Researchers find Neanderthal remains in many locations in Europe and Asia, providing a wealth of crucial information about their anatomy, behavior, and culture. The Catalog of Neanderthal Remains lists key Neanderthal man habitation sites with brief descriptions, major findings, and their approximate ages.
This catalog does not include archaeological sites where there are only material traces of the presence of Neanderthals, but no remains of these ancient people were found. Also, not mentioned here are the remains of hominids that exhibit clear features of hybridization between Neanderthals and Sapiens, which are difficult to attribute to the genus Homo neanderthalensis (for example, Oase, “Denny” from the Denisova Cave, Lagar Velho).
Since the exact classification of several specimens has not yet been established, this catalog may include some pre-Neanderthals as well as Neanderthals with sapient features. One hypothesis suggests that throughout the Middle Pleistocene, about 350,000 years ago or slightly earlier, two evolutionary lines existed in Europe: one from late Homo erectus and the other from pre-Neanderthals.
Neanderthal sites and remains, from the most ancient to 130 thousand years
Isernia La Pineta
The Isernia La Pineta site in south-central Italy (province Isernia, region Molise) is located along the left bank of the Cavaliere Stream. This expansive open-air site is notable for its rich lithic industry and abundant faunal remains.
The find from the site is a deciduous human incisor, labeled IS42, likely identified as a deciduous maxillary left lateral incisor (di2). The tooth is well preserved, comprising both the crown and a single, heavily resorbed root measuring 4 mm in length. The root is straight and oval in cross-section. There is wear along the incisal edge. The labial surface shows extensive striations, likely from functional wear.
The find from the site is a deciduous human incisor, labeled IS42, likely identified as a deciduous maxillary left lateral incisor (di2). The tooth is well preserved, comprising both the crown and a single, heavily resorbed root measuring 4 mm in length. The root is straight and oval in cross-section. There is wear along the incisal edge. The labial surface shows extensive striations, likely from functional wear.
The age of the child is estimated to be 5–7 years. It is difficult to determine exactly which species the specimen belongs to. Possible options are Homo heidelbergensis, Homo antecessor, and Homo neanderthalensis. The greatest similarity is observed with Neanderthal dental variability, but most fossil evidence from this time period matches those of Homo heidelbergensis. Acheulean technology. Middle Pleistocene, the end of MIS 15 (Marine Isotope Stage), 583,000–561,000 years ago.
Neanderthal remains: Isernia La Pineta deciduous incisor (IS42). Li: lingual; La: labial; I: incisal; D: distal; M: mesial. Scale bar: 2 mm. ©Peretto et al.
Atapuerca
Atapuerca, Sima de los Huesos (Pit of Bones), Cueva Mayor. Northern Spain. Province of Burgos. Judging by the accumulation of skulls and hundreds of bones, bodies were thrown into the cave whole, below the level of human habitation, as part of a ritual. Researchers found the remains of 29 individuals from the same population: 13 individuals over 20 years old, 16 adolescents aged 12–20 years, and 3 children under the age of 12. Among them:
A fragmentary skull of the specimen “Agamemnon” (skull-4), cranial capacity of 1390 cm³, 430,000 years old;
A complete skull of “Miguelon” (skull-5) of the same age with a cranial capacity of 1125 cm³. The mandible of this skull was found almost intact several years later, not far from the same place;
Also, the specimen “Rui” (skull-6), an immature individual with a cranial capacity of 1220 cm³;
A Neanderthal child aged 10–11, “Benjamina” (skull-14), 530 thousand years ago;
A young individual (skull-17), with two penetrating injuries on the frontal bone above the left eye, the skull composed of 52 fragments found over 20 years of excavations (1990–2010), 430 thousand years ago;
And other specimens. Two hyoid bones were also preserved, indistinguishable from modern ones. These are early Neanderthals (not Homo heidelbergensis), according to DNA analysis.
Visogliano
Visogliano Rock Shelter. Northeastern Italy. Aurisina, near Trieste, Friuli-Venezia Giulia, Duino. Researchers found a fragment of an exceptionally robust right mandible, five teeth, and a root, as well as two fragments of crowns. Recent studies indicate early Neanderthals. Acheulean industry, the Italian Pebble Culture, and Tayacian (a version of the Clactonian industry). 458 thousand years ago.
Fontana Ranuccio
Fontana Ranuccio. Central Italy. Sacco River Valley, Frosinone municipality, 50 km from Rome, Latium. Discoveries include 4 dental remains: one lower left incisor, two lower molars (left and right first molars), and probably the upper left canine. Studies have shown that the teeth exhibit clear Neanderthal characteristics. These fossilized teeth from Italy, among the oldest human remains on the Italian Peninsula, indicate that Neanderthal dental features evolved around 450,000 years ago. Acheulean. 450–430 thousand years ago.
Galeria Pesada (Gruta da Aroeira)
The cave of Galeria Pesada (Gruta da Aroeira). Portugal. Torres Novas, Almonda Karstic System. The designation Galeria Pesada corresponds to the internal channels of the karstic system that currently remain unconnected to the external environment. The designation Gruta da Aroeira is applied to the discovered collapsed entrance, sealed with deposits.
