The Process of UngulateDomestication atÇayönü,SoutheasternTurkey: AMultidisciplinary Approachfocusing on Bos sp. and Cervus elaphus
Hitomi HONGO School of Advanced Science,GraduateUniversity forAdvanced Studies ShonanVillage,Hayama, Kanagawa240-0193(Japan) [email protected]
Jessica PEARSON School of Archaeology,Classicsand Egyptology,University of Liverpool 12-14 AbercrombySq.,Liverpool L697WZ (United Kingdom) [email protected]
BanuÖKSÜZ Faculty of Prehistory,IstanbulUniversity Beyazit,Istanbul(Turkey) [email protected]
Gülçin l˙ LGEZDI Institut für Ur-und Frühgeschichteund ArchäologiedesMittelalters Eberhard-Karls-UniversitätTübingen Schloss Hohentübingen,72070Tübingen (Germany) [email protected]
Hongo H.,Pearson J.,Öksüz B.& I˙ gezdi G.2009. —The Process of UngulateDomestica- tion atÇayönü,SoutheasternTurkey:AMultidisciplinary Approachfocusing on Bos sp. and Cervus elaphus.Anthropozoologica 44(1):63-78.
ABSTRACT Bos and Cervus remainsfrom Prepottery andPottery Neolithiclevelsat ÇayönüTepesiareexamined employing amultidisciplinary approach, combining theanalysisofmorphology,ageprofiles,andstableisotopesin bonecollagen. Theresults showthattheprocess ofcattledomestication started atabout thesametimeasthree otherungulatetaxa(sheep,goats and pigs),bytheChannelled Building Subphase(endofEarlyPPNB/beginning ofMiddlePPNB). Twostagesareevidentin theprocess ofdomestication: theinitialappearance ofdomesticanimalscouldbedetected in thefaunal
ANTROPOZOOLOGICA •2009•44 (1) ©PublicationsScientifiquesduMuséumnationald’Histoirenaturelle,Paris. 63 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
KEYWORDS remainsbytheappearance ofsomesmall-sized individualsandsubtlechanges ÇayönüTepesi, in thekill-off patterns,aswell asin thechangesin stableisotoperatiosof SoutheasternTurkey, Prepottery Neolithic, carbon andnitrogen. Whilehunting ofwildprogenitors continued, there Domestication, wasanoverall decreasein theproportion ofmiscellaneous wildtaxain the Pigs,Sheep, Goats,Cattle, assemblage.Thesecondstageofthedomestication process beginsin thelate- Red deer, finalPPNB,suggested bymarked sizereduction anddemographicchange, Kill-off patterns, namelytheincreaseoffemalesin theassemblage, aswell asdramaticincrease Sizeindexanalysis, stableisotopeanalysis in sheepandgoats.
RÉSUMÉ Leprocessusde domestication desOngulésàÇayönü,TurquieduSud-Est: approchemultidisciplinaireàpropos de Bos sp. etde Cervus elaphus Lesrestesde Bos etde Cervus provenantdesniveaux néolithiques précéramiquesetàpoterieàÇayönüTepesiontétéétudiésensuivantune approchemultidisciplinaire, combinantl’analysemorphologique, lesprofils d’âgeetlesisotopesstablesdanslecollagènedesos.Lesrésultats montrent queleprocessus de domestication dubétail acommencé enmêmetempsque troisautresOngulés(moutons,chèvresetcochons)parlaphase Channelled Building (fin duPPNBancien/début duPPNBmoyen). Deux étapes sontmisesenévidence dansleprocessus de domestication :lespremières tracesd’animaux domestiquespeuventêtredétectéesdanslesrestesde MOTS CLÉS fauneparl’apparition de quelquesindividus de petitetailleetde subtils ÇayönüTepesi, changements danslesmodèlesd’abattage, évolution attestée parl’observation TurquieduSud-Est, Néolithiqueprécéramique, de changements danslesratiosdesisotopesstablesducarboneetde l’azote. domestication, Bienquedesanimaux sauvagesaientcontinuéàêtrechassé, on noteune cochons,moutons, bétail, diminution globaledesindividus sauvagesdanslesassemblages.Ladeuxième chèvres, étapeduprocessus dudomestication commence danslePPNBtardif, comme Cerfélaphe, lesuggèrentunenetteréduction de lataille, unchangementdémographique, profilsd’abattage, indicesde taille, àsavoirl’augmentation desfemelles,etaussil’augmentation considérabledes analysesisotopiquesstables moutonsetdeschèvres.
INTRODUCTION ofTaurus Mountainsin southeasternAnatolia. RecentresearchatEarlyandMiddlePPNBsites Domestication offour importantlivestockspecies in southeasternAnatoliaandnorthernSyria, such (sheep,goats,cattleandpigs)progressed in the asCaferHöyük,NevalıÇori,Tell Halula, aswell northernandeasternpart ofthe“FertileCrescent” asÇayönüsuggests thatthedomestication process during theEarlyandMiddlePPNB.Evidence from ofpigsandcattlemightalsohavebegunatabout faunalremainsfrom archaeologicalsitesin the thesametime(Helmer1988, Hongo &Meadow area suggests thatthemanagementofgoats began 1998, 2000; Rosenberg&Redding 1998, Peters in thenorthernpart oftheZagrosMountainsby etal. 1999,2005;Ervynk etal. 2001,Hongo etal. about 10000 years ago (Hesse1978, Zeder1999, 2002,2004;Helmer etal. 2005). 2005,2006; Zeder&Hesse2000). Theprocess of Morphologicalchanges,including reduction ofbody sheepdomestication alsobeganatabout thesame size, andkill-off patternsareimportantandcom- time, probablyin thearea including thefoothills monlyused markers in zooarchaeologicalresearch
64 ANTHROPOZOOLOGICA •2009•44 (1) The Process of Ungulate Domestication at Çayönü, Southeastern Turkey
