HAL Id: hal-02367214
https://hal.archives-ouvertes.fr/hal-02367214
Submitted on 9 Jan 2021
HAL is a multi-disciplinary open access archive for the deposit and dissemination of sci- entific research documents, whether they are pub- lished or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers.
L’archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d’enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.
The Upper Pleistocene brown bear (Carnivora, Ursidae) in the Zagros: Evidence from Wezmeh Cave,
Kermanshah, Iran
Hervé Monchot, Marjan Mashkour, Fereidoun Biglari, Kamyar Abdi
To cite this version:
Hervé Monchot, Marjan Mashkour, Fereidoun Biglari, Kamyar Abdi. The Upper Pleistocene brown
bear (Carnivora, Ursidae) in the Zagros: Evidence from Wezmeh Cave, Kermanshah, Iran. Annales
de Paléontologie, Elsevier Masson, 2019, 106 (2), pp.102381. �10.1016/j.annpal.2019.102381�. �hal-
02367214�
The Upper Pleistocene brown bear (Carnivora, Ursidae) in the Zagros:
Evidence from Wezmeh Cave, Kermanshah, Iran
Les Ours brun (Carnivora, Ursidae) du Pléistocène supérieur dans le Zagros : l’exemple de la grotte Wezmeh, Kermanshah, Iran
Hervé Monchot
a,, Marjan Mashkour
b, Fereidoun Biglari
c, Kamyar Abdi
daUMR8167«OrientetMéditerranée »,ParisSorbonneuniversités, 27,ruePaul–Bert, 94204Ivry-sur-Seine cedex,France
bCP55,archéozoologie,archéobotanique (UMR7209), CNRS,MNHN, UPMC,ParisSorbonneuniversités, 55,rueBuffon,75005Paris,France cDepartmentofthepaleolithic, National MuseumofIran,30,TirstreetImamavenue,Tehran, Iran
dDepartment ofarchaeology andhistory, university ofShiraz,Shiraz,Iran
a r t i c l e i n f o
Keywords:
Brown bear Ursusarctos Skeletal representation Ageing
Biometry ZagrosMountain
a bs t r a c t
While bears (Ursidae) arewell represented inWestern Europe andtheCaucasus during thePleistocene, bearremains from this period arerare inSouthwest Asia. Only alimited number ofsites, bothnatural andarchaeological, haveyielded evidence ofbrown bear(UrsusarctosLinnaeus, 1758). Skeletal remains forthis species are often represented byalimited number ofelements. The discovery of192 remains identified asbrown bearinWezmeh Cave(Kermanshah Province, Iran) isexceptional. Thispaperpresents adetailed description oftheWezmeh osteological assemblage, which confirms thatU. arctoswas already present inZagros during theUpper Pleistocene.
Motsclés: Oursbrun Ursusarctos
Représentation squelettique Profil demortalité Biométrie
Montagnes duZagros
r é s u mé
Silesours (Ursidae) sont bien représentés enEurope occidentale, ycompris dansleCaucase, pendant le Pléistocène, ils sont plus rares enAsie du Sud-Ouest. Seul un nombre limité desites naturels ou archéologiques ontlivré des ossements d’ours brun (Ursus arctos Linnaeus, 1758) etgénéralement les restes squelettiques decette espèce sont souvent représentés parpeudespécimens. Ladécouverte de 192restes identifiés comme appartenant àdesoursbrunsdanslagrottedeWezmeh (province deKerman- shah,Iran) constitue uneexception. Cetarticle apourbutdedécrire endétail cetassemblage ostéologique quiconfirme laprésence déjàd’UrsusarctosauPléistocène supérieur dans le Zagros.
1. Introduction
The brown bear (Ursus arctos Linnaeus, 1758) is aterrestrial mammal species with avery broad geographic distribution across Europe, AsiaandNorth America. TheSyrian brown bearsubspecies (Ursus arctos syriacus Hemprich and Ehrenberg, 1928), which is
Corresponding author.
E-mail addresses: herve.monchot@wanadoo.fr (H.Monchot), mashkour@mnhn.fr (M.Mashkour), fbiglari@gmail.com (F.Biglari), Kamyar.Abdi@gmail.com (K.Abdi).
smaller than the nominate U.arctosarctos, isnative toSouthwest AsiaandisfoundinTurkmenistan, Iran, IraqandTurkey. Itisextinct in Palestine and more recently, in Syria, although a few Syrian brown bearsstill exist alongtheborder between Lebanon andSyria (e.g. Hatt, 1959; Heptner andNaumov, 1998; Masseti, 2009). The Iranian brown bear isfound in the present day in the north and west of Iran, primarily in the Zagros Mountains from the Azer- baijan border tothe Shiraz region inFars, and also inthe Alborz Mountains from Astara toeastern Golestan. Generally found inthe mountainous areas throughout itshome range, theSyrian brown bears appear toden and hibernate in caves and tree hollows in high elevation forests (e.g. Lay, 1967; Etemad, 1985; Ziaie, 2008;
Farhadinia andValizadegan, 2015).
