Cour. Forsch.- Inst. SCllckenberg I 259 I 99 - 110 I -I Figs I Frankfun a. M .. 13. 12. 200 7

Biostra tigraphic succession of the Early and Middle Pleistoccnc mammal faun as ofthe Atapuel'ca cave sites (Burgos, Spain)

With 4 figs

Gloria CUENCA-BESC6s & Nuria G AKCiA

Abs t r:lc\

The At3pu~rc3 C:l\~ 1000:l1il;~S {GrJn Doliua - TD. Trinchcr.l Galeri~ - TG. Trill~hcm flef.1ntc - TE. Sim3 de los Huc;;os - SH) rcprcS<.'nt onc of thc beSI records Of lhc Europe,1n Plcistoccnc f.1un'l. nlc siles are located in long and str;lIigrJphically "ell·documented S<:\;lions. ranging from the Early to lhe I\ l iddle PleiSlocenc. They yielded ,In imprcssh e amount of fossils but also of lilhi~ industries. "Iuch provide an insight ;nto Man's earliest records and acti, ities 111 Westwl Europe Icannibalism. earliest burial. food pro..:cssing). The SlrJtigrJphic distribution ofthl' Alapul:rca n1ll11unals rc"eals 6 difli.·renl local faunal assembkLg~"S. "hkh arc here dl'Signated the At:.pucrca 1:llIn<l1 unils lATA FU). ddinl-d by lhe co-occurrence of p.1r1i~ukLr manullalian t'IX3. 1\ tol1:u' ·er. the Sierra de At:lpuerca Sill"S arc well knOWIl for Ih"ir Early to Middle PleistOC" ne hominid· bearing localities. Le' cl 6 oflhe Gmn Dolill3 sil(." has yiclded Ihe oldcst lossil hominid in WeSlcm Europe. IIrmw (/1II."('(."s.OI·. daled 1l.1ial"Om:lgnelicall)". mdiolllelrically as '1dl as bioSlraligrJphi""Uy 10 around IWO kJ BY. This 1c,"c1 TD6 characlerizes Ihe ATA FU 4. Thc SicTrJ de Alapu~rc" also yield_,> the best collection of MiJdk I'icisloccnc fossil hominid remains, d~s igl1al~d IlfJnlQ /re;ddbe'"gt'I/.I'i.\", Iro", Sim3 d" los 1-1l1e;;os 3nd Trinchem G31~ria. nle Sima de los Huesos has ,m ag" 01"53U b 13.1'. SII ,md TG arc ref(."rr~-d 10 ATA FU 6. Nl1merous fossils from other AlapUtn:a siles are excellenlly preserved. Large cami,.on:s and herbh-orcs. smalll11ammals. as well "s birds. liards. Hshc~. and rrogs :Ifi; fOll nd in :tlmost c,-cry Ie,-d . To dale. a tuwl of I U3 la."1 belonging 10 9 OT<krs. 27 f3milics ,md 61 genera 111\' I\~of(kd lrom Ihe Awpucrca ca"e loc:llities.

Key w ord s: lliOSlr:tligrJphy, Mammal;a. Early :.nd Middle Pkistoccne. t\t;lpuen:a. Sp.,in. Europe

Introduct ion

The last d ecade ofrese:lreh on Atapllcrca H ill (Sierra de

Alapllerca. Burgos. North-west Spai n) has contributed

sign ificantly to our underslanding of human evoluti on

in Europe during the Early and Middle Pleistoccne. The

earliest Western European foss il human remains were

found in 1994 in Gran O o lina (TO ) levcl 6 (also na med

Trinchera O o lina in somc papers) tCARIlONELL ct al. 1995.

1999. B EKMimEz DE CASTKO et a l. 1997a. b. 1999). Gran

Dolina level 6 (T06) has yie lded an illlponant collection

of Homo allfece.\·SQI" remains (BEKMUDEZ 1)1, C ASTRO et al.

I 9971t) and lithic anef3cts (CARBONELL et al. 2001). Th is

layer lies just llrlder the MatuyamalBrllnhes palaeomag­

netic boundary and is therefore older than 780 ka B.P_

(PAKI:!; & Pl, KEZ G ON7.AL£l 1995. 1999. P tREZ-GOl'zALfL

ct al. 1995.2001). This locali ty has al so produced abun-

dant micro- and m acrom alTlll1al remains (CUENcA-Bm;c6s

e t al. 20 01. G AKciA & AII.SUAGA 20 0 1a. b, V .... N DIOR M ADE

200 1). Human presence has a lso been revea led in other

levels of the sequence (T04, TD5. TOlO :lnd TOI I) by

s tone tools as we ll as by human-modified ungulate re­

m ains (C,\IUlONELl e l al. 2001 ).

Sima de los I-I uesos (S I-I ) and Trinc hem Ga1crb (TG )

also yielded the best co llect ion of Midd le Pl eislocene

Homo heide/bergel/Sis rell1:1ins ever ulleanhed. Given the

importance of these palaeontological and archaeological

discoveries, the Atapuerca learn has foc used in paniclllar

on the geology. stratigraphy. geomorpho logy. magne­

lostrati graphy. radi ometric ages and b iostraligraphy of

these sites (AII.SUAGA et al. 1993. 1997. 1 999a. b. CAKBON­

ELL et a1. 1995. CUENCA-B ESCOS et al. 1995. 1997. 1999a.

b. 200 1.2005. PARES & PEREZ G ONz.,\LEZ 1995. 1999. PE­

KEZ G ONzALEL et al. 1995.2001. R OSAS et al. 1999.2001.

AUlhurs' mldrc~scs: G. CuE"<c ... ·I3L'iC'". Arc:. de 1'"kt)111010gia del Dcpan3m~nlO de Cienc;:.s ,ok b Ti,·rra. Facuit:,d de Ciendas. Uni\'ersidad de i.Jr:'l!OlJ'. 500()<) Zar:lgOZll. Spain. <cucnc:.gw U11l1ar.cs>: Dp\. 01" Anthropology, UNM. Albuqucrquc. l\7131 NM. USA: N. G"~(l". Dp\. I'alcon· lologia. Uni\'c~id:,,1 COl11phnens.: de Mlldrid. F. C. Gcologicas. Ciudad Unhcrsilar;a sfn. 2S()40 l\1adrid. Spain: CcnlTO (Uc:\t·ISCIII) de Evolu· cion ). C0f11ponamiCl110 Ilumanos. Cl Sincslo Ddgado 4. Pabcllon 11. 18019 Madrid. Spain. <ngJrci~ 1Z! isciii .e$>; Ilul11all E\Olul lon Rc.stJrch Cenler. Muscum of Vcnebmh: Zoology. Uni\ersilY ofC:,litOmia. 3 I 01 Valley Life Scicnc,·s Building. Bt."rkelcy. C;.lifomi:l 94720-3 160. USA

, E. Sch"el7.crb.1n",che Ver1ag~buehhandlllng (Nag<,,1e u. Olx·rmilkr). 2007. ISSN 0341-111 tl

CUENo.-l3rscos & GM(cLoI.: Mammal faunas of the Atapuerca cave sites ! Spain)

Duero BaSin

Fig. 1: Th<.' Al~pu<.'r<.'a si tuation.

FRANCE

2004. BI SCHOFf et al. 1997. GARc lA et al. 1997. G ,\RciA & ARSUAGA 1998. 1999.2001 a. b, AGU IRRIO 1999. 2004. FAL­GUER£S et al. 1999, 2001. V,\N OER MAOE 1999a. b. 200 I. LMLANA & CUENCA-BEsc6s 2000, L6p£z Am ONANZAS & CUENCA-BESC6s 2002. GARCiA 2003. LAPlANA CONESA et al. 2004. C UENCA-BEsc6s & ROI' ES 2007).

Th e mammalian ta.>;a represented at the Atapuerca cave locali ties belong to 9 diOe rent orders: Eulipotyphla (Insectivores). Chiroptera. Primates. Carnivora. Probos­cidea. Perissodactyla. Artiodactyla. Rodentia. and Lago­morpha. The aim of this work is to update and analyse the biostratigraphic distribution of mammalian faunas in the Atapuerca sites and to define the Atapuerca Faunal Units: ATA FU.

The Sierra de A tapuerca Pleistocene sites and their Faunas

The Sicrra de Atapuerca is a small prominence in the otherwise 113t Mioeene valley of Castill a. near Burgos city in North-west Spain (fig. I). The Sierra is composed of Upper Cretaceous limestone. which started to be af­jected by karSlic dissolution at the end of the Miocene. An abandoned railway (Trinchera del Ferrocarri l) cuts through the Sierra de Atapuerca. e;"posing cave depos­its at both si des [Trinchera Penal (TP). Gran Dolina

100

Trlllchcru Dolona * r.;;,,,,,,¥' l ,,"cllem Z~'lI":ros _T""d,~," G:otcrin*

c"".·. dol''''''"'' romp'''''''-

The Atapuerco sites

'lt l~ClI£RA

Dlil. Trinchera del Elethnte FEIIM()CA RRII • .I

~"'<"'..,"'" "'c"'.· •. , .. ) ....

-- S,ma dc !os J Jut'SOs *

Fig. 2: The sites at the Atapuerca railway cutting.

(TD). Trinchera Galeria cave comp lcx (TG). including Trinchera Zarpazos (TZ) and Trinchera Norte (TN). and Trinchera del Ele lilnte (TE): fig. 2]. The karstic caves and ga llcries. which are of phre3tic+vadose origin. con­tain sediments dated between at least 1.5 Ma and ca. 200 ka tFALGU I~Ht,S et a1. 2001). w hi ch represent excellent examples of well-dated. long and compl ete European continental Pleistocene sections. Their geology is well­known thanks to PARr-oS & Pt REZ Gml7..At.Ill. (1995. 1999) and PERIiZ GON7.ALEl. et al. (2001).

The cave sed imen ts yielded a rich record of Pleis­toccne fau na and lithics. associated with a large amount of allochthonous materials. Atapuerca has also some ac­tive caves, such as Cueva Mayor-Sima de los Huesos and Cueva del Silo cave complex (see ARSUAGA et al. 1997). Within the Cueva Mayor complex the "Trinchera del Fer­rocarril" sites have been distinguished from the "Sima de los Huesos" site. The Trinchera del Ferrocarril sites TD and TG have yielded late Early to Middle Pleistocene hu­man and fauna) remains. as well as lithic industries. The TG complex and TO are not connected with the Cueva Mayor-Cueva del Silo caves. and probably represent separate karst systems (fig. 2). The Trinchera del Elefilntc site belongs to the Cueva Mayor cave system and out­crops at the "Trinchera" railway corridor.

