, 30 (1), 127-164.

GRAY J.E. (1869). – Catalogue of carnivorous, pachydermatous, and eden-tate Mammalia in the British Museum, London. – British Mu-seum (Natural History), VII, 1-398.

HAY O.P. (1921). – Descriptions of species of Pleistocene Vertebrata: typesor specimens of most of which are preserved in the United Sta-tes National Museum. – Proc. US Nat. Mus., 59, 599-642.

HENDEY Q. B. (1974). – The late Cenozoic Carnivora of the southwesternCape Province. – Ann. South African Mus., 63, 1-369.

HENDEY Q. B. (1978). – Late Tertiary Hyaenidae from Langebaanweg,South Africa, and their relevance to the phylogeny of the family.– Ann. South African Mus., 76 (7), 265-297

HOWELL F.C. (1987). – Preliminary observations on Carnivora from the Sa-habi Formation, Libya. In: N. T. BOAZ, A. EL-ARNAUTI, A. W.GAZIRY, J. HEINZELIN DE, D. DECHANT BOAZ, Eds., Neogene pa-leontology and geology of Sahabi. – Alan R. Liss, New York,153-181.

KHOMENKO I.P. (1932). – Hyaena borissiaki n. sp. from the Roussillon fau-na of Bessarabia. – Travaux de l’Institut paléozoologique del’Académie des Sciences del’URSS, 1, 81-134 (in Russian withGerman summary).

KOUFOS G.D. (1987). – Chasmaporthetes bonisi, a new hyaenid (Carnivo-ra, Mammalia) from the Late Miocene of Macedonia (Greece). –Bull. Soc. géol. Fr., (8), III, 5, 913-920.

KURTÉN B. & WERDELIN L. (1988). – A review of the genus Chasmaporthe-tes HAY, 1921 (Carnivora, Hyaenidae). – J. Vert. Paleontol., 8(1), 46-66.

PEIGNÉ S., BONIS L. de, LIKIUS A., MACKAYE H.T., VIGNAUD P. & BRUNET

M. (2005). – A new machairodontine (Carnivora, Felidae) fromthe Late Miocene hominid locality of TM 266, Toros-Menalla,Chad. – C.R. Palevol (Acad. des Sciences), 4 (3), 243-253.

PILGRIM G. (1931). – Catalogue of the Pontian Carnivora of Europe. – DeptGeol.-British Museum (Natural History), London, 1-174.

QIU ZHANXIANG (1987). – Die Hyaeniden aus dem Ruscinium und Villa-franchium Chinas. – Münchn. Geowiss. Abh., 9, 1-110.

ROOK L., FERRETTI M.P., ARCA M. & TUVERI C. (2004). – Chasmaporthetesmelei n.sp., an endemic hyaenid (Carnivora, Mammalia) fromthe Monte Tuttavista fissure fillings (late Pliocene to early Pleis-tocene, Sardinia, Italy). – Riv. Ital. Pal. Strat., 110 (3), 707-714.

SCHAUB S. (1941). – Ein neues hyaenidengenus von der Montagne de Per-rier. – Eclogae Geol. Helv., 34 (2), 279-286.

STIRTON R.A. & CHRISTIAN W.G. (1940). – A member of the Hyaenidaefrom the Upper Pliocene of Texas. – J. Mammal., 4, 445-448.

TURNER A. (1987). – New fossil carnivore remains from the Sterkfonteinhominid site (Mammalia, Carnivora). – Ann. Transvaal Mus.,34, (15), 319-347.



VIGNAUD P., DURINGER P., MACKAYE H.T., LIKIUS A., BLONDEL C., BOIS-

SERIE J. R., BONIS L. de, EISENMANN V., ETIENNE M.E., GERAADS

D., GUY F., LEHMANN T., LIHOREAU F., LÓPEZ-MARTÍNEZ N.,MOURER-CHAUVIRÉ C., OTERO O., RAGE J.-C., SCHUSTER M., VI-

RIOT L., ZAZZO A. & BRUNET M. (2002). – Geology and palaeon-tology of the Upper Miocene Toros-Menalla hominid locality,Chad. – Nature, 418, 152-155.

VIRET J. (1954). – Le loess à bancs durcis de Saint-Vallier (Drome) et safaune de mammifères villafranchiens. – Nouv. Arch. Mus. Hist.Nat. Lyon, 4, 1-200.

WERDELIN L. & SOLOUNIAS N. (1991). – The Hyaenidae: taxonomy, syste-matics and evolution. – Fossils & Strata, 30, 1-106.

WERDELIN L., TURNER A. & SOLOUNIAS N. (1994). – Studies of fossil hyae-nids: the genera Hyaenictis GAUDRY and Chasmaporthetes HAY,with a reconsideration of the Hyaenidae of Langebaanweg,South Africa. – Zool. Jour. Linn. Soc., 3, 197-217.

ZDANSKY O. (1924). – Jungtertiäre Carnivoren Chinas. – Paleontol. Sin., 2(51), 1-149.



Documents

First occurrence of the 'hunting hyena' Chasmaporthetes in the Late Miocene fossil bearing localities of Toros Menalla, Chad (Africa)