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PresenceofthegenusCuoninupperPleistoceneandinitialHolocenesitesoftheIberianPeninsula:newremainsidentifiedin...

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Journal of Archaeological Science 37 (2010) 437–450

Contents lists avai

Journal of Archaeological Science

journal homepage: ht tp: / /www.elsevier .com/locate/ jas

Review

Presence of the genus Cuon in upper Pleistocene and initial Holocene sitesof the Iberian Peninsula: new remains identified in archaeological contextsof the Mediterranean region

Manuel Perez Ripoll a, Juan V. Morales Perez a,c, Alfred Sanchis Serra b,c,*, J. Emili Aura Tortosa a,Inocencio Sarrion Montanana b

a Departament de Prehistoria i Arqueologia, Facultat de Geografia i Historia, Universitat de Valencia, Blasco Ibanez 28, 46010 Valencia, Spainb Museu de Prehistoria de Valencia, SIP (Servei d’Investigacio Prehistorica), Diputacio de Valencia, Corona 36, 46003 Valencia, Spainc Sociedades Cazadoras–Recolectoras Prehistoricas Research Group, Universidad de La Laguna, Tenerife, Spain

a r t i c l e i n f o

Article history:Received 20 July 2009Received in revised form1 October 2009Accepted 6 October 2009

Keywords:CuonCanidaeTaxonomyTaphonomyMorphologyOsteometry

* Corresponding author at: Museu de Prehistoria dvestigacio Prehistorica), Diputacio de Valencia, CoronTel.: þ34 963 883 597; fax: þ34 963 883 536.

E-mail address: alfred.sanchis@dival.es (A. Sanchis

0305-4403/$ – see front matter � 2009 Elsevier Ltd.doi:10.1016/j.jas.2009.10.008

a b s t r a c t

The dhole (Cuon alpinus) has been documented on the Iberian Peninsula throughout the Middle andUpper Pleistocene in archaeological and palaeontological contexts. However, this is an uncommonspecies among the Iberian quaternary faunae as a whole. This paper focuses on the remains of the genusCuon recovered in three different sites of the Iberian Mediterranean region and their morphological andosteometrical study. By comparing these with fourteen complete skeletons of Canis lupus, as well as withother remains of C. alpinus, we intend to highlight some anatomical and metrical differences betweenCuon and Canis that, added to those already published, could be a functional base-line for the identifi-cation of this genus. The identification of this species is complex, especially in archaeological contexts,and in some cases the dhole has been identified as Canis or Canidae. Furthermore, this paper explores thechronology of the presence of the dhole in the Iberian Peninsula and the different contexts where theseanimals have been recovered from.

� 2009 Elsevier Ltd. All rights reserved.

1. Introduction

The dhole, Asiatic wild dog, Indian wild dog, or red dog (Cuonalpinus Pallas 1811) is a middle-sized Canidae, shorter but sturdierthan a wolf. Nowadays, the dhole is present in various types ofbiotopes in various regions of India, western China, and most ofSoutheastern Asia including the western islands of Indonesia(Durbin et al., 2004). Throughout the Pleistocene the geographicaldistribution of this genus was wider, and it is found in central andeastern Europe (for instance, Adam, 1959; Baryshnikov, 1996;Wolsan, 1993) and even in western Europe (for instance, Bonifay,1971; Bouchud, 1951; Boule and de Villeneuve, 1927; Cordy, 1983;Cregut-Bonnoure, 1996; Garcıa, 2003:145), where three differentspecies have been described for this period (Bonifay, 1971; Cregut-Bonnoure, 1996; Garcıa, 2003:146): Cuon stehlini, Cuon priscus andC. alpinus, being latter the most recent and the smallest one. While

e Valencia, SIP (Servei d’In-a 36, 46003 Valencia, Spain.

Serra).

All rights reserved.

some authors have considered these three species as part of thesame phylogenetic line (Adam, 1959), others have argued that theyare independent (Bonifay, 1971).

On the Iberian Peninsula, this species was identified for the firsttime by Schlosser (1923), who called the species as Cuon bourreti(Altuna, 1983). The oldest remains of C. alpinus have been docu-mented in the Trinchera–Galerıa Middle Pleistocene site in Sierrade Atapuerca (Burgos) (Cervera et al., 1999; Garcıa, 2003:146;Garcıa and Arsuaga, 1998). During the Upper Pleistocene thisspecies was present in several sites from the central (Altuna, 1973),western (Cardoso, 1992, 1993), southern (Barroso et al., 2006),northern (Altuna, 1981, 1986; Castanos, 1987), northeastern andeastern peninsular regions (Blasco, 1995; Martınez, 1996, 2001;Martınez and Sarrion, 2001; Vinas and Villalta, 1975). Furthermore,other remains have been recovered from palaeontological siteswhich are devoid of a clear chronological ascription or entirelydecontextualised (Altuna, 1983; Castanos, 1988; Schlosser, 1923).However, this can be considered as an uncommon species amongstthe Iberian quaternary faunae as a whole.

This paper presents several osseous remains of Cuon recoveredfrom three sites from the central area of the Iberian Mediterraneanregion with a chronological sequence that covers almost all the

Fig. 1. Sites of the Iberian Peninsula where Cuon alpinus is present.

Table 1Description of the morphological characters of the cranium of Cuon alpinus as compared

Cranium Cuon alpinus

Frontal–Dorsal Medial depression of the frontals (a) Flat or convexTemporal line (b) Less markedZygomatic process of the frontal (c) Short and bluntFronto-nasal suture (d) Wide, with the v

zygomatic procesNasal process Short, wide and tNasals (e) Wide

Nuchal Nuchal crest (a) Wide, triangularpronounced occi

Mastoid crest (b) Slightly developeNuchal tubercles (c) Slightly develope

is hardly markedZygomatic process of the temporal (nuchalregion) (d)

Robust

Lateral Sagittal crest (a) Slightly markedExternal occipital process (b) Less pronouncedZygomatic process of the frontals andsupraorbital border (c)

In a more basal-l

Nasal shape (d) ConvexMaxillary-nasal region (e) Short and wideLenght Infraorbital border-foramen (f) ShortIntraorbital border (g) Wide and bluntFacial crest of zygomatic (h) High and slightlyZygomatic arch (i) FlatRetroarticular process (j) Small

Basal Basal region of the occipital (a) Well-marked lon

Oval foramen and pterygopalatine crest (b) Pterygopalatine c

Basisphenoid (c) WideMaxillae and palatines (d) Short and wide. TDiastema C1-P1 (e) ShortDiastemas between premolars (f) AbsentPalatine fissure (g) Long and wideL between I1 alveolus-oral border of the palatinefissure (h)

