RESEARCH ARTICLE
Ch onological eassessmen o he Middle o
Uppe Paleoli hic ansi ion and Ea ly Uppe
Paleoli hic cul u es in Can ab ian Spain
Ana B. Ma ı
´n-A oyo
1,2
*, Joseba Rios-Ga aiza
3
, Law ence G. S aus
4
, Jenni e R. Jones
1
,
Ma co de la Rasilla
5
, Manuel R. Gonza
´lez Mo ales
1
, Michael Richa ds
6
, Jesu
´s Al una
7
,
Ko o Ma iezku ena
7
, Da id Ocio
8
1Ins i u o In e nacional de In es igaciones P ehis o
´ icas de Can ab ia, (Uni e sidad de Can ab ia,
San ande , Gobie no de Can ab ia), San ande , Spain, 2Le e hulme Cen e o E olu iona y S udies,
Depa men o A chaeology, Uni e si y o Camb idge, Camb idge, Uni ed Kingdom, 3Cen o Nacional de
In es igacio
´n de la E olucio
´n Humana, Bu gos, Spain, 4Depa men o An h opology, MSC01 1040,
Uni e si y o New Mexico, Albuque que, New Mexico, Uni ed S a es o Ame ica, 5Depa amen o de His o ia,
Uni e sidad de O iedo, O iedo, Spain, 6Depa men o A chaeology, Simon F ase Uni e si y, Bu naby,
B i ish Columbia, Canada, 7Cen o de Conse acio
´n e In es igacio
´n de los Ma e iales A queolo
´gicos y
Paleon olo
´gicos de Gipuzkoa, Donos ia/San Sebas ia
´n, Spain, 8Depa men o Ci il and En i onmen al
Enginee ing, Impe ial College London, London, Uni ed Kingdom
*[email p o ec ed]
Abs ac
Me hodological ad ances in da ing he Middle o Uppe Paleoli hic ansi ion p o ide a be e
unde s anding o he eplacemen o local Neande hal popula ions by Ana omically Mode n
Humans. Today we know ha his eplacemen was no a single, pan-Eu opean e en , bu
a he i ook place a di e en imes in di e en egions. Thus, local condi ions could ha e
played a ole. Ibe ia ep esen s a signi ican mac o- egion o s udy his p ocess. No he n
A lan ic Spain con ains e idence o bo h Mous e ian and Ea ly Uppe Paleoli hic occupa-
ions, al hough mos o hem a e no p ope ly da ed, hus hinde ing he chances o an
adequa e in e p e a ion. He e we p esen 46 new adioca bon da es conduc ed using ul a-
il a ion p e- ea men me hod o an h opogenically manipula ed bones om 13 si es in he
Can ab ian egion con aining Mous e ian, Au ignacian and G a e ian le els, o which 30
a e conside ed ele an . These da es, alongside p e iously epo ed ones, we e in eg a ed
in o a Bayesian age model o econs uc an absolu e imescale o he ansi ional pe iod.
Acco ding o i , he Mous e ian disappea ed in he egion by 47.9–45.1ka cal BP, while he
Cha
ˆ elpe onian las ed be ween 42.6k and 41.5ka cal BP. The Mous e ian and Cha
ˆ elpe o-
nian did no o e lap, indica ing ha he la e migh be ei he in usi e o an o shoo o he
Mous e ian. The new ch onology also sugges s ha he Au ignacian appea s be ween
43.3–40.5ka cal BP o e lapping wi h he Cha
ˆ elpe onian, and ended a ound 34.6–33.1ka
cal BP, a e he G a e ian had al eady been es ablished in he egion. This e idence indi-
ca es ha Neande hals and AMH co-exis ed <1,000 yea s, wi h he ca ea ha no diagnos-
ic human emains ha e been ound wi h he la es Mous e ian, Cha
ˆ elpe onian o ea lies
Au ignacian in Can ab ian Spain.
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 1 / 20
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OPEN ACCESS
Ci a ion: Ma ı
´n-A oyo AB, Rios-Ga aiza J, S aus
LG, Jones JR, de la Rasilla M, Gonza
´lez Mo ales
MR, e al. (2018) Ch onological eassessmen o
he Middle o Uppe Paleoli hic ansi ion and Ea ly
Uppe Paleoli hic cul u es in Can ab ian Spain.
PLoS ONE 13(4): e0194708. h ps://doi.o g/
10.1371/jou nal.pone.0194708
Edi o : John P. Ha , New Yo k S a e Museum,
UNITED STATES
Recei ed: Augus 11, 2017
Accep ed: Ma ch 8, 2018
Published: Ap il 18, 2018
Copy igh : ©2018 Ma ı
´n-A oyo e al. This is an
open access a icle dis ibu ed unde he e ms o
he C ea i e Commons A ibu ion License, which
pe mi s un es ic ed use, dis ibu ion, and
ep oduc ion in any medium, p o ided he o iginal
au ho and sou ce a e c edi ed.
