Impactos y Vulnerabilidad I. Martínez1*, G. Cuenca-Bescós2+ 1 Departamento de Ciencias de la Vida, Área de Antropología Física, Universidad de Alcalá Departamento de Ciencias de la Tierra, Grupo Aragosaurus-IUCA, Área de Paleontología, Universidad de Zaragoza * Correo electrónico: ignacio.martinezm@uah.es + Correo electrónico: cuencag@unizar.es 2 1 Adaptación, vulnerabilidad e impacto en la evolución humana Resultados clave /DV PRGL¿FDFLRQHV HQ OD YHJHWDFLyQ GHELGDV D XQ FDPELRFOLPiWLFRTXHFRPHQ]yKDFH¶PLOORQHVGHDxRV HQHOHVWHGHÈIULFDIXHURQGHWHUPLQDQWHVHQODDSDULFLyQ GHOJpQHURHomo. /RV GDWRV VREUH OD GLYHUVLGDG UHODWLYD GH ODV FRPXQLGDGHV GH PLFURPDPtIHURV HQ HO \DFLPLHQWR GH *UDQ'ROLQD6LHUUDGH$WDSXHUFDVHxDODQXQSHULRGRGH GHVSREODPLHQWRGH(XURSDOLJDGRDXQFDPELRFOLPiWLFR KDFHHQWUH¶\¶PLOORQHVGHDxRV /DH[WLQFLyQGHORVQHDQGHUWDOHVHVWXYRHVWUHFKDPHQWH UHODFLRQDGD FRQ ODV UiSLGDV ÀXFWXDFLRQHV FOLPiWLFDV VXIULGDVHQ(XURSDD¿QDOHVGHO3OHLVWRFHQRVXSHULRU (OFDPELRGHHFRQRPtD\GHPHQWDOLGDGGHELGRVDOD UHYROXFLyQ QHROtWLFD VRQ ODV FDXVDV IXQGDPHQWDOHV GHO SURIXQGR LPSDFWR GH ODV DFWLYLGDGHV KXPDQDV HQ ORV HFRVLVWHPDV GH WRGR HO PXQGR GHVGH KDFH DOUHGHGRU GH DxRV Contexto 'HVGH VX RULJHQ KDFH DOUHGHGRU GH PLOORQHV GH DxRV 0D ORV VHUHV YLYRV VH KDQ YLVWR DIHFWDGRV SRU ODV YDULDFLRQHV FOLPiWLFDV TXH VH KDQ VXFHGLGRDORODUJRGHODKLVWRULDGHOD7LHUUD/RVIDFWRUHV FOLPiWLFRV VRQ GHWHUPLQDQWHV HQ ORV SDUiPHWURV ItVLFR TXtPLFRV FRPR VRQ OD WHPSHUDWXUD DPELHQWH HO JUDGR GHUDGLDFLyQVRODURODDFLGH]\FRPSRVLFLyQGHODJXD \WDPELpQHQODVYDULDEOHVGHQDWXUDOH]DELROyJLFDFRPR VRQHOWLSRGHYHJHWDFLyQ\GHIDXQDGHODVTXHGHSHQGHQ ODYLGDRODPXHUWHGHORVRUJDQLVPRV3RUHOORORVVHUHV YLYRV HVWiQ VXMHWRV D XQ LQFHVDQWH SURFHVR HYROXWLYR FX\R UHVXOWDGR HV VX DGDSWDFLyQ D ORV GLVWLQWRV FOLPDV GHFDGDPRPHQWR(QHVWHVHQWLGRVHSRGUtDGHVFULELUOD 9LGDFRPRXQVLVWHPDH[SHUWRTXHFRQYLHUWHODHQHUJtD HQ LQIRUPDFLyQ D WUDYpV GH XQ SURFHVR GH JHQHUDFLyQ DOHDWRULDGHYDULDELOLGDG\VHOHFFLyQGHODVYDULDQWHVPiV IDYRUDEOHVSDUDODVXSHUYLYHQFLDGHORVLQGLYLGXRVHQODV FRQGLFLRQHVDPELHQWDOHVGHFDGDPRPHQWR 1XHVWUR OLQDMH QR KD VLGR LQPXQH D HVWD FLUFXQVWDQFLD \ GXUDQWH OD KLVWRULD HYROXWLYD GH QXHVWUR JUXSR ]RROyJLFR ORV VHUHV KXPDQRV \ VXV DQWHSDVDGRV KDQ VLGR DIHFWDGRV SRU ODV YDULDFLRQHV DFDHFLGDV HQ HO FOLPD )LQOD\VRQ 1XHVWUD YXOQHUDELOLGDG D ORV HYHQWRVGHFDPELRFOLPiWLFRKDQGHWHUPLQDGRODH[WLQFLyQ GH DOJXQDV HVSHFLHV GH QXHVWUD IDPLOLD HYROXWLYD \ WDPELpQKDQFRQGLFLRQDGRQXHVWUDKLVWRULDELRJHRJUi¿FD HVSHFLDOPHQWH HQ OR UHIHULGR DO SREODPLHQWR GH ODV WLHUUDVGHO9LHMR0XQGR3RURWUDSDUWHORVFDPELRVHQ HOFOLPDWDPELpQKDQVLGRGHWHUPLQDQWHVHQODDSDULFLyQ GHDGDSWDFLRQHVHVSHFt¿FDV\HQODDSDULFLyQGHQXHYDV HVSHFLHV \ PX\ VLJQL¿FDWLYDPHQWH MXJDURQ XQ SDSHO FUXFLDOHQHOVXUJLPLHQWRGHOSURSLRJpQHURHomo. /D HYROXFLyQ ELROyJLFD GH QXHVWUD HVWLUSH GHVHPERFyHQXQIHQyPHQRQRYHGRVRHQODKLVWRULDGH OD9LGDODHYROXFLyQFXOWXUDOIUXWRGHODFXDOORVVHUHV KXPDQRV FRQVLJXLHURQ XQD LQGHSHQGHQFLD FUHFLHQWH GH ORVIDFWRUHVFOLPiWLFRV)LQDOPHQWHHQHO~OWLPRWUDPRGH QXHVWUDKLVWRULDHYROXWLYDODKXPDQLGDGVHKDFRQYHUWLGR HQXQIDFWRUFDGDYH]PiVLQÀX\HQWHHQHOFOLPDSULPHUR DHVFDODVORFDOHV\UHFLHQWHPHQWHDHVFDODSODQHWDULD (OHVWDEOHFLPLHQWRGHODVYDULDFLRQHVFOLPiWLFDV GHO SDVDGR \ GH VX LQÀXHQFLD HQ ODV FRPXQLGDGHV ELROyJLFDV HQ JHQHUDO \ GH OD HYROXFLyQ KXPDQD HQ SDUWLFXODUHVXQSUREOHPDTXHSXHGHVHUDERUGDGRDSDUWLU GHOUHJLVWURJHROyJLFR\WDPELpQGHVGHHOUHJLVWURIyVLO (QHVWDOtQHDHOHVWXGLRGHORVJUDQRVGHSROHQIRVLOL]DGRV SURSRUFLRQD XQD LQIRUPDFLyQ GH JUDQ LPSRUWDQFLD SXHV 81 QRVSHUPLWHDSUR[LPDUQRVDOFRQRFLPLHQWRGHYDULDEOHV WDQ GHSHQGLHQWHV GH ORV IDFWRUHV FOLPiWLFRV FRPR VRQ HO WLSR GH ÀRUD \ ODV SURSRUFLRQHV HQWUH YHJHWDFLyQ DUEyUHD DUEXVWLYD \ KHUEiFHD 'HVDIRUWXQDGDPHQWH ODV FRQGLFLRQHV GH IRVLOL]DFLyQ SUHVHQWHV HQ PXFKRV \DFLPLHQWRV FRQ SUHVHQFLD KXPDQD HVSHFLDOPHQWH HQ FXHYDVQRIDYRUHFHQODFRQVHUYDFLyQGHOSROHQ\KDFHQ TXHHOUHJLVWURSROtQLFRHQGLFKRV\DFLPLHQWRVHVWpDYHFHV DXVHQWHRVHDPX\GLVFRQWLQXR8QDYtDFRPSOHPHQWDULD GHDSUR[LPDFLyQDOSUREOHPDODFRQVWLWX\HHOHVWXGLRGH ODIDXQDDFRPSDxDQWHHQGLFKRV\DFLPLHQWRV /RV UHVWRV GH JUDQGHV PDPtIHURV VRQ PX\ DEXQGDQWHV HQ ORV \DFLPLHQWRV DUTXHROyJLFRV SHUR VX YDORU HQ OD UHFRQVWUXFFLyQ SDOHRDPELHQWDO HVWi PX\ PHQJXDGR SRU GRV FLUFXQVWDQFLDV (Q SULPHU OXJDU HO iUHD GH YLGD GH GLFKRV DQLPDOHV VXHOH DEDUFDU XQD JUDQ H[WHQVLyQ \ VX DVRFLDFLyQ D DPELHQWHV FRQFUHWRV UHVXOWD OD[D HQ OD PD\RUtD GH ODV HVSHFLHV SUHVHQWHV HQ ORV \DFLPLHQWRV $Vt HV IUHFXHQWH HQFRQWUDU DVRFLDGRV HQ ORV \DFLPLHQWRV IyVLOHV GH HTXLQRV KDELWXDOPHQWH UHODFLRQDGRV FRQ DPELHQWHV DELHUWRV FRQ RWURV GH FpUYLGRV FRQVLGHUDGRV FRPR KDELWDQWHV GH PHGLRV PiVDUERODGRV3RURWUDSDUWHORVUHVWRVGHORVJUDQGHV PDPtIHURV OOHJDURQ D ORV \DFLPLHQWRV DUTXHROyJLFRV GHELGRDVXFRQVXPRSRUSDUWHGHORVKXPDQRV\IXHURQ VXV SUHIHUHQFLDV FLQHJpWLFDV SRU HQFLPD RWURV IDFWRUHV PHGLRDPELHQWDOHVODVTXHGHWHUPLQDURQODSUHVHQFLDGH XQDVXRWUDVHVSHFLHV 6LQHPEDUJRHVWRVSUREOHPDVSXHGHQVRVOD\DUVH VLVHHPSOHDQORVIyVLOHVGHPLFURPDPtIHURV/DFRUWDYLGD GHORVLQGLYLGXRVVXUHGXFLGDiUHDGHYLGD\ODHVWUHFKD DGDSWDFLyQDDPELHQWHVPX\FRQFUHWRVGHPXFKDVGHVXV HVSHFLHV FRQYLHUWH D HVWRV SHTXHxRV KDELWDQWHV GH ORV ERVTXHV ULEHUDV \ SUDGHUDV HQ H[FHOHQWHV LQGLFDGRUHV PHGLRDPELHQWDOHV$GHPiVORVPLFURPDPtIHURVQRHUDQ SUHVDVGHORVKXPDQRVGHOSDVDGRSRUORTXHVXSUHVHQFLD HQORV\DFLPLHQWRVQRUHVSRQGHDODVDFWLYLGDGHVKXPDQDV\ QRPXHVWUDSRUWDQWRHOVHVJRDQWUySLFR&XHQFD%HVFyV HWDO(QODPD\RUtDGHORVFDVRVORVUHVWRVGHORV PLFURPDPtIHURVOOHJDURQDORV\DFLPLHQWRVHQHOVHQRGH HJDJUySLODVGHDYHVGHSUHVDHVSHFLDOPHQWHGHOHFKX]DV (VWDVDYHVVRQGHSUHGDGRUHVJHQHUDOLVWDVTXHUHDOL]DQXQ PXHVWUHRQRVHVJDGRGHODVHVSHFLHVGHPLFURPDPtIHURV /RVSDOHRQWyORJRVKDQDSUHQGLGRDGLVWLQJXLUODVKXHOODV GH OD GLJHVWLyQ GH HVWDV DYHV HQ ORV KXHVRV GH VXV YtFWLPDV \ SXHGHQ DVt VHOHFFLRQDU ORV FDVRV HQ ORV TXH OD DFXPXODFLyQ GH UHVWRV IyVLOHV GH PLFURPDPtIHURV HV UD]RQDEOHPHQWH UHSUHVHQWDWLYD GH ORV HFRVLVWHPDV GHO SDVDGR (QWUH ODV GLVWLQWDV DSUR[LPDFLRQHV UHDOL]DGDV D SDUWLU GHO HVWXGLR GH ORV IyVLOHV GH PLFURPDPtIHURV HO DQiOLVLVGHODGLYHUVLGDGUHODWLYDGHODVFRPXQLGDGHVGH PLFURPDPtIHURV GHO SDVDGR HVWi RIUHFLHQGR UHVXOWDGRV GH JUDQ LQWHUpV \ VH KD UHYHODGR HQ ORV ~OWLPRV DxRV FRPR XQD H[FHOHQWH KHUUDPLHQWD HQ OD UHFRQVWUXFFLyQ SDOHRDPELHQWDOHQORV\DFLPLHQWRVDUTXHROyJLFRV (Q HVWD OtQHD HV LQWHUHVDQWH GHVWDFDU TXH XQ PHMRU FRQRFLPLHQWR GH OD HVWUXFWXUD GH ODV DFWXDOHV FRPXQLGDGHV GH PLFURPDPtIHURV HQ ORV GLIHUHQWHV DPELHQWHV QRV SHUPLWLUi UHDOL]DU UHFRQVWUXFFLRQHV SDOHRDPELHQWDOHVPiVSUHFLVDV\¿GHGLJQDV3RUHOORXQR GHORVFDPSRVGHLQYHVWLJDFLyQSDOHRHFROyJLFDGHPD\RU LQWHUpV HQ OD DFWXDOLGDG FRQVLVWH HQ OD UHFRSLODFLyQ \ DQiOLVLVGHHVWRVGDWRVDSDUWLUGHOHVWXGLRGHHJDJUySLODV HQORVGLIHUHQWHVELRWRSRVLEpULFRV 82 Adaptación en la evolución humana: el origen del género Homo $XQTXH HO UHJLVWUR IyVLO HV PX\ IUDJPHQWDULR SDUHFHFODURTXHORVSULPHURVUHSUHVHQWDQWHVGHQXHVWUR OLQDMH YLYLHURQ HQWUH KDFH \ 0D 0DUWtQH] )LJXUD/HVFRQRFHPRVSRUXQSXxDGRGHIyVLOHVTXH SURFHGHQGHXQDH[WHQVDUHJLyQGHOFRQWLQHQWHDIULFDQR TXH VH H[WLHQGH HQWUH OD DFWXDO 5HS~EOLFD GH &KDG (WLRStD \ .HQLD /D HVSHFLH GH OD TXH WHQHPRV PiV GDWRVHVArdipithecus ramidusTXHYLYLyKDFHDOUHGHGRU GH 0D 6H WUDWDED GH SULPDWHV FX\D H[LVWHQFLD WUDQVFXUUtD HQ HO VHQR GHO ERVTXH WURSLFDO \ FX\R WLSR GH YLGD \ GH DOLPHQWDFLyQ QR GLIHULUtDQ GH ODV GH ORV DFWXDOHVFKLPSDQFpV$XQTXHVXWLSRGHORFRPRFLyQHUD IXQGDPHQWDOPHQWH DUERUtFROD HUDQ EtSHGRV IDFXOWDWLYRV TXH VH HUJXtDQ VREUH VXV H[WUHPLGDGHV SRVWHULRUHV SDUD GHVSOD]DUVHSRUHOVXHORGHOERVTXHWURSLFDO +DFH HQWUH \ 0D QXHVWUD IDPLOLD ELROyJLFD JR]DED GH XQ FLHUWR HVSOHQGRU HQ FXDQWR DO Q~PHUR GH HVSHFLHV GH ODV TXH OD PHMRU FRQRFLGD HV Australopithecus afarensis (Q JHQHUDO VH WUDWDED GH IRUPDV SOHQDPHQWH DGDSWDGDV D OD ORFRPRFLyQ EtSHGD DXQTXH D~Q FRQVHUYDEDQ LPSRUWDQWHV FDSDFLGDGHV DUERUtFRODV /D DQDWRPtD FUDQHDO PDQGLEXODU \ PX\ HVSHFLDOPHQWH GH OD GHQWLFLyQ DSXQWDQ D TXH HVWRV KRPtQLGRVKDEtDQFDPELDGRODGLHWDGHVXVDQWHSDVDGRV SDUDKDFHUODPiVYDULDGDLQFOX\HQGRSURGXFWRVYHJHWDOHV TXHUHTXHUtDQXQDPDVWLFDFLyQDOJRPiVLQWHQVD$XQTXH VXYLGDVHJXtDYLQFXODGDDORVDPELHQWHVIRUHVWDOHVHUDQ FDSDFHVGHYLYLU\PHGUDUHQODVOLQGHVGHORVERVTXHV\ HQODVSUDGHUDVDUERODGDV (VWD UHODWLYD GLYHUVLGDG FRPHQ]y D PHQJXDU FRPR FRQVHFXHQFLD GH XQ FDPELR FOLPiWLFR TXH FRPHQ]yKDFHXQRV0D\TXHDIHFWySURIXQGDPHQWH D OD YHJHWDFLyQ GH ORV HFRVLVWHPDV HQ ORV TXH