Short

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450438

Upper Pleistocene and up to the Initial Holocene (MIS 4-1). We havecarried out a study and a revision of, on the one hand, some remainsfrom old excavations deposited in the Museu de Prehistoria deValencia. For instance, the hemimandible from the Mousterianlevels of Cova Negra (Xativa, Valencia), firstly identified as Canislupus (Perez, 1977) and currently as Cuon cf. alpinus, and severalcranial and post-cranial bones from Cova del Parpallo (Gandia,Valencia), remains that were partially described in a preliminarwork (Martınez and Sarrion, 2001) and identified as C. alpinus. Onthe other hand, we present two remains recently discovered in theCoves de Santa Maira site (Castell de Castells, Alacant): a posteriorhalf of a cranium recovered from the E chamber levels in 2007 andclassified as C. alpinus, and an upper canine discovered in theEpipaleolithic levels of another chamber from the karstic complexwhich had been previously classified as Canidae (Aura et al., 2006)and currently as cf. Cuon (Fig. 1).

This paper focuses on the newly identified remains of Cuon andtheir morphological and metrical study. Moreover, by comparing itwith C. lupus we intend to highlight some anatomical differencesbetween the genera. Added to those already published, thesedifferences could be a functional base-line for the identification ofthe genus. Furthermore, this paper explores firstly, the chronologyof appearance and disappearance of the genus Cuon from thearchaeological and palaeontological register in the Iberian Penin-sula; and secondly, the different contexts, natural or anthropogenic(cultural), from which these animals have been recovered.

to present-day Canis lupus.

Canis lupus

ConcavePronouncedRobust and pronounced

ertex opposing the frontals.

Narrow, with the vertex in a lower position.

riangular Long and narrowLong and narrow

and flat, with a slightlypital protuberance

Narrow and oval. The occipital protuberance iswell-developed.

d Pronounced and sharpd. The sinus between tubercles.

Larger and well-developed. The sinus is deeper.

Thin

Well-developedthan in Canis Well-developed

ateral position In a more dorsal-frontal position

ConcaveLong and thinLongThin and sharp

developed. On a basal position and well-developedArched and highWell-developed. It reaches the base of theauditory bulla.

gitudinal crest Smooth or with a poorly developed longitudinalcrest

rest ends on the oval foramen Pterygopalatine crest continues in a crest overthe foramen and reaches the retroarticularprocessThin and long

he palate is triangular Thin and longLargePresentShort and narrowLong

Fig. 2. Crania of Canis lupus (female and male) (up) and Cuon alpinus from Obarretaand Parpallo (down). Dorsal view. Letters refer to Table 1.

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 439

2. Materials and methods

Comparative anatomy has allowed us to identify the remainsfrom Cova Negra as Cuon cf. alpinus, those from Cova del Parpalloand the cranium from Coves de Santa Maira as C. alpinus, and thecanine from Santa Maira as cf. Cuon. As a reference for this study wehave used the complete skeleton of the C. alpinus from Obarreta(Altuna, 1983) and a hemimandible (BI/2848) from the Durangue-sado region (Castanos, 1988), both deposited in the MuseoArqueologico, Etnografico e Historico Vasco of Bilbao. The collec-tion has also been compared with fourteen complete skeletons of C.lupus – four females (EBD23272, MNCN16167, MNCN16162 andMNCN21561), seven males (EBD15903, MNCN16166, MNCN16241,MNCN16225, MNCN16239, MNCN16242 and MNCN16153) andthree of undetermined sex (MNCN16235, MNCN16218 andMNCN16219) – from the Estacion Biologica de Donana and theMuseo Nacional de Ciencias Naturales. All these animals have thedefinitive dentition with different wear stages and they have theirlong bones ossified. The measurements obtained from these wolvesare very similar to the published data from the Upper Pleistocene

and Initial Holocene wolves from the Iberian Peninsula (Altuna,1972; Castanos, 1987; Nadal, 1998). All these measurements havebeen included in the graphs. The metrical study has been carriedout using the measurements proposed by von den Driesch (1976).Osteometric tables show the measurements of the new remains ascompared to those of the Cuon remains from Trinchera–Galerıa(Cervera et al., 1999; Garcıa, 2003:153, 155, 164, 165; Garcıa andArsuaga, 1998), Moros de Gabasa (Blasco, 1995), Boquete de Zafar-raya (Barroso et al., 2006), Los Casares (Altuna, 1973), Cova Negra(Martınez, 1996, 2001), Gruta do Escoural (Cardoso, 1992), La Riera(Altuna, 1986), Rascano (Altuna, 1981), Bolinkoba (Castanos, 1987),Amalda (Altuna, 1990) Obarreta (Altuna, 1983) and Duranguesado(Castanos, 1988).

2.1. Sites and remains

2.1.1. Cova Negra (Xativa, Valencia)This Mousterian site is located in Xativa (Valencia) and has one

of the most complete Upper Pleistocene sequences (MIS 5-3) of theIberian Mediterranean region (Aura et al., 2002; Villaverde, 1984;Villaverde et al., 1996). Some studies have pointed out the presenceof C. alpinus on this site. It was identified on the Wurm III (MIS 3)levels excavated in the 1980s (Martınez, 1996, 2001; Martınez andSarrion, 2001).

A Canidae hemimandible was recovered from the 1950s exca-vations carried out by F. Jorda and was first classified as C. lupus(Perez, 1977). Since then, it has been reanalyzed by the authors andhas been identified as Cuon cf. alpinus. We prefer to maintain thespecies as cf. alpinus due to the large size of the mandible and to thefact that M2 has two roots (archaic features). It is a well-preservedfragment of a left hemimandible (catalog no. 30246; CN 9830; taludentrada, capa 7), with complete premolar and molar series (P1-M2).M3 is nonexistent, but there is a space between M2 and the anterioredge of the ramus. A small microalveolus-like hole is visible in thisspace. The anterior part of the mandible is fragmented in the canineregion, and the posterior one in the base of the ramus. Both anteriorand posterior regions seem to present post-depositional fractures.This bone was recovered from a Wurm I (MIS 5d-b) context and wasfound associated to Paracharentian industries and Stephanorhinushemitoechus, Equidae, Bos primigenius, Caprini, Cervus elaphus,Capreolus capreolus, Ursus arctos, Felis sylvestris and Oryctolaguscuniculus remains as well as Crocuta crocuta coprolites.

