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ANTHROPOLOGY
Re ised age o Schöningen hun ing spea s indica es
in ensi ica ion o Neande hal coope a i e beha io
a ound 200,000 yea s ago
Ja od M. Hu son1,2*†, Felix Bi mann3,4, Pe e Fische 5, Alejand o Ga cía- Mo eno1,6,
Sabine Gaudzinski- Windheuse 1,7, Ellie Nelson8, José E. O iz9, Ki s y E. H. Penkman8,
Zo an M. Pe ić10, Daniel Rich e 7,11,12, T inidad To es9, Elaine Tu ne 1, A i za Villaluenga1,13,14,
Dus in Whi e8, Ola Jö is1,7*†
The Schöningen 13II- 4 a chaeological si e in Ge many holds i le o he mos comple e Paleoli hic wooden hun -
ing spea s e e disco e ed, ye i s age has ne e been p ope ly se led. Ini ial es ima es placed he si e a a ound
400,000 yea s; his age was la e e ised o oughly 300,000 yea s. He e, we epo age es ima es o he “Spea
Ho izon” based on amino acid geoch onology o ossils ob ained di ec ly om he ind- bea ing deposi s. Toge he
wi h a eassessmen o egional Middle Pleis ocene ch onos a ig aphy, hese da a place he Schöningen spea s
a ~200,000 yea s. This e ised age posi ions he Spea Ho izon alongside o he si es ha collec i ely eco d a shi
owa d communal hun ing s a egies. The Schöningen a chaeological eco d exempli ies his beha io al ans o -
ma ion ha a ose wi hin he inc easingly complex social en i onmen s o Middle Paleoli hic Neande hals.
INTRODUCTION
The Schöningen 13II- 4 “Spea Ho izon” si e (Landk eis Helms ed ,
Lowe Saxony, Ge many) (Fig. 1) lays claim o he wo ld’s oldes com-
ple e wooden hun ing weapons (1), including nine spea s, one lance,
and six double- poin ed s icks (2). This a senal is joined by a small
assemblage o abou 1500 lin a i ac s (3) sca e ed a ound he
bu che ed emains o mo e han 50 ho ses ( able S1) (4), al oge he
ep esen ing he spoils o epea ed ambush hun ing episodes along
he sho eline o a o me lake (4–9). Nowhe e else in he Middle
Pleis ocene eco d a e such hun ing encoun e s so well p ese ed in
undis u bed con ex s. Acco dingly, he si e has come o se e as a
ma ke ho izon in he de elopmen o p emode n human hun ing
abili ies (10). Gi en his, i is c ucial o asc ibe an accu a e age o he
si e o place he echnologies and beha io s e ealed om he Spea
Ho izon in he b oade imeline o human beha io al e olu ion.
Qua e na y exposu es a he Schöningen open- cas ligni e mine
ep esen some o he mos comple e e es ial a chi es co e ing he
la e hal o he Middle Pleis ocene (see Supplemen a y Tex and igs.
S1 o S3) and con ain nume ous Paleoli hic a chaeological locali ies,
mos no ably he 13II- 4 Spea Ho izon (11). The Spea Ho izon i sel
has ne e been di ec ly da ed success ully. Ins ead, age es ima es o
he spea s ha e elied on a gene alized co ela ion o he s a ig aphic
sequence o he global paleoclima ological eco d in combina ion
wi h c oss- da ing o o he ossil le els wi hin he Schöningen mine
complex (see Supplemen a y Tex ).
The s a ig aphic sequence a Schöningen spans om he Els e i-
an glacia ion o he Holocene (Table 1; see also Supplemen a y Tex ,
igs. S2 and S3, and able S2) (12–14). The Els e ian ep esen s he
sou he nmos ad ance o he Fennoscandian ice shee in o no he n
cen al Eu ope and da es o ma ine oxygen iso ope s age (MIS) 12
(15–18). A Schöningen, Els e ian glacial ills and clays a e ollowed
by a se ies o in e sec ing sedimen - illed channels (Cycles I o VI,
om bo om o op and old o young). Channels con aining Cycle I
o III deposi s a e a gued o ha e o med du ing pe iods o majo
clima ic and en i onmen al change, mos likely a ound he e mina-
ion o he majo glacial cycles, and subsequen ly illed by del aic-
lacus ine deposi s du ing he ensuing in e glacial pe iods. The basic
assump ion ha oughly 100,000 yea s (ky ) long in e glacial- glacial
cycles s a and end wi h majo glacial e mina ions, du ing which
he channels o med, p o ides he backbone o he ela i e ch onol-
ogy a Schöningen, bu such a s ic co ela ion has no been es ab-
lished di ec ly. This la e Middle Pleis ocene sequence is sealed by
g ound mo aines o he D en he ice ad ance o he Saalian glacia ion
(12–14) da ed o MIS 6 (Table 1) (19–21). D en hian ills a e suc-
ceeded by MIS 5e Eemian in e glacial deposi s in he Schöningen
no he n mining ield (13) and by loess- paleosol sequences o Cy-
cles IV o V and Holocene Cycle VI deposi s in he sou he n mining
ield (14).
The Els e ian (MIS 12) and D en hian (MIS 6) ice ad ances ep-
esen he mos se e e cold pe iods o he Eu opean Middle Pleis o-
cene (Fig. 1), which b ough abou subs an ial landscape emodeling
1MOnRePOS A chaeological Resea ch cen e and Museum o Human Beha iou al
e olu ion, leibniz Zen um ü A chäologie, 56567 neuwied, Ge many. 2depa -
men o Paleobiology, na ional Museum o na u al His o y, Smi hsonian ins i u-
ion, Washing on, d.c. 20013, USA. 3lowe Saxony ins i u e o His o ical coas al
Resea ch, 26382 Wilhelmsha en, Ge many. 4ins i u e o Geog aphy, Uni e si y o
B emen, 28359 B emen, Ge many. 5ins i u e o Geog aphy, Johannes Gu enbe g
Uni e si y Mainz, 55099 Mainz, Ge many. 6MUPAc Museum o P ehis o y and A chae-
ology o can ab ia, 39009 San ande , Spain. 7ins i u e o Ancien S udies, depa men
o P ehis o ic and P o ohis o ic A chaeology, Johannes Gu enbe g Uni e si y Mainz,
55099 Mainz, Ge many. 8neaa labo a o y, depa men o chemis y, Uni e si y o
Yo k, Yo k YO10 5dd, UK. 9labo a o y o Biomolecula S a ig aphy, e. .S.i. Minas y
ene gía, Uni e sidad Poli écnica de Mad id, 28003 Mad id, Spain. 10depa men o
Geology, lund Uni e si y, Söl ega an 12, Se- 223 62 lund, Sweden. 11Zen um ü
Bal ische und Skandina ische A chäologie, leibniz Zen um ü A chäologie, 24837
Schleswig, Ge many. 12depa men o Human O igins, Max Planck ins i u e o e olu-
iona y An h opology, 04103 leipzig, Ge many. 13consolida ed Resea ch G oup on
P ehis o y: Human e olu ion, clima e change and cul u al Adap a ion in P eindus ial
Socie ies (GiZAPRe i - 1435- 22), Uni e si y o he Basque coun y (UP /eHU), 01006
i o ia- Gas eiz, Spain. 14depa men o P ehis o y, Ancien His o y, and A chaeology,
complu ense Uni e si y o Mad id, 28040 Mad id, Spain.