Da a A ailabili y S a emen : All ele an da a a e
wi hin he pape and i s Suppo ing In o ma ion
iles.
Funding: This esea ch has been suppo ed by he
Eu opean Commission h ough FP7-PEOPLE-
2012-CIG (Re : 322112), he Spanish Minis y o
Economy and Compe i i eness (HAR2012-33956
and RYC-2011-00695), Can ab ia Campus
In e na ional and Uni e si y o Can ab ia o ABMA.
The Ins i u o In e nacional de In es igaciones
In oduc ion
The Middle o Uppe Paleoli hic ansi ion is gene ally associa ed wi h he eplacemen o
local Neande hal popula ions by Ana omically Mode n Humans (AMH) in wes e n Eu asia
[1]. One o he mos deba ed issues is he p ecise ch onology o he eplacemen o he popula-
ion and i s po en ial ela ionship wi h majo biological, en i onmen al and cul u al e en s.
Due o ecen e isions o a chaeological, ch onological and geog aphical da a, he spa io- em-
po al delimi a ion o he Middle o Uppe Paleoli hic ansi ion in cen al and wes e n Eu ope
has imp o ed [1–5]. Me hodological de elopmen s in adioca bon da ing ha emo e young
ca bon con amina ion om old o ganic samples [6–7] ha e been c ucial o es ablishing accu-
a e ch onologies, especially a he adioca bon limi a ound ~50,000 yea s ago, when he Mid-
dle o Uppe Paleoli hic ansi ion ook place. Thus, he e-da ing o se e al key si es in Eu ope
has p oduced olde ages han p e iously hough [1]. The iming o Neande hal disappea ance
in no hwes e n Ibe ia (~48-46ka uncal BP) and I aly (~44-42ka cal BP) and he age o so-
called “ ansi ional indus ies” (Cha
ˆ elpe onian and Uluzzian, bo h 44 o 40ka cal BP) ha e
been ecen ly delimi ed [1,3]. While he a ibu ion o Neande hals o he Cha
ˆ elpe onian
in no he n Ibe ia and sou hwes e n F ance is seldom con es ed (bu see [8]), he ques ion o
he make s o he Uluzzian is s ill deba ed. Human deciduous ee h ound in Sou he n I aly
(G o a del Ca allo) sugges ha AMH we e he make s o he Uluzzian [9], whe eas o he
esea che s con inue o a gue o Neande hal au ho ship [10]. Those “T ansi ional echno-
complexes” (especially he Cha
ˆ elpe onian), ha e been in e p e ed as an e idence o a sho
pe iod o coexis ence be ween local and immig an human popula ions [1,11], wi h he ea ly
phases o he Au ignacian complex (P o o-Au ignacian and ea ly Au ignacian) being a ib-
u ed o he i s mode n humans in Eu ope, as ea ly as 42 ka cal BP, du ing GI10. In Ibe ia, he
end o he Mous e ian has been shown o ha e been ea lie in he No h han once hough ,
while he Ea ly Uppe Paleoli hic in he Sou h (G a e ian) is olde han was p e iously
belie ed [12]. Thus, he hypo hesis o a e y ea ly appea ance o Au ignacian in no he n Ibe-
ia is no longe suppo ed [2,4].
He e, we ocus on he Can ab ian egion in no he n Ibe ia, whe e he a chaeology, s a ig-
aphy and ch onology o se e al key si es—El Cas illo, Labeko Koba, La Viña, Esquilleu and
Mo ı
´n—ha e been ecen ly e iewed [2,5,13], p o iding an ini ial amewo k o he Middle
o Uppe Paleoli hic ansi ion. Howe e , u he esea ch a mo e ep esen a i e si es in his
egion wi h Mous e ian, Cha
ˆ elpe onian, Au ignacian and G a e ian ma e ials, ha ei he
lacked adioca bon da es o had da es ob ained in he la e 20 h cen u y be o e he de elop-
men o igo ous p e- ea men me hods, is needed o secu ely es ablish he ch onology by
including mo e a chaeological si es. As pa o a wide p ojec o econs uc he clima e and
local and egional en i onmen al condi ions ha la e Neande hals and ea ly AMH aced in
he egion du ing Ma ine Iso opic S age 3 (MIS3), using s able iso opic analyses (δ
13
C, δ
15
N,
δ
18
O and δ
34
S), i s we pe o med a comple e e iew o he ch onological da a om 13
egional si es ha included 28 Middle and Ea ly Uppe Paleoli hic a chaeological le els. Ou o
51 samples ha we e p ocessed o adioca bon da ing, 5 ailed and he emaining 46 ecei ed
an AMS measu emen using new ul a il a ion me hod ha is mo e igo uous in emo ing
possible bone collagen con amina ion, om con empo aneous si es which ei he lacked da es
o had p e iously p oduced p oblema ic da es.