Galeria Pesada
Neanderthal remains consist of large teeth: the left canine of the lower jaw (Pesada 1) and the left third molar of the upper jaw (Pesada 2). The teeth are large and robust. They have a thick enamel layer and a well-developed root. The teeth are associated with stone tools from the Acheulian industry that have elements of proto-Levallois and Micoque technology (Keilmessergruppe). Late Middle Pleistocene, 300–278 thousand years ago.
(A and B)—The Aroeira 1 (Pesada 1) left mandibular canine (A) and the Aroeira 2 (Pesada 2) left maxillary third molar (B) in mesial (Mes), distal (Dist), buccal (Bucc), and lingual view (Ling). Scale bar: 2 cm. (C)—occlusal views of these teeth. Scale in millimeters. ©Daura et al.
Gruta da Aroeira
The majority of the right half of the cranium (excluding the occipital bone), a fragmentary right upper jaw, a preserved part of the nasal floor, and two fragmentary molar roots represent the skull of an adult individual. The cranial volume is 1,100 cm³. Burnt bones suggest the use of fire. Archaic Neanderthals. Acheulean. Middle Pleistocene, 418–390 thousand years ago.
Swanscombe
Swanscombe gravel quarry. United Kingdom. County of Kent, Barnfield Pit. The two parietal and occipital bones are from a young woman. The cranial volume is 1250–1325 cm³. Early Neanderthals or pre-Neanderthals. Associated with the Acheulean and Clactonian industries. 400–300 thousand years ago.
Occipital/foramen magnum of Swanscombe Fossil, ca. 400,000 years old, replica, Museum Tautavel (Perpignan region), France. Young female. In 1935–1936, Alvan T. Marston discovered two fragments, then A. Gibson and J. Wymer discovered the third fragment in 1955. ©Gerbil
Occipital/foramen magnum of Swanscombe Fossil, ca. 400,000 years old, replica, Museum Tautavel (Perpignan region), France. Young female. In 1935–1936, Alvan T. Marston discovered two fragments, then A. Gibson and J. Wymer discovered the third fragment in 1955. ©GerbilOccipital/foramen magnum of Swanscombe Fossil, ca. 400,000 years old, replica, Museum Tautavel (Perpignan region), France. Young female. In 1935–1936, Alvan T. Marston discovered two fragments, then A. Gibson and J. Wymer discovered the third fragment in 1955. ©Gerbil
Qesem
Qesem Cave. Israel. The cave is situated on the western slopes of the Judean Hills, near the town of Kafr Qasim. There are a total of 8 isolated teeth: 6 permanent mandibular teeth and 2 milk teeth. They belong to at least six different individuals. Possibly, ancestors of the Skhul-Qafzeh group. The site exhibits numerous traces of regular fire use. Qesem stone tools belong to two industries: the Amdian and Acheulo-Yabrudian cultural complexes (pre-Mousterian). 380–220 thousand years ago.
Teeth of primitive people discovered in Qesem. Photo: Hershkovitz I. et als. / AJPhA
Multiple views of the lower left canine found in the older (300,000 years+) part of Qesem Cave. ©Hershkovitz et al.
Petralona
Petralona Cave. Northern Greece. Katsika Hill, Chalkidiki province. Remains of at least 15 individuals. The human skull was found in the cave, embedded in the rock 30 cm above the floor. The skull was covered with a calcite crust. The lower jaw was missing. Later, the rest of the skeleton was discovered, which has not yet been described. Petralona 1—age around 30 years, height 155 cm, brain volume 1220 cm³, according to another estimate, 1155 cm³. The cranial vault bones are thick, with a powerful brow ridge. It combines archaic, Neanderthaloid, and even sapient features. Early pre-Neanderthal, archaic Neanderthal. Traces of fire and bone tools. Mousterian. 200 thousand years ago, possibly 350 thousand years ago.
The Petralona skull after the cleaning of its facial area by Prof. J. Melentis. © Museum of Geology-Palaeontology
Orgnac 3 (Mattecarlinque)
Orgnac 3 (also known as Mattecarlinque). Southeast France. This Neanderthal remains site is located halfway between the Ardèche and Cèze rivers in the Ardèche department of the Rhône-Alpes region. There are 7 hominid teeth: the upper right canine of an adult individual; two milk incisors from a teenager around 5 years old; 2 deciduous molars belonging to adolescents about 9 years old; and another 2 deciduous molars for which there is no information. Early Neanderthals. The transition from the late Acheulean industry to the Mousterian and Levallois. 340–298 thousand years ago.
Karain
Karain Cave. Turkey. Near Yagca Village, municipality of Antalya, southern coast. Initially, two isolated Neanderthal teeth were found: the left upper second milk tooth and a broken root of another tooth (possibly a lower incisor). Later, three more teeth (one obviously sapiens) and two Neanderthals (left upper first molar and left lower molar) were discovered. Subsequent findings include the lower jaw, three first phalanges, the third phalanx of the left hand, the radius and cuboid bones, and a fragment of a fibula bone (it is unclear whether it belongs to a human). 250–200 thousand years ago. Then were found two vertebrae and a fragment of the diaphysis of the femur, 340–250 thousand years ago. A partial skull, possibly belonging to a modern human from the late Paleolithic or Mesolithic. Mousterian, Levallois.
Karain. Fusion of distal epiphysis. a) A femoral metaphysis and a distal epiphysis are not totally fused (see under the yellow lines). b) A totally fused intercondylar portion with the shaft in Karain. c) A lateral condyle zone of Karain is not totally fused. d) A posterolateral zone of the Karain metaphysis is not totally fused. ©Chevalier et al.
