Neanderthal Sites are found throughout Europe, except its northern regions, and further east, through the Levant, Caucasus, and Central Asia up to Altai. The catalog of the most significant sites of finds of the remains of Neanderthal man gives their brief description and characteristics.
Neanderthal sites and remains, ranging in age from 75 thousand years to 56 thousand years
Boccard
Grotte Boccard (Boccard Cave). France. Municipality of Créancey in the Côte-d’Or department, Burgundy. There are three permanent teeth: the left upper lateral incisor, the upper second right premolar, and the lower second right molar. Remains of hearths. Mousterian with Levallois elements. 75,000 years.
Boccard Cave. GB 78 B14 C5ccm 26 (left I2), 1: mesial view, 2: lingual view, 3: distal view, 4: occlusal view, 5: X-ray through the distal surface, 6: details of the crown illustrating gaps in the enamel and the oval-shaped fossa. ©Maureille
Vergisson
Site de Vergisson II (Grotte de Ronzevau or de la Maréchaude). Eastern France. Mâconnais region, Saône-et-Loire department, Burgundy. Several human phalanges (not yet published). Sixteen teeth belong to an individual around twenty years old who can be attributed to Neanderthals. 75,000–65,000 years ago.
In the deposits of Vergisson I “les Tasnières”, and Vergisson IV, Neanderthal remains were also discovered. Vergisson IV (Vg 4-83) is an isolated left upper central incisor of a young individual aged 7–9 years. This tooth is typical of Neanderthals, but with a predominance of left-sided striations, indicating left-handedness in the individual. In general, however, Neanderthals exhibited pronounced right-handedness. Classic Neanderthal. Mousterian of Quina type. 42,000–40,000 years ago.
Vergisson. (a) The original tooth showing the incomplete root and intact crown with some chips removed from the occlusal border; (b) the central region of the crown with the striations; (c) a magnification of the region. ©Condemi et al.
Obi-Rakhmat
Obi-Rakhmat Cave. Northeast Uzbekistan. Altai, Talas Alatau Range of the Tian Shan Mountains. The name means “Thanks for the water” in Uzbek. 6 isolated upper left teeth (OR-1) and 121 skull fragments from one adolescent aged 8–12 years. All teeth had intact crowns with developed roots, and exhibited taurodontism. A study of mitochondrial DNA points Obi-Rakhmat-1 to classic Neanderthals. Mousterian-Levallois. Middle Paleolithic, 75,000–70,000 years ago.
Obi-Rakhmat. Parietal reconstruction of OR-1 composed of 27 cranial fragments. (Modified after Glantz et al., 2008 and Krivoshapkin)
The left maxillary dentition of the Obi-Rakhmat hominin (OR-1). Left: lateral incisor, canine, third, and fourth premolars (upper and lower teeth, respectively), first and second molars (upper and lower teeth, respectively). ©Smith et al. Right: (a) upper left lateral incisor (I2) in mesial view and (b) lingual view; (c) upper left canine (C) in mesial view and (d) lingual view; (e) upper third premolar (P3); (f) upper fourth premolar (P4); (g) upper first molar (M1); (h) upper second molar (M2). ©Bailey et al.
Bisceglie
Bisceglie (Santa Croce di Bisceglie, Grotta di Santa Croce), Santa Croce caves. Italy. Bisceglie municipality, Barletta-Andria-Trani province, Bari, Apulia. Biscegli 1 is the diaphyseal body of the right femur (including the lesser trochanter and part of the neck) and other fragments from an adult individual. Classic Neanderthal. Charentian Mousterian of Quina type. Approximately 75,000–65,000 years ago.
Neanderthal right femur from Santa Croce di Bisceglie.
Archi
Archi. Italy. San Francesco, Calabria. Mandible with five teeth of a 3-year-old Neanderthal child (Archi 1): right lower canine; first and second molars, right and left. Only deciduous teeth had erupted. As in the mandibles of other juvenile Neanderthals, this specimen is broad across the symphyseal region, which lacks a central keel, mental fossae, and thickened inferior margin. The mandibular symphysis was incompletely fused. Classic Neanderthal. 75,000–55,000 years ago.
Archi 1 is a Neanderthal child. The slight indentation of the inferior margin at the midline probably reflects an incomplete fusion of the symphysis. Bar=1 cm. Istituto Italiano di Paleontologia Umana. ©Schwartz et al.
Fossellone
Grotta del Fossellone (Fossellone Cave). Central Italy. Mediterranean coast, village of San Felice Circeo, Monte Circeo, province of Latium.
Fossellone 3 (previously designated as Circeo 4) is a fragment of the mandibular symphysis and three left teeth (premolar and first and second molars) from a 9–10-year-old child, whose bones have not yet reached maturity. There is a tendency for root separation. The root of the second left lower premolar has developed more than half its length. This stage of development corresponds to an average age of 9 years. There are no signs of wear on the cusps. The first left molar is almost complete; only two mesial cusps of the root are chipped. The wear is moderate, not yet exposing the dentin. The left second lower permanent molar is well preserved. The occlusal surface shows no wear. The roots have only reached half their height, corresponding to an age of 10 years. It is likely that the mandible belonged to the same individual as the three teeth. Classic Neanderthal. Mousterian. 74,000 years ago.
Vaufrey
Grotte Vaufrey (or Grotte XV). France. Falaise du Conte, Cénac-et-Saint-Julien, Dordogne. Vaufrey 1 is the second lower left molar of an adult individual. Classic Neanderthal. Mousterian of Quina type. Middle Pleistocene, 74,000 years ago.