2.1.2. Cova del Parpallo (Gandia, Valencia)The collection studied here consists of an almost complete

cranium, two hemimandibles, several fragments of the rightscapula, the left radius, the right femur, and an ilium fragment(catalog no. 24210; galerıa, ultima capa). Leaving aside recentfractures, the bones are almost all complete and belong to the sameadult individual, who probably died of natural causes; they do notshow anthropogenic marks. Some of the remains were alreadyknown and had been partially described (Martınez and Sarrion,2001).

The collection was recovered from the basal levels of a smallgallery within the cave during the excavation carried out by Ll.Pericot in 1930; levels which must be prior to Gravettian levels.Although the deepest of Gravettian artifacts reach a similar depth,the clear Gravettian occupations overlap with these levels (Aura,1995; Pericot, 1930, 1942).

The cranium is complete, except for the right zygomatic arch,the anterior half of the nasal bones, and the base of tympanicbullae. The upper teeth – right C1, P4, M1 and left M1 – are held inthe maxilla alveoli. The crowns of P4 and M1 show post-deposi-tional fractures. Left P3 and right P4 have been recently fractured

Fig. 3. Crania of Cuon alpinus from Parpallo (left, up), Santa Maira (left, down) and Obarreta (down right), and Canis lupus (up right). Nuchal view. Letters refer to Table 1.

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450440

and their roots are held in the alveoli. The rest of the dentition waslost post-mortem.

The left hemimandible is fragmented in the ramus region.Dentition is partially preserved. P2, P3, P4 and M2 were lost post-mortem. P1 and M1 present recent fractures. The right mandible iscomplete. P2 and M1 are complete and held in the alveoli. M2 waslost post-mortem, and P3 and P4 present recent fractures as do theincisors and the canines.

2.1.3. Les Coves de Santa Maira (Castell de Castells, Alacant)This is a karstic complex with several entrances located on the

right bank of the Famorca ravine. The two main entrances showdifferent but complementary archaeological sequences. Thedeposit in entrance W spans the final Upper Paleolithic, Epi-paleolithic, Mesolithic and Neolithic, with Bronze Age, Roman andmedieval artifacts at surface levels. An upper right canine identifiedas cf. Cuon (SM04 AA5/1 3.23) has been recovered from the lowerlayers of Unit 3 (ca. 8500–9000 years BP) (Aura et al., 2006). Thetooth does not present any kind of wear, and should belong toa young or subadult individual.

In chamber E (called Corral del Gordo), the majority of theHolocene sequence is missing. Several Neolithic fumier levels (UnitI) have been described here which are superimposed to Pleistocenelevels (Unit II) in erosive contact. Magdalenian and Solutreanremains have been recovered from this unit. The sequence,however, is still under study. A complete Cuon neurocranium(SM07M7/6 II.2 inv. 5) was discovered in this unit covered byyellow sands sediment and next to the external limit of a concen-tration of small and middle-sized stones and blocks that could bethe remains of a structure located in the central layers of Unit II.

There are no direct datings for this layer, but this sequence is datedbetween 11 and 14 Ky BP (Aura et al., 2006). The cranium is frag-mented on the zygomatic process of the frontal bones and thezygomatic arches are also missing.

3. Discussion

3.1. Anatomical and metrical characteristics of the Cuon remains.Comparison with C. lupus

3.1.1. Cranium from Cova del ParpalloThis cranium is short, mainly in its anterior half, and broad in

dorsal and basal views with wide open zygomatic arches whichgive it a sturdy appearance. When viewed laterally, the cranium ismore convex than the Canis ones, with the eye sockets in a lowerposition. The outline of the fronto-nasal region is convex. All thesefeatures are characteristics of the genus Cuon, and have beendescribed and compiled in previous studies (Altuna, 1983; Cregut-Bonnoure, 1996).

3.1.2. Cranium from Coves de Santa MairaOnly the neurocranium is preserved here, its features being

similar to those observed in the Cranium from Parpallo.Both crania have been compared with those of C. lupus in order

to observe their distinctive features which have been compiled inTable 1. This compilation allows a detailed description of thedifferences between both genera (Table 1; Figs. 2–5).

The cranium recovered in Parpallo and the one from Obarretaare the best preserved in the whole of the Iberian Peninsula. Thecranium from Parpallo is longer than that from Obarreta, but clearly

Fig. 5. Crania of Canis lupus female (up) and Cuon alpinus from Obarreta, Santa Mairaand Parpallo (down). Basal view. Letters refer to Table 1.

Fig. 4. Crania of Canis lupus famale and male (up) and Cuon alpinus from Santa Mairaand Parpallo (down). Lateral view. Letters refer to Table 1.

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 441

shorter when compared to the crania of both males and females C.lupus. The length of the Parpallo cranium is significantly longerthan that obtained from the Obarreta cranium, but clearly smallerthan those from males and females C. lupus. The differences in sizebetween Parpallo and Obarreta individuals could be related tosexual dimorphism. The Cuon from Obarreta was identified asa possible female (Altuna, 1983), although studies in living pop-ulations demonstrate that there are no significant differencesbetween male and female (Durbin et al., 2004). Measurements ofthe cranium from Santa Maira are closer to those from the one fromParpallo. Differences in width between the crania of the Cuon andthe C. lupus are slighter and, in some cases, the Cuon cranium iswider when compared to those of the C. lupus (for instance, Table 2,n� 29). The cranial height measurements show that the crania of C.lupus are higher than those of Cuon due to the greater developmentof the sagittal crest in C. lupus (Table 2, n� 38 and 39). Thus, C.alpinus crania are shorter than those belonging to C. lupus, butsimilar in width, and their appearance is sturdier and more glob-ular, ‘‘more feline’’ as described by Boule and de Villeneuve (1927)(Table 2).

3.1.3. Cranium from Cova del Parpallo: upper dentitionLeft C1 shows an important wear facet on the mesial area. The

occlusal wear facet is typical of aged individuals. The posterior edgeof the crown is fragmented and affects the greater part of the distalridge. A wear facet is observed on the mesial–lateral edge of thecrown and root contact area. The posterior edge is quite straight(half-moon outline), and the base of the crown is straight, regular

and quite horizontal. The crown is shorter when compared to thosefrom the C. lupus (Tables 3 and 4; Fig. 6).

Left P4. The anterior side is partially fragmented. The deuter-ocone is more displaced distally than the paracone, as has beenobserved in the Cuon from Obarreta (Altuna, 1983) (Table 5).