to monitor initial domestication. Changes observed Grill Subphase corresponds to the Early PPNB in each of these markers, however, are in many cases (ca. 8500–8300 cal. BC). The following Channelled subtle and often allow multiple interpretations. Building Subphase goes into the beginning of the Therefore, it is useful to examine the faunal assem- Middle PPNB (ca. 8300-8200 cal. BC), which conti- blage employing more than one analytical method, nues with the Cobble-paved (ca. 8200-7800/7500 in order to see whether the changes observed in cal. BC) into the Cell Building Subphases. The Late different kinds of analysis coincide in timing, and PPNB includes most of the Cell and Large Room whether there is a trend that might suggest a kind Building (ca. 7800/7500-6800 cal. BC) Subphases, of human intervention on the animal population but with at least part of the latter continuing into that might lead to domestication of the species. the Final PPNB (or “PPNC”: ca. 6800-6300 cal. In this paper, we examine Bos remains from Çayönü BC) (calibrated dates based on the CANeW Upper in southeastern Turkey using multiple analytical Mesopotamia 14C chart, updated Feb. 2006, www. approaches in order to document the timing and canew.org). This Prepottery sequence is immedia- process of the domestication of cattle. The trend tely followed by early Pottery Neolithic (Özdoğan observed in Bos remains through time is compared 1995, 1999; Hongo & Meadow 2000, Ervynck et with that in Cervus , another large ungulate exploited al . 2001), which is also contemporary with the Final at the site, as well as in caprines and pigs. PPNB in the Levant. Although it is problematic to apply the terms used in Prepottery Neolithic chro- nology in the Levant to Anatolia, in this paper the THE SITE terms are used strictly as the periodization without reference to material culture. Çayönü Tepesi is located on one of the small tri- butaries of the Tigris, about five kilometers from the foot of the Taurus Mountains in southeastern TRENDS OBSERVED IN THE RELATIVE Turkey (Özdoğan & Özdoğan 1990). Like other PROPORTION OF ANIMAL PPN sites in the region, the site is situated at the TAXA THROUGH TIME foot of the mountains along an important route connecting the mountain area and the plains and Pigs are the most abundantly represented taxon steppes to the south. The occupation of the site during the Prepottery Neolithic at Çayönü, and spans about 3000 years, from Late PPNA into the always represent close to 40 percent of the identified early Pottery Neolithic. Thus, the faunal assemblage specimens, especially in the earlier three subphases from Çayönü provides us with the opportunity to (Table 1) (Hongo & Meadow 1998, 2000; Hongo examine changes in the animal exploitation pat- et al . 2002, 2005). Four species of domestic lives- terns through time at a single site over the critical tock (pigs, cattle, sheep, and goats) and their wild period when the process of animal domestication progenitors are grouped together in Table 1. Sheep progressed in the region. and goats were insignificant in the early phase at The Prepottery Neolithic period at Çayönü is divi- the site, but their proportion, especially of sheep, ded into six subphases, with each period named increases steadily. A wide range of wild taxa were for characteristic architectural types (Özdoğan & exploited in early subphases, including red deer, Özdoğan 1990, 1998; Özdoğan A. 1995, 1999; gazelle, roe deer, Asiatic wild ass, brown bear, leo- Bıçakçı 1998, Erim-Özdoğan 2007). The Round pard, red fox, hare, and a few other small mammal Building subphase (ca. 10200-8500 cal. BC, with species. Some birds and tortoise were also exploited, possible hiatus between 9300-8700 BC) corresponds and fish remains exist, though rare. The proportion to the Prepottery Neolithic A (PPNA) period of of these miscellaneous wild taxa, however, decreases the Levant. The early part of the Grill Building significantly after the Channelled Building Sub- subphase is also considered to be contemporary phase. On the other hand, the proportion of the with the Prepottery Neolithic A (PPNA) period four ‘pro-domestic’ taxa, by which we mean pig, (Özdoğan A. 1995, 1999). The remainder of the sheep, goat, and cattle, steadily increases through
ANTHROPOZOOLOGICA • 2009 • 44 (1) 65 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
T ABLE 1.— Relativeproportion of pigs,sheep and goats,cattle,and miscellaneous wild taxain eachsubphaseatÇayönü (percentage based on NISP).
Totalofthe four Building Sheep Other Pigs Cattle “pro-domestic” subphase and Goats wild taxa taxa Round 35,96,917,960,735.9 Grill 44,610,89,464,831.9 Channelled 37,914,75,958,537.9 Cobble-paved 31,322,913,968,226.4 Cell 31,924,217,974,119.2 Large Room 21,953,613,188,79.4 EarlyPN 35,446,611,393,36.2
Notes:“Otherwild taxa”include gazelle,cervids,equids,bear,fox,hare,and miscellaneous small mammals,birds,and amphibia. Wild and domesticformsof pigs,sheep,goats,and cattle arenotdifferentiated. Animaltaxanotreflected in thistable aredomesticdogsand unidentified mediumand large bovidsorcervids. time.Sheepandgoats,especially,increasein the respectively.Whileintensivelyexploiting themost latersubphasesandtheirproportion reached over accessibletaxon,whichgenerallycomprised more 50percentofthetotalNISP bytheendofPPNB. thanonethirdanduptoasmuchas60percentof Thus,twoimportanttrendsthrough timecanbe thefaunalremainsatasite, awide variety ofwild observed atÇayönü:thefirst isthedecreasein the animalswerealsohunted.Theshiftin subsistence proportion ofmiscellaneous wildtaxaaround8300 from suchbroad-spectrumstrategytoincreasing calibrated BC.Thesecondistheincreaseofsheep dependence on four “pro-domestic”taxaoccurred andgoats,whichinthebeginning wasgradual, byMiddlePPNB(Hongo etal. 2002,2005). Sheep becoming dramaticin theLatePPNB. andgoats,whichwereinsignificantin theearly ThesetwotrendsareuniversalinsoutheasternAna- subphasesatthesite, becamedominantbyLate tolia.Theinitialsign ofchangein thecontinuum PPNB.Thisphenomenon wasprobablyrelated to ofthedomestication process ismanifested asa theinitialkeeping ofdomesticanimals. shiftfrom abroad-spectrumanimalexploitation Thesecondshiftin animalexploitation patterns strategycombined withtheintensiveexploitation atarchaeologicalsitesin theregion isadramatic ofadominanttaxon toastrategyconcentratingon increasein sheepandgoats in theLatetoFinal sheepandgoats around7500-7000calibrated BC. PPNB.AtsiteslikeHayazHöyük(Buitenhuis During thePPNAandEarlyPPNB,settlements in 1985),GürcütepeII (Driesch&Peters1999), southeasternAnatoliaspecialized in theexploitation Çayönü(LargeRoom andearlyPottery Neolithic ofoneparticularanimalspeciesthatwasthemost levels),andGritille(Stein 1986),theproportion of accessibletaxon atthesite(Table2). Wildpigs sheepandgoats,themajority ofwhichareconsi- wereheavilyexploited atÇayönuandwildsheepat dered domesticbased on bothbonesizeandkill HallanÇemi. Located closertothehigherelevation off patterns,jumpsto50-70 percentofthefaunal ofTaurus mountains,wildgoats wereabundantat remains(Table2). CaferHöyük,whilegazelleswerethemain gameat siteslocated closetotheHarranPlains.Proportions ofsheepandgoatbonesatmost sitesareonlyabout COMPARISON OF CATTLE O BOS SP.U 10percentuntil theendoftheEarly/ beginning AND RED DEER O CERVUS ELAPHUS# oftheMiddlePPNB.TheexceptionsareHallan Çemi (Rosenberg etal .1995;1998)andCafer Comparison ofchangesobserved in various mor- Höyük(Helmer1998),wheretheremainsofwild phologicalandnon-morphologicalmarkers in Bos sheepandwildgoats dominatefaunalassemblages, anddeerthrough timeprovide us withauseful
66 ANTHROPOZOOLOGICA •2009•44 (1) TheProcess ofUngulateDomestication atÇayönü,SoutheasternTurkey
T ABLE 2.–DominantspeciesatNeolithicsitesin southeastern specieswouldhelp us distinguishing humaninduced Anatolia. factors from theenvironmentalfactors thatmight haveaffected themorphology,demography,and SiteDominantSpecies% behaviorofthesetwospecies. HallanÇemi wild sheep 43.0 Çayönür pig 36.5 P ROPORTION OF B OS AND C ERVUS Göbekli Tepe gazelle 43.0 Figure1AandBshowtheproportion of Bos and Nevali ÇoriI/II gazelle 63.0 Cervus in eachlevelatÇayönü.Theproportion Çayönügpig 44.7 of Bos iscloseto20%ofthetotalNISP in the Çayönüchpig 37.9 PPNARoundBuilding Subphase, but decreases Nevali ÇoriIII gazelle 59.0 during theEarlyPPNBandearlypart ofMiddle CaferHöyükwild goat42.9 PPNB(Grill andChannelled Building Subphases), Çayönücppig 31.3 thenturntoasuddenincrease.Theproportion of Nevali ÇoriIV gazelle 42.0 Cervus increasesdramaticallyin theChannelled Çayönücpig 31.9 Subphase, asifcompensating forthedecreasein Gritille sheep and goat71.0 Bos byintensivered deerhunting. Thismighthave HayazHöyüksheep and goat64.0 resulted in thenearextermination ofred deerin Gürcütepe II sheep (and goat)65.0 thevicinity ofthesite.Theproportion of Cervus Çayönülrsheep and goat53.6 begantodecreasein thefollowing Cobble-Paved ÇayönüPN sheep and goat46.6 Subphase.Thedecreasein wildtaxain generalis observed starting aftertheChannelled Building Subphase, andtheaurochs,thelargest mammal insightin theprocess ofdomestication. Bothaurochs speciesin theregion,mighthavebeenthefirst andred deerarelargeungulatesadapted toforest wildmammalthatwasaffected byhunting and environments andwereactivelyhunted atÇayönü themodification oftheenvironmentin thevicinity during thePrepottery Neolithicperiod.Only Bos, ofthesite, namely,thedecreaseofforests because however,wasdomesticated atsomepointduring ofcontinuous occupation ofthesiteandother thePPNB.Therefore, comparison ofthesetwo humanactivities.Thisprocess issuggested bythe
%A:Cattle % B:Red deer 20 20
15 15
10 10
5 5
0 0 RGCH CP CLRPN rgch cp cLRPN
subphase subphase
Fig. 1. —Proportion of cattle and red deerin eachsubphaseatÇayönü. R: round;G: grill;CH: channelled;CP: cobble paved;C: cell;LR: large room;PN: Pottery Neolithic.
ANTHROPOZOOLOGICA •2009•44 (1) 67 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
0.2 SubphaseatÇayönü,aswell asthatatÇatalhöyük, whichareconsidered tobe wild(Russell &Martin 2005;Russell etal. 2005) fitin thehypothesized size 0.1 LSI n=21 n=20 n=24 rangein theaurochs(Fig. 2).Thesizedistribution of n=17 n=15 n=18 Bos atPPNAlevelofKörtikTepe,alsolocated on the 0.0 upperTigris(Arbuckle&Özkaya2006),alsoshows wild asimilarrange, though slightlylarger.WhentheLog
-0.1 max SizeIndices(LSI)of Bos areplotted in theformof
domestic min 75% histogram,thesizeof Bos in theRoundBuilding 25% median Subphaseshows abimodaldistribution (Fig. 3A), outlier -0.2 RGCHCP CLRPN whichprobablyreflects thesexualdimorphismof Subphase aurochsin theregion.Thefemaleaurochsin Anatolia tendtobe smallerthantheDanishfemaleused as F IG.2.–Sizeofcattle atÇayönü. R: round;G: grill;CHCP: channelled and cobble paved;C: cell; thestandardanimal. LR: large room;PN: Pottery Neolithic. Theoverall rangeofsizedistributionsdidnotchange in thefollowing Grill-Building Subphase, wherethe Bos samplescamemostlyfrom EarlyPPNBlevels. sharpdecreasein thenumberof Bos remainsin Thereseemtobe morefemalesamong thesamples theassemblageduring theEarlyandearlyMiddle in thissubphase(Figs2&3B),but small sample PPNB.Finallythesuddenincreasein Bos in the sizemakesthisobservation tentative.Thespecimens Cobble-paved Subphasemayindicatetheshiftto from theChannelled (n =8)andCobble-paved keeping domesticcattleatthesite. (n =7)Building Subphaseswerecombined because ofsmall samplesizeandbecausethesizerangesof B ODYSIZE OF B OS AND C ERVUS Bos in thesetwosubphasesaresimilar.Thereisa Inordertoassess whetherornotashiftfrom aurochs slightdecreasein thesizeof Bos in theseearlyMiddle hunting tocattlekeeping occurred bytheMiddle PPNBsubphases(Figs2&3C). Atthesametime, PPNB,thebodysizeof Bos and Cervus from Çayönü thesizerangebecamemuchbroader,withafew wereexamined.Thesizesof Bos long bonesin each specimensfalling belowthesizerangeofaurochs. subphaseatÇayönüwerecompared against themea- Thesmallest specimeninFigure3C,withLSI value surements ofastandardanimal,whichisaDanish -0.14,andanothersmall onewithLSI value-0.077 femaleaurochs(Fig. 