Vaufrey 1. (A) buccal, (B) mesial, (C) lingual, (D) distal, and (E) occlusal views. ©Garralda et al.
Guattari
Guattari Cave, Mount Circeo (San Felice Circeo village). Italy, near Rome. One of the most well-known locations among Italian Neanderthal sites. Remains of 11 Neanderthals. Well-preserved skull (Guattari 1) and mandible with a molar (Guattari 2) of an adult male individual—68,000–50,000 years ago. Partial mandible with teeth from an adolescent (Guattari 3)—74,000–60,000 years ago. Skull of a probable adult male (Guattari 4). Female skull (Guattari 5). Anterior part of mandible with chin, unidentified sex (Guattari 6). Right femur diaphysis of a probable adult male (Guattari 7). The metacarpal bone of the right thumb (Guattari 9). Most of the bones have hyena teeth marks. The oldest of the remains dates from between 100,000 and 90,000 years ago, while the others are believed to be younger, dating from 68,000 to 50,000 years ago. Typical Mousterian.
Neanderthal finds, Guattari Cave, Circeo, Italy, excavated in 2019–2021. ©Mac9
The Italian Culture Ministry said the Guattari Cave in San Felice Circeo was “one of the most significant places in the world for the history of Neanderthals.” ©Emanuele Antonio Minerva/Italian Culture Ministry via AP
Guattari-1 ©Museo Nazionale Preistorico Etnografico
Guattari-5 female skull.
Cotencher
La Grotte de Cotencher (Cotencher Cave). Western Switzerland. Areuse Canyon, Rochefort, near the city of Neuchâtel, Swiss Jura.
Upper jaw of an adult woman around 40 years old. Characterized by gracile form and pronounced microdontia. Gracile (Mediterranean) Neanderthal. Charentian Mousterian of Quina type. Middle Paleolithic, 73,000–68,000 years ago.
Cotencher. Mandible of a Neanderthal woman. ©Bill Harby.
Shanidar
Shanidar Cave. Iraq. Erbil, Zagros Mountains, Kurdistan. At least 10 individuals, including adults and three children. An adult specimen, Shanidar Z, is over 70,000 years old. His remains consist of a crushed but complete skull, upper thorax, and both hands. Specimen Nandy (Shanidar 1) is the partial skeleton of an adult male aged 35–45 years. He had part of his right arm amputated. This injury had healed, indicating care by other group members. The brain volume is 1600 cm3. The skeletal remains of Shanidar 2 are upper and lower jaws and teeth, skull fragments, and non-cranial bones. 60,000–45,000 years ago. Shanidar 3, a male approximately 40–50 years old, has a height of 1.69 m. Teeth are heavily worn. He had a penetrating wound to the left ninth rib. At the time of his death, the wound on his bone had started to heal. Shanidar 4, a male weighing 72 kg, was buried with flowers. Shanidar 5 is the partial skeleton of a man aged 35–50 years, weighing 68 kg. 50,000–46,000 years ago. Shanidar 7 is the skeleton of a child. 129 hearths have been counted in the Mousterian layer. 73,000–40,000 years ago.
Rivaux
Le Rivaux, open-air site. Southeastern France. Near the city of Puy-en-Velay, Espaly, Haute-Loire department. Lower part of a tibia bone. Mousterian industry. 70,000 years ago.
Rivaux. Fragment of the tibia. ©Raynal.
Nadale
Nadale (De Nadale Cave), a small cave. Northern Italy. Southern slope of the Berici Hills, Veneto region. Nadale 1—a shed first deciduous right lower molar of a Neanderthal. Complete crown and an almost completely resorbed root. Hearth remains. Mousterian of La Quina type. Middle Paleolithic, 70,000 years ago.
This is a 3D reconstruction of the exfoliated Neanderthal milk tooth found in the De Nadale Cave. Nadale 1 is a lower right first deciduous molar. Right: Nadale 1 tooth in buccal (B), lingual (L), occlusal (O), mesial (M), and distal (D) views. ©Arnaud et al.
Regourdou
Regourdou. France. Montignac-sur-Vézère, near (800 m) the famous Lascaux Cave, Dordogne. Regourdou 1—mandible and many skeletal elements. Later, numerous additional skeletal elements were found (vertebrae, ribs, parts of the upper and lower limbs, sternum, etc.). This is possibly the burial of a young, almost adult Neanderthal. This is possibly the burial of a young, almost adult Neanderthal (25 years old). This Neanderthal was about 162–172 centimeters tall and weighed approximately 64–66 kilograms. The reconstructed pelvic bone showed that the skeleton belonged to a male. The second individual (Regourdou 2) is only a calcaneus (heel bone). Mousterian. 70,000 years ago.
The skeleton of Regourdou 1. ©Collections Ville de Perigueux, Musee d’Art et d’Archeologie du Perigord
Occlusal view of Regourdou 1 mandible. The apparent malpositioning of the teeth is due to reconstruction. ©P. Semal, Royal Belgian Institute of Natural Sciences, Brussels.
Diverse rib fragments. ©Maureille et al.
The new right calcaneus is attributed to Regourdou 2. Medial, lateral, and dorsal views. ©Maureille et al.
Roc de Marsal
Roc de Marsal Cave. France. Near Campagne-de-Bugue, Dordogne. A fairly well-preserved partial skeleton of a child aged 3–4 years, height 80–100 cm. The skeleton was inside the pit (possibly of natural origin), suggesting burial. In 2019, a review of the museum’s collection revealed three additional fragments that further complement this already very complete skeleton of a Neanderthal child. Gracile Neanderthal. Mousterian. 70,000 years ago.