The fossils from these sites arc very well preserved and numerous. Carnivores and herbivores. but also small vertebrates such as bats. rodents. shrews. rabbits. birds. lizards. fish. and frogs arc found in almost every level of the cave-filli ngs (AGU tRR E 1999.2004. CUI,NCA-D ESCOS ct a1. 1999:1. b. 2001. GARC iA & ARSUAGA 1999. 2001a. b. VAN DIOR MADE 1999a. b 2001. L61'EZ ANTONAN7..AS & CUENCA-B ESC6s 2002. CUE!'JCA-BESCOS 2003a. b. GARCiA

2(03). T 0 5, for instance. provided as much as lOO micro­mammal mandibles in less Ihan 100 kgs o f sediment. and in the upper T O leve ls (TOI O. 11) nearly a thousand firs t mo lars o f Te,.ricola were collected from o nc less than 500kg o f sediment.

T hl' Trinch{'ra d cl EI{'fa nlc sil {'

The TE is an ancient opening 10 the Cueva Mayor karst subsystem. T he s tratigmphie section o fTE is 25 III thi ck : about 19 III o f the cave infill contains all ochthonous rub­ble from the construction of the ra ilway. Six add itional meters of sediment were discovered excavat ing under the current fie ld level during the first years of sampl ing in the trench (II.OSAS et al. 2004).

The section of TE incl udes Lower and Middle Pleis­tocene layers (ROSAS et al. 2001 .2004. CU£NcA -I3I,sn'ls & ROFl:s CI I"VEZ 2004 ). The site comprises 2 1 stratigraphic Icvels whieh have been grouped up into the fo llowi ng three sedime ntary phases: Phase 1. represen ted by the Lowcr Red Un it (T ELR U); Phase 11. which [cd to the deposition of the Upper Red Vn it (TE VRU): and Phase [[1. when Ihe uppemlOst infill accumu lated (the distribu­tion o f the T E mammals is shown in figs 3 and 4):

Phase J o f TE: Because the TE LR U levcls TE8 to TE I4 show an inverse polarity. the lower part o f the TE succession is referred 10 the Mat uyama Chron (ROSAS et aJ. 200 1). TE LR U is rich in faunal remains: birds. rep­tiles. as well as mammals. T he latter include: (I ) ElI lipo­typhla (Insectivores) - Erillace/ls c r. elllupaell.\·, Ta/pa ct: elllupat!(I. Oesmani nae illdet. . SOI¥!X sp., Oert!melld ia ji.1'lidell.\". AsoriCII/US gibbemdoll . Cmcidlll'(/ a fT. kOI"ll­fe/di. and Crocid/lra cf. I1ISSII/(I (CUENCA-BESC6s & ROfF.s 200 .. k ROH'S & C UE"{CA-BESCOS 2006): (2) Carnivora ­C(lIIil' cr. {/mellsis/mol·bochellsis. Vu/pes c f. %pecoide.,·, Urm,,' c f. dolillellsis, e r. O(ll'(lIIogale amiqlla, M IISfe/O c r. palerlll illeo/praellil!a/is, Pl/III /Ullicfis ef. I/el'fii (GARcIA & HOWFLL in press). Pallfltel'lI gOlllblls=oege/lsis, Lyllx cf. issiQ(lo/'el/si.~ ; (3) Perissodactyla - Sleplwllorhilllls ell'us­ellS, and Eqllidae indet.: (4) Art iodactyla - HiPJXJPotallllls sp .. Suidae indet.. Cervidae indet. , £uclodocerus gilllii/ Mega/oceIVl' S{willi. Bodt/(le indet.. 8 isoll sp.; (5) Roden­tia - Scilll'lIS c f. II'lwrtae, Eliomys ljllercilllls. CmlOrjiber. UlIga/"olll.l:I 1I(lIIIIS. Pliomys cf. l'implicior. IIIf0plwiolll.\'l· {C/I"ol'llli. A. bllrgolldiae. A. IlII liellsi.~ (formerly described as /bel"Omys aIT. IlIIescm¥!lIsis). Arvicolidae novo gen. novo sp .. C{lslillomys rh'as, 1I,)Ode1l1115 sp. (LAI'LA NA & CUF.JI;.CA-Br:scOs 2000) and (6) Lagomorpha - Leplls sp. :md Oryclo/aglls sp. Other venebrates: (7) Aves - Car­dllelis ch/oris. Fa/co sp .. Pertli.\' paleoperdix, CoIl/mix COIl/mix. Cul llmbo /il·;a. /-{(I/i(leellls a/b;cill(l, Ci"c(lell/s galf;cl/s, Fa/co fi IlIllIC/I/IIS, TII,.(/us sp .. Alaudid3e indel.. and AI1IIS sp.: (8) fi shes. amphibians and reptiles - Sa lmo­nidac inde!.. Bllfo bufo. B. c{l/lIIl1illl. and Chelonia indet. (3. 4 and 6 to 8 after ROSAS et al. 2004, 5 by LAPI..ANA & CUE"{CA-B[scOs 2000. CUENCA-BESCOS & Ron.s CUAV£$

Cour, FOJ'1)ch.- Insl. St'nckenbcrg, 259. 2007

2004. and 2 by GAlI.ciA). In previous studies the faunal assemblage of the TE Lower Red Unit is shown to be the oldest at Atapuerca ( LMLANA & Cun<cA-BESCOS 2000 . ROSAS et al. 200-L ROfES & CUFNCA-B ESCOS 2006), In partic ular. the AllupiJaiolllys ( Rode ntia) associa1i on is sim il ar to that recorded in other South-Europea n local i­ties. as Les Valerots (France), Bagur 2 (Spain). Pietrafitla (Italy,. Monte Peglia ( Italy) or Le Vallonnet (France). which makes T ELRU older than the deepest exposcd lev­els o f Gran Oolina (T03 to TD6. ca. 0.78-0.9 Ma, . The arvicolid and illsectivore assemblage shows 1hatth is unit may be at least as o ld as the localities o f Fuente Nueva 3 ( FN3) and Barranco Leon (BL ) in lhc G ranada basin (ca. 1.2 \0 1.5 Ma: AGusrl t llf!. 200 .... ROFES & CUf ..... CA­BESCOs 20(4).

The carnivore assemblage fro m T ELR U is :llso very ric h and referable to the Epivillafranchian. as indicatcd by Pallllollicli.\" (GARCiA & HOWI' Ll.. in press) which occurs at a number of Pli o-Pk' istocene sites in Eurasia : Casa Sgherri ( Italy) (MARCOUNl Cl al. 2000). Upper Va ldarno (Italy) (FICAlI.t'l..l..l & TOII.KE 1967. ROOK 1995), Vill:iny­Kalkberg (= Vi llany 3). Csam6ta I. Beremend (Hungary) (J;'NOSS ~' 1986), Khapry ( Russia) and Etul ia (Mo ldova) (SOTNIKOVA el a!. 2002). j ust to quotc somc. Palll/uniclil' is recovered in some late Early Pleistocene s ites like Pietrafi Ua (ROOK 1995): it d isappeared around the Jar..l­mi llo Subchron. There arc at least four other European loca li ties with fa unas similar to those of thc T E LR U. also assoc iated with li thic ane facts: Fuente Nueva 3 and Barranco Le6n in Guadix- Baza. Soulh Spain, Le Va llon­net in Francc and Pirro Nord in Ital y ( Lu~u.EY et 31. 1988. AGUSTi et OIL 2004 . ROSAS et 31. 200 .. 1-, MOULt.El e t 31. 2006. ARJ..A R£LLO el 31. 2007).

Ph3SC 11 o fTE: the T EU RU comprises levels T EI5 to TE I9. Level TE I9 conta ins a small number of rodents which can be o f Middle Pleistocene age b.1sed on si mi­larit ies with those from the levels T OIO and TDII of TD. as well as fro m TG and SI--I. Among the carni vores from T E 19. some (i. c .. Cmcllfll CIVCllflI and I'lI lpel' 1'/Ilpel') are ei ther Midd le or Latc Pleistocene in age. [n cont ras t. the bear rema ins present cave-bear lineage traits. which arc not as derived as those of the Late Plcistocene Urslls spell/ellS. r • .lI her as those of the Midd le Pl cistQCene U. del/ illgeri (RosAS et at. 2004). Stone tools recovered in TEI 9 indica te a latest Middle Pleistoeene age fo r thi s Unit ( ROSAS et at. 2004). However. because no T:Jdi01llet­ric dat ing corroborates this yet and because o f the scarcity ort he sma ll mammal record. the age attributed to T E I9 is st ill tcntativc. The herbivore associat ion incl udes only one taxon of biostratigraph ic interest. Sft!IJ/UlI1odl;III1S iJemiloecllll.,· (ROSAS et a l. 2004 ) that gi ves us a Midd[e to ea1'ly Late Pleis toeene age and which is also present in the Trinchcra Galeria un its TG Il and TGIII (ROSAS et al. 1998).

Phase III ofTE: the third phase comprises lhe upper­most infill sediments. namely the levels TE20 and T E2 1 (RoSAS et al. 2004).

10 1

CUl "CA-IlL\COS & G Ait" ' ... : Mammal faunas of the Alapucrea cave sites (Spain)

The TELRU is providing essential information on human activi ty in Europe belween approximately 1.1 to 1.5 Ma . This chronological all ribul ion is supponed by Ihe li lh ic tools from TE LR U (ROSAS et a l. 2001. 200.t). as well as by Ihose fro m the other European localit ies men tioned above (AGUSTi et a1. 2004. MouLL!. Cl a1. 2006. AItLAIIHLO cl <11. 2007). The T ELRU and Ihe Gua­dix-Ba7..<1 lilhic tools. correspond tcchnologica lly to thc Oldowan Industry or Mode I. [n addition. TELRU has yielded severa l large herbivore bones with CUI-marks. which indicate that hu man agents played a role in lorm­ing this n13m malian acculllul;llion. Moreover. the rich and d iverse TELRU venebrate assemblage is character­ized by taxa typical of waml and moist wooded areas and open lands.

