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Nyctereutes lockwoodi, n. sp., a new canid (Carnivora: Mammalia) fromthe middle Pliocene of Dikika, Lower Awash, EthiopiaDenis Geraads; Zeresenay Alemsegeda; René Bobeb; Denné Reedc

a Department of Anthropology, California Academy of Sciences, San Francisco, California, U.S.A. b

Department of Anthropology, University of Georgia, Athens, Georgia, U.S.A. c Department ofAnthropology, University of Texas at Austin, Austin, Texas, U.S.A.

Online publication date: 19 May 2010

To cite this Article Geraads, Denis , Alemseged, Zeresenay , Bobe, René and Reed, Denné(2010) 'Nyctereutes lockwoodi, n.sp., a new canid (Carnivora: Mammalia) from the middle Pliocene of Dikika, Lower Awash, Ethiopia', Journal ofVertebrate Paleontology, 30: 3, 981 — 987To link to this Article: DOI: 10.1080/02724631003758326URL: http://dx.doi.org/10.1080/02724631003758326

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Journal of Vertebrate Paleontology 30(3):981–987, May 2010© 2010 by the Society of Vertebrate Paleontology

SHORT COMMUNICATION

NYCTEREUTES LOCKWOODI, N. SP., A NEW CANID (CARNIVORA: MAMMALIA) FROM THEMIDDLE PLIOCENE OF DIKIKA, LOWER AWASH, ETHIOPIA

DENIS GERAADS,*,1 ZERESENAY ALEMSEGED,2 RENE BOBE,3 and DENNE REED4; 1UPR 2147 du CNRS, 44 rue del’Amiral Mouchez, 75014 Paris, France, [email protected]; 2Department of Anthropology, California Academy ofSciences, San Francisco, California 94118, U.S.A. , [email protected]; 3Department of Anthropology, University ofGeorgia, Athens, Georgia 30602, U.S.A. , [email protected]; 4Department of Anthropology, University of Texas at Austin, Austin,Texas 78712, U.S.A. , [email protected]

The Dikika Research Project (DRP), led by one of us (Z.A.),explores Lower Pliocene to early middle Pleistocene sedimentsin the Lower Awash Valley, Ethiopia, mostly on the right bankof the Awash River, south of the sites of Hadar and Gona (Ger-aads et al., 2004; Alemseged et al., 2005, 2006; Wynn et al., 2006,2008). In the eastern part of the DRP area, exposed on the rightside of the Awash River, the exposures belong primarily to theSidi Hakoma and Basal Members of the Hadar Formation, whichrange in age from >3.8 to <3.2 Ma. The sediments of the SidiHakoma Member (dated from 3.42 to 3.24 Ma; Campisano, 2007)are highly fossiliferous, and have yielded several hominin fos-sil specimens, including an almost complete skeleton of a childAustralopithecus afarensis nick-named ‘Selam’ (Alemseged et al.,2006). Not unexpectedly, hyenas are the most abundant carni-vores, but canids are also common. In this paper, we describe allcanid specimens found to date at Dikika and assign them to asingle new species.

Institutional Abbreviations—CCEC, Centre de Conservationet d’Etude des Collections, Lyon; MNHN, Museum Nationald’Histoire Naturelle, Paris; NME, National Museum of Ethiopia,Addis Ababa; UCBL, Universite Claude Bernard, Lyon.

SYSTEMATIC PALEONTOLOGY

Order CARNIVORA Bowdich, 1821Family CANIDAE Gray, 1821

Genus NYCTEREUTES Temminck, 1838

Type Species—Nyctereutes procyonoides (Gray, 1834).

NYCTEREUTES LOCKWOODI, n. sp.

Type—Nearly complete skull, missing most of the incisors,tips of the canines, and parts of the auditory bullae, DIK-31–1(Fig. 1). The specimen is housed in the NME. It was discoveredby Ali Buh in the Andedo drainage (coordinates WGS 84, UTMzone 37: 0674950, 1226058), at the base of the Sidi Hakoma Mem-ber of the Hadar Formation, and it dates to ca. 3.35 Ma.