The skeleton of a Neanderthal child, discovered at the Roc de Marsal site in Dordogne. ©Leo Fyllnet
Skull of the infant. Le Musee National de Prehistoire, Les Eyzies-de-Tayac. ©Don Hitchcock
The model of a Neanderthal child from Roc de Marsal. The child is based on the three-year-old found at Roc de Marsal. Sculptor: Elisabeth Daynes. ©Don Hitchcock.
Mandible of the infant. ©Hitchcock
Macassargues (Verrerie)
Grottes de Macassargues (also known as La Verrerie, Bragassargues). France. Near the village of Robiac, commune of Montmirat, Canton of Saint-Mamert, Nîmes district. Neanderthal remains: the second permanent lower left molar of an adolescent aged 13 to 15 years old, and the upper third of the right radius of a child approximately 10 years old. Classic Neanderthal. Mousterian. 70,000 years ago.
Macassargues. Second left molar (Homo neanderthalensis).
Teshik-Tash
Teshik-Tash, Man-Kuan cave. Uzbekistan. Bajsuntau Mountain Range. Teshik-Tash means “holey stone” in Uzbek. Boy aged 8–9 years old. It was possibly a burial, as he was surrounded by five pairs of wild goat horns. The horns were dug into the ground. The skull was reconstructed from about 150 fragments. The mtDNA sequence of Teshik Tash is closely related to the mtDNA sequence of a Neanderthal from Scladina. Cranial capacity: 1490–1500 cm3. Classic Neanderthal. Mousterian. 70,000 years ago.
The Teshik-Tash skull, discovered in 1938 and estimated to be 70,000 years old, is believed to have been from a nine-year-old boy. This specimen was excavated in Uzbekistan and represents one of the easternmost Neanderthal finds.
Reconstruction of a burial site Teshik-Tash 1, with a skull and bones from Neanderthals in Uzbekistan, where remains were discovered. ©Natural History Museum, London/Science Photo Library.
Portel-Ouest
Le Portel-Ouest (West Cave), also known as “Cap del Saut”. France. Ariege, Pyrenees. Seven children, one or two adolescents, and one adult individual. 23 teeth, fragments of the parietal bone. Charentian Mousterian type. 70,000–65,000 years ago.
Parietal fragment of an adult from sublevel F2 at Portel-Ouest with a quasi-close sagittal suture. External, internal, and anterior views. ©Becam, Chevalier
Mismatching of interproximal facets between right dm1 and dm2 from Portel-Ouest. 2. Internal anatomy of permanent teeth from Portel-Ouest. M: Mesial; B: Buccal; L: Lingual. Blue: enamel; yellow: dentin; red: pulp cavity. All are left teeth. ©Becam, Chevalier
Ein Qashish
Ein Qashish, open-air site. Northern Israel. The site is situated on the left bank of the Qishon River, east of Mount Carmel. There are three partial skeletons. EQH1, a nondiagnostic skull fragment; EQH2, an upper right third molar (RM 3); and EQH3, lower limb bones of a young Neanderthal male. Seasonal hunting camp. Mousterian, Levallois. 70,000–60,000 years ago.
Oliveira
Gruta da Oliveira (Torres Novas). Portugal. The cave is part of the Almonda karst system. Fragments of Neanderthals. Teeth: a heavily worn partial premolar or molar of an adult individual (Oliveira 6); distal right mandibular molar (Oliveira 8) of a 9-year-old child; and first right premolar of the lower jaw (Oliveira 9) of a 13-year-old adolescent. Skeletal remains: several cranial elements, rib, etc. Shoulder bones, phalanx, and left tibia 70,000–60,000 years ago; another phalanx and right ulna 47,000–46,000 years ago. Seafood was consumed. About 10 hearths remains. Mousterian. Middle Paleolithic.
Neanderthal remains in Gruta da Oliveira (the Olive Tree Cave). a. right humerus; b. distal right humeral diaphysis; c. phalanx of hand; d. right manual proximal phalanx 2; e. right ulna; f. left tibia; g. humerus. ©Willman et al.; ©Zilhao et al.
a) Oliveira 6 postcanine partial crown and root in external (below) and occlusal-external (above) views.
b) Oliveira 8 right mandibular molar (probably M2 or M3). Left: distal root and crown in mesial view. Upper right: distal crown in occlusal view. Lower right: the crown in distal view.
c) The Oliveira 9 right P3 in mesial, distal, buccal, lingual, and occlusal views. Brackets on the occlusal view indicate the positions of the interproximal facets. Scale bar: 1 cm. ©Willman et al.
Mezmaiskaya
Mezmaiskaya Cave. Russia. Adygea, North Caucasus. Neanderthal Sites: the remains of three individuals were found here. Specimen Mezmaiskaya 1 is the almost complete skeleton of a child aged from two weeks to two months, 70,000–60,000 years ago. Mezmaiskaya 2: 24 skull fragments of a child (male) aged 1–2 years, 45,000–42,000 years ago (calibrated). Mezmaiskaya 3: teeth found dating back 45,000–43,000 years. Classic Neanderthals. The remains of hearths are lined with stone tiles, bone tools. Mousterian, Micoquian.
Left: A virtual reconstruction of the Mezmaiskaya Neanderthal infant. (A) Skeleton. (B) Skull in right lateral view. (Scale bars, 5 cm.) ©Ponce de Leon et al.