The new
14
C adioca bon da es ob ained in his p ojec , in combina ion wi h o he a ailable
ul a il e ed da es om he egion, ha e allowed he de elopmen o an independen , adio-
me ic ch onology o p e iously unda ed si es in he Can ab ian egion, which had un il now
been a ibu ed only h ough s a ig aphic posi ion and/o ma e ial cul u e cha ac e is ics o
he Mous e ian, Cha
ˆ elpe onian, Au ignacian o G a e ian echno-complexes. Re-da ing o
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 2 / 20
P ehis o
´ icas de Can ab ia is sponso ed by he
Uni e si y o Can ab ia, he Go e nmen o
Can ab ia and Banco San ande . This pape was
mainly w i en du ing a esea ch s ay by one o us
(ABMA) a he Le e hulme Cen e o E olu iona y
S udies, Depa men o A chaeology, Uni e si y o
Camb idge unded by Spanish Minis y o
Educa ion, Cul u e and Spo s (Jose de Cas illejo
P og am, Re no. CAS15_00054). The unde s had
no ole in s udy design, da a collec ion and
analysis, decision o publish, o p epa a ion o he
manusc ip .
Compe ing in e es s: The au ho s ha e decla ed
ha no compe ing in e es s exis .
sequences ha had p e iously been da ed using con en ional o ea ly AMS
14
C me hods was
unde aken, p oducing signi ican changes in p e ious in e p e a ions o ele an si es, some-
hing ha has also been e i ied by e iewing he li hic echnocomplexes. The esul s p o ide
g ea e empo al p ecision o he Middle o Uppe Paleoli hic ansi ion in he Can ab ian
Region by building a Bayesian model using samples ea ed wi h a obus p e- ea men
me hodology. The model includes new Mous e ian modelled da es wi hin he ange o he
calib a ion cu e and de ines he bounda ies o he s a and end o each o he Ea ly Uppe
Paleoli hic echno-complexes.
This esea ch p esen s new possibili ies o add essing he iming o he c i ical p ocesses o
cul u al change, local ex inc ions and popula ion eplacemen ha happened du ing he Mid-
dle-Uppe Paleoli hic ansi ion in no he n Spain and mo e gene ally in Eu ope. The da ing
esul s epo ed he e will also se e o discuss he s a ig aphic in eg i y and cul u al a ibu-
ion o he si es analysed.
Ma e ial
Thi een si es we e da ed as pa o his p ojec , including a o al o h ee Middle Paleoli hic
and wen y- i e ea ly Uppe Paleoli hic a chaeological le els. These si es a e loca ed in he
mode n-day egions o As u ias in he wes , Can ab ia in he cen e and he Basque Coun y
in he eas o he Can ab ian Region (Fig 1). These a chaeological le els we e selec ed because
o hei a ibu ion o he Mous e ian (n = 3), Cha
ˆ elpe onian (n = 1), Au ignacian (n = 11)
and G a e ian (N = 13) echno-complexes, p e iously de e mined based on hei s a ig aphic
posi ion, ma e ial cul u e, a ailable da ing o a combina ion o hese ac o s. The Mous e ian
le els da ed we e Amalda VII [14], Axlo IV [15] and Llonı
´n VI [16–20], while he only Cha
ˆ el-
pe onian le el was Ekain Xa [21]. The Au ignacian le els we e Ai zbi a e (Ca e III-en ance
a ea) Vb cen e [22], Ekain IXb, Cob an e V and VI [23], El Ruso Ca e I IVb [24], El O e o IV,
V and VI [25], Mo in 7c [26], Co alejos B (2) and C (3) [27]. Finally, he G a e ian le els
Fig 1. Loca ion o si es in he Can ab ian egion, no he n Spain. 1: La Viña, 2: La Gu¨elga, 3: Llonı
´n, 4: Esquilleu, 5: El Cas illo, 6: Co alejos, 7: Ruso I, 8:
Mo ı
´n, 9: Cob an e, 10: El O e o, 11: Mi o
´n, 12: El Cuco, 13: A illo , 14: Axlo , 15: Bolinkoba, 16: Labeko Koba, 17: Ekain, 18: Amalda, 19: Ai zbi a e Ca e III.
h ps://doi.o g/10.1371/jou nal.pone.0194708.g001
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 3 / 20
we e Ai zbi a e (Ca e III-en ance a ea) IV, Va and Vb uppe , Amalda V and VI, Bolinkoba
VI/F [28–29], El Cuco III and Vb [30–31], La Viña VII, VIII, IX and X [32–36] and Llonı
´n V.
