Catalog of Neanderthal Remains Sites, 2 of 4

Neanderthal Remains Sites research is bringing more and more knowledge about past eras and the life of Homo neanderthalensis during the Ice Ages. We learn about the places where stone tools were made, the hunting of large herbivores, and shelters from dangerous predators.

The number of known Neanderthal remains sites is constantly increasing. Sometimes, new habitation sites and hunting grounds of Neanderthal man are discovered. Researchers explore already known findings using new scientific methods, including molecular biology methods, which often leads to their reclassification. Excavations of old sites continue, bringing forth new discoveries.

Neanderthal remains sites, ranging in age from 130 thousand years to 75 thousand years

Bolomor

Bolomor Cave. Spain. Near Tavernes de la Valldigna, La Safor, Valencia province. Neanderthal remains were found in this cave. A central part of the fibula, and an almost complete parietal bone (one piece and four more fragments found later) of an adult Homo neanderthalensis. Two isolated teeth: a lower molar from a 3–5-year-old child (HCB-02) and an upper canine from a 5–6-year-old child (HCB-05). Later, a part of the left maxilla of a young individual was found. Early Neanderthals. 15 hearths were found. Acheulean and Mousterian. 130,000 years ago.

Parietal and other bones of Homo neanderthalensis from the Cova del Bolomor. Prehistoric Museum of Valencia.

Bolomor. Two views of the fragment of the maxilla and its location in the hominid skull.

Bolomor. Left: Four fragments of the parietal bone of Homo neanderthalensis. Right: Lower left first molar (m1) of a child of ca. 5 years of Homo neanderthalensis. It has five cusps: three in the buccal part and two in the lingual part.

Combe Grenal

Combe Grenal Cave. Southwest France. Near Domme, Dordogne. At level 25, 27 cranial and postcranial specimens from 8 individuals were found, including two children, two or three adolescents, and several adults of various ages. Among these discoveries are a juvenile mandible and deciduous molar, as well as an adult cranium with a mandible, two molars, a humerus, three phalanges, and some hand bones. The age of the findings is approximately 75–65 thousand years. In layer 60, a part of an incisor, belonging to a child around 3 years old (Combe-Grenal Hominid 31), was found. The age of this fossil is estimated at 130,000 years. Traces of cannibalism. Hearths. Mousterian.

Rozhok

Rozhok I. Russia. Northeastern Azov Region. A permanent second left molar, belonging to an adult Neanderthal, was found. The crown of the molar is well-preserved, but the roots are absent. Among the Neanderthal remains sites, this location is considered a seasonal hunting camp. Ancient hearths. Micoquian industry. Middle Paleolithic, 130–115 thousand years.

Molar found on the Rozhok-1 site. Alisa Zubova et al. / Journal of Human Evolution, 2022

Krapina

Krapina Cave, also known as Hušnjakovo Brdo (Hušnjak Hill). Northern Croatia. Near Zagreb. Several dozen individuals of varying gender, ranging in age from 2 to 40 years, with over a thousand remains. Researchers found five informative partial skulls in Krapina, as well as around eleven upper and twelve lower jaws (excluding sixteen rami) of varying completeness. The skull volumes range from 1326 to 1359 cm³. The partial skull of a teenager aged 6–8 years (Kr 1, “A” skull) is well known. Kr 3 (“C” skull) is the most complete skull found at Krapina. This individual died as a teenager. There is evidence of the use of feathers as ornamentation. The question of cannibalism remains debatable. Mousterian industry. Typical Neanderthals. Deposits accumulated over about 20 thousand years; the age of the findings estimated to be 130–110 thousand years.

Krapina 59. Left (anterior view) and right (left lateral view). Hravatsku Prirodoslovni Muzej, Zagreb. Bar =1 cm. ©Schwartz et al.

Sima de las Palomas

Sima de las Palomas del Cabezo Gordo, Rock-Dove Hole. Southeastern Spain. Municipality of Torre-Pacheco, Murcia region, Valencia province. Initially, fused upper and lower Neanderthal jaws were found (SP1). A total of 232 remains from at least 15 individuals have been discovered (8 mandibles and other bones). This includes an adult individual (SP92) and the skeleton of a young woman under 20 years old (SP96, nicknamed “Paloma”), possibly buried with a child aged 5 or 6 years (SP97). Their bodies were covered with stones (possibly intentionally). “Paloma” had a height of only 1.5 meters. Also found were 4 Neanderthal teeth and the ascending branch of the mandible. Mousterian-Levallois. 130–90 thousand years.

Sima de las Palomas del Cabezo Gordo. Left: right lateral view of the Palomas 96 crushed skull. The squamous frontal bone and the parietal bone are evident in pieces. Scale bar: 5 cm.
Right: Neanderthal child skull SP97 (“Paloma’s child”). ©Walker et al.

Sima de las Palomas. Neanderthal post-cranial remains of adult female skeleton SP96, “Paloma”. ©Walker et al.

Malarnaud

Malarnaud Cave. Southwest France. Near the town of Montseron, Ariège department. Partial mandible of a child aged 12–14 years, an atlas, and an isolated tooth. The third molars of the Malarnaud specimen are partially formed, which is appropriate for its age. Classical Neanderthal. 129–122 thousand years.

Malarnaud-1. Mandible of Neanderthal. ©Heim, Granat.
Color insert in the image on the right: occlusal view of the alveoli of the anterior mandible (white bar is 1 cm). ©Lacy.

Quinzano

Quinzano Quarry. Italy. Cava Vecchia near Ca’ Rotta di Quinzano, Verona, Veneto. Occipital bone. Possibly pre-Neanderthal. Archaic Mousterian and La Quina-type Mousterian. 128–115 thousand years.