*co esponding au ho . email: ja od. hu son@ leiza. de (J.M.H.); ola . joe is@ leiza. de
(O.J.)
† hese au ho s con ibu ed equally o his wo k.
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(16–22). Th oughou he no he n Eu opean Lowlands, smalle de-
p essions le a e he e ea o he ice shee s we e illed du ing he
succeeding Hols einian (MIS 11 peak) and Eemian (~MIS 5e) in e -
glacial pe iods (Table 1; see also ables S2 and S3). Wi h qui e uni-
o m ege a ional successions, he Hols einian In e glacial is closely
linked o he Hoxnian In e glacial in he B i ish Isles and he Maso ian
In e glacial known om Poland (Table 1; see also Supplemen a y
Tex ) (23). Simila ly, he Eemian o no he n Cen al Eu ope co e-
la es wi h he Ipswichian on he B i ish Isles (24).
Pollen p ese ed a di e en exposu es in he Schöningen ligni e
mining complex be ween he Els e ian and D en hian ills shows
ha mos o he del aic- lacus ine deposi s o Cycles I o III p edom-
inan ly o med unde in e glacial o in e s adial clima ic condi ions,
wi h he landscape shi ing be ween o es ed and mo e open, s eppic
ege a ion. On he basis o hei palynology, he peak in e glacial
phases o Cycles I o III we e locally named Al e sdo , Reinsdo ,
and Schöningen (13,25) and a e belie ed o co ela e wi h he wa m-
es pe iods o MIS 11, 9, and 7, espec i ely (13,26–28) (Table 1; see
also able S2). This co ela ion o ms he basis o he Schöningen
s anda d ch onos a ig aphic model applied in mos ecen publica-
ions (27–29). Each o hese in e glacial phases we e succeeded by
deposi s o o es ed in e s adial cha ac e , lacking he mophilous plan
species. Simila in e s adials, locally named O leben I, O leben II,
and Esbeck, also p eceded he Al e sdo In e glacial and ha e been
assigned a la e Els e ian age (12). He e, hese p e- Al e sdo deposi s
a e e e ed o as “Cycle 0” (see igs. S2 and S3).
A chaeological inds om he Schöningen 13II- 4 Spea Ho izon
o igina e om laye s 4a o 4c wi hin he la e pa o he Cycle II
(Reinsdo ) sequence, a he ansi ion om bios a ig aphic uni C
o D (BU- C o BU- D) (27), o me ly assigned o he Reinsdo - B
In e s adial (Fig. 2 and Table 1; see also igs. S2 and S3 and ables S1
and S2). A his ime, he local landscape consis ed o woodland o
open woodland o alde , bi ch, and willow, wi h meadows and aised
bogs lanking he o me lakesho e (27). Hominin occupa ion o he
Fig. 1. Loca ion o Schöningen. Map showing he loca ion o Schöningen (whi e
do ) ela i e o he maximum ex en o majo Middle (Anglian, els e ian, and Saalian
ad ances) and Uppe (de ensian and Weichselian) Pleis ocene inland glacial ad-
ances ac oss no he n eu ope (100,101). Map based on SR M30_PlUS da a.
Table 1. La e Middle Pleis ocene o Holocene sequence a Schöningen. he Schöningen s a ig apic sequence compa ed wi h MiSs and egional
subdi isions o his pe iod based on he MiS age o in e glacial and glacial deposi s (see Supplemen a y ex ). he la e Middle Pleis ocene Schöningen cycles i
o iii a e b acke ed be ween els e ian and d en hian ills o MiS 12 and 6 age. he e o e, Schöningen cycles i o iii a e o be placed be ween MiS 12 and 6, bu
hei co ela ion wi h MiS s ages emains poo ly es ablished and is based on he assump ion ha he Al e sdo in e glacial o cycle i co ela es o he
Hols einian in e glacial o MiS 11c. Bolded and unde lined ex shows he posi ion o he Schöningen 13ii- 4 “Spea Ho izon.” lGM,las glacial maximum;
iG,in e glacial.
Uni s o he
Qua e na y
B i ish Isles Wes e n Cen al
Eu ope
Schöningen Eas e n Cen al
Eu ope
MIS
Holocene Holocene Holocene
cycle i
Holocene Holocene 1
Uppe
Pleis ocene
lGM
ad ances
lGM
ad ances
Weichselian
lGM
ad ances
2
3
de ensian Weichselian
cycle
cycle i
Weichselian 4
5d- 5a
ipswichian eemian eemian eemian 5e
la e Middle
Pleis ocene
Poo ly
esol ed
Wa he ad ance Wa he ad ance 6
d en he ad ance d en hian Ode ad ance
Saalian complex
cycle iii
13 II- 4*
cycle ii
cycle i
Schöningen iG
13 II- 4*
Reinsdo iG
Al e sdo iG
Poo ly
esol ed
7
8
9
10
Hoxnian Hols einian Maso ian 11c
Anglian els e ian els e ian†San- 2 12
* he Schöningen 13ii- 4 Spea Ho izon is con ained in a la e phase o cycle ii deposi s [ ansi ion om bios a ig aphic uni s BU- c o BU- d (27)], se e al me e s
abo e he Reinsdo in e glacial peak sedimen s. †A Schöningen, se e al o es ed in e s adials o p esumably la e els e ian age ha e been desc ibed om
he no he n mining ield, p eceding he Al e sdo in e glacial (12), which a e e e ed o cycle 0 in he ex .
