One hundred years later: new evidence supporting an intentional Neandertal

burial at La Chapelle-aux-Saints (Corrèze, France)

William Rendu, Cédric Beauval, Isabelle Crevecoeur, Priscilla Bayle, Antoine Balzeau,

Thierry Bismuth, Laurence Bourguignon, Géraldine Delfour, Jean-Philippe Faivre, François

Lacrampe-Cuyaubère, Carlotta Tavormina, Dominique Todisco, Alain Turq, Bruno Maureille.

SUPPORTING INFORMATION (SI) SUPPORTING TEXTS ............................................................................................................................. 2 

TEXT S1: Bouffia 118 zooarchaeological analysis ............................................................................ 2 

TEXT S2: New Neandertal remains discovered during the 2011‐2012 excavations ....................... 2 

SUPPORTING FIGURES ......................................................................................................................... 7 

FIGURE S1: Site location and picture of the site entrance. ............................................................. 7 

FIGURE S2: Map of the La Chapelle‐aux‐Saints cavity complex and main stratigraphic 

information. ..................................................................................................................................... 8 

FIGURE S3: Mousterian sites with faunal spectra dominated by Bison or Reindeer attributed to 

the MIS 4 or the MIS 3 ..................................................................................................................... 9 

FIGURE S4: Human dental remains discovered during the 2011‐2012 excavations and during the 

1920’s. ........................................................................................................................................... 10 

FIGURE S5: Human osseous remains discovered during the 2011‐2012. ..................................... 11 

FIGURE S6: First excavation stage of the pit structure in 2011. .................................................... 12 

FIGURE S7: Picture of the excavators nearby the burial pit, after its excavation ......................... 12 

FIGURE S8: Picture of the pit after the 2012 excavation. ............................................................. 13 

FIGURE S9: Skeletal part representation of LCS1 after the 2011‐2012 discoveries. ..................... 14 

SUPPORTING TABLES ......................................................................................................................... 15 

TABLE S1: Faunal spectrums from Bouffia 118. ............................................................................ 15 

TABLE S2: Taphonomic modifications of the Alpha and C2 faunal remains from Bouffia 118. .... 15 

TABLE S3: Faunal spectra of the Bouffia Bonneval from the 2011‐2012 excavations. ................. 16 

TABLE S4: Faunal spectrum of the Bouffia Bonneval from the Bouyssonies’ excavations. .......... 17 

TABLE S5: Taphonomic alterations of the faunal remains from the Bouyssonie excavation. ...... 18 

TABLE S6: Taphonomic alterations of the faunal remains from the recent excavation. .............. 18 

TABLE S7: Potential primary and secondary Western European Neandertal burial. .................... 19 

REFERENCES FOR SUPPORTING INFORMATION ................................................................................ 20 

SUPPORTINGTEXTS

TEXTS1:Bouffia118zooarchaeologicalanalysis

Two archaeological levels have been identified in Bouffia 118: Alpha and underlying C2.

Alpha has yielded a large quantity of bifacial thinning flakes linked to the MTA techno-

complex (see 1). The faunal assemblage from level Alpha is largely dominated by Bison

remains (Table S1) presenting a high frequency of weathering modifications and root

etchings (Table S2). All the skeletal elements seem to have been introduced into the site. On

the other hand, the faunal spectrum of layer C2 is largely specialized on reindeer (Table S1)

and is associated with a Levallois technology produced on local and exogenous raw

materials. While deer remains are very well-preserved, they are systematically stained by

natural manganese deposits (Table S1). Skeletal part profiles suggest a highly selective

pattern with more than 90% of the NISP being limb fragments. A statistically significant

relationship exists between the skeletal part representation and the quantity of marrow and

grease contained within the different imported elements.

Although the material is yet to be radiocarbon dated, comparisons with other well-dated

sequences from Southwestern France containing the same faunal associations allowed a

biochronological approach to the two layers (Fig S3). During the Middle Paleolithic of

Southwestern France, reindeer dominated faunal spectrums occur only during MIS 4 and the

very beginning of MIS 3 and are followed by a 'Bison phase' at around 45ky represented by

an overwhelming dominance of this large bovine in faunal assemblages recovered from sites

attributed to this period (for a complete synthesis see 2).

TEXTS2:NewNeandertalremainsdiscoveredduringthe2011‐2012excavations

Thirteen new hominin remains have been discovered during our field work in 2011 and 2012.

