Catalog of Neanderthal Remains Sites, 4 of 4

Neanderthal fossils

Changes in temperature, water flows, and geological erosion gradually destroy the remnants of Neanderthals and traces of their material culture. Neanderthal fossils aged 50–40 thousand years differ from more ancient ones in terms of better preservation of remains. This allows for the investigation of DNA from such findings and the acquisition of previously unavailable information.

The website Dinoera presents a catalog of Neanderthal fossils sites with brief descriptions and informative illustrations of significant discoveries. The study of the history of Neanderthal man continues.


Šipka cave. Czech Republic. Near Štramberk, Moravia. Fragment of the mandible of a Neanderthal child aged 9–10 years. The finding was destroyed during World War II. Traces of fire use are present. Mousterian industry. Late Pleistocene, 55 thousand years ago.

Mandible from Šipka cave

Mandible fragment. ©Hitchcock.


Amud Cave. Israel. A cave near the Sea of Galilee (Lake Tiberias). Amud 1 is an almost complete skeleton of an adult male. Cranial volume is 1740 cm3, which is the largest value known for all hominids. Height: 174–178 cm, weight: 70 kg. Amud 7 is an articulated skeleton of a 10-month-old child. Amud 9 is the distal part of the right leg of a woman. Her height reached 160–166 cm and her weight—60 kg. Other scattered parts of several more children and infant skeletons were also found, totaling 19 individuals. 55 thousand years ago.

Amud-1 skull

Adult Neanderthal man from Amud Cave (specimen Amud-1).

Amud-7 mandible

Amud-7 is a Neanderthal child. The mandible is broad. This specimen (Amud-7) lacks a central keel, mental fossae, and a thickened inferior margin. Bar=1 cm. ©Schwartz et al.

Neanderthal fossils: Amud 7 burial

The Amud 7 infant burial in situ (modified from the original photograph made by E. Hovers).

Amud-9 leg
Remains of a Neanderthal woman Amud-9 fossils in situ. Found in Israel, near Lake Tiberias. ©Pearson et al.
Amud-9 metatarsal
Right first metatarsal of Amud-9: A) lateral; B) medial; C) superior; D) inferior; E) distal view. ©Pearson et al.

Aman Kutan

Aman Kutan. Uzbekistan. Western foothills of the Zarafshan Range, Lion’s Cave. Fragment of the femur of a young Neanderthal. Remnants of hearths. Late Mousterian. The End of the Late Tashkent Epoch, 55 thousand years ago.

Lion's Cave, Aman Kutan
Entrance to Lion’s Cave, Aman Kutan, where Neanderthal fossils were found.


Cova del Gegant. Spain. The site is located near the town of Sitges, on the promontory known as Punta de les Coves, Garraf Massif, Barcelona, Catalonia. So far, Neanderthal fossils have been found here, corresponding to a minimum of four individuals.

Gegant-1 is a large part of the edentulous mandible. Corresponds to an individual over 15 years old. Gegant-2 is a permanent lower lateral incisor that could correspond to a Neanderthal adolescent between 8 and 10 years old. Gegant-3 is the germ of a permanent central incisor, possibly from a younger individual than those mentioned above. This specimen remains unpublished. New findings: Gegant-4 corresponds to the distal portion of a left humerus, and Gegant-5 to a fragment of a right mandible with two teeth. Both remains could correspond to the same individual for about 4.5–6 years. Mousterian. The latest Neanderthal to date discovered in Catalonia. Upper Pleistocene. 55–52 thousand years ago (calibrated).

Gegant-1 mandible
Neanderthal mandible (Gegant-1). The fossil is 52,300 years old and represents the latest Neanderthal discovered so far in Catalonia. ©Daura et al.
Gegant-2 incisor
Neanderthal incisor (Gegant-2) from the “La Cova del Gegant” in buccal (a), lingual (b), distal (c), and mesial (d). Scale = 5 cm. ©Daura et al.
Gegant-4 humerus
Gegant-5 mandible

Neanderthal humerus (Gegant4) and jaw (Gegant-5). ©Quam et al.

Gegant G5 teeth

Gegant G5. The dm2 and M1 are fully erupted and in situ in their alveolar sockets. The canine and premolars are still unerupted and housed within the body of the mandible. ©Quam et al.

Prado Vargas

Prado Vargas Cave. Northern Spain. Burgos, Cantabria. Isolated deciduous lower left first molar of a Neanderthal. Child: 9–10 years old. The crown is complete, worn, but well-preserved, with resorbed roots. A small group of Neanderthals inhabited this cave seasonally. Late Mousterian, Levallois. 55–43 thousand years ago.

Deciduous molar from Prado Vargas Cave

Prado Vargas. Different views of PV-1360 tooth: A: mesial; B: occlusal; C: distal; D: buccal; E: radicular. ©Ruiz et al.


Stajnia (Stable Cave). Southern Poland. Northern Polish Carpathian Mountains. Three isolated molars from three different individuals: S5000 upper second heavily worn permanent molar of a man around 20 years old; S4300 lower first or second permanent molar (corresponding to 17–18 or 23–24 years old, respectively). The older of these individuals died at the age of 30, and the other was just over 20 years old. S4619 is the upper right unerupted molar of a 6-year-old child. Micoquian industry, Levallois. 54–52 thousand years ago, possibly more than 100 thousand years ago.

Molars from Stajnia

Stajnia Cave. Upper left: layer D2 (specimen 5000—Neanderthal upper second molar). ©Zarski et al. Upper right: A virtual 3D model of the S4619 tooth. ©Nowaczewska et al. Bottom: 3D digital model of the Stajnia S5000 molar. ©Stefano Benazzi.

Neanderthal molar S4619

S4619, Neanderthal right molar. a) Occlusal view. b) Lingual aspect of the protocone e Ca (Carabelli’s trait) is visible. c) Inferior view. d) Outlines depicting individual cusp base areas. e) The occlusal polygon described by the cusp apices of the protocone (A), paracone (B), metacone (C), and hypocone (D). f) Moire contourogram of S4619 M crown. Abbreviations: Pr: protocone; Pa: paracone; Me: metacone; Hy: hypocone; C5: metaconule; MAT: mesial accessory tubercle. ©Nowaczewska et al.


Cueva de El Esquilléu (Esquilleu cave or Big rock shelter). Northern Spain. Right side of the Deva River valley, Castro-Cillorigo, Cantabria. Neanderthal tooth. Remains of hearths with charcoal and charcoal from the burning of bones. Quina-type Mousterian, Levallois. 54–44 thousand years ago.

Neanderthal fossils: Cueva de El Esquilléu

Neanderthal remains from Cueva de El Esquilleu are on display at MUPAC, the Museum of Prehistory and Archeology of Cantabria in Santander, Cantabria, Spain. ©Thilo Parg.


Le Moustier. France. Village of Peyzac-le-Moustier, Dordogne. Two Neanderthal fossils of young individuals were found here. The complete skeleton (Mousterian-1), belongs to a Homo neanderthalensis adolescent aged about 15.5 years. Cranial capacity in the range 1500 см3–1565 см3. The skull of Mousterian-1 suffered significant damage after discovery. The nearly complete skeleton Mousterian-2 belongs to a 4-month-old infant. Both findings represent burials. The Mousterian tool culture is named after the Le Moustier site. 53–43 thousand years ago (calibrated).

Moustier I skull
Moustier I skull. The skeleton was discovered in 1908. Photo: Facsimile, Carl Bento, © Australian Museum.
Moustier I dentition
Le Moustier I dentition. Museum fur Vor- und Fruhgeschichte, Berlin, 2002. Photo: MiranC.
Moustier 2 skull

Virtual reconstructions of Le Moustier 2. Parts obtained via mirror-imaging are plotted in a darker shade; semitransparent surfaces were estimated based on complete reference crania. ©Gunz et al.

