|An approximate reconstruction of a Neanderthal skeleton. The central rib cage, including the sternum, and parts of the pelvis are from modern humans.|
|Known Neanderthal range in Europe (blue), Southwest Asia (orange), Uzbekistan (green), and the Altai Mountains (violet).|
Currently the earliest fossils of Neanderthals in Europe are dated between 450,000 and 430,000 years ago, and thereafter Neanderthals expanded into Southwest and Central Asia. They are known from numerous fossils, as well as stone tool assemblages. Almost all assemblages younger than 160,000 years are of the so-called Mousterian techno-complex, which is characterised by tools made out of stone flakes. The type specimen is Neanderthal 1, found in the Neander Valley in the German Rhineland, in 1856.
Compared to modern humans, Neanderthals were stockier, with shorter legs and bigger bodies. These are often explained as adaptations to conserve heat in a cold climate, but these could also be adaptations for sprinting in a warmer, forested landscape which they seemed to have preferred based on fossil distribution. Male and female Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively, within the range of the values for anatomically modern humans. Average males stood around 164 to 168 cm (5.4 to 5.5 ft) and females 152 to 156 cm (5.0 to 5.1 ft) tall.
There has been growing evidence for admixture between Neanderthals and anatomically modern humans, reflected in the genomes of all modern non-African populations but not in the genomes of most sub-Saharan Africans. The proportion of Neanderthal-derived ancestry is estimated to be around 1–4% of the modern Eurasian genome. This suggests that some interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration of around 50-70,000 years ago. Recent admixture analyses have added to the complexity, finding that Eastern Neanderthals derived up to 2% of their ancestry from an earlier wave of anatomically modern humans who left Africa some 100,000 years ago.
- 1 Name and classification
- 2 Evolution
- 3 Demographics
- 4 Anatomy
- 5 Culture
- 6 Interbreeding with archaic and modern humans
- 7 Extinction
- 8 History of research
- 9 Specimens
- 10 In popular culture
- 11 See also
- 12 Notes
- 13 References
- 14 Further reading
- 15 External links
Name and classification
Neanderthals are named after one of the first sites where their fossils were discovered in the mid-19th century in the Neander Valley, just east of Düsseldorf, at the time in the Rhine Province of the Kingdom of Prussia (now in North Rhine-Westphalia, Germany). The valley itself was named for Joachim Neander, Neander being the Hellenized form of the surname Neumann ("new man"). The German spelling of Thal (valley) was current in the 19th century; it is now spelled Tal.[a]
Neanderthal 1 was known as the "Neanderthal cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man". The binomial name Homo neanderthalensis—extending the name "Neanderthal man" from the individual type specimen to the entire group—was first proposed by the Anglo-Irish geologist William King in a paper read to the British Association in 1863, although in the following year he stated that the specimen was not human and rejected the name. King's name had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus. Popular English usage of "Neanderthal" as shorthand for "Neanderthal man", as in "the Neanderthals" or "a Neanderthal", emerged in the popular literature of the 1920s.
Since the historical spelling -th- in German represents the phoneme /t/, not the fricative /θ/, standard British pronunciation of "Neanderthal" is with /t/ (IPA: /niːˈændərtɑːl/). Because of the usual sound represented by the digraph ⟨th⟩ in English, "Neanderthal" is also pronounced with the voiceless fricative /θ/ (as /niːˈændərθɔːl/).
The spelling Neandertal is occasionally seen in English, even in scientific publications. Since "Neanderthal", or "Neandertal", is a common name, there is no authoritative prescription on its spelling, unlike the spelling of the binominal name H. neanderthalensis, which is predicated by King 1864. The common name in German is invariably Neandertaler (lit. "of the valley of Neander"), not Neandertal, but the spelling of the name of the Neander Valley itself (Neandertal vs. Neanderthal) has been affected by the species name, the names of the Neanderthal Museum and of Neanderthal station persisting with pre-1900 orthography.
Ever since the discovery of the Neanderthal fossils, expert opinion has been divided as to whether Neanderthals should be considered a separate species (Homo neanderthalensis) or a subspecies (Homo sapiens neanderthalensis) relative to modern humans. Svante Pääbo (2014) described such "taxonomic wars" as unresolveable in principle, "since there is no definition of species perfectly describing the case." The question depends on the definition of Homo sapiens as a chronospecies, which has also been in flux throughout the 20th century. Authorities preferring classification of Neanderthals as subspecies have introduced the subspecies name Homo sapiens sapiens for the anatomically modern Cro-Magnon population which lived in Europe at the same time as Neanderthals, while authorities preferring classification as separate species use Homo sapiens as equivalent to "anatomically modern humans".