The G ran I)olin:\ Sile

Alapuerca's longest sequence is the Gran Oolina lTO) site.:lII appro.-.:imately 19 m cave infilling that comprises 11 str:lIigraphic levels. Almost all oflhem (T03- TOI I) are rich in fossi l fa una and anefacts of Early 10 Middle Pleistocene age (C.AIWONELL et 31. 1995. 1999. GAltciA & AItSUA GA 2001 a. b. GAltciA 2003. VAN OER MAOE 200 [. LOI'I;.l ANTONANZAS & CUENCA-BESC6s 2002. CU£NCA-BI:­scos et al. 2005). ESR dating :md Uranium series dates place Ihe Gr:.m Dolina fossiliferous levels between about 300 and SOO ka (FALGUER[S Cl al. 1999.200 1). These re­sui ts are consiSlent with the results o r the palaeomagnelic studics donc by PARES & PI:ItEZ GONzAuiZ (1995. [999). which correlate the Gran Oo[ina [eve r 1'5 1l0 m 13 [ polarity wilh the Jar:1.I11ill0 Subc hron. The Maluya ma·Brun hes reversal occurs in T07 and therefore thc fossil irerOlls levels T03 to T0 6 correspond to the younger pan of the Matuyama Chron. between the Jaram ill0 and Brunhes normal palacomagnclic zones. The level T06. which contains the HOlllo (lllI eces.WJI' re mai ns (T06b in figs 3 . .t). lies under T07. and is thus older Ihan 0.7S Ma. Near the top of T0 6 there arc 15 cm of a brownish·rcd clay (Aurora stratum) which have yielded, in less than 9 square meters excavated between Ihe 1994 and 1995 fie ld seasons. lilhic aneracls. abundant raunal remains but also more Ihan SO fossils of Homo (ll/Iece~':iOI' (CARIIONI'_LL et al. 1995). By its fossi l content leve l T06 can be divided inlO T0 6a and T0 6b (the latter comprises Ihe Aurora level). Also the le\'el T0 8 is subdivided inlo the levels TOSa and T Ogb and ranges from 0.7 to 0.5 Ma . Levels TOIO- 11 arc dated from.tOO to 300 ka (FALGU[R);S et al. 1999).

UH'I:Z ANlONAN7.J\S & CUENCA-BESC6s (2002). BLA IN (2005). ,md CUENCA- BES<'6s et al. (2005) ana lysed the d istr ibution of small vcnebrates in the Gran Oolina's 11 stratigraphical levels. presenting some pal:teocn viron­mental interpretations. Despite some connict between the resul ts obtained tram the analysis of the herpctofauna and the IlHulHlla[ian fauna. in general a continental. dry and

102

cold climate is inrerred at the beginning oflhe succession (T0 3 to lower T05). The fa una in the upper pan ofle\'cI T0 5 and in T0 6 renects a complex interglacial period wilh nuctualions in the degree of re [at ive humidity. The species Microtus afl mfficepoides appears in T0 7 and in the lower pan of T0 8 (T08a). indicat ing a relalively wet period. Data from upper levels (TOSb. TOI O. TOi l ) re­Aeet a change in Ihe p.1laeocnvironmental conditions. and a dry period with slight osci llations in relalivc moislure appears to have takcn place (CUENCA-Br:scos cl al. 2005). The TO [evels arc also corre lated wit h oxygcn isotope stagcs (M IS): TO]. T04. and T 05 with M[S 22. and T0 6 with M[S 21. 10wer T DS (TOSa) is corrclated with MIS IS. upper TOS (TOSb) is correlated with M [5 13 or 15. and fina lly TOI O and TOll are correlated with MIS 9 or 11. The succession of fau na[ assemblages at T O reflects landscape and environmental changes over the past mil­[ion years (CUENCA-BI~c6s et al. 2005). Shins belween wood land. open land and moorland are indicated by the relativc abundance of small lll:llllmal species (in terms of Minimal Number of Individuals). Using its fauna l distri­bution. we divide the Gran Oolina site into five differenl mammal un its (unils I to 5 of TO) indicated in fig . 3 as Alapuerca's faunalunils 210 6.

Thc Trinchcn t G a lcri:t sit e

The Trinchera Galeria (TG) complex is another group of cavilies and pils located at Trinchera del Ferrocarril. TG is fi lled up wit h sedi ments conlaining abundant fauna I and archaeological remai ns (ROSAS et a1. 1998. [999. OR­lEG" 1999) and comprises Ihree areas: Trinchera Galeria. Trinchera None and Cueva de los Zarpazos (TG. TN. TZ). Five phases of deposition of clastic sediments havc been recognized in the silc (G l to GV by PI~KEZ-GON7.A LI::Z

el a1. 1995) and as many as twelve "human li ving floors" in the G II[ phase. Many stone tools (q uartzi te pebbles, possibly uscd to brcak boncs. and l1int tools) :lIld her­bivore bones with cut marks in the lower levels indicate an earlier. recurrent human occupalion of the cavc. Man used the cave for shon periods of time. with occasional consumption of large mammal s tCAIUJONELL et a1. 1999. Dh:l. el al. 1999). A frag ment of a human mandiblc was recovered in 1976 and years later. in the 1995 fie ld sea­son. a human parietal bone was found associated wilh a handaxe and other Achculian tools (ORTtiGA 1999). Those human remains were assigned to HOlllo heidelbergellsis.

Ihe same species recorded at Ihe Sima de los "iucsos (AR­SUAGA et al. I 999a). The chronology of the TG deposil (G I[ and Gill fossiliferous uni ts) ranges between 350 10 200 ka (CAKIlONELL el al. 1995. 1999. GAll-cIA & AItSUAG" 1998. ROSAS cl al. 1998. 1999. CUF"'CA - B I~c6s et al. I 999a. GAll-CiA 2003).

The rodents from the TG complex sites (Units GII and G ill) are characteristic of the Middle Pleistoccne : Manllota sp .. £Iiomy.\· qUel'cillll.\'. f~l':>· t"ix (AC(llIfltioll)

I'il/ogradOl'i. Allocricefl/.\· corre=emis, PfioIllY.\· lenki. Arl'ico lo arr. sapidlls. Microlllsj(lllsoni. Micro/lis w ·· \"{I/is. TI.',./"icola lI/opllel./llellsis. IberolllYl' b/"ecciells;.f, Apodel/l ll !> sp. Note that the large rodent ACCllllhioll is exclusive or the TG and TZ sites. Arricola <lIT . . wp­it/m from TG is slightl y larger than Ar\"ico la all'. s(lpidl/~'

from the Acheulian site Aridos I (Spai n). Large size is a derived c haracter in the Arr;("ola .wpidus lineage. but we nevertheless round some individuals rrom TG thal show the primitive "Mimomys- K<llIte" on their occl usal surf3ce. The Arl"icola species from Galeria likely W3S an early rorm related to the e."l: tant A. sapielm. The origin of this Iberi3n water molc TCmains unclear, and we believe it derived rrom an Early Pleistocene Mimolllp,' stock dis­tinct rrom the Mill/omys sOI'ini lineage th<lt leads to the An'icola terres/ris species. This is the subject or a work in preparmion by one of us (GCB).

The Micmtlls from TG (also at TO lD. TOIl) is as­signed to M. jClIIsolli for its similarity to CIIAUl>oE'S (1972) subspecies or M. agresri.~ (asymmetry. SA and ACC) rrom Sai nt Estcve Janson (France). The morphology or M. jclIIsoni from TG is intemlcdiate between those ort he Micmfll.\· from Saint Estcve Janson and La Fage (France). The species /ht!ltlmys brecciellsis (round also in the Ooli­na Icvels TOSb to TOil : see fig. ) is morphologically similar but slightl y larger th:1Il that round at Ambron3 (CUI.Nl'A- BESCOS ct al. 1999a). It is also larger than the Ibemlll",s from Aridos I (Aridos I and Ambrona are both in Spain) (LOI'EZ MARrlNI;.l 1980, StSt 1986) while it is similar in size to the 1ht!ltllllp" rrom Sai nt Estc:vc Janson. Porcupines together with other Mediterrancan taxa. sueh as /berum ... .I' bl¥!Cciem'is and An'ico la a(r. sapidlls, may indicate a humid and warm cl imatc during TG's uni ts GII and G ill l CUf.l'.("A-BES('6s et <11 . 1999:1).

The carnivore assembl<lge rrom TG (Units GlI and Gi l l) includes typical Middle Pleistoeene taxa : COllis IlIpIIS. 0 /01/ alpillll.l" elll'OpaellS and r1dpe.,· " Idpe~· . Meles mele.,·. "'"stela lIimlis. Urs lls sp. (dellillge"i-.~pel(lelll·

lineage). Palllhemleo cf /ossilil'. L.lnr pon/illlls .\]H!laelll· and Felil .~i/n's/ris. The TG Callis has a p~ and M1 mor­phologically suggesti ve or the Callis IUPlls group. but its dimensions arc inlermediate between COllis II/o.~bllchel/­

sis and CCllli.\ IlIpl/.~ (modem Iberian sample). The TG specimen belongs to the first .. true wolves" as those from Lunel Vie! (France) and Heppenloch (Gennany) (GARC1A 200): I ~ I). The transition rrom CClllis mosbachellsil' to the first true wolves (Cm/is IlIpu'\") occurred during the I-Iol stcin pt!riod. between MIS 9 and II (about 0.4 Ma). A minimum number of five Cl/Oil cllpilllll' (c. a. ellltlpaells) adult indi viduals have been recovered from TG. likel y dcrived from a ca tastrophic event or a natural trap. 0/011

(llpillllS is rairly abundant in Iberia n Late P1cistoecllc sites (especially between 100 and 15 ka). The presence of C. a/pilllls in the TG Middle Plei stQCene site represcnts the first occurrence of this canid on the Iberian Peninsula (Gi\RClA 200) ; 178). Bear rClll<lins were previou sly 3S­signed to UrSIlS spe!(lell.\· (GARCJA & AItSUAGA 1998) due

Cour. Forsch.-lnsl. Scntkcnbcrg. 259. 1007

to the size of onc p4. Ilowever. the TG ursid renl3ins 3fe scaree and cannot be assigned to species with certainty because of the overlap of the P~ vallles with those of U. dellillgeri and U. lIrctol' samples. Nevertheless, this premolar is morphologically allributHble to the speleoid linc:lgc. Pall/hero /eo. Vlllpes I'IIlpes. MII.I·leI(1 lIi\'{llis and Meles lIIele~' arc also represented in TOlO Hnd TOil (GAI<CiA 2003).