Scladina

Scladina (Sclayn Cave). Belgium. Village of Sclayn, Andenne municipality, Namur province, Wallonia. 19 remains of young Neanderthal. Almost complete mandible, fragment of the right maxilla, and 16 deciduous teeth of the same child aged 9–12 years (“Scladina child”, probably a girl). There is no retromolar diastema (absent in most Neanderthal children). Also included are one first metatarsal bone, three permanent upper molars, and one tooth from another Neanderthal infant, along with other elements. Mousterian. 127–120 thousand years.

Left: 2 Image of the Scladina Neanderthal first molar. Note a small portion of the root was removed prior to sectioning for ancient DNA analysis. (Adapted from Smith et al. 2014)
Right: This is a tooth that laboratory tests have already established that it belonged to a Neanderthal man at least thirty years old.

Hyene

Grotte de l’Hyène (“Hyena Cave”). France. Arcy-sur-Cure, Yonne, Burgundy. Part of a very robust adult mandible, the maxilla of an elderly individual, part of a child’s forehead, six teeth from adolescents and young adults, a fragment of the fibula, and a metacarpal bone. The chin of a modern type. Relatively small cranial capacity, a very long face base, and very large teeth. Two Neanderthals were named “Augustin” and “Augustine”. Mousterian. 125–75 thousand years.

Hohlenstein-Stadel

Hohlenstein-Stadel Cave. Southern Germany. Lone Valley, Swabian Jura, Baden-Württemberg. Almost complete right femur with traces of carnivore bites. Both epiphyseal ends are missing in this specimen, but the diaphysis is well-preserved. A man is 1.6 meters tall. Represents an isolated population of Neanderthals that separated from the main lineage around 270,000 years ago. Mousterian. 124–120 thousand years.

Femur of a male Neanderthal from Hohlenstein Stadel Cave: (1) different views of the fossil and (2) a detailed image of the proximal zone. ©Camaros

Naulette

La Naulette Cave. Belgium. Village of Chaleux, commune of Houyet, province of Namur, Wallonia. Left half of the mandible, ulna, third metacarpal bone, lower canine, and other teeth and fragments. A young woman, aged 17. Neanderthal with some archaic features. Mousterian. 124–119 thousand years.

La Naulette: mandible from different views, ulna, and third metacarpal. ©Schwartz, Tattersall

Artenac

Artenac. Central-western France. Commune of Saint-Mary, department of Charente. Two fragmented skull bones: the maxilla (Artenac 1) and the frontal bone (Artenac 2), evidently belonging to different individuals. Mousterian industry of the Ferrassie type. 124–119 thousand years.

Left: Artenac 1. Right maxillary fragment: a. occlusal; b. superior; c. lateral; d. frontal. Right: Artenac 2. Frontal bone: a. superior; b. lateral; c. inferior; d. frontal. ©Mann et al.

Cueva del Angel

Cueva del Ángel. Spain. Aras Mountains (southern slope of the Sierra de Araceli), near the town of Lucena, province of Cordoba, Andalusia. A 15 cm long diaphysis of a female femur, evidently belonging to a pre-Neanderthal. Also, the right lower third molar. Based solely on the nearby Acheulean industry, its age is estimated at 300,000 years, which seems doubtful. Lower third molar of a Neanderthal. Remains of hearths, charred bones. Late Acheulean without Levallois. Middle Pleistocene. 121–104 thousand years.

Human right lower third molar from Cueva del Angel: root fragment (cervical end) embedded in the sediment block (left) and crown (right). ©Bermudez et al.

Masloukh

El Masloukh Cave. Northern Lebanon. South of Madfuna, along the Mediterranean coast. Researchers found one tooth—the second upper molar. Acheulean Yabrudian. Riss-Würm. 120 thousand years.

Moula-Guercy

Baume Moula-Guercy, Moula-Guercy Cave. Southeastern France. 80 meters above the modern Rhone River, North of Marseille, near the town of Valence, Soyons, Ardèche department. 119 human remains were discovered, representing at least 6 individuals: two adults—a large mature man and a smaller individual, possibly a woman (18–21 years old), two adolescents, likely female, and two children aged four to seven. Fragments of bones and teeth. Traces of cannibalism. Mousterian and Levallois industries. 120–100 thousand years.

Moula-Guercy Neanderthal mandibular (left panel) and maxillary (right panel) dental material. Mesial is up, except (A) and (I).
Mandible: (A) M-G2-419, (B) M-D1-230, occlusal (C) M-L4TNN5, buccal/labial (D) M-G3-251, lingual (E) M-J5-TNN4, buccal/labial (F) M-*-TNN2, lingual (G) MG4-144, lingual (H) M-S-TNN1 buccal/labial;
Maxilla: (I) M-I4-55, (J) M-G2-117, distal (K) M-F3-215, occlusal (L) M-H3-73, lingual (M) M-I4-TNN3, lingual (N) M-D2-588, buccal/labial (O) MD1-259, lingual (P) M-S-27 lingual. ©Hlusko et al.

Guercy 1 represents the remains of a late-stage adolescent (?15–16.0 years) female. It comprises a conjoining set of 11 cranial vault fragments. ©Richards et al.

Ohaba Ponor (Bordul Mare, Sura Mare)

Ohaba Ponor (also Bordul Mare). Romania. Ohaba-Ponor village, Pui commune, Hunedoara County. Şura Mare Cave. Three phalanges of the lower limb of an adult individual. Later, another phalanx was discovered. This phalanx is from a different individual, but it is also from an adult Neanderthal. All of them have not been properly described to date. Neanderthals brought spherical geodes into the cave and painted them with ocher. The geodes may have had symbolic or ritual significance to the Neanderthals. Cult of bear skulls. Classical Neanderthal. Mousterian. 120–100 thousand years.