Neanderthal fossils: Moustier 2 skeleton

Le Moustier 2. Source: Original, le Musee National de Prehistoire, Les Eyzies-de-Tayac. ©Philippe Jugie.


Cueva de Covalejos. Spain. The site is located near Arce, in the municipality of Velo de Piélagos, Cantabria. Neanderthal fossils are three teeth. CV-1 is poorly preserved deciduous upper right first molar. Corresponds to an individual aged 6–10 years. CV-3 is the deciduous lower left first molar of a child aged 2–3 years. La Quina-type Mousterian, age around 53 thousand years. CV-4 is a poorly preserved tooth. It appears to be a deciduous upper central incisor. CV-2 was found at the Aurignacian level and belongs to Homo sapiens. There are traces of fish consumption. Bone tools. Mousterian. 45–40 thousand years ago.

Covalejos Neanderthal remains
Neanderthal remains from Cueva de Covalejos are on display at MUPAC, the Museum of Prehistory and Archeology of Cantabria in Santander. ©Thilo Parg.
Neanderthal teeth from Covalejos
Human teeth from Covalejos CV-4, CV-3, and CV-1. These three teeth belong to Neanderthals. Source: Montes Barquin R. and Sanguino Gonzalez.


Denisova Cave. Russia. Ayu-Tash, Altai. In addition to Denisovans, scientists found several Neanderthal fossils here. Altai Neanderthal, or Denisova 5, is the bone from the toe of an adult woman (the phalanx of the little finger of the left foot), as well as a small, indeterminate fragment of a man’s bone (Denisova 17). 52 thousand years ago.

Neanderthal phalanx

The toe phalanx of a Neanderthal found in the East Gallery of Denisova Cave in 2010. The high-quality genome sequence was generated from this small Neanderthal toe bone. ©Prüfer.

Bone fragment Denisova-17

Bone Denisova-17, found in 2011 in Denisova Cave. Samantha Brown et al./Nature Ecology & Evolution, 2021.


Cova de les Teixoneres. Spain. Village Moia, karst complex Les Coves del Toll, Barcelona. Scientists have discovered the archaic Neanderthal fossils of four humans. Among them were two children aged about 6–7 and 11 years old and one adult, as well as two fragments of the occipital bone of a Neanderthal youth and an upper child’s canine with moderate wear. This last child was 7–9 years old and was nicknamed “Moià child”. There are traces of cannibal tools on one clavicle. Mousterian, Levallois. Middle Paleolithic. 52 thousand years ago.

Teixoneres occipital bones

Two fragments of the occipital bone of a Neanderthal skull. Maria D. Guillen / IPHES

Teixoneres clavicle
A fragment of a previously found clavicle with traces of tools. Leandro Zilio/CONICET; Ruth Blasco/IPHES
Teixoneres tooth and parietal fragment
Neanderthal child tooth and parietal fragment discovered at Teixoneres Cave in Spain. ©IPHES
Teeth from Teixoneres Cave

Some human remains were found on level IIIb of the Teixoneres Cave. Maria D. Guillen/IPHES-BUSQUEDA


La Ferrassie, rock shelter. France. Near Le Bugue and Les Eyzies, Savignac-de-Miremont, Dordogne department. Neanderthal fossils of seven individuals.

La Ferrassie 1 is an adult male 45 years old with a nearly complete skull, a brain volume of 1641 cm3, a height of 172 cm, and a weight of about 85 kg. This specimen is considered a classic example of Neanderthal anatomy. The age of the fossil is 52–40 thousand years. La Ferrassie 2—an incomplete skull and skeleton of a female Neanderthal 25–30 years old, weight 67 kg, dated 50–44 thousand years ago. The next find is La Ferrassie 3—an incomplete skeleton of a 10-year-old child. La Ferrassie 4—partial skeleton of a newborn baby (12 days old). La Ferrassie 5—the seven-month-old fetus of a child. The fossils of La Ferrassie 6 are an almost complete skeleton of a child 3–5 years old.  And finally, La Ferrassie 8 is a 2.5-year-old child with well-preserved teeth. The age of the find is 45–40 thousand years. Burial rituals. Mousterian. The rest of the specimens are about 52–45 thousand years old.

Ferrassie I Neanderthal
The adult is based on La Ferrassie I. ©Elisabeth Daynes. Photo: Don Hitchcock.
“Old man” of Ferrassie
Skull of Ferrrassie I, the “old man” of Ferrassie. ©Jc Domenech.
Ferrrassie I skeleton

Ferrrassie I. Collections du Museum national d’Histoire naturelle, Paris. ©Hitchcock.

Neanderthal fossils: Ferrrassie skeleton in situ

View of the skeleton at the time it was found. It was lying on the Acheulean beds, surrounded and covered by Mousterian deposits. Photo: Capitan and Peyrony (1910). Photographer: Almost certainly Monsieur Lucas, l’institution aux Eyzies.

Ferrassie 2 foot
Ferrassie 2 foot replica

The foot of the skeleton of La Ferrassie 2 Neanderthal is still conserved in its block of sediment. Left: ©Pablos; right: ©John Reader/Science Photo Library.


Rochers-de-Villeneuve. Central-western France. Lussac-les-Chateaux, Vienne. Neanderthal partial diaphysis of the left femur (RdV 1) with traces of carnivore teeth. Belongs to an adolescent or, more likely, an adult. Morphological features of the femur and ancient DNA extracted from it indicate that it belongs to Neanderthals and differs from early modern humans. Classic Neanderthal. Traces of hearths. Mousterian, Levallois. The End of the Middle Paleolithic, 51–45 thousand years ago (calibrated).

Femur from Rochers-de-Villeneuve

RdV 1 left femur fragment. Shown are the anterior (A), posterior (B), medial (C), and lateral (D) areas. (E) Details of the carnivore toothmarks on the distal end of the RdV 1 human femur diaphysis. ©Beauval et al.


Pech-de-l’Azé caves. Southwest France. Commune of Carsac-Aillac, Dordogne department, Aquitaine. The remains of Pech I include a nearly complete cranium and mandible of a young Neanderthal child that has been aged between 2.5 and 3 years. The mandible of Pech I is remarkably complete, although only on the right side of the jaw are the teeth well-preserved. All the deciduous teeth are fully erupted. There are numerous hearths. Mousterian of Acheulean tradition. Middle Paleolithic, 51–44 thousand years ago.

Pech I  skull
Child skull and from Pech-de-l'Azé

Child skull and mandible from Pech-de-l’Aze, Dordogne. ©Hitchcock.


Grotte du Bison, Bison Cave. France. Commune of Arcy-sur-Cure, Yonne, Burgundy. Located near Grotte du Renne. Two teeth—one deciduous tooth (T14) and one permanent tooth (J2. U14-2931)—were discovered during early excavations. Later, ten more fossil human specimens were found, including a fragment of the maxilla of an adult individual (P11.8) with 6 teeth and 9 isolated teeth (temporary and permanent). The latest findings: the second lower right deciduous molar of a 3-year-old child (BIS-I S6-2); the second lower right deciduous molar of an 8–10-year-old child (BIS I Q5.1). The roots of this tooth are in the initial stage of resorption. Remains of hearths, grease lamps, traces of ocher. Late Mousterian. 51–44 thousand years and 48–42 thousand years.