2,Degerbøl 1970,but following belong totheChannelled Building subphase.The thecorrected measurements byGrigson1989),using smallest specimenfrom theCobble-paved Buil- the“Difference ofLogs” technique(Uerpmann 1979; ding SubphasehastheLSI value-0.086.Thereis Meadow1981,1983,1999). Although overlapin areboundin sizein thefollowing Cell Building thesizeoffemaleAurochsandmaledomesticcattle Subphase, withall thespecimensfrom thissubphase oftenmakesthecleardistinction betweendomestic falling in thesizerangeofwild Bos (Fig. 3D).This andwildpopulations,estimated rangesofthesizeof reboundmightpartlybe caused bygreaternumber wildanddomestic Bos areindicated byarrows on the ofolderindividualsin theassemblagewhosebreadth leftside ofthechart.Theseestimated sizerangesare dimensionscanbe largerthanyoungerindividuals based on theBronzeAgeaurochsanddomesticcattle (see thesection on ageprofilesbelow). Giventhe remainsfrom DidiGorain Georgia(H.-P.Uerpmann, small numberofsamples,itisalsopossiblethatjust personalcommunication 2007). Theproposed size bychance all themeasured specimensfrom this rangeforaurochsisnarrowerthanthatindicated subphasecamefrom wildindividuals.Aclearshift byGrigson (1989). Eventhough chronologically tosmallersizeisobserved in theLate-FinalPPNB apart,becauseofgeographicalcloseness,theaurochs Large-room Subphase, whichisatleast in part due population in Georgiamightbe comparablein its totheincreaseoffemalesin theassemblage(Fig. 3E), sizerangetotheNeolithicaurochsin easternAna- Furthersizediminution occurred in thefollowing tolia.Thesizeof Bos in thePPNARoundBuilding Pottery Neolithic, manifested in theoverall shift
68 ANTHROPOZOOLOGICA •2009•44 (1) TheProcess ofUngulateDomestication atÇayönü,SoutheasternTurkey
25 25 A. round (n=20) D. cell (n=24)
20 20
15 15
% %
10 10
5 5
0 0 -0,1 -0,1 -0,1 -0,1 -0,1 -0,1 -0,1 -0,1 -0,1 -0,1 -0 -0 -0 -0 00,010,020,030,040,050,06 -0,14-0,13-0,12-0,11 -0,1 -0,09-0,08-0,07-0,06-0,05-0,04-0,03-0,02-0,010 0,01 0,02 0,03 0,04 0,05 0,06 LSI LSI
25 25 B. grill (n=21) E. large room (n=18)
20 20
15 15
% %
10 10
5 5
0 0 -0,14-0,13-0,12-0,11 -0,1 -0,09-0,08-0,07-0,06-0,05-0,04-0,03-0,02-0,010 0,01 0,02 0,03 0,04 0,05 0,06 -0,14-0,13-0,12-0,11 -0,1 -0,09-0,08-0,07-0,06-0,05-0,04-0,03-0,02-0,010 0,01 0,02 0,03 0,04 0,05 0,06 LSI LSI
25 25 C. channelled &cobble-paved (n=15) F. Pottery Neolithic (n=17)
20 20
15 15
% %
10 10
5 5
0 0 -0,14-0,13-0,12-0,11 -0,1 -0,09-0,08-0,07-0,06-0,05-0,04-0,03-0,02-0,010 0,01 0,02 0,03 0,04 0,05 0,06 -0,14-0,13-0,12-0,11 -0,1 -0,09-0,08-0,07-0,06-0,05-0,04-0,03-0,02-0,010 0,01 0,02 0,03 0,04 0,05 0,06 LSI LSI
Fig. 3.–Sizeof Bos atÇayönü. A.round;B.grill;C.channelled and cobble paved;D.cell;E.large room;F.Pottery Neolithic.LSI: Log SizeIndex. in therangeofthesizedistribution andincreasein Thesizerangeof Bos from PPNAandPPNBlevelsat females(Fig. 3F). Themajority ofthespecimensin Çayönücorrespondsvery well withthatatcontem- Late-FinalPPNBandPottery Neolithiclevelsare porary sitesin southeasternAnatolia, suchasfrom well within thesizerangeofdomesticcattle. Göbekli Tepe, Nevali Cori,andGürcütepe(Peters Sex-specificchangesin slaughterpatterncouldnotbe etal. 2005:fig. 11). examined withthedataathandmainlyduetosmall Forthesizecomparison of Cervus ,measurements of samplesize.Although wedidmeasurebothfused amodernfemalecollected in Turkeywereused as andunfused specimens,thenumberofmeasurable thestandard(İlgezdi2002). Incontrast to Bos,the unfused specimenswasvery small,andtheirsizeall sizeof Cervus wasremarkablystablethroughout the fell in thesmallest endofthesizedistribution:unfused Prepottery NeolithicandPottery Neolithic(Fig. 4). specimenswithLSI valuesfarbelowtherangeofLSI Ifthesizediminution observed in Bos werecaused distribution ofotherspecimensfrom thesamelevel byenvironmentalchanges,suchasunfavorablecli- werenotincluded in thecharts.Anunfused distal mateordeterioration in vegetation, Cervus should humerusfrom theLarge-room Subphasehad an alsohavebeenaffected.Thefactthattheshiftin LSI value-0.079,whichwasincluded in thechart. sizeisobserved onlyin Bos suggests thattherewere
ANTHROPOZOOLOGICA •2009•44 (1) 69 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
0.2 Building Subphaseshowhigh survivalratesatboth n=15 n=16 juvenileandsubadultagestages.Inthefollowing n=15 n=51 n=7
0.1 Grill Building Subphase, thekill-off schedulebecame earlier,withonlyabout 60 %oftheanimalssurvi- ving thesubadultagestage.Thiscontinuesin the 0.0
LSI following subphases,with Bos being slaughtered mainlybetweenlatejuvenileandsubadultagestage,
-0.1 max exceptin theCell Building Subphasewherethe min 75% survivalrateisratherhigh in all agestages.Inthe 25% median casesofsomesubphases,a“rebound”atthejuvenile outlier -0.2 R-G CH CP CLR-PN agestageoccurs,whichiscaused primarilybylarge Subphase numbers offused distaltibiae andmetapodials, theelements thattendtobe preserved betterthan Fig. 4. –Sizeofred deeratÇayönü. otherskeletalparts.Slaughteratevenyoungerages R-G: round and grill;CH: channelled;CP: cobble-paved;C: cell; isobserved in theLate-FinalPPNBandPottery LR-PN: large room and Pottery Neolithic. Neolithic.Thesurvivalrateattheinfantileage stagein theselatesubphasesislowercompared to otherfactors involved in thereduction ofsizein thatin theearlierphases,andonly30to45%of Bos,probablyhumaninduced ones. Bos survived intofull adult.Again,in contrast to Bos,therewaslittlechangein thekill-off patterns K ILL- OFF PATTERNS FOR B OS AND C ERVUS of Cervus through time(Fig. 5B). Themajority Figure5Ashows thekill-off patternsof Bos in each ofdeerremainsin theassemblagebelong toadult subphaseatÇayönü. Bos in thePPNARound individualsthroughout theoccupation ofthesite.