Neanderthal Remains Sites: Sura Mare Cave (Ohaba Ponor)

Soulabe-las-Maretas

Soulabé-las-Maretas. Southwest France. Montseron, near Malarno, Ariège. 3 left lower molars (first, second, and third) and 1 incisor of an adult individual. There is taurodontism. Classical Neanderthal. Mousterian. 120–100 thousand years.

Montgaudier

Grotte de Montgaudier (Montgaudier Cave). France. Near the village of Les-Eyzies, close to the town of Montbron, Charente, Aquitaine: Dordogne. This Neanderthal remains site includes the rock shelter Abri Lartet. Incomplete mandible of a child with 3 permanent teeth. No chin protrusion, symphysis straight or slightly inclined backwards. A girl aged 12–14 years. Mousterian industry. 120–80 thousand years.

An adolescent female neandertal mandible from Montgaudier Cave.

Šala

Šaľa, open-air site. Southwest Slovakia. On the banks of the Váh river, Galanta District, 50 km northeast of Bratislava, Moravia. Šala I—the well-preserved frontal bone of a young woman; Šala II—two parts of the cranium (left parietal and frontal bones) of an adult male aged 40–59 years. Neanderthals of the late classical “Spy” group. 120–80 thousand years.

Frontal bone of the Neanderthal specimen Šala. The frontal bone is probably female. ©Matejka.

The left frontal and parietal bones of the Neanderthal specimen Šala 2. ©Jakab.

Chateauneuf

Chateauneuf, open-air site, and the caves of Châteauneuf. France. The commune of Châteauneuf-sur-Charente, near the city of Angoulême, department of Charente. Several Neanderthal remains sites are located here:

Grotte-a-Melon

In the Grotte-à-Melon cave, among Mousterian stone tools, an incomplete lower Neanderthal jaw of a child aged 2–3 years was found (according to Patte).

L’abri de Hauteroche

Chateauneuf 2: At the foot of the cliff, in the Hauteroche (Haute-Roche) shelter, fragments of a child’s bones were discovered. An incomplete and flattened skull, part of the right ramus of the mandible, and several isolated teeth. The left deciduous molars are more worn than the right ones. The roots of the deciduous molars are undergoing resorption. The age is estimated to be between five and six years. Additionally, an incisor of an adult individual and an isolated child’s tooth from the group of microdont Mediterranean Neanderthals were found. These remains belong to the classical Neanderthals, carriers of a stone industry related to the denticulate Mousterian. There is a circular hearth of more than one square meter surrounded by two rows of limestones. 120–70 thousand years. IMAGE-пещера

Skhul

Skhul Cave, or rock shelter, Mugharet es-Skhul, “Cave of the Kids”. Israel. Western slope of Mount Carmel. 14 individuals, both Homo sapiens and Homo neanderthalensis (for example, Skhul 4 and 9). Skhul 4: Male, 40–50 years old; the forehead is sloping, the mandible is very massive, and the nose protrudes strongly. Skhul 9: Male, about 50 years old; the skull vault is low, and the brow ridge is strong; there are traces of wounds on the skull. The identity of individual Skhul 5 is questionable. Levallois-Mousterian industry, decorations made from drilled shells. 119–100 thousand years and 80–40 thousand years.

Skhul-4. The skull is clearly visible, but part of the skeleton is still embedded in the matrix. ©Danielle

Jaskinia

Jaskinia Ciemna (Dark Cave). Poland. Małopolskie Voivodeship, Ojcowski National Park. Ciemne 1 is the right lower incisor of an individual aged 20–40 years. The root is broken. Tiny phalanges—bones of the fingers of a 5–7-year-old Neanderthal child. The bones show traces of digestion, apparently from a large predatory bird. Micoquian. 115,000 years.

Sarstedt

Sarstedt, sand quarry. Northwestern Germany. Hildesheim District, Leine River Valley, south of Hanover. Skull fragments, including: the complete right temporal bone (Sst I) of a girl aged 2 to 4 years; the occipital fragment (Sst II); and the left parietal fragment (Sst III), which has some sapient features but is still Neanderthal. There are also unpublished findings. Classical Neanderthal. Keilmesser-gruppen techno-complex. Middle Paleolithic, late Pleistocene, 115 thousand years.

Sst I, Sst II, and Sst III. Hominid skull fragments from Late Pleistocene layers in Leine Valley (Sarstedt, District of Hildesheim, Germany). ©Czarnetzki A., Gaudzinski S., et Push C.M.

Sst I, Sst II, and Sst III. Hominid skull fragments from Late Pleistocene layers in Leine Valley (Sarstedt, District of Hildesheim, Germany). ©Czarnetzki A., Gaudzinski S., et Push C.M.

Qafzeh

Qafzeh Cave, Djebel Kafzeh. Israel. South of Nazareth. About 25 skeletons were found, including Neanderthals. Qafzeh-11 is the burial of a Neanderthal adolescent aged 12–13 years. In his hands, he held the antlers of a large red deer to his chest. Brain volume: 1530–1560 cm³. Levallois-Mousterian industry. 115–92 thousand years.

A skull cast of the Qafzeh 11 child. Natural History Museum, London.

Burial 11 from Qafzeh cave, Israel. On the left is a photo taken part-way though excavating the grave, showing the upper portion of the skeleton and the deer antlers clasped in the hand. On the right is a drawing of the burial showing the same. Image from Vandermeersch & Bar-Yosef 2019.

Qafzeh 11. Mandible of adult specimen from Djebel Kafzeh. ©Schwartz et al.