https://doi.org/10.1016/j.annpal.2019.102381
The majority of the Pleistocene sites in Southwest Asia that contain ursid bones are located along the Levantine coast from northern Palestine totheLebanese mountains (Table 1;Fig.1).The earliest evidence ofbear was found inthe Lower Pleistocene site ofUbeidiya intheJordan valley where 13remains ofUrsusetruscus (Cuvier, 1823) were recovered (Martínez-Navarro etal.,2009).The onlysites during theMiddle Pleistocene with thepresence ofursid remains areCaveBearintheUpper Galilee (Tchernov andTsoukala, 1997)andDarband CaveintheAlborz (Biglari andShidrang, 2006;
Biglari et al., 2007). These comprise large assemblages of teeth and bones identified as Ursus deningeri (von Reichenau, 1904) (Argant unpublished report; interested readers mayrefer inpartic- ulartoArgant andPhilippe (2002) foranoverview ofthecomplex phylogeny of Plio-Pleistocene Eurasian Ursidae). Finally, during the Upper Pleistocene only U. arctos has been identified, from remains found inMount Carmel, innorthern Levant/Galilee, inthe Lebanon Mount (e.g.Kurtén, 1965; Hooijer, 1961; Garrard, 1980) and in Northwest Syria (Griggo, 2004). In the Zagros Mountains
Table 1
ListofPleistocene ursid remains fromSouthwestern Asia(notexhaustive).
Listedesrestesd’ours duPléistocène duProcheetduMoyen-Orient (liste nonexhaustive).
Site(Level) Country Species Age/Culture NISP(MNI) Reference
Antelias Lebanon Ursusarctos UpperPalaeolithic Notcounted Garrard, 1980
Amud (B2) Israel Ursussp. UpperPleistocene/Middle Palaeolithic 1 Rabinovich andHovers, 2004
AbriBezez(B) Lebanon Ursusarctos UpperPleistocene/Middle Palaeolithic 3 Garrard, 1983
AbriZumoffen (9–3) Lebanon Ursusarctos Middle Palaeolithic 3 Garrard, 1983
AbriZumoffen (21–11) Lebanon Ursusarctos Middle Palaeoilthic 1 Garrard, 1983
Azokhcave Nagorno-Karabakh Ursusspelaeus LatePleistocene 265(8) VanderMadeetal.,2016
Bear’scave Israel Ursusdeningeri Middle Pleistocene (Early Toringian) 63 Tchernov andTsoukala, 1997
Darband cave Iran Ursusdeningeri Middle Pleistocene 62 Biglari etal.,2007
Dederiyeh Syria Ursusarctos Middle Palaeolithic 1 Griggo,2004
El-Wad (B–C) Israel Ursusarctos UpperPalaeolithic-Epipalaeolithic Notcounted Garrard, 1980
Emirkaya (2) Turkey Ursusdeningeri Middle Pleistocene Notcounted Senetal.,1991
Hayonim Israel Ursusarctos UpperPaleolithic/Mousterian-Kebaran 2 Stiner, 2006
Hovk-1 Armenia Ursusspelaeus UpperPleistocene 186(12) Bar-Oz etal.,2012
Karain (E) Turkey Ursussp. UpperPleistocene/Middle Palaeolithic 73 Otteetal.,1998
Kebara Israel Ursusarctos UpperPleistocene/Middle Palaeolithic 2 Dayan, 1994
Kéoué Lebanon Ursusarctos UpperPleistocene/Middle Palaeolithic Notcounted Garrard, 1980
Ksar Akil Lebanon Ursusarctos UpperPleistocene/Middle Palaeolithic Not counted Hooijer, 1961
Maslouk Lebanon Ursusarctos UpperPleistocene/Middle Palaeolithic 1 Monchot andGauthier, unpublished
NahrIbrahim Lebanon Ursusarctos UpperPleistocene/Middle Palaeolithic 31 Monchot andGauthier, unpublished
NahrelJoz Lebanon Ursusarctos Middle Palaeolithic Notcounted Garrard, 1980
Ras-El-Keb Lebanon Ursusarctos Middle Palaeolithic 49(11) Garrard, 1998
Shanidar Iraq Ursusarctos UpperPleistocene/Middle Palaeolithic 5(3) Evins, 1982
Tabun Israel Ursusarctos Midldle/Early UpperPalaeolithic Notcounted Garrard, 1980
Ubeidiya Israel Ursusetruscus EarlyPleistocene 13 Martínez-Navarro etal.,2009
Wezmeh Iran Ursusarctos UpperPleistocene 192(8) Mashkour etal.,2009 (thisstudy)
Zuttiyeh Israel Ursusarctos Acheulo-Yabrudian complex (3/4) Bate,1927
Yarimburgaz Turkey Uarctos/deningeri Middle Pleistocene 3920 Stiner etal.,1998
NISP: number ofidentified specimens; MNI: minimum number ofindividuals/ NISP:nombrederestesdéterminés;MNI:nombreminimum d’individus.