The herbivore association has some taxa in common wi th TO IO and TOll , i. e. Eqlll/s C(lbal"/~· . Sill' c r . .ft·m/a. Cen·/I.\ elaplllls prisclI.\·, DWI/a dallla atl'. claetonimw. Hemitragus bOIl(l/i and Stephallorhillll.\· hemiloeelllls. whi le others, such as Megll/ocems dowkinsi as well as :I

smaIl8i.\·01l (instead of the large Bosl Bi.wlI from TO). are only present in TG I 0 and TG 11. This association is typi­calor the middle Middle Plcistoce ne (ROSAS ct al. 1998. VAN J)FIt MADE I 999a, 200 I ).

T he S imil de los Uuesos site

The Sillla de los Huesos ( 5 1-1) cave is located deep inside Cueva Mayor-Cue\'a del Si lo comple.I( (around halra km from the surrace entrnnce). SH contains an extraordin<lry accumulation or approxim3tel y thiny human indi vidu­:lls (1/01110 heildelbergellsis) in <I ll ancient mud-breccia. together with Hn 3ssemblage of carnivores. rodents Hnd insecti vores (CUL'ICA- BESCOs ct a1. 1997. 2001. GAl<ciA et al. 1997. GARCi/\ & AI<SUAGA 2001b. CUENCA-B(sc6s 200)a. GAI<CIA 200): ( I) Eulipotyphla ( Insecti vofes) - Cmcit/II/"O sp.: (2) Chi roptera - Rhitwlop/lIIs lIIehe~l ·i.

/I,~I'o/is lIIyO/il/ M. b~I'/"i and Milliop/eru.\· schreiber.,·i: (3) Carn ivora - Callis sp .. 1111pes mlpes. MlIrle.\· lII(1rle.\/ /tI. /011/(/. Meles metes. MIIl 'le/a IIil"(llis. MI/Me/o (PlIIorills) putorills, UrSIlS dellillgeri. Lyw: IXlrdillllS .~pellle/ls. Felis silwsrr;J, P(llJIhera leo cr. /os.I'ilil"' PalJlhera sp. (jag­uar/leopard sizc): (4) Rodentia - Eliom.l:\· qlle"(:il/ll~·. AI­locricet/ll' corre=ells;.\", Pliomys lellki relicf/l.\·. ;"·licrot inae indct. and Apoelell/II,\ syh·mic/ls. Some fi sh vcrtebrae and teeth (Sollllo sp. and LellcisclIs sp.). undetemlincd small reptile bones, rrngments or gastropods, arthropods, and grecn fresh water algae (charophytes) arc also present.

U-scrics analyses of a speleothem (SRA- ) covering the /101110 heidelbel'gellsis-bcaring level yield a minimum age of 530 ka (BISCl/OfF et al. 2007) for the human accumulation. The presence or Pall/hem leo cr./cm·ilis as well as Clelhri­/JIIOllly~' acm/"hbl in the SI-! (GARciA et al. 1997. Blso lol'l' et a1. 2007) suggests 3 maximum age of around 600 ka. which is the oldest ror POl/fhel"Cl loo in Western Europe. and the s.me round at lsemia La Pineta (Italy) (GAl<ciA & AItSUA(;A 1999). This large relid is 3150 found in the Middle Plcistoccne TDIO and TDII levels from the nearby Gr.1Il Dolin;! and in all layers orTrinchcra Galeria. Corrd:ltion or the rodent fa una of the SI-! with that rrom Gr.m Dolina pro­vides runher chronological indications. PliOIl~I 'S lenki and Allocrice/lIs cO/w=.em·i~· are ch:lr.tcteriSlic elements or Mid­dlc Pleistocene rodent asscmblllges and occur both in the

103

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Cour. Forsch.- Inst. Scnckcnbcrg. 259, 2007

;~; ;; ~ 2~2 22 ~ -......... '" .. Ata~ Faunal Units

""" Plerslocene Age

!.. J., · . n·

•

• • • •

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",,'''' Homo anleoessor Homo heidebefVensis VillPes d. V. 8IopecoIdes

Canis sp (C. arn.nStSfmosNdlen$l$J l)'tlf cl. issIOdotenSlS

Pannooi:tis cf. nesti

cf. Baranogale antlC/ua Ursus doIinenSls Lynlf sp.

Mustela palermmea • Panthera gombaszoegensis

•• Camdae indet.

• · ..

• Ctocula Ct'OaJta

V~s ptaegIaaaI$ Canis mosbachen$l$ Fe/is sp.

HomoIhe/"I(Jm latidens

Melesmeles •• • L)'tlf pardlnus spelaeus

••• • Ursus sp. • • • •• Vu/pes vulpes

• • • Panthtlraleo

CaniS lupus

Cuon a~s europaeus Mattes d . M. forna.1m.wIes MuSle/a II/'oIlMs

• MuSlela puloros • "" lI!'5Us deningen

• Ursus sp. IU. rfemngerilspelaeusJ • Febs sNeSlns

• Cervidae i1det.

• • Hippopotamus sp • • • • • • (Jama "neSlii" vallonnetensis ,," ' •• • Stephanorllinus ellllscus ,," ' •• • Equus (stenonliln)

• • • •• " EucJadoceros gwIi • Bow:Iae fidel .. . • • • Bison d . W>If1Istedrensrs

• •• • Cervus elaphus cf a(:Ofl)tlatus

• Suidae Jldet.

D, • D, Sus serofa Mammulhu$ sp.

"" PraeovIDm pnscus • •• "" Equu$ (caballoid) D, Q. . " Slephanorltinus hemltoechus

• •• CeMls elaphus pnscu$

• Megaloceros dawfunsi? • • Bison sp. (smal) • • • Hen'iragus bonaJ

• •• Dama llama all. ciactoniane

f !

I

105

CUENC,,-lkscOs & GMICi,,: Mammal faunas of the Atapuerca cave sites (Sp3in)

SH and in the Gran Dolina TOl O and TOIl levels (CUENCA­BEsc6s et al. 1997.2001) which yielded combined ESR-U­series dates on mammal teelh of 308 10 418 ka (F ALGUEItES Cl al. 1999). Fossil remains of Apodemlls ~T/I"lI(iclI.\· are com­mon from Ihe early Middle PleiSlocene onwards in Europe. and Ihe late Middle Pleislocene field mouse is particularly large. Plioll~\".\· /ellki with crown cementum occurs in Middle Pleistocene localities as [semia la Pineta (SAt ... (983). Old Plio/ll)"~· individuals usually present crown ccmentun in the re-entrant angles tRABEDER 1981).

The abundant U. dellillgai sample from Sima de los Huesos is of special interest for establishi ng the evolu­tionary status or the Pleistocene bears. The chronology of the tr:.Ulsition of U. dellillgeri to U. spelaells is difficult to establish: ursids from Biache-Saint-Vaast (France). dated to the end orl he isotopic stage 7 (250 ka: GIlIIIARL> ct al. 2005). are considered to be transitional between the two species (A UGUSTE [992): level VII of Lezetxiki (Spai n) dated to between 200 and 300 ka (G,\RCiA et al. (997) also presents transitional traits (GAItCI;\ 1003) while in level VI (younger in age). ursids with more speleoid traits are found. The pattem that emerges is a terminal Middle Plcistocene transition from U. delliJlgeri to U. spe1aells. The metric and morphological pattems that characterize the U. deJlillgeri population from SH do not correspond with transitional forms of U. spe/aells (BONlfAY & Bus­Sli' ItE 1989). but with a typical representative of U. dellill­ge/"i tGARCIA et al . 1997. GARCiA 2003: 155).

Defi nition of the Alapllcrca Local Faunal Uni ts

The distinct fauna I assemblages frOlllthe variolls levels in the Atapuerca cave sites form a complex system of local fau nal units (figs 3. 4). which are defined both by the oc­currence and by the evolutionary status of particlllar taxa.

A tapuc rca ICa un a l Unit) (ATA FU I )

The ATA FU I is characterized by the co-occurrence of two Allop/wiolll)"s species. Cas/illoIIIY.v and AsoriclI/lIs. together with Pmlllollic/is cr. nes/ii . The TELRU assem­blage characterizes the oldest faunal unit of the Atapuerca sites (ATA FU I). This unit indicates a wet and temperate period. presumably short aller the arid Ebllronian phase (1.7 Ma). which would have opened the way to the distri­bution ofAlloplwioll~I's in Central. Western and Southem Europe (here and furthe r on. the chronostratigraphical correlations arc drawn from GIIlIlARD et al. 2005). This warm phase has been tentatively correlated to the Waal­ian (1.5- 1. 3 Ma) on the basis of its distinctive insectivore assoc iation (CUENCA- BESCOS & ROFES 2004. 2007). as well as of other temperate taxa. such as the large herbiv­ore J-liPPOPO/WIIIIS. the sci urid ScillrtlS wan/we and the murid Cas/illolll)"S rimJ· (figs 3. 4).

106

Ata puerca F:mllal Unit 2 (ATA f U 2)

The ATA FU 1 is chamcterized by the occurrence of Ibel"O-111.1'.\· hllescarensi.\·. Allocricellls and CroC/I/a. This unit is represented by the oldest faunal asscmblage of the Gran Oolina si te (TD I): this faunal assemblage is represented in the Gran Oolina levels 3 and 4 (T03-4). It is character­ised also by the occurrence. in the Atapuerca sites. of the rodents Alarmola sp .. H)"s/n~\" rejo.j·sa. Allocric(!IiIS bursae. P/iomy.\· episcopalis. Mimolll.I'J· SOl!illi, SfellO("l"(lI/illS gre­galoides. Terrico", (//1"a/ideIlS. MicI"QIIIS seseae. /bel"OlI~l"s hll(!scarellsis. and Apodemus sp. Among the insectivores. afT. Beremelldia sp. and Sorex mill/lms appear here for the first time. The most ancient fauna I asscl11blrtge from the Trinchera del Elefante Lower Red Unit. ATA FU I. almost disappears. with the exception of Allophaiomys cha/illei. CaslOI: Eliomy.\·. Erill(lct'IIS and ullpa. The most remarkable events concemi ng the Cami vora from T03. 4 are: a) the earliest. stratigraphically controlled occurrence in Europe o f CroCII/a croCUf(I (GARCIA & ARSUAGA 1999), regularly present from around 900 ka to around halfa mil­li on years ago. it is constantly associated with humans: b) TD4 bear remains belong to U"m.~ do/illellSis (GAkCiA & AItSUAGA 100 I a, b). Some of the postcranial propor­tions and dental traits of this ursid suggest an ancestral position within the cave bear lineage (V. dellillgeri - U. spell/em·), very close 10 the common ancestor of Ur.ws al"c/os. Comparative studies reveal significant si milarities to Ihe Le Vallonnet ursid (GARClA 2003: 466) but also to the Untennassfeld bears. now under review (GARciA 2004). However, new remains of UI"S/IS er dolillem·is are recorded from TELRU (ATA FU I). and this might lower its first oc­currence into FU I. As for the resl of the TD4 camivores. Calli.~ mosbacllemis and VII/pes praeglacia/is, appear for the first time in this unit. whi le Pall/hem gombas::oegellsis was already present in TELRU (ATA FU I).