Teeth from the Grotte du Bison
Isolated teeth from the Grotte du Bison. 3: Upper central deciduous incisor (tooth I.Q8.121). 4: Teeth found in 1963: a and b: tooth T14; c, d, e: tooth U.14.2931 (© MNP Les Eyzies-Dist. RMN-photo Ph. Jugie). 5. Upper permanent incisor and (upper and lower) first molars. a: tooth S15-1; b: teeth P8-31 and I.P8. 6. Second upper permanent molars: a: P8-101 and b: P7-48. 7. Upper permanent third molar (P8-63): a, b. 8. Lower permanent third molar (P7-78): a and b. ©Tillier et al.
Maxilla P11.8 from Grotte du Bison
Maxilla fragment P11.8 from Level I of the Grotte du Bison, Arcy-sur-Cure. Incomplete right adult maxilla (P11-8) from the Grotte du Bison. a: medial view, showing a steeply sloping anterior nasal floor (Photo J. Enloe); b: occlusal view, illustrating the tooth morphology and dental wear (Photo C. Tolmie); c: lateral view, showing the crenulated tooth marks on the upper margin of the maxillary bone suggest carnivore modification of this specimen (Photo C. Tolmie). Scale in cm.
Neanderthal fossils from Grotte du Bison
Neanderthal fossils from Grotte du Bison

“What also characterizes Neanderthal teeth is the size of the crowns, the roots, and the complexity of the patterns on the surface of the enamel. It is also a much finer enamel than ours”. Photos by Juliette Henrion.

Bronzovaya (Bronze)

Bronzovaya Cave (Bronze Cave). Western Georgia. Near Tsutskhvati or Cuckhvati, left bank of the Shabatghele River. Upper left first permanent molar of a child aged 8–13 years. According to more recent data, it is a deciduous tooth of a 3-year-old child, exceptionally well preserved, and only slightly worn. The tooth exhibits taurodontism. Remnants of hearths. Mousterian, Levallois. 50 thousand years ago.

Molar from Bronze Cave

Upper first left molar from the Bronze Cave in the village of Tsutskhvati (Cuckhvati). A: occlusal view. ©F. Rivals and T. Chevalier. B: lateral view. ©Gabunia et al.


Kulna Cave. Czech Republic. The municipal territory of Sloup, Blansko District, north of Brno, South Moravian Region. Neanderthal fossils include: the right part of a maxilla with four teeth (canine, both premolars, and the first molar) of an adolescent boy (14–15 years old); the right parietal bone of a man; other fragments; and 3 teeth. There are tools made of crystalline quartz. Mousterian, Micoquian. 50 thousand years ago.

Maxilla from Kulna

A fragment of the upper jaw of a Neanderthal boy was found by archaeologists in 1965 in the Kulna cave. ©Skokan.

Neanderthal fossils from Kulna Cave

Kulna Cave (near Sloup, Blansko District): 1–maxila fragment; 2–right parietal bone; 3–milk teeth. ©Zde.

Noisetier (Peyrere)

Grotte du Noisetier, also known as Grotte de Peyrère or Grotte de Serrat de la Toue (Noisetier Cave, Hazelnut Tree Cave). Southern France. In the commune of Ardengost, town of Fréchet-Aure, Hautes-Pyrénées department in the Occitanie region. Three Neanderthal teeth were found, including the upper central deciduous left incisor (Ns 06 D13 c1). They exhibit significant wear of the crown and incomplete roots, as in a child 5–10 years old. They probably belong to the same individual. Mousterian-Levallois. 50 thousand years ago.

Neanderthal incisor from Noisetier

Neanderthal deciduous incisor Ns 06 D13 c1 – 185. Photo: B.M.


Vanguard Cave. Gibraltar. The third Neanderthal of Gibraltar. Upper right deciduous canine of a child aged 4–5 years. The find corresponds to the time when the cave was a hyena den. A layer of ash and mussel shells likely indicates that Neanderthals prepared seafood using fire to open the shells. 50 thousand years ago.

Vanguard (Gibraltar-3)

Vanguard Cave. Upper right canine of a Neanderthal 4–5 years old (Gibraltar-3). ©The Gibraltar Museum.


Grotta del Cavallo, also known as Grotta delle Giumente (Mares) or Uluzzo A, “Cave of the Horse”. Southern Italy. Uluzzo Bay, commune of Nardò, province of Lecce, Apulia. Two Neanderthal teeth. Cavallo A—second deciduous left lower molar; Cavallo D—right lower incisor of a young individual. Classic Neanderthal. Mousterian, Uluzzian (the Italian equivalent of the French Châtelperronian). 50–47 thousand years ago.

Teeth from Grotta del Cavallo

A. left dM2 referred to layer L of Cavallo (the A-tooth); B. left dM1 referred to level E-III of Cavallo (the B-tooth); C. dM2 referred to level E-II/I of Cavallo (the C-tooth). ©Benazzi. D. Neanderthal right L1 of Cavallo (the D-tooth). D—not to scale. ©Moroni et al. The white rectangles mark the tuberculum molare of the B-tooth. B=buccal, D=distal, L=lingual, and M=mesial.

Neanderthal tooth from Cavallo

Tooth from Grotta del Cavallo. ©Fabbri.


Riparo Broion. Northern Italy. Berici Mounts, Venetian Pre-Alps. Broion 1 is a shed upper right canine, heavily worn. Approximately a quarter of the root is preserved, suggesting an age of around 11–12 years. The tooth is characterized by a stocky crown, bulging buccally, and a lingual cervical eminence, producing an asymmetrical outline. Overall data concur to align Riparo Broion 1 with Neanderthals. Mousterian. 50–46 thousand years ago.

Riparo Broion deciduous tooth

An upper canine milk tooth that belonged to a Neanderthal child, aged 11 or 12, that lived between 48,000 and 45,000 years ago. B = buccal; L = lingual. Credit: Journal of Human Evolution

Neanderthal tooth from Broion Cave

This is a 3D reconstruction of the Neanderthal milk tooth found in the Broion Cave. ©University of Bologna. Riparo Broion 1 (RB1), right dC1, in different views. Right: the enamel-dentin junction of the tooth shows the mesiodistal convexity (a), the lingual crests and fossae (b), and an enamel fracture on the distal side of the crown (c). Abbreviations: O = occlusal; D = distal; M = mesial. Scale bar = 1 cm. ©Romandini


La Crouzade. France. Municipality of Gruissan, department of Aude. Crouzade V (frontal bone) and Crouzade VI (maxilla) belong to Homo sapiens. Six Neanderthal fossils were found in three different Mousterian levels. Clavicle CX; two humeri CIX (infant) and CI (adolescent); three hand phalanges CII and CIII from one three-month-old infant, as well as CIV from an adolescent. Classic Neanderthals. Mousterian. 50–45 thousand years ago.

Neanderthal fossils from Crouzade

Neanderthal remains from La Crouzade cave. ©Saos et al.

Fonds-de-Foret (Bay Bonnet)

Fonds-de-Forêt, also known as Première Caverne du Bay Bonnet caves. Belgium. Left bank of the Magne stream, Municipality of Trooz, province of Liège. There are two Neanderthal fossils: a left femur from an adult individual and a left upper molar. Classic Neanderthals, similar to those from Spy. Charentian of the Quina type. 50–44 thousand years ago. 

Femur from Fonds-de-Forêt

The Neanderthal left femur from Fonds-de-Foret: (a) anterior; (b) medial; (c) distal; and (d) dorsal views. Image by G. Abrams, modified after pictures from E. De Wamme (Copyright RBINS) and Camaros et al.


Grotta del Capelvenere. Southeast Italy. Near the church of S. Caterina di Nardò, not far from Torre dell’Alto, Lecce, Apulia. The first upper right molar of an adult individual. Mousterian of La Quina type. 50–40 thousand years ago.