% A: cattle B: reddeer 100 90 80 70 60 50 40 30 20 10 0 infantile juvenile subadult adult infantile juvenile subadult adult Age Stage Age Stage
Round (n=52) Round (n=24) Grill (n=56) Grill (n=22) Channeled (n=55) Channeled (n=136) Cobble-paved (n=32) Cobble-paved (n=26) Cell (n=53) Cell (n=32) Large-room (n=46) Large-room (n=8) Early PN (n=51) Early PN (n=5)
F IG.5.—Age profile of cattle and red deeratÇayönü.
70 ANTHROPOZOOLOGICA •2009•44 (1) TheProcess ofUngulateDomestication atÇayönü,SoutheasternTurkey
12 10 12 10
10 δ#" N‰ 12 10 12
8 14 PDB)
Y δ&%N‰ IR) (AIR) ( 8 14 PDB) 6 16 (A Y 16 ( N‰
C‰ 6 " 4 δ#$ C‰ 18 $ %N‰ δ# δ# 2 20 4 δ&'C‰ 18 'C‰ δ& 0 22 2 20 δ& Round Grill Channel Cobble Cell Large Room 0 22 Subphase Round Grill Channel Cobble Cell Large Room Subphase
F IG.6.—Carbon and nitrogen isotope ratio of cattle from Çayönü. Fig. 7.—Carbon and nitrogen isotope ratio of red deerfrom Çayönü. Slightlyyoungerkill-off patternduring theChan- nelled Building Subphasewasobserved, coinciding -19.29‰ (1.27‰variation) andin δ 15Nfrom withwhentheproportion of Cervus in theassem- 4.80to8.17‰(3.37‰variation),doesnotshow blagesuddenlyincreases,suggesting higherhunting anytrendsin lighterorheavierisotopevaluesover pressureon Cervus during thisperiod. time(Fig. 7)furthersuggesting thetrendseeninthe Bos isnotaclimate-drivenphenomenon. Secondly, thetrendsin isotopevaluescontradicteachotherif ANALYSIS OF CARBON interpreted in termsofclimatechange.Specifically, AND NITROGEN ISOTOPE RATIOS higher δ 15Nvalueshavebeenidentified asaresponse toincreasing aridity in bothplants (Schwarcz etal. Carbon andNitrogenisotoperatiosin humanand 1999) andanimals(SchoeningerandDeNiro1984; animalboneremainsfrom Çayönüwereanalyzed Gröcke etal .1997). Therefore, thetrendin Bos byoneoftheauthors (J.Pearson). Evaluation of mightsuggest decreasing aridity overtime.However, thephysiologyofplants identified from thesite the δ 13 Cvaluesbecomeheavierovertime, whichis showed thatall plants preserved ascharred material normallyindicativeof increasing aridity overtime areC3plants (Table3,see Appendix). (Tieszen1991). Asaresult,asimplemodelofcli- The Bos spp.results vary in δ 13 Cfrom -20.48 maticchangecannotexplain theobserved trend, to-17.2‰(3.28‰) andin δ 15Nfrom 6.31to andwethereforepostulatethatthesetrendsare 10.20‰(3.89‰ variation) withbothisotopes human-induced changesin diet. revealing atrendtowardslighter δ 15Nandheavier Since thelast aurochsbecameextinctin Europe δ 13 C(Fig. 6)overtime.Anumberofindividuals in the17 th century,wehavevery littleknowledge probablyhad access toC4plants starting in the about thehabitat,behavior,ordietofthisanimal, ChannelBuilding Subphasealthough whichplants whichhinders our interpretation. We assumethat mayhavebeenconsumed isunknownsince no C4 aurochswereadapted toforest environments,but plants havebeenidentified in thearchaeobotanical theremighthavebeenseveralsubspeciesofaurochs assemblage(VanZeist &deRoller1992). Agradual within Southwest Asia, wichwhereadapted todifferent trendin isotopevalues,asshownhere, mightbe environments (Uerpmann 1987). Generallyspeaking, indicativeofclimaticchange.However,thisseems morepositive δ 13 Cvaluesmayindicatemoreopen unlikelyfortworeasons.Firstly,iftheclimatewas vegetation anddrierenvironments (Tieszen1991). changing duetovariationsin precipitationand/or Thus,oneexplanation forthemorepositive δ 13 C temperaturethiswouldhaveknock-on effects on valuesin someofthe Bos bonesin theChannelled animalphysiologyacross all generaaswell ason Building Subphaseandlateristhatitindicatesthe plantcommunities(in termsofbothphysiology deterioration oftheaurochshabitat,perhapsbecause andtheappearance anddisappearance ofparticular oftheimpactoflong-termhumanoccupation and generaofplants). Anexamination ofthe Cervus deforestation. Itispossiblethat,priortodomesti- isotopevalues,whichvary in δ 13 Cfrom -20.56to cation,someaurochsbecameadapted tomoreopen
ANTHROPOZOOLOGICA •2009•44 (1) 71 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
areas(withdiffering δ 15Nvaluesunderpinned bya analyzed faunalremainsfrom Nevali Çoriandpoint separateplantbiomass)neartheedgeoftheforest out thatsmaller-sized specimensofsheep,goats and andtothesecondary vegetation created byhuman pigshavelower δ 15Nvaluesin thebonecollagen activities,whichlaidfoundation fordomestication. thanlarge-sized specimens.Asapossibleexplana- Red deermaynothavebeenasadaptiveasaurochs tion forthedifference in δ 15Nvalues,theysuggest andretreated withtheforest,resulting in theunchan- thatsmaller-sized animals(presumablyrepresenting ged δ 13 Cvaluesin theirbones.Thefactthatthe individualsunderculturalcontrol) mighthavebeen proportion of Cervus in theassemblagedecreased givenlegumesasfodderwhichcouldcauselower aftertheChannelledBuildingSubphasemayalso δ 15Nvalues.Thisassumption mightalsoexplain the suggest thatitbecameincreasinglydifficulttohunt lower δ 15Nvaluesin somecattlespecimensfrom the deerin thevicinity ofthesite. Channelled Building andlatersubphasesatÇayönü. Furtherevidence fordifferencesbetween Bos across thesubphasesofoccupation atthesiteisprovided T IMING OF DOMESTICATION through evaluation ofthenitrogenisotopevalues. OF FOUR LIVESTOCK ANIMALS AT Ç AYÖNÜ Drucker etal. (2003)hasdiscussed howmorenegative Changesin overall proportion,kill-off schedule, δ 13 Candmorepositive δ 15Nvaluescanbe theresult andisotoperatiosin thebonesof Bos occurred in ofcarbon andnitrogenrecirculation in organisms theChannelled Building Subphase, whichwere living on thefloorofdensecanopied woodland.In notobserved in Cervus .Asonly Bos wasdomesti- theearliest subphaseofoccupation atÇayönü(Round cated atsomepointduring thePPNB,itislikely Building Subphase)the Bos specimenshavethemost thatthesechangesareatleast partlyduetohuman negative δ 13 Candmost positive δ 15Nvalues.Over intervention on aurochspopulationsthateven- time δ 13 Cbecomesmorepositiveand δ 15Nvalues tuallyled tofull domestication. Based mainlyon arelower.Thus,weinterprettheÇayönü Bos isotope bodysize, Peters etal .