O he egional si es wi h con empo aneous cul u al a ibu ions, bu unclea s a ig aphy,
insu icien a chaeological in o ma ion, o hose ha a e cu en ly s ill unde s udy we e no
included he e. We also ook in o conside a ion he adioca bon da es ecen ly published o
he egional si es o La Viña, Esquilleu, La Gu¨elga, El Cas illo, Mo ı
´n, El Mi o
´n, A illo and
Labeko Koba [2–5,37–38] ha ha e been achie ed using he same ul a il a ion me hod as
his s udy, making hem compa able and app op ia e o include in he Bayesian models. We
ha e ini ially accep ed he cul u e-s a ig aphic designa ions o indi idual le els p oposed by
he exca a o s/analys s o he a ious si es. This b ings wi h i he consequence o inco po a -
ing di e ing classi ica o y c i e ia (including di e ing app oaches o “lumping” e sus “spli -
ing” in such designa ions) among he many di e en esea che s who s udied he a ious
a i ac assemblages, some o which con ain mo e de ini i e empo ally/cul u ally bounded
and hus diagnos ic a i ac s han o he s. Howe e , he Bayesian app oach applied he e has he
capaci y o challenge hese p io a ibu ions, p omp ing a ein e p e a ion o he li hic assem-
blages and he s a ig aphy. Whe e misma ches we e iden i ied, si es we e e isi ed, and i nec-
essa y, a ibu ions changed acco ding o ou e iews. In case o doub , da es we e disca ded
o a oid any bias in he conclusions.
51 samples we e p ocessed o adioca bon da ing on bones wi h e idence o human
manipula ion such as an h opogenic ac u es and/o cu ma ks. Sample in o ma ion o each
specimen, including si e, a chaeological le el, con ex , sample numbe , animal species and
ana omical elemen is included in S1 Table. The loca ion o he a chaeological ma e ial in
egional museums and ins i u ions is indica ed in he desc ip ion o each si e in S1 File. This
esea ch implies a ema kable inc ease in he numbe o eliable adiome ic da es in he
egion and pe iod o s udy, which cu en ly comp ises 61 de e mina ions [4,5,12–13] [S3
Table]. The selec ion o he auna samples in museum collec ions was based on he ollowing
c i e ia: 1) s a ig aphic posi ion o he emains a he si e; 2) selec ion o bones p e e ably
di ec ly labelled o assu e hei p o enance wi hin he si e; 3) selec ion o animal bones axo-
nomically and ana omically iden i iable and 4) selec ion o ana omical elemen s wi h clea
an h opogenic ma ks.
Me hods
Radioca bon da ing
P io o adioca bon da ing, a subsample o all specimens was analysed o ob ain δ
13
C and
δ
15
N alues, which also es ablished he s a e o collagen p ese a ion. O he bones p e iously
analysed o s able iso opic analyses, specimens yielding >1% collagen we e selec ed o da ing
o maximise he likelihood o achie ing a success ul adioca bon da e. All samples me he
quali y assu ance c i e ia as shown in S1 Table. Collagen ex ac ion and analysis o adioca -
bon da ing we e unde aken a he Ox o d Radioca bon Accele a o Uni (ORAU). Collagen
was ob ained ollowing he me hod de ailed by B ock e al [39], which in ol es he demine ali-
sa ion o he mine al componen (and any exogenous ca bona es) o d illed bone powe using
0.5M HCL a 5˚C o e nigh , be o e emo al o o ganics and humic acid using 0.1M NaOH
solu ion o 30 minu es a oom empe a e (RT), and a inal wash in 0.5M HCL o 1 hou a
RT. The collagen was gela inised in a 0.001M HCL solu ion o 20 hou s a 70˚C. EZEE™bio-
logical il e s (45–90 μm) we e used o emo e smalle soluble componen s, be o e ul a il a-
ion using cleaned 30 kDa MWCO Vi aspin™15 ul a il e s. Combus ion o he collagen using
an elemen al analyse (ANCA-GSL), linked o an iso ope a io-mass spec ome e (Se con 20–
20) p oduced Ca bon and Ni ogen s able iso ope da a, and samples we e da ed using an
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 4 / 20
Accele a o Mass Spec ome e ollowing con e sion o excess CO
2
in o g aphi e using an
i on ca alys [40].