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Spea Ho izon coincided wi h he ansi ion om an open wood-
land s eppic phase (laye 4c; BU- C) o he second o es ed in e s a-
dial phase ha succeeded he Reinsdo In e glacial (laye s 4a/b;
BU- D), p esumably close in ime o he onse o he nex glacial pe-
iod (28,29).
Age es ima es o he Spea Ho izon ha e been amended o e he
yea s om ~400,000 o ~300,000 yea s ago (ka) (1,30–32), MIS 11
o MIS 9, depending on age es ima es o he p eceding Cycle I de-
posi s and al e na i e co ela ions wi h iso ope clima e eco ds (30).
The moluminescence (TL) da ing o hea ed lin om he Schöningen
13I- 1 (Cycle I) a chaeological si e esul ed in an age o 321±14 ka,
placing he si e in o MIS 9 (31) and he Al e sdo peak in e gla-
cial deposi s in o MIS 9e. The Schöningen 13II- 4 Spea Ho izon
inds come om close o he op o he nex younge sedimen ologi-
cal cycle and should he e o e da e o MIS 7. U anium- se ies da es o
290±5 ka o pea ho izons wi hin he Cycle II sequence we e wi h-
d awn because o hei ecognized open- sys em beha io (33,34).
Op ically s imula ed luminescence (OSL) da a assign le el 13II- 2c1
(Cycle II) a maximum age o ~300 ka (32). Le el 13II- 2c1 o med
du ing cold clima ic condi ions (BU- A) immedia ely a e he Rein-
sdo In e glacial, indica ing deposi ion o he sedimen in MIS 9d
(27,32). On he basis o hese es ima es, bo h Cycle I and Cycle II
sequences would da e o MIS 9. The TL and OSL da es om uni s
13I- 1 and 13II- 2c1, howe e , o igina e om samples loca ed s a i-
g aphically below uni 13II- 4 and can he e o e only se e as maxi-
mum age es ima es o he Spea Ho izon. He e, we p o ide age es ima es
o he Schöningen spea s and associa ed a chaeology based on ami-
no acid geoch onology (AAG) o ossils eco e ed di ec ly om he
Spea Ho izon deposi s.
RESULTS
Sys ema ic sampling o AAG inco po a ed mul iple ossil ypes e-
co e ed om among he denses concen a ions o a chaeological
Fig. 2. Schöningen 13II- 4 Spea Ho izon exca a ion plan. Map showing he loca ions o ope cula, os acod, and ho se oo h samples selec ed o AAG analysis (sample
numbe s e e o able 2 and ables S4 and S5). Fou Bi hynia ope cula we e analyzed om wo loci; 13 os acod samples we e analyzed om h ee loci; nine ho se den ine
samples we e analyzed om eigh indi idual mola s. S a ig aphically, he Spea Ho izon wi hin si e Schöningen 13ii- 4 measu es some 20 o 40 cm in hickness, encom-
passing laye s 4b (lamina ed o ganic sil ) and 4b/4c ( ansi ion o calca eous sil ) (102) a he ansi ion om Reinsdo BU- c o BU- d (27). Some a chaeological emains
om laye s 4a (o ganic sil ) and he op o 4c (calca eous sil ) a e also included in he Spea Ho izon. he si e has been almos comple ely exca a ed, co e ing some 3900 m2
and yielding nea ly 15,000 a chaeological inds (11). Spa ially, he main ind concen a ion occupies a 60 m–by–10 m band s addling he no h- Sou h–o ien ed sho eline
o he o me lake (68,102–105). no e ha exca a ion squa es shown a e 10 m by 10 m.
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inds in he Schöningen 13II- 4 Spea Ho izon (Fig. 2; see also ig. S4).
AAG da a we e ob ained using a e e se- phase high- pe o mance
liquid ch oma og aph (RP- HPLC) om he in ac ys alline ac ion
o ou Bi hynia en acula a ope cula (Table 2). As in ac ys alline
p o ein decomposi ion (IcPD) in ope cula ep esen s an e ec i e
closed sys em o ossil p o ein (35), hese da a we e he p ima y
means o de i e an age es ima ion. Addi ional independen nume i-
cal AAG age es ima es om os acods and ho se ee h (Fig. 2; see
also ables S4 and S5) a e conside ed explo a o y gi en he po en ial
open- sys em na u e o he samples and he sca ci y o he equi ed
compa a o ma e ial.
Measu ed agains an AAG amewo k es ablished o Bi hynia
ope cula on he B i ish Isles (35), he Schöningen 13II- 4 Spea Ho-
izon ope cula da ase clea ly alls wi hin he younge ange o si es
a ibu ed o MIS 7 (Fig. 3 and Table 2). Howe e , he same in e-
g a ed empe a u e his o y canno be assumed o di e en egions
(36), so compa ison wi h geog aphically close si es is c ucial. Com-
p ehensi e ope cula aminos a ig aphies o con inen al Eu ope a e
now unde de elopmen , and compa isons wi h ini ial da a can al-
eady be made.
The Spea Ho izon ope cula D/L alues ( a io o igh - o le -
handed amino acids; see Ma e ials and Me hods) a e simila o hose
om he MIS 7 in e glacial locali y a T ianglen in Copenhagen,
Denma k (37). Because T ianglen (~450 km o he no h- no heas )
likely expe ienced sligh ly colde in eg a ed empe a u es han Schönin-
gen h oughou he la e Middle Pleis ocene, D/L alues om T ianglen
would be sligh ly lowe han a Schöningen o samples o a simila
age. Compa ed wi h he las in e glacial Eemian deposi s (MIS 5e) o
he nea by si e o Neuma k- No d 2, Ge many (38,39), he Spea Ho-
izon ope cula show sligh ly highe D/L alues in he as e - acemizing
alanine (Ala) and aspa gine/aspa ic acid (Asx), he wo amino ac-
ids ha p o ide he bes empo al esolu ion o si es o his age (Fig.
3 and Table 2). Thus, he Spea Ho izon is olde han he Eemian
deposi s a Neuma k- No d 2. As he Cycle II sequence a Schöningen
is o e lain by a glacial ill a ibu ed o he D en hian s age o he
Saalian glacia ion (Table 1), he Spea Ho izon mus also p e- da e
MIS 6. O e lapping D/L alues o he slowe - acemizing aline
(Val) and glu amine/glu amic acid (Glx) om he Spea Ho izon
and om Neuma k- No d 2 (Fig. 3 and Table 2) indica e ha he
Schöningen samples canno p e- da e he Eemian In e glacial (MIS
5e) by se e al clima ic cycles. Con e sely, obse ed le els o IcPD in
he Spea Ho izon ope cula da ase a e subs an ially lowe , and
he e o e younge , han hose o he Hols einian Paludinenbank de-
posi s in Be lin (40) and he “pos - Els e ian” in e glacial lu ial
deposi s a he bo om o he Bilzingsleben (Ge many) a e ine
a ibu ed o MIS 11 (41). Conside ing ha hese compa a o si es
ha e expe ienced a simila empe a u e his o y, an age o MIS 7 is
indica ed o he Schöningen 13II- 4 Spea Ho izon, which is sub-
s an ially younge han all ea lie es ima es (1,30–32).