Seven of them are attributed to the LCS 1 individual based on their morphology and the

preservation of the skeleton. Additional dental remains, an upper third molar from the

collections of the Musée de l’Homme (Paris, France), a molar root fragment, and an upper

second premolar, could belong to additional adult individuals (18). Finally, three deciduous

teeth represent additional juvenile specimens.

New LCS 1 remains:

1) A right upper second premolar (LCAS 2011 R73 # 12) - Fig S4A&B

This tooth is limited to the root (S4). The virtual extraction of the left upper first premolar on

LCS 1 shows that this tooth is two rooted and exhibits a taurodont morphology. The

morphological similarities between the tooth still in place on the LCS 1 maxilla and LCAS

2011 R73 # 12 doesn't exclude the association of the new dental root to the same individual.

Fig S4B provides a virtual extraction and reconstruction of the first premolar still in place on

the left upper dental arch of LCS 1 allowing a comparison with LCAS 2011 R73 # 12.

2) A fragment of a right scapula (LCAS 2011 S73 #322) - Fig S5A

The piece is part of a right mature scapula preserving most of the glenoid cavity, 3

centimeters of the axillary border and 70 mm2 of the adjacent body. The roots of the coracoid

and acromion processes are broken. The morphology of the broken parts indicates ancient

breaks. As scapular elements are missing from the LCS 1 individual, we consider that the

simplest hypothesis is that this scapula fragment belongs to the LCS 1 skeleton. Note the

clear insertion of the M. teres minor on the dorsal surface (Fig S5A).

3) A distal extremity of a right ulna (LCAS 2011 S73 #62) - Fig S5B

This piece is 33 mm long and represents a well- preserved radial articulation with part of the

diaphysis. The proximal part of the fragment has an irregular and recent break. The antero-

posterior and transverse diameters of the articular extremity are respectively of 23.7 mm and

22.1 mm. The minimal circumference of the diaphysis, measured just above the radial

articulation, is 33.5 mm and the maximum height of the articular circumference is 9.5 mm.

No direct refitting is possible between the broken distal extremity of the right LCS1 ulna and

the newly discovered fragment as the original right ulna was repaired using different kinds of

materials. Nevertheless, the shapes of the two pieces suggest that this distal extremity

derives from the same bone.

4) A distal extremity of the left third metacarpal (LCAS 2012 S73 #449) - Fig S5C

This piece is part of the distal extremity of a metacarpal. The radial part of the articular

surface is well-preserved, while the palmo-ulnar side is broken. The comparison of this distal

extremity with the original material allows to attribute this fragment to the left third metacarpal

of LCS1, whose distal extremity is missing and reconstructed. The size and morphology of

the new fragment correspond with this missing area. Moreover, there is a perfect secondary

anatomical connection with the proximal articular surface of the right third proximal phalanx.

Finally, a shallow ovoid depression is present on the articular surface. The same

characteristic can be observed on the distal articular surface of the right second metacarpal

from the original collection.

5) A proximal phalanx of the right thumb (LCAS 2011 R74 #109) - Fig S5D

This piece is 30.7 mm long and is well-preserved with the exception of the palmar surface of

the diaphysis which is broken. Two small fragments of compact bone from this area are

however still preserved. The phalanx is slender and the lateral crests are not marked which

is typical for most of the Neandertal proximal pollical phalanxes (19). No pathological

conditions are perceptible.

Following the morphology and size of the other hand phalanxes associated to LCS 1

individual (20), we consider that the simplest hypothesis is that this piece belongs to the

same specimen.

6) A distal phalanx of the right thumb (LCAS 2012 S73 S2 Rem 25-30 seau 2) – Fig S5E

This piece is partially broken with the medial side of the apical tuft and part of the diaphysis

missing. Given the taphonomy of the break, it could results from the 1908's excavation. This

phalanx presents a large proximal extremity in relation to its maximum length following the

Neandertal pattern (19). The insertion area of the muscle pollicis longus is well marked. Only

the apical tuft shows few Neandertal affinity with a gracile distal extremity. Finally, the

proximal articular surface fits perfectly with distal articulation of the new right proximal

phalanx.

7) A distal extremity of the left third metatarsal (LCAS 2012 S73 #568) – Fig S5F

This piece is highly deformed by bone remodelling. It is however possible to identify it as the

distal extremity of a metatarsal. The articular surface is eburnated and flattened. Its

comparison with the original collection allowed to bring it closer to the right third metatarsal.

The symmetry is not straightforward because of the pathological state of this distal extremity,

but its dimension and morphology closely resemble the antimeric bone from the original

collection.