Bayesian modelling
Indi idual Bayesian age models could be buil o La Viña (S1 Fig), Co alejos (S2 Fig), El
Cuco (S3 Fig), Ekain (S4 Fig) Amalda si es (S5 Fig) and Ai zibi a e (Ca e III) (S6 Fig) using
OxCal4.2 so wa e [41] and he INTCAL13 calib a ion cu e [42]. No ch onological models
we e buil o he o he si es, because o he limi ed numbe o da es. The Bayesian model
enables o modi y he calib a ed P obabili y Dis ibu ion Func ion (PDF) o indi idual da es
based on he exis ing ela i e s a ig aphic and o he ela i e age in o ma ion. Bo h new ul a-
il a ion da es and p e ious ones ecen ly published o he pe iod o s udy in no he n Spain
we e conside ed o comple eness (see S2 Table). The p esence o p oblema ic de e mina ions
ha do no ag ee wi h he p io amewo k was conside ed by adop ing a - ype ou lie model
wi h an ini ial 5% p obabili y o each de e mina ion o be an ou lie . Likewise, a s- ype ou lie
model was used o es he cohe ence o wo adioca bon da es ob ained om he same bone
emain [43]. A esolu ion o 20 yea s was assumed, being a easonable balance be ween
equi ed accu acy and compu a ional cos s, and a sensi i i y es on he ou lie model and
p io bounda y es was conduc ed o ensu e he obus ness o he esul s. A discussion o each
a chaeological si e and cul u al a ibu ion acco ding o he da es a e discussed indi idually in
S1 File. CQL codes a e also epo ed he e (S1 and S2 Codes). An O de unc ion in he OxCal
was used o calcula e he p obabili y ha one PDF p eda ed ano he , p o iding in o ma ion o
assess synch onici y and empo al o e lap o indi idual a chaeological le els and cul u al
phases in each o he six indi idual si es modelled. In all cases, con e gence was g ea e han
95% and he model ag eemen index close o 100% excep in he case o La Viña, whe e i is
s ill clea ly abo e he 60% alidi y h eshold.
Once models o indi idual si es we e conduc ed, modelled da es om each indus y (pos-
e io PDFs) we e g ouped wi hin a single cul u al phase (inse ed as p io s) wi hou assuming
any o de . This was achie ed by placing hem wi hin a single Phase wi h s a and end bound-
a ies. Indi idually calib a ed da es coming om si es we e no indi idual si e model could be
conduc ed (such as Cob an e Le el VI, Llonı
´n V, Bolinkoba VI/F and Mo in 4), as well as
o he exis ing eliable modelled da es in he egion ( om Labeko Koba [4], El Cas illo [5] and
La Viña [13]), we e included. These new uno de ed Bayesian age models enabled de e mining
a egional PDF o he empo al bounda ies be ween Mous e ian, Cha
ˆ elpe onian, Au igna-
cian and G a e ian, as well as an easie compa ison be ween si es. The ad an age o his analy-
sis in compa ison wi h indi idual models is ha i enables u he cons aining he unce ain y
o he bounda ies by assuming ha he onse and disappea ance o cul u es we e egional p o-
cesses a he han localised phenomena. This is easonable gi en he known le el o egional
in e connec ion du ing La e Pleis ocene in no he n Ibe ia. Likewise, i educes he po en ial
bias a one pa icula si e, as pooling oge he se e al si es in one egion se es o inc ease he
eliabili y o he esul s, which can be in e p e ed egionally wi h mo e con idence (see S11
Figu e in [3] and see Figu e 8 in [5]). The di e ence be ween he PDFs o he s a and end
bounda ies was also calcula ed o es ima e he likely du a ion o he phase. The esul s we e
compa ed wi h he G eenland Oxygen Iso opic eco d (NGRIP) [44,45] as a global clima ic
eco d o de ine he di e en cul u al pe iods in he Can ab ian egion.
All Bayesian models we e un 4–5 imes and esul s compa ed o check consis ency. They
disclosed accep able le els o ep oducibili y when compa ed, al hough key bounda y pa ame-
e s we e usually wi hin 50–100 yea s o one ano he wi h epe i ion o he model. This is he
usual accu acy expec ed when using his app oach, and consequen ly, all da es epo ed he e
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 5 / 20

ha e been ounded o he nea es cen u y. Finally, he younges and oldes da es o each cul-
u al pe iod we e emo ed and Bayesian models we e e un o es he dependence o he con-
clusions on hem.
Resul s and discussion
Ou o he 51 da es p ocessed o adioca bon da ing, 46 p oduced success ul AMS measu e-
men s as hey had enough collagen o p o ide iable esul s. Fou samples ailed due o low
collagen yields and one due o a high C:N alue. Collagen was gene ally well p ese ed and
only one sample om El Cuco yielded less han 1% collagen, bu i had a C:N a io o 3.39 was
wi hin he accep able ange o in i o collagen [46]. O he 46 da es, 3 da es we e beyond he
adioca bon limi and 13 showed misma ches wi h hei de ined cul u al a ibu ion p o iding
a ch onology younge han ini ially a ibu ed ei he by cul u al a e ac s ound wi hin ha
le el o p e ious da ing (S1 Table). Consequen ly, in his s udy, a subs an ial da ase o 30 alid
new da es was ob ained and only wi h hose da es Bayesian models we e gene a ed.
The 13 da es ha show disc epancies be ween he cul u al a ibu ions de e mined by he
new da es ob ained and hose based on adi ional analyses o he a chaeological ma e ials co -
espond o 7 o he 13 s udied si es (11 o he 28 le els). This migh be caused by one o se e al
easons such as exca a ion me hods (e.g., ailu e o see o iden i y s a ig aphic di e ences,
a ibu ions based on e y small, no e y diagnos ic a i ac collec ions om limi ed exca a-
ions o es pi s) o cu a o ial issues wi h he aunal ma e ial, as se e al o he egional muse-
ums ha e been mo ed epea edly o di e en loca ions o e se e al decades. Fo each si e,
possible easons o hose inconsis encies a e discussed in S1 File.