Etymology—Named after the late Charles Lockwood, for hiscontribution to our knowledge of the genus Australopithecus inSouth and East Africa as well as his role in the exploration ofthe morphological temporal trends of A. afarensis in the HadarFormation (Kimbel, 2009).

Referred Specimens—DIK-1–22a is a fragment of leftmandibular corpus with m2 and the distal half of m1, both heav-ily worn. DIK-1–22b is a piece from the right side, including theposterior part of the corpus and most of the ramus (Fig. 2). Lo-cality DIK-31 belongs to the same stratigraphic level as DIK-1,where the juvenile Australopithecus skeleton comes from (Alem-seged et al., 2006; Wynn et al., 2006). DIK-70–2 is an incomplete

*Corresponding author.

skull from the slightly younger Denen Dora Member, ca. 3.20 Ma(Fig. 3). All fossils are housed in the NME.

Diagnosis—A canid about the size of a small modern black-backed jackal (Canis mesomelas) with a short and narrow snout;infra-orbital foramen close to the orbit; anterior zygomatic rootabove the back of M2; small teeth, especially the incisors, canines,and premolars; small to vestigial protocone on P4; M1 and M2with paracone and metacone low and sub-equal in size; mandiblewith a well-developed sub-angular lobe.

Description—The type cranium (Fig. 1) is that of an old in-dividual with heavily worn molars; it has suffered minor defor-mation, being slightly compressed dorsoventrally. Most of thebones of the snout are fragmented, but not displaced relative toeach other. The skull, which is the best preserved specimen of aPliocene African canid, is about as large as those of C. aureus orC. mesomelas, slightly smaller than modern Canis adustus, whichare all found in Ethiopia today. Its measurements are given inTable 1.

In lateral view, the skull is more like Vulpes or Nyctereutes thanlike an adult jackal in that the occiput is low because there is notrue sagittal crest; however, the dorsal profile rises less steeplyfrom the nuchal crest than in foxes, because the maximum brain-case width is more anterior. The zygomatic arch is robust; theventral orbital border is not laterally everted, in contrast to thatof foxes, but the skull is broader at this level than at the alve-olar level. The posterior border of the anterior zygomatic rootdescends above the distal part of M2, in contrast to almost allmodern canids, where it descends more rostrally, with most ofM2 being inserted on a caudal expansion of the palate.

In dorsal view, the braincase is less expanded caudally than it isin foxes. The frontal is inflated by sinuses extending into the post-orbital processes, although a hint of a dorsal depression remains.The post-orbital processes are broad, and the skull has a less pro-nounced post-orbital constriction. The temporal lines convergeregularly caudally; they meet near the fronto-parietal suture, butdo not fuse completely. The surface of the braincase is rugose, asis often seen in modern canids. The muzzle is short and narrow,especially behind the canines; the arrangement of its bones doesnot differ from other Canis, Vulpes, or Nyctereutes. The nasalsend caudally at the same level as the maxilla.

The occipital is less triangular than in many adult Canis, wherethe width usually decreases sharply and regularly from mastoidlevel to the sagittal crest; here the nuchal crests converge lessstrongly upwards in their middle part, giving the occipital a morebell-shaped outline (Fig. 1D). The paroccipital process is wellseparated from the bulla, to which it is connected by a bonyflange, but it is short, as it does not extend more ventrally thanthe bulla. No notch separates it from the mastoid.

The auditory bulla is large and somewhat spherical; rostrally, italmost reaches the level of the foramen ovale. The post-glenoidprocess is broad; it extends almost to the lateral side of theglenoid fossa. The palate is narrow.

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FIGURE 1. Nyctereutes lockwoodi, skull DIK-31–1 in A, lateral, B, dor-sal, C, ventral, and D, postero-ventro-lateral views. Scale bar equals 5 cm.