Ata pucrca f aun a l Unit 3 (ATA FU 3)

TO Unit 2 represents the ATA FU 3. which is character­ized by the Gran Dolina levelS. and the lower part of level 6 (T05 and T D6a). [t is characleri sed by the co-oc­currence of the rodents Alloplwiom)'s c/wlillei and MiclV­IIIYs minI/IllS. the camivore flomOlherillm cr. /(I(idell.~ and the wild boar Sus scrop,.

Ata puerC:I F:mllal Unit 4 (ATA FU 4)

TD Unit 3 (ATA FU 4) is represented by the levcl T06b (lllso ca ll ed Aurora s tratum) and is the interval after the co-occurrence of Allophaiomys chali/1ei. and the large. red-Iooth shrews (CUI:.NCA-BESC()S & ROPES 2007). and before the occurrence of UicJ"Ol/Is raflicepoides. Moreo­ver the ATA FU 4 is characteri sed by the occurrence of Homo (lII/eces.)"Or.

At:ll)Ucrca Faunal Unit 5 (ATA FU 5)

TD Unit4 (ATA FU 5) is represented by the Gran Oolina levels T7 and T08a. It is defined by the co-occurrence of M icm fllS rllllicelJoides and cf. PI"(leOl'ibo.\· prisc /ls. Allocr icefll.\' bllrsal!, IbI! IVIIIY.~ /lII escarel/sis, Pliomys episcoJXIIi.~ and Mill/omy:; sM i lli . This is the latest occur­rencc of e{lI/is lIIosbachel/sis and Cmcl/fa croCI/W in TD, although still occurs in the TEUR U (see figs 3, 4).

Ata pu erca F:1I111l11 Unit 6 (ATA FU 6)

The ATA FU 6 is characterized by the co-occurrence of HOII/o heidelbergellsis. deri ved microtines. and Sfeplul/lo­l'hillllS hemifoec/IllS among other large and small mam­mu Is. The Atapuerca FtlUnal Unit 6 is well represented by Grun Dolina's unit fi ve (ATA ru 6), which includes levcl s 8b. 10 and 11 (TD8b. TDIOand T011 ),as well as the t:1unal assemblages of T EURU_ TG (G ll and G ill ) and SI-I. Moreover, the lower limit o f the ATA FU 6 is characteri sed by the occ urrence o f A llocricefll~' COI'­

re=ellsis. Terl'icola (If(lpller(llIellsis and Ibemmys brec­dells is. while its upper lil11it is unknown at the momcnt . A Canis /lIplIS M I from TOIO is fair ly small as thosc fi rs t "truc wolves", which are intennediate betwecn C. 1II0s/)achel/sis and modcrn C. IlIpIIS. The FAD of the true wol ves (Collis I/lpm') could be in 'I'D 10. dated to Mound 400 ka. or contemporaneous wi th TG (350 to 200 ka) whl're e{mis IlIplls is actuall y prese nt (Uni l Gil l). The same case ::applics to P{mtheraleo whosc first appearance cou ld either be at the basc ofTOIO. ::at TG or at S I-I. The base of TO la is characteriscd by the latest occurrcnce of I-!olllother i llm cr. ImidellS . Thc co-occurrence of thesc large-sized relids is uncommon: TO 10 could wcll repre­sent the presence of the sabre-tooth cat and of diffusion of Ihe lions. The Atapuerca s ites of Trinchcnl Galcria (TG ), Trillchera Zarpazos (TZ). Trinchcra del Elefantc Uppcr Red Unit or phase 11 (TE 19) and Sima de los I-Iuesos (SH ) share se veral small mamilla I species with unit 5 of the Gran 00lin3 sequence, thereforc we includc them in the same unit (ATA FU 6 of figs 3. 4). The Sima de los Huesos is not rich in its small mammal content but it shares the most common and characteristic rodcnts of Ihi s unit. i. e . Allocricetlu' COI'IY:!=el1sis and Pliomys lel1k1 withTD8b--TDI I andTG- TZ.

Atapuerca's fauna I SL"{jucnce can therefore tenlatively be proposed as a biostratigraphic framework for the Eurly to Middle PleislOcene in Spain (figs 3, 4 ).

Conclusions

The main conclusions of thi s study are that : I) Ihc fi rs l human acti vit ies in Wcstern Europe are recorded al­ready in the Early Plci stocene in the Iberian Penin sula

Cour. Forseh . -ln~1. Senckcnberg. 259, 2007

sites of Barranco Leon. Fucnte Nueva 3 in Granada and Trinchera dcl Elcfante Lowe r Red Unit, at Atapuerea . More precisely, the Trinchera dcl Elefante Lower Red Unit as well as the ATA FU I faunal assemblage date this early human evidence between 1.5 and 1. 1 Ma. The ATA FU I fauna indicales warm and wet scttings. probably referable to the Waalian. The European Early Pleistocene locali ties with lithic tools, as Fuente Nueva 3, Barranco Lean, Le Vallon nel. and Pirro Nord. can be included in ATA FU 1.

[n Gran Oolina (TO). the fi rs t human s from West­em Europe (Homo (ll/Iecessol') lived in warm. wel and wooded landscapes. probably corresponding to the Ma­rine lsolope Slages 21 to 19 (ATA FU 4). Thc trnnsition from the Early to Middl e Pleistocenc al Atapuerea is characteri sed by a palaeocnvironmelltal change recorded between the levels T0 5-6 (ATA FU 3. 4) :lI1d thc levels TDS- IO (ATA FU 5. 6).

The gencral opcning of the landscape at the begin­ning of the Middle Plci stoccne, ind icated by the ATA ru 6 evidence (levels Gran Dolina 8b to 11, Trinchera Elefante Upper Red Unit. Trinehera Galeria and Sima de los Huesos). likely promoted the dispersal of HOl/lo heidelberge,u is across Westem Europe.

Acknowledgements

Wc thank Eva BARCELONA, Ignacio CANUI>O and Hell ry GllllERT for their help: the Atapucrca team thal e;'(ca­vatc the sites every year. the MCYT research projects PB 96- 1026.C03- 02 . BXX 2000- 1258~C03-02 . and 60S 2003-08938- C03-0 1: the Junta dc Castilla y Lcon, thc Atapuerca and Ouques de Soria Foundations. and INAEM for their human and financial support . Drs. Lutz Chri stian MAUL and E\'elyne CRt GUT-BoNNOU RI' greatl y improved thc manuscript with their comments.

Refe rences

AGI.!I MK~. E. ( 1999): Molar mmaduro de M~mul en Gmn Dotina lAta. ptlcrca, Burgos). - Rc, ista E.sp;,"ol~ dl' P~ toonlolog.l~ . Ilorn· enaje at Prof.}. Tru)'ols, n. eXl .: 135-1-42: (h iedo.

AGU MM[. E. (200·,,: The Atapuerc:. (Sp.ll n ) fo~si l Silcs: [Xltenl1at , pro~rcss and queslions. - In: Mllul, L. C. & KII III.to. f , K.·I). (E.ds.): Late Neogene and QU:llcm:.1)' biodh crs ilY alld e\olu· lion: Rcg imml de"e lopnlcn1S :II,d inlcrrcg ional corre bllOI' S, 18· International Scnckenb.:1l: Confl'rcnce. VI Inlernal lonat I'alaconlologica l ColloqUium in Weimar. Weimar\GermanYI. 251h - 30th Apri l, 20001. - Term No~ tm , 200-4 (2): 61- 63: Berlin.

ACt'. '!. 1.. TOIt~). I., (hl ~ , O. & MIIKrl,[i~.-NII' \uo, 13. t2~I: Ilu· man presence In the Eart )' I'le istoce nl' o r Ihc Guad l.\-Uaza BaSin IGrJnada, Spain): Ihe 5 1tl'S o f Fuenlc Nuc\'a 3 and Oar­mnco u:oo. - In: MAUl , L. C. & KIIIII_H . R.-D. I Edq: Lalt NL'<IgC11C alld Quaternary biodl \'&:fSl ty and <:\o lulion: Rcgionlll ,k "dopmcnls lInd illl l·rrcgional corrcl:uiolls. t8'" Inlcmati'>nal

107

CutSCII-llLscos & GIII!.Ci,,: t\.l anunlll faunas of the Atapucrca cave si tes (Spain)

Scnd,C'nbc:~ Coofcreoce_ VI International Palaeontological Colloquium 111 Welmar. Welmar (Germany). 25th 30th Apn l. 20(14. - Terr-.. Noslr:I. 20(4 12): 64; Bcrlll1.

A~'I;II' "'' J. L.. GKACM. A .• IAKl'<lO. C. MAKli ... u. I. &: I" ·.MI.I.. I'. J. (1 9')<):11: RC'st\) er-meal hUlllallo de Ga lerialCue';1 de lo~ Zar­pal.O); ($it'rra d~ AI:IPUCrc: •. Burgosl. - In: CII KII('~ rl.l. E .• R(O!>,,~. A. & Oil" . J. C. (&ls. l: Al3puerca: Ocup:lclonc~ hu­m:1l1as y pak-oo..~olOl!:ia del )-JCUllfCnlo de Galcri:l. Memorias.

Arqurologia .'11 C3511113 y LeOn. 7: 233-235; lDrnor.. AM\t1lC11I. J . L.. /l1"'MI''''l7.. I .• GkIIO~.A .. C~KKETERO. J. /It & CUItO'­

Ill. E. [ 1993): Thn.'(' nc" human s~ulls from Ihe 5nna de los Il llesos Middl .. Pki~IOo:ene ~'I~ 111 Sierra tk AI<lpIlCre;I. Spall1.

N;lhll\', .' 62: 534 537; I.ondon. I\x'L'1IC11I. J . L.. MAK ljo;.l~ . I.. GkA(·III. A .• C AkkEn;kO. J. ~l .. Lt 'H f'<l.O.