YLYtDQ ORV KRPLQLQRV (Q HVD pSRFD FRPHQ]DURQ D SURGXFLUVH HQ QXHVWUR SODQHWD PDUFDGDV RVFLODFLRQHV FOLPiWLFDV DVRFLDGDV D OD H[WHQVLyQ GH ORV PDQWRV GH KLHOR HQ HO KHPLVIHULRQRUWHGXUDQWHODVpSRFDVIUtDVGHWHUPLQDQGR XQ FDPELR HQ HO UpJLPHQ GH OOXYLDV TXH VH IXHURQ KDFLHQGR SDXODWLQDPHQWH PiV HVFDVDV HQ OD UHJLyQ RULHQWDO GH ÈIULFD &RPR FRQVHFXHQFLD ORV ERVTXHV IXHURQUHGXFLpQGRVHPLHQWUDVTXHODVSUDGHUDV\VDEDQDV VHH[WHQGLHURQSRUDPSOLDVUHJLRQHV(QORV\DFLPLHQWRV GH HVD pSRFD VH UHJLVWUD HO UHHPSOD]DPLHQWR GH IDXQDV HVSHFLDOPHQWHGHPLFURPDPtIHURV\DQWtORSHVDVRFLDGDV D PHGLRV DUERODGRV \ ULFRV HQ DJXD SRU RWUDV PiV FDUDFWHUtVWLFDVGHSUDGHUDVKHUEiFHDV%REHHWDO (VWD FULVLV FOLPiWLFD \ HFROyJLFD LQÀX\y GHFLVLYDPHQWH HQODKLVWRULDHYROXWLYDGHORVKRPLQLQRVFRQWULEX\HQGR DODGHVDSDULFLyQGHWRGRVORVUHSUHVHQWDQWHVGHOJpQHUR Australopithecus \ SURSLFLDQGR OD VHOHFFLyQ GH QXHYDV IRUPDV PHMRU DGDSWDGDV D ORV PHGLRV PiV DELHUWRV ORV SULPHURV UHSUHVHQWDQWHV GH ORV JpQHURV Paranthropus ORVSDUiQWURSRV\HomoORVKXPDQRV /RV SDUiQWURSRV PHGUDURQ D OR ODUJR GH XQD DPSOLD ]RQD JHRJUi¿FD TXH DEDUFDED GHVGH ODV ULEHUDV GHO 0DU 5RMR HQ OD DFWXDO (WLRStD KDVWD HO H[WUHPR PHULGLRQDO GHO FRQWLQHQWH HQ OD DFWXDO 6XGiIULFD 6H WUDWDEDGHKRPLQLQRVFRQXQJUDQGHVDUUROORGHODSDUDWR PDVWLFDGRU TXH HVWDEDQ HVSHFLDOL]DGRV HQ HO FRQVXPR GHOWLSRGHSODQWDVPiVGXUDV\¿EURVDVTXHFUHFHQHQ ODV SUDGHUDV \ VDEDQDV DIULFDQDV 8QJDU 6SRQKHLPHU /D HVSHFLHParanthropus boisei TXH YLYLy HQ OD Figura 1 Figura 1. Distribución temporal de los homininos. Fuente: Elaboración propia. WLHUUDV RULHQWDOHV GH ÈIULFD HQWUH KDFH \ 0D IXH OD TXH OOHYy DO H[WUHPR HVWD HVSHFLDOL]DFLyQ \ VH KD GHWHUPLQDGRTXHODVSODQWDVGHWLSR&FDUDFWHUtVWLFDVGH ORVPHGLRVDELHUWRVFRQVWLWXLUtDQKDVWDHOGHVXGLHWD &HUOLQJHWDO$SHVDUGHVXHVSHFLDODGDSWDFLyQ D ORV QXHYRV DPELHQWHV DELHUWRV ORV SDUiQWURSRV VH H[WLQJXLHURQ¿QDOPHQWHKDFHDOUHGHGRUGHXQPLOOyQGH DxRV6RORQXHVWURJpQHURIXHFDSD]GHVREUHYLYLUHQXQ PXQGRGHVDUERODGRPHUFHGDXQUHYROXFLRQDULRFDPELR TXH OH KDELOLWy SDUD RFXSDU XQ QLFKR HFROyJLFR LQpGLWR HQ OD KLVWRULD GH ORV SULPDWHV HO FRQVXPR KDELWXDO GH FDUQH (OSULPHUIyVLODWULEXLEOHDQXHVWURJpQHURHVXQ UHVWRGHKXHVRPD[LODU\WLHQHXQDDQWLJHGDGFHUFDQDD ORV0DDXQTXHQRHVKDVWDKDFH0DTXHORVIyVLOHV VRQORVX¿FLHQWHPHQWHDEXQGDQWHV\VLJQL¿FDWLYRV$OD SULPHUD HVSHFLH KXPDQD VH OH GHQRPLQD Homo habilis \ SUHVHQWD FRPR SULQFLSDO UDVJR DQDWyPLFR XQ FHUHEUR QHWDPHQWHPD\RUTXHHOGHVXVDQWHSDVDGRVQRKXPDQRV $UVXDJD0DUWtQH] /D FODYH GH OD VXSHUYLYHQFLD GH ORV SULPHURV KXPDQRV QR HVWXYR HQ XQ FDPELR DQDWyPLFR TXH OH IDFXOWDUDSDUDFDPELDUVXGLHWD6HWUDWyGHXQLQYHQWR HO SULPHUR GH QXHVWUD KLVWRULD DO TXH FRQRFHPRV JHQpULFDPHQWHFRPRWDOODGHODSLHGUD)XHODREWHQFLyQ GH ODVFDV FRQ DULVWDV TXHEUDQGR ORV FDQWRV OR TXH KDELOLWy D ORV SULPHURV KXPDQRV SDUD