Right: A virtual reconstruction of Mezmaiskaya. Parts obtained via mirror imaging are plotted in a darker shade. The scale bar is 10 mm. ©Gunz et al.
View from the west of the Mezmaiskaya 1 skeleton, which is lying in anatomical arrangement; the black arrow points to the north. ©Golovanova et al.
Neanderthal incisor (Mezmaiskaya 3) from Mezmaiskaya Cave. ©Andreeva et al.
Forbes’ Quarry
Forbes’ Quarry Cave. Gibraltar (British Overseas Territory). Specimen Gibraltar 1: skull of an adult woman around 40 years old (nickname “Nana”). Genetically close to individuals from Scladina (Belgium), Hohlenstein-Stadel (Germany), “Thorin” from Mandrin Cave (France), and Mezmaiskaya (Russia). Teeth are heavily worn. Brow ridges are prominent but lightly built. Large and protruding nose. This is the second find among all Neanderthal sites, but initially it was not recognized as Homo neanderthalensis. 70,000–60,000 years ago.
The adult female Neanderthal cranium discovered at Forbes Quarry, Gibraltar.
Petit-Puymoyen
Petit-Puymoyen. Southwest France. Valley of the Eaux-Claires stream, commune of Petit-Puymoyen, Charente. The site includes the rock shelter L’abri Commont, located in front of the cave. At the Commont shelter, two fragments of mandibles and a fragment of the maxilla of adolescents were found. In the cave, the right temporal bone of an infant (PPm5), the right capitate and left hamate bones (PPm6), a fragment of an occipital bone, the left cuboid and left medial cuneiform bones, as well as two incisors and several other teeth, were found. In total, there are the remains of six people. Mousterian (Quina and Ferrassie variants). 70,000–60,000 years ago.
The Petit-Puymoyen 1 mandible in occlusal (a), left lateral (b), and symphyseal (c) views. (d) Close-up photograph of the exposed root system in the Petit-Puymoyen 3 mandible, indicating the mesiobuccal (MB), distobuccal (DB), and distolingual (DL) root tips. Scale=1cm. ©Quam et al.
The Petit-Puymoyen 7 occipital fragment in endocranial (upper left) and external (upper right) views and detail of the suprainiac region (inset). Note the roughened area of bone surface in the suprainiac area, as well as the emissary foramen above it. Scale=2cm. Inset scale=1cm. ©Quam et al.
Simard
L’abri Simard, Grotte Simard (also known as René Simard). France. The cave is located near the border of the municipalities of Puymoyen and Torsac, in the valley of the Eaux-Claires stream, at a place called Les Prés-Marty, Charente, Nouvelle-Aquitaine. During excavations, a large number of human bones and fragments (children and adolescents), totaling one hundred and five elements, were discovered. Several individual teeth, vertebrae, fragmentary iliac and ischial bones, clavicles, bones of the arms, legs, feet, and femurs, as well as a long bone, identified three children around 12 years old and two children aged 2 years. Classic Neanderthal. Mousterian of Quina type. First half of the Middle Paleolithic, 70,000–60,000 years ago.
Grotte Simard. Rocks of the Eaux-Claires valley, Puymoyen. ©Jack ma
Manie
Arma delle Mànie, Cave of Manie. Italy. Near the town of Finale Ligure, just 500 meters from the Fate Cave, in the province of Savona. Le Mànie 1 is the lower incisor of an adult individual. Mousterian, Levallois. 70,000–60,000 years ago.
Cave of Manie.
Barakaevskaya
Barakaevskaya Cave (Barakai). Russia. Gubskoe Gorge, near the city of Maykop, North Caucasus. In Barakaevskaya Cave, remains of Neanderthals were found—a robust mandible (a child aged 2–3 years), cranial bones, and 10 isolated teeth with taurodontism. Classic Neanderthals. Hearths and fireplaces remains. Grinding stones for grains. Mousterian and Micoquian industries. Middle Paleolithic, 70,000–57,000 years ago.
The Neanderthal child’s mandible from Barakaevskaya Cave. 1–4 views from four sides; 5 the position in the Mousterian deposits in situ. ©Levkovskaya
©Haritonov
Shukbah
Shukbah Cave (Shovakh, Shubabiq, Shuqba). Northern Israel. Hebron Hills, north of Jerusalem, Palestinian West Bank, 8 km from the Sea of Galilee. Lower molar with mild taurodontism (EM 3869) and cranial fragments of a nine-year-old child. The molar is very large, its roots are relatively short, and it has four branches fused along most of its length. Neanderthal features predominate. Hearth remains. Nubian Levallois technology. 70,000–56,000 years ago.
Photographic and virtual renderings of surfaces of the external morphology, dentin, and pulp are illustrated in occlusal, mesial, distal, buccal, and lingual views. ©Natural History Museum, London
Bacho Kiro
Bacho Kiro. Bulgaria. Municipality of Dryanovo. The Bacho Kiro cave is primarily known for the findings of early Homo sapiens, among the first to arrive in Europe. However, more ancient fossils related to Neanderthals have also been found there. These are mainly fragmentary remains of cranial bones, jaws, teeth, and other small bone fragments. This includes a fragment of a mandible (numbered 1124). Classic Neanderthal. Middle Paleolithic, 70,000–50,000 years ago.
Fragment of the mandible of a Neanderthal from Bacho Kiro and other remains.
Remete Felso
Remete Felsö (Remete Upper Cave). Hungary. Near the town of Mariaremete, outskirts of Budapest. Upper left canine from an adolescent, and two right lower incisors. The teeth are large and heavily worn. Classic Neanderthal. Szeletian industry. 70,000–50,000 years ago.