C .• G"kcIA. N. & Okln;". A. J. (19'97): Smla d,' los Huesos (Sierra de Atapllerca. Spam). Th .. ~ I t e. - JOllrnal of Iluman E'ollllion. 33: 109-127: 1.00<100.

AIl'1 "la. J. L.. ~hMI1'U. J.. LOIH.VU. C .. Guu~. A .. \ IL·;;'W. A .• AI ,I"'."o, O. & G 11 I l.vU. J .( I9'}<)hl: The human cr:lIl1al retnall1S from Gr.on Dolina Lu"Cf l'lcl stCH: .. ne site (Siem. de- Alapllerca. Spall". - Joum~1 of Human Evolul ion • . H (3-1): 43 1-57: l.oudnn.

AIlZ"KlUU. M .• MII~COl-''''I. 1' •• p"VI", G .• J'''\ 't~. fo.t.. l' I,TK()'<lo. C .. l'uMI{x'I. !t t. . KOOIo.. L. 8: S,.nul.ll. R (2007): E\ldcrK"e of e~rh<'Sl hum3n oc~um:II""e In Europe; the Site of l'mo Nord (Southern Ital y). - Nalu!'\Oi~nseh3Ik'11. 9-': 107 112: 111."1-d.:I~

I)J;M ltllM1IM C ~"II.'), J. M . & NIl"l)l.ll'. M. E. (1997b): P3ra<:<Xklllogr~­phy ofth..., I\tapuerca SH Middle plcisl0,:elle hom ll1id SlItnple.

Jllunml ofl luman 1:' ·o lut ion. 3J: 333-355; I.llndon, 131 K,IILIlIJ Ill' CA';WO). J. M .• i{Vl>~'. A. & NIl<JI.ll~. M. E. \ I'm): IXn­

lal Il:malll~ from AlapUt'reu-TD6 \GrJn Duliml s ite. Bu~os. Sp:tin. - JOtlmal of IlunuIII E'olulloo. 31: 523 66; Lundtm.

01 RIIlllt"1 Ill. CWJll(l. J . M .• A.'I 0\(;.\. J. 1... C"Il", ... ru. E .• R (}\,,~.

A •• \1"1111'1:7_ I. & MO .... )ulll\. M. \ 1997a ): A 11011111111.1 from the WIlier l' lclStl)<.'.:nc of Alapuerea. Spall1 : " oss lblc Ances.­tor to Nealldertals 3nd Mo(km Humans. - SCIence. 276: 13<)2· 1395; W~~hinglOlI D.C.

BI~-II"n. J. 1... I'nz~.ITKIC"~. L.. l.nm. 1... AKW"O.t. J. L.. F"l(;lTkI'~. C.

13;\11111". J. J. 8: IlLu r .... T. (1'J'J7 ); Goolo~y mid prcltllll11ary d:tllllll of the homlllld-bc:;lrtn~ sedlmcntary fi ll oflhc S.ma de l.~ Iluesos C hamb..--r. CIIC"a Mayorofthe Sierra de Atapucll:a. Burgru.. Sp:tin. - Journal of I luman E'olulloo. 33' 129--154: London.

!31 ~Ol .. n. J . L . WltLIA~I~. K. W. R'")<;l .... ~t'[It. R. J .. AMI'I"l'Itt!. A.. AK~lM,~. J. 1... G~IllIIl. N. & Cl ENCII -Bl\(( .... G . (:::0117 ): II ll!h-..... solulion U-scnes d~le~ front Ihe Sima de los lI uesos homln ids yie lds k~ : Implicalions tor Ihe ... olulion or lhc <'arl y Ne~ndertha ls. Joomal of An:haL'tJlo~lcal Sciem:e. 34: 7(,3 770: Lanea>l .. r.

II I " .... H.-A. 12005): ConlnhulIo n de la l'ah:nherpCtofaune IAm­plllhia & Squamal.:l):i 13 coonal:;sance de I'r,olulIOll du chmal <'1 \lu pa}'sa~e du l'hocene superoeur au 1'1<' 1$IOCCne mo)en d·E.'pagnc . - l'hO The~IS. 1-102: P3 roS (Museum n~hon<ll d' II I~I();rc lI alUl\'lle. lnstllul dc P:dcontologlc Ilulllallle).

1l<'~II "'. M. I'. & IlUSSII,RI .. J. 1'. [1989): Lcs gr .. ndes 1'lUn<'~ de b groue d'Aldcne ~ Ursado.'s). - l3 uliclln Musec AlI lhropolOJ::le pr.:hls­hlt"1t.tue. 32: 11-19. Mormco.

C ~MI!O.,,,n I. 1: .. M()SQI., I.k". M . O Ll!. A .• R<JIlkK;LU.. X.-l' .. S .II I"')I. 'I. M . S\L~. R •• Vlk{.I\. J. M . (2001): Strul·tUn..· morphotel·h· 1I1quc de l'mJustne Itlhl'-juc du l'lcl~l~ne mfmeur Cl mO)'cn ... ·Alapuerca (Buq;os. Espa~ne). - l.·Anlhropolol:l~. 10S: 259- :::1\1): l"lroS_

CII MU,N.I I. F .. R!.HM ~ IlU Ill. C ~'I KO). J. M .• AIl,t " .. A. J. l. ., Dh.;.. J. C. R.-... ,,\. A.. Cll,,( .,·B ,-~"U\. G .. S,\(.II. i{ .. MU\()( U A. M. & i{'OM Kot 11.. X. 1'.11'.195 ): umer Plcblocclle hommids ;l11d ane­f:...-t, from Atapucn:(I-T06 (Sp:t,nl. - Sclenee. 269: 1126--83(): Wa,htngton D.e.

C~Mlkl"'.ru .• E .• E.q(H~". M .• ~h~II' N\!fkA.A.. MU~"JLlkl\.. M .• Ro­Ilkkot'l1_ X. 1' .. OuL. A .• S~I~. R .• W.ItGH. J. M .• IlIRIIlI>F.7 IM. C",lIIO. J.I\ \. & OMlI ()~. A. I. \ Iml: The Plcl~t ocene ~'I~

108

or Gr:ln Dol ina. Sic-mI <k I\tapllcrca. spa,": a hlSh'>!') o f the archal'QIOi;ical in,csugallOOs. - Journal of l-l uman E\\'IUhon. 37: 12l}-1S4: London.

CIt~I.I"' . J . (I 972 1: u~ rongellrs du l'lcbtoccnc moyen el SUp-Crleur de I'r:ulce . - C~hicrs dc P:.1 ':01l1010gie. 1-110; I':.ris ICNRS).

Cl.l~l .. -111 <;("t". G . \ 2003a l: J\n~hsl~ lilog .. netico de , tllocri(wtt:( dcl I'lclstoccno [Cricetidae. Rooenlla. ~l3mlllaha). - In: Wf.7-

M~Il"!I"u. N .. PEL;"ll-C"M ...... H ... [~. p. & II t~"lIs[)t7 Ftll" .... ' ­L1El. M . (&ls.l: En tomo a FOsil~ de Mamifcl"I»: 1);1(ac1oo. E"oluciOri ) P~I,'tJ(Itnbjen\e . - Coloqulos oc l>ak'lMllologia. \"01. e\t .• I : 95-113; Madnd.

C[;f"l ~-n l-.c:\J~. G. (2t103b): The Microlll:mHnal reeord as pro~)' of l'al~eOo:n';ronlllenml changc~ il11hc I'lclslocellc oflhe Slcm dcAI:lpucrea (Bu~()s. Sp:ll nl. - In: RUII-z.,.~AI .\. M. B .. Dv· kIIlJO-V "-l-'~o. M .. VAI IM'tIt ,tII.I.O,-ROtllC!GI.t:I. 1\ .. Gn ·G~~a~. M. J .• 1l ~IlLlI\.JI-Azt"R.IT£. T. tit .• Bl'~n~I "'ll-Gl'Ilfklll . l. I. & M"'HiM'Z-M E.. ... UIlAII.\t. I. {I'd~.): QU31C'mary e hlll~hc chang­e' and enllronmelllal croses In lhe Mediterranean Rcgion . - ! 330--1380: Alea!.i de IIcnares (UniHTSldad de Aka!:i de Iknare~).

CUENC,,-BI.-<;OJS. G. & Rorr:s CII .... "El. J. (2004): r nS<.~tit oros ( Mam­mai m). d ima y pa lSiljt· dc los ni-'eles inferioll:s de Trll1~her:l

Elditnlc (Pleis\oceno Inferior. Atapuercal. - In: Il"VI-'liM'<V. E. & RUIIIlI J" R", 5. (Eds.): Mlsccll\nca cn homenaJc a Emil­i~no AgUlrTc. - Zona Arquc:ol~ica. -': 151- 156: AIe:II:' de Iknal"l.'S.

Ct;["u-B(S(:o-;. G . & Ron;. •• J. (2(07): First e\ Id"nee o f POlSOllOIIS ~h",,,, s With an en,,'11O<I1311011 apparntus . - Nalurn l»cnschar­t~n. 94 : 113- 116: lk ldelbc:rg.

CUl."("~· lk-.c:-..)<;. G •. C""LI)O. J. I. &: l.1I ~I .IINII. C. (19')5): Los J\l"\ico­lidos I Roocntia. M~nllna l ia) de !os nil elt-os infenord dc Gmn Dohna (l' lcisux~no Inferior. Alupuerca. Ilurgos. Espai'ta) .

Ke' ISt3 Esp:tMla de Pak~mtu!ogia. 10: 202- 218: 0, .c-.Io. C L l'-<,,-Br'>I.tl!.. G .• CII"U)(). J. I . & LA~L.-\~". C ( Ima): An:ilis is

hloc~lr.otigr:ilico de los rocdo:>fcs del Pklsloceno Medlo del yaemllemo de Galcrla (Sier ..... de Alapuerca. llurg(lsl. - In: C~NIIO'lll_ E .• Ro<~s. A & 1)itz. J. C . [ Eds. l: AI~pucrca: OCllp:u:"ioncs hurnunn.s y paleoccologi3 de l Y:IClIlllelllu d<: Ga leria . M~mori~s. - Ar'-jlleol()gi~ ell C,,~till:t y Le611. 7: 1 i!9 210; 2;1Inor ...

Cl't'-/( \·!3c, .. ·( .... G .. C"NU)(). J. t. &: 1.,,'l.A'lIl. C. (2001): L:a ~ucnee d,'~ I\lngeurs [Mamm~lial des ~ II<'S du l'1';IS!oc .. ne mfcncur et IllO)'en d·Al:lp"'-'ll:a \Bu~os. Espa£M). - l·Anthropologic. 105: 11 ~130: Pan s.

