New Neanderthal remains from the Châtelperronian attributed layer X of the Grotte du Renne (Arcy-sur-Cure, France)

Short Communication

Juliette Henrion a,*, Jean-Jacques Hublin b, Bruno Maureille a

a PACEA UMR 5199, Univ. de Bordeaux – CNRS – Ministère de la Culture, 33600, Pessac, France

b Paléoanthropologie, Collège de France, 75231, Paris, France

E-mail address: Juliette.henrion@u-bordeaux.fr (J. Henrion)

**Keywords:** Dental morphology; Tooth crown; EDJ; Premolar; Molar; Hyoid

# 1. Introduction

## 1.1. *The Grotte du Renne location and geological environment*

The Grotte du Renne (GR) is located on the township of Arcy-sur-Cure (Yonne, France). The Cure River, which crosses the town, contributed to the weathering of the Arcy-Saint-Moré limestone formation and to the development of complex karstic systems. Among these karstic systems, the Arcy-sur-Cure caves open southwards on the left bank of the Cure. The caves have been visited since the 17th century (Baffier and Girard, 1997, 1998; Girard, 2019). Research into Arcy's prehistoric remains began with A. Parat's excavations between 1897 and 1901. (Parat, 1901). However, scientific investigations started with A. Leroi-Gourhan's excavations (1948–1963), which uncovered Upper Pleistocene human remains within four caves: the Grotte du Loup, the Grotte du Bison, the Grotte du Renne and its attached Schoepflin gallery, and the Grotte de l'Hyène.

The GR, documented by P. Poulain in 1938, was excavated by A. Leroi-Gourhan's team between 1949 and 1963 (Leroi-Gourhan and Leroi-Gourhan, 1964; Girard, 2019). The cave extends over 80 m² and is filled by a seven-meter-thick sedimentary sequence,

<footnote>*Corresponding author.</footnote>
characterized by clayey and sandy horizons with embedded limestone slabs and clasts (Baffier and Girard 1997, 1998; Roblin-Jouve, 2019). Below the Holocene soil, 13 archaeological layers have been identified and attributed to five typo-chronological units, including from the base to the top: the Mousterian (XV to XI), the Châtelperronian (X to VIII), the Aurignacian (VII), the Gravettian (VI to IV), and the Magdalenian (III; Girard, 1980). Materials from layers XII (ca. 46 ka cal BP) to layer V (ca. 27.5 ka cal BP) have been dated via various radiocarbon methods. The Châtelperronian attributed layers date to between 44.5 and 41 ka cal BP for layers X–IX and to between 41 and 39.5 ka cal BP for layer VIII (Higham et al., 2010; Zilao et al., 2011; Hublin et al., 2012; summarized and reassessed by Banks and d’Errico, 2019). One hominin bone fragment (AR-14, from layer X) has been directly dated to 42.56–40.46 ka cal BP (OxCal, 95.4%; Welker et al. 2016). The GR Châtelperronian unit has undergone more thorough investigations than the Mousterian layers (Leroi-Gourhan, 1961; David, 1990, 2001; Rocca et al., 2017), and this with a particular focus on the Middle-to-Upper Paleolithic transition due to the numerous hominin remains associated with long-term Châtelperronian occupations—occupation floors with traces of habitats and hearths, as interpreted by Leroi-Gourhan (1961) and Hublin et al. (2012). However, the later hypotheses and the attribution of the Neandertal remains to the Chatelperronian material have been challenged due to vertical post-depositional movements, evidenced by numerous outliers in radiocarbon dating of the faunal remains (Bertran et al. 2010; Higham et al. 2010; Discamps et al. 2015).

## 1.2. Grotte du Renne human remains

The first human remains were found in the GR in 1949 (Leroi-Gourhan, 1958), and as of 1963, 76 human fossils have been recovered. These fossils are curated at the Musée National de Préhistoire (Eyzies-de-Tayac, France), along with the majority of Arcy-sur-Cure's Paleolithic collections resulting from Leroi-Gourhan's fieldwork efforts. Of the 76 human fossils, 73 have been partially studied and/or inventoried in five publications. In 1958, Leroi-Gourhan described seven teeth (i.e., including two teeth later determined to be cave bear lower incisors; Bailey and Hublin, 2006; Maureille and Hublin,
2019). A temporal bone (Hublin, 1996) and 30 isolated teeth (Bailey and Hublin, 2006) were
each the subject of morphological analysis with taxonomical considerations. Welker et al.
(2016) identified 28 cranial (likely belonging to the same individual) and infracranial
fragments among the faunal and human remains via a zooarchaeology by mass spectrometry
analysis, along with proteomic (on AR-16 and AR-30) and genetic (on AR-14 and AR-30)
analyses.