Grotta del Capelvenere

Finding site of a Neanderthal molar in Grotta del Capelvenere.

Ortvale Klde

Ortvale Klde rock shelter. Georgia. The site is located near Chiatura (Tchiatoura), not far from the villages of Abas-Zodi and Haut-Rgani, in the gorge of the Cerula River. Second lower permanent molar. The degree of wear and the type of tooth suggest that it belongs to an adult individual. The roots are not fused. Levallois. Middle Paleolithic, 50–40 thousand years ago.

Ortvale Klde molar

Ortvala Klde left M2 occlusal. Scale = 1 cm. Photos by F. Rivals and T. Chevalier.

Devil’s Tower

Devil’s Tower Cave. Gibraltar (British Overseas Territory). The site is situated in a cleft in the North Face, not far east of Forbes’ Quarry. Devil’s Tower Child, or Gibraltar 2—consists of five fragments of the skull (maxillary, parietal, temporal, cranial, and mandibular) of a 4-year-old boy. The child had large front teeth and a long, wide face. There were traces of hearths and parts of burned bones found at the site. Mousterian. 50–42 thousand years ago.

Devil's Tower child skull

The Devil’s Tower child skull. ©Guerin Nicolas.

Devil's Tower (Gibraltar 2) Neanderthal fossils

The Devil’s Tower (Gibraltar 2) Neanderthal child fossil in pieces. ©Buck/NHM.

Devil’s Tower mandible

Gibraltar 2 (Devil’s Tower), a Neanderthal juvenile mandible, and 2 isolated molar germs. Left: occlusal view. ©Tanya M. Smith, Paul Tafforeau. Right: top (anterior view), middle (right lateral view). Bar=1 cm. ©Schwartz et al.

Trou Al’Wesse (Modave)

Trou Al’Wesse. Belgium. Municipality of Modave, Petit-Modave, right bank of the Hoyoux River, province of Liège. From sedimentary deposits where the bones of Neanderthals themselves were not preserved, geneticists extracted Neanderthal mitochondrial DNA. Mousterian. 50–41 thousand years ago (the lower date is an uncalibrated value).

Trou Al’Wesse

View of the archaeological site of Trou Al’Wesse, Belgium. (Credit/Quelle: Monika V. Knul)

Abri Moula

Abri Moula. Southeastern France. Slope of Serre de Guercy, Soyons, Ardeche department. 13 Neanderthal fossils were discovered (seven cranial fragments, two teeth, and three postcranial fragments). There are signs of cannibalism (cutmarks on the bones from stone tools). Typical Mousterian with elements of Levallois. 49 thousand years ago.

Abri Moula radius
Cutmarks on the bicipital tuberosity of the radius. ©Defleur et al.
Abri Moula parietal
Cutmarks on the parietal bone from Abri Moula. ©Defleur et al.


Cova Simanya (Sant Llorenç Savall). Spain. The site is located in the Sant Llorenç del Munt i de l’Obac Natural Park, Barcelona. 54 remains of skeletons: upper and lower dentition, a fragment of the mandible, vertebrae, the left and right upper limbs (humeri and hand bones), and foot bones. The most complete individual included a whole undamaged humerus (SI-1), another humerus, two wrist bones, and a vertebra, all attributed to the same adult individual aged 30–40 years. The estimated height was 154 cm, indicating a female. Individuals aged 13 years (SI-60), 11.5 years (SI-7), and 8 years (SI-29) were also found, as well as an isolated third molar. Late Homo neanderthalensis. Traces of hearths. Mousterian. 49–42 thousand years ago.

Neanderthal fossils from Simanya

Foremost Neanderthal remains found at the Simanya Gran gallery in Cova Simanya (Barcelona, Spain). (A) Left humerus; (B) shaft, caudal view; (C) bones of the foot; (D) bones of the right hand; (E) upper P3 and M3; (F) fragment of the atlas and first phalanx of the hand; and (G) fragment of the ascending branch. Author: MNCN-CSIC / IPHES-CERCA / Frontiers in Earth Science.

Neanderthal tooth from Cova Simanya
Neanderthal tooth from Cova Simanya

Detail of the Neanderthal tooth recovered in Cova Simanya. Author: Juan I. Morales/IPHES-CERCA

St Brelade

St. Brelade (La Cotte de St. Brelade). Jersey, Channel Islands. 13 Neanderthal teeth from two individuals. Later, a number of bone fragments were found. The teeth show evidence of taurodontism. Two teeth suggest that they originated from both Neanderthal and modern human ancestry. There are also traces of hearths present. Levallois. 48 thousand years ago.

Neanderthal fossils: St. Brelade tooth

Two of the teeth recovered from La Cotte de St. Brelade (Jersey). It’s thought they could be evidence of a hybrid population of Neanderthals and Homo sapiens. ©Credit Societe Jersiaise Photographic Archive.


Monasheskaya (Monk’s Cave). Russia. Left bank of the Gubs River, Krasnodar Krai, North Caucasus. Two phalanges, fragments of vertebrae, ribs, two incisors, and eleven fragments of teeth were found at Monasheskaya. These remains show a number of archaic features. Monasheskaya Cave was inhabited later than the neighboring Barakaevskaya Cave, around 48 thousand years ago.

Monasheckaya Cave

View of the left, northern side of the Gubs River canyon (arrow indicates location of Monasheckaya Cave).


Vindija Cave. Northern Croatia. Municipality of Donja Voća, near the town of Varaždin. The oldest Neanderthal fossils from the cave consist of parts of the cranial vault, brow ridges, upper and lower jaws, isolated teeth, and limb bones from both young and adult individuals. There are at least 12 individuals. They represent a transitional stage from the classic, robust form to a more gracile one. Mousterian, with elements of the Aurignacian industry. 48–45 thousand years ago (calibrated).

Vindija supraorbital fragments

Frontal view of Vindija supraorbital fragments, shown on the same approximate scale. A: Vi 262 and B: Vi 224; C: Vi 202 and D: Vi 279; and E: Vi 260 and F: Vi 264. Vi 224 and 279 may not be fully adult. ©Wolpoff et al.

Neanderthal bone Vi-*28
High-resolution photographs of the Vi-*28 Neanderthal bone found at Vindija Cave, Croatia. The bone yields evidence for a probable cut and gauge marks (right upper part of the bone). Credit: Thomas Higham.
Long bones from Vindija Cave
Three fragments of long bones represent three different female Neanderthals who occupied the Vindija Cave. DNA from these bones was analyzed for genome sequencing. Max Planck Institute for Evolutionary Anthropology.
Vindija maxillae
Maxillae Vi 225 (left) and Vi 259 in inferior, frontal, and lateral views. Specimens are shown on the same approximate scale. Note the relative size of the incisor roots, the morphology of the lower nasal margin, and the low position of the maxillary sinus. ©Wolpoff et al.
Vindija mandibles
Lateral views of the three Vindija mandibles. Vi 231 (A), Vi 250 (B), and Vi 206 (C). Right: Occlusal views of Vi 231 (D), Vi 206 (E), and Vi 226 (F). The specimens are shown on the same approximate scale. Note the noticeable variation in toothrow shape. ©Wolpoff et al.
Vindija anterior teeth

The isolated Vindija anterior teeth are shown on the same approximate scale. There are Vi 289 (I?), Vi 287 (maxillary canine), Vi 290 (I1), all in lingual view; Vi 288, a mandibular canine in lingual view; next are lingual and labial views of the Vi 201 I, (note the chipping surrounding the occlusal surface on both sides); and Vi 286 (L). We believe that each tooth represents a different individual. ©Wolpoff et al.