Of the incisors, only the bases of the right I2 and I3 are pre-served. The latter is significantly larger than the former, andseparated from it by a short diastema. The tips of the canines aremissing but these teeth were mesio-distally short and, from theconvergence of their mesial and distal borders, it can be deducedthat they were low-crowned. The anterior premolars are small,simple, and separated by long diastemas. P1 is peg-like, with nohint of a distal cusp. An incipient distal cusp is present on P2,and is more distinct on P3, but remains weak. The carnassials andmolars are also small relative to skull size. P4 is almost twice aslong as P3; its protocone extends farther mesially than the para-cone, but the mesial border of the tooth is not concave in occlusalview, and is surrounded by a well-marked cingulum. All premo-lars are slightly worn, but most of the crown of M1, lingual to theparacone and metacone, is worn off. These cusps are sub-equalin size; labially, they are bordered by a continuous cingulum. M2

TABLE 1. Cranial measurements (in mm) of Nyctereutes lockwoodi.

DIK-31–1 DIK-70–2

E1 Condylo-basal length 143.2 —E7RA6 Length from rostrum to

rear of M274.8 —

RA4 Length of nasals 53 —RA21 Length from orbit to

infra-orbital foramen11.5 —

E17RA12 Bi-mastoid width 52.2 —E18RA10 Bi-condylar width 29.8 —E21RA9 Bi-zygomatic width 77.6 —E22RA15 Width over post-orbital

constriction31 —

E24RA14 Width over post-orbitalprocesses

42 44

E25RA13 Minimum bi-orbital width 27 28.4E9RA8 Length P4–M2 27.2 28.3RA5 Length P1–M2 52 ca. 52

The first column is the measurement number in Eisenmann and van derGeer, 1999 (E), and/or in Rook and Azzaroli-Puccetti, 1996 (RA).

FIGURE 2. Nyctereutes lockwoodi, mandible DIK-1–22: A, photo-graphic reconstruction based upon left (to the right of the dashed line)and right pieces, lateral view; B, detail of the angular process, medialview, with muscle scars for the superficial part of the medial pterygoid(1), and for its deeper part (2). Scale bar equals 2 cm.

FIGURE 3. Nyctereutes lockwoodi, incomplete skull DIK-70–2, occlusalview of P4–M2. Scale bar equals 2 cm.

TABLE 2. Tooth measurements (in mm) of Nyctereutes lockwoodi.

P1 P2 P3 P4 M1 M2

L W L W L W L ext L max W L W perp W max L W

DIK-31–1 4.0 2.7 6.7 2.8 7.5 3.5 12.2 13.6 6.0 9.2+ 11.9 13.1 6.6 8.5DIK-70–2 — — — — — — 12.4 13.1 6.2 10.6 12.1 13.4 6.2 8.6


SHORT COMMUNICATIONS 983

is also much worn and rather large compared to M1. Tooth mea-surements are given in Table 2.

The partial skull DIK-70–2 is less complete than DIK-31–1, butits molars are well preserved and unworn (Fig. 3). We refer it tothe same species, as most of its features match those of the holo-type: (1) overall size is comparable, and measurements are simi-lar (Table 1); (2) the post-orbital processes are strong and salient,with a hint of a dorsal depression, but they are well inflated, anda break across the frontals shows them to be extensively pneu-matized; (3) the posterior border of the zygomatic arch descendsabove the posterior root of M2, instead of anterior to it as in mostcanids; and (4) the infra-orbital foramen opens immediately infront of the anterior root of P4, and is closer to the orbit than it isin most canids.

Differences from the holotype are that: (1) the temporal linesform a low sagittal crest; (2) the orbit is less anterior with respectto the tooth-row, so that the distance between it and the infra-orbital foramen is greater than in the type; and (3) the protoconeof P4 is extremely small, being in fact little more than a slightinflation at the base of the paracone.