Balver (Balve)

Balver Höhle, Balve Cave. Northwestern Germany. Sauerland Karst, Hönnetal, North Rhine-Westphalia. Human remains were discovered. Classical Neanderthal. Micoquian, Mousterian industry. Middle Paleolithic, 110–85 thousand years.

Subalyuk

Subalyuk (Suba-lyuk) Cave. Hungary. Near the village of Cserkpfalu, Borsod-Abacj-Zeinplén county, Bükk National Park. The remains of two individuals. Subalyuk 1 includes a partial mandible, teeth, and some postcranial elements. It is unclear whether these adult remains belong to the same individual. Subalyuk 2 consists of the cranial vault, maxilla, and isolated teeth of a child aged approximately 3–7 years. Classical Neanderthals. Late Mousterian industry of the Quina type. 110–80 thousand years.

A: The adult Subalyuk 1 mandible in superior view; B: Anterior view of the Subalyuk 1 mandible illustrating the symphyseal morphology; C: The Subalyuk 1 jaw in left lateral view. The arrow indicates the development of the incurvatio mandibulae anterior. ©Ildiko Pap et al.

Masque

La Masque. Southern France. Vaucluse. At this site, numerous remains of a single adolescent were found, including skull fragments, vertebrae, bones of the hands and feet, ulna, humerus, radius, patella, and premolar. Classical Neanderthal. Mousterian industry. Middle Paleolithic, 110–80 thousand years.

Moulin du Milieu

Moulin du Milieu Cave. France. On the right bank of the Lède River, Gavaudun commune, Lot-et-Garonne, Agen, Haut Agenais. Hominid remains. Mousterian of Quina type. Middle Paleolithic, 110–80 thousand years.

Figueira Brava

Figueira Brava Cave (Gruta da Figueira Brava). Portugal. The foot of the southern slope of Serra ela Arrabiela, between Alpertuche and Portinho da Arrábida. The remains correspond to an individual aged 16 to 17 years, most likely male. Discovered and collected were a Neanderthal tooth and phalanx, the second premolar, metacarpal bone, and phalanx. Classical Neanderthal. Evidence of hearths. Consumption of seafood. Cannibalism. Late Mousterian-Levallois. 106–86 thousand years.

Comparison of a Homo neanderthalensis premolar from Gruta da Figueira Brava (left) with an identical Homo sapiens tooth. Sequentially: occlusal, distal, and buccal views. Greatest length: 2.3 cm. According to M. Telles Antunes

Premolar (left) and molar (right) teeth of Homo neanderthalensis from the Figueira Brava cave. Note the wear grooves on the molar. ©JE Egocheaga et al., 2004.

An idealized reconstruction of the Figueira Brava cave, inhabited by Neanderthals. Internal view. Source: drawing by Claudia Matos Pereira, 2015.

Ganovce

Ganovce. Slovakia. Near the town of Poprad. Ganovce-1 is a cast of the brain cavity with a part of the cranial vault of an adult woman, with a volume of 1320 cm³. Ganovce-2 is the gracile radius and fibula bones of a child. Atypical Neanderthals. 105 thousand years.

The travertine endocast with bone rests of the Neanderthal woman Ganovce 1 (from top to bottom and left to right: right lateral, left lateral, anterior, superior, inferior, and posterior views). Photographs by Martin Frouz.

Travertine casts of the left fibula – specimen Ganovce 2: a) the positive plaster cast, b) the negative cast. ©Vlcek

Merveilles (Castel-Merle)

Abri des Merveilles. Southwest France. Castel-Merle Valley (also known as Vallon des Roches, a small stream) on the banks of the Vézère River, Sergeac commune, Dordogne. The first or second lower molar of a hominid was discovered. The molar is severely worn during life, but otherwise it is practically undamaged. There is moderate taurodontism. Rock crystal tools. Mousterian. 105–100 thousand years.

Buccal, lingual, mesial, distal, and occlusal views of the Abri des Merveilles lower left molar. ©Trinkaus.

Djruchula

Djruchula (Djroutchoula) Cave. Western Georgia. Village of Kvemo-Zodi, right bank of the Djruchula River, Chiatura district, Imereti, Transcaucasia. In the second Mousterian cultural layer of Djruchula Cave, near a hearth, the upper left first permanent molar of an adult individual was found. It exhibits taurodontism, and the roots are fused. By all indications, the tooth is Neanderthal and morphologically similar to remains from the Skhul Cave. There are remains of the hearths. Mousterian, Levallois. Middle Paleolithic, 105–90 thousand years.

Djruchula left M1 occlusal view. Scale = 1 cm. Photos by F. Rivals and T. Chevalier.

Matuzka

Matuzka Cave. Russia. On the right bank of the Matuzka River, near the hamlet of Guamka, Krasnodar Krai, in the Northwest Caucasus. A large, well-preserved left lower canine; only the very tip of the root is broken off; and a fragment of the upper right lateral incisor. Both teeth have a shovel-shaped form and exhibit characteristics typical of Neanderthals. Hearth remains. Mousterian. Middle Paleolithic. 105–90 thousand years.

Matuzka. Human tooth. Layer 5B. © Golovanova et al.

Pesturina

Pešturina (aka Jelašnica Cave). Southeastern Serbia. Near the village of Jelašnica, close to the city of Niš. Atlas fragment (PES-1). Diaphysis (shaft) of the left radius bone of a young individual (PES-2)—92–44 thousand years. Well-preserved permanent right first molar of a child (PES-3)—102 thousand years. Charentian Mousterian industry.

Pesturina. A 3D re-creation of a Neanderthal tooth unearthed in Serbia shows the finer details of the 100,000-year-old fossil. Joshua Lindal in Radovic et al.