Fig.1. Mapofthemain Pleistocene Ursid sitesofSouthwest Asia(notexhaustive).
Carte des principaux sitesàursidés du Pléistocène en Asie du Sud-Ouest (liste non exhaustive).
Fig.2. A.General viewofthenarrow valley attheQazivand Mountain wheretheWezmeh Caveislocated. Thesiteisindicated byanarrow. B.LocationofWezmeh caveand mapofthesitewith boneassemblage location (after Abdietal.,2002).
A.Vuegénérale del’étroite vallée delamontagne Qazivand aveclocalisation delagrotteWezmeh (indiquée paruneflèche). B.Localisation delagrotteWezmeh etcartedu siteavecl’emplacement del’assemblage osseux (d’aprèsAbdietal.,2002).
thebrown bearhasonly beenidentified inShanidar (Evins, 1982), although abear tooth was also collected onthesurface oftheMar Koulian cave, an Upper Pleistocene/Holocene site near Rawansar (Biglari andTaheri, 2000).Todate, noremains ofcave bear(Ursus spelaeus Rosenmüller, 1794) have been described in Southwest Asia. Although common inthe Caucasus (e.g. Baryshnikov, 1998;
Bar-Oz etal., 2012; Van der Made etal., 2016),it seems that this species never reached the Zagros-Taurus and Alborz mountain ranges.
Several recent studies have investigated the phylogeny ofthe Syrian brown bear (Murtskhvaladze et al., 2010; Calvignac etal., 2009).Agenetic study onmitochondrial DNA shows thatallbrown bears occurring in the Caucasus appear tobemonophyletic, and are a subspecies of the Eurasian brown bear (U. arctos arctos) (Murtskhvaladze etal., 2010; Lortkipanidze, 2010). However, the taxonomic status ofthespecies inthisregion remains unclear and itsdistribution andourunderstanding ofits evolution during the Pleistocene in this part of the world remains fragmentary. The
discovery of192 brown bearbones inWezmeh cave intheZagros Mountains istherefore animportant contribution tothehistory of thistaxon inSouthwest Asia.
2. Wezmeh Cave
This site was discovered in1999 during anarchaeological sur- veyunder thedirection ofKamyar Abdi intheareaofIslamabad-e Gharb (Province of Kermanshah, Iran), a region with numerous Palaeolithic sites (Smith, 1986). The cave is located about 10km southeast ofthecity ofIslamabad-e Gharb and3.5km northeast of thevillage ofTajar-e Akbar. Thecave isatanelevation of1,430m a.s.l., approximately 100mlower thanthesummit oftheQazivand Mountains, andabout 60mabove thevalley floor onasteep slope with a36 dip. The cavity is almost horizontal, formed between geological layers ofkarstic limestone. Itsentrance facesnorth andis 2mwide and1.20mhigh.Thecaveisabout27mlongandabout45 m2(Fig.2,Abdietal.,2002).Thecave waslooted during the1990s,
resulting inpiles ofdisturbed deposits infront ofthecaveentrance above thesteep slope. After acareful inspection ofthesite, geoar- chaeological studies indicate that the disturbed sediments came from therearofthecavewhere sedimentation reaches amaximum of3mdeep.These disturbed deposits ontheexterior slope yielded anabundant faunal assemblage inexcellent preservation, particu- larly rich incarnivoran andungulate remains. Aninitial faunal list was established, andnotably all ofthe carnivore families known tobepresent during theUpper Pleistocene inSouthwest Asia are present inthe Wezmeh assemblage: Hyaenidae (spotted hyena), Canidae (wolf andredfox), Felidae (lion, leopard, lynx/caracal and wild cat), Mustelidae (badger, stonemarten andpolecat), Herpesti- dae(mongoose) andofcourse Ursidae (brown bear). This lasttaxon represents 11.1% of the total NISP (carnivore and ungulate) and 13.9%ofthecarnivore remains found inWezmeh site(Table2).The first series ofpalaeontological andtaphonomical studies suggestan Upper Pleistocene origin forthisassemblage, with asimilar skele- talelement profile totheMiddle Palaeolithic assemblages from the southern Levantine (Mashkour etal., 2009).
Due to the disturbed context of this assemblage, and to the notable diversity of the mammalian species present, it was extremely important torun adirect dating analysis onthe prin- cipal species present in the cave. Several bones were selected from thecarnivore andherbivore remains, including bear, spotted hyena andwild boar. Aseries ofU/Th dates (CNRS, Gif-sur-Yvette) were performed on tooth enamel, providing ages of 63.1±12.6 and67.3 ±11.8 Kya forthe brown bear, 14.4±1.3, 23.4 ±4.9 and 26.6±9.4 Kya for the spotted hyena and finally 13.8±3.4 Kya for the wild boar (Mashkour et al., 2009: tab. 2). The oldest occupation ofthe cave may therefore bedated tothe Late Pleis- tocene. Three hyena coprolites were datedbytheAMS radiocarbon method, producing calibrated 14Cagesof19228 ±352calBPand 12744 ±370cal BP, aswell asaHolocene date of405±59cal BP (Djamali etal.,2011).Several fragmented human bones andteeth were discovered in the cave among the faunal remains, includ- ingtheNeanderthal premolar “Wezmeh 1”(Trinkaus etal., 2008;
Zanolli etal., 2019).One ofthe human remains was dated tothe early Neolithic, around 9300 cal BP (Broushaki et al., 2016). No
Table 2
Species distributions oflarge mammals inNISP (Number ofIdentified Specimens) andMNI(Minimum Number ofIndividuals) forWezmeh cave.