El Salt. Spain. Near the city of Alcoy, in the province of Alicante. It is an open-air rock shelter. 6 upper jaw teeth belonging to a young individual: two molars (Salt 4 and 5) and remnants of a third tooth (Salt 6), as well as two premolars (Salt 2 and 3) and a fragment of an incisor (Salt 1). Classic Neanderthals. Hearth remains. Levallois. 47–45 thousand years ago.

Salt 2

Salt 2: Right P3. A: buccal; B: lingual (the arrow in A and C indicates the hypoplasia line); C: mesial; D: distal; E: occlusal. White line = 10 mm. ©Garralda et al.

Salt 3

Salt 3: Right P4. A: lingual; B: buccal; C: mesial; D: distal; E: X-ray of the distal side; F: occlusal. White line = 10 mm. ©Garralda et al.

Salt 4

Salt 4: Right M1. A: mesial; B: distal; C: lingual; D: buccal; F: occlusal. White line = 10 mm. ©Garralda et al.


La Baume Néron (Neron Cave). France. Soyons commune, Ardèche department. Slightly worn permanent upper first molar, deciduous right canine of the mandible. Taurodontism. Adolescent. Mousterian of Quina type, Neronian (large, thick flakes). 47–45 thousand years ago (calibrated).

Molar Baume Neuron

Upper left permanent molar Baume Neuron (top): 1–Distal view; 2–Mesial view; 3–Occlusal view; 4–X-ray. Right decidual canine of Baume Neuron (below): 1–Vestibular view; 2–Lingual view; 3–Occlusal view; 4–X-ray. ©Defleur et al.

Trou de l’Abime (Couvin)

Trou de l’Abîme (Abyss Hole). Also known as La caverne de l’Abîme and Couvin Cave. Belgium. Near Couvin, Namur province, Wallonia. The crown of the deciduous right second molar of the mandible. The child was 7–11 years old. A distinctive stone industry corresponding to the transition between the Middle and Upper Paleolithic techno-complexes “Couvin facies”. 47–44.5 thousand years ago.

Fragment of the mandible. Trou de l’Abime

Neanderthal deciduous tooth. The tooth is still partially embedded in the breccia. Photo: Joel Eloy, AWEM.


Zafarraya. Cueva del Boquete de Zafarraya Cave. Spain. Alcaucín, Granada province, Andalusia. 16 bones were found: femur, two additional femur fragments, toe phalanx, pubic bone, fragment of the tibia, scapula, several teeth, and three jaws, including a well-preserved mandible of a woman aged 20 to 30 years (Zafarraya 2). In this jaw, 13 teeth out of 16 are preserved. Such good preservation is exceptional. Classic Neanderthals. Hearth remains. There are traces of cannibalism. Mousterian. 46 thousand years ago.

Neanderthal fossils from Zafarraya

Mandible and femur of a Neanderthal from Boquete de Zafarraya.


Anghilak Cave. Southeastern Uzbekistan. Qashqadaryo (Kashkadariya) region, on the foothills of the Zerafshan mountain range, 45 km south-west of Samarkand. The fifth right metatarsal bone (AH-1) of an adult individual. Classic Neanderthal. The cave inhabitants consumed turtles as food. Mousterian. Middle Paleolithic, 46 thousand years ago (calibrated).

Metatarsal bone from Anghilak Cave

AH-1: metatarsal bone from Anghilak Cave. ©Glantz et al.


Les Cottés. Southwest France. Located on the southwest edge of the Paris Basin. Upper molar tooth of an adult Neanderthal woman. Analysis of tooth tissues indicates that its diet consisted mainly of the meat of large herbivore mammals, including reindeer and horses. Mousterian of La Quina type. There is a Châtelperronian industry dated to 39,000 years, but whether it is related to Neanderthals is unknown. 46–44 thousand years ago (calibrated).

Les Cottés molar

The upper molar of a late-adult Neanderthal woman discovered during excavations at Les Cottes Cave in France. Left: MPI f. Evolutionary Anthropology/A. Le Cabec. Right: M. Soressi/S. Schstz.


Goyet Caves. Belgium. Troisième caverne (“Third cave”), Namur Province, Wallonia. Neanderthal fossils from five individuals: 4 adolescents or adults and one child. In total, there were about 96 bone samples and 3 isolated teeth. Individual Goyet Q56-1 (right femur) was female and is genetically closest to the Spy Neanderthals. There are tool marks on the bones, probably indicating signs of cannibalism. It is evident that the Neanderthals from the Troisième cave in Goyet were killed and used as a food source. 46–44 thousand years ago (calibrated).

Neanderthal jaw from Goyet

Neanderthal jaw from Goyet. ©AFP News Agency.

Neanderthal fossils from Goyet

Neanderthal remains from the Troisieme caverne of Goyet (Belgium). The remains have scrape marks, indicating that they were butchered, with cannibalism being the “most parsimonious explanation”. Scale=3 cm. ©Rougier.

Goyet. Neanderthal fossils

Goyet. Human remains different specimens. ©IRSNB – Eric Dewamme.


Arrillor (La Cueva Arrilor). Spain. Municipality of Cigoitia or Zigoitia (Álava), near the town of Murua, Gorbea Mountain (Araba, Basque Country). Neanderthal tooth—a shed upper second right molar of an individual aged 9 to 13 years old. Mousterian, Levallois. Middle Paleolithic. 46–43 thousand years ago.

Deciduous molar from Arrillor
Neanderthal tooth from La Cueva Arrilor

Neanderthal tooth from La Cueva Arrilor. ©Arabako Arkeologia Museoa

Roccia San Sebastiano

Roccia San Sebastiano. Southern Italy. Mondragone-Caserta. Lower left second deciduous molar (RSS1). The tooth is worn and has a complete crown and a quarter of the root. The child was 9–12 years old. The second tooth found belongs to Homo sapiens. Late Mousterian. 45–44 thousand years ago.

Child tooth from Roccia San Sebastiano

Three-dimensional digital models of lower left second deciduous molar RSS 1. On the right side, there is the enamel-dentin junction (EDJ) of the teeth. AntFov–anterior fovea; MTC–middle trigonid crest; MET–Metaconid; ENT–Entoconid; HYP–Hypoconid; HYPL–Hypoconulid; PRO–Protoconid; B–buccal; D–distal; L–lingual; M–mesial; O–occlusal. ©Oxilia et al.

Ksar Akil

Ksar Akil (Ksar Aqil) rock shelter. Lebanon. Located 10 km north of Beirut. It is a large stone shelter beneath a steep limestone cliff. Ksar Akil 2: “Ethelruda”, teeth and a fragment of the right part of the maxilla of a young adult woman. It exhibits Neanderthal features as well as some characteristics of archaic Homo sapiens. Aside from Ksar Akil 2, the Early Ahmarian level of XVII yielded the fossils of two children. One of them was represented by a maxilla and a few ribs, but was probably never excavated. The beginning of the Late Paleolithic, Mousterian-Levallois industry. 45.9 thousand years ago, possibly older.

Fragment of the maxilla Ksar Akil

View of the partial right maxilla of Ethelruda (Ksar Akil 2). ©Douka et al.

Castaigne (Caminero)

Castaigne Cave (also known as Caminero). France. Located in the valley of the Eaux-Claires stream, municipality of Torsac, near Angoulême, Charente. Neanderthal fossils (41 elements), including the remains of at least six (possibly eight) individuals. These include isolated teeth, cranial fragments (parts of the skull cap, parietal, temporal, and occipital bones), and postcranial elements (femur, humerus, clavicle, phalanges, etc.), belonging to individuals of different ages. Classic Neanderthal. Mousterian and Levallois industries. 45–42 thousand years ago.