Foradada

Cova Foradada (Forada, La Safor). Spain. Calafell, near the village of Oliva, Valencia province. Initially, almost complete maxilla with teeth, a fibula, and four cranium fragments from an adult individual aged 34–45 years were found; a molar from a child aged 2.5 years. Later, the most complete Neanderthal skeleton on the Iberian Peninsula was discovered. This is a man aged 30–40 years, a classical Neanderthal. Possibly, it is a burial. Eagle claw necklace (Châtelperronian layer). 100 thousand years.

Neanderthal remains have been found at Cova Foradada in Oliva and are on display at the Museu de Prehistoria de Valencia. ©Biel Alino

Necklace. Phalanx 1 of the first digit (the thumb or Hallux) from the left foot of a large eagle. Dorsal, medial, plantar, and lateral views of the phalange (from left to right, respectively). ©Rodriguez-Hidalgo

Taddeo

Taddeo cave. Southwest Italy. Marina di Camerota coast (La Calanca beach), Salerno province, Campania. 3 Neanderthal teeth. Taddeo 1 is the lower left canine. Taddeo 2 is the upper right premolar. Taddeo 3 is the upper right molar. T2 and T3 belong to the same individual. Taddeo 4 (lower molar) has characteristics of both Neanderthal and Sapiens, so its classification is uncertain. Mousterian. Classical Neanderthals. Middle Paleolithic, 100–80 thousand years.

Taddeo 1 (LC1): a) Labial view; b) Lingual view; c) Occlusal view; d) Distal view; e) Mesial view. The white bar in the figure is equivalent to 1 cm. ©Benazzi et al.

3D digital model of Taddeo 2. a) Mesial view of the tooth without the wax used for filling the holes. b) Enamel-dentin junction (EDJ) of Taddeo 2. c) Enamel-dentin junction (EDJ) of Taddeo 3. B—buccal; L—lingual; D—distal; M—mesial. ©Benazzi et al.

Altmuhl

Altmühl Valley. Southern Germany. Neuessing area, Altmühltal, Bavaria. Neanderthal remains sites—three adjacent caves:

Klausennische

Klausennische, rock shelter. Located between Mittlere and Untere Klause caves. Crown of the first right upper deciduous incisor of a Neanderthal child (Neuessing 1). The root is missing. The current location of the fossil is unknown. Atypical Neanderthal. Late Acheulean, Micoquian, transition to Mousterian. Tools of the Wedge knife type. 100–70 thousand years.

Untere Klause

Untere Klause (Lower Klause). The lowest of the Klausen caves (Klausenhöhle). The acromial half of a typical, strongly S-shaped Neanderthal clavicle (Neuessing 3). Middle Paleolithic.

Sesselfelsgrotte

Sesselfelsgrotte, rock shelter site. Several finds of immature Neanderthals. Two shed deciduous teeth from children around 12 years old (Sesselfelsgrotte 2 and 3). The remaining 12 finds include skull fragments, a fragment of the mandible, some vertebrae, ribs, part of the humerus, ulna, and femur belonging to a newborn (Sesselfelsgrotte 1). Classical Neanderthal. Mousterian, Micoquian. 90–60 thousand years (second tooth) and 60–46 thousand years (skeleton and one tooth).

The neonate Neanderthal fossils from Sesselfelsgrotte. ©Rathgeber.

Leuca

The Leuca Caves, Capo di Leuca. Italy. Near Santa Maria di Leuca, Salento province, southern Italian region of Apulia. The site is located a few meters away from Grotta dei Giganti. Leuca I was found in the Grotta del Bambino (Child’s Cave), also known as the Grotta delle Tre Porte (Cave of the Three Doors). An unworn second upper left molar of a 10-year-old child. The crown is well preserved, but the root is incomplete. Classical Neanderthal. Mousterian. 100–70 thousand years.

Grotta Titti is located nearby, in the same cave. Neanderthal remains were also found here.

Left—Leuca I (LM2): (a) buccal; (b) lingual; (c) distal; (d) occlusal; and (e) mesial views. Scale 1 cm. Right—3D digital model of the reconstructed crown (a) and enamel-dentin junction EDJ (b).
PRO 5 protocone; PAR 5 paracone; MET 5 metacone; HYP 5 hypocone; C5 5 cusp fifth; AC-CO 5 accessory cusplet on the crista oblique; AC-PAR 5 accen#shy;tuated accessory ridge on the paracone; AC-PRO 5 accentuated accessory ridge on the protocone; MAT 5 mesial accessory tubercles. ©Benazzi et al.

Ras el Kelb

Ras el Kelb (Ras el-Kalb). Lebanon. The site is located on the coast, 8 km north of Beirut. Four teeth were found here. Two of them were discovered in layer 3 of level K; one was never studied, and the other, the upper left third premolar of an adult approximately 20 years old, was studied by Vallois and attributed to Neanderthals. Today, it is kept in the National Museum of Beirut (?). The second was found and extracted from a piece of breccia during the analysis of material in the National Museum of Beirut. Its identification should be carried out among the materials stored in the Department of Antiquities.

Two more teeth were found among the faunal remains from the site during the analysis of material in London. Both are upper left second molars, one permanent and one deciduous. The first is an upper molar belonging to an adult about 25 years old. The second is also an upper deciduous tooth, and it may belong to a child about six years old. These likely belong to Neanderthals, yet their preservation does not allow them to be reliably classified taxonomically. Mousterian, Levallois. 100,000–70,000 years ago.

Neanderthal, left upper premolar, Ras el-Kalb I. Photo Archives DGA. ©Yazbeck

Contrada Ianni (Nicotera)

Contrada Jannì (Ianni) di San Calogero. Southern Italy. Commune of Nicotera, near the city of Vibo Valentia, province of Catanzaro, Calabria. Almost complete left parietal bone (Nicotera 1) and a small free wormian bone, likely belonging to the right portion of the lambdoid suture of a child approximately 2–3 years old. Classical Neanderthal. Mousterian. Late Pleistocene. 100–60 thousand years.