Presumed new adult specimens:

1) A left upper second premolar (LCAS 2012 R73 #221) - Fig S4C

This tooth presents a mesio-distal fracture and its root is apically broken. The crown shows a

very high wear degree, the lingual aspect of the enamel being completely worn and the pulp

cavity open. Only the maximal bucco-lingual diameter of the root can be estimated to 11 mm.

Compared to the right upper second premolar LCAS 2011 R73 # 12 likely belonging to

LCS1(see above), this new tooth has a shorter root with a morphologically different apex.

Thus, this tooth does not belong to LCS1, and represents a new adult individual at La

Chapelle-aux-Saints.

2) A right upper third permanent molar discovered in 1920 – Fig S4D

This right upper third molar was discovered September 7th, 1920 by a cousin of Bouyssonie

visiting the bouffia Bonneval who entrusted it to Jean Bouyssonie. It was recognized by the

latter as a human tooth. As he suspected it might belong to the LCS 1 fossil, he sent it to H.

Breuil (letter available at the Central library of the Muséum national d’Histoire naturelle, Paris

- Henri Breuil archives - classification mark Br 28) to give it to M. Boule. The tooth has been

in the fossil collections of the Musée de l’Homme (Paris) ever since (18).

Unfortunately, the upper alveolar arch of LCS 1 is poorly preserved and it is not possible to

test if this large third molar could refit.

3) A distal root of a lower permanent molar (LCAS 2011 R73 #26) - Fig S4E

This root of a human permanent molar bears a recent fracture on one face. The outward

orientation and distal curvature indicate that this root is a distal one of a lower molar. The

asymmetry of the lingual and the buccal edges (the buccal one is higher and less convex

than the lingual one) and the orientation of the apex suggest a left root. The slightly oblique

angle of the root relative to the inferred occlusal or cervical plane suggests that this root

cannot be a third molar and is probably more likely to be a second or a first molar.

This root belongs to either an adult or a child older than 12-13 years. There is substantial

resorption of the back part of the alveolar bone on both sides of the LCS 1 mandible. Such a

resorption is associated with a severe parodontopathy. No lower permanent molar would still

have been in place in the mouth of this Neandertal for at least several months before its

death. Thus, this fragment of a root of a lower left permanent molar cannot be attributed to

LCS1 and therefore represents another individual, like the one represented by the left upper

second premolar LCAS 2012 R73 # 221 (see above).

New juvenile specimens:

1) A right upper deciduous central incisor (LCAS 2011 S73 Rem 10-15 seau 1) – S4F

This tooth presents a complete functional crown and an apically broken root. The labial

surface is convex in both the vertical and horizontal planes. The mesial border is less convex

than the distal one which is also shorter. The lingual surface is shovel-shaped presenting a

well-developed mesial (higher) and distal (wider) margin and a clear lingual tubercule. The

mesial margin almost reaches the cingulum while the distal one is separated from it by a

deep surface. The crown occlusal border presents light attrition and there is also a very small

mesial interproximal contact facet at the mid-height of the crown. Considering the occlusal

wear, this tooth could have belonged to a child, circa 3 years old, younger than the Roc-de-

Marsal 1 and older than the Pech-de-l’Azé 1 specimens (21, 22). The crown mesio-distal

diameter is 8.17 mm and the bucco-lingual is 6.25 mm.

2) A right upper deciduous second molar (LCAS 2012 T74 S3 Rem 15-20 seau 1 & T73

S2 Rem 10-15 seau 2) - Fig S4G

This new tooth consists of two fragments from adjacent squares, one from T74 representing

the three quarters of the lingual crown, and the other one a small buccal portion from T73.

There are large fractures between the two fragments. The crown shows high occlusal wear

(6-1 following 23), and large interproximal contact facets. The beveled edge of the small root

fragment suggests that the tooth has been lost during the life of the individual. Following 24,

this tooth could have belonged to a 11-12 years old child. The crown mesio-distal diameter is

estimated to 8.83 mm, and the bucco-lingual to 11.50 mm.

Given their different wear patterns, this new tooth and the right upper deciduous central

incisor LCAS 2011 S73 Rem 10-15 seau 1 (see above) cannot have belonged to the same

juvenile individual.

3) A right lower deciduous first molar (LCAS 2012 Q74 S1 Rem 50-55 seau 1) - Fig S4H

This is a well-preserved crown showing a high occlusal wear (5-2 following 23), and

pronounced interproximal contact facets. A small fossa on the mesial aspect of the

protoconid recalls the morphology shown by Rescoundudou 2 (25), and Pech-de-l'Azé 1

(26). Following 24, this tooth could have belonged to a 10 years old child. The crown mesio-

distal diameter is 8.18 mm, and the bucco-lingual is 7.67 mm.