Liste desgrands mammifères enNISP (nombre despécimens identifiés) etMNI (nombre minimal d’individus) pourlagrottedeWezmeh.
NISP %NISP MNI
Carnivora
Crocutacrocutaspelaea Spotted Hyena 437 25.2 10
Ursusarctos Brown Bear 192 11.1 8
Pantheraleo Lion 16 0.9 2
Pantherapardus Leopard 2 0.1 1
Caracal/Lynx/Felis chaus Caracal/Lynx/Jungle cat 4 0.2 2
Felissilvestris Wildcat 8 0.5 2
Canislupus Wolf 176 10.1 6
Vulpesvulpes RedFox 492 28.4 19
Melesmeles Badger 42 2.4 7
Mustelaputorius Polecat 7 0.4 2
Martesmartes(foina?) Stonemarten 2 0.1 1
Herpestessp. Mongoose 5 0.3 3
Perissodactyla
Dicerorhinus Rhinoceros 2 0.1 1
Equuscaballus Horse 1 0.1 1
Hemionus/E. asinus Hemione/Donkey 4 0.2 1
Artiodactyla
Bosprimigenius Aurochs 13 0.8 2
Susscrofa Wild boar 97 5.6 4
Cervuselaphus Reddeer 59 3.4 3
Gazellasp. Gazelle 17 1.0 1
Ovisorientalis Mouflon 127 7.3 6
Capra aegagrus Wild goat 31 1.8 2
evidence of human activity (e.g. industry, burned bone, cutting traces) was observed.
3. The brown bear assemblage
The species identification of brown bear (Ursus arctos) ver- sus cave bear (Ursus spelaeus and Ursus deningeri) was based on theshape (morphology andsize) ofcertain skeletal elements (e.g.
Kurtén, 1955; Stiner, 1998; Stiner etal.,1998; Petronio etal.,2003;
Schweizer, 1999, 2005).Allthebones (N=192) present atWezmeh caveundoubtedly belong tothe brown bear(Figs.3–5).
3.1. Thebearboneconservation
The surfaces of the bones are in an excellent state ofpreser- vation; inmost cases they are smooth without cracks orfissures, characteristic of unweathered bones. Nevertheless, some pieces (n=9)present verylimited surface weathering with acracking par- allel tothe fibre structure (longitudinal) (Todisco and Monchot, 2008:Tab. 2).While thesmall bones arecomplete oralmost com- plete, the long bones exhibit recent fragmentation related tothe looting activities orpost-depositional processes. Thefracture char- acteristics of the long bones clearly indicate a complete shaft circumference, atransverse fracture outline andasmooth fracture edge.Allthese criteria suggest that these longbones were broken when they were dry, i.e.anindication ofpost-depositional break- age(Villa andMahieu, 1991).Finally, thebonesurfaces show some traces ofmanganese (n=3),concretions (n=3)andcarnivore tooth pits(n=3).
3.2. Thedistribution ofskeletal elements
Among skeletal elements there is a dominant presence of isolated teeth (20.8%) and extremity elements (45.8%) including carpal, tarsal, phalanxes andmetapodial; trunk elements, i.e.verte- brae and ribs, contribute a smaller percentage (16.7%) (Table 3, Fig.6).Thelongbonesarenever complete andareveryfragmented.
Humerus elements are represented bytwo distal andone proxi- malfragment, tibia byonedistal fragment, femur byfour proximal andfour distal fragments, the ulna byfour proximal andonedis- tal fragments, and the radius by three shafts, onedistal and one proximal fragments. There isnomarked predominance ofasingle anatomical region ormodule (e.g.forelimb vshindlimb) orinbone laterality ( 2=19.09;df=22;p=0.639), which excludes anyselec- tive transport (i.e. anthropogenic, carnivore, hydraulic) ofbrown bear bones within thecave (cf. Quiles, 2003).This type ofassem- blage, which consists primarily ofisolated teeth and extremities of the appendicular skeleton (i.e. carpal, tarsal, metapodial and phalanges), isfound inmany caves ordens ofUrsus denigeri (e.g.
Château, Scharzfeld, Westbury, Vertesszöllos II,Fosseetal., 2002: Fig. 3) and Ursus spelaeus (e.g. Ekain, Azé 1–3, Divje Babe, Fosse etal.,2002:Fig.4)inEurope andintheCaucasus (Hovk-1 cave,Bar- Oz etal., 2012). This type ofskeletal representation isalso found in the Mesolithic rock shelter of Kostias Klde in the Republic of Georgia where brown bears have been hunted byhumans (Bar-Oz etal., 2009),highlighting thedifferential survivorship ofdifferent skeletal elements.