Nicotera I left parietal. A: exocranic view; B: endocranic view. ©Bonfiglio et al.

Axlor

Axlor. Spain. Near the village of Dima, Biscay, Basque Country. In Axlor, the remains of four hominids were found: 7 teeth of young individuals and a fragment of the left parietal bone. 5 teeth from one individual, two of which are currently lost (descriptions and photos have been preserved), exhibit morphological features and an overall size indicative of sapiens. Two shed deciduous teeth from different individuals (right or left dm2 and left di1) and the parietal fragment of an adult found at the oldest level (VIII) show Neanderthal features. They were found in an undisturbed Mousterian context. Evidence of the use of fire. Classical Neanderthals. Mousterian-Levallois of the Quina type. Middle Paleolithic, 100–50 thousand years and 47–42 thousand years.

The hominid remains from Axlor. Neanderthal deciduous tooth. ©Gomez-Olivency et al.

Axlor VIII. Neanderthal parietal fragment. ©Gomez-Olivency et al.

Fondo Cattie (Maglie)

Fondo Cattìe. Italy. Municipality of Maglie, Lecce Province, Apulia. Neanderthal permanent (second or third) isolated molar from the right mandible (Maglie 1). The tooth shows evidence of taurodontism. Classical Neanderthal. Mousterian industry of the Quina type. Middle to Late Paleolithic. The dating is questionable, ranging from 100,000 to 40,000 years.

Maglie 1. ©Silvana M. Borgognini Tarli

Hora

Cueva Horá. Spain. Municipality of Darro, province of Granada. A well-preserved fragment of the frontal bone. This fragment was broken off from the main part after fossilization. It is difficult to determine the sex of the individual. This is either an adolescent male or an adult female. Classical Neanderthal. Typical Mousterian, not Levallois. Middle to Late Paleolithic. The dating ranges from 100,000 to 40,000 years.

Fragment of the frontal bone of Cueva Hora. Lateral, external, and internal views. ©Uriarte

Kalamakia

Kalamakia. Southern Greece. Western coast of the Mani Peninsula, northwest of Areopolis, Peloponnese. The site yielded 14 finds identified as human remains. These include isolated teeth, cranial, and postcranial remains. Among these, 8 isolated teeth belonged to individuals aged 30–50 years, and 2 teeth were from children around 6 years old, possibly from the same individual. The minimum number of individuals is 8. KAL3: Left upper third molar, the most ancient specimen on the site. KAL8: Right upper second molar. KAL10: Right upper lateral incisor, with a shovel-shaped crown. KAL11: Left upper central incisor, belonging to a 40–50-year-old Neanderthal. KAL1: Occipital fragment. KAL4: A subadult lumbar vertebra (L-3 or L-4) belonging to a juvenile aged between 6 and 18 years. KAL7: Right fibula fragment. KAL14: Left navicular bone. There are several hearths. Mousterian with elements of Levallois. Middle Paleolithic, dating from 100,000 to 39,000 years.

KAL4 vertebra in (a) superior, (b) inferior, and (c) posterior views. Scale in centimeters.

KAL14 navicular. (a) proximal, (b) distal, and (c) dorsal views. Scale in centimeters.

Rescoundudou

Rescoundudou. France. Sébazac-Concourès, Aveyron. Five Neanderthal remains: three deciduous teeth from the upper and lower jaws (one of them fragmentary), a fragment of a permanent lower molar, and the germ of a permanent upper lateral incisor (Resc V). They represent at least two different individuals: an adult with worn teeth and an adolescent around eight years old. Mousterian La Ferrassie type, Levallois. It dates to 96,000–82,000 years ago.

Le Rescoundudou I in place during the excavation. Photo: L. Batreau.

The Le Rescoundudou teeth. a: Resc I occlusal view; b: Resc II occlusal view; c: Resc III buccal view; d: Resc IV occlusal view; e: Resc V mesial view; f: Resc V occlusal view.

Valdegoba

Valdegoba Cave. Northern Spain. Near Huermeces, north of the city of Burgos. 13 remains from five individuals. Two mandibular fragments of a male adolescent aged 13–14 years, VB1, preserves an almost complete set of teeth (11) and exhibits a combination of both archaic and specific Neanderthal features. The second individual, VB2, has a set of ten deciduous teeth, indicating an age of 6–9 months at the time of death. VB3 is a proximal finger phalanx; VB4 is the fourth right metatarsal of an adolescent (around 15 years); and VB5 is the fifth left metatarsal of an adult. Classical Neanderthals. Mousterian–Levallois. It dates to 95,000–73,000 years ago.

The Valdegoba postcranial remains. a) VB4; b) VB5; d) VB3.

The Valdegoba 1 (VB1) mandible. A—right lateral view; B—superior view; C, D—another fragment of the same jaw; E—anterior view; F—internal symphysis; G—superior view of the left fragment; I—VB1 left P3 in mesial (right) and distal (left) views, showing bifurcation of the root. Scale bar = 2 cm. ©Quam et al.

Tossal de la Font

Tossal de la Font (La Cova de Dalt del Tossal de la Font). Eastern Spain. Castellón, Vilafamés province, Valencia region. A total of three human fossils were found. These include a distal fragment of the left humerus (CTF-1) and a fragment of the right ilium (CTF-2), both from a young woman. The third specimen is a fragment of the upper left molar from an 8-year-old child (CTF-3). Typical Mousterian industry. Date to 90,000–70,000 years ago.