The molars of DIK-70–2 are large relative to the carnassial.The M1 is not much broader than it is long, with only a shallowdistal concavity; the paracone and metacone are low and sub-equal, both in surface and height and are bordered by a stronglabial cingulum. The crescent-shaped hypocone is thick and ex-tends mesially and distally into a faint cingulum. Paracone andmetacone are also of similar size on M2, and the hypocone of thistooth is also large.

The teeth of the mandible pieces DIK-1–22 (Fig. 2) are tooworn to offer any useful detail, except that the metaconids ofm1 and m2 are still high, but the occurrence of a cristid betweenhypoconid and entoconid of m1 cannot be ascertained. The m3is represented by its alveolus only. This mandible is remarkablefor the expansion of the sub-angular area, as occurs in someother fossil and modern canids where its role is to provide alarger area for insertion of the digastric muscle. In DIK-1–22, itcan be called a lobe, but is less expanded than it is in modernNyctereutes or Otocyon. Although the incomplete nature of themandible precludes precise orientation, it is also clear that theanterior edge of the ascending ramus is less vertical than it is innormal canids. However, on the medial side of the rather largeangular process (Fig. 2B), the (ventral) scar for the insertion ofthe superficial medial pterygoid is large, as it is in canids with asub-angular lobe, whereas it is narrower than the (dorsal) scarfor the deeper part of this muscle in normal canids (Gaspard,1964; Tedford et al., 1995).

There is no definite evidence that DIK-1–22 belongs to thesame species as DIK-31–1, but they were found not far from eachother in the same layers, they match in size and articulate satis-factorily, and the sub-angular lobe of the mandible fits with thereduced upper premolars. Pending evidence of a second similar-sized canid in the lower Sidi Hakoma Member at Dikika, we as-sign these specimens to the same species.

Comparisons and Discussion—The Dikika canid is clearly notrelated to the specialized carnivorous Cuon and Lycaon. It is alsomuch smaller than Canis of the wolf group, so that comparisonscan be restricted to Vulpes, jackals (Canis adustus, C. aureus,C. mesomelas), the living C. simensis of Ethiopia, canids of theEucyon group, and Nyctereutes.

In size (condylo-basal length) it is intermediate between thesmaller Vulpes and the larger Canis, although these genera dooverlap somewhat in this respect. It is also intermediate in sev-eral other measurements, such as length of the P4–M2 toothrow (for which there is almost no overlap between Canis andVulpes), and most measurements of width: muzzle width acrossbases of canines, bi-zygomatic width, and widths across the or-bits, the post-orbital processes, the post-orbital constriction, thebraincase, the mastoids, or the occipital condyles (measurements

FIGURE 4. Plot of distance from orbit to infra-orbital foramen (mea-surement 21 of Rook and Azzaroli-Puccetti, 1996) versus condylo-basallength. Fossil Nyctereutes are N. donnezani from Calta (Ginsburg, 1998;MNHN), N. megamastoides (or N. vulpinus according to Monguillonet al., 2004) from Saint-Vallier (Viret, 1954; CCEC), and N. sinensisfrom Nihowan (Teilhard and Piveteau, 1930; MNHN). All measurementstaken by the authors, except some from Rook and Azzaroli-Puccetti(1996).

taken according to Eisenmann and van der Geer, 1999, and Rookand Azzaroli-Puccetti, 1996). It is significantly larger than livingNyctereutes procyonoides, but similar in size to fossil forms of thisgenus.

The Dikika canid differs from most or all modern canids ofsimilar size in four distinctive characters:

1. The short distance between the orbit and the infra-orbitalforamen, which is very close to P4. In other species, the infra-orbital foramen is more anterior, between P4 and P3 or evenabove P3, and it is farther from the orbit (Fig. 4).

2. The lack of posterior extension of the palate behind the zy-gomatic root. In most canids, the anterior root of the zygomaoriginates above the anterior root of M2 or between M1 andM2, so that most of M2 is borne by a posterior expansion ofthe palate. In the Dikika canid, it originates above the poste-rior border of M2.