These closed- sys em AAG ope cula da a a e suppo ed by low
D/L alues and age calcula ions o os acod and ho se ee h sam-
ples om he Schöningen 13II- 4 Spea Ho izon (see ables S4 and
S5) ha place he si e nea he end o he la e Middle Pleis ocene,
al hough we concede ha he age calcula ions de i ed om os a-
cod and ho se ee h a e s ill expe imen al and lack a obus com-
pa a o da ase . S a ig aphically, he Spea Ho izon mus p e- da e
he MIS 6 ice ad ance o he D en hian (13,14,19–21), and all
da a om ope cula, os acod, and ho se ee h samples ag ee wi h
his ch onology.
DISCUSSION
Ch onos a ig aphic implica ions
The AAG da a b ing abou a se ies o implica ions o unde s anding
he la e Middle Pleis ocene ch onos a ig aphy a Schöningen and o
es ablishing a mo e p ecise age o he 13II- 4 Spea Ho izon. A MIS 7
age pushes he a chaeology o he Spea Ho izon o a da e oughly
100 ky younge han p e iously conside ed. Consequen ly, his im-
plies ha channel o ma ion and cycles o deposi ion a Schöningen
would no in e e y case co espond o ull in e glacial- glacial cycles,
whe e each s a ed and ended wi h a majo glacial e mina ion. In-
s ead, we p opose a e ised Schöningen ch onos a ig aphic model
whe e co ela ions o he Schöningen Cycles wi h iso ope ch onolo-
gies a e conside ed a subs age le els, and Cycles II and III a e bo h
placed in o MIS 7, wi h Cycle III also ex ending in o ea ly MIS 6 (Fig. 4;
see also able S2).
Wi h Cycle II da ing o MIS 7, he p eceding Cycle I Al e sdo
In e glacial can be placed in o he peak phase o MIS 9 (i.e., MIS 9e),
which is in ull ag eemen wi h he TL age es ablished o he
Schöningen 13I- 1 si e (Fig. 4) (31). Howe e , his con adic s ea lie
age models ha co ela e he Cycle I Al e sdo deposi s o he
Hols einian o MIS 11. The basic a gumen o his co ela ion was ha
Cycle I in e glacial deposi s ep esen he i s pe iod cha ac e ized
by ully in e glacial condi ions ollowing he Els e ian glacial se ies a
Table 2. P o enance and amino acid da a o Schöningen ope cula samples. e o e ms ep esen 1 Sd abou he mean o he duplica e analyses o an
indi idual sample. each sample was bleached (b), wi h he ee amino acid (FAA) ac ion signi ied by “F” and he o al hyd olysable ac ion by “H*” (see Ma e ials
and Me hods). neaa , no h eas Amino Acid Racemiza ion.
Schöningen ID P o enance Laye NEaa no. Ala D/L Asx D/L Glx D/L Se D/L Val D/L
9807 709/2 4b and 4b/4c
10606bF 0.300±0.000 0.708±0.004 0.169±0.006 0.986±0.000 0.170±0.002
10606bH*0.262±0.002 0.641±0.000 0.166±0.000 0.772±0.011 0.142±0.001
10607bF 0.305±0.001 0.717±0.004 0.202±0.004 0.992±0.004 0.173±0.001
10607bH*0.255±0.001 0.612±0.001 0.176±0.002 0.736±0.008 0.135±0.000
11912 724/15 4b and 4b/4c
10608bF 0.306±0.000 0.696±0.000 0.162±0.002 0.983±0.004 0.177±0.006
10608bH*0.236±0.001 0.587±0.002 0.140±0.002 0.622±0.003 0.152±0.005
10609bF 0.294±0.002 0.700±0.001 0.190±0.003 0.982±0.007 0.165±0.001
10609bH*0.249±0.001 0.616±0.001 0.168±0.002 0.759±0.000 0.131±0.000
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Schöningen. On he basis o i s palynology, he Al e sdo In e gla-
cial was a gued o be simila o he Hols einian (25), bu i s sligh ly
di e en pollen successions sugges ed ha i could ep esen a local
a ian o he Hols einian (26). Gi en he o e all simila i ies be ween
Hoxnian, Hols einian, and Maso ian In e glacial a chi es (23), such
local a ia ion o he Hols einian a Schöningen is unlikely. Ra he ,
he Al e sdo In e glacial ep esen s an in e glacial di e en om
and younge han he Hols einian sensu s ic o (see able S3).
In no he n cen al Eu ope, o es ed in e s adials, like he Cycle 0
O leben I, O leben II, and Esbeck In e s adials ha p eceded he
Al e sdo In e glacial deposi s in he no he n mining a ea o
Schöningen (25), ypically co ela e o he la e subs ages o he
Fig. 3. Resul s o IcPD analysis on Bi hynia ope cula. F ee (FAA) e sus o al ( HAA) d/l alues o alanine (Ala), aline ( al), glu amic acid/glu amine (Glx), and aspa ic
acid/aspa agine (Asx) om Schöningen 13ii- 4 bleached B. en acula a ope cula ( able 2) compa ed wi h shells om Ge man and danish si es: oig s ed (A e n in e gla-
cial, ~MiS 29- 21) (84); da a om he la es c ome ian, p e- els e ian si e o Miesenheim i (MiS 13) (83) (see able S6); he pos - els e ian (MiS 11) si e o Bilzingsleben (see
able S6); he Hols einian (MiS 11) Paludinenbank deposi s in Be lin (see able S6); copenhagen ianglen (MiS 7) (37); and neuma k- no d 2 (MiS 5e) (38,39). Schöningen
and compa a o si es lie be ween 50.4°n and 52.5°n la i ude, excep o ianglen, which is loca ed u he o he no h a 55.7°n la i ude. While he in eg a ed empe a-
u e his o ies will be sligh ly di e en , he anspa en blue ellipses show he ange o alues exhibi ed by ope cula om B i ish si es (35) wi h age co ela ion o he MiS
eco d (78). no e he di e en ly scaled axes o each g aph; each amino acid acemizes a a di e en a e, p o iding di e en le els o empo al esolu ion.