Remete-Felso. Right lower permanent teeth: buccal view 1—i2, 2—i1, 3—C; lingual view: 1—i2, 2—i1, 3—C. Right lower canine: the occlusal view illustrates the high degree of attrition. ©Tillier et al.
Placard
Le Placard, Grotte du Placard. France. Village of Rochebertier, commune of Vilhonneur, Moulins-sur-Tardoire, Charente. Several isolated teeth (premolars and molars) and a jaw fragment belong to Neanderthals. Several hearths. Classic Neanderthal. Charentian industry, including Mousterian of Quina type. Middle Paleolithic, 70,000–50,000 years ago.
The Placard Cave is located in Vilhonneur, in the heart of the Tardoire valley. ©Jack ma
Krijn
Krijn. Netherlands. Coastal waters of the North Sea, province of Zeeland. A fragment of the skull (left frontal bone with a segment of the supraorbital torus) of a young Neanderthal who lived in Doggerland, a prehistoric landscape now submerged underwater off the Dutch coast. This Zeeland Ridge Neanderthal was discovered in sediments extracted from the bottom of the North Sea. It is the first Pleistocene fossil hominin found at the bottom of the sea. The partial frontal bone shows a lesion caused by an epidermoid cyst. Age: 70,000–50,000 years.
The fossil of the frontal bone and its location on the Neanderthal skull. ©Servaas Neijens
Dederiyeh
Dederiyeh Cave. Northern Syria. Afrin District, Mount Simeon. Burial of two children aged 1.5–2.5 years. Dederiyeh 1—the nearly complete skeleton of a 2-year-old child; estimated height 80 cm. Dederiyeh 2—a similar partial skeleton. Additionally, separate bones of at least four Neanderthals—adults, children, and infants—were found. Remains of three children’s burials. Classic Neanderthals. Mousterian. 70,000–50,000 years ago.
Dederiyeh 1. A two-year-old Neanderthal child from Northern Syria.
Virtual reconstruction of the Dederiyeh 1 (left) and Dederiyeh 2 (right) skulls. (Scale bar, 5 cm.) ©Golovanova et al.
Dederiyeh 2 Neanderthal skull.
Wezmeh
Wezmeh. Western Iran. Zagros Mountains, near Eslamabad, Central District of Eslamabad-e Gharb County. Several bone fragments and teeth were found. Wezmeh Child (Wezmeh 1) is an isolated, unerupted right upper premolar of a Neanderthal child aged 6–10 years. 70,000–40,000 years ago.
Unerupted Neanderthal maxillary right premolar tooth (P3 or possibly P4) of an individual between 6 and 10 years old from Wezmeh Cave. ©Zanolli et al.
Painting of a 9-year-old Neanderthal child by Tom Bjorklund.
Warendorf
Warendorf, sand quarry. Germany. Next to the the Kottruper lakes, near Neuwarendorf, not far from Münster, North-Rhine-Westphalia. An incomplete right parietal bone, nearly identical to specimens from La Chapelle-aux-Saints and Feldhofer. Technocomplex of wedge knives (Keilmessergruppen). Classic Neanderthal (confirmed by DNA analysis). 70,000–40,000 years ago.
Neanderthal skull fragment from Neuwarendorf. Internal (with imprints of blood vessels) and external views of the parietal bone.
Mandrin
Mandrin Cave (Mandrin Rock Shelter). Southern France. Located in the Rhône Valley, Drôme department, near Montélimar and Malataverne, 130 km north of Marseille. It resembles more of a shelter than a cave. A total of 9 typical Neanderthal teeth have been found, although the attribution of one tooth fragment is in question. There are traces of soot from fires on the ceiling. 69,000–46,000 years ago.
In the latest archaeological level of the Mandrin cave, the remains of an adult male Neanderthal skeleton were found. The scientific group named the specimen “Thorin” in honor of the last king of the dwarves in Tolkien’s works. This individual is the only nearly complete Neanderthal found since the discovery of “Pierrette” 42 years ago (in Saint-Cesaire). Several hundred very fragmentary remains were found—wrist elements, five phalanges of the left hand, a series of cranial elements, a fragmentary mandible, teeth, and others. This individual had supernumerary lower molars (fourth molars). Thorin’s fourth molars had an irregular shape and only one root. 52,000–49,000 years ago. Mousterian of the La Quina type, Neronian.
The discovery of Thorin. Fossils of a Neanderthal nicknamed Thorin in situ. ©Ludovic Slimak et al.
The Grotte Mandrin human remains. Plate illustrating the nine dental elements. LLdm2, deciduous lower left second molar; LLM1, permanent lower left first molar; LRM3, permanent lower right third molar; URdm2, deciduous upper right second molar. m–mesial and o–occlusal views, except Man 15 m–mesial and d–distal views. ©Slimak
Finds of Neanderthal teeth in the Mandrin Grotto. 3D reconstruction of Thorin’s jaw and teeth. ©Ludovic Slimak et al.
Banyoles
Banyoles (Catalan), Bañolas (Spanish), Pla de la Formiga (Porqueras) open quarry. Northeastern Spain. Girona. Archaic mandible of a 40-year-old woman. The taxonomic affiliation of the jaw is still being discussed. There is evidence of fish consumption. 66,000–45,000 years ago.
The jaw of a Neanderthal found in 1887 by Pere Alsius in a quarry of the Pla de la Formiga (Porqueras). It is believed from a woman of about 40 years prior to the last glaciation (Wurm).