3. The small size of its premolars (Figs. 1, 5, and 6). Only someCanis adustus, a single specimen of N. megamastoides, and thelarger C. simensis may have premolars as small as at Dikika.

FIGURE 5. Plot of sum of the lengths of P2, P3, and P4 versuscondylo-basal length. Fossil Nyctereutes are N. donnezani from Perpignan(Deperet, 1890; CCEC), N. megamastoides from Dafnero (Koufos, 1993),Perrier (Boule, 1889; MNHN), La Puebla (Kurten and Crusafont, 1977),Saint-Vallier (Viret, 1954; CCEC), and N. sinensis from Nihowan (Teil-hard and Piveteau, 1930; MNHN). All measurements taken by the au-thors, except some from Kurten and Crusafont (1977) and Koufos (pers.comm.).



FIGURE 6. Plot of labial length of P4 versus length of M1 + length ofM2. Fossil Nyctereutes as in Figures 4 and 5, plus N. abdeslami from Ahl alOughlam (Geraads, 1997), N. donnezani from Layna (Rook et al., 1991),and N. sinensis from Zhoukoudian (Pei, 1934).

It should be noted, however, that the often-used ratio of thelength of P4 to those of the molars (Fig. 6) separates most C.mesomelas and C. aureus from C. adustus, but not the latterfrom Nyctereutes.

4. The small size of the protocone of P4. In other canids, the sizeand position of the protocone is variable (it may or may notreach farther anteriorly than the paracone), but it is never asreduced as in DIK-70–2. Even Dusicyon australis, said by Ted-ford et al. (1995) to have the smallest P4 protocone of all mod-ern canids, has a larger one than DIK-70–2. This cusp is larger,although still small, on DIK-31–1, but even on this specimen,the mesial border of the carnassial lacks the concavity betweenparacone and protocone that is virtually constant in modernCanis, Vulpes, and Nyctereutes.

In contrast to Tedford et al. (1995) and Zrzavy and Ricankova(2004), we found no consistent difference between Vulpes (wehave examined the two species of comparable size, V. vulpes andV. rueppelli) and Canis in the relative positions of the bulla andparoccipital process, but the latter is definitely short in DIK-31–1,as in Vulpes, whereas its mastoid is definitely laterally expanded,as these authors have reported for Canis. The Dikika canid alsodiffers from Vulpes in: (1) the lack of eversion of the ventral or-bital border; (2) the absence of conspicuous dorsal depression ofthe post-orbital processes of the frontal; (3) the much weakerwaist behind the latter; (4) the absence of a notch between theparoccipital process and the mastoid process; (5) the large I3 rel-ative to I2; (6) the low (short) canine; (7) the longer M1 relativeto P4; and (8) the larger metacone of M1 and M2. No synapo-morphies are shared by Vulpes and the Dikika canid, so that itsassignment to that genus can be ruled out.

In addition to the four distinctive characters mentioned previ-ously, jackals further differ from the Dikika canid in their broadermuzzle, in the larger and higher paracone of M1, and in the P4that is usually longer relative to M1. Canis adustus, which is thespecies most similar to the Dikika canid in its small premolars(both relative to the molars and to skull length), has a deep facewith an inflated fronto-nasal area quite unlike the Dikika canid, aslender zygomatic arch, a broad palate, a long canine, a large P4protocone, and is thus clearly distinct.

The Ethiopian C. simensis is phenetically close to jackals(Rook and Azzaroli-Puccetti, 1996), but is probably closely re-lated to wolves and coyotes (Gottelli et al., 1994; Lindblad-Tohet al., 2005). It resembles the Dikika canid in its long M1 relativeto P4, short anterior premolars relative to skull size, and zygo-matic arch descending above the back of M2, but differs in its

much larger size, long and narrow muzzle, broad incisor arch, tri-angular occipital, and strong sagittal crest.