In 2019, an exhaustive inventory was conducted on the GR anthropological
collection, including the brief description of 61 specimens from the Châtelperronian layers
(Maureille and Hublin, 2019). Since then, the GR faunal collections were inspected on the
initiative of B. M. to ensure the absence of unidentified hominin remains. This work led to
the identification by B.M. and C. Beauval of two teeth (Fig. 1) and a greater horn of a hyoid
(Fig. 2) from the lithostratigraphic horizon Xb2 (where most of the human remains in the
Châtelperronian layer X have been found; Leroi-Gourhan, 1952; Leroi-Gourhan and Leroi-
Gourhan 1964). A left P4 germ (Arcy 1952 R Xb2 Y13) was found within the hominin
collection of the Musée national de Préhistoire (Eyzies-de-Tayac, France). Another left dP4
germ (Arcy 1963 R Xb2 B6 460) and the greater horn of a hyoid were identified among the
few remains still stored at a private property in Venoy (France).

The aims of this paper are to describe the three newly identified human fossils and to
evaluate their taxonomic affinities. We compare the two fossil teeth to published GR fossils,
focusing on their morphological and metrical similarities. The pairing of the new left dP4
with a previously published tooth (Bailey and Hublin, 2006) also allows us to discuss the
taphonomic integrity of the GR Châtelperronian layers. Lastly, we introduce the third greater
horn of a hyoid known from the Eurasian Early Upper Pleistocene besides the right and left
ones from the Kebara 2 Neanderthal (Arensburg et al., 1989).

**2. Materials and methods**

2.1. *Comparative samples*

The two newly discovered teeth (Arcy 1952 R Xb2 Y13 and Arcy 1963 R Xb2
B6 460) were morphologically compared to their homologs within the GR collection. The
metric comparative samples include published teeth from Arcy-sur-Cure (n = 12), in addition
to a Neanderthal (n = 130) and an Upper Paleolithic modern human sample (n = 163) gleaned
from the literature and PACEA scientific members' database (Supplementary Online Material
[SOM] Table S1). Comparative material for a 3D geometric morphometric analysis of the P4
enamel-dentine junction (EDJ) consisted of a sample of semi-landmark coordinates shared by
C. Zanolli. The semi-landmarks were acquired from digital reconstructions of microcomputed
tomographic (μCT) scans of Neanderthals (n = 9) and modern humans (n = 9;
SOM Table S2). For the greater horn of the hyoid, our metric comparative sample includes
Late Neolithic modern humans from France, housed at PACEA ('L'Aven de la Boucle', n =
13; 'Le dolmen des Peirières', n = 9; Duday, 2009) and the CT scan of the original Kebara 2
hyoid bone from the Max Plank Institute for Evolutionary Anthropology (Leipzig, Germany)
database for the qualitative comparisons.

<figure><img src="image_1.png" /><figcaption>**Figure 1.** The two new teeth from the GR Châtelperronian sub-layer Xb2. The left dP4 germ (Arcy 1963 R Xb2 B6 460), showing a) from left to right the occlusal enamel surface (OES) in occlusal, distal, lingual, mesial, and</figcaption></figure>
buccal views, b) the enamel-dentine junction (EDJ) in occlusal (left) and distal (right) views, c) the mesial histological section, and d) the hypoconulid (C5) twinned dentin horn detail (white arrows). The left P4 germ (Arcy 1952 R Xb2 Y13) with e) from left to right the OES in occlusal, lingual, mesial, buccal, and distal views, and f) the EDJ in occlusal (left) and lingual (right) views, g) the histological section, and h) the close-up of the OES taphonomic alterations. Scale bar of 5 mm.

<figure><img src="image_2.png" /><figcaption>**Figure 2.** A left greater horn of the hyoid (Arcy 1962 R Xb2 C10 1) from the GR Châtelperronian sub-layer Xb2 in a) lateral and b) medial views. Abbreviations: P = proximal; D = distal. Scale bar of 5 mm.</figcaption></figure>

## 2.2. *Data acquisition*

Micro-computed tomographic scans were acquired at PLACAMAT (University of Bordeaux and CNRS, France), with an isometric voxel size of 21 µm for the two newly discovered teeth (100 kV, 300 µA, 0.1mm copper filter) and 22 µm for the greater horn of the hyoid for the Arcy and the modern human sample (100 kV, 120 µA, 0.1mm copper filter).

The published human remains from the GR were scanned at the Max Plank Institute with a 16 µm resolution (100 kV, 62 µA, 0.1mm aluminum filter), and the Kebara 2 hyoid at 31 µm (130 kV, 100 µA, 0.25mm copper filter). We segmented the µCT scans semi-automatically in Avizo v. 8.1 (Visualization Sciences Group Inc., Berlin). We edited the segmentation manually because of the low mineralization of tooth enamel and the presence of sediments (with grey levels similar to the dentine and bones).

Descriptions of the teeth include the preservation, health status, determination of age at death, and morphologies of the occlusal enamel surface (OES) and EDJ. Virtual histological sections following Smith et al. (2011) were taken with VG StudioMax v. 3.5.0 (Volume Graphics GmbH, Heidelberg), through the mesial cusps of the teeth to observe the taphonomic, health, and developmental status of the teeth. The age at death of the fossil individuals was determined in accordance with modern human standards for bone maturation
(Gray et al. 2021), dental development, and dental eruption (Moorrees et al., 1963; Liversidge and Molleson, 2004; AlQahtani et al. 2010, 2014; Roksandic and Armstrong, 2011; Shackelford et al. 2012). Occlusal enamel surface and EDJ morphologies were described following the terminology of Scott and Irish (2017). Discrete OES and EDJ characters were scored and compared to published Neanderthals and Upper Paleolithic modern humans (for LM2, LdP4 and LP—excluding the P4 EDJ; Bailey, 2002a, b, 2006; Bailey et al., 2011; Martinón-Torres et al., 2012, 2014; Martínez de Pinillos et al., 2014; Martin et al., 2017; Scott and Irish, 2017; Davies et al., 2019). The greatest mesiodistal and buccolingual diameters at the OES (measured three times each by J.H. and B.M.) and the crown index were acquired (Martin and Saller, 1957). We used 3D geometric morphometrics to compare the shape of the P4 marginal ridge at the EDJ to the modern human and Neanderthal samples (SOM Table S2). With the B-spline function in Avizo v. 8.1, we placed two sets of semilandmarks:

one on the mesial and the other on the distal part of the marginal ridge, starting and ending with the horn tips (SOM Fig. 1). Each interpolated curves from the semilandmark sets were used to resample in R v. 4.2.2 (R Core Team, 2022) a total of 177 semi6 landmarks around the dentin ridge (Pan et al., 2017, Zanolli et al., 2019). Principal component analysis (PCA) was used to consider the comparative group structure along the PC scores and to assess the position of Arcy P4 in this morphospace, with a discriminant analysis used to investigate group attributions probabilities.

The description of the greater horn of the hyoid was conducted following Gray et al. (2021) and included six measurements (SOM Fig. S2). The hyoid measurements were taken three times each by two of the authors (J. H. and B. M.) using a 150 mm Mitutoyo digital calipers (0.01 mm accuracy) and the means were used for metric comparisons.

## 3. Results

### 3.1. *The left dP4 germ* (Arcy 1963 R Xb2 B6 460)

The crown (Fig. 1a) is well preserved except for the small chips missing from the cervico-buccal edge and the protoconid apex. We observed some taphonomic demineralization of the enamel through the cracks and the intercuspal grooves. Due to the
taphonomic alteration of the crown surfaces we were unable to visually distinguish any pathological manifestation (pitted or linear hypoplasia), but on the tomographic acquisitions a stress line was noticeable near the cervical edge (Fig. 3a, b). Moreover, the cusp apices have pit-like deformations and the marginal ridges have groove-like deformations (Fig. 3c), which are not present on the EDJ (Fig. 1b). These defects occurred during the early stage of development or the enamel's maturation process.