UDVJDU OD JUXHVD SLHOGHORVJUDQGHVXQJXODGRV\SDUDFRUWDUORVP~VFXORV \WHQGRQHVGHODVFDUFDVDVGHORVDQLPDOHVPXHUWRV/D WDOODGHODSLHGUDSURSLFLyHOFDPELRGHQLFKRHFROyJLFR TXHSHUPLWLyODVXSHUYLYHQFLDGHQXHVWUROLQDMHHLQLFLy OD HYROXFLyQ FXOWXUDO GH OD +XPDQLGDG 6H WUDWD GH XQ IHQyPHQR H[FOXVLYR GH QXHVWUD HVWLUSH TXH KD LGR JDQDQGRLPSRUWDQFLDIUHQWHDODHYROXFLyQELROyJLFD\HV HOTXHKDSHUPLWLGRDORVVHUHVKXPDQRVKDELWDUFRQp[LWR HQWRGRVORVHFRVLVWHPDVWHUUHVWUHVGHOSODQHWD Vulnerabilidad en la evolución humana: crisis en el poblamiento de Europa +DVWDORVGHVFXEULPLHQWRVUHDOL]DGRVHQOD6LHUUD GH$WDSXHUFDDPHGLDGRVGHORVDxRVGHOSDVDGRVLJOR ;;0DUWtQH]SDUHFtD¿UPHPHQWHHVWDEOHFLGRTXH ODKXPDQLGDGQRKDEtDDUULEDGRDQXHVWURFRQWLQHQWHKDVWD HO¿ORGHOPHGLRPLOOyQGHDxRV/RVUHVXOWDGRVGHPiV GHXQVLJORGHH[FDYDFLRQHVHQGHFHQDVGH\DFLPLHQWRV DUTXHROyJLFRV DVt OR DWHVWLJXDEDQ 6LQ HPEDUJR HO GHVFXEULPLHQWR GH IyVLOHV KXPDQRV H LQGXVWULD OtWLFD DVRFLDGDHQ$WDSXHUFDKDGHPRVWUDGRTXHORVKXPDQRV OOHJDURQD(XURSDKDFHDOPHQRV0D/DYDQJXDUGLD GHORVGHEDWHVVREUHHOSREODPLHQWRGHQXHVWURFRQWLQHQWH VHFHQWUDDKRUDHQGHWHUPLQDUVLGLFKRSREODPLHQWRIXH FRQWLQXRHQHOWLHPSRRVLVHLQWHUUXPSLyHQRFDVLRQHV (Q ORV \DFLPLHQWRV HXURSHRV FRPSUHQGLGRV HQWUHKDFHFHUFDGH0D\DOUHGHGRUGH0DSXHGHQ GLVWLQJXLUVH GRV IDVHV FDUDFWHUL]DGDV FDGD XQD SRU XQD PDQHUDGLIHUHQWHGHSURGXFLUODLQGXVWULDOtWLFDR0RGR WHFQROyJLFR(O0RGRUHSUHVHQWDODIRUPDPiVVHQFLOOD GHWDOODUODSLHGUD\DSDUHFLyHQÈIULFDKDFHPiVGH0D 3RUVXSDUWHHO0RGRHVPiVHODERUDGR\VHRULJLQy WDPELpQHQHOFRQWLQHQWHDIULFDQRKDFHDOJRPHQRVGH 0D$PEURVH (O SUREOHPD UDGLFD HQ TXH HO 0RGR DSDUHFH V~ELWDPHQWH HQ (XURSD KDFH DOUHGHGRU GH DxRVDVRFLDGRDODSUHVHQFLDGHHomo heidelbergensis PLHQWUDV TXH ORV \DFLPLHQWRV PiV DQWLJXRV SUHVHQWDQ ~WLOHVGHO0RGR\HVWiQDVRFLDGRVDHomo antecessor ([LVWHSXHVXQDPSOLRGHVIDVHHQWUHODOOHJDGDGHDPERV PRGRVWHFQROyJLFRVD(XURSD$GHPiVHQWUHORV~OWLPRV \DFLPLHQWRV FRQ 0RGR KDFH DOUHGHGRU GH DxRV \ ORV SULPHURV FRQ 0RGR VREUH ORV 83 DxRVKD\XQLQWHUYDORGHFDVLDxRVVLQUHJLVWUR GHRFXSDFLyQKXPDQDHQ(XURSD 6LQHPEDUJRH[LVWHRWURSXQWRGHYLVWDVHJ~QHO FXDOH. heidelbergensisQRVHUtDXQDHVSHFLHLQPLJUDQWH VLQR TXH VH KDEUtD RULJLQDGR SRU HYROXFLyQ ORFDO GHVGH ODVSREODFLRQHVGHH. antecessor/DV~ELWDDSDULFLyQGHO 0RGRHQ(XURSDKDFHPHGLRPLOOyQGHDxRVHVGLItFLO GHH[SOLFDUGHVGHHVWDKLSyWHVLV\KDEUtDTXHDGPLWLUXQ GREOH RULJHQ LQGHSHQGLHQWH HQ ÈIULFD KDFH FHUFD GH 0D\HQ(XURSDKDFH0DSDUDHVWDPDQHUDGHWDOODU ODSLHGUD3DUDORVSDUWLGDULRVGHHVWDVHJXQGDH[SOLFDFLyQ OD DXVHQFLD GH UHJLVWUR DUTXHROyJLFR \ SDOHRQWROyJLFR GHODKXPDQLGDGHQ(XURSDHQWUHKDFH\0DQR LQGLFDUtD HO GHVSREODPLHQWR GHO FRQWLQHQWH VLQR TXH VH GHEHUtDVHQFLOODPHQWHDXQVHVJRGHPXHVWUHRGHGLFKRV UHJLVWURV D OD pSRFD HQ OD TXH ORV KXPDQRV SDUHFHQ HYDSRUDUVH GHOUHVWRGHOFRQWLQHQWHWDPSRFRKD\SUHVHQFLDKXPDQD UHJLVWUDGD HQ *UDQ 'ROLQD 'H