Indi idual Bayesian models we e c ea ed o si es whe e samples belonged o mo e han
one laye , such as La Viña (S1 Fig), Co alejos (S2 Fig), El Cuco (S3 Fig), Amalda (S5 Fig) and
Ai zbi a e III (S6 Fig). Fo Ekain wo da es o Le el IXb we e also modelled (S4 Fig). CQL
codes a e p esen ed in S1 and S2 Codes. This allowed, conside ing he s a ig aphy o he si es
as p io in o ma ion o he da es, leading o mo e obus esul s. The obus ness o he esul s
has also been e i ied by means o a sensi i i y es (see S4 Table), which esul ed in ma ginal
changes o he ele an bounda ies be ween cul u es.
In se e al a chaeological le els, ypological ch onologies ha e been challenged by he new
da ing e idence. Thus, in As u ias, he si e o Llonı
´n Le el V, a ibu ed o he G a e ian [16–
20], yielded wo e y di e en esul s: one cohe en wi h an Ea ly G a e ian a ibu ion (~28k
uncal BP) and ano he da e o 20k uncal BP, which could co espond o an ea ly Solu ean o a
e y la e phase o he G a e ian, younge han Mo ı
´n Le el 4 [47] and simila o he AMS
da es o Le el III om Ai zbi a e III [48]. These esul s sugges ei he a much longe o ma-
ion o Le el V (mo e han 8,000 yea s o only 9 cm o s a ig aphy), al hough we canno dis-
miss he possibili y o some dis u bance o admix u e wi h o e lying Solu ean Le el IV which
emains unda ed. In he Au onomous Communi y o Can ab ia, he new ch onology o El
O e o, which was p e iously unda ed, indica es a Magdalenian ch onology (~15–10 ka uncal
BP, and he ea e ) ins ead o an Au ignacian one as adi ionally p oposed [25]. Despi e he
p esence o appa en ly cha ac e is ic a i ac s [25,49–50], his esul u ges cau ion in he da ing
o le els solely using a i ac s ha a e no empo ally diagnos ic. In Le el V o Cob an e, iden i-
ied as Au ignacian, despi e i s lack o cha ac e is ic ma e ial cul u e [51] he ob ained da es
co espond o he Solu ean ins ead (~18k uncal BP). The appa en absence o cha ac e is ic
Au ignacian li hics, and he p esence o a leas one piece wi h pa ial in asi e (Solu ean)
e ouch [51], suppo his a ibu ion. Cob an e Le el VI was in e p e ed as A chaic Au igna-
cian because o he p esence o cha ac e is ic li hic ools, such as Du ou bladele s, la ge “Au i-
gnacian” blades and ca ina ed end-sc ape s, which appea ed alongside Mous e ian-looking
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 6 / 20
ools [51]. New wo da es show an incohe ence be ween hem: one wi h a da e beyond he
adioca bon limi and ano he wi h a con empo aneous egional Ea ly Au ignacian da e.
Cu a ional o s a ig aphical issues could be he eason o his incohe ence as explained in S1
File. In El Cuco, he Le els III and Vb p e iously de ined [30] bo h as G a e ian p o ided
Au ignacian (~35k uncal BP) and Mous e ian (~49k uncal BP) da es, espec i ely. The lowe
le els o his sequence (VII-XIII) ha e been ecen ly eassessed h ough new 14C AMS da ing
on shells and a echnological li hic s udy. These new analyses con i m a Mous e ian a ibu ion
o le els p e iously de ined as Au ignacian and an Au ignacian ch onology o le els de ined
as G a e ian [31]. In El Ruso, Le el IVb o iginally de ined as E ol ed Au ignacian [52], p o-
ided a G a e ian da e (~28k uncal BP). Al hough he small li hic assemblage is no e y
diagnos ic, he p esence o a la -nosed end-sc ape and a double ca ina ed end-sc ape was
men ioned.
In Bizkaia, he G a e ian Le el VI o Bolinkoba p o ided a con empo aneous da e o his
egional pe iod, while a second one is much younge (~10k uncal BP), a ibu able o he
egional Azilian, sugges ing signi ican mixing o cu a ional p oblem as no diagnos ic Azilian
li hic ma e ial has been eco e ed o iden i ied in Le el VI assemblage [53]. In Amalda (Gipuz-
koa), h ee da es p o ided younge da es han expec ed [54]. One sample om Mous e ian
Le el VII p o ided a da e wi hin a G a e ian ime ange (~28k uncal BP), possibly e lec ing a
limi ed admix u e be ween Le els VI and VII [55]. Ano he sample om he G a e ian Le el
VI ga e a medie al da e. I s loca ion close o he ca e en ance, whe e he uppe le els we e
exposed, poin s owa ds he possibili y ha ma e ials om his o ic imes we e inse ed in o
Las Glacial sedimen s. A u he sample om he G a e ian Le el V p o ided a Magdalenian
age (~14k uncal BP), maybe de i ing om unknown p oblems du ing exca a ion o cu a ion.