The tooth crown is complete, but we observed no root development (Fig. 1a, c). We thus estimated the age at death of this individual between 4.5–10.5 months (infancy).

The crown outline is oval. The lingual cusp tips are internally compressed with the metaconid oriented towards the sagittal sulcus. The fissure organization follows a Y pattern.

The anterior fovea is deep (grade 3) and is bordered by a high marginal ridge mesially (including a lingual tuberosity) and an interrupted mid-trigonid crest (grade 1) distally. The latter has a developed accessory cuspule near the protoconid. The five main cusps are well developed (protoconid > metaconid > entoconid > hypoconid > hypoconulid). A sixth cusp is also present and is larger (grade 2) than the hypoconulid (grade 4). There is a cuspule between the C5 and C6. No C7 is observed, but the metaconid mesial ridges and the entoconid lateral ridges are complex. Although there is no protostylid, the buccal aspect of the protoconid has an incipient shelf (Skinner et al., 2008).

At the EDJ, the mid-trigonid crest is continuous but dips in the middle (grade 2). The accessory cuspule on the mid-trigonid crest has a high, sharp dentine horn. The metaconid dentine horn is strongly internally oriented. The C6 has a distinct dentine horn that is lower than the main cusps. Moreover, the hypoconid dentine crest has a distal protrusion and the hypoconulid has a twinned horn tip (Fig. 1d; Martin et al., 2017). No C7 is present at the EDJ, however there are weak ridges on the distal aspect of the metaconid. Lastly, the taloninid is formed by the disto-lateral ridges of the entoconid and hypoconulid, which are separated by the central sulcus (OES and EDJ talonid expression: grade 0).
<figure><img src="image_3.png" /><figcaption>**Figure 3.** From left to right, the Grotte du Renne left dP4 (Arcy 1963 R Xb2 B6 460) and right dP4 (Arcy 1963 R Xb2 C8) with a) mesial developmental slices showing stress lines (white arrows) and b) buccal faces showing hypoplasia corresponding to the stress lines (white dotted lines). The bottom figure (c) highlights the groove and pit-like deformation on the apics of the crown structures for both molars. Scale bars of 1 mm.</figcaption></figure>

## 3.2. *The left P4 germ* (Arcy 1952 R Xb2 Y13)

The enamel surface appears blunt and the crown is cracked (Fig. 1e, h), with considerable chipping on the protoconid marginal ridges. All the cracks are oriented along the Hunter-Schreger bands and dentine tubules (Fig. 1g). This condition is probably the result of taphonomic desiccation of an immature enamel, noticeable due to low enamel density and light grey line at the EDJ. A stress line is also visible near the EDJ, especially in the center of the crown. The cervical part of the crown is incompletely developed (Fig. 1e, g). Thus, the age at death of this specimen is estimated between 4.5–5.5 years (early childhood).

The crown outline is fairly symmetric in occlusal view, with a round lingual edge, almost squared, and a trapezoidal buccal edge. Buccally, the crown is superoinferiorly flat,
nearly concave, and without any groove or ridges. The protoconid has a short and wide
mesiomarginal ridge (cusp-like) and a distomarginal ridge that extends nearly to the
distolingual corner of the crown. The metaconid is in mesial position (grade 1), and a small
distolingual accessory cusp is present (grade 2). The lateral ridges of these two main cusps
are well developed, but there is no discernable transverse crest (grade 0).

At the EDJ (Fig. 1f), both the protoconid and metaconid have faint crests (grade 1a
transverse crest). The mesiolateral ridges of the buccal and lingual cusps (especially
developed on the metaconid) define the distal border of a deep anterior fovea. The central
groove is narrower than the mesial and distal foveae. On the EDJ, two accessory ridges are
noticeable on the central part of the distomarginal ridge, and two horn-like tuberosities are
present on the mesiobuccal and distobuccal part of the protoconid marginal ridges. The
marginal ridge is almost continuous, joining at the protruding metaconid dentine horn.