Pradayrol
Pradayrol Cave. France. Caniac-du-Causse, Lot department. The right upper permanent incisor of an adult individual is known here. Early Homo neanderthalensis. Mousterian. 335–250 thousand years ago.
The Pradayrol upper permanent central incisor: buccal, lingual, and occlusal views. ©Seronie-Vivien, Tillier
Castel di Guido
Open-air site Castel di Guido. Italy. Rome, Lazio. Fossilized hominid bones include 7 different cranial and postcranial elements, from a minimum of 2 to a maximum of 6 adult individuals. Two fragments of femurs (Castel di Guido-1 and 2) from two different adult female individuals. Fragment of the right occipital bone (CdG-3). Right maxilla without teeth (CdG-4). Part of the right parietal bone (CdG-5). Almost the entire right temporal bone (CdG-6) and a fragment of the left temporal vault (CdG-7). Acheulean. Late Middle Pleistocene, 330–320 thousand years ago.
Castel di Guido. (a) Medial view of the CdG-1 right femoral shaft fragment. (b) Posterior view of the CdG-2 left femoral shaft fragment. ©Mariani-Costantini et al.
Views of the Castel di Guido hominid skull fragments: CdG-3 (occipital fragment), (en)—endocranic; CdG-4 (right maxillary fragment), (b)—buccal; (i)—inferior; CdG-5 (right parietal fragment), (en)—endocranic; CdG-6 (right temporal), (ex)—exocranic, (en)—endocranic. Squares: 1 cm. ©Mariani-Costantini et al.
Lezetxiki
Lezetxiki Cave. Northern Spain. City of Arrasate, province of Gipuzkoa, Basque Country. Neanderthal remains are the right humerus of a woman and two isolated teeth (including the third premolar). The individual’s height is 1.63 meters. Found shells that could have served as “ornaments” for Neanderthals. Early Neanderthal. Mousterian or pre-Mousterian. The minimum age of the teeth is around 164 thousand years; the age of the humerus is 310–280 thousand years.
Neanderthal teeth found in the Lezetxiki cave in Arrasate (Gipuzkoa). EFE/Alvaro Arrizabalaga ©Lopez-Onaindia et al.
Pictured is a Homo neanderthalensis humerus found in the Lezetxiki cave in Arrasate (Guipuzcoa). EFE/Alvaro Arrizabalaga
Neanderthal-decorated shell remains found in the Lezetxiki Cave in Arrasate (Gipuzkoa). ©Alvaro Arrizabalaga
Vertesszolles
Vertesszölles. Hungary. Located 15 km west of Budapest, on the terrace of the Ataler River, municipality in Komárom-Esztergom county. Vertesszöllös II is a massive male occipital bone. Vertesszölles I include the teeth of a 7-year-old child (fallen lower canine and second molar). Remains of hearths, Clactonian industry. The cranial volume is 1400 cm³. 300 thousand years ago.
Vergranne
Aven de Vergranne. France. Vergranne, 10 km north of Baumé-les-Dames, Doubs department, Bourgogne-Franche-Comté region, Besançon. The worn deciduous upper left tooth of a child between 7 and 8 years old. Early Neanderthal. Middle Pleistocene (Mindel). 300 thousand years ago.
A deciduous tooth from Vergranne is kept in the Montbeliard Museum. Photo: DR
Ciota Ciara
Ciota Ciara (Monte Fenera) cave. Northern Italy. Ciota Ciara, Ciutarun, and Belvedere Caves on the slopes of Mount Fenera, near the town of Borgosesia, province of Vercelli, Piedmont. An occipital bone and the second lower incisor with an incomplete root, indicating affiliation with a young individual, were initially discovered. Subsequently, three more teeth were found: Fenera 2, the second right lower molar with two complete roots; Fenera 3, the right upper premolar; and Fenera 4, the right lower canine (Ciutarun cave). Mousterian. 300 thousand years ago.
Occipital bone (part of the back of the skull) and a second lower incisor, probably belonging to a young adult.
Ciota Ciara. These are two teeth, a canine (Fenera 4) and a lower molar tooth (Fenera 2), identified at the same stratigraphic levels where human remains were found in 2019.
Bad Cannstatt
Bad Cannstatt. Germany. “Haas” Quarry, near the Neckar River, not far from Stuttgart, Baden-Württemberg. A hunting site near a lake with a hot spring. Two tooth fragments were initially considered the incisors of a red deer. However, the first was later identified as a fragment of the crown of a pathological lower left human canine (BC 1). The second fragment represents the root of a non-pathological upper molar (BC 2). These fragments resemble those from the Steinheim skull. Pre-Neanderthal or early Neanderthal. There are remnants of wooden tools made from maple and an elephant shin hoe. Microlithic chips and stone tools. Middle Pleistocene, 295 thousand years ago.
Monte delle Gioie
Monte delle Gioie. Central Italy. Right bank of the Aniene River, northeast suburb of Rome, Latium. Neanderthal remains: fragments of human bones. The site was leveled to make way for the railway yard of Portonaccio station. Mousterian, Levallois (Protopontinian type). 295–290 thousand years ago.