Both left and right M1 are preserved, but the left one is frag-mented on its lingual side. It is triangular in shape with a furtherdevelopment of the sharp area (paracone and metacone) whencompared with the talon, which is a common characteristic of theCuon genus (Garcıa, 2003:156). The paracone is larger than themetacone and both are surrounded by a cingulum which is well-developed in the labial and mesial edges. The paracone is moreworn than the metacone. The protocone is poorly defined. There isa highly worn central-lingual depression among the cusps, distallylimited by an enamel ridge facing the labial cusp of the metacone(Table 6).

Table 2Measurements in mm of the cranium of Cuon from Parpallo and Santa Maira as compared to other crania of Cuon from the sites of Obarreta and Gabasa, and present-day Canislupus. L: length; B: breadth; H: height.

Cranium Cuon Canis lupus

Parpallo Santa Maira Gabasa g Obarreta N Mean min Max S

(1) Total L 197 176 14 241.73 228 258 9.05(2) Condylobasal L 188 189.7 179 14 224.36 208 243.62 9.14(3) Basal L 179 183 159.5 14 213.24 196.15 228.82 8.60(7) Upper neurocranium L 98.63 91 14 114.80 101.24 126.26 6.70(8) Viscerocranium L 100.49 84.5 13 121.26 111.74 128.9 5.82(9) Facial L 109.42 98.3 14 137.36 123.09 150.05 7.14(15) L of cheektooth row 63.66 64.7 57.5 14 80.69 72.42 85.56 3.50(16) L of the molar row 19.26 15 14 20.87 18.7 23.42 1.09(17) L of the premolar row 50.64 47.6 14 62.75 57.72 67.77 3.09(19) L of P4 20 18.64 14 23.22 20.7 25.02 1.36(23) Greatest mastoid B 75.6 76.19 64.5 14 79.72 73.2 86.33 3.61(25) Greatest B of the occipital condyles 39.48 40.16 36.5 14 43.87 38.88 48.62 2.63(27) Greatest B of the foramen magnum 18.96 18.93 19.9 14 22.10 18.99 25.03 1.65(28) H of the foramen magnum 15.6 12.71 12.8 14 18.81 17.05 20.99 1.22(29) Greatest neurocranium B 75.3 74.27 63 14 69.99 64.5 74.02 2.96(30) Zygomatic B 126 120.1 13 135.43 119.63 148.75 8.58(31) Least B of skull 41.21 35.31 14 42.11 40.34 45.22 1.37(32) Frontal B 52.56 53 14 63.75 54.8 74.74 5.34(33) Least B between the orbits 44.9 37.5 14 44.68 40.38 51.33 3.29(34) Greatest palatal B 68.91 64 14 75.69 69.4 82.66 3.11(35) Least palatal B 46.93 34.5 14 41.58 36.4 47.54 2.76(37) Greatest inner H of the orbit 32.45 31.09 14 36.11 32.14 38.88 1.73(38) Skull H 66.16 62.8 56 14 73.04 65 80.91 4.04(39) Skull H without sagittal crest 61.22 57.34 54.6 14 58.84 53.66 66.85 2.93(40) H of the occipital triangle 55.25 45.74 49.8 14 57.13 54.35 60.87 2.16B of auditory bullas 70.28 66.66 62.72 13 63.37 59.8 69.1 2.62Least B of auditory bullas 21.6 19.87 17.68 14 18.36 14.92 20.97 1.75

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450442

The two M2 are missing, but the left alveolus shows that the leftM2 had its roots fused.

3.1.4. C1 from Coves de Santa MairaThe distal and mesial crown tubercles indicate that we are

dealing with an upper left canine (Hillson, 1986). It has neitherwear surfaces nor fractures. The ridges on the posterior edge of thecrown and on the lateral side are well-developed and sharp. Thecrown is lower than that of the C. lupus canines. Its morphology issimilar to the upper canine of C. alpinus from Cova del Parpallo, andshows clear differences when compared to the upper canines of C.lupus (Tables 3 and 4; Fig. 6). However, we have classified it as cf. C.since this is an isolated tooth recovered in a recent context. Futurefindings will confirm or reject the presence of the genus in Holo-cene chronologies.

Table 3Measurements of the left C1 of Cuon from Parpallo and Santa Maira as compared topresent-day Canis lupus. L: length.

Left C1 Cuon Canis lupus

Parpallo SantaMaira

N Mean Min Max S

Total L 42.42 38.72 2 52.84 51.38 54.3 2.064752Total L of the

crown13.31 14.73 2 17.58 16.7 18.4 1.2

Total L of theroot

29.11 23.99 2 35.275 34.65 35.9 0.883883

Antero-posteriorL of the root

13.65 11.4 3 14.40333 13.4 15.21 0.920887

Transversal L ofthe root

8.27 6.83 3 9.356667 8.54 10.33 0.905226

Antero-posteriorL at the base ofthe crown

11.94 10.97 12 13.42417 12.65 14.06 0.501352

Transversal L atthe base of thecrown

7.72 6.69 13 8.413846 7.5 9.06 0.448303

The length and the antero-posterior diameter at the base of thecrown of the upper canines from Parpallo and Santa Maira aresmaller when compared to those of the C. lupus. On the contrary,the transversal diameter at the base of the crown is sometimeswider in Cuon. Therefore, the canines of Cuon are more circular insection. The length and width of the P4 and M1 from Parpallo arevery similar to other Cuon teeth from other sites in the IberianPeninsula (La Riera, Gabasa, Boquete de Zafarraya and Bolinkoba),and always smaller when compared to those of the C. lupus, excepta few teeth of female wolves which show slightly smallermeasurements than the ones of Cuon (Figs. 7–9).

3.1.5. Mandible from Cova del ParpalloThis mandible is shorter than that of the C. lupus, which is

a characteristic feature of the Cuon (Altuna, 1983; Garcıa,2003:149). The foramina are displaced towards the distal area, andthe diastema are missing (Table 7, Fig. 10).

Dentition:P2 has two roots and one slightly worn cusp (protoconid).M1 has two well-developed roots. The paracone is horizontally

arranged and very worn. The high protoconid shows an important

Table 4Description of the morphological characters of the C1 of Cuon as compared topresent-day Canis lupus. H: height.

C1 Cuon Canis lupus

Basal line of the enamel (a) Right Curved, with a notchon the oral side

H of the crown (b) Lower HigherLine between the distal apex

of the root and the tip ofthe crown (b)

Almost straight Curved crown isaborally projected

Shape of the root (c) Nearly globular More slender in shapeDistal extreme of the root Distinct in shape and

aborally projectedAlmost indistinct

Fig. 6. C1 of Canis familiaris (up left, lingual view; down left, anterior view), Canis lupus (center right, lingual view) and Cuon from Santa Maira (center left, anterior view; up right,lingual view) and Parpallo (down right, lingual view). Letters refer to Table 4.