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Fig. 4. Majo s a ig aphic uni s o he Schöningen Pleis ocene sequences. Schöningen s a ig aphy is co ela ed wi h global MiSs and he deu e ium eco d o em-
pe a u e change documen ed in he An a c ic dome c ice co e (106) o e he pas 500 ky (see also able S2). G een a ow in he igh column indica es he s a ig aphic
posi ion o he Schöningen 13 ii- 4 Spea Ho izon wi h he app oxima e age o ~200 ka. colo s o cycles i o i co espond o he colo coding in igs. S1 o S3. da a om
Schöningen no he n mining a ea modi ied om (12,25); da a om Schöningen sou he n mining a ea modi ied om (13,26); l age es ima es o Schöningen 13i- 1 om
(31). colo coding o ice co e phases and MiS ollows an ex apola ion o en i onmen al condi ions in no he n cen al eu ope (42). - i o - ma ks a he le deno e he
e mina ions o majo glacial cycles. iG,in e glacial; iS,in e s adial; U,majo uncon o mi ies.
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longe empe a e pe iods [e.g., MIS 5a and 5c in Uppe Pleis ocene
eco d (42)]. In e s adials equi alen o he O leben I, O leben II,
and Esbeck a e no documen ed elsewhe e om la e Els e ian depos-
i s (43). On he con a y, whene e s a ig aphic da a a e a ailable,
he Hols einian succession lies immedia ely and conco dan ly on op
o he Els e ian glacial se ies (18,19, 44,45), indica ing he apid
change om ully glacial o ully in e glacial condi ions, lea ing no
ime o he es ablishmen o in e s adial o es ed en i onmen s (Fig.
4). Ou e ised age model (see able S2) no only ea i ms he MIS 9e
age o he Al e sdo In e glacial (Cycle I) bu also implies ha no
dis inc ly Hols einian (MIS 11 peak=MIS 11c) deposi s ha e ye
been iden i ied a Schöningen. As a esul , he O leben I, O leben II,
and Esbeck In e s adials would co espond o he la e wa ming
phases wi hin MIS 11 (i.e., MIS 11a) a he han ep esen ing la e
MIS 12 wa ming in e als (25). The Cycle I Al e sdo In e glacial o
MIS 9e was hen ailed by u he mild in e s adial phases (Missaue
I, Missaue II, and SU A), likely co ela ing wi h MIS 9c–a (Fig. 4).
Mo ing h ough he s a ig aphic sequence, he Cycle II Reinsdo
In e glacial peak deposi s would co ela e o MIS 7e and he 13II- 4
Spea Ho izon o a mild in e al owa d he end o MIS 7. This is
no con adic ed by OSL da a o laye 13II- 2c1 ha ha e been ex-
plici ly desc ibed as maximum age es ima es (32), wi h single aliquo s
co e ing a wide ange o indi idual measu emen s be ween ~500
and~200 ka (see ig. S5).
The palynologically de ined Schöningen In e glacial o he no h-
e n mining a ea succeeds Cycle II o he sou he n mining ield and
would co ela e o he inal phase o MIS 7 and he ensuing o es ed
in e s adials o he MIS 6 ansi ion (Fig. 4; see also able S2). Cycle
III deposi s in he sou he n mining a ea do no p ese e any sedi-
men s o clea ly empe a e cha ac e bu equen ly display c yogen-
ic ea u es, placing hem pa ly in o MIS 6 p eceding he D en hian
ice ad ance.
In line wi h he egional ch onology o he no he n cen al Eu opean
lowlands (see Supplemen a y Tex ), ou da a now p o ide ou ch on-
ological e e ence poin s in he Schöningen sequence (Fig. 4 and
see also able S2): (i) he Els e ian ills a he base o he sequence
da e o MIS 12 (15,17); (ii) he Al e sdo In e glacial ep esen ed a
Schöningen 13I- 1 da es o MIS 9 (31) bu does no co espond o he
Hols einian sensu s ic o (MIS 11); (iii) he amino acid da a sugges a
MIS 7 age o Schöningen 13II- 4; and (i ) he la e Saalian D en he ills
belong o MIS 6 (13,14,19–21). Wi h e e ence o he local bios a i-
g aphic e idence (27), he Spea Ho izon co esponds o he second
o es ed in e s adial phase (BU- D) a e he Reinsdo In e glacial
peak, which likely co ela es o a phase o clima ic de e io a ion a he
end o MIS 7c o wi hin MIS 7b (Fig. 4; see also Supplemen a y Tex ,
ig. S6, and able S2). In consequence, we sugges a la e MIS 7 age a ound
~200 ka o he Spea Ho izon and he wo ld’s oldes comple e hun ing
weapon y, oughly 100 ka younge han p e iously assumed.
Wi h hese ch onos a ig aphic implica ions, he ou empo al
e e ence poin s in he Schöningen sequence and e ised age o he
Spea Ho izon can se e as he wo king ounda ions o a ee alua-
ion o Middle Pleis ocene mammalian bios a ig aphy ac oss sho -
and long- e m clima e cycles.
Hominin beha io al implica ions
The Schöningen spea s will be celeb a ed o e e as spec acles o Pa-
leoli hic echnology, ye hei impo ance o unde s anding hominin
beha io al e olu ion can only be ully app ecia ed wi hin he wide
con ex o he la e Middle Pleis ocene a chaeological eco d.
The small li hic assemblage made o local lin unde sco es he si e’s
unc ional cha ac e and epheme al use o aw ma e ials in he Spea
Ho izon. Blank p oduc ion, o any o m o p ima y li hic educ ion,
played a mino ole a he si e. E idence o he applica ion o Le allois
laking s a egies is absen . Ins ead, he assemblage consis s la gely o
inished ools b ough o he si e and an abundance o small lakes,
chips, and esha pening deb is, wi h use wea hin ing a ac i i ies e-
la ed o bu che y, woodwo king, and hide sc aping (3). O e all, he
li hic assemblage is no indica i e o any pa icula ime pe iod bu is
o en a ibu ed he Lowe Paleoli hic owing o he absence o p epa ed-
co e echnology and he o iginal olde age es ima es o he si e (3).