(2005) suggested thatin the dataasindicating thegradualmovementofthese upperEuphratesregion Bos weredomesticated by animalsout ofcanopied woodlandandintoamore theMiddlePPNB.Helmer etal .(2005) proposed openenvironment.Thesecondinterpretation isthat Bos domestication atevenearlierdate, in theEarly wemightbe seeing aninflux ofanother Bos popula- PPNB(8800-8300 cal. BC),based on thesizeand tion from adifferentecologicalnicheunderpinned by degree ofsexualdimorphismofcattleatD’jade in plants withslightlydifferent δ 13 Cand δ 15Nvalues. northernSyria.AtÇayönü,ashifttowardearlier AtÇayönü,specimenswith δ 13 Cvaluesover-20 and slaughterscheduleandpossibleincreaseoffemale δ 15Nvaluesbelow8appearstarting in theChannelled Bos in thesamplesareindicated in thelateEarly Building Subphase.Since wehavealreadyidentified PPNB(Grill Building Subphase),although these sizediminution morphologicallyitistempting to observationsarehindered bysmall samplesize. suggest thatthedietary variationssupport theidea Moreclearchangesin othermarkers,bodysizeand ofchangesin therelationship specificallybetween isotoperatio ofbones,took place in thefollowing humansand Bos during theChannelled Building Channelled Building Subphase.Thus,theresults of Subphaseofoccupation atÇayönüTepesi,which analysisatÇayönücorroboratetheevidence from mayindicatetheappearance ofdomesticated Bos. othersitesin theregion. Bycombining multiple Noe-Nygaard etal. (2005) made anobservation on linesofevidence, wecanconclude thatdomestic aurochsin Denmarkthataurochsgenerallyshows cattlewerepresentalsointheupperTigrisregion morenegative δ 13 Cvaluesandmorepositive δ 15N probablybytheendofEarlyPPNB,andcertainly valuesthandomesticcattle.Ifweassumethese bythebeginning ofMiddlePPNB.Thisobservation observationsalsoapplyforAurochspopulationsin leadstotheconclusion thatthedomestication pro- Anatolia, theappearance of Bos withmorenegative cess ofall four importantlivestockanimalsbeganat δ 13 Cvaluesandmorepositive δ 15NvaluesatÇayönü Çayönüatabout thesametime, andprogressed in wouldalsosupport thehypothesisthatdomesticcattle parallel(Hongo etal. 2004,2005). Withthedata appeared during theChannelled Building Subphase. athand, however,wecannotexclude thepossibility AsforsoutheasternAnatolia, Lösch etal .(2006) thatdomesticcattlewereintroduced intotheupper
72 ANTHROPOZOOLOGICA •2009•44 (1) TheProcess ofUngulateDomestication atÇayönü,SoutheasternTurkey
Tigrisregion from elsewhere.Whetherornotthe becameearlierthrough time, bothsheepandgoats upperTigrisregion wasoneoftheprimary centers showed highersurvivalratein latersubphases, wherethedomestication process for Bos progressed especiallyin theLateandFinalPPNB(Hongo et locallyshouldbeinvestigated further.Ifsuccessful, al. 2005:figs5&6). geneticanalysismighthelp clarifythisissue. We havealreadyreported in detail about thesize andkill-off patternsofpigsfrom Çayönü(Hongo & SUMMARYAND CONCLUSION Meadow1998, 2000; Ervynck etal. 2001;Hongo etal. 2002;). Boththebodysizeandthelength Thedomestication process ofungulatesatÇayönü ofmandibularthirdmolarofpigsfrom Çayönü consisted oftwostages.Theinitialsignsincluded showagradualdiminution overtime, withsome theappearance ofasmall numberofsmaller-sized smallerindividualsstarting toappearasearlyasin pigsandasubtlechangein theslaughterschedule theGrill Building Subphase.Hunting ofwildpigs ofsheep,goats,cattle, andpigs(but notofred alsocontinued throughout thePrepottery Neolithic, deer)bytheendofEarlyPPNBorbeginning of resulting in increasing variability in thesizeofpigs theMiddlePPNB.Thiscoincideswiththeshift through time.Most ofthepig M3s from Prepottery in thesubsistence patterns,namelythedecreasein Neolithiclevelsarecomparablein sizetothatof thevariety ofhunted taxaandgradualincreasein modernwildpigs,but someteeththatfall in the caprines,beginning in theMiddlePPNB(around rangeofsizeoverlapforwildanddomesticpigs 8200 calibrated BC). Agrowing socialcomplexity at appearfrom thelateMiddlePPNB.Aclearshift thesiteisalsosuggested in theendofEarlyPPNB in thesizerangeofpost-cranialbonesofpigsdid andearlyMiddlePPNB,bytheappearance ofspe- nottakeplace until Cell Building Subphase, and cialized workshopareasandtheimportance given clearreduction ofthelengthofM3isobserved only tocommunalspacewithin thesite.However,the in thesamplesfrom Pottery Neolithic(Hongo & subsistence patternsbased on along tradition of Meadow1998, 2000; Hongo etal. 2004). Another sedentary hunter-gatherers in theregion still per- lineofevidence, kill-off patterns,showprogressively sisted.Whiletherewasagradualbut steadyincrease earlierslaughterschedulein latersubphases.The in sheepandgoats through timeatthesite, itseems trendsobserved in thesizeandkill-off patternsof thatanimalkeeping (andfarming) wasinitially pigsaregradual,but onedirectional,andwehave onlyanadditionaloption in abroad-spectrum concluded thatatleast small numberofdomestic subsistence strategy.Acontinuous andincreasin- pigsexisted atthesiteasearlyastheEarlyPPNB. glyintensiveexploitation ofwildresourcesbythe Thesizesofbothsheepandgoats in thePPNAand sedentary villagers,however,graduallyexhausted EarlyPPNBlevelsatÇayönüwererelativelylarge, theresourcesaroundthesiteoverthenext seve- suggesting thehunting ofawildpopulation. More ralhundred years.Thisprocess ofenvironmental small animalsappearbytheendofEarlyPPNBor deterioration isevidentin thesteadydecreaseofthe thebeginning ofMiddlePPNB.Aclearshiftin the relativeproportion ofmiscellaneous wildanimals sizedistribution towardsmallersizeisobserved in in thefaunalassemblage. theLate-FinalPPNBLargeRoom Subphase, which Asthesecondstage, adrasticshiftin thefaunal wasatleast partlyduetoanincreasein thenumber recordisobserved during theLatetoFinalPPNB, offemalesin themeasured assemblage(Hongo etal. characterized byasharpincreasein sheepandgoats 2005:figs3&4). Forthecaseofsheepandgoats, andbyheavy reliance on domesticanimals.The small samplesizesofthelate-fusing skeletalparts, traditionalsubsistence patterncollapsed andthe whichcause“rebounds” in thesurvivorship curves socio-economicbasisofthesitewasdrasticallytrans- atsubadultandadultagestagesmakestheanaly- formed.Aclearshifttowardsmaller-sized animalsis sisofkill-off patternsdifficult.We have, however, observed in all four taxa, including sheep,goats,pigs compared thesurvivorship ratesatthejuvenileage andcattle.Anincreasein femalesin themeasured stagein eachsubphaseandreported that,in con- assemblageisalsosuggested bythesizedistribution. trast topigsandcattlein whichslaughterschedule Clayfigurinesdepicting sheeporgoats arefound