Human remains. a) Anterior, posterior, and distal views of humerus fragment CTF-1; b) Medial and lateral views of coxal fragment CTF-2. ©Gusi et al.

Madonna dell’Arma

Madonna dell’Arma (also known as Nostra Signora Annunziata dell’Alma) cave. Northwestern Italy. Mediterranean coastal area, near San Remo and Taggia, Imperia province, Liguria. The cave yielded four skull fragments. Two articulated fragments of the occipital bone are articulated. Also, a fragment of the frontal bone with the edge of the left eye socket. All three are probably from the same individual. The fourth fragment comes from a more ancient layer. Remains of hearths are present. Classical Neanderthal. Mousterian, Levallois. It dates to 88,000–73,000 years ago.

Rochelot

Grotte de Rochelot. France. Saint-Amant-de-Bonnieure, Charente department, Nouvelle-Aquitaine region. The site is a hyena den with Neanderthal remains. During excavations, six worn permanent teeth (two upper and four lower) of an adult Neanderthal were found. The teeth, apparently belonging to the same individual, are very well preserved. The roots are intact, but the crowns are worn to the extent that dentin is often exposed. A fragment of the humerus diaphysis with hyena bite marks was also unearthed. The deltoid tuberosity of the bone is narrow, characteristic of Neanderthals. Mousterian. It dates to 85,000–80,000 years ago.

Presentation of the dental remains of Rochelot. ©Couture, Tournepiche.

The two lower second molars of Rochelot: vestibular and occlusal views. ©Couture, Tournepichee.

Fragment of humerus diaphysis with traces of hyena impact. ©Couture, Tournepichee.

Merdivenli

Merdivenli Cave, “Staircase Cave”. Turkey. Near the village of Mağracık, Samandağ district, Hatay province, at the foot of the Amanos-Musa mountain range. Found a lower molar and three fragments of human bone. Subsequently, in the lower layer of Middle Paleolithic deposits, two upper permanent molars and one lower permanent molar of a human were discovered. Levallois-Mousterian stone tools, similar to Near Eastern ones (Levantine), and Mousterian artifacts resembling those from Europe were identified. It dates to 85,000–75,000 years ago.

Merdivenli cave, Turkey.

Genay

Genay. France. Armançon Valley, Genay et Créancey, municipality of Côte d’Or, Burgundy. Remains of two adults and one child. Genay 1 has twenty-five teeth and several skull fragments, including the left part of the supraorbital torus, as well as fragments of parietal, occipital, and temporal bones. Genay 2 is an isolated tooth. The tooth is not well preserved, but it appears to be from an adult. Genay 3 is the germ of the lower left molar of a 2–3-year-old child. Mousterian. 82 thousand years ago.

The human remains of Genay 1 in situ. Note the position of the teeth, which generally respects their anatomical arrangement (collection and copyright DRAC Bourgogne).

Genay 3, occlusal, and lingual faces. Scale: 10 mm. ©Garralda et al.

Ramandils

Ramandils Cave. Southern France. “Le Rec Mendil”, Port-la-Nouvelle commune, department of Aude, 30 km from the city of Narbonne. 3 fragmented human teeth. Classical Neanderthal. Evidence of rabbit and shellfish consumption. Typical Mousterian. Middle Paleolithic, 82–80 thousand years ago. IMAGE-пещера

At the Les Ramandils cave in southern France, many rabbit bones were found burned, suggesting the animals were eaten by humans. ©Eugene Morin.

Fate

Fate Cave (Le Fate, Caverna delle Fate). Northern Italy. Finale Ligure, province of Savona Savona, Liguria. 16 fragments of bones from Neanderthal adults and children, attributed to at least five individuals. They include skull fragments, 10 isolated teeth, and postcranial fragments. The fragment of the frontal bone (Le Fate I) and the left lower half-jaw with three teeth (Le Fate II) belong to an 8–10-year-old child. The fragment of the mandible with the right third molar (Le Fate III) belongs to an adult individual. Classical Neanderthals from the Hortus group. Levallois industry, typical Mousterian. 82–75 thousand years ago.

Fate II and Fate III. Neanderthal remains from a Mousterian deposit in Northern Italy (Caverna delle Fate, Finale Ligure). ©Giacobini et al.

Calascio

Grottoni di Calascio, Rocca Calascio. Italy. Province of L’Aquila in Abruzzo. The head of the right femur (proximal epiphysis) of a boy aged 12–14 years. Classical Neanderthal. Traces of hearths. Mousterian. Middle Paleolithic, 80,000 years ago.

Grottoni di Calascio.

Peyrards

Les Peyrards (also Baume des Peyrards). France. Buoux commune, Apt, region Provence-Alpes-Côte d’Azur, department of Vaucluse. Remains of Neanderthals: four teeth from three young individuals and a child around ten years old. Hearth sites. Classical Neanderthal. Mousterian, Levallois. 80,000 years ago.

Lakonis

Lakonis I, rock shelter. Southern Greece. Mani peninsula, southern Peloponnese. Left lower third molar of a Neanderthal, approximately 35 years old. Late Paleolithic, 80,000–75,000 years ago.

Lakonis I (LKH 1), a Neanderthal left M 3. (A), occlusal view; (B) radiograph of lateral view; (C) buccal view; (D) lingual view. ©Harvati et al.

Kiik-Koba

Kiik-Koba. Ukraine. Crimea, Zuia River shore. Kiik-Koba 1: Adult male (alternatively reported as female) aged 40, with remains of the leg, foot, and more (canine, 17 hand elements, well-preserved right patella, tibia, and fibula, and 53 bilateral foot remains). Kiik-Koba 2: An appendicular skeleton (scapula, humerus, ulna, radius, metacarpals, phalanges, pelvic bone, femur, tibia, fibula) of a 12-month-old infant. Both findings represent burials. Classical Neanderthals. Kiik-Koba Mousterian. 80,000–70,000 years ago.