Lacking di ec da es o he Spea Ho izon, he e had long been a
ce ain ai o unce ain y su ounding he deep age o he si e. Wi h
claimed ages o ~400 o ~300 ka o he Spea Ho izon, Schöningen
s ood as an ou lie among si es a ibu ed o he Lowe Paleoli hic
and o Homo heidelbe gensis (46), he p obable las common ances o
o ea ly mode n humans ha e ol ed in A ica and he Neande hals
o wes e n Eu asia (47). Ou da ing e idence o he Spea Ho izon
co ec s his misma ch and aligns he Schöningen spea s wi hin he
ime ame o Eu opean Neande hals and he Middle Paleoli hic.
A la e MIS 7 age does no diminish he signi icance o he si e—
he Spea Ho izon a chaeological eco d and associa ed beha io s
s and unchanged— a he ou e ised age assignmen posi ions he
Schöningen indings in line wi h o he e idence om his pe iod o
eme ging Neande hal beha io al complexi y (48,49).
The Spea Ho izon p o ides unequi ocal e idence o he ecu -
en use o he Schöningen lakesho e as an ambush hun ing g ound
a which en i e ho se amily g oups we e a ge ed, killed, and bu ch-
e ed (4–9). This scena io o epea ed hun ing e en s gene a ed a zoo-
a chaeological signa u e simila o many Middle Paleoli hic si es wi h
la ge, mono- speci ic aunal assemblages a ibu ed o Neande hal
hun ing (50–52). Whe eas he Schöningen hominins specialized in
hun ing ho ses, o he si es show an equally s ong p e e ence o a
single p ey axon, including au ochs a Biache- Sain - Vaas (F ance;
MIS 7) (53), ed dee a G o e du Laza e (F ance; MIS 6) (54), hi-
noce os a Taubach (Ge many; MIS 5e) (55), bison a Mau an (F ance;
MIS 5) (56), ho se a Zwoleń (Poland; MIS 5a/4) (57), and eindee a
Salzgi e Lebens ed (Ge many; MIS 5/4) (58,59), o men ion only
a ew. By MIS 5e, Neande hals egula ly ook down e en he la ges
p ey animals, as e idenced by he nume ous bu che ed elephan ca -
casses om Neuma k- No d 1 (Ge many) (60). Occasionally, mono-
speci ic hun ing has been in e p e ed om Lowe Paleoli hic si es,
mos no ably eindee in le el L a La Caune de l’A ago (Tau a el,
F ance) (61) and bison in TD 10.2 a G an Dolina (A apue ca, Spain)
(62). Howe e , he aphonomic his o ies o hese ca e si es a e a
mo e complica ed, blu ing in e p e a ions o any disc e e hun ing
e en s a hese locali ies. Gi en his, i appea s ha mono- speci ic
hun ing coalesced in o egula p ac ice du ing MIS 7 and became
widesp ead in la e Middle Paleoli hic con ex s (63). To achie e his
le el o ou ine success, hun ing o ays a Schöningen and o he
mono- speci ic si es mus ha e been wholly collabo a i e and g oup-
minded e o s mo i a ed by a communal se o objec i es (64,65).
The g owing body o e idence o high le els o hun ing collabo a-
ion implies ha such Neande hal social en i onmen s became a
powe ul e olu iona y o ce du ing MIS 7, allowing o g ea e con ol
o e hei en i onmen s and ul ima ely leading o inc eased impac s
on he ecosys em by he la e Pleis ocene. The age es ima e o ~200 ka
ealigns he Schöningen 13II- 4 Spea Ho izon wi h his o ma i e
pe iod o Neande hal hun ing specializa ion.
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MATERIALS AND METHODS
Amino acid geoch onology
AAG elies on he ime- dependen deg ada ion o p o eins wi hin
ossils. The e a e 20 na u ally occu ing amino acids in li ing o gan-
isms, and all bu one ha e a chi al cen e , meaning ha amino acids
can exis in wo chemically iden ical mi o image o ms (enan io-
me s), like le and igh hands. In li ing o ganisms, p o eins a e al-
mos exclusi ely made om he le - handed o m. A e dea h, o
upon cessa ion o issue u no e , a spon aneous acemiza ion eac-
ion occu s g adually o ed ess he imbalance un il he e is a ace-
mic equilib ium o le - (L) and igh - handed (D) amino acids. The
p opo ion o D- o- L amino acids is he e o e a measu e o he ex-
en o p o ein deg ada ion and, i his is p edic able o e ime (e.g.,
wi hin a closed sys em), can be used o es ima e he age o a sam-
ple (66,67).
Sample collec ion
Ope cula (B. en acula a) and os acod (He pe ocyp is ep ans and
P ionocyp is zenke i) samples we e collec ed om sedimen blocks
li ed om he o iginal exca a ion conduc ed by Ha mu Thieme
and he Niede sächsisches Landesam ü Denkmalp lege (NLD) be-
ween 1992 and 2008 (Fig. 2; see Supplemen a y Tex and ig. S4)
(11,68). These blocks we e ini ially in ended o p ese e impo an o
agile aunal specimens and now ep esen he las emaining sedi-
men a y eco d om he main ind concen a ion o he Spea Ho i-
zon. O e he cou se o he exca a ion, nea ly 50 sedimen blocks
we e anspo ed o he NORDFROST GmbH & Co indus ial eeze
acili y nea Ba singhausen, Ge many and s o ed unde subze o (de-
g ees Celsius) empe a u e un il p ocessing began in 2014 a he NLD
and la e a he Paläon Fo schungs- und E lebniszen um Schöninge
Spee e (Fo schungsmuseum Schöningen) in Schöningen, Ge many.
The blocks we e hawed slowly unde con olled condi ions inside a
labo a o y ume hood o p ohibi mold g ow h. A e consolida ing
and ex ac ing he bones o in e es , hal o he emaining po ion o
each block was exca a ed, and he sedimen was wa e - sie ed by laye
h ough nes ed sc eens; sedimen columns we e also p epa ed o mi-
c omo phology and pollen analyses (69).
Samples om ho se (Equus mosbachensis) pe manen mola s o igi-
na ed om he 1992–2008 NLD exca a ions a Schöningen 13II- 4
(Fig. 2; see also Supplemen a y Tex ) (11,68). Den ine powde was
ex ac ed om he buccal su aces o he ho se ee h by diamond-
headed d ill bi a he Cu - Engelho n- Zen um A chäome ie gGmbH
in Mannheim, Ge many.