Sedia del Diavolo
Open-air site Sedia del Diavolo. Central Italy. Left bank of the Aniene River, northeast suburb of Rome (Monte Mario area of Rome), Via Salaria, Latium. The Neanderthal remains were found in a layer of gravel and sand. There are femur diaphysis and the second right metatarsal of an adult Neanderthal. Today, Sedia del Diavolo is the square of Elio Callisto. Archaic Mousterian (Protopontinian type). 295–285 thousand years ago.
Velika Balanica
Velika Balanica (Great Balanica). Southeastern Serbia. Village of Sićevo, city of Niš. Two found teeth belong to an adult individual: the permanent upper third molar (BH-2) and upper central incisor (BH-5). A fragment of the right upper jaw (BH-4) with one intact first right molar belongs to a young individual. Milk upper fourth premolar (BH-3) is a 6-year-old child. Taurodontism. Charentian industry, some elements are similar to those of the Acheulo-Yabrudian. Remains of hearths. Middle Pleistocene (Chibanian), 295–285 thousand years ago.
Remains of Neanderthal teeth from Velika Balanica. ©Mirjana Roksandic
Neanderthal remains: teeth from Velika Balanica. ©profimedia
Ponte Mammolo
Open-air site Ponte Mammolo. Italy. Right bank of the Aniene River, Rome. The diaphysis of the femur of an adult male. Proximal and distal metaphyses are absent. The third trochanter is discernible. Pre-Mousterian and Levallois. Archaic Neanderthal. Middle Pleistocene, 295–245 thousand years ago.
Ponte Mammolo femur shaft fragment: a) anterior view; b) lateral view; c) posterior views d) medial view; e) middle shaft view; f) subtrochanteric shaft view. ©Rubini et al.
Reconstruction of the insertion muscular area in the Ponte Mammolo fossil femur. ©Rubini et al.
Casal de’Pazzi
Open-air site Casal de’Pazzi (also Rebibbia — C. de’Pazzi). Italy. Aniene River Valley, municipality of Rome. The Neanderthal remains found at this site contain a fragment of the right parietal bone of an adult (Casal de’ Pazzi 1) and other elements, totaling the remains of 6 individuals. Early Neanderthal. Acheulean–Pre-Mousterian. Late Middle Pleistocene. Approximately 295–245 thousand years ago.
Parietal bone of early Neanderthal from Casal de’Pazzi.
Reconstruction of the landscape along the lower Aniene valley about 200,000 years ago.
Cova Negra
Cova Negra (Cueva Negra del Estrecho, Cova Negra de Xàtiva), karstic cave. Southeastern Spain. Left bank of the Albaida River, Murcia region, Valencia province. Very thick parietal bone, other bones (one metacarpal, one metatarsal, two femurs, a radius, a fibula), and teeth of seven individuals, two of whom were adults and one was an adolescent, and four were children under 10 years old. The outer surfaces of two recovered cranial remains have marks left by a cutting tool. This may be related to cannibalism. Early Neanderthals. Mousterian, Paracharentian. According to 2019 data, the age is 273–261 thousand years.
The Cova Negra Parietal in external (left) and endocranial (right) views. The sagittal suture is visible along the superior margin of the specimen. ©Arsuaga et al.
Cova Negra Femur I (CN 42168 and CN 42169) and Cova Negra radius (CN 42165). ©Arsuaga et al.
Saccopastore
Saccopastore Quarry. Italy. Aniene River Valley, left bank, Rome, Latium. Two adult individuals, one male and one female, totaling at least six individuals. Initially, an almost entirely preserved Neanderthal adult female skull with teeth was found (Saccopastore 1). Subsequently, a skull with teeth but without the upper part from a 30-year-old adult male was discovered (Saccopastore 2). Early atypical “Ehringsdorf” group. Brain volumes of 1280 and 1300 cm³. Mousterian. Approximately 250 thousand years ago.
Reconstruction of the landscape of the Saccopastore site (about 250,000 years ago).
Reilingen
Reilingen, gravel quarry. Germany. Near Heidelberg, Baden-Württemberg. In the gravel deposits of the Rhine, researchers found: the back part of the skull, consisting of two complete parietal bones; a major portion of the right temporal bone; and 70% of the occipital bone from an adult individual. It exhibits a bilaterally convex occipital torus. None of the sutures are completely closed. The cranial volume is 1430 cm³. An archaic Neanderthal, resembling Swanscombe and Atapuerca, possibly a pre-Neanderthal. Approximately 250 thousand years ago.
Reilingen skull, left lateral, and posterior views. ©Staatliches Museum fur Naturkunde
Reilingen skull, right lateral, and dorsal views. Temporal bone: inferior view. The top is anterior. Photo: Fred Spoor ©Staatliches Museum fur Naturkunde
Balzi Rossi (Grimaldi)
Grotte dei Balzi Rossi (Grimaldi Caves), “Red Stones”. Italy. Municipality of Ventimiglia, Liguria. The complex comprises 15 caves and grottoes situated right on the Franco-Italian border: seven caves, three shelters, and two open-air sites. In these caves overlooking the sea on Mount Bellinda in Grimaldi, near the village of Ventimiglia, stone tools dating back to the Mousterian period were discovered. A few fossil remains found in these caves belong to pre-Neanderthals or early Middle Pleistocene Neanderthals.