4. Thanatocoenosis structure and mortality profile
Two main methods were utilised inorder toestimate the age atthe time ofdeath. The first method through the estimation of thestage oftooth eruption, andbyanalysing dental wear. Follow- ing sequential eruption, teeth become progressively more worn and distinctive wear patterns are formed on the different enamel
Fig.3. TeethofUrsusarctosfromWemezh cave.LeftupperP4(WZ 109,1–2=lingual view; 3=occlusal view); rightupper M1(WZ104, 4=lingual view; 5=occlusal view);
rightupperM2(WZ101,6=lingual view; 7=vestibular view; 8=occlusal view); leftuppercanine (WZ111,9=lingual view; 10–11=buccalview); rightlowerM3(WZ1598, 12–14 =occlusal view; 13–15 =vestibular view); lower M2right(WZ 1594,16–18 =occlusal view; 17–19 =vestibular view).
Dentsd’Ursus arctosde lagrottedeWemezh. P4supérieure gauche(WZ 109,1–2=vuelinguale ;3=vueocclusale) ;M1supérieure droite(WZ 104,4=vuelinguale ;5=vue occlusale) ;M2supérieure droite (WZ 101, 6=vue linguale ;7=vuevestibulaire ;8=vueocclusale) ;canine supérieure gauche (WZ 111, 9=vuelinguale ;10–11 =vue vestibulaire) ;M3inférieure droite (WZ 1598, 12–14=vue occlusale ;13–15 =vue vestibulaire) ;M2 inférieure droite (WZ 1594, 16–18 =vue occlusale ;17–19 =vue vestibulaire).
facets. According tothe work ofQuiles (2003, 2004),ageestima- tion iscorrelated tonine agegroups (see adescription ofeachage group inQuiles, 2004:tab. 5): infantile, juvenile, sub-adult 1,sub- adult 2, adult 1,adult 2,adult 3,old adult and very adult. These
age-classes have unfortunately not yet been calibrated with real ages known for present day brown bear. The second method is based ontheepiphyseal fusion (Weinstock, 2009),andislessreli- able asaresult ofthe various taphonomical processes that affect
Fig.4. Post-cranial elements ofUrsusarctosfromWemezh cave.Rightproximal radius(WZ465,1=posterior view,2=proximal view); Rightdistalradius (WZ466,3=posterior view, 4=anterior view); right proximal ulna (WZ 471, 5=lateral view); left proximal femur (WZ 1560, 6=anterior view, 7=posterior view); leftdistal femur (WZ 459, 8=posterior view); leftdistal femur (WZ460,9=posterior view); leftcalcaneus (WZ456,10–12 =anterior view,11–13 =posterior view).
Eléments post-crâniens d’Ursus arctosde lagrottedeWemezh. Radius proximal droit(WZ465, 1=vuepostérieure ;2=vueproximale) ;radius distaldroit(WZ 466,3=vue postérieure, 4=vueantérieure) ;ulnaproximale droite(WZ 471, 5=vuelatérale) ;fémur proximal gauche (WZ1560, 6=vueantérieure, 7=vuepostérieure) ;fémur distal gauche(WZ 459,8=vuepostérieure) ;fémur distalgauche(WZ 460,9=vuepostérieure) ;calcanéum gauche(WZ456,10–12 =vueantérieure, 11–13 =vuepostérieure).
skeletal remains, especially those ofimmature individuals andthe types ofskeletal elements with ahighmarrow andspongiosa con- tent. Only four imprecise age-classes may be defined with the appendicular epiphyseal fusion method: foetus ornewborn, juve- nile, sub-adult andadult.
Thus, basedon40identifiable teeth found atWezmeh, 29teeth (12 lower and 17 upper teeth) were assigned an age group, and when combined with tooth type aminimum of eight individuals were identified: one juvenile, three sub-adults and four adults (Table 4).Theonlyunfused bonespresent inthebrown bearassem- blageareonedistal metapodial (anelement which fusesaround 4–5 years) andone proximal phalanx (which fuse around 2–3 years).
This confirms theabsence ofyoung andoldbears from theassem- blage.
5. Osteometry: The Wezmeh brown bear
The Wezmeh brown bear metrical analysis is presented in Table 5forthe teeth andTables 6–9 for thepost-cranial material.
The measurements were taken following theprocedures outlined by Schweizer (1999) .The tooth dimensions ofU.arctos from the Wezmeh cave were compared with those from other Southwest Asian sites, and the post-cephalic dimensions were compared with material from various European assemblages. However, due to the small sample size and to the lack of published data on brown bears, the dimensions of the Wezmeh brown bear bones appear to fall within the size range of both present day and fossilised European brown bears (Altuna, 1973; Ballesio, 1983).
For example, if we calculate the ratio of the length of the first
Fig.5. Metacarpals (MC)andmetatarsals (MT) ofUrsusarctosfromWemezh cave:leftMCII(WZ325,1=lateral view; 2=medial view), rightMCIII(WZ324,3=medial view.