Phalanx and femoral diaphysis from Castaigne

Castaigne. Left: Photos of the palmar, medial or lateral, and dorsal views of a proximal phalanx, probably the 3rd or 4th finger of the hand. Scale = circa 30 mm. ©Philippe Jugie (MNP). Right: photo of the posteromedial view of the left femoral diaphysis (proximal extremity at the top). Scale: 10 cm. Modified from Duport (1968, unpublished).

Castaigne isolated teeth

Castaigne. Photos of the isolated teeth that identified the six different individuals within the paleoanthropological collection. Scale = 5 cm. 1 = lower left lateral deciduous incisor germ, distal view; 2 = lower right deciduous canine, mesial view; 4 = lower left permanent I1, distal view; 6 = lower left P3, distal view; 7 = lower right permanent canine, distal view; 8 = M1 or M2 permanent upper left, distal view. Photos: Bruno Maureille or Maryelle Bessou (Universite de Bordeaux, UMR PACEA).

Toothpick grooves on the molar
Toothpick grooves on the mesial (left) and distal (right) views on the lower left M1 or M2 of a young adult. Photos by Maryelle Bessou (Universite de Bordeaux, UMR PACEA).
Toothpick grooves on the incisor
Castaigne Cave (Caminero). Toothpick grooves on the distal views on the lower right lateral incisor. Photos Maryelle Bessou (Universite de Bordeaux, UMR PACEA).


La cueva de los Casares. Spain. Located in the village of Riba de Saelices, Guadalajara province. A metacarpal bone belonging to a classic Neanderthal was found. Anthropogenic charcoal was also discovered. Mousterian industry. Middle Paleolithic, 45–42 thousand years ago.

Neandertal metacarpal from Casares

Neandertal metacarpal found in Seno A (bar is 5 mm). ©Alcaraz-Castano, modified after Basabe.


Feldhofer Cave (Kleine Feldhofer Grotte), Fedhof Cave. Germany. Neander Valley, Düsseldorf. Initially, 15 bones were found: a skull cap, two femurs, three bones of the right arm, two bones of the left arm, the ilium, as well as fragments of the scapula and ribs. Then, 73 more bones were discovered, belonging to Neanderthal 1 and two other individuals (a child and, presumably, a woman—the right humerus, right ulna, and other fragments). For Fedhofer 1, the found parts of the left jugal and maxilla (NN 34) are particularly informative. Its entire anatomy is clearly Neanderthal, emphasizing once again the “classic” Neanderthal type of the individual. Fedhofer 1 is a type specimen for the species Homo neanderthalensis. The weight of the individual is 79 kg. Micoquian. Late Middle Paleolithic, 45–42 thousand years ago (calibrated).

The cranium of Neanderthal 1
The cranium of Neanderthal 1, the first fossil recognized as Neanderthal. This find became the type specimen for the Neanderthal genus.
The zygomatic-maxilla and Feldhofer 1 cranium
The left zygomatic-maxilla (NN 34) articulated with the Feldhofer 1 calotte. Illustration by M. Cartmill.
Feldhofer skeleton

Partial Neanderthal skeleton from the Kleine Feldhofer Grotte, Germany.


Grotta Breuil. Italy. This is one of the caves in Monte Circeo, Lazio province. Several Neanderthal fossils were found in the cave. The fragment of the skull Breuil 1 is a uniformly curved flat bone corresponding to the mastoid process of the left parietal bone of an adult. Breuil 2 is a permanent tooth of the mandible, probably the first left lower molar. The crown is heavily worn, and the root is absent. The specimen belonged to an individual no younger than 30 years old. Breuil 3 is a permanent third molar of the left mandible, which has not yet developed two-thirds of the root. The crown is already formed, while the root is still developing. This tooth belonged to an adolescent 13–14 years old. Classic Neanderthals. Traces of fire use. Mousterian, Pontinian variant. 45–40 thousand years ago.

Breuil 2
Breuil 2, left first mandibular molar. Top, vestibular (left) and lingual (right) views; center, mesial (left) and distal (right) views; bottom, occlusal view (left) and X-ray (right). ©Manzi, Passarello.
Breuil 3
Breuil 3 is a permanent third molar on the left half of the mandible. It has not yet developed two-thirds of its root. It is the tooth of a 13–14-year-old Neanderthal adolescent. ©Manzi, Passarello.
Breuil 1

Internal (left) and external (right) views of Breuil I parietal fragment (drawn by M. Mei).


Riparo Mezzena (Mezzena Rockshelter). Northern Italy. Near Verona, Avesa Valley. The occipital bone and mandible of a Neanderthal, a total of 14 bone fragments, three of which belong to the postcranial skeleton. Obviously, there are two adult individuals. Analysis of nuclear DNA indicates light skin color and belonging to Neanderthals, despite some assumptions about hybridization. Mousterian with small elements of Charentian, Levallois. 45–40 thousand years ago.

Mezzena. Neanderthal mandible

Mandible from Riparo Mezzena; upper; bottom; lateral; frontal views. ©Condemi et al.

Ortvala Cave

Ortvala Cave. Georgia. Near the village of Didi Rgani, Imereti. Two teeth from adult individuals were found in Ortvala Cave, near Sakajia Cave. One of them is the upper third left molar, and the second tooth may be the second upper incisor. The teeth are too worn to describe the occlusal surface. Mousterian, Levallois. 44.5–42.5 thousand years ago.

Ortvala Cave teeth

A—Ortvala Cave left M3 occlusal; B—Ortvala Cave I2 (?) occlusal. Scale = 1 cm. Photos by F. Rivals and T. Chevalier.


Engis Cave (Schmerling Caves). Belgium. Village of Les Awirs, municipality of Flémalle, province of Liege, Wallonia. Two cranial vaults were found. Engis 1—an adult individual from the Neolithic period (Homo sapiens). The specimen Engis 2 consists of a partially preserved cranial vault with associated fragments of the maxilla and jugale, as well as the upper and lower teeth. The individual was a child aged 4–6 years. Typical Mousterian of the Levallois facies. 44,000 years ago.

Neanderthal Child Engis 2

The Neanderthal Child Engis 2, found by Schmerling in the second cave during the winter of 1829–1830. (A) Skullcap and (B) second lower right deciduous molar (LRdm2). Image by G. Abrams after pictures from G. Focant (A, Copyright SPW) and A. Le Cabec (B).


Okladnikov Cave (Sibiryachikha Cave). Russia. On the bank of the Sibiryachikha River, Altai, Southern Siberia. 168 bones and fragments, including teeth, fragments of the humerus and femur, and the phalanx. Among them, 17 fragmentary remains from the Okladnikov Cave can be attributed to two children, one adolescent, and one adult. Neanderthal Okladnikov 2 turned out to be closer to European Neanderthals than Asian ones. Mousterian. Several findings date to 44,000 years ago.

Neanderthal fossils from Okladnikov cave

Neanderthal tooth and bones from the Okladnikov cave. Scale 1 cm. ©Bence Viola.


Grotte de Spy (Spy Cave). Belgium. Located on the bank of the Orneau River, in the municipality of Jemeppe-sur-Sambre, Grotte de la Betche-al-Rotche, Namur province, Wallonia. A very incomplete skeleton of a woman, Spy I, with a cranial volume of 1300 cm3; and a young man, Spy II, weighing 83.6 kg, with a cranial volume of about 1500 cm3. The remains of both individuals consist of cranial vaults, upper and lower jaw elements, as well as a series of often incomplete postcranial remains. Spy III is a small child; Spy VI (two mandibular corpus fragments) is an 18-month-old child; and there are several other adults. Spy94a, the upper right molar, probably belongs to Spy I. Charentian subtype of the Mousterian industry, of the Quina type. 44–42 thousand years ago.