Grotte du Prince
Grotte du Prince (Grotta del Principe, Grimaldi) (named after Prince Albert I of Monaco). Balzi Rossi Cave Complex. Pelvic bone of an adult woman, weighing 75 kg. Early Neanderthal. Acheulean, Mousterian. 250–200 thousand years ago.
Grotte du Prince. The pelvic bone of a woman.
Pontnewydd
Pontnewydd Cave (Bontnewydd), “New Bridge”. United Kingdom. Near the town of Rhyl, close to St. Asaph, Denbighshire, North Wales. Bones and 17 taurodont teeth from five individuals—an adult, two adolescents, and two older children, including teeth and part of the jaw of a Neanderthal boy approximately 11 years old. Pontnewydd 2 consists of fragments of the upper and lower jaws of an 8–9-year-old child. Archaic Neanderthals. Late Acheulean, Levallois. 250–200 thousand years ago.
Pontnewydd Cave. Left: The fragment of Neanderthal bone, something rarely found in Britain, is a piece of upper jaw from an eight-year-old boy, with the upper first and second molars still in place. Right: Early Neanderthal tooth (bottom) and X-ray (top). The X-rays show the enlarged pulp cavity, which has helped archaeologists identify the Pontnewydd teeth as belonging to Neanderthals. ©Amgueddfa Cymru – National Museum Wales.
Neanderthal premolar tooth found in Pontnewydd Cave, north-east Wales. Premolar tooth with a complete crown and mostly complete root; just the basal tip is missing. ©Amgueddfa Cymru
Poggio
Grotta del Poggio. Italy. The cave and shelter are located directly east of the town of Marina di Camerota, Salerno, Campania. Grotta del Poggio 1 is a left upper molar tooth of an adult individual. Grotta del Poggio 2 is the talus of a female individual. Early Neanderthal. Mousterian, Tayacian, and Clactonian. Middle Pleistocene, 240–190 thousand years ago.
Left M1 referred to layer 6 of Poggio (the D-tooth). The white rectangles mark the Carabelli cusp of the D-tooth. ©Benazzi
Tourville
Tourville-la-Rivière. Northern France. Seine-Maritime Department, Normandy. Three fossilized bones of the left arm from one Neanderthal individual (humerus, ulna, and radius), Tourville 1. They may have belonged to a late adolescent or adult male. Mousterian, Levallois. 236–183 thousand years ago.
Neanderthal remains. The Tourville left upper limb remains. 1. humerus; 2. ulna; 3. radius. For all the bones: A: anterior view; M: medial view; P: posterior view; L: lateral view. ©Denis Gliksman
Taubach
Taubach. Southern Germany. In the Ilm River Valley, near the city of Weimar, Thuringia. Neanderthal remains include two molars: the first lower left permanent molar from an adult individual or adolescent aged 12–14 years; and a large first lower left deciduous molar from a 9-year-old child. The surface is more worn than usual for modern deciduous teeth of this type. Hearth remains. Mousterian. Atypical Neanderthals. 123 thousand years ago (Eemian interglacial); according to other data, 234–230 thousand years ago.
The first molar of the human mandible. Found in Deluvium, near Taubach. Inner and outer sides of the tooth. (From Nehring). The grinding surface of the first right lower molar tooth from Taubach. The letters denote several small prominences called cusps.
Ehringsdorf
Ehringsdorf Quarry, open location. Germany. Suburb of Weimar, Ilm Valley, Thuringia. Found 35 fragments of at least nine early Neanderthals. Ehringsdorf H (IX)—the skull was discovered with the accompanying remains of a female skeleton aged 20–30 years. Brain volume: 1450 cm³. Ehringsdorf G (VII)—the left half of the lower jaw of a child aged 10.5–14 years. Ehringsdorf F (VI)—a primitive mandible of an adult individual. Ehringsdorf E (V)—part of a massive, strongly curved left femur. The teeth do not exhibit taurodontism. Remains of hearths. Cannibalism. Atypical Neanderthals. Late Acheulean, early Mousterian. 230–205 thousand years ago.
Payre
Payre. Southeastern France. The site is located at the confluence of the Rhone and Payre rivers, in the Ardèche department. 8 teeth, a fragment of the left parietal bone, and half of the lower jaw were discovered, totaling five individuals, both adults and children. 4 teeth found nearby may belong to a child around 7 years old. 4 lower teeth, upper tooth, and fragment of the parietal bone likely represent two or three individuals. A robust upper molar, very worn, could represent one adult individual. This worn molar cannot belong to the same individual as the other teeth. Two children showed signs of lead exposure. Middle to early Late Pleistocene, 230–200 thousand years ago.
Photos of the Neanderthal tooth from Payre. Scan areas to determine age are indicated.
Some of the human remains at Payre (with number and level). ©Moncel, Condemi
Mollet
Cova de Mollet (Mollet Cave). Northern Spain. Reclau Caves, municipality of Serinyà, Pla de l’Estany, approximately 5 km north of the city of Banyoles and 30 km from the city of Girona, Northeastern Iberian Peninsula, Catalonia. Isolated archaic upper right first molar of a child (Mollet I). Occlusal wear is absent. The morphological and morphometric characteristics of the tooth correspond to those of Homo neanderthalensis. The site also served as a den for hyenas, but predators did not digest the human tooth. This is the oldest human remains found in Catalonia. Early Neanderthal. 215,000 years.