In Ai zbi a e ca e III Le el Vb (cen e ) yielded a single da e ha is olde han he no ma i e
a ibu ion (likely Ea ly Au ignacian a he han E ol ed Au ignacian) [56], e idencing a pos-
sible p esence o an Ea ly Au ignacian a he base o he le el. In Ekain (Gipuzkoa), Le el Xa,
unda ed, was in e p e ed as a Cha
ˆ elpe onian hun ing camp, wi h a small li hic collec ion,
including a single ypical Cha
ˆ elpe onian poin , and h ee o he backed blades and bladele s
[57]. The new da e o Le el Xa is 34 ka uncal BP, which is a oo young o he egional Cha
ˆ-
elpe onian, da ed a 37-38k uncal BP in nea by Labeko Koba [4]. The loca ion o he bone
sampled a he si e and he signi ican p esence o ca e bea s do no ule ou he possibili y o
admix u e caused ei he by hose ca ni o es o by o he pos -deposi ional p ocesses no iden i-
ied du ing he exca a ion o Ekain.
These disco dances a e pa icula ly ele an o si es p o usely used in he pas o in es i-
ga e he cul u al adap a ions du ing he Can ab ian Uppe Paleoli hic and e lec he complexi-
ies o a ibu ing a chaeological le els based solely on li hic ypology o echnology. I also
e eals he p oblems caused by he a ibu ions based on single da es wi hou aking in o
accoun he cha ac e is ics o he a chaeological assemblage.
The egional ch onological amewo k o each cul u al pe iod as ob ained om he Bayes-
ian age model is p esen ed in Figs 2–4. No egional model o he Cha
ˆ elpe onian is included
as only ul a il e ed da es a e a ailable o Le el IX lowe o Labeko Koba and his si e has been
modelled elsewhe e (see Figu e S20 in [3]). Rega ding he ch onology o he La e Mous e ian
in he Can ab ian egion, he da es om Axlo go beyond he adioca bon limi and samples
om Llonı
´n Le el VI ailed. Howe e , wo new da es, bo h om Le el VII in Amalda, add
impo an new e idence o he exis ing eco d. The upda ed Bayesian model o he Uppe o
Middle Paleoli hic ansi ion shows ha he end bounda y PDF o he Mous e ian in he
egion is 47.9–45.1ka cal BP (all p obabili y anges a e exp essed a 95,4% he ea e ), con i m-
ing ha i occu ed ea lie in he A lan ic zone han in he no heas e n Ibe ia and in wes e n
and no he n Eu ope [3]. The appa en end o he Mous e ian came immedia ely be o e and
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 7 / 20
du ing GI12 (Fig 2). No Cha
ˆ elpe onian da es we e ob ained in his p ojec , despi e Le el Xa
o Ekain ha ing o iginally been de ined as Cha
ˆ elpe onian based on li hic ypology. Howe e ,
he da e ob ained om Ekain was oo young o he egional ange o he Cha
ˆ elpe onian,
which is ound in he Can ab ian Region in Mo ı
´n and Labeko Koba, bu is only eliably da ed
in he la e a 42.6–41.4ka cal BP (see SI pp.47 in [3]).
The ea lie disappea ance o hose Mous e ian g oups in he Can ab ian egion (no he n
A lan ic Ibe ia), while hey s ill su i ed in Ca alonia (no h-eas e n Medi e anean Ibe ia)
o a ew mo e millennia un il jus be o e 42 cal BP [2,58], as da ed in Romani and L‘A b eda,
migh ha e been caused by as ye unknown ac o s and equi es u he esea ch.
The age model da a e eals ha he e was an in e al o 2,700 o 5,800k (95.4% p obabili y),
wi h a median o 4,400 yea s be ween he end o he Mous e ian and he s a o he Cha
ˆ elpe -
onian in he Can ab ian Region (Table 1). The Cha
ˆ elpe onian appea s o s a se e al
millennia a e he Mous e ian comple ely disappea ed in he egion, while in F ance his “ an-
si ional” echnocomplex appea s be o e he end o he Mous e ian [2]. I Neande hals we e
esponsible o bo h Mous e ian and Cha
ˆ elpe onian echnocomplexes in he Can ab ian
Fig 2. Radioca bon da es o he uppe mos da es o he Mous e ian assemblages om he Can ab ian egion (As u ias, Can ab ia and
he Basque Coun y) calib a ed agains In Cal13 [42] in OxCal .4.2 [41] assuming each sample has a 5% p io p obabili y o being and
ou lie wi hin he gene al - ype ou lie model [43]. All da es ha e been ob ained using he ul a il a ion p o ocol [40]. Re e ences a e
gi en in S1 Table. A
model
= 99.8. deno es p e iously unmodelled da es in oduced as R-da es while he o he s a e modelled da es a
indi idual si es in oduced as P io PDFs.