4=lateral view), leftMCIV(WZ1930, 5=lateral view; 6=medial view), leftMCV(WZ323, 7=lateral view, 8=medial view); rightMTI(WZ 327,8=medial view, 9=lateral view); leftMTII(WZ326, 10=lateral view, 11=medial view); rightMTIII(WZ1932, 13=lateral view, 14=medial view), rightMTIV(WZ328,15=medial view, 16=lateral view).
Métacarpiens (MC) etmétatarsiens (MT) d’Ursus arctosde lagrotte deWemezh :MCIIgauche(WZ 325, 1=vuelatérale, 2=vuemédiale) ;MCIIIdroit (WZ 324, 3=vue médiale, 4=vuelatérale), MCIVgauche(WZ1930, 5=vuelatérale, 6=vuemédiale) ;MCVgauche(WZ 323,7=vuelatérale, 8=vuemédiane) ;MTIdroite (WZ327, 8=vue médiale, 9=vuelatérale) ;MTIIgauche (WZ 326, 10=vuelatérale, 11=vue médiale) ;MT IIIdroite (WZ 1932, 13=vuelatérale, 14=vuemédiale) ;MTIVdroit(WZ 328, 15=vuemédiale, 16=vuelatérale).
metatarsal asa percentage of the length of the fifth, we obtain 72.3%fortheWezmeh material. This index ranges between 71.1%
and 74.2% for Ursus arctos, and between 62.3% and 66.9% for Ursus spelaeus and Ursus deningeri (Ballesio, 1983: Tab. 19). The
teeth (Fig. 7) ofthe Wezmeh specimens are larger than those of the present day Syrian brown bear (U. a.syriacus) and are sim- ilar tothose found at Middle Palaeolithic sites such as Ksar’Akil or Tabun C, which are prehistoric sites with later dates than
Table 3
Census ofbrown bear(Ursusarctos)remains onWezmeh caveinnumber ofidenti- fiedspecimens (NISP)andinminimum numberofindividual ofcombination (MNIc)/
nventaires desrestesd’oursbrun(Ursus arctos)de lagrottedeWezmeh ennombre despécimens identifiés (NISP) etennombre minimal d’individu decombinaison (MNIc).
Skeletal element NISP %NISP MNIc
Left Right Ind/Imp Total
Mandible – 1 – 1 0.52 1
Upperisolated teeth 8 9 1 18 9.38 5
Lowerisolated teeth 13 8 1 22 11.46 8
Thoracic vertebra – – 13 13 6.77 –
Lumbar vertebra – – 1 1 0.52 1
Rib – – 17 17 8.85 –
Sternebra – – 1 1 0.52 1
Humerus 1 2 – 3 1.56 2(D)
Radius 1 2 2 5 2.60 3(SH)
Ulna 1 4 – 5 2.60 3(P)
Carpal 3 – 1 4 2.08 2
Metacarpal II 2 1 – 3 1.56 2
Metacarpal III 1 2 – 3 1.56 2
Metacarpal IV 2 1 – 3 1.56 2
Metacarpal V 2 1 – 3 1.56 2
Pelvis 3 3 – 6 3.12 3
Femur 4 4 – 8 4.17 3
Patella 1 – – 1 0.52 1
Tibia – 1 – 1 0.52 1(SH)
Fibula 2 – – 2 1.04 2(D)
Tarsal 4 5 2 11 5.73 2
Metatarsal I 1 1 – 2 1.04 1
Metatarsal II 1 1 – 2 1.04 1
Metatarsal III – 2 – 2 1.04 2
Metatarsal IV 1 3 – 4 2.08 3
Metatarsal V – 2 – 2 1.04 2
Metapodial – 1 6 7 3.65 –
Phalange I – – 22 22 11.46 –
Phalange II – – 9 9 4.69 –
Phalange III – – 11 11 5.73 –
TOTAL 51 54 87 192 100 8
SH:diaphysis; D:distalextremity; P:proximal extremity/ SH:diaphyse;D:extrémité distale;P:extrémitéproximale.
the Acheuleo-Yabrudian site ofZuttiyeh (Tchernov and Tsoukala, 1997).
Likeotherursids, thebrown beardisplays astrong sexual dimor- phism, which is particularly notable in the transverse diameter of the canines (Koby, 1949; Kurtén, 1955). However the overlap
Table 4
Minimum number ofindividuals according tothe9classes ofage.
Nombre minimum d’individus selonles9classes d’âge.