Human first upper right molar (M1) from Layer 5 of Mollet Cave. Left: buccal view. Scale: 10 mm. Universitat de Girona. ©Maroto et al.
Montmaurin
Montmaurin Cave. Southwest France. North of the village of Saint Gaudens, Haute-Garonne department.
La Niche
La Niche Karst Pipe. The remains of a female mandible with 6 molars, a left tibia, a lower tooth, and a vertebra were found at the site. The jaw is missing a bony chin. Late Acheulean. Approximately 200–150 thousand years ago.
Upper view of the Neanderthal mandible Grotte de la Niche. Different morphological aspects. a) Internal aspect of the symphysis, showing the fossa genioglossa; b) Left view of the mandible. Note the receding wall of the symphysis, the position of the mental foramen, the flat surface of the corpus and ramus, and the regular gonion profile.
Coupe-Gorge
Coupe-Gorge cave. 8 Neanderthal remains, including a fragment of the front part of the lower jaw (symphysis) of a growing child with two incisor buds and an isolated tooth, date to 125 thousand years ago. The right upper jaw, with two premolars in place and two isolated teeth, was also found. In 2022, a human shoulder bone was found during excavations, and its attribution is yet to be established. Late Mousterian industry. The end of the Middle Pleistocene, 200 thousand years ago.
Montmaurin (Coupe-Gorge)-3. Fragment of the symphysis of the mandible of a child 4.5–5 years old in different views: A, vestibular view with chin elements; B, posterior view with 1) the planum, 2) the transverse torus, 3) the genian fossa, 4) the imprint of the missing permanent incisor germ, 5) the genioglossal apophyses, and 6) the geniohyoidal apophyses. ©Granat, Peyre
The maxillary part of the Coup-Gorge. 1—external view with the peri-nasal swelling. The canine is in place. 2—internal side. 3—Coupe-Gorge maxilla positioned on a current skull. It integrates very well. On the actual skull, the foot of the ascending ramus is concave. ©Granat, Peyre
Fontechevade
Fontechevade Cave. Western France. Department of Charente, Nouvelle-Aquitaine. Two fragments of skulls. Fontechevade 2 (or Homo II) is a large fragment of an archaic cranial vault (the upper part of the frontal bone and parts of the left and right parietal bones). The bones are quite thick. The cranial volume is 1350 cm3. Initially, it was mistakenly believed that both findings lacked developed brow ridges and therefore belonged not to Neanderthals but to some presapiens. However, one of the fossils (Homo I) belongs to later deposits (OIS 3) and evidently represents Homo sapiens, while the other has not preserved the area of brow ridges at all, but other characteristics correspond to Neanderthals. Tayacian industry. 200,000 years ago.
Fontechevade II: fragment of a cranial vault. ©Paleo Core and OSA
Biache-Saint-Vaast
River terrace of Biache-Saint-Vaast. Northern France. Department of Pas-de-Calais. Two partial skulls, resembling early and classic Neanderthals. Biache-Saint-Vaast 1—occipital bone, fragment of the upper jaw, and isolated teeth. Biache-Saint-Vaast 2—fragments of the frontal, parietal, and temporal bones of a female individual. Cranial volume: 1200 cm3. Mousterian, Levallois. 196–159 thousand years ago.
Aubesier
Rock shelter Bau de l’Aubesier. Southeastern France. Gorges de la Nesque, near the town of Monieux, department of Vaucluse. Eight isolated deciduous and permanent teeth. Two isolated teeth from adult individuals (Aubesier 4 and 10). Aubesier 11 is the mandible of an adult individual with worn teeth. The chin is beveled. All the alveoli were in varying degrees of healing, meaning that the teeth were lost during life. Aubesier 12 is an upper jaw molar with large dental caries from the end of the middle to the beginning of the Late Pleistocene. Neanderthal with archaic features. Mousterian and Levalloisian facies. 191–169 thousand years ago.
Middle Pleistocene human dental remains from the Bau de l’Aubesier. From left to right: Aubesier 4 upper incisor lingual, and distal; Aubesier 10 upper molar mesial, and distal. Scale in mm. ©Lebel, Trinkaus
Aubesier 11 mandible in right lateral and superior views. Scale in cm. ©Lebel, Trinkaus
Behistun
Behistun site (also known as Bisetoun, Behistoun, or Bisitun). Northwest Iran. Zagros Mountains, Harsin County, Kermanshah Province. A fossilized specimen is the right radius proximal diaphysis of an adult individual, with breaks on each end at an angle. Mousterian with Levallois components. Middle Paleolithic. 190–60 thousand years ago.
Four views of the Bisitun right radial diaphysis. From left to right, anterior (A), posterior (P), medial (M), and lateral (L). Note the faunal damage marks on the posterior surface and the prominent pronator teres tuberosity on the lateral side. © Trinkaus
Kozarnika
Kozarnika Cave, known as the “Goat Shed”. Northwest Bulgaria. Approximately 3 km from the village of Oreshets in the Dimovo district. The radius of a Neanderthal child is about 6 months old. 183,000–128,000 years ago.
Kozarnika. The radius of a Neanderthal child.
Altamura
Altamura (Lamalunga Cave). Southern Italy. Karst region of La Murgia, near the city of Altamura, Province of Bari, Apulia. A very complete and well-preserved skeleton of an adult male individual, known as “Altamura Man”. The cranial volume is measured at 1190 cm³. DNA analysis from the scapula confirmed that the individual was a Neanderthal. This specimen represents an archaic Neanderthal and dates back to the period between 172,000 and 128,000 years ago.