KHFKR VRQ ORV ~QLFRV QLYHOHVGHO\DFLPLHQWRVLQHYLGHQFLDVGHGLFKDSUHVHQFLD (VWHKHFKRFRQFXHUGDFRQODLGHDGHTXHHQHVDpSRFDORV KXPDQRV KDEtDQ GHVDSDUHFLGR GHO FRQWLQHQWH HXURSHR $XQTXH WDPELpQ SRGUtD RFXUULU TXH MXVWR HQWRQFHV OD FXHYDQRIXHUDDFFHVLEOHSDUDORVKXPDQRV\TXHSRUHVR VHDQ ORV ~QLFRV QLYHOHV VLQ RFXSDFLyQ KXPDQD GH *UDQ 'ROLQD (QHVWHFRQWH[WRHVPX\LQWHUHVDQWHGHVWDFDUTXH DXQTXHQRKD\DUHJLVWURGHRFXSDFLyQKXPDQDGHODFXHYD HQGLFKRVQLYHOHVQLDSHQDVUHVWRVGHPDFURPDPtIHURV VXVVHGLPHQWRVVRQULFRVHQIyVLOHVGHPLFURPDPtIHURV /DV LQYHVWLJDFLRQHV UHDOL]DGDV PXHVWUDQ TXH MXVWR HQ HVRV QLYHOHV VLQ RFXSDFLyQ KXPDQD VH UHÀHMD XQ LPSRUWDQWH FDPELR HQ OD HVWUXFWXUD GH ODV FRPXQLGDGHV GHPLFURPDPtIHURVTXHVHKDFHQDOJRPHQRVGLYHUVDV\ SDVDQGHHVWDUGRPLQDGDVSRUHVSHFLHVWtSLFDVGHPHGLRV DELHUWRV \ K~PHGRV D SUHGRPLQDU RWUDV FDUDFWHUtVWLFDV GH PHGLRV WDPELpQ DELHUWRV SHUR PiV VHFRV &XHQFD %HVFyVHWDO)LJXUD(VPX\VXJHVWLYR SHQVDU TXH HVWH FDPELR HQ ODV FRQGLFLRQHV DPELHQWDOHV ELHQ SXGR HVWDU UHODFLRQDGR FRQ OD GHVDSDULFLyQ GH H. antecessor GH (XURSD \ VX SRVWHULRU VXVWLWXFLyQ SRU H. heidelbergensis. (Q HVWH FRQWH[WR HO \DFLPLHQWR GH *UDQ 'ROLQDHQOD6LHUUDGH$WDSXHUFDRIUHFHXQDH[FHOHQWH RSRUWXQLGDG SDUD FRQWUDVWDU DPEDV RSLQLRQHV (VWH \DFLPLHQWR FRQVWLWX\H XQ FDVR H[FHSFLRQDO HQ (XURSD SXHV FRQWLHQH XQ UHJLVWUR HVWUDWLJUi¿FR PX\ FRPSOHWR TXHDEDUFDGHVGHKDFHFHUFDGH0DKDVWDKDFHDOUHGHGRU GHDxRVHLQFOX\HQLYHOHVFRQRFXSDFLyQKXPDQD FRQ HO 0RGR \ QLYHOHV GH RFXSDFLyQ FRQ HO 0RGR 3XHV ELHQ HQ ORV QLYHOHV LQWHUPHGLRV FRUUHVSRQGLHQWHV 3HUR HVWD FULVLV QR IXH OD ~QLFD TXH KXELHURQ GHDIURQWDUODVSREODFLRQHVKXPDQDVHXURSHDVDORODUJR GHO3OHLVWRFHQR+DFHXQRVDxRVFRPHQ]yHQHO KHPLVIHULRQRUWHODIDVHPiVIUtDGHOD~OWLPDJODFLDFLyQ 'XUDQWH HVWD pSRFD ODV HVWHSDV LQYDGLHURQ ODV WLHUUDV GHODPD\RUSDUWHGH(XURSDTXHVXIULHURQLPSRUWDQWHV FDPELRVHFROyJLFRV'XUDQWHHVHSHULRGRVHH[WLQJXLHURQ ORVQHDQGHUWDOHV\HOHomo sapiensVHHQVHxRUHyGHODV WLHUUDVGH(XURSD 8QD SRVLEOH H[SOLFDFLyQ SDUD HVWD VLWXDFLyQ HV DGPLWLUTXHKXERGRVHSLVRGLRVGLIHUHQWHVGHSREODPLHQWR GH(XURSD(OSULPHURSURWDJRQL]DGRSRUH. antecessor TXH WUDtD FRQVLJR HO 0RGR \ HO VHJXQGR OOHYDGR D FDERSRUH. heidelbergensisFRQHO0RGR(ODPSOLR LQWHUYDORWHPSRUDOVLQDSHQDV\DFLPLHQWRVFRQSUHVHQFLD KXPDQDH[LVWHQWHHQWUHDPERVHSLVRGLRVFRUUHVSRQGHUtD DXQPRPHQWRGHGHVSREODPLHQWRGH(XURSD Figura 2 Figura 2. Riqueza relativa de especies de micromamíferos en el yacimiento de Gran Dolina (Sierra de Atapuerca, Burgos). NMI: Número Mínimo de Individuos. Fuente: Elaborado a partir de Cuenca-Bescós et al. (2005). 