CLI[",- ~·llf'o(.(I'., G .. Ltl't.ll"~. C. & C""llkJ. J. t. I 1mh): Blochrooo­log.c:" IIn plic~hon s of the AI"\ icoltda .. (Rodenlla. M:llnmalia) fn.)nl the Lowcr 1' lcisIO,:<.'I1C h011lIlud-b..·arinl! !clel ofTrindlcr;. [)ohm, 6 [TD6. J\lapucrca. Sp:. in). - Journal of I !tunall E,·o.­Iulion. 37: 353-373; l.ondon.

Cn",-"-Ik...:. ,,,,. G .• RorES. J. & G~M(ill·I'lMIUlTII. J. C. [20(5): Early Europeans and enlll"Ol1","cnlal change across the E:orl}-\lld­d1c Plcl~tocene 1r:lIlSHlon small mammalian c'locn-ce from TnncherJ Dolma ca'c. Atapuerca. Spain. - In : Ih ,,1). M. J . & Gill" INII. I'. L IEd'. I: Early Middlc I'lelstocene Ir~I1~lllons: The Lal1d-Oct~n El Idene~ . Ge('lol,lical 50,.-iely of London. Spcei~l l' ubli~~lions. 2-'7: 277· 28(1: London.

Ct.f'-/{ ~-B I ~.O<;. G .• l.1IPl.A~ I C~I'<£'II. C .• C"Nl'IX>. J . l. & I\K'-IJMl~. J. L. (1997 1: Small lII~mnul ls from Sun:! de I()~ Ilueso<. Jour-nal oflluman E\olullon. j3: 175-190; London.

Dtv. J . C. M' >III.--...o. v.. R ...... l(.t.:u. J .• R'N U.. J .• C KU£'. l. & 11l(""-'IT. R. \ 19'}9): EstudlO arqucolbglco de Ios restos de macro'erteb­r.odm de la Unid:ld III Ut: Galena (Sierra deAwpuen:al. - In: C" Mnv'£tl.. E .• i{,-",~,. A. & 1)11'1. J. C (Ed~.): All1pllcr,a : Ocupa"oncs hurnun:,s y p:lku~eologi:t del yaclmlCnh) dc G;lkri~ . M"tnorias. J\rqucolulli:. <'11 C:lslIll:! y Leon. 7: 265 2112; l:lInora.

l'''lt.t I M"~. C. BIII.\I". J. J .• Y'''''''· ''I ,~. Y .• AIt'l~l,lI. J. 1... BlMln '''f! IH; CM'lllt). J. M .• C"IlIlO"':I.I .• E .. BI-.£IM,n. J. L & I:>tM.)' J . M. (1'I9'}): Earliest humans rn Europe:: the all" of T06 GrJn 1)011113. Alapucrca. Spalll. Journal of l1umall E,'ulullon. 37' 343 352: Lundon.

FAI "l.tkF_~. C. BAIIAI". J .• J .• Y\J+..OI·,\\M. Y .• BISCIIOIT. J . L.. AkSUA(;~. J. L.. BrRh lU1l17 Ilf. CAHRO. J. ~'I. . CAMIIONF.I.I.. E. & DOLO. J .''''1. P.UUII; D~lalion p"r RPE Cl U-Th l.kS siles plc is· l<)1:"n('S ,r Alapl"'re~; Silll~ de los II0l·:iOS. TrinchcrJ Dolina el Trin,',ht'ra Gak-d~. - L' Anlhropologic. 105; 71- 8 1: Paris.

FI("CARll.ll. G. 8; TOItMf". D, (I %7): 1Ill1us ldid~ En/lydriCII .• ' ga/iC/Qhk. ,1.:1 PlciSlo.:cne d,·lI~1 S .. rdegna, - P~tlat'Qnlogl';lphia ltalica. 33; 131J-!6(J: PiS.1.

GARn ... . N. (2003): Osos y olros eami,oros de la SielTll de Alapllerea. F\lnd~ciOI1 OSQ f\ Sluri~ s . - )- 575; Ovkdo (Fundadon Oso de f\ slurius).

GAMeIA, N. (2\)().l1; N~"' RcsullS on Ih~ r~lllaills o(Ursidae frolll Ulltcr­llI:J ss J"dd: compar:llions" ilh Unms (/QI/lIi!II"i)' from Alapuerca 'lIld Olhcr Earl)' and Middk Pleistoccne siles. - In: MAUl .• L. C. & K)l llI . ~L R.-D. (Eds,): Late Nt"Qgene and Qualcmaf), biodi'ersily and ~"nlulion : Region,11 dc,l"Iopmems and inler­regional cOn"clalions. IS" Inlcmalion~tI Senckcnberg Confer­ence. Vllnlcmalionai p" lal'(Jnl010gical Colloquiulll in Wdm'lr. Wcimnr (GemwIlY). 2SIh - )Olh April. 2t~. - Teml NOSlr:t. 2004 (2): 11 2- 113: Berlin.

GAMCiA. N. & ARS\lA(iA. J. L. (1998): Thl' Clrni"ore remains frum Ihe homillid·bcaring TrinchcrJ Galeria. Sieffil de Alapucrca. 1\'I,d· die I'ki~locc llc sile (Spain). - GcoblOs. 31; 65'11---67-1; Lyon.

GARCI,I . N. & AH';UA(;A . J . L. (IW':I): Cam;,orcs fromlhe Early Pkis· loccnl.' hUllllllid·bc<lring Trinchcm Dolilla (, (Sierra de Al3' pUCTca. Sp"inl . - Journal of Human E\olulion, 37: -11 S- -I30; Lundun.

G,IKCi ... . N. & ARSlJAGA. 1. L. I~UOI a) : Ur.fll.1 <lo lilll:II';.I' : a nc" ,pceil's ofLo"er Plcistocellc un-id from Trinehl.·r:t [)(>linJ. Alapucrcn (Spain). - Comptes Rcndus de I'Aendcmic des Seicnc~"S. 332: 717-725; Paris.

G"kCI", N. & AII.'UA(lA. J. L. (2001 b): Lcs camiwres (I\-bmmali,,) d .. s si l~S du I'I~i sloci:nc ,tnl.'icn d mOY"'1l d'Alapuerea IEspagllcj. - L'Anlhropologi ... IllS: 83- 93: I'"ri s.

GAkti". N. & Il t'WI:ll .. F. C. ~il1 prcs.s ): Nc\\ di~o,""ryura I3rge mUSlcl­id - P"IIII(III1(;li)' er. 1/eJ'lii - (C"mil'or,,: r-.1:mul1ali,,) from the Early 1'lcistocel1c local ilY ofSima del ElcfanlC ISierra de Ala· puerca. Sp"in). - 1'"I:leonlogrJphka Ablcilung A: SlUUgan.

G,IRCIA. N .• AKSU)lGA. J. L. & TOIIRI". T. (1997); The carnivore remains from Ihe Sinm de los lIuesos r.. l iddlc Plcistoccnc site (S;Cffil de Atapucrca. Spain). - Journal of Human EI'olutiol1 , 3.1: 155- 17-1: Lond"n.

GI1I11.'kl). 1'. L.. Il-c'J-IlllIA.\I. S .. COIlt:N. K. r-. 1. & l\'IIIl>lAII.IFI.I.\J, A. (200S ): Global chronostrJligrJphieal correlation tnblc for thc 1:.51 2.7 million years. - Boreas. J 4: I; Oslo.

lI .. u,SY. D. (1 9)\6): Plcistocenc VI.·r!Cbr:llC Falln3S or Hunga ry. - 1-20~; Bnda~st (Ak:ldcmi"i KiadOi.

LAI'LA:'A. C. & Cl'U .. CA ·BLo;{'(X. G. (20011): Un" nucl'a eSiXele de M,,:;m. I,,; ( .. //Iv/llwill /llY;'! \Anieolidac, RO<l~nlia. Mammalia) Cll cl l' I~ISII)(:Cn() Inkrior dc' I:t Sil.'rrJ de Al~pu~re3 (Burgos. Espa~a).

RCI isl" Es pallokt d,' Palconlologia. 1 S: 77-1<7: Ovil·d\,. LAI'L'''A CONL'<A. c.. Co~wrF SAkI, A. & ClJ~.NCA -B Il.'\(l.s . G. (~O(W):

l'uparios de dipleros del 1'leiSlocel)() M~dio en 13 SicffiI de Alapuc· ... :" (Burgos, Esp,ula). - In BAQUlI)Mo'O. E. & Run,o JAM", S. (Eds. ): MiS<:c'l:\nc3 "n humcnaj~ a Emiliano Aguirre. - bma Arqueologic". -4 : 2-1(,-~5-1: Alenlo de Ilenares.

L",u ANI(>)oIM>:I .... ~, R, & CUI."c,, -B~-"c()s, G, (2002 ): The Grllrl Do lin" Sile {Lm,cr 10 I\ l iddk Plcis1occlle, Al;lpuerc". l3urgos. Spainl: n .. " p"Ia~O.·Il\"ironnll"nlal dala b:lsed on Ihe di slribulion of ,mall llla"unals. - I'alaeogeography. 1'"lacoelill1<l\ology. p"lncrJ/:;:ok'gy. 186: 3 11 - 334: AmSIl·rd'lln.

L')!'I:7-MAkll~U .• N. (19~n): Los microm,llniferos (Rodenlia. lusec· li,ora. L~gomurphJ y Chiroplera) de l Silio de OC UP<lCillll "chc1cnse de t\ridos-l ( Argalld". r-.l"drid) . - In: S...,..-ro"JA. M .. UJI'U·M AKnNE1. N. & 1'1:1112 GO:<l.ILE7.. A. (Eds.): OcUJXI' clvnes t\~hclcnscs en cl "alk del Jamllla IArg'Ulda. Mndrid). - 1(, 1- 202: I- ladrid tl>ipllt:le ilm Pro, illci"I ).

LU ll" . H. Df. KM!LK~. II. -D .• MOIG:.£. A.·"'I . & MOlJLl~. P.·E. (1988): Lcs l"aunes dc gmnds lllalllmifcrcs de b grUllc du "aIlUIll1CI. Roqllebrllne·Cap·M:lrlin. Alpes-Marilllllcs. -L·A nlhropvlogie. 91: 465--169: Pari"

Cour. Forsch.- Insl. Scnckcnberg. 259. 2007

MADE, J. VA" l>I:R 1 19'/')a); Artiodnclyla del ync il11iell1o InCSl'plcisH>CI.·IH> de Galeria (Sicrra de Alapllcrca), - In : C.'kl«OS[ll. E .. RO),,~s. A. & Diu. J. C (Eds.): At~pllcrca: Ocllp.wioncs hllm,lIlas y palco«ologia del yacil11icnh> de Galcria. MenlOTlas. -Arque· o logia CIl C~s1illa y Loon. 7: 1-13--167: Z;tmorJ.