Altamura Man, surrounded by limestone deposits (a thick layer of calcite). ©di Nicola de Paulis
Altamura skeleton in situ and three-dimensional topographic distribution of some elements of the human skeleton. Top view.
Apidima
Apidima Cave. Southern Greece. Situated in Lakonia on the Mani Peninsula. Researchers found the remains of 6 or 8 individuals, primarily fragments of skulls. In the excavation, a male skull, referred to as Apidima I, was discovered. Apidima II is a robust female skull with a sloping forehead and prominent brow ridges. The brain volume of the latter measures 1290–1454 cm³. It shares similarities with skulls from sites like Sima de los Huesos, Swanscombe, Biache St. Vaast, and Lazaret.
Some researchers consider Apidima 1 as an older fossil (210,000 years ago), representing an archaic Homo sapiens. However, both skulls were found positioned vertically against the cave wall, in the same sedimentary layer, with a 15 cm distance between them, facing in opposite directions. These remains exhibit similar fossilization conditions. The absence of other human remains, stone tools, or faunal remains suggests an anthropogenic ritual associated with death. Both individuals were adults, around 30 years old, but Apidima 1 appears slightly younger than Apidima 2. These early Neanderthals lived around 170,000 years ago.
Apidima 1. The skull fragment and its reconstruction from side and posterior views. ©Harvati et al.
Apidima 2 frontal view (©Harvati et al.) and lateral view (©Brauer et al.).
Ochtendung
Ochtendung (Wannen). Germany. The site is located near the city of Koblenz in the Rhineland-Palatinate region. Neanderthal remains consist of three elements of the cranial vault (the frontal bone and anterior parts of the parietal bones). These remains belong to a male individual aged between 30 and 45 years. The antemortem modifications on the skullcap indicate that it was used as an artifact by other individuals. It was discovered along with Pleistocene mammal bones and stone tools that represent traces of Neanderthal groups who used this volcanic mountain range for hunting. This strong and thick skullcap is an example of the transition of Homo heidelbergensis to the classic Neanderthal. Early or typical Neanderthal. 170,000 to 160,000 years ago.
Skullcap of an early Neanderthal from Ochtendung near Koblenz, Germany. Found in 1997 at the hill Schweinskopf/Karmelenberga near Ochtendung/Koblenz in an industrial Lava mine by the archeologist Axel von Berg. According to the stratigraphy, the find comes from the glacial phase of 170,000 years ago, at a time from which only very few hominid fossils in Europe are known.
Abri Suard
Abri Suard (Suard Cave). Southwestern France. The cave located in the commune of La Chaise-de-Vouthon, in the Charente department, has yielded 52 findings. These include fragmented cranial bones (frontal, left parietal, and occipital bones), fragments of two mandibles, a phalanx, and teeth from several individuals, both adults and children. One of the mandibles, as well as the parietal and occipital bones, belongs to a child aged 4–5 years. Classic Neanderthals. The archaeological industry is associated with Mousterian and Châtelperronian artifacts. 163,000 years ago.
Cranium Abri Suard S16
Virtual reconstructions of a Neanderthal mandibular left first molar crown (Abri Suard S5) in standard views. The inset shows the enamel cap and underlying dentin core separately.
Lazaret
Le Lazaret Cave (Grotte du Lazaret). Southern France. Cave situated near a suburb of Nice, Alpes-Maritimes, Provence. Neanderthal remains are cranial fragments, including the right parietal bone, several teeth from a nine-year-old adolescent, and seven other bones. Later discoveries included the central fragment of the left humerus (Laz 14) and a naturally shed lower molar from a child aged 10 to 12 years. These two remains correspond to two individuals, an adult and a child. They were found in a Late Acheulean layer with two identified hearths. Archaic Neanderthals. 160,000–150,000 years ago.
Lazaret-24. Human frontal bone from the Lazaret cave, Nice, UA28. Discovered in August 2011. ©Miniwark
Diaphyseal fragment of the left humerus, Laz 14. a—anterior view, b—internal view, c—posterior view, d—lateral view. ©Cauche
The environment of Lazaret Cave during cold periods. ©Departement des Alpes-Maritime
Qal-e Kord
Qal-e Kord Cave (Ghale Kord). West-Central Iran. Zagros Mountains, near the village of Hesar Valiasr, Avaj County, northwestern province of Qazvin. The site has yielded a fossilized Neanderthal child’s tooth. This site represents the oldest human settlement discovered in Iran, dating back to approximately 155,000 years ago.
Neanderthal child’s tooth
Bourgeois-Delaunay
Bourgeois-Delaunay cavern. Southwestern France. Commune of La Chaise-de-Vouthon, Charente department. The excavation found 23 bones, including fragments of skulls, jaw pieces, and other bones and teeth, from multiple individuals, both adults and children. BD1 is the mandible of an adult individual, complete with all teeth. BD22 is the fragmented skull of an adult individual. Classic Neanderthals. 151,000–126,000 years ago.
Bourgeois-Delaunay mandible BD1. Microtomographic-based 3D rendering in semi-transparency of the specimen in anterior (A) and slightly obliquely-oriented lateral left (B) views. Photograph in superior view (C) and in superior view from the left side (D), showing the reconstructed portion of the inferoanterior aspect of the Neanderthal mandible. ©Zanolli et al.