In o al, ou ope cula, 15 os acod al es, and nine den ine col-
lagen samples om eigh ho se mola s we e subjec ed o AAG anal-
ysis (Fig. 2). All samples o igina ed om he Schöningen 13II- 4
Spea Ho izon, laye s 4b, 4b/4c, and 4c.
Ope cula analysis
Analysis o B. en acula a ope cula samples combined he isola ion
o an “in ac ys alline” ac ion o amino acids by bleach ea men
(70–73) wi h RP- HPLC (74), pe o med a he No h Eas Amino
Acid Racemiza ion Labo a o y, Uni e si y o Yo k, UK. This combi-
na ion o echniques gene a es D/L alues o amino acids om he
chemically p o ec ed p o ein wi hin he biomine al, he eby en-
abling dec eased sample sizes and inc eased analy ical eliabili y.
Fo p e ea men , indi idual ope cula we e sonica ed, insed mul-
iple imes in HPLC- g ade wa e , and ai - d ied. To isola e he in a-
c ys alline ac ion, each cleaned ope culum was ans e ed o a
plas ic mic ocen i uge ube and c ushed, and 50μl o 12% NaOCl
was added pe millig am o he powde ed sample. The ubes we e
shaken, es ed o 24 hou s, eshaken, and soaked again o 24 hou s,
and he NaOCl was pipe ed o . The emaining powde was insed
wi h H2O, cen i uged, and insed again; his p ocess was epea ed
i e imes. Fo comple e emo al o he bleach, HPLC- g ade me ha-
nol was added, le o es o a ew minu es, cen i uged, and pipe ed
o , and he bleached powde was ai d ied o e nigh . The bleached,
d y powde was spli in o wo subsamples: one o he analysis o he
ee amino acid (FAA) ac ion and one o he o al hyd olysable
amino acid ac ion (THAA). The FAA subsamples we e demine al-
ized wi h 10μl o 2 M HCl pe millig am o CaCO3 and d ied
o e nigh in a cen i ugal e apo a o ; he THAA subsamples we e
demine alized in 20μl o 7 M HCl pe millig am o CaCO3. The sub-
samples we e sealed unde N2 and hyd olyzed by hea ing a 110°C o
24 hou s. Following hyd olysis, ials we e placed in a cen i ugal e ap-
o a o o d y o e nigh .
Samples we e ehyd a ed wi h 0.01 mM HCl and l- homo- a ginine
(in e nal s anda d syn he ic amino acid) and analyzed in duplica e
by RP- HPLC using luo escence de ec ion (75). A sample solu ion
olume o 2μl was mixed online wi h 2.2μl o de i a izing eagen
(260 mMN- isobu y yl- l- cys eine and 170 mM o- ph haldialdehyde
dissol ed in 1.0 M po assium bo a e bu e , adjus ed o pH 10.4 wi h
KOH) immedia ely be o e injec ion. De i a ized amino acids we e
sepa a ed wi h C18 Hype Sil BDS C18 e e se- phase columns (5 μm;
250 mm leng h by 4 mm diame e ) a 25°C using a linea g adien elu-
ion o sodium ace a e bu e (eluen A; 23 mM sodium ace a e ihy-
d a e wi h 1.5 mM sodium azide and 1.3 μM EDTA, adjus ed o
pH 6.00±0.01 wi h 10% ace ic acid and 10 M sodium hyd oxide),
HPLC- g ade me hanol (eluen B), and HPLC- g ade ace oni ile
(eluen C).
Du ing p epa a i e hyd olysis, bo h aspa agine and glu amine
unde go apid and i e e sible deamina ion o aspa ic acid (Asp)
and glu amic acid (Glu), espec i ely (76). I is he e o e impossible
o di e en ia e hese amino acids om hei de i a i es, and hey
a e epo ed oge he as Asx and Glx. The D/L alues o aspa ic
acid/aspa agine (Asx), glu amic acid/glu amine (Glx), se ine (Se ),
alanine (Ala), and aline (Val), as well as he (Se )/(Ala) alue, p o-
ide eliable es ima es o p o ein decomposi ion. Se ine is geochem-
ically uns able, wi h one o i s decomposi ion p oduc s being alanine
(77). This enables he a io o he concen a ion o se ine (Se ) o he
concen a ion o alanine (Ala) o be used as a use ul indica ion o
he ex en o p o ein decomposi ion. The D/L alue o an amino acid
will inc ease wi h inc easing ime, while he (Se )/(Ala) alue will de-
c ease. Each amino acid acemizes a di e en a es and he e o e is
use ul o e di e en imescales. The D/L o se ine is less use ul as a
geoch onological ool o samples o his age, bu abe an alues may
indica e con amina ion. In a closed sys em, he amino acid a ios o
he FAA and he THAA subsamples should be highly co ela ed, en-
abling he ecogni ion o comp omised samples (35,78,79). Amino
acid da a om he Schöningen ope cula a e consis en wi h closed-
sys em p o eins and he e o e can be eliably used o age es ima ion.
The ex en o IcPD in bo h he FAA and THAA inc eases wi h ime,
wi h inc eased le els o p o ein b eakdown du ing wa m clima ic
s ages and a slowing in he a es o deg ada ion in cold s ages. O e a
small geog aphical a ea, i can be assumed ha he in eg a ed em-
pe a u e his o ies a e e ec i ely he same. Gi en a simila empe a u e
his o y, his hen allows an aminos a ig aphic amewo k o an
a ea o be de eloped, plo ing he FAA agains he THAA da a, wi h
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younge samples plo ing owa d he bo om le - hand co ne o he
g aph and olde samples owa d he op igh - hand co ne o he g aph.
In his sense, IcPD ep esen s a ela i e ch onology, no wi hs anding a
egional he mal g adien (80). Independen geoch onology allows
hese clus e s o be co ela ed o he ma ine oxygen iso ope eco d;
such a amewo k has been de eloped o ope cula om he Uni ed
Kingdom (35,78). The Schöningen samples ha e amino acid a ios
ha a e sligh ly highe han hose o he same species om si es co -
ela ed wi h he B i ish Ipswichian and he e o e he Eemian and MIS
5e (Fig. 3 and Table 2) (81). Howe e , a di ec co ela ion mus be
aken wi h cau ion owing o he likely di e ences be ween he em-
pe a u e his o ies o con inen al and B i ish Pleis ocene si es. To ad-
d ess his, a compa ison was made be ween he Schöningen samples
and an ini ial da ase o o he Ge man and Danish si es wi h mo e
compa able empe a u e his o ies: he MIS 5e si e o Neuma k- No d
2 (38,39,82); he independen ly da ed MIS 7 si e a Copenhagen
T ianglen (37); IcPD da a om he Hols einian Paludinenbank laye
in Be lin (MIS 11) (see able S6); he pos - Els e ian (MIS 11) in e gla-
cial g a els a he base o he Bilzingsleben a e ine (see able S6); he
la es C ome ian, “p e- Els e ian” si e o Miesenheim I [MIS 13 (83)]
(see able S6); and he A e n In e glacial si e o Voig s ed [~MIS 29-
21 (84)].