Neanderthal mandible from Banyoles. ©Roberto Saez.
Pelenos
Las Pelenos (Monsempron). France. Monsempron-Libos, Lot-et-Garonne, Agen. Remains of approximately 14 individuals, including teeth, fragments of the mandible, and two parts of the cranial vault. Mousterian. Around 65,000 years ago.
Las Pelenos. Upper right second permanent molar. D distal view, V buccal view, M mesial view, L lingual view, O occlusal view, A apical view. ©Scolan
Mas-Viel
Le Mas-Viel (Mas-Viel Cave). France. Commune of Saint Simon, Lot, Quercy. Classic Neanderthal. Mousterian of the Quina type. Middle Paleolithic. 65,000 to 50,000 years ago.
Jonzac (Chez-Pinaud)
Jonzac (Chez-Pinaud). Southwest France. This Neanderthal site is located just outside the city limits of Jonzac in the CharenteMaritime department. Neanderthal tooth—upper right premolar (P4 or P3). The surface of the tooth was significantly damaged, and the crown was heavily worn during the individual’s lifetime. The root is bifurcated. Classic Neanderthal. Bone tools. Mousterian of the La Quina type, Levallois, and Mousterian of Acheulian tradition. 65,000 to 48,000 years ago.
Jonzac/Chez-Pinaud: human tooth CPN05 E14-Z13 SW-07. From left to right: distal, mesial, buccal, lingual and occlusal views. Dentin is segmented in gray, enamel is segmented in white. ©Jaubert et al.
Pinilla del Valle
Pinilla del Valle. Spain. Lozoya River Valley, municipality of Pinilla del Valle, province of Madrid. On the hill of Calvero de la Higuera, there are 5 Neanderthal sites, of which 3 contain Neanderthal remains. A total of 9 teeth have been found.
Cueva de la Buena Pinta
Cave of the Good Pinta, The Promising Cave. Two molars belong to one individual. Remains of hearths. Mousterian. 60,000 to 50,000 years ago.
Cueva del Camino
2 molars of an individual aged 30 to 35 years. There are marks on the teeth left by a sharpened object used for tooth cleaning. 63,400 years ago.
Des-Cubierta Cave
(Cave Des-Covered), Descubierta Cave. In the hearth, 4 teeth were found, 3 of them deciduous, and a jaw fragment belonging to the “Lozoya girl” aged 2–3 years. Possibly a burial. Classic Neanderthals. Mousterian. 42,000 to 38,000 years ago.
This specimen is from the Pinilla del Valle site, Cueva del Camino, near Madrid, Spain. These teeth are molars from a Neanderthal individual aged 30 to 35. The teeth include markings made by a pointed object used to clean teeth. ©Trueba
Neanderthal remains from Pinilla del Valle. Left: Cueva Des-Cubierta; right: Cueva de la Buena Pinta. ©Roberto Saez
Kebara
Kebara (Mugharat al-Kabara). Israel. Cave on the western slope of Mount Carmel. Two skeletons of six-month-old infants and the specimen nicknamed “Moshe” (Kebara 2), a nearly complete skeleton of an adult Neanderthal, male, height 173 cm, weight 76 kg (burial). The hyoid bone, indicative of anatomical capacity for speech, has been preserved. Hearth remains. 62,000 to 60,000 years ago.
Kebara 2. Cast of a Neanderthal burial in Kebara Cave, Kebara Cave, Mt. Carmel, Israel, from around 60,000 years ago. The position of the upper limbs suggests the body was deposited in the grave before rigor mortis set in. The head is absent. Some scientists believe it was removed after burial, but we don’t know why.
Chapelle-aux-Saints
La Chapelle-aux-Saints cave (also Bouia Bonneval). Central France. Bouffia Bonneval Cave, Correze department. A quite complete skeleton of an adult male around 40 years old, suffering from severe arthritis and having lost most of his teeth. The discovered remains included a skull with the lower jaw, a clavicle, two nearly complete humeri, two incomplete radius bones, several arm bones, fragments of ilium, two incomplete femurs, parts of tibia bones, several foot bones, and numerous vertebrae and ribs. Nicknamed: “The Old Man of La Chapelle”. Cranial capacity: 1625 cm3, weight: 77 kg. Burial ritual. Classic Neanderthal. Late Mousterian. 60,000 years ago.
Skull of La Chapelle aux Saints (France). ©Mounier, ©Bone Clones.
Reconstruction of the Neanderthal burial at La Chapelle-aux-Saints, France, on display at Musee de La Chapelle-aux-Saints, Correze, France. ©Mourre.
Eguisheim
Eguisheim (German: Egisheim). Northeastern France. Located 7 km from the city of Colmar, in the department of Haut-Rhin (Upper Rhine), Alsace. Two fragments of the skull (frontal and parietal bones). Found in 1865, it still remains dated as belonging to the “Mammoth Epoch”. The age of the find can only be estimated at around 60,000 years.
Comparison of the Neanderthal skull and the Eguisheim skull. Profile outlines of the Feldhofer Neanderthal skull and the Eguisheim skull.
Abric Romani
l’Abric Romaní, rock shelter. Spain. The cliff of Cinglera del Capelló, municipality of Capellades, along the Anoia River, 50 km from Barcelona, Catalonia. Fragmented Neanderthal skull (part of the parietal, temporal, and jugale bones). Some calcified wooden structural elements and their imprints have been preserved, along with about a hundred wooden tools. Only at level Q were the remains of 300 hearths and accumulations of ash and charcoal found. Middle Paleolithic, 60,000 years ago.