Table 5Measurements of the P4 of Cuon alpinus from Parpallo as compared to other archaeological remains of Cuon alpinus and present-day Canis lupus. L: length, B: breadth.

P4 Cuon Canis lupus

Parpallo (left) La Riera Gabasa e Gabasa g B. Zafarraya B. Zafarraya Amalda VII N Mean Min Max S

(18) L 21.53 21.5 21.6 22.7 20 19.4 20.5 14 24.798571 22.54 26.1 1.091526(18a) B 9.65 11 – – 10.4 10.3 10.3 14 12.930714 11.96 14.25 0.657881

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 443

wear facet that reaches the paraconid. The metaconid is very smallin the lingual side, with only one worn cusp (hipoconid) in a centralposition. The entoconid is missing, which is a usual feature of theCuon genus (Altuna, 1981, 1983; Barroso et al., 2006; Blasco, 1995;Castanos, 1987; Cervera et al., 1999; Garcıa, 2003:148).

3.1.6. Left hemimandible fragment from Cova NegraIt exhibits similar characteristics to that from Cova del Parpallo

(Table 7, Fig. 10). This fragment of mandible preserves all thepremolar and molar series. All the teeth, excepting P1, have tworoots.

P1 has one cusp (protoconid) slightly worn.P2 is characterized by a high protoconid with a small cusp on the

labio-distal side. This tooth has no anterior cusp.P3 exhibits a main central cusp in a central position, and a small

anterior cusp on the mesial edge of the protoconid. There isa smaller cusp towards the central cusp, and a lower cingular cuspon the distal edge.

P4 has a large and only slightly worn protoconid. It has a smallanterior cusp and two posterior ones, the mesial one being larger thanthe distal. These posterior cusps are also present in the individualsfrom Trinchera–Galerıa (Cervera et al., 1999; Garcıa, 2003:148).

The carnassial, M1, has a very horizontal paraconid and a highand well-developed protoconid. These two cusps are slightly worn.The metaconid is small, flat and not too well-defined (in contrast toC. lupus) on the lingual side. A small depression appears between

Table 6Measurements of the M1 of Cuon alpinus from Parpallo as compared to others from differ

M1 Cuon alpinus

Parpallo (left) La Riera Bolinkoba Gabasa Amalda VII B. Zafarraya

(20) L 13.45 12.6 13.6 12.7 12.6 12.6 10(20) B 15.82 16.2 14.3 15.7 13.8 12.9

the posterior edge of the metaconid and the distal ridge of theparaconid. The talonid has only one cusp (hipoconid) and is worn inthe central area. On the lingual area the entoconid is represented bya cingulum, not by a real cusp, between the lingual edge and thedistal ridge of the hipoconid. These features have been described ina similar way for Obarreta and Trinchera–Galerıa individuals(Altuna, 1983; Garcıa, 2003:155) (Fig. 10b).

M2 is a subrectangular tooth, with a slightly worn main cusp(protoconid) in the central area of the trigonid. In the lingual area,the metaconid is not well-developed and is aligned with theprotoconid (in the Canis it is on a more oblique position). These twocusps are joined by a ridge. On the distal area, the tooth has a smalland severely worn cusp (hipoconid).

The length of the mandible from Parpallo is a lot lower than thatof the C. lupus, even in the females, but higher than that one fromObarreta. The measurement of the alveolar series coincides withwhat has already been exposed above. The differences, however,are not that clear concerning the height and thickness of themandibular body with regard to the dimensions of the teeth. Theanterior part of the mandibular body of Cuon is higher (between P2

and P3), whereas the mandibles of C. lupus are higher in theposterior part (P4–M1 and M1). The mandibles of C. lupus are mostlywider than those of Cuon. If we compare the average alveolar P1–M2

length between the C. alpinus and the C. lupus by taking themeasurements that appear in Table 7, we find that the length is 18%shorter in the Cuon than in the Canis, whereas this ratio is reduced

ent sites of the Iberian Peninsula and present-day Canis lupus. L: length, B: breadth.

Canis lupus

TG-AT1 TG-AT2 N Mean Min Max S

13.2 14.5 13.9 23 16.5369565 14.91 19.57 1.0962723414.7 14.1 23 18.2543478 15 20.95 1.46013333

M1

9

10

11

12

13

14

15

16

17

18

19

12 14 16 18 20 22 24

B

L

Parpalló

La Riera

Bolinkoba

Zafarraya

Trinchera Galería

Canis lupus

(male)

Canis lupus

(female)

Canis lupus

(unknown)

C. lupus Letzetxiki

C. lupus Marizulo

Fig. 9. Scatterplot of the breadth (B) and length (L) of the M1 of Cuon from differentPleistocene sites (Parpallo, La Riera, Bolinkoba, Zafarraya and Trinchera–Galerıa)compared with present-day Canis lupus (male, female and unknown sex) and UpperPleistocene wolves from Lezetxiki and Marizulo sites.

C1

base of the crown

6

6,5

7

7,5

8

8,5

9

9,5

10 11 12 13 14 15A-P lenght

T len

gh

t

ParpallóSanta MairaC. lupus (male)

C. lupus (female)

C. lupus (unknown)

Fig. 7. Scatterplot of the antero-posterior (A-P) and transversal (T) lengths of the C1 ofCuon from Cova del Parpallo and Santa Maira compared with present-day Canis lupus(male, female and unknown sex).

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450444

to 10% when calculated by taking the mandibular thicknessbetween P4-M1 and to 4% when calculated taking the heightbetween P2-P3. This comparison shows that Cuon mandibles areshorter than Canis mandibles but rather sturdier (Table 8).

The measurements of M1 (Fig. 11) and M2 are clearly larger inCanis, whereas the dimensions of P2, P3 and P4 are closer. P1 isbetter developed in Cuon and is considerably larger, probably due tothe fact that this tooth is less functional in Canis. The measurementsof the lower canines are larger, mainly the antero-posterior diam-eter, as it has been observed for the upper canines.