Os acod and oo h analysis
Os acod al es we e emo ed om he sedimen samples wi h a
needle, cleaned ul asonically in dis illed deionized (DDI) wa e ,
and insed wi h DDI wa e o emo e sedimen . Some al es we e
addi ionally cleaned wi h a small b ush unde a binocula mic o-
scope o elimina e ine deb is. The cleaned al es we e subsequen ly
subme ged in 3% H2O2 o 2 hou s o emo e seconda y o ganic
molecules adso bed o he shells (74,85). Only anslucen speci-
mens we e selec ed o analysis, as hese a e o en be e p ese ed
and as a po en ial means o excluding ewo ked samples. Den ine
powde was ex ac ed om a hole (2 mm in diame e , 5 o 10 cm in
dep h) bo ed in o he buccal su aces o ho se ee h wi h a diamond-
headed d ill bi . A e demine aliza ion, he den ine samples we e
pu i ied and concen a ed by dialysis (3.5 kDa) (86,87). All os a-
cod and den ine samples we e p epa ed and subjec ed o amino acid
analysis using an RP- HPLC (75,85) in he Biomolecula S a ig a-
phy Labo a o y (Labo a o io de Es a ig a ía Biomolecula ) a he
Uni e sidad Poli écnica de Mad id, Spain.
The samples we e sealed unde N2 in 20μl o 7 M HCl pe mil-
lig am o den ine o 7μl o 6 M HCl pe millig am o os acod and
hyd olyzed by hea ing a 100°C o 20 hou s o elease pep ide-
bound amino acids. The hyd olysa es we e e apo a ed in acuo, e-
hyd a ed in 7μl o 0.01 M HCl wi h 1.5 mM sodium azide and 0.03 mM
l- homo- a ginine (in e nal s anda d syn he ic amino acid), and analyzed
using an Agilen - 1100 HPLC equipped wi h a luo escence de ec-
o . Exci a ion and emission wa eleng hs we e p og ammed a 230
and 445 nm, espec i ely. De i a iza ion ook place be o e injec ion
by mixing 2μl o sample solu ion wi h 2.2μl o p ecolumn de i a iza-
ion eagen , consis ing o 260 mM isobu y yl- l- cys eine chi al hiol
and 170 mM o- ph haldialdehyde dissol ed in 1.0 M po assium bo a e
bu e solu ion a pH 10.4. De i a ized amino acids we e sepa-
a ed wi h Hype sil BDS C18 e e se- phase columns (5 μm; 250 mm
leng h by 4 mm inne diame e ) using a h ee- sol en linea g adien
elu ion o sodium ace a e bu e (eluen A: 23 mM sodium ace a e
ihyd a e wi h 1.5 mM sodium azide and 1.3 μM EDTA, adjus ed
o pH 6.00 wi h 10% ace ic acid and 10 M sodium hyd oxide),
HPLC- g ade me hanol (eluen B), and ace oni ile (eluen C). The
linea g adien was pe o med a 25°C wi h a low a e o 1.0ml/
min, om 95% eluen A and 5% eluen B upon injec ion o 76.6%
eluen A, 23% eluen B, and 0.4% eluen C a 31 min.
Fo os acods, he analysis o mul iple amino acids yields la gely
edundan in o ma ion on sample age (88); he e o e, only he as-
pa ic acid/aspa agine (Asx) and glu amic acid/glu amine (Glx)
cons i uen s o he os acod al es we e analyzed, as Asx and Glx
accoun o ~50% o he amino acid con en in mos os acod al es
(85,89,90). To es ablish he ages o he os acod samples, D/L
alues o Asx and Glx we e en e ed in o he equa ions es ablished o
Cyp ideis o osa and H. ep ans om cen al and sou he n pa s o
he Ibe ian Peninsula (D/L Asx<0.401 and D/L Glx<0.140) (91)
Since H. ep ans was included in he analyzed samples, he D/L
alues we e di ec ly compa able wi hou a con e sion ac o ; also,
C. o osa and H. ep ans show simila acemiza ion a es (91). How-
e e , P. zenke i analyzed om Schöningen may show some di e ences
in acemiza ion pa e ns when compa ed wi h hose o H. ep ans.
A cu o alue o 0.8 in he concen a ion o l- se ine o ha o
l- aspa ic acid was used o de e mine anomalous samples as se ine
decomposes apidly (92), and excessi e amoun s can indica e
con amina ion by mode n amino acids. Two analy ical samples
(Schöningen ID 11965) su e ed his kind o con amina ion and we e
ejec ed o he age calcula ion.
Fo he ho se oo h samples, only aspa ic acid (Asx; as due o he
p epa a ion me hod i is composed o bo h aspa ic acid and aspa a-
gine) was used in he analysis as i acemizes as e and has been
shown o p o ide be e age es ima es o oo h samples (93–98). In
many cases, he D enan iome s o o he amino acids could no be
iden i ied in he ch oma og ams o appea ed in such low amoun s
ha he D/L alues we e no sui able o di e en ia ing be ween sam-
ples (e.g., leucine and isoleucine).
Nume ical ages o he ho se den ine samples we e calcula ed
om D/L alues o aspa ic acid (Asx) wi h he equa ion es ablished
o den ine collagen o mammals om he Ibe ian Peninsula (D/L
Asx<0.40) (98)
Al hough amino acid acemiza ion depends on he axon con-
side ed (99), hese algo i hms we e used on he assump ion ha col-
lagen acemiza ion a es a e no a ec ed by genus (98).
Supplemen a y Ma e ials
This PDF ile includes:
Supplemen a y ex
Figs. S1 o S6
ables S1 o S6
Re e ences
√
Age (ka)=−3.586+19.745Ln
�
1+D∕L Asx
1−D∕L Asx
�;
=0.993, P=0.001
√
Age (ka)=−3.186+58.972Ln
�
1+D∕L Glx
1−D∕L Glx
�;
=0.989, P=0.001
Age (ka)=−10.4+868.1 ∗D∕L Asx; =0.97, P
<
0.001
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