h ps://doi.o g/10.1371/jou nal.pone.0194708.g002
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 8 / 20
Fig 3. Radioca bon da es o he Au ignacian assemblages om he Can ab ian egion (As u ias, Can ab ia and he
Basque Coun y) calib a ed agains In Cal13 [42] in OxCal .4.2 [41] assuming each sample has a 5% p io
p obabili y o being and ou lie wi hin he gene al - ype ou lie model [43]. All da es ha e been ob ained using he
ul a il a ion p o ocol [40]. Re e ences a e gi en in S1 Table. A
model
= 84.2. deno es p e iously unmodelled da es
in oduced as R-da es while he o he s a e modelled da es a indi idual si es in oduced as P io PDFs.
h ps://doi.o g/10.1371/jou nal.pone.0194708.g003
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 9 / 20
S6 Fig. Radioca bon da es om Ai zbi a e III modelled in OxCal4.2 [41,43] agains
INTCAL13 [42].
(TIF)
S1 Code. CQL indi idual codes.
(DOCX)
S2 Code. CQL egional codes.
(DOCX)
Acknowledgmen s
This esea ch has been suppo ed by he Eu opean Commission h ough FP7-PEOPLE-2012-
CIG (Re : 322112), he Spanish Minis y o Economy and Compe i i eness (HAR2012-33956
and RYC-2011-00695), Can ab ia Campus In e na ional and Uni e si y o Can ab ia o
ABMA. The Ins i u o In e nacional de In es igaciones P ehis o
´ icas de Can ab ia is sponso ed
by he Uni e si y o Can ab ia, he Go e nmen o Can ab ia and Banco San ande . The
au ho s wan o acknowledge he Museo de P ehis o ia y A queologı
´a de Can ab ia, Museo
A queolo
´gico de Bizkaia (specially o D. Ga a e) and he Cen o de Pa imonio Cul u al del
Depa amen o de Cul u a del Gobie no Vasco o he acili ies p o ided du ing he sampling
and s udying ma e ial. They wa mly app ecia e he signi ican help o a chaeologis P. Rasines,
R. Mon es and E. Muñoz du ing sample selec ion. This pape was mainly w i en du ing a
esea ch s ay by one o us (ABMA) a he Le e hulme Cen e o E olu iona y S udies, Depa -
men o A chaeology, Uni e si y o Camb idge unded by Spanish Minis y o Educa ion,
Cul u e and Spo s (Jose de Cas illejo P og am, Re no. CAS15_00054). ABMA and DO a e
g a e ul o P o . Robe Foley and P o . Ma a Mi azo
´n-Lah o hei ins i u ional suppo .
Au ho Con ibu ions
Concep ualiza ion: Ana B. Ma ı
´n-A oyo.
Da a cu a ion: Ana B. Ma ı
´n-A oyo.
Fo mal analysis: Ana B. Ma ı
´n-A oyo, Joseba Rios-Ga aiza , Da id Ocio.
Funding acquisi ion: Ana B. Ma ı
´n-A oyo.
In es iga ion: Ana B. Ma ı
´n-A oyo, Joseba Rios-Ga aiza , Jenni e R. Jones.
Me hodology: Ana B. Ma ı
´n-A oyo.
P ojec adminis a ion: Ana B. Ma ı
´n-A oyo, Jenni e R. Jones.
Resou ces: Ana B. Ma ı
´n-A oyo, Ma co de la Rasilla, Manuel R. Gonza
´lez Mo ales, Michael
Richa ds, Jesu
´s Al una, Ko o Ma iezku ena.
So wa e: Ana B. Ma ı
´n-A oyo, Da id Ocio.
Supe ision: Ana B. Ma ı
´n-A oyo.
Valida ion: Ana B. Ma ı
´n-A oyo, Law ence G. S aus, Da id Ocio.
Visualiza ion: Ana B. Ma ı
´n-A oyo, Joseba Rios-Ga aiza , Da id Ocio.
W i ing – o iginal d a : Ana B. Ma ı
´n-A oyo, Joseba Rios-Ga aiza , Law ence G. S aus,
Jenni e R. Jones, Da id Ocio.
Ch onological eassessmen o he Middle o Uppe Paleoli hic ansi ion in Can ab ian Spain
PLOS ONE | h ps://doi.o g/10.1371/jou nal.pone.0194708 Ap il 18, 2018 16 / 20

W i ing – e iew & edi ing: Ana B. Ma ı
´n-A oyo, Joseba Rios-Ga aiza , Law ence G. S aus,
Jenni e R. Jones, Ma co de la Rasilla, Manuel R. Gonza
´lez Mo ales, Michael Richa ds,
Jesu
´s Al una, Ko o Ma iezku ena, Da id Ocio.
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