3.1.7. Post-cranial skeleton from Cova del ParpalloWhen compared to the Canis ones, the Cuon post-cranial

remains show – except for a smaller bone size (length) which is,however, difficult to observe in fragmented archaeological material– little anatomical differences (Cervera et al., 1999; Garcıa,2003:161), being in occasions a matter of details difficult to assessdue to their own intraspecific variability. Differences observed inremains of wolves and gathered in Table 9 should be consideredmore as comparative descriptions to be confirmed by futureresearch than as clear discriminating criteria (Figs. 12–14).

The shortage of post-cranial remains and, therefore, of post-cranial measurements deprives us from making assessments aboutthe body dimensions of Iberian dholes. The Parpallo measurementsare higher than those of Obarreta and Cova Negra (which are closerto each other) but they are mostly inferior to those of the subactualspecimens of C. lupus, except for the minimum measurementsprovided by several female wolves (Table 10).

P4

18

20

22

24

26

28

30

9 11 13 15B

L

ParpallóLa RieraZafarrayaAmaldaC. lupus (male)

C. lupus (female)

C. lupus (unknown)

C. lupus Lezetxiki

C. lupus Bora Gran

Fig. 8. Scatterplot of the breadth (B) and length (L) of the P4 of Cuon from differentPleistocene sites (Parpallo, La Riera, Zafarraya and Amalda) compared with present-dayCanis lupus (male, female and unknown sex) and Upper Pleistocene wolves fromLezetxiki and Bora Gran sites.

3.2. Diachronic distribution of the genus Cuon in the IberianPeninsula during Upper Pleistocene

C. alpinus has been identified in the Iberian Peninsula from theMiddle Pleistocene at the Trinchera–Galerıa site in Atapuerca(Cervera et al., 1999; Garcıa, 2003:146; Garcıa and Arsuaga, 1998),although the greater part of the references belongs to the UpperPleistocene. The Cuon sp. humerus from the site of Tossal de la Font(Gusi et al., 1980) belongs to the beginning of this phase, althoughit has never been described nor have any measurements beenprovided. In Mousterian contexts it is present from North to Southof the mainland in Gabasa (Blasco, 1995), Cova del Gegant (Vinasand Villalta, 1975), Amalda (Altuna, 1990), Los Casares (Altuna,1973) and Boquete de Zafarraya (Barroso et al., 2006). During theUpper Paleolithic, the species has been documented from North toSouth of the Peninsula in Amalda (Altuna, 1990), Rascano (Altuna,1981), La Riera (Altuna, 1986) and Bolinkoba (Castanos, 1988). Themandible from Cova Negra presented here was recovered ina Mousterian context, in the slope of the entrance corridor to thecave within strata that belong to the last stages of the Wurm I (MIS5d-b). The Cuon remains from this same site belonging to laterexcavations carried out by V. Villaverde are placed in the Wurm III(MIS 3) (Martınez, 1996). The remains recovered in Cova de Par-pallo are more problematic since they are not related to a culturalcontext as are those from Obarreta (Altuna, 1983) or Durangue-sado (Castanos, 1988). As to the C. alpinus mandible remain fromthe Gruta do Escoural, even when belonging to an archeologicalsite with a Mousterian sequence, the remain was found in a gallery(I) of unclear chronology (prior to the Holocene). The remainsfrom Parpallo were located in a crevice on the lower part of thewestern wall, placed in a sandy stratum or a stratum of karsticdissolution, associated with some remains of Capra pyrenaica withno archaeological material and with no evidence of anthropicprocessing. We could consider these remains to be prior to thehuman occupation of the cavity, which would place them inchronologies prior to the Gravettian. As to the cranium fromentrance E in Santa Maira, even though the context in which itappears is still under study, it surely belongs to the end of theUpper Paleolithic. These remains confirm, together with thosefrom the Boquete de Zafarraya and Cova del Gegant, and for theMediterranean context, a situation similar to the one observed in

Table 7Description of the morphological characters of the mandibles of Cuon as compared to present-day Canis lupus.

Mandible Cuon Canis lupus

Diastema C1-P1 (a) Reduced LargerDiastema P2-P3 (b) Very small Larger in 75% of the studied casesMiddle mental foramen (c) Under P2 (Parpallo)

Under P2 anterior (Cova Negra)Under P1–P2 diasteme

Posterior mental foramen (d) Under P3 posterior (Parpallo)Under P3 (Cova Negra)

Under P3 anterior

M3 (e) Absent PresentAngle between coronoid process and

the body of the mandible (f)Smaller Larger

Aboral border of the ramus (g) Facing the front Facing the rearAnterior border of the coronoid process (h) Well-developed and rounded ThinBasal region of the masseteric fossa (i) Closed and well-delimited Open to the mandibular bodyAngular process (j) The process continues in a crest

towards the masseteric fossaThe crest is absent or less marked in a oblique position

Relation between angular process andmandibular condyle (k)

The process is larger in the aboral directioncompared to the mandibular condyle

The process is shorter when compared to the condyle

Muscular insertions of the angularprocess (l)

The inferior pterygoid fossa is better-developed The upper pterygoid fossa is better-develped

Oclussion of C1 With I3 With C1–P1 diasteme

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 445

the Bay of Biscay. Nevertheless, the canine from the W entrance inSanta Maira, located at an epipaleolithical level, belongs to thelatest European context in which remains of this species appear.This would mean that it lasted until at least the first stages of theHolocene, beyond the reference of Rascano (Magdalenian) in the

Fig. 10. Mandibles of Cuon from Cova Negra, Duranguesado, Obarreta and Parpallo, and Canocclusal view (detail).

context of Cantabric region (Altuna, 1981). Nevertheless, we areaware of the fact that this is the only remain and it is possible that,since it is a canine, it was perhaps collected due to its possibleornamental use (although is not perforated) and might come froma different and unknown context.

is lupus (down). Letters refer to Table 7. Up right: M1 from the mandible of Cova Negra,

Table 8Measurements of the mandibles of Cuon from Parpallo and Cova Negra (CN) as compared to other mandibles of Cuon from the sites of Los Casares (LC), Obarreta (Obar),Duranguesado (Dur), Amalda (Ama), Escoural (Esc) and Trinchera–Galerıa (TG), and present-day Canis lupus. L: length; B: breadth; H: height.