Given its wear degree, this new tooth could belong to the same individual as the right upper

deciduous second molar LCAS 2012 T74 S3 Rem 15-20 seau 1 & T73 S2 Rem 10-15 seau

2.

At least two juvenile individuals are recognized at La Chapelle-aux-Saints, one represented

by an upper deciduous central incisor, and the other one by an upper deciduous second

molar and a lower deciduous first molar.

SUPPORTINGFIGURES

FIGURES1:Sitelocationandpictureofthesiteentrance.A: Map From Geoatlas. B: the picture was taken from the base of the slope in front of the cave entrance before the 2011 excavation.

FIGURES2:M

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FIGURES3:MousteriansiteswithfaunalspectradominatedbyBisonorReindeerattributedtotheMIS4ortheMIS31: Saint-Césaire, Layer EGPF (3); 2: Marillac-Les Pradelles, Layers 9 and 10 (4); 3: Jonzac, Layers 22, 20, 18, 16, 14, 12 and 9 (5-6); 4: La Quina, Layer 8 (7), Layers 6a and 6c (8-9); 5: Roc-de-Marsal (10); 6: Combe Grenal, Layers 17 and 16 (11); 7 : Pech de l’Azé IV, Layers I and H (12); 8: Sous-les-Vignes (13); 9: Puycelsi-La Rouquette, Layer 1 (14); 10: Mauran, (15-17). Cultural and radiochronological attribution are discussed and synthesized by (2).

FIGURES4:Humandentalremainsdiscoveredduringthe2011‐2012excavationsandduringthe1920’s.A, C, E-H dental remains discovered during the 2011-2012 excavation. D: M3 discovered in 1920. B : virtual extraction and reconstruction of the first premolar still in place on the left upper dental arch of LCS 1 allowing a comparison with LCAS 2011 R73 # 12

FIGURES5:Humanosseousremainsdiscoveredduringthe2011‐2012.

FIGURES6:Firstexcavationstageofthepitstructurein2011.The dark sediment corresponds to the Bouyssonie’s backfill; the light corresponds to the marl. The dotted line marks the pit limit.

FIGURES7:Pictureoftheexcavatorsnearbytheburialpit,afteritsexcavationLeft: Jean Bouyssonie and Father Robert visiting the site just after the discovery; Right: current excavators in 2011. The white line on the two pictures underlines the same relief of the cavity roof. The pictures were taken from the entrance of the cavity. The 1909 picture comes from the Fonds Bouyssonie-Ecole Bossuet.

FIGURES8:Pictureofthepitafterthe2012excavation.Backfill can only be seen now at the extremity of the cavity and the substratum is exposed on the totality of the excavated area. The picture was taken from the entrance of the cavity in September 2012.

FIGURES9:SkeletalpartrepresentationofLCS1afterthe2011‐2012discoveries.Grey: Skeletal elements identified in 1908; Red: Skeletal elements from the 2011 excavation. Drawing from: URA 376 CNRS after S.T. Constandse - Westermann and C. Meikeljohn, modified by M. Guillon, P. Sellier and P. Courtaud. Digitized by M. Coutureau (INRAP).

SUPPORTINGTABLES

TAXA ALPHA C2

Fox 7 2

Hyena 1

Wild cat 1

Reindeer 35 605

Roe deer 1

Cervid 1

Bovine 140 78

Horse 2

Avifauna 3

Marmotte 1 1

NISP 189 689

Medium sized ungulates 2 11

Large sized ungulates 226 73

Mammals 50 46

Total 467 819

TABLES1:FaunalspectrumsfromBouffia118.Faunal spectra established on the two main layers from bouffia 118. Alpha is the upper layer, dominated by bovine remains; C2 is the lower layer, dominated by reindeer.

Taphonomic

modifications

Alpha C2

NR %NR NR %NR

Longitudinal cracks 104 22,27% 237 28,94%

Exfoliation 163 34,90% 169 20,63%

Flaking of the outer

surface 65 13,92% 72 8,79%

Smoothed bone 98 20,99% 58 7,04%

Manganese deposits 11 2,36% 115 14,05%

Carnivore modifications 26 5.07% 51 6.23%

TABLES2:TaphonomicmodificationsoftheAlphaandC2faunalremainsfromBouffia118.Significant differences can be observed in the preservation pattern between the upper layer (Alpha) and the lower one C2 particularly if we considered the frequencies of smoothed bones and manganese deposits.