Neanderthal fossils from Spy Cave

There are four Neanderthal specimens from Spy Cave. (A) the right scapula (Spy 572a). (B, a) the maxillary fragment and upper third molar (Spy 94a) associated with a maxillary fragment (B, b) (Spy 11a) and a mandibular fragment (B, c) (Spy 12a). (C) a first upper right deciduous incisor (Spy 589a). (D) the lumbar vertebra (Spy 737a). Image by G. Abrams after pictures from M. Toussaint (A, D) and E. De Wamme (B, C; copyright RBINS).

Spy 1
Spy 2

Skulls of Spy 1 (left) and Spy 2 (right), right lateral view, Belgium. ©Institut Royal des Sciences Naturelles de Belgique.

Mandible of Spy 1
Neanderthal mandible: left (anterior view) and right (left lateral view) of Spy 1. Institute Royal des Sciences Naturelle de Belgique, Brussels. Bar=1 cm. ©Schwartz et al.
Jaws of the Spy 2
The reconstructed jaws of the Spy II Neanderthal, displaying where the dental calculus was removed from the impeccably white back molars. ©Royal Belgian Institute of Natural Sciences.
Neanderthal fossils: Spy 2 skeleton

The Spy 2 skeleton (a male) was discovered during the first excavations in 1886. ©Royal Belgian Institute of Nature Science.

Spy IV
A lower jaw and teeth of a Neanderthal infant (Spy IV) were unearthed in a Belgian cave. ©Crevecoeur.
Molar Spy 94a
Upper right molar of a male Neanderthal, Spy 94a, from Spy Cave, Belgium. ©Isabelle Crevecoeur.
Mandibular fragments of the Spy VI

Preserved mandibular fragments of the Spy VI child. Spy 646a: right mandibular corpus fragment. Spy 194a: left mandibular corpus fragment. ©Isabelle Crevecoeur.


Walou Cave. Belgium. Located on the left bank of the Magne River, in the municipality of Trooz, Liège province. The lower left first premolar of an adult individual, found in a Mousterian context in the cave. The tooth has one root and a robust crown. Mousterian. 44–42 thousand years ago.

Premolar from Walou Cave

Walou Cave: premolar from layer C sup.


Grotte du Renne (Reindeer’s Cave). Northeastern France. Arcy-sur-Cure, Yonne department, Burgundy. The site includes the Schoepflin gallery. Neanderthal fossils: 76 bone elements, including teeth (29 teeth from 6 individuals, including 14 deciduous teeth) and a fragment of a child’s temporal bone. All teeth have Neanderthal characteristics. Also, scientists found two tooth buds in the Châtelperronian layer belonging to Neanderthal children. Both findings are tooth buds of left premolars. The first belonged to an infant aged between 4.5 and 10.5 months, and the second belonged to a child aged 4.5–5.5 years. Later, 9 more Neanderthal teeth were found. Additionally, they identified a large horn of the hyoid bone (cornu majus) of an adult. Level X contains symbolic decorations, awls, drilled animal teeth, and pendants made of ivory. Châtelperronian industry; more ancient layers are Mousterian. 44–41 thousand years ago.

The jaws from Grotte du Renne

A—Mandible II and B—maxilla III from layer 20 of Grotte du Renne (Leroi-Gourhan, 1958). Photos by B. Maureille.

Grotte du Renne hip bone
A picture of the front and back of the hip bone fragment (AR-63) found in the Grotte du Renne. ©Gicqueau et al.
Neanderthal remains from Grotte du Renne
Grotte du Renne, Chatelperronien layer. Up = lower left canine, bottom = temporal bone. ©Thilo Parg.
Grotte du Renne hyoid bone

A digital model of the hyoid bone greater horn fragment found at the Grotte du Rennes. Juliette Henrion et al./Journal of Human Evolution, 2023

Dental remains from Grotte du Renne

An example of dental remains from the Châtelperronian layers of the Grotte du Renne (Arcy sur Cure, France) displays Neanderthal features. Photo: Shara Bailey.


Saint-Cesaire (La Roche-à-Pierrot), open-air site. Southwest France. Left bank of the Coran River, Charente-Maritime Department. The remains of a young female adolescent named “Pierrette”, a typical Neanderthal. There is evidence of a healed cranial fracture. Later, phalanges and metacarpal elements were found here. For the first time, an early Late Paleolithic culture, the Châtelperronian, was discovered among Neanderthals. In this region, Neanderthals were likely contemporaneous with the first Homo sapiens. 43–42 thousand years ago (calibrated).

Reconstruction of the St. Cesaire 1 Neanderthal skull
Woman of Saint Cesaire

Left: Computerized reconstruction of the St. Cesaire 1 Neanderthal skull (mirror-imaged completed parts are transparent) showing the bony scar in the right apical cranial vault. Scale bar = 5 cm. ©Zollikofer et al. Right: a photo of the woman of Saint Cesaire. ©Don Hitchcock.

Neanderthal fossils from Saint-Cesar

Skull and long bones of a Neanderthal from Saint-Cesar (France). ©Kenneth Garrett.

Neanderthal fossils of Saint Cesaire

Reconstruction of the discovery of the Neanderthal skull of Saint Cesaire. Paleosite: Photo by Kroko for

Bawa Yawan

Bawa Yawan, rock shelter. Western Iran. Nawdarwan Valley, Kermanshah, Central Zagros. A shed deciduous tooth (BY1) of a six-year-old child was found at a depth of 2.5 meters. It is the lower left deciduous canine, in which the crown is very well preserved, along with one-fourth of the tooth root. There were no caries or enamel hypoplasia on the tooth, but the surface was slightly worn, revealing part of the dentin. After radiocarbon dating, it was determined that the child lived approximately 43.6–41.5 thousand years ago (calibrated).

Deciduous tooth from Bawa Yawan

The new Neanderthal lower left deciduous canine (BY1) from Bawa Yawan Rockshelter. a) BY1 in different views. b) The enamel-dentin junction of the tooth shows the cervical eminence and lingual ridges. Abbreviations: B: buccal, L: lingual, M: mesial, D: distal, O: occlusal. ©Saman Heydari-Guran et al.

Neanderthal child and his tooth

The baby tooth of a Neanderthal child was found in Iran.


Zaskalnaya VI (Kolosovskaya), settlement. Ukraine. Eastern Crimea, near Ak-Kaya rock, close to the village of Vishennoye and the town of Belogorsk. Disarticulated bones of several Neanderthal children and adolescents were found, including a complete right arm, foot bones, vertebrae, and teeth. Some of them may have been the remains of a destroyed triple burial of children aged 1.2–3, 5–6 years. Also found were the lower jaws of two young Neanderthals, along with other isolated remains of at least 5–6 individuals ranging from a few months to 12–14 years old. Classic Neanderthals. There are hearths. Micoquian of Ak-Kaya industrial tradition. Middle Paleolithic, 43–41 thousand years ago.

Burial of children from Zaskalnaya VI

Triple dismembered burial of Neanderthal children from Zaskalnaya VI (Kolosovskaya), archival photo of excavation area with skeletal remains after cleaning. ©Kolosov et al.

Mandible Zsk VI-72

A mandible Zsk VI-72 from Zaskalnaya VI (Kolosovskaya), excavations of 1972. ©Stepanchuk et al.

Molars from Zaskalnaya
Morphology of the right М1 and М2 of the mandible Zsk VI-72. ©Stepanchuk et al.
Premolars from Zaskalnaya
Morphology of the left P1 of the mandible Zsk VI-72. ©Stepanchuk et al.