Moros (Gabasa)
Cueva de los Moros de Gabasa. Spain. The cave is located in Gabasa, Peralta de Calasanz commune, Huesca province, Catalonia. Neanderthal remains are three teeth (an upper premolar and two lower molars) from an 11–12-year-old adolescent, an adult male, and an elderly individual. Additional findings include the first right metatarsal bone, the first phalanx of the left foot, and an incomplete clavicle. These Neanderthal bones are part of a burial site that scavengers disturbed. Studies of tooth enamel suggest a predominantly meat-based Neanderthal diet. Moustier of the Acheulean tradition. 150,000 years ago.
Carihuela (Pinar)
Carihuela (Cariguela) cave, Pinar. Southern Spain. Municipality of Pinar, province of Granada, Andalusia. Two fragments of the left parietal bones from two adult Neanderthals (Pinar 1 and Pinar 2) were discovered. Additionally, an almost complete frontal bone from a 6-year-old child (Pinar 3) was found, along with 6 upper left teeth (including 2 milk teeth) from a non-adult individual aged around 8–9 years (Pinar 7). All teeth exhibit large Neanderthal morphologies. The findings suggest evidence of cannibalism. These are classic Neanderthals of the l’Hortus type. The assemblage is associated with the Mousterian industry and Levallois technology. Dating from 146,000 to 117,000 years ago. Earlier dating, proposed by some sources (50,000–46,000 years ago), is not confirmed, according to Wood et al. (2013).
The frontal bone of a neanderthal child from the Cave of La Carihuela del Pinar; it is dated 146,000 to 117,000 years ago.
On the left is the upper deciduous canine (buccal and lingual projection). In the center is the second deciduous molar (occlusal projection). On the right is a buccal view of the upper left permanent central incisor: linear enamel hypoplasia. Scale = approximately 1 cm. ©Manuel et al.
On the left are the left permanent central and lateral incisors (lingual side). On the right is the left first premolar (occlusal view). Scale = 1cm approx. ©Manuel et al.
Molare
Grotta Il Molare rock shelter (Molare di Scario, Riparo del Molare, Grotta della Molara). Italy. Scario, San Giovanni, province of Salerno, Campania. Molare 1 is a very archaic lower jaw with 4 milk molars from a child aged 3–4 years. The individual represents a classic Neanderthal. The site also revealed remnants of seven hearths. Typical Mousterian technological tradition. 140,000 years.
Juvenile Neanderthal mandible from Molare (Italy), with the lower-left second deciduous molar virtually extracted and dissected using X-ray microtomography. The transparency of the enamel shows the dentin and the pulp chamber.
Sirogne (Rocamadour)
Grotte Sirogne (Rocamadour). France. Grotte situated on the right bank of the Alzou River near the locality of Merle in the commune of Rocamadour (Lot), Quercy. Researchers found approximately 84 human remains, representing at least 10 Neanderthal individuals. The collection includes half of the mandible of a juvenile individual, elements from an adult individual, more than 80 isolated teeth, and postcranial skeletal elements. The site dates to the end of the Middle Pleistocene, estimated to be between 140,000 and 125,000 years ago.
Half of the mandible from Grotte Sirogne (Rocamadour).
Nesher Ramla
Nesher Ramla. Israel. This open-air karstic depression located southeast of Jerusalem, near the city of Ramallah. 5 fragments of cranial vault, including the right parietal part (NR-1), and an almost complete mandible (NR-2) exhibiting typical Neanderthal characteristics. The archaeological horizon at Nesher Ramla is associated with the Levallois technique, and the site dates to the Middle Pleistocene, approximately 140,000 to 120,000 years ago.
Fossil hominin material from Nesher Ramla unit VI. Parts of a jaw (left) and a braincase (right), found at Israel’s Nesher Ramla site, represent an ancient hominid population that contributed to the evolution of European Neanderthals and possibly some ancient Homo groups in East Asia, researchers say. ©Levin et al.
Tabun
Mugharet et-Tabun (Tabun Cave). Israel. The deep rock shelter of Tabun is situated on the southwestern slope of Mount Carmel. Found here are Neanderthal remains, including a partial skeleton, mandible, some premolars, and isolated limb bones from several individuals. Tabun I (C1) represents the skeleton of a woman younger than 30 years, measuring 151–154 cm in height and weighing 63 kg. Tabun II is a large, extremely robust lower jaw of a man aged 30–35 years. The jaw exhibits typical Neanderthal features, such as a broad and somewhat arcuate front.
As is typical of Neanderthals, the front of the jaw is very broad and somewhat arcuate from side to side. As in some Neanderthals, the symphyseal region is very deep and is elevated slightly along its inferior margin. There are also inferior marginal tubercles beneath the very posteriorly placed mental foramina. These findings suggest that Neanderthals and early modern humans alternately inhabited this region. Classical Neanderthals. Moustier with the Levallois technique. 140,000–112,000 years ago (calibrated).
Female Homo neanderthalensis skull discovered at Tabun Cave at Mount Carmel in Israel. Known as Tabun 1, this Neanderthal specimen is around 130,000 years old.
Mandible of Tabun II. Bar = 1 cm. ©Schwartz et al.