Taxa Backfill C4 C2sup C5 Wolf 3 1 Fox 61

Hyena 3 Felid 3

Badger 60 Polecat 0 1

Carnivore 2 Hyduntinus 0 1

Horse 56 7 3 Ovicapra 9

Capra 1 Bovine 675 87 1 1

Roe deer 5 3 Reindeer 4225 439 98 19 Red deer 7 1 Giant deer 1 Wild boar 13 Marmot 8 7

Lagomorph 101 NISP 5233 538 111 20

Large size ungulate 751 84 3 4

Mammal 457 36 2

Avifauna 78 NR 6519 658 114 26

TABLES3:FaunalspectraoftheBouffiaBonnevalfromthe2011‐2012excavations.C4: material from the non previously excavated area; C2sup: material moved into the marl by the cyroturbation. C5: faunal remains from the cryoclastic feature cut by the pit. The Backfill assemblage corresponds to what had been discarded by the Bouyssonies during their excavation.

Bouyssonies' excavation TAXA NISP Bear 1 Wolf 1 Fox 4

Horse 25 Rhinoceros 3 Ovicapra 5

Capra 3 Bovine 915

Reindeer 529 Red deer 12 Giant deer 8

Cervid 19 Wild boar 1 Marmot 1

Lagomorph 6 NISP 1533

Large size ungulate 255

Mammal 53 NR 1841

TABLES4:FaunalspectrumoftheBouffiaBonnevalfromtheBouyssonies’excavations.We have here to keep in mind that the sampling was highly selective and large taxa might be over-represented in this collection. It probably explains the main differences between the Bouyssonies collection and the material yielded by the backfill excavation.

Taphonomic alterations Reindeer Bison Neandertal NR % NR % NR %

Longitudinal craks 220 41,59 439 47,98 5 6,49 exfoliation 233 44,04 241 26,33 0 0 freezing modification 46 8,69 116 12,68 0 0 smouthed bone 196 37,05 180 19,68 4 5 Manganeze deposit 42 7,94 34 3,72 6 8

Bonesurfacealteration0 - 1 260 55,32 85 41,67 77 100

Bonesurfacealteration2 89 18,94 60 29,41 0 0

Bonesurfacealteration3 75 15,96 45 22,06 0 0

Bonesurfacealteration4 46 9,79 14 6,86 0 0

Carnivoremodifications 21 4 47 5,1 0 0

TABLES5:TaphonomicalterationsofthefaunalremainsfromtheBouyssonieexcavation.Bone surface alteration pattern refers to the stage of preservation of the cortical surface of the remains (following (16)). 0-1: less than 25% of the remain surfaces are altered, 2: less than 50% of the remain surfaces are altered. 3: less than 75% of the remain surfaces are altered, 4: more than 50% of the remain surfaces are altered. Major differences can be observed between the different analyzed taxa. Human remains are significantly less altered than the faunal ones. Bison and reindeer exhibit also two different patterns of preservation: with for instance frequencies of exfoliation and manganese deposits higher on reindeer remains than on bison ones.

Taphonomic alterations Reindeer Bison NR % NR %

Longitudinal craks 150 11,82 46 20,91 exfoliation 418 32,94 73 33,18 freezing modification 111 8,75 17 7,73 smouthed bone 81 6,36 18 8,18 Manganeze deposit 180 32,07 19 19,09 Bone surface alteration 0 - 1 730 62,07 115 59,89 Bone surface alteration 2 211 17,94 32 16,67 Bone surface alteration 3 99 8,42 23 11,98 Bone surface alteration 4 136 11,56 22 11,46 Carnivore modifications 30 2,4 9 4,1

TABLES6:Taphonomicalterationsofthefaunalremainsfromtherecentexcavation. Sample made from the 2011 material. Differences between the two collections might be explained by the fact that part of the recent one is composed by very small fragments which have a smaller cortical surface and, thus, have a lower probability to exhibit carnivore marks.

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(34

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(3

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s P

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9)

TABLES7:PotentialprimaryandsecondaryWesternEuropeanNeandertalburial.

The

tab

le c

ompi

les

the

avai

labl

e in

form

atio

n fo

r th

e W

est

Eur

opea

n si

tes

whe

re t

he h

ypot

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s of

Nea

nder

tal b

uria

l has

bee

n ad

vanc

ed b

y pr

evio

us r

esea

rche

rs. C

olum

n 11

list

s th

e an

alys

es fo

r w

hich

re

sear

ch m

ust

be u

nder

take

n (a

nthr

o =

mis

sing

info

rmat

ion

abou

t th

e sp

atia

l di

strib

utio

n of

the

hum

an r

emai

ns).

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