3.3. *The greater horn of the hyoid (Arcy 1962 R Xb2 C10 1)*

The greater horn of the hyoid is incomplete (Fig. 3); the distal extremity is missing,
and the inferior edge is slightly damaged. The proximal articular facet is well preserved and
represents the tallest sagittal section of the horn. The superior edge of the greater horn is
concave. The inferior edge is flat from the proximal extremity until the mid-section of the
bone, where the distal end of the thyrohyoid membrane insertion marks an upward deflection
of the horn.

The thyrohyoid membrane insertion produces a concavity on the lateral side of the
bone while the medial face is flatter. Thus, the greater horn can be identified as belonging to
the left side. There are no attachment sites for the hyoglossus and middle pharyngeal
constrictor muscle insertions. Considering the variability of the greater horn of the hyoid
within extant humans (Arensburg et al., 1989), we propose that this bone has reached its full
adult size. The absence of secondary bone modifications frequently seen on extant human
older adults suggests that the individual may have been a young adult.

**4. Discussion and conclusions**
The size of the dP₄ (Arcy 1963 R Xb2 B6 460) is within the range of both Neanderthals and Upper Paleolithic modern humans (SOM Tables S3 and S4). However, its discrete features clearly align with Neanderthals (e.g., Bailey, 2006; Bailey and Hublin, 2006). These features include a Y cusp pattern (82.1% for *n* = 28; Martinón-Torres et al., 2012); a continuous mid-trigonid crest (53.3% for *n* = 30; Martinón-Torres et al., 2012); a large anterior fovea (87.5% for *n* = 32; Martinón-Torres et al., 2012); the presence of a sixth cusp (48.3% for *n* = 29; Martinón-Torres et al., 2012); the anterior fovea expression at the EDJ (Martinón-Torres et al., 2012; Martínez de Pinillos et al., 2014); and twinned dentine horns on the hypoconulid (e.g. Martin et al., 2017). Based on this combination of traits (SOM Table S5), we consider this newly found dP₄ as belonging to a Neanderthal individual.

The new dP₄ was found in the south-east area of the cave (SOM Fig. S5), two meters from previously described teeth attributed to two individuals who died within their first year of life (Bailey and Hublin, 2006; Maureille and Hublin, 2019). They include a right dP₄ (Arcy 1963 R Xb2 C8), which is very similar in size and morphology to the new left dP₄ (Fig. 3). In particular, these two teeth have the same unusual enamel deformations on the cusp apices and ridges (Fig. 3c) as well as a linear hypoplasia in the same area (Fig. 3a, b). Therefore, it is highly probable that those teeth are antimeres from the same Neanderthal individual.

The left P₄ crown (Arcy 1952 R Xb2 Y13) is not fully developed; thus, its dimensions are considerably smaller than they would be if the crown were complete (SOM Tables S3 and S6). Nonetheless, we observe strongly expressed features such as developed buccal marginal ridges, and marked anterior fovea and distal groove (SOM Table 5). Such features are expressed in Neanderthal P₄s (Bailey, 2006), along with a high frequency of mesially-placed metaconid and multiple lingual accessory cusps (96.9% for *n* = 32 and 90.6% for *n* = 31; Bailey, 2006). The new Arcy premolar combines a mesially placed metaconid (Neanderthallike)

with a symmetrical outline and no transverse crest, both observed at low frequencies in Neanderthals (both at 6.5% for *n* = 31 in Bailey 2006). The previously published lower second premolar from the GR ('Tooth no. 4' in Bailey and Hublin, 2006) presents those traits, with a highly asymmetric crown (its distolingual accessory cusp is as large as the main cusps). But despite its crown symmetry, the new P₄ OES morphology is closer to the
Neandertal P4.

There are no published inter- or intraspecific comparative data on the P4 EDJ
morphological traits, only a geometric morphometric study by Pan and collaborators (2020).
The PCA analysis we conducted on dentine marginal ridge shape (SOM Fig. S3) places the
new premolar within the Neanderthal sample distribution, with a posterior probability of
100% based on the discriminant analysis. Its morphology is closest to two Krapina teeth:
KRD26, which has a rounder marginal ridge, and KRD30, which has a squared outline that is
asymmetric buccodistally (SOM Fig. S4a, b). The new GR premolar distolateral ridges are
similar to KRD30 for both main cusps, and its lingual horn tip is high relative to the buccal
one, similar to 'Tooth 4' from the GR (SOM Fig. S4c). However, these comparative teeth
(SOM Fig. S4) have a transverse crest that connects to the lingual margin and flattens at the
occlusal basin (Type 1a, adapted from Davies et al. 2019).