Khudji, an open-air site. Northern Tajikistan. Foothills of the Gissar (Hisar) Range, Shahrinav District, 40 km from Dushanbe. The second deciduous incisor has a broken root, possibly lower. Child aged 6–9 years. Classic Neanderthal. Mousterian. Levallois technology is absent. Late Middle Paleolithic, 42 thousand years ago (calibrated).

Khudji, Tajikistan

Excavations at the Khudji site.


Gruta Nova da Columbeira. Portugal. Near Bombarral, Leiria. Neanderthal tooth—fragment of the lower left molar. The inhabitants of the site also consumed rabbits and turtles. Classic Neanderthal. Late Mousterian, Levallois. 41.5 thousand years ago.


Gruta Nova da Columbeira.


Üçağızli cave. Turkey. Hatay province, on the Mediterranean coast of southern Turkey, 6 km from the border with Syria. One of the large, isolated teeth found has Neanderthal traits. 41.4–40 thousand years ago.

Üçağızli cave

Ucagizli cave.

Trou Magrite

Trou Magrite (Magritte). Belgium. Anhée commune, Walzin, Pont-à-Lesse, near the municipality of Dinant and the city of Anseremme, Namur, Wallonia. Dupont excavated the already identified human remains of six individuals (bones of three adults, one adolescent, and two children). They belong to the Mesolithic period. In addition, two more ancient finds were discovered: the diaphysis of the left femur of a male Neanderthal child aged 8–10 months. A permanent right upper canine of an adult or adolescent. Late Neanderthal (according to paleogenetic DNA analysis). Mousterian. 41 thousand years ago.

Trou Magrite

Entrance to Trou Magrite. ©Groupe Speleo la Corde Dinant.

Saint-Brais II

Saint-Brais II. Switzerland. Municipality of Saint-Brais, district of Franches-Montagnes, canton of Jura. This is part of a group of three caves. Upper left first incisor. The tooth has distinct cut marks caused by accidental contact between a cutting tool and the enamel of the labial surface of the teeth. The site is a temporary settlement. Late Neanderthal. Final Mousterian. Middle Paleolithic, 40 thousand years ago.

Neanderthal incisor from Saint-Brais

A Neanderthal tooth discovered in 1955 in the Saint-Brais II cave. An example of mass-additive dental traits in archaic humans. The complex and “robust” morphology of a Neanderthal incisor. ©Willman

Striations on the Saint-Brais tooth

Extensive instrumental striations on the Saint-Brais Neanderthal (left I1). The cutmarks are so distinct that they are visible macroscopically (left image). Right image: SEM micrograph showing extensive striations at 50x magnification. ©Willman.


La grotte de Jaurens, the Jaurens Cave. France. Nespouls, Corrèze. Robust upper Neanderthal canine. Minor wear suggests that this tooth belonged to a young individual. Late Neanderthal. 40 thousand years ago.

Neanderthal canine from Jaurens

Upper canine from Jaurens. Mesial and palatal views. ©Bouvier.


Grotte du Loup (Cosnac), “Wolf Cave”. France. Arcy-sur-Cure cave complex, Burgundy. The tooth is a right lower first or second molar. It is so worn that the relief of the molar is completely lost. Two incisors were also found here, as well as two fragments of a heavily damaged cranial vault. Classic Neanderthal. A sandstone plaque with an engraving. Final Mousterian. Transition from the Middle to the Late Paleolithic. 40 thousand years ago.

Molar from Grotte du Loup

Hominid tooth. ©Leroi-Gourhan.

Grotte des Fees

Grotte des Fées, “Fairy Cave”. France. Arcy-sur-Cure, Yonne department, Burgundy, administrative region Bourgogne-Franche-Comté. Mandible, right premolar. Mousterian, Châtelperronian. The attribution of the finding to Neanderthals or Homo sapiens remains unclear. 40 thousand years ago.

Grotte des Fees

Grotte des Fees, France. ©Lyne Sarrazin.

Neanderthal Fossil Sites with limited data

There are several sites with Neanderthal fossils for which only very scant and insufficient information is available. The publications about these caves are quite old, and the presented information is often unclear, especially regarding their chronology. In some cases, the sites may have been discovered long ago, when archaeological techniques were less sophisticated. As a result, the excavations may not have been conducted in a way that would preserve all the available evidence.

This includes a number of findings from the 19th and early 20th centuries that were described improperly, inadequately, or even erroneously. During excavations, material from different layers was often mixed, which further complicates the situation. The majority of the descriptions are already outdated. Fossil human remains discovered decades ago remain largely unknown to the scientific community. Moreover, in recent years, many new Neanderthal fossil discoveries have accumulated that remain undescribed.

Information about all these findings is insufficient to draw any conclusions or make any deductions. However, they need to be mentioned, at least in the briefest form.


Font-de-Gaume Cave. Southwest France. Near Les Eyzies-de-Tayac-Sireuil, Dordogne department. The cave is known for Magdalenian cave paintings, but long before the Cro-Magnons, Neanderthals inhabited it. The Neanderthal fossils include teeth and bones. Classic Neanderthal. The site dates between 50,000 and 40,000 years ago.

Font-de-Gaume Cave

Font-de-Gaume Cave. ©Patrick Bergeot


Genista 1. Gibraltar (British Overseas Territory). The cave is located beneath Windmill Hill. Genista Cave 1—the first of four caves. A hominid deciduous, unerupted molar was found at the site. This is probably a left second lower molar. The molar is of a classical Neanderthal. Middle Pleistocene. The molar has been dated to between 50 and 40 thousand years ago.

Gibraltar. Genista Cave 1

Genista 1 Cave.


Gruta da Salemas. Portugal. Located in the Rio de Lousa Valley, near the village of Lousa, in the municipality of Loures in the Lisbon District. A second lower deciduous molar of a Neanderthal was found in a Mousterian layer. It is a complete molar of a classical Neanderthal. However, there is a problem with dating the find. The molar was found in a Mousterian layer (Middle Paleolithic), which suggests that it is between 50 and 40 thousand years old. However, radiocarbon dating of bone samples from the site has produced a date of 25 thousand years ago. It is likely that the bone samples used for radiocarbon dating entered the cave from the outside. Perhaps subsequent dating attempts will be more successful.

Gruta da Salemas

Gruta da Salemas.

Angles-sur-l’Anglin (Roc-aux-Sorciers)

Angles-sur-l’Anglin (Abri du Roc aux Roc-aux-Sorciers, “Wizard’s Rock”). Western France. Municipality of Angles-sur-l’Anglin, right bank of the Anglin River, Vienne department, Nouvelle-Aquitaine region. Classical Neanderthal. Middle Paleolithic. The charcoal found on the site dates back between 50 and 40 thousand years ago. The Angles-sur-l’Anglin site is also famous for the rock art located on a cliff face overlooking the Anglin River. The creators of this rock art are obviously Homo sapiens.


Angles-sur-l’Anglin (Abri du Roc aux Roc-aux-Sorciers) site.


Kanal Cave. Turkey. Çevlik village, Hatay province. Left upper deciduous canine from the Mousterian layer. It is of a young Neanderthal. Levallois-Mousterian. Middle Paleolithic.

Kanal Cave

Kanal Cave, Turkey.


İncili Cave, İncirliin Mağarası (Big Cave). Southwest Turkey. District of Milas in the province of Muğla, on the northern slope of Mount Manastır overlooking the Gökçeler Canyon. The human remains include the lower jaw, upper jaw, femur, tibia, several vertebrae, and foot bones. The remains are thought to represent a single male individual aged about 50 years old. A number of features indicate Neanderthal; however, such classification is still under question.

İncili Cave

Incili Cave, Turkey.

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