84 &OiVLFDPHQWHODGHVDSDULFLyQGHORVQHDQGHUWDOHV VHKDDWULEXLGRDXQDVXSXHVWDLQIHULRULGDGPHQWDO\VRFLDO IUHQWHDODVSREODFLRQHVGHH. sapiensTXHSURFHGtDQGH ÈIULFD\DQWHODVTXHQRSXGLHURQFRPSHWLU6LQHPEDUJR HVWDLGHDVHKDYLVWRGHVD¿DGDHQORV~OWLPRVDxRVSRUXQ QXHYR SXQWR GH YLVWD TXH VRVWLHQH TXH ODV FDSDFLGDGHV PHQWDOHV GH ORV QHDQGHUWDOHV QR IXHURQ PX\ GLIHUHQWHV GH ODV GH ORV DFWXDOHV KXPDQRV \ TXH IXHURQ IDFWRUHV HFROyJLFRVGHWHUPLQDGRVSRUFDPELRVFOLPiWLFRVUiSLGRV HLQWHQVRVORVTXHRFDVLRQDURQHOGHFOLYH\GHVDSDULFLyQ GH ORV QHDQGHUWDOHV )LQOD\VRQ \ &DUULyQ 2WURV DXWRUHVD~QUHFRQRFLHQGRODLPSRUWDQFLDGHORVIDFWRUHV FOLPiWLFRV\HFROyJLFRVHQHOSURFHVRGHH[WLQFLyQGHORV QHDQGHUWDOHV VLJXHQ UHVHUYDQGR XQ SDSHO SURWDJRQLVWD D OD FRPSHWHQFLD FRQ ORV KXPDQRV PRGHUQRV SXHV RSLQDQ TXH VLQ OD SUHVLyQ GH pVWRV ORV QHDQGHUWDOHV KDEUtDQVREUHYLYLGRDODVFULVLVDPELHQWDOHVGHOD~OWLPD JODFLDFLyQ%DQNVHWDO (Q FXDOTXLHU FDVR VHD FRQ HO FRQFXUVR GHO H. sapiensRVLQpOQDGLHGXGDHQODDFWXDOLGDGGHTXH IXHURQODVH[WUHPDV\UiSLGDVÀXFWXDFLRQHVFOLPiWLFDVGH ¿QDOHVGHO3OHLVWRFHQRVXSHULRUODFDXVDSULQFLSDOGHOD GHVDSDULFLyQGHODVSREODFLRQHVQHDQGHUWDOHV Impacto en la evolución humana: la revolución neolítica H. sapiensHVTXL]iODPiVSHFXOLDUGHWRGDVODVHVSHFLHV KXPDQDV&RPSDUDGDFRQODVGHPiVGHVWDFDSRUWHQHUHO FXHUSRPiVHVWUHFKRHOFUiQHRPiVHVIpULFR\VREUHWRGR OD FDUD PiV SHTXHxD \ DSHQDV VREUHVDOLHQWH 1XHVWUR FRPSRUWDPLHQWRWDPELpQSUHVHQWDDVSHFWRVWDQQRYHGRVRV FRPRVRQHODUWH\XQDJUDQYHUVDWLOLGDGWHFQROyJLFDTXH LQFOX\HODLQYHQFLyQGHDUPDVGHODUJRDOFDQFHFRPRORV SURSXOVRUHV \ ORV DUFRV $GHPiV QXHVWUD RUJDQL]DFLyQ VRFLDO HV VHJXUDPHQWH WDPELpQ PXFKR PiV FRPSOHMD TXHODGHOUHVWRGHHVSHFLHVKXPDQDV7RGRHOORQRVKD IDFXOWDGRSDUDSREODUHQPHQRVGHFLQFXHQWDPLODxRV WRGRVORVHFRVLVWHPDVGHOSODQHWD)LQOD\VRQ(VWD JUDQYHUVDWLOLGDGHFROyJLFDLQGLFDXQDUHODWLYDOLEHUDFLyQ GH OD LQÀXHQFLD GH ODV FRQGLFLRQHV DPELHQWDOHV 3RU SULPHUD YH] HQ OD KLVWRULD GH OD 9LGD XQD HVSHFLH VH H[WLHQGHFRQp[LWRGHVGHORVERVTXHVGHOHFXDGRUKDVWD HOÈUWLFR +DFH DOUHGHGRU GH GLH] PLO DxRV FRLQFLGLHQGR FRQ HO ¿QDO GH OD ~OWLPD JODFLDFLyQ ORV KXPDQRV UHDOL]DURQ OD PD\RU UHYROXFLyQ GH VX KLVWRULD GHVGH OD LQYHQFLyQGHODWDOODGHODSLHGUD6HWUDWDGHOFRPLHQ]R GHODDJULFXOWXUD\GHODJDQDGHUtDTXHVXSXVRHOLQLFLR GHODHFRQRPtDGHSURGXFFLyQ\HO¿QDOGHODHFRQRPtD EDVDGD HQ OD FD]D \ OD UHFROHFFLyQ$ HVWH PRPHQWR GH OD KLVWRULD GH OD KXPDQLGDG OR OODPDPRV 1HROtWLFR (Q ODUHYROXFLyQQHROtWLFDVHHQFXHQWUDQHOFRPLHQ]RGHOD VHGHQWDUL]DFLyQ \ HO RULJHQ GH ODV FLXGDGHV \ VH LQLFLy HQWRQFHV OD H[WUDRUGLQDULD H[SDQVLyQ GHPRJUi¿FD TXH FRQWLQ~DHQQXHVWURVGtDV (O QXHYR WLSR GH HFRQRPtD GHWHUPLQy TXH ORV KXPDQRVFRPHQ]iUDPRVDH[SORWDUVLVWHPiWLFDPHQWHORV WHUULWRULRV PRGL¿FiQGRORV GUiVWLFDPHQWH SDUD SRQHUORV D QXHVWUR VHUYLFLR (O IXHJR SDUD DFODUDU ORV ERVTXHV ODV REUDV KLGUiXOLFDV SDUD GRPDU ODV DJXDV GH ORV UtRV OD FUHFLHQWH DFXPXODFLyQ GH GHVKHFKRV OD H[WLQFLyQ VHOHFWLYD GH HVSHFLHV FRPSHWLGRUDV \ HO IDYRUHFLPLHQWR GHRWUDVFRQVLGHUDGDV~WLOHVIXHURQ\VRQDOJXQDVGHODV KHUUDPLHQWDV FRQ ODV TXH ORV KXPDQRV KHPRV DOWHUDGR