I Juv SA1 SA2 A1 A2 A3 OA VOA Total
Lowerteeth 0 1 1 1 1 1 0 0 0 5
Upperteeth 0 1 2 1 2 1 1 0 0 8
Total(MNIc) 0 1 2 1 2 1 1 0 0 8
I:infantile; Juv:juvenile; SA1: sub-adult 1;SA2: sub-adult 2;A1:adult1;A2:adult 2;A3:adult3;OA: oldadult; VOA: veryadult/I:infantile ;Juv:juvénile;SA1:sub- adulte1;SA2:sub-adulte 2;A1:adulte1;A2:adulte2;A3:adulte3;OA:adulte âgé;VOA:adultetrèsâgé.
between the two sexes is such that it is impossible toseparate them only from univariate andbivariate diagrams (Kurtén, 1955;
Schweizer, 2004).Itistherefore preferable tousemore reliable sta- tistical methods, such as mixture analysis (Quiles and Monchot, 2004; Quiles etal., 2005). However thesmall sample size andthe general lackofdataonsexual dimorphism inbrown bearsalso pre- vents thetypesofanalyses thatarenecessary inorder todistinguish male andfemale remains.
6. Discussion
6.1. Structure oftheWezmeh bearpopulation
The mortality profile indicates that sub-adult and adult males appear tohave preferentially occupied the cave. The absence of very young juveniles may beexplained bythefactthatthey donot hibernate andareexclusively nursed bytheir mother (Nelson etal., 1983).Inaddition, they areprobably underrepresented because of the fragility ofthe tooth buds andbone remains, which are sub- jecttoextensive destruction, exacerbated bytheabsence ofsieving during the excavation. The sub-adult period, which includes the first two hibernations, is critical toindividual survival. Foraging by themselves, and still physically immature and inexperienced, young adults have the greatest probability ofdying during hiber- nation (McLoughlin et al., 2003). An abundance of young adult remains is found for instance on the sites of Fate, Basura, Hor- tus and Arbreda (Quiles, 2003). The presence of adult 1remains in the Wezmeh assemblage, mostly males (e.g. Badalucco), is probably due tothese factors, especially as the dominant older
Fig.6. Anatomical distribution ofWezmeh brown bearremains established as%ofthenumber ofidentified specimens (NISP).
Profil anatomique de l’ours brun de Wezmehétablie en %du nombre de spécimens identifiés (NISP).
Table 5
Toothmeasurements ofUrsusarctosfromWezmeh cave.
Mesuresdesdentsd’Ursus arctos delagrottedeWezmeh.
L B Bpost HC Wear stage
WZ101 UpperM2 35.7 19.4 15.8 8.1 4
WZ100 UpperM2 39.2 21.1 16.8 9.1 4
WZ102 Upper M2 38.2 20.7 – 8.5 5
Bant
WZ104 Upper M1 24.6 17.1 16.7 9.0 6
WZ109 UpperP4 15.0 11.6 7.7 4
WZ1596 UpperP4 16.2 12.3 8.3 4
WZ1597 UpperP4 15.5 11.2 9.0 2
WZ116 UpperI3 9.3 10.2 – 4
WZ119 Upper I3 9.8 12.5 3.3 5
WZ118 Upper I3 10.4 13.1 7.9 4
WZ120 Upper I3 11.6 13.1 3.3 6
WZ1592 UpperI3 11.2 12.9 3.0 6
WZ125 UpperI1 7.0 8.1 8.1 3
B DTrac
WZ110 UpperC 17.1 21 – 2
WZ111 Upper C 16.3 19.1 – 3
L Bpost Bant
WZ108 LowerM2 23.7 14.4 13.9 8.0 5
WZ105 LowerM2 26.6 17.0 15.7 7.5 4
WZ1594 LowerM2 24.7 15.1 14.4 9.3 4
L B Bpost
WZ102 LowerM3 22.4 16.0 – 5.7 3
WZ106 Lower M3 20.7 15.7 – 8.7 4
WZ107 LowerM3 22.4 16.0 – 5.7 3
WZ1598 LowerM3 21.7 16.0 – 6.5 3
WZ113 LowerM1 – – 13.9 – 5
WZ82 LowerP4 14.2 7.3 – 7.3 3
WZ1589 LowerP4 13.9 7.1 – 7.1 3
WZ126 LowerI2 6.6 9.3 – 4.7 3
WZ124 Lower I2 6.8 9.5 – 2.9 5
WZ1591 LowerI3 8.6 9.7 – 6.2 5
Allmeasurements areinmm.L:Antero-posterior lengthnearthebaseofthecrown;
B:transversal breadth near thebaseofthecrown; HC:height ofthecrown; Bant:
anterior transversal breadthnearthebaseofthecrown; Bpost: posterior transversal breadth nearthebaseofthecrown/Touteslesmesuressontenmm.L:longueurantéro- postérieureàlabasedelacouronne;B:largeurtransversale àlabasedelacouronne; HC:hauteur delacouronne ;Bant:largeur transversale antérieure àlabasedela couronne;Bpost:largeurtransversale postérieure àlabasedelacouronne.
males exert a social pressure that results in restricted access to food resources (Stokes etal., 1981; Nelson et al., 1983). Adult 1 males are also typically forced tolive inless favourable environ- ments (see thedispersion ofsub-adult males, McLellan andHovey, 2001), as dominant males already occupy the highest quality territories.