The genus Eucyon was erected by Tedford and Qiu (1996) forEucyon davisi of North America and China. The original diagno-sis of this genus listed only, as distinctive characters from Canis,the lack of transverse cristid between the hypoconid and ento-conid of m1, and the presence of a second posterior cusplet onp4, the single apomorphic feature of the genus. The Dikika caniddiffers from the type-species, and from another Chinese species,E. zhoui, in that its infra-orbital foramen is much closer to theorbit, in its much smaller anterior premolars, which are as largein Eucyon as in Vulpes, in its less marked post-orbital constric-tion, broader post-orbital processes, and less expanded zygomaticarches.

In recent years, a number of other Pliocene Old World canidshave been assigned to this genus (Koufos, 1997; Kostopoulosand Sen, 1999; Morales et al., 2005; Spassov and Rook, 2006;Howell and Garcıa, 2007; Garcıa, 2008) but none of these re-cent studies have attempted to delineate the genus; significantly,no revised diagnosis has ever been provided, and the result isthat, in the Old World, ‘Eucyon’ has become a waste baskettaxon for early canids. We consider the scanty African mate-rial from Lukeino (Morales et al., 2005) and Lemudong’o (How-ell and Garcıa, 2007) unidentifiable below family level. A max-illa from Aramis, ca. 4.4 Ma, referred by Garcıa (2008) to hernew species Eucyon wokari, is more significant. It shares withthe Dikika canid a posteriorly situated infra-orbital foramen andsimilar tooth measurements and morphology, especially the lowlabial cups on M1. Differences are the more anterior zygomaticroot, P3 closer to P4, and occurrence of an M3, probably an in-dividual atavistic feature. These differences are not great but weprefer not to assign the Dikika form to ‘wokari,’ because: (1) thetype mandible of this species shows no deepening under m2 andthus certainly it lacked a subangular lobe; (2) there is no evidencethat the Aramis form had small teeth for its skull size; and (3) thesuspicion of co-specificity that would have been provided by con-temporaneity is lacking. In any case the Dikika canid cannot beincluded in Eucyon, as the apomorphic character of this genus,the second posterior cusplet of p4, would not fit the reducedP4 protocone and reduced anterior premolars of the Dikikacanid.

Barry (1987) assigned to aff. Canis brevirostris Ewer, 1956 (aspecies defined at Sterkfontein), a partial skull LAET-3562 (withmost of the teeth) and associated mandible LAET-3522, fromLaetoli, a site slightly earlier than DIK-31. The size and pro-portions of the teeth are similar to those of DIK-31–1, but theinfra-orbital foramen is more anteriorly placed (above P3), theanterior premolars are much more crowded (length from the dis-tal border of the canine to the anterior border of P4 is 84% thatof the length P4–M2, versus 104% at Dikika); the labial cuspsof M1 are also more unequal in size, and the protocone of P4 isstronger. The Laetoli canid was first considered closely relatedto Eucyon by Werdelin and Lewis (2000), but later the same au-thors (2005) pointed to its affinities to Nyctereutes, although themandible lacks a sub-angular lobe, another difference with theDikika form.

From South Turkwell, a site roughly contemporaneous withthe SH Member of the Hadar Formation, Werdelin and Lewis(2000) assigned two mandibular fragments of a canid much largerthan the Dikika form to a new unnamed species of Canis. FromOlduvai Bed I, Petter (1973) described a canid that is also largerand has much longer P4s than the Dikika form as Canis mesome-las and it is indeed closer to modern jackals,

From Ahl al Oughlam in Morocco, dated to ca. 2.5 Ma, Ger-aads (1997, 2008) described a canid the size of modern jackalsand slightly larger than the Dikika form as Canis aff. aureus. Al-though its anterior upper premolars are unknown, it is very dis-tinct from the Dikika specimen in its normal-sized protocone on



P4, its large paracone, and reduced lingual half of M1, and in theposition of M2 behind the zygomatic root, but the infra-orbitalforamen is above the tip of the anterior root of P4, and is thusmore posterior than in modern forms.