GUiVWLFDPHQWHORVHFRVLVWHPDVGHOSODQHWD3HURTXL]iHO FDPELRPiVLPSRUWDQWHQRIXHUDGHtQGROHHFRQyPLFR6H SURGXMRRWUDWUDQVIRUPDFLyQPiVSURIXQGDTXHKDWHQLGR JUDYtVLPDV FRQVHFXHQFLDV SDUD HO UHVWR GH FULDWXUDV GHO SODQHWD &RQ OD UHYROXFLyQ QHROtWLFD GHMDPRV GH YHUQRV FRPR KLMRV GH OD7LHUUD \ SDVDPRV D FRQVLGHUDUQRV VXV SURSLHWDULRV«&UHFHG\PXOWLSOLFDURV 1XHVWUDHVSHFLHGHVFHQGLHQWHGHXQODUJROLQDMH FX\D KLVWRULD HVWXYR GHWHUPLQDGD SRU ODV FRQGLFLRQHV PHGLRDPELHQWDOHV VH FRQVLGHUD KR\ DO PDUJHQ GH OD 1DWXUDOH]D \ KD SDVDGR D HQJURVDU OD PDJUD OLVWD GH HVSHFLHVFDSDFHVGHSURGXFLUXQLPSDFWRHFROyJLFRJOREDO VREUHHOSODQHWD,QFOXVRGHPRGL¿FDUODFRPSRVLFLyQGH VXDWPyVIHUDWDOFRPRKLFLHURQHQORVDOERUHVGHQXHVWUR PXQGRODVFLDQREDFWHULDV3HURKD\DOJRIUXWRGHQXHVWUD KLVWRULDHYROXWLYDTXHQXQFDKDEtDHVWDGRDODOFDQFHGH QLQJ~QRWURRUJDQLVPRHQODKLVWRULDGHOD9LGD6RPRVOD ~QLFDFULDWXUDFRQODFDSDFLGDG\ODUHVSRQVDELOLGDGGH HOHJLU(QQXHVWUDPDQRHVWiGHFLGLUTXHQXHVWURIXWXURHO GHQXHVWUDVKLMDVHKLMRV\HOGHQXHVWURPXQGRVHDPHMRU TXHHOSDVDGR 5HIHUHQFLDVELEOLRJUiÀFDV $UVXDJD-/0DUWtQH],/DHVSHFLHHOHJLGD7HPDV GHKR\0DGULG $PEURVH6+3DOHROLWKLF7HFKQRORJ\DQG+XPDQ (YROXWLRQScience %DQNV:(G¶(UULFR)3HWHUVRQ$7.DJH\DPD06LPD $6iQFKH]*RxL0) 1 H D Q G H U W K D O ([WLQFWLRQE\&RPSHWLWLYH([FOXVLRQPLoS ONE H %REH5%HKUHQVPH\HU$.&KDSPDQ5()DXQDO FKDQJHHQYLURQPHQWDOYDULDELOLW\DQGODWH3OLRFHQH KRPLQLQ HYROXWLRQ Journal of Human Evolution &HUOLQJ7(0EXD(.LUHUD)00DQWKL).*ULQH)( /HDNH\0*6SRQKHLPHU0 8QR.7'LHW RIParanthropus boiseiLQWKHHDUO\3OHLVWRFHQHRI (DVW$IULFDProceedings of the National Academy of Sciences of USA &XHQFD%HVFyV * 0DUtQ$UUR\R $% 0DUWtQH] , *RQ]iOH]0RUDOHV056WUDXV/*5HODWLRQ VKLS EHWZHHQ 0DJGDOHQLDQ VXEVLVWHQFH DQG HQYL URQPHQWDO FKDQJH 7KH PDPPDOLDQ HYLGHQFH IURP (O 0LUyQ 6SDLQ Quaternary Intern tional &XHQFD%HVFyV*0HOHUR5XELR05RIHV-0DUWtQH], $UVXDJD-/%ODLQ+$ /ySH]*DUFtD -0 &DUERQHOO ( %HUPXGH] GH &DVWUR -0 7KH (DUO\± 0LGGOH3OHLVWRFHQHHQYLURQPHQWDODQG FOLPDWLFFKDQJHDQGWKHKXPDQH[SDQVLRQ LQ:HVWHUQ(XURSH$FDVHVWXG\ZLWKVPDOOYHUWH EUDWHV*UDQ'ROLQD$WDSXHUFD6SDLQJournal of Human Evolution &XHQFD%HVFyV * 5RIHV - *DUFLD3LPLHQWD - (QYLURQPHQWDO FKDQJH DFURVV WKH HDUO\PLGGOH 3OHLVWRFHQHWUDQVLWLRQVPDOOPDPPDOLDQHYLGHQFH 85 IURPWKH7ULQFKHUD'ROLQDFDYH$WDSXHUFD6SDLQ (Q+HDG0-*LEEDUG3/HGLWRUHVEarly-Middle Pleistocene Transitions: The Land-Ocean Evidence*HRORJLFDO 6RFLHW\ RI /RQGRQ 6SHFLDO 3XEOLFDWLRQ/RQGUHVSS )LQOD\VRQ&El sueño del neandertal. Por qué se extinguieron los neandertales y nosotros sobrevivimos(GLWRULDO&UtWLFD%DUFHORQD )LQOD\VRQ & &DUULyQ -6 5DSLG HFRORJLFDO WXUQ RYHU DQG LWV LPSDFW RQ 1HDQGHUWKDO DQG RWKHU KX PDQSRSXODWLRQVTrends in Ecology and Evolution 0DUWtQH],El primate que quería volar(VSDVD 0DGULG 8QJDU 36 6SRQKHLPHU 0 7KH 'LHWV RI (DUO\ +RPLQLQVScience 86