The partial remains of a Neanderthal skull were discovered at the Abric Romani site in Capellades, Spain. ©Maria D. Guillen
Marillac
Marillac (also known as Les Pradelles). Southwest France. Site is located in the municipality of Marillac-le-Franc, 5 km east of La Rochefoucauld, near the Ligonne River, Charente.
About 30 Neanderthal remains exist, including fragments of skulls and jaws. Marillac 3—incomplete right hemifrontal bone (with perimortem manipulation scars) of an adult individual. The inner surface of this skull fragment is very rough and uneven. Three postcranial skeleton fragments: adults—incomplete diaphysis of the right radius (M24) and left fibula (M26). Also, the diaphysis of the right femur (M25) of a child (7–10 years old) is similar to that of Teshik-Tash. In 2010, a well-preserved fragment of the left femoral diaphysis of an adult Neanderthal was found (LP10-D13). Teeth from nine individuals (14 permanent and 2 deciduous teeth). This Neanderthal site is a hunting camp without traces of hearths, and there is no evidence of residential use of the site. Bone with notches. Traces of cannibalism. Mousterian of the La Quina type. 60–58 thousand years ago.
Human remains discovered during 2001 to 2004 field activities at Marillac. ©Maureille et al.
Fumane
Fumane (also known as Riparo-grotta di Fumane, Stazione della Neve, Riparo Solinas). Stone shelter expanding into a cave. Northern Italy. Valpolicella, at the foot of the Venetian Pre-Alps, in the province of Verona. Fumane 1—a fallen-out second deciduous molar of a Neanderthal child aged 10–11 years. Fumane 2 and 3 belong to Homo sapiens. The teeth Fumane 4 and 5 are the right upper and lower incisors of a 6-year-old child. Caregivers first gave solid food to both humans and Neanderthals at around six months of age, as revealed by the analysis of ancient teeth. The study shows that Neanderthal children grew and were weaned as babies at a similar time to that of modern humans. Remains of hearths. Painted seashells were found, possibly used as decoration (perhaps a necklace). Classic Neanderthal. Mousterian, Levallois. Middle Paleolithic. 60–50 thousand years ago.
Neanderthal tooth Fumane 1, a lower left second deciduous molar. This is a 3D reconstruction of the Neanderthal exfoliated milk tooth found in the Fumane Cave.
Jarama
Jarama VI site, Jarama. Central Spain. Left bank of the Jarama River, Valdesotos, Guadalajara. The first metatarsal bone was embedded in fluvial sands and silts. Mousterian. The end of the Middle Paleolithic, 60–50 thousand years ago.
Dorsal, medial, plantar, lateral, and proximal views of the Jarama VI first metatarsal (scale in cm). ©Lorenzo et al.
Sakajia
Sakajia Cave (Sakažhia). Western Georgia. Located near the village of Godogani, Terjola Municipality in the Imereti region. A fragment of the left maxilla with 4 teeth and an isolated left first lower molar. The presence of alveolar incisor sockets and dental row indicates that approximately half of the palate is preserved. The canine, premolars, and first molars are in anatomical position. An adult individual, not younger than 30 years old. Neanderthal with progressive (Near Eastern) features. Apparently, seasonal settlements were located here. Mousterian, Levallois. Microlithic tools. The middle stage of the Upper Paleolithic, 60–50 thousand years ago.
Sakajia. 1—left maxillary lateral; 2—left M1 occlusal. ©Rivals, Chevalier
San Bernardino
San Bernardino. Italy. Berici Hills, near the Broion Cave, Mossano municipality, Veneto. San Bernardino 5, a lower right second deciduous incisor of a Neanderthal. Classic Neanderthal. Late Mousterian. 60–40 thousand years ago. The lower right third molar (San Bernardino 4) and distal phalanx (San Bernardino 3) belong to Homo sapiens of the medieval age.
San Bernardino site and incisor of a Neanderthal.
Salzgitter-Lebenstedt
Salzgitter-Lebenstedt, open-air site. Northern Germany. Near Braunschweig, Lower Saxony. Five Neanderthal remains have been identified here, including one occipital bone, two articulated fragments of the right parietal bone, and two fragments of the left femurs (one adult and one infant). Among the finds at Salzgitter-Lebenstedt are numerous animal bones and tools made from them. Micoquian industry, Levallois. Middle Paleolithic, 58–54 thousand years ago.
Sidron
Sidron Cave (Cueva del Sidron). Northern Spain. Near the village of Borines, municipality of Piloña, Asturias. The cave contains jaws and bone fragments, totaling over 1,600 bones, including more than 200 teeth. In the Galería del Osario, 13 individuals have been identified, including seven adults (three men, four women), three teenagers, two minors, and one child. They constituted a closely related kin group. It is likely that cannibals placed them there. One of the individuals, El Sidrón J1, was between 7 and 8 years old at the time of death, with a cranial capacity of 1,330 cm3. The remains of El Sidrón Adult 2 (A2) is maxilla and mandible. Also, a left fully mature foot composed of in situ articulated bones was recovered. The mitochondrial DNA indicates the patrilocality of this Neanderthal group. Late Pleistocene, 58–45 thousand years ago (calibrated).
Fossil remains of a Neanderthal child who lived 49,000 years ago, discovered in the El Sidron cave in Asturias. ©MNCN-CSIC.
A Neanderthal left fully mature foot from El Sidron, still embedded in the rock and after cleaning. ©Tom Higham, ©Rios et al.