In Europe, Canis adoxus Martin, 1973, from Saint Esteve inFrance is roughly contemporaneous with the Dikika canid, andresembles it in its small anterior premolars separated by di-astemas (Martin, 1973; Rook, 1992). The type-skull is dorsoven-trally crushed, but it certainly differed in its larger size, narrowskull with stronger post-orbital constriction and especially nar-row anterior muzzle, orbit far from the infra-orbital foramen,posterior root of zygomatic arch above the middle of M2, largercanine (the difference is much greater that what could be ex-plained by sexual dimorphism), protocone of P4 of normal size,and high paracone of M1. Except for its narrow muzzle, it resem-bles C. simensis.

The occurrence of a subangular lobe in the DIK-1–22 mandiblewarrants special comparison with Nyctereutes, of which the livingrepresentative, the Palaearctic N. procyonoides, is more closelyrelated to foxes than to Canis (Bardeleben et al., 2005; Lindblad-Toh et al., 2005). The Dikika canid resembles N. procyonoides inhaving a cranium that is not very constricted behind the orbits, ashort paroccipital process, and a short P4 relative to M1 and M2(Fig. 6), but the Dikika canid differs in the four characters pre-viously mentioned (the small P4 protocone contrasts with a verylarge one in the raccoon-dog), as well as in its larger size, longerand narrower skull, weaker post-orbital processes, broader post-glenoid process, more curved zygomatic arch, longer premaxillae,more extensive frontal sinus, larger I3 relative to I1 and I2, shortcanine, and broader M1. Except the small canine, unique to N.lockwoodi, n. sp., these characters (several of which were notedby Tedford et al., 1995) also distinguish Canis as a whole frommost Nyctereutes.

The fossil forms of this genus are larger than the living one.Late Pliocene to Pleistocene forms, N. megamastoides from Eu-rope (Viret, 1954; Kurten and Crusafont, 1977; Rook et al., 1991),N. abdeslami from Morocco (Geraads, 1997), and N. sinensisfrom China (Teilhard and Piveteau, 1930; Pei, 1934; Tedford andQiu, 1991) share the same characters as N. procyonoides shareswith the Dikika form, but also the same differences, except size.They have small anterior premolars (perhaps a negative allome-try by comparison with N. procyonoides), but other dental andcranial features (including the sub-angular lobe, although it isusually smaller) indicate close relationships with the living form.

In South Africa, Ficcarelli et al. (1984) referred to Nyctereutesthe associated type skull and mandible of Thos terblancheiBroom, 1948, from the Lower Pleistocene of Kromdraai A, on thebasis of its sub-angular mandibular lobe. Other features of thisspecimen (of which we have seen a good cast and photographs)are: the post-orbital process is strong, but with a slight dorsaldepression; the upper canine is low; the anterior premolars aresmall; the crushing part of the dentition is large, with m2 broaderthan m1, and a large m3 (some of these characters were noted byEwer, 1956). These are all features consistent with Nyctereutes,and we therefore support assignment of the Kromdraai A speci-men to this genus, in spite of its large size (larger than any otherspecimen of this genus) and the fact that the size of the protoconeof P4 is unknown.

The earlier N. donnezani (Soria and Aguirre, 1976; Ginsburg,1998), which probably includes some specimens described asN. sinensis (Czyzewska, 1969) or N. tingi (Koufos, 1997), has alonger skull and at most a poorly expressed sub-angular lobe,but two characters show that it is related to the previous species:strong post-orbital processes and strong protocone on P4. Thiswould rank these characters above the large size of the crush-ing part of the dentition (which is also seen in C. adustus and C.simensis among living forms) as the best diagnostic features ofNyctereutes, but they are less clearly expressed in another primi-

tive species, Nyctereutes tingi Tedford and Qiu, 1991, from China.The latter has a poorly expanded sub-angular lobe, premolarsthat are not so reduced as at Dikika, and a short, high skull,so that this species is certainly distinct from the Dikika canid.However, it also suggests that the long flattened post-orbital pro-cesses and enlarged P4 protocone were absent in some basalNyctereutes, thereby removing these from the list of differencesbetween the Dikika canid and this genus, provided that N. tingireally belongs here.