<figure><img src="image_4.png" /><figcaption>**Figure 4.** Digital reconstructions of a left greater horn of the hyoid of a) a modern human (BC93 H17 C4b 328), b) Arcy (A1962 R Xb2 C10 1), and c) Kebara 2 Neanderthal, and their morphological comparison by manual alignments (with the superior edge of the proximal articulation, the angle, and the distal extremity) with d) a</figcaption></figure>
modern human (BC93 H17 C4b 328) and Arcy (A1962 R Xb2 C10 1), 26 e) modern human (BC93 H17 C4b 328) and Kebara 2, and f) Arcy (A1962 R Xb2 C10 1) and Kebara 2. Abbreviations: P = proximal; D = distal.
Scale bar of 5 mm.

Hyoid bone dimensions and morphology have remained stable since the Middle Pleistocene, according to previous studies on individuals from La Sima de los Huesos, Kebara, and Castel di Guido (Martínez et al., 2008; D'Anastasio et al., 2013; Capasso et al., 2016). The only greater horn of the hyoid preserved within the Middle to Upper Pleistocene fossil record are the left and right greater horns of the Kebara 2 Neanderthal (Arensburg et al., 1989). Compared to the GR greater horn, their thyrohyoid insertion sites are more prominent, with more concave lateral faces (Fig. 4a-c.). The dimensions of the Kebara 2 hyoid bone fall within the range of modern human variation according to Arensburg et al. (1989); we reached the same conclusion for the GR greater horn of the hyoid (SOM Table S7; Fig. 4d-f). Hence, the taxonomical status of the GR greater horn cannot be determined from morphological and metric data.

In conclusion, we ascribe the newly rediscovered human dP4 and P4 from the base of the Châtelperronian sequence (layer Xb2) as belonging to Neanderthals. The pairing of isolated dP4s as antimeres can be considered as supporting evidence for the taphonomic integrity of layer X, suggesting that there has not been significant disturbance in the area where most of the immature remains were discovered. Lastly, based on the taxonomic attribution of the dental remains and the taphonomic evidence presented above, we believe it is most parsimonious to consider the newly discovered left great horn of the hyoid, the second known among the Late Pleistocene fossil record (i.e., MIS 5 to 3), as belonging to a Neanderthal. However, there are no morphological or metric characteristics that allow us to distinguish it from those of our modern human sample.

## Acknowledgments

We are grateful to N. Fourment, head of the Musée National de Préhistoire, for the access to Arcy-sur-Cure human collection and to C. Beauval for his expertise in the investigation of the faunal collection of Arcy-sur-Cure caves. We would like to thank H. Duday for the access to a comparative sample of greater horns of the hyoid from his
fieldwork. We are thankful to N. Vanderesse (PLACAMAT) for the scanning of the new
hominin remains, and to H. Temming (MPI) for the CT scans of the Grotte du Renne fossils.
For the access to Kebara 2 hyoid bone CT scan we thank Y. Rak, A. Barash, E. Been, A.
Winzer, and A. Le Cabec. We are also thankful to A. Le Cabec and C. Zanolli for their help
and support. For access to comparative data for the geometric morphometric analysis of the
EDJ, we sincerely thank C. Zanolli. This research is a part of a Ph.D. project supported by a
MITI CNRS grant and is funded by the mission for transverse and interdisciplinary
initiatives of the CNRS and by the Nouvelle-Aquitaine regional project: ‘ADNER’,
convention no. AAPR2021-2020-11779310. This research benefited from the scientific
framework of the University of Bordeaux's IdEx ‘Investments for the Future’ program/GPR
‘Human Past’, and the UMR5199 EuraPal and EvoDiBio teams. Finally, we are deeply
thankful to the Editor-in-Chief and the three reviewers for their useful comments and
suggestions which have greatly enhanced the quality of our original manuscript.

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