Lateral view of the lower and upper jaws (Mandibles I and II) of adult hominids from El Sidron Cave in anatomical connection. ©J. Fortea
Hortus
Hortus Cave (Grotte de l’Hortus). France. The site is located in the Pic Saint-Loup massif, near Montpellier, commune of Valflaunès (Hérault). Remains of 20–33 individuals, adults, and children.
Hortus II includes the mandibular symphyseal region and isolated permanent right and left first molars with moderate wear. The age of the child is about 6.5–8 years old. The Hortus II mandible is associated with a partial maxilla from Hortus III. Hortus IV consists of the mandibular corpus, ascending ramus, and several teeth. The age of this late subadult or young adult is 18–30 years old. Hortus V includes a small fragment of the left distal corpus of the jaw and several isolated teeth, aged up to 18–25 years. A right third lower molar from a young adult estimated to be 22–30 years old represents Hortus VI. Hortus VIII is a well-preserved upper first molar, age 26–34 years old. Hortus XI is the right upper third molar, age 45–50 years old. Mousterian. 57,000 years ago.
Left: Hortus IV mandibular. Right: Neanderthal maxillary canine with a “pronounced” lingual cusp. ©Hublin
Left: On the right mandibular permanent first molar (M1) of Hortus II, dental features can be observed, such as a metaconulid (a) and an anterior fovea (b).
Right: The left (a) and right (b) mandibular permanent first and second molars (M1 and M2) of Hortus IV exhibit well-demarcated anterior foveae.
Tour of La Chaise
Tour of La Chaise Cave (Grotte de la Tour). Western France. Located 30 km from Angoulême, near La Chaise-de-Vouthon, Charente. Partial diaphysis of the left femur of an adult male, approximately 50% of its estimated original biomechanical length. There are multiple tooth marks on the bone. 57,000 years ago.
CDV-Tour 1 (left femoral shaft) in anterior (a), posterior (b), medial (c), and lateral (d) views (proximal is upper). Scale bar: 1.5 cm. ©Puymerail
Dzerava skala Cave (Palfy)
Dzeravá skála Cave (also Pálfy, Pálffybarlang). Western Slovakia. Mokrá dolina, municipality of Plavecký Mikuláš, Malacky District, Bratislava Region. Isolated teeth, including the right lower second molar. Wood charcoal. Classical Neanderthal. 57–50 thousand years ago.
Dzerava skala Cave. ©Doronenko
Quina
La Quina. Southwest France. The Neanderthal site is located approximately 5 km from the village of Villebois-Lavalette, Gardes-le-Pontaroux, near the Voultron River, in Charente. Fossil remains of about 27 Neanderthals, adults and children, have been found: skulls, isolated teeth, and bones. 2 astragalus, apparently belonging to two legs of the same individual. They were a few centimeters apart and had the same proportions. La Quina 9 is left hemimandible with the preserved symphysis. 2 nearly complete skulls: La Quina 5 (adult female, with a brain capacity of 1350 cm³) and La Quina 18 (child aged 7–8 years). The Quina stone industry gave its name to a special facies known as Mousterian of the La Quina type. 57–44 thousand years ago (calibrated).
Skull of the La Quina adult woman H5. Reconstruction of her appearance based on the remains found. Artist: Elisabeth Daynes, Paris. Source: Neanderthal Museum, Mettmann, near Dusseldorf. ©Don Hitchcock
Buca del Tasso
Buca del Tasso, “Badger’s Den”. Northern Italy. Near the village of Metato, left side of Carpigna Creek, district of Camaiore, province of Lucca, Tuscany. Incomplete femur of a 9-year-old child. Classic Neanderthal. Traces of hearths. Mousterian, Levallois. 57–40 thousand years ago.
Buca del Tasso. A human femur, which can be referred to as Homo neanderthalensis.
Melpignano (Nuzzo)
Melpignano (Cava Nuzzo). Southern Italy. Nuzzo quarry, district of Melpignano, province of Lecce, Salento region, Apulia. Right upper premolar of an adolescent or adult. Classic Neanderthal. Late Pleistocene. 57–40 thousand years ago.
Tagliente
Riparo Tagliente. Italy. Village of Stallavena, municipality of Grezzana, province of Verona, in the pre-alpine massif of Monti Lessini. A phalanx and two typical Neanderthal deciduous teeth: the upper right second molar (Tagliente 3) and lower left canine (Tagliente 4). Classic Neanderthal. Late Mousterian. Middle and late Paleolithic. 57–40 thousand years ago.
3D digital model of the enamel-dentin junction in occlusal view of Tagliente 3 and Tagliente 4 (scale = 0.5 cm). B, buccal; D, distal; L, lingual; M, mesial views. ©Fontana et al.
Balauziere
Grotte de la Balauzière. Southeast France. Commune of Vers-Pont-du-Gard, district of Saint-Privat, Vallée du Gardon, department of Languedoc-Roussillon: Gard. The cranial remains belong to Homo sapiens. Out of six isolated teeth, only two can be attributed to Neanderthals (LB XIII is too incomplete to be identified). La Balauzière X: the left upper canine. The occlusal surface is worn, and the tip is worn down. La Balauzière XII: lower left third molar. It is characterized by a small crown and has four cusps. Two vestibular cusps are particularly worn. These teeth are outside the range of variation of late Paleolithic humans. Mousterian of the La Quina type. Classic Neanderthal. 57–40 thousand years ago.
Grotte de la Balauzière. LB X: A—vestibular view, B—lingual view. LB XII: C—vestibular view, D—lingual view, E—occlusal view. ©Gambier