MAN. J. VA:' I}[R (l999b): Ungulales from At:lpucn::1 TD6. - Joumal of l-l uman Evolutioll. 37: 389--113: London.

MAIIE. J. \'AN l'l:lt (2001): Les Ongulcs d·AI"pu~rc<l. SIr:llIgraphie el biogcogl';lphie, - L' An!hropologic. IllS: 95-11 3; JJ"ris.

MAKCQ\.INI. F. . Bow.I)()lo;~A. F. I' .• z...:'OIl'TI A. G .. K0TSA~I'. T. & MAl.7,'. p, (2000): 1'reliminary dala on the mkro- and rn"~ rolll,,,nlllal

remains from Casa Sghcn"i. Lo\\cr Vald"mo ITuscany. haly ). - Bollclino dcll" Soc iet3 P;tlconl010gica 11ali,ma. 39 (2): 243- 252; MI)lkna.

MQULLl, P.-E .. r:CUAsSOllX, A .. LlCO"I~\T. E. Dl'.SCU.lI~. E. & BAILON. S. (2006): Apport des gr:lllds m:llmnilc res dc III grolle dn " "lIon· net (Roquebnme·C~'p-M'U1;n. Alpes-M:trilimcs. Fmn.::cl;) la connaiss:mce du eadre biochronoJOlliquc de la s~"\:onde moil;':" du l'l~iS1''IC"nc ;nleriellr (I' Europe. - L·Anlhrupolugic. I1 U:

837-849; Pari •.

ORIf\;A, A. I. (1999): El pcimien\O de G"krb (Skrr.1 de Alapllcn:a). Cr6nic" de una inlerl'l.'n~i()n. - In; CAkIlONHL, F. .. RvsAs. A. & DIU, J. C. tEds.): f\taf!,,~rea: Ocllpa~ioncs h1lt11"lla S y palcOl'Cologia del y"cimienlo de G,lIcria . Mcmorias. - Arque­ologia en Caslil1a )' Leon. 7: 13--- 17: 7"'1I110r:i.

l'AlI.l~ . J. M. & P l~t:Z·GIJNZAL~Z. A.(1995): 1':llcomagl1c!ic Agl.· lo r Hominid Fossi ls m Alapuerca Archaeological Site, Spain. - Science. 269: 830-832; Washington D.e.

PARf_~. J. M. & P!.Hll-GONlAU.l. A. (1 999): I-bgnctochronoJogy :md slr;ltigrnphy M GrJn Dolin3 scc llun. Awpuercn Arch:leolugicnl I BUTgos. Spain). - JOllrnal of IllIm;1ll E'olulion. 37: 32S- 342: Lo ndon.

l' i:Rf:7-·G6:.1 .... I.U .• A .. AL~I .~A'll)lIe" T .. PI'''ILLA. A .. GAl.l .... M(IO. J .. B Ui"'­

VAS, J .. M IM11:<U_ M. J. & OHIHlA, A. 1. (1 995): Aprox im;tcion a la e~!rJligr"lb de G"kria en 1;1 Trineher;1 de la Sicml de Al"­pucn;a tBurgOS). - In: BL~Ml'lJtl.lll C"'~"MV. J. r..) .• AR~UM;A.

J. L. & CAMIKr.'~LL. E. (Eds.): EI'o lucion human;1 Clt Eump'1 y los y;lcimicmos dc la Sicffil de At~putrea. Worhhop Caslll!o d,' b MOla. - I: 99-146; "al1adoll<.l.

1" .MEl-GOI'.I.Alll. A .. 1'AMI~S . J. 11-1.. CAMIWNEtL. E .. ALEl.xA~OkL. T .• ORn:GA. A. J.. B t:1i!T<J. A. & MAkHI'> M[RI'o. M. A. (2001 ); Geologic de 1;1 SicrrJ de A t aJlIICre~1 el slrali!lrnpilic de~ I"I:m· pli sS<lges kaTSliquc$ de G"kria Cl Dolina (Bllrgos. Esp.11;11e). - L·A nlhropologie . 105: 27--1); I'aris.

R,\l!uJI:II., G. ( 198 1): Die An Icolidell (Rodent;". I\ lamtn:JliaJ ;IUS dcm Pli07.~n und tl~l1l :lll<:rell PleiSlu7.!in ,"011 Nit-derOsten"elch. -Bcllr:igl":tor I'al:iunlologic von 6s1ern:ich. 8: 1- 373: Wiell ,

R<Jff.s. J. & C UE"<" ... -B"'-~. G. \2()()4): Smalll\bmmals fromlhe Span· ish I'lci.1 C>CCIlC s ites of AI:lpllcn.":t. - Joumal of Vcrlcbr~lc PnlcolllolollY. 2-' : 10SA; Northhrwk.

RlJn:s. J. & CU~~A-Br"or... G. (2006): Firsl c,'idenee oflhe Soricidae (Mammnlia) ..r~(Jric"I,, < t:ibhemt/'III (l'cl':nyi. ]1\64) in lhe I'JcislOCc'nc' oC Nonh Iberia . - Ri, i,l:l 11alian" di I'alconlolo­gia e Str:lli!lr:I!;3. 112: 301 - 315: Fi ren7.e.

Root. . L. (1,-,,)5): Pmll/()/IICI"· lIe..-l1l (Camilora Mal11l11:lli~llrom lhc

law Villafmnchi311 of !'il.·lmrl1la (Umbria, 11"ly). I'rcliminary 1100e. - Ec10gac geologlca HelI'Clia, 118 (3 ): llB- I\M; Bnsd.

Ro,,,,. A .. " l.kf.l-GOS1.A11:l.. A .. C AR II-USEtL. E .• MAin .. J. \,,,~ O~K.

SANCll r.l. A .• LMt .... "A. C. CL'f.>I( \· IlES(·us. G .• PAHe.. J. M. & HU\,Ul l. R. (2{)OII: Lc gisemenl plcisloc~ne de la "Sima del Elefanlc·· (SicTJ"J de Alapuerca. Espagll~). - L·Amhmpulogic. I (IS: )UI-3 12; )' ;'fis,

R"sA~, A" CAMIK)lo,I'LI .. E .. CUfNl"A. G .. GAKCiA. 1'1" FI,W'l"WIJ J"'1.I<), Y .. MAllf, J. ,"''l OCR, Olll, A .. 1' ''Mts. J. Ill.. P>.Kl7 G0:'U.Lll, A .. SA'lCllf;: MAII.C(l, A .. SA"CIIU Otll.I.IJN. B. & V ... l.\'!:lIll1. J . 11991\): CrOllologia. b;<lcs!r"ligralia y p,tlco<:coiogi" .Id l'lcis!t)CI.·no Medio .I,' G"I"ria (SicrrJ de Alapucn:a. Esp:tila) , - Rc\ iSla Esp:l!lola de PalconloIO!li;I. 13: 7J - :SO; O"k-<lu,

RUM.', A .. CARIIIY.'ELI., E" OI.Ll. A" I'lkl'.l. G()'l/..\UL A., VAU.I'I·R I)l. J .. HUGU£T, R .. CA(tH". I .. R()SlLl .• L SAl .... . R .. Mu:<l)l.:I.K A. M ..

109

Cu[:<'cA·B£scos & G l\lId,,: Mammal faunas ofthc Atapucrca cavc sitcs (Spain)

M AD(, J. us 01:11.. S;'SCllEZ M AM(.Xl. A .• C Uf.-.G\. G., Rt,lIlIO:iGtJu,

J . P. & ROI>ItKiut:l. J. \ 1999): ContnDucion del yacinllcnto de G:llcria ISierr:1 de Alapucrca) al CU31cnmrio IbCrico. - In: C.'NIIO:<'EL.l, E., Ros ... s. A. & DI~z. J. C. (Eds.): Ampucrca: Ocupaeionl'S humanas y palcoceologia del yacimlcnto de Galeria. Mtmonas. - ArqutOlogia en Castilla '1 lloOn. 7: 377- 390: UmOr:l.

Ru:.. •. \. A .• Il tJQJ£I", R .. PI.Rf.7.-(i(r;l.ALEZ, A .. C ... M_ELL, E .. V"'U.I'[MIJU,

J .. M"DE, J. V ... S llf.k . AI.l.UE, E .. G ARel .... N .. MAkTINEl· P~M£1.,

R .. ROOklc;uf.l .. R .. SAI./I, R .. 5,..1./101(. 1' .. SIMON, G .• M"' MTh~EZ·

M .I1.A. C. I3.I~Tlk. M .. S ... "«:HU. A. & P"llls. J. M . (200-1 ): Inllial approoch I() the si te formallon and Palcoecology of the "Sima del Elcfanlc": a Plcistoc .. nc karst locahl'1 al Ala­puen::1 Hill . - In: B ... QCWAMI. E. & RUKIO J ... R ... , S. (Eds.): ~l iSl:cI:inc3 en homcnajc a Emilinno Aguirr ... - Zona Arquco--10gica . ..J : 134-155: Alea l:l de I knare~.

S ... G\. B. (1983): u. fauna dt:1 giacimcnto dl Iscmla la PmCla (nOla prdiminarc). Iscmlu la Pincla. un acc:nnp"mcnlo pni anlico di 700.000 Illln;' C:II:.logo dclla mOSlrJ . - 71 - 79: Bologn!l. lEd. Calderini).

SESl. C. \ 19);6): InSIXti-oros. Rocdorcs '1 lagomOl'"fos (Mammalia) del sitio de ocupac ion Aehclen~ de AlIlbrona (Soria. F...sp:lfia). - Esludios GtOlOgicos . . '1: 355-359: Madrid.

5ol1'< IIWl· .... M . V" B ... K'.lNln .... V. S. & TIIOI', V. V. (2002): Camil'~s

of Ihe Khapry f~un:tI assemblage and the;r strJI;grJphic im· plications. - Stratigraph'1 and Geologic~l Correlation. 10: 375-390: Mosco".

M:lIIuseripl submillcd 2005- IO--{).l Revised manuscripl :ll'cepttd 2007-06-08