The Kiik-Koba 2 right scapula in dorsal, glenoid, and axillary views. ©Mednikova et al.

Pecheurs

Abri des Pêcheurs, rock shelter. France. The site is located on the left bank of the Chassezac, a tributary of the Ardèche River, Casteljau, Ardèche. Robust lower canine of an adult Neanderthal. Mousterian. Middle and Upper Paleolithic, 80,000–70,000 years ago.

Lower canine from Abri des Pecheurs. Distal and lingual views. ©Bouvier.

Rotches de D’Gennly (Montignies)

Rotches de D’Gennly Cave. Belgium. Commune of Montignies-le-Tilleul, on the right bank of the Sambre River, Hainaut province, Wallonia. Three left metacarpals belonging to one individual and a fragment of the left femur (broken at the proximal end near the second trochanter and, slightly above the condyles, at the distal end). Mousterian. Middle Paleolithic. 80,000–60,000 years ago.

Rigabe

Grotte de Rigabe. Southern France. Commune of Artigues, Var department, Draguignan district, Rians canton. One tooth. A hearth lined with stones. Classical Neanderthal. Mousterian, Levallois. 80,000–60,000 years ago.

Grotte de Rigabe

Chagyrskaya

Chagyrskaya Cave. Russia. On the left bank of the Charysh River, Northwestern Altai. 74 bone fragments and teeth, the remains of a total of 13 individuals. Discoveries from Chagyrskaya-8 (the phalanx of a woman) and Chagyrskaya-12 belonged to the individual Chagyrskaya-F. The age of this fossil is 80 thousand years. The mandible Chagyrskaya-6 and incisor Chagyrskaya-14 belong to the same individual (Chagyrskaya-C). The deciduous tooth Chagyrskaya-19 and two molars Chagyrskaya-13 and Chagyrskaya-63, despite different stages of development, belong to the same boy (Changyrskaya-G), who lost them in this cave between the ages of nine and fifteen. Bone tools. Mousterian and Micoquian industries. 59,000–51,000 years ago.

Some 74 Neanderthal fossils have been found at Chagyrskaya since 2007, including Chagyrskaya-9. Picture: Maria Mednikova

Right lower jaw fragment (Chagyrskaya 6) with canine, premolars, and 2 molars (tooth 43–47). It was discovered in 2011 in the Chagyrskaya Cave in Siberia. ©Thilo Parg, ©Salazar-Garcia et al.

Ochoz (Svedul Stul)

Ochoz (Sveduv Stul Cave). Czech Republic. Central Moravia, Prostějov District. Skull fragments, including parietal and temporal bones (Ochoz II), the mandible of an adult individual (Ochoz I), the right molar, and postcranial fragments. Mousterian. 80,000–45,000 years ago.

Sveduv Stul cave. The mandible of a Neanderthal man, also known as the Ochoz mandible. This jaw found in 1905 by Karl Kubasek in the rear of the cave under a fallen ceiling. ©Hitchcock

Hunas

Hunas. Southern Germany. Located 40 km from Nuremberg, southeast Bavaria. Isolated third (or second) right lower molar. The tooth is moderately worn and nearly intact. Taurodontism is absent. Classical Neanderthal. Mousterian. Middle Paleolithic, 79,000–76,000 years ago.

Crvena Stijena

Crvena Stijena, (Red Cliff), rock shelter. Montenegro. Municipality of Nikšic, near the village of Petrovici, Podgorica, near the border of Montenegro and Bosnia and Herzegovina. Canine tooth. This Neanderthal tooth was found during the examination of fragments that fell from the lower layers. Remains of stone-lined hearths. The traditional facies (Denticulate Mousterian, Micromousterian, etc.) do not directly attribute stone tool assemblages. Instead, an undifferentiated (non-Levallois) facies of the typical Mousterian with Uluzzian elements is attributed to them. Middle Paleolithic, 78,000–48,000 years ago.

Left: Neanderthal tooth found while screening debris fallen from lower strata in Crvena Stijena (photo: Jamie Clark)
Right: Circular hearth basin, with charcoal and ash fill removed, near the base of Layer 2, in the in situ Mesolithic deposits in Area I, 2005 excavations at Crvena Stijena (photo: Dusan Mihailovic)

Castillo

El Castillo Cave. Spain. Near the village of Puente Viesgo, Cantabria. Two findings consist of postcranial skeleton bones. One is a fragment of the proximal phalanx of the right index finger of an adult individual (Castillo-416), dating back to 77,000 years ago. The second is a fragment of the head of the right femur, which belonged to a child approximately eight years old (Castillo-228). About 70 thousand years.

Castillo-1466 (ULP4) is the second left upper premolar of an adult individual. This tooth has a thick layer of enamel and an unusual pulp structure, dating back 44.2 thousand years.

Three isolated crowns of deciduous teeth from three children. The non-erupted upper left central incisor (ULdi1, Castillo 924) of a 4–5-year-old child; the erupted upper left second molar (ULdm2, Castillo 492) of a 10-year-old child; the erupted lower right second molar (LRdm2, Castillo 292) of a 10–13-year-old child. Middle to Late Paleolithic, 44–43 thousand years ago. Classical Neanderthals. Mousterian of Acheulean Tradition.

Left: Castillo-416 hand phalanx. A: dorsal; B: palmar; C: proximal; and D: distal views. Right: Castillo-228, the femoral head. A: medial side; B: lateral surface towards the metaphyseal surface on the proximal femoral end. Scale = 50 mm. ©Maria Dolores Garralda et al.