Furthermore, it’s interesting tonote that bear MNI based on teeth isabout twice the MNI based onpost-cranial boneelements (3 to8), emphasizing the dramatic loss of bone material in situ in Wezmeh Cave. Relatively speaking, the bones from Wezmeh Cave are in good condition, and the conditions for fossilisation appear to befavorable; the lack of certain skeletal parts there- fore cannot be readily explained. The cavity has been used by many carnivores, including the spotted hyena (Crocuta crocuta spelaeaGoldfuss, 1823), which hasthereputation ofbonecrusher (Monchot, 2008). We may suppose that some ofthe bear bones were therefore consumed while thetissue was still fresh andedi- ble.There isevidence ofgnawing by carnivores onmaterial from thecave, suggesting thatsomefragile boneelements (such asthose that are thin, flat and trabecular) may have been partly orcom- pletely destroyed bybiological agency soon after death(Mashkour et al., 2009: 689).
Table 6
Measurements ofthecalcaneus, patella, femur andradius andulnaofUrsusarctos fromWemeh cave/
Mesures ducalcanéum, delapatella, dufémur,duradiusetducubitusd’Ursus arctos delagrottedeWemeh.
GL DT DAP Lmanu DTfacdist DAPst DDPst
WZ456 Calcaneus 76.5 50.9 38.5 42.9 26.5 19.4 15.5
GL DT DAP DAPart. Lart.
WZ458 Patella 49.7 35.8 22.6 20.5 38.9
DAP DT DTtroch DTfossa DAPepimed
WZ460 Femur distal (66.2) 81 43.2 15.4 (50.5)
WZ459 Femur distal 55.1 80 42.0 15.9 50.3
DAPcol DTcol Bcaput Dcaput
WZ463 Femurproximal 21.6 33.9 – –
WZ462 Femurproximal – – 49.0 48.1
WZ1560 Femurproximal 20.5 32.7 42.7 41.2
DAPcol DTol
WZ471 Ulnaproximal 69.2 45.1
WZ1561 Ulna proximal 63.1 –
DTd DAPd
WZ468 Ulnadistal 41.4 24.2
WZ472 Fibula distal 25.6 15.8
WZ1566 Fibula distal 25.9 16.0
WZ466 Radius distal – 58.4
Allmeasurements areinmm. GL:greatest length; DT:transversal diameter; DAP:
antero-posterior diameter; Lart.: length ofthearticulation; DAPart.: articulation antero-posterior diameter; Lmanu: lengthofthemanubrium;DTfacdist: transversal diameter ofthedistalarticulation; DAPst:antero-posterior diameter ofthesusten- taculum tali; DDP st: dorso-plantar diameter ofthesustentaculum tali;DTtroch:
maximum transversal diameter ofthetrochlea; DTfossa: maximum transversal diameter ofthefossa between thetwo condyles; DAP epimed: antero-posterior diameter ofthemedial condyle; DAPcol:antero-posterior diameter ofthediaph- ysisjustundertheproximal epiphysis; DTcol:transversal diameter ofthediaphysis just under theproximal epiphysis; Bcaput: Greatest breadth ofthecaputfemoris;
Dcaput: greatest depth ofthecaputfemoris; DTol:olecranon transversal diame- ter/Touteslesmesuressontenmm.GL:longueurmaximale;DT:diamètretransversal ; DAP:diamètreantéro-postérieur ;Lart.:Longueurdel’articulation ;DAPart.:diamètre antéro-postérieur del’articulation ;Lmanu :longueurdumanubrium ;DTfacdiste: diamètre transverse del’articulation distale ;DAPst:diamètre antéro-postérieur du sustentaculum tali;DDPst:diamètredorso-plantaire dusustentaculum tali;DTtroch: diamètre transverse maximal delatrochlée ;DTfosse:diamètre transverse maxi- maldelafosseentrelesdeuxcondyles;DAPepimed:diamètre antéro-postérieur du condyleinterne;DAPcol:diamètreantéro-postérieur deladiaphyse justeendessous del’épiphyseproximale ;DTcol:diamètretransversedeladiaphysejusteendessousde l’épiphyseproximale ;Bcaput:laplusgrandelargeurducaputfemoris;Dcaput:plus grandeprofondeurducaputfemoris;DTol:diamètre transverse del’olécrane.
6.2. AnupdateonthegeneticsofbrownbearsintheSouthwest Asia
Thegenetics ofSouthwest Asian bears have notyet been stud- ied indetail despite alarge number ofpublications (Taberlet and Bouvet, 1992; Talbot and Shields, 1996,Miller etal., 2006). Most recently, Calvignac and colleagues have analysed samples from Syria, Lebanon andIran (Calvignac etal., 2009). Thephylogenetic reconstruction basedonmtDNA hasrevealed ahighgenetic diver- sity, with thepresence ofthree distinct clades (Miller etal.,2006).
The Syrian specimens display a genetic proximity to the east- European bears, whereas theLebanese specimens arecloser tothe west-European bears (Talbot andShields, 1996).Iranian bearsform athird unique groupthat forthemoment cannot begrouped with any ofthe others (Miller etal.,2006; Calvignac etal., 2009). The history of the species inthis area is therefore much more com- plex than previously thought, and it may have involved several