Mandible Cuon Canis lupus

Par(left)

Par(right)

CN LC Obarright

Dur AmaVII

Esc TG1 TG2 TG3 N Mean Min Max S

(1) Total L 153 130 13 176.350769 160.81 196.22 8.77636263(2) L ang. proc.-Infradent. 157 132 13 174.51 162.1 185.95 6.5822324(3) L indent. Condyle/ang proc-Infradent. 147.5 124.5 13 167.342308 155.4 184.55 7.34864858(4) L cond. proc.-aboral border C1 alveolus 134.61 114.5 121.8 13 154.428462 140.84 171.86 8.00867847(5) L indent. Condyle/ang. proc.-aboral

border C1 alveolus129.75 110 118 13 145.474615 135.6 160.24 6.57069455

(7) L aboral border alveolus M3-aboralborder C1

75.44 76.15 68.5 69.97 13 93.2276923 87.6 102.04 3.72928088

(8) L P1-M2 alveolar 72.5 71.86 77.1 67 67.61 72.8 73.9 13 87.1661538 81.52 94.33 3.3145702(9) L P2-M2 alveolar 66.29 64.66 69.6 63.02 13 81.0976923 75.5 87.51 3.1783097(10) L M1-M2 alveolar 29.74 30.54 34.1 32.5 31.51 31 32.4 13 38.2553846 35.44 40 1.17911418(11) L P1-P4 alveolar 42.41 41.73 44 37.5 42.3 43.1 13 49.7092308 46.26 56.44 2.68200628(12) L P2-P4 alveolar 36.17 34.22 37.45 33.7 13 43.5138462 38.63 50.15 2.63637864(13) L of M1 22.21 24.8 23 22.89 22.7 22.2 23.6 24.2 14 28.005 26.05 29.62 1.09490252(13a) B of M1 9.81 9.7 8.6 9.17 9 8.7 8.9 9.6 14 11.1714286 10.51 12.05 0.47991758(14) L of M1 alveolar 22.13 22.38 23.65 22.53 13 27.1807692 24.6 29.21 1.4592947(15) L of M2 9 7 8.2 8.2 8.4 13 11.1676923 10.05 11.89 0.49721145(15a) B of M2 7.1 6.2 6.6 13 8.33384615 7.23 8.93 0.43521141(18) H vertical ramus 67.99 59.5 63.86 13 74.7338462 65.9 82.4 5.05338094(19) H of the mandible behind M1 27.78 28.88 28.12 24.8 24 27.87 13 30.6823077 27.48 34.16 2.12726019(20) H of the mandible between P2y P3 24.4 24.32 23.92 19.9 23.17 12 24.0491667 22.68 26.22 0.96678905H of the mandible between P4 and M1 26.88 25.94 26.43 21.5 24.38 13 28.7338462 26.46 32 1.66486005B of the mandible between P4 and M1 11.96 11.96 12.92 11.7 12.2 13 13.5107692 12.19 15.33 0.89096821L of P1 6.5 6.15 11 5.71909091 4.64 6.7 0.58764708B of P1 4 4.03 11 4.36 4.1 4.91 0.25187298L of P2 8.73 10 8.7 10.05 10.4 10 14 11.6828571 10.46 12.55 0.65493897B of P2 4.82 5.2 4.2 5.31 5 5.3 14 5.85642857 5.3 6.91 0.41181787L of P3 12 11.2 11.9 12 13.48 12.39 14.4 0.63031017B of P3 5.7 5.6 5.7 12 6.59 6 6.98 0.34295507L of P4 14.6 14.94 13 13.6 14.8 15.2 14 15.3485714 14.4 16.32 0.53784266B of P4 7 7.26 6.5 7.3 6.7 7.5 14 7.59785714 6.88 8.41 0.37613783L of the antero-posterior base of the

crown of C1

11.94 12.22 12 13.225 12.25 13.8 0.50618897

L of the transversal base of thecrown of C1

8.18 8.78 12 9.76916667 8.69 11.02 0.67559345

L condyle of the mandible 23.03 21.75 13 31.5884615 27.91 37.5 2.36293901

M1

20

22

24

26

28

30

32

8 9 10 11 12 13

B

L

Parpalló

Cova Negra

Obarreta

Duranguesado

Trinchera Galería

Amalda VII

C. lupus (male)

C. lupus (female)

C. lupus (unknown)

C. lupus Urtiaga

C. lupus Ermitia

C. lupus Vizcaya

C. lupus En Parco

Fig. 11. Scatterplot of the breadth (B) and length (L) of the M1 of Cuon from differentPleistocene sites (Parpallo, Cova Negra, Obarreta, Duranguesado Trinchera–Galerıa andAmalda) compared with present-day Canis lupus (male, female and unknown sex),with Upper Pleistocene wolves from Urtiaga, Ermitia and several sites of Vizcaya (themean and a 5% error is indicated), and with the Initial Holocene site of En Parco.

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450446

3.3. Appearance contexts: collections of cultural and natural origin

Given the data put forward by this paper it can be asserted thatthe remains of the genus Cuon provide several forms of contribu-tion. In palaeontological contexts, the remains are of natural originand could have been caused by predation or the natural death of ananimal in a chasm or cavity. The latter would be the case with theCuon from Obarreta. Its remains do not show any kind of alterationrelated to the consumption or transport by other predators: themajority of the skeleton is available and the bones are mainlycomplete (Altuna, 1983).

We should also consider archaeological contexts where twokinds of accumulation can be distinguished. On the one hand, thoseof natural origin, with similar characteristics to those described forpalaeontological contexts. One of these cases would be the Cuonfrom Parpallo which shows no evidence of human activity or fromother predators and which, leaving out those fractures causedduring its recovery, is mostly complete. Furthermore, theirdiscovery took place in the last layer of the sequence of one of thecavity galleries, probably in a context prior to the human occupa-tion of the cave. Both this animal and the one recovered in Obarretaare adults. The case of the one found in Boquete de Zafarraya ismore difficult because, even if it is true that there seem to becontributions from other carnivores (or perhaps other dholes), thepossibility of these remains being placed in the cavity by a humanbeing should not be dismissed. They mostly appear at the end of thecave interspersed between archaeological levels of higher density.

Table 9Description of the morphological characters of the scapula, the radius and the femur of Cuon alpinus compared to present-day Canis lupus.

Post-cranial skeleton Cuon alpinus Canis lupus

Parpallo _/\

Scapula Infraglenoid tubercle (a) Slightly developed Well-developedAcromion (b) Well-developed. The suprahamatus

process is larger.Less developed. The suprahamtusprocess is smaller and distally situated

Spina (c) The transition between spina andacromion is curved

Spina and acromion form an almost right angle

Dorsal edge of the glenoidcavity (d)

The notch is absent With a marked notch

Articular cavity (e) Subcircular SubrectangularCoracoid process (f) Well-developed Less developed

Radius Lateral side of the distalepiphisis (a)

Flat, slightly pointing laterally,with a small notch between it andthe lateral crest

Curved laterally, and continues in the lateral crest.