The generic assignment of the Dikika canid is not straightfor-ward. Its unusual features raise the possibility that it belongs toa new genus, but there would be little support for defining it,and we prefer to hold a conservative view, although the pastgeneric diversity of this family is probably underestimated. Thetooth proportions, thick but low sagittal crest, short and narrowsnout, and (if the mandible really belongs to the same species)sub-angular lobe and somewhat expanded angular process are allfeatures of Nyctereutes, even though most if not all of these char-acters are functionally related (increase of chewing versus slic-ing and especially regarding the role of the front teeth in seizingpreys). Still, most of these characters are not found in Canis, andtip the balance in favor of Nyctereutes, so that assignment to thisgenus is the most satisfactory option. It also explains the sharedcharacters with Nyctereutes terblanchei: small canine and premo-lars, large size of the molars, and subangular lobe. The Dikikacanid is earlier, and more primitive in its smaller size, less salientpost-orbital processes, weaker sub-angular lobe, and less uprightascending ramus, but they may be phyletically related.

Conclusion—Nyctereutes lockwoodi, n. sp., belongs to a typeof canid unknown in Africa today. Its small canines and pre-molars beside large crushing teeth strongly suggest an omnivo-rous diet like that of modern N. procyonoides and of the AfricanC. adustus, which have comparable dental adaptations. Its nichemay have been similar to that of this jackal, but its relativeabundance compared to other contemporaneous carnivores sug-gest that it might have extended to niches today occupied bylarger (civet, honey badger) or smaller (Otocyon) forms; the guildof medium-sized generalist carnivores is still too poorly docu-mented at Dikika for detailed ecological assessments.

Its assignment to Nyctereutes also raises interesting biogeo-graphic and phylogenetic issues. It is certainly not closely relatedto the contemporaneous European and Turkish N. donnezani,nor to the later N. abdeslami from Morocco, and we must ratherimagine an immigration from Southern Asia of a form related toN. tingi, perhaps also ancestral to N. terblanchei (whose more def-inite Nyctereutes features would therefore be parallel acquisitionswith Palaearctic forms?).

It is also interesting that, like modern Nyctereutes, it is phe-netically more different from Vulpes than it is from Canis, al-though genetic data (Wang et al., 2004; Bardeleben et al., 2005;Lindblad-Toh et al., 2005) unambiguously show the latter to beremote from Nyctereutes. Other discrepancies between molecularand morphological data (such as the early divergence of the side-striped and black-backed jackals from other Canis s.l., includingdhole and wild dog) show that the various cladograms based upongenetic data should not be taken at face value, but also highlightthe amount of parallelism in the history of the family, still verypatchily documented in Africa.

ACKNOWLEDGMENTS

We are grateful to Jara Hailemariam (ARCCH), MamituYilma (NME), and the regional states for allowing us to workin the field and at the Museum. We thank the Max Planck So-ciety for partly supporting field work at Dikika. We also thankJ. Wynn for his invaluable help in the field and comments onthe manuscript, C. Argot, J. Lesur-Gebremariam (MNHN), D.Berthet (CCEC), and A. Prieur (UCBL) for providing access



to collections in their care, J. Barry, B. Engesser, G. Koufos, S.Potze, N. Spassov, F. Thackeray, and J. Zrzavy for providing un-published data, casts, or photos, and the reviewers, X. Wang andL. Werdelin, for their enlightened comments on the manuscript.

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Submitted April 21, 2009; accepted October 20, 2009.