ANTHROPOZOOLOGICA •2009•44 (1) 73 Hongo H., Pearson J., Öksüz B. & İlgezdi G.
only from the Late PPNB subphases, which also Jura (France). Palaeogeography, Palaeoclimatology, suggests a change in the relationship between these Palaeoecology 195: 375-388. animals and humans. As discussed elsewhere (Hongo E#*Q-Ö_Lta?\ A. 2007. — Çayönü, in Ö_Lta?\ M. & B?;_iAi\ N. (eds), Türkiye’de Neolitik Dönem. et al. 2004, 2005), this major shift in subsistence Arkeoloji ve Sanat Yayınları, Istanbul: 57-97. took place together with a fundamental change in E#PV\!= A., Dte\iV K., Ht\_t H. & Mi?LtR the social system. In other words, the shift from the R.H. 2002. — Born Free? New Evidence for the tradition of sedentary hunter-gathers to the socio- Status of Sus scrofa at Neolithic Çayönü Tepesi economic (and perhaps also psychological) system (Southeastern Anatolia). Paléorient 27(2): 47-73. of agro-pastoralists, which laid the foundation for G#*_4t\ C. 1989. – Size and Sex: Evidence for the Domestication of Cattle in the Near East, in the Pottery Neolithic tradition, occurred in parallel M*AAi4 A., W*AA*?Q4 D. U G?#L\i# N. (eds), The with the major shift in subsistence. Beginnings of Agriculture . BAR International Series As for cattle the domestication process for Bos began 496. Archaeopress, Oxford : 77-109. by the end of Early the PPNB at about the same G#{!=i D. R., Bt!yi#i\4 H. & M?#*tbb* A. 1997. time as for the other important livestock species and — Annual rainfall and nitrogen-isotope correlation progressed in a parallel manner. Although initial signs in macropod collagen: application as a palaeopre- cipitation indicator. Earth and Planetary Science of domestication in different aspects of the faunal Letters 153: 279-285. record are usually subtle, they can be detected by HiAQi# D. 1988. — Les animaux de Cafer et des sites combining more than one marker to examine the précéramiques due Sud-Est de la Turquie: Essai de timing and trend of changes over time. Comparing Synthèse . Anatolica 15: 37-48. prodomestic species and other wild taxa has also proved HiAQi# D., Gt/#*!yt\ L., Mt\!ytb H., Pibi#4 J. to be a useful tool in detecting human-induced shifts & S?p? Si_/* M. 2005. — Identifying early domestic cattle from Pre-Pottery Neolithic sites on in exploitation patterns that lead to domestication. the Middle Euphrates using sexual dimorphism, in Pibi#4 J. & HiAQi# D. (eds), The First Steps of Animal Domestication. Oxbow Books, Oxford: REFERENCES 86-95. Hi44i B. 1978. — Evidence for husbandry from the A#e/!=Ai B.S. & Ö_=?V? V. 2006. — Animal exploita- early Neolithic sites of Ganj Dareh in Western Iran. tion at Körtik Tepe: An early Aceramic Neolithic site PhD Dissertation. Columbia University, New York. in southeastern Turkey. Paléorient 32(2): 113-136. Ht\_t H. & Mi?LtR R.H. 1998. — Pig Exploi- Bg-?=-g E. 1998. — An essay on the chronology of tation at Neolithic Çayönü Tepesi (Southeastern the PrePottery Neolithic settlements of the Tau- Anatolia), in NiA4t\ S.M. (ed.), Ancestors for the rus Region (Turkey) with the building remains Pigs: Pigs in Prehistory . MASCA Research Papers in and 14C dates, in A#4ie5= G., MiAA*\= M.J. & Science and Archaeology 15. University of Pennsyl- S!y*#Qi# W. (eds), Light on Top of the Black Hill. vania, Museum of Archaeology and Anthropology, Ege Yayınları, Istanbul: 137-150. Philadelphia: 77-98. B/*bi\y/*4 H. 1985. — Preliminary report on the Ht\_t H. & Mi?LtR R.H. 2000. — Faunal faunal remains of Hayaz Höyük from the 1979- remains from Prepottery Neolithic levels at Çayönü, 1983 seasons. Anatolica 12: 62-74. southeastern Turkey: A Preliminary Report Focusing Di_i#e?A M. 1970. — I. Zoological Part, in Di_i#e?A on Pigs ( Sus sp.), in B/*bi\y/*4 H., M?4y=t/# M. M. & F#iL4=*AL B. (eds), The Urus (Bos primige- & CytV=i A.L. (eds), Archaeozoology of the Near nius Bojanus) and Neolithic domesticated cattle (Bos East IVA . ARC-Publications, Groningen: 121-140. taurus domesticus Linné) in Denmark . Det Kongelige Ht\_t H., Mi?LtR R.H., Ö=4/_ B., İA_i_L* G. Danske Videnskabernes Selskab, Biologiske Skrifter 2002. — The process of ungulate domestication in 17(1). Munskgaard, København: 5-178. Prepottery Neolithic Çayönü, southeastern Turkey, D#*i4!y A. Pt\ Li\ U Pibi#4 J. 1999. — Vorläu- in AA-Sy*V?e A.H., CytV=i A.M., & B/*bi\y/*4 figer bericht über die archäozoologischen Untersu- H. (eds), Archaeozoology of the Near East V .ARC- chungen am Göbekli Tepe und am Gürcütepe bei Publication, Groningen: 153-165. Urfa, Turkei. Istanbuler Mitteilungen 49: 23-39. Ht\_t H., Mi?LtR R.H., Ö=4/_ B., İA_i_L* D#/!=i# D., Bt!yi#i\4 H., B#*L?/Ab A. & B*AA*t/ G. 2004. — Animal Exploitation at Çayönü D. 2003. — Carbon and nitrogen isotopic compo- Tepesi,Southeastern Anatolia, in Featuring Com- sition of red deer ( Cervus elaphus ) collagen as a tool plex Societies in Prehistory: Studies in Memoriam for tracking palaeoenvironmental change during the of the Braidwoods. TÜBA-AR [ Turkish Academy of Late-Glacial and Early Holocene in the northern Sciences Journal of Archaeology ] 7:107-119.