Styloid process (b) Shorter, less angular and blunter Longer and pointedDorsal crest (c) Well-developed Slightly developed or rounded

Femur Intertrocanteric crest (a) More oblique and straight More vertical and curvedCaput femorae (b) Rounded The lateral side of the caput is obliquely prolonged

towards the trochanteric fossaUpper border of the neck (c) Long and flat Shorter and concave

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 447

It is also the site where more remains and individuals have beenrecovered (NMI¼ 19) in the Iberian Peninsula, a fact that could berelated to the characteristics of the cavity environment and itsposition with regard to the sea level (1100 msnm) (Barroso et al.,2006). A different type of contribution in an archaeological contextis the anthropic or cultural context. This could be the case for manyof the remains recovered in peninsular archaeological sites,although it is not often possible to clearly establish this origin.Unlike the contributions described before, these are isolatedelements or either fragments of them. The mandible from CovaNegra and the neurocranium from Santa Maira are the only remainswith anthropic marks related to the butchering process. In thelateral part of the mandible, we observe a number of lithic marksbelonging to short and oblique scrapes which, considering theirposition, can be related to the removal of the skin according to theobservations carried out in the treatment of the skin of Vulpesvulpes in subactual trappers (Perez and Morales, 2008). In the

Fig. 12. Scapulae of Cuon alpinus from Obarreta and P

cranium we find a long, thin and oblique incision over the leftparietal. Consequently both in the case of the mandible from CovaNegra and the cranium from Santa Maira the ultimate aim was theobtainment of the skin (Fig. 15).

4. Assessment

The morphological and metrical study of the remains of Canidaeput forward by this study and their comparison with the remains ofthe Canis and Cuon genera have allowed us to identify the former asbelonging to the species C. alpinus (Parpallo and the cranium fromSanta Maira), Cuon cf. alpinus (Cova Negra) and cf. Cuon (C1 fromSanta Maira). Thus, the presence of this genus in the Mediterraneancontext of the Iberian Peninsula during the Upper Pleistocene inMousterian contexts and the Upper Paleolithic is confirmed. Theappearance of the canine tooth recovered in the W entrance of theSanta Maira site could represent the identification of this genus in

arpallo, and Canis lupus. Letters refer to Table 9.

Fig. 13. Radius of Cuon alpinus from Obarreta and Parpallo, and Canis lupus. Letters refer to Table 9.

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450448

an archaeological context of the early Holocene, which could meanthat C. alpinus lasted longer in this area.

The Cuon appears both in contexts of natural and archaeologicaldeposits. The former are related to natural deaths and in these casesthe skeleton appears almost complete, as in Obarreta, or there arewhole bones with no alteration marks, as happens in Parpallo,which even being an archaeological site includes remains which are

Fig. 14. Femorae of Cuon alpinus from Obarreta and P

prior to the human occupation. The latter are related to the humanactivity according to the cultural contexts in which they appear andthe lithic marks they display, as would be the case with themandible from Cova Negra and the cranium from Santa Maira.

The comparison with C. lupus has highlighted the difficulties andlimitations met with when trying to distinguish between the Cuonand Canis genera, especially when it concerns post-cranial remains,

arpallo, and Canis lupus. Letters refer to Table 9.

Table 10Measurement of different post-cranial bones of Cuon alpinus from Parpallo as compared to other bones of Cuon from the sites of Cova Negra (CN), Obarreta, Boquete deZafarraya (BZ) and Trinchera–Galerıa (TG), and present-day Canis lupus.

Post-cranial skeleton Cuon alpinus Canis lupus

Parpallo Obarreta CN BZ TG N Mean Min Max S

Scapula GLP 35.07 30.5 29 10 40.36 36.96 44.98 2.474SLC 28.17 23.3 24.6 10 33.22 29.45 38.51 2.442BG 22.22 19.9 18.2 10 24.36 21.75 28.4 2.049LG 29.30 25.5 24.7 10 32.99 30.17 36.66 1.901

Radius Bp t 22.50 18.2 18 9 24.09 22.64 26.82 1.36Bp ap 14.83 12.7 9 16.52 15.65 18.08 0.884Bd t 29.35 25.5 24 9 31.69 28.17 34.05 1.872Bd ap 17.10 14.21 9 17.53 15.94 19.14 1.025GL 175.4 152 151 9 203.6 196.4 222.4 7.767SD 15.17 12.5 12 9 16.61 14.65 18.93 1.392

Femur Bp t 47.99 39 (44) 9 50.41 47.25 57.02 2.972DC 23.88 20 9 23.88 22.76 26.44 1.071Bd t 40.81 33.5 9 41.14 36.54 44.84 2.456GL 219.5 184 9 225 213.5 246.2 9.493

M. Perez Ripoll et al. / Journal of Archaeological Science 37 (2010) 437–450 449

fragmented in many cases, as is pointed out by some earlier studies(Blasco, 1995). Many of the identified remains of Cuon are dentaland cranial remains which allow much more obvious diagnoses. Onthe other hand, the post-cranial skeleton is normally less repre-sented and mostly identified in contexts of natural contribution,like Obarreta or Parpallo (whole or half skeletons), or either whenthere is an accumulation of several individuals, as is the case withBoquete de Zafarraya (Barroso et al., 2006). This leads us to raisea thought: during the Upper Pleistocene the references from thegenus Canis or from the Canidae family occur more often and thoseof the genus Cuon are a lot less frequent, is it thus possible that

Fig. 15. Lithic marks on the mandible from Cova Negra (up) and on the cranium fromSanta Maira (down).

a large part of the Cuon remains might have been identified asCanidae or even as Canis sp.?

Acknowledgements

We would like to thank J. Altuna and P. Ma. Castanos for theinformation provided on Cuon in Euskadi and E. Ma. Barriocanal(Museo Arqueologico, Etnografico e Historico Vasco) for her kind-ness and help during our visit to Bilbao, R. Soriguer (Estacionbiologica de Donana) and J. Barreiro (Museo Nacional de CienciasNaturales) for the specimens of wolves they lent for the compara-tive study and, also, B. Martınez (ICREA-Universitat Rovira i Virgili),A. Arribas (IGME), C. Dıez (Universidad de Burgos) and J.Ph. Brugal(MMSH-CNRS) for their useful comments. We are grateful to N.Gomez Handford for her indispensable help in translating thispaper, and J. Vives for the revision of the text. We thank twoanonymous reviewers and the editor for their helpful comments onearlier drafts of the manuscript.

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