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WlG#0}L.&DlppzcG#M.J.1992onwards.— HkpwkxD.(eds), TheFirstSteps ofAnimalDomes- TheFamiliesofFloweringPlants:Descriptions,Illus- tication.OxbowBooks,Oxford: 125-146. trations,Identification,andInformation Retrieval. ZkWkxM.A.2006.—Acriticalassessmentofmarker Version:19thAugust 1999.
Submitted on 8February2008; accepted on 24October2008.
76 ANTHROPOZOOLOGICA •2009•44 (1) TheProcess ofUngulateDomestication atÇayönü,SoutheasternTurkey
APPENDIX Table 1. –Archaeobotanicallist identified atÇayönüTepesi(aftervanZeist &de Roller1992).
Keytophotosyntheticpathwayreferencecolumn 1:Watson and Dallwitz 1992onwards 2:Sage and Monson 1999 3:pers.comm.
FamilyGenera/SpeciesCommon Photosynthesis REF Ranunculaceae Adonis sp. pheasant’seyeC3(all species)1 Poaceae Agrostis type bentgrass C3(all species)1 Rosaceae Amygdalus sp. wild plum/almond C3(all species)1 Primulaceae Anagallis sp. scarletpimpernel C3(all species)1 BoraginaceaeAnchusasp. bugloss C3(all species)2 Leguminosae/Fabaceae Astralagus sp. milk vetchC3(all species)1 Poaceae Bromus sp. brome grass C3(all species)1 Ulmaceae Celtis sp. Hackberry C3(all species)1 Euphorbiaceae Chrozophora sp. Dyer’scroton? unknownx Fabaceae Cicer sp. chickpeaC3(all species)1 Poaceae/Graminae Cynodon sp. bermudagrass unknownx Lilicaeae Echinaria sp. ?unknownx Moraceae Ficus sp. fig C3(all species)1 Fumariaceae Fumaria sp. fumitory NoC4recorded forfamily2 Rubiaceae Galium sp. bedstraw/cleavers C3/CAM, Galium =C31 Cistaceae Helianthemum sp. Frost weed/RockroseC3(all species)1 PoaceaeHordeumspontaneum2rowhulled C3(all Hordeum sp.) 1 Leguminosae/FabaceaeLathyrus cicera/sativus vetchling C3(all species)1 Fabaceae Lens sp. Lentil C3(all Lens sp.) 1 Linaceae Linum sp. flaxC3(all species)1 BoraginaceaeLithospermumtenuiflorumgromwell? C3(all Lithospermum sp.) 1 PoaceaeLoliumrigidum/perenne ryegrass C3(all species)1 Solanaceae Lycium type honeythorn? unknownx Malvaceae Malva sp. mallowC3(all species1 Fabaceae Medicago sp. medickC3(all species)1 Fabaceae Melilotus sp. cloverunknownx Poaceae Phalaris sp. canary grass C3(all species)1 Anacardinaceae Pistacia sp. Pistachio C3(all species)1 FabaceaePisumpeaC3(all species)1 Plantaginaceae Plantago lagopus type Plantain/Ribwort C3(all species)1 Polygonaceae Polygonum sp. knotweed C3(all species)1 Fagaceae Quercus sp. oakC3(all species)1 Ranunculaceae Ranunculus spp. Buttercupgenus C3(all species)1 CyperaceaeScirpus maritimus seaclub-rushC3(all species)3 CaryophyllaceaeSilene campion unknownx Poaceae/Graminae Stipa spp. Needle grass C31 Thymelaeaceae Thymelaea sp. thymelaeaC3(all species)1
ANTHROPOZOOLOGICA •2009•44 (1) 77 Hongo H.,Pearson J.,Öksüz B.&İlgezdiG.
FamilyGenera/SpeciesCommon Photosynthesis REF Fabaceae Trigonella sp. trigonel C3(all species)1 PoaceaeTriticumaestivum/durumbread/durumwheatC3(all species)1 PoaceaeTriticumboeoticumwild einkornC3(all species)1 PoaceaeTriticumdicoccumemmerC3(all species)1 PoaceaeTriticumdicoccoideswild emmerC3(all species)1 PoaceaeTriticummonoccumeinkornC3(all species)1 PoaceaeTriticumspp. wheatspeciesC3(all species)1 CaryophyllaceaeVacarriaCowbasil/cowherbC3(all species)2 ScrophulariaceaeVerbascumMullein? C3(all species)2 Verbenaceae Verbena spp. vervain C31 Fabaceae Vicia cf. ervilla bittervetchC3(all species)1 Vitidaceae Vitis spp. grape C3(all species)1 Lamiaceae/LabiataeZiziphoraMintfamilyNon C4anatomy1
78 ANTHROPOZOOLOGICA •2009•44 (1)