During the early 20th century, a prevailing view of Neanderthals as "simian", influenced by Arthur Keith and Marcellin Boule, tended to exaggerate the anatomical differences between Neanderthals and Cro-Magnons. Beginning in the 1930s, revised reconstructions of Neanderthals increasingly emphasized the similarity rather than differences from modern humans. From the 1940s throughout the 1970s, it was increasingly common to use the subspecies classification of Homo sapiens neanderthalensis vs. Homo sapiens sapiens. The hypothesis of "multiregional origin" of modern man was formulated in the 1980s on such grounds, arguing for the presence of an unbroken succession of fossil sites in both Europe and Asia. Hybridization between Neanderthals and Cro Magnon had been suggested on skeletal and craniological grounds since the early 20th century, and found increasing support in the later 20th century, until Neanderthal admixture was found to be present in modern populations' genomes in the 2010s.
It is largely thought that H. heidelbergensis was the last common ancestor of Neanderthals, Denisovans, and modern humans after populations became isolated in Europe, Asia, and Africa respectively. The taxonomic distinction between H. heidelbergensis and Neanderthals is mostly due to a fossil gap in Europe between 300,000 and 243,000 years ago (Marine isotope stage 8). "Neanderthals", by conventions, are fossils which date to after this gap. The quality of the fossil record greatly increases from 130,000 years ago onwards. Specimens younger than this date make up the bulk of known Neanderthal skeletons and were the first whose anatomy was comprehensively studied. In morphological studies, the term "classic Neanderthal" may be used in a narrower sense for Neanderthals younger than 71,000 years old (MIS 4 and 3).
Numerous dates for the Neanderthal/human split exist. The date of around 250,000 years ago cites the Florisbad Skull ("H. helmei") as being the ancestral species, and the split is associated with the Levallois technique of making stone tools. The date of 400,000 years ago uses "H. rhodesiensis" as the ancestral species. 600,000 years ago claims that H. heidelbergensis only inhabited Europe and directly evolved into Neanderthal, classified as a chronospecies. 800,000 has H. antecessor as the ancestral species, however, different variations of this model would push the date back to 1 million years ago. DNA studies have yielded various results on divergence time, such as 182–592 kya, 321–553 kya, 475–654 kya, 550–690 kya, 550–765 kya, 520–800 kya, before 800 kya, and so forth.
Neanderthals and Denisovans are more closely related to each other than they are to humans, meaning the Neanderthal/Denisovan split occurred after their split with humans. When counting the number of derived alleles–the alleles which were not present in the last common ancestor–the human and Neanderthal/Denisovan lineages split around 765–550 kya.
As for the Neanderthal/Denisovan split, using a mutation rate of 1x10-9 or 0.5x10-9 per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190 kya or 473–381 kya respectively. Using 1.1x10-8 per generation with a new generation every 29 years, the time is 744 kya. Using 5x10-10 nucleotide site per year, it is 644 kya. Using the latter dates, the split had likely already occurred by the time hominins spread out across Europe, and unique Neanderthal features had begun evolving by 600–500 kya.
Early Neanderthals, living before the Eemian interglacial (130 ka), are poorly known and come mostly from European sites. From 130 ka onwards, the quality of the fossil record increases dramatically, recorded from Western, Central, Eastern, and Mediterranean Europe, as well as Southwest, Central, and Northern Asia up to the Altai Mountains in Siberia. The southernmost find was recorded at Shuqba Cave, Levant, and the easternmost as 85°E at Denisova Cave, Siberia.
The limit of their northern range appears to have been south of 53°N (Bontnewydd, Wales), although it is difficult to assess because glacial advances destroy most human remains, the Bontnewydd tooth being exceptional. Middle Palaeolithic artefacts have been found up to 60°N on the Russian plains.
It is unknown how the rapidly fluctuation climate of the last glacial period (Dansgaard–Oeschger events) impacted Neanderthals, as warming periods would produce more favorable temperatures, but encourage forest growth and deter megafauna; whereas frigid periods would produce the opposite. Populations may have peaked in cold but not extreme intervals, such as marine isotope stages 8 and 6. It is possible their range expanded and contracted as the ice retreated and grew respectively to avoid permafrost areas, residing in certain refuge zones. However, Neanderthals may have preferred a forested landscape.
Analysis of the genomic DNA from three locations suggests that about 120,000 years ago there were genetically distinct Neanderthal populations in Western Europe and Siberia. By 90,000 years ago, the western population had spread east into Siberia.
Like modern humans, Neanderthals probably descended from a very small population with an effective population–the number of individuals who can bear children–of 3,000 to 12,000 approximately. However, Neanderthals maintained this very low population, living in small, isolated, inbred groups. Various studies, using mtDNA analysis, yield varying effective populations, such as about 15,000 individuals (corresponding to a total population of roughly 150,000 individuals); 5,000 to 9,000 remaining constant; or 3,000 to 25,000 steadily increasing until 50,000 BCE before declining until extinction. However, all agree on low population, which may have been up to 10 times smaller than contemporary human populations in Western Europe. This may be explained in the context of the "Boserupian Trap": a population's carrying capacity is limited by the amount of food it can obtain, which in turn is limited by its technology. Innovation increases with population, but if the population is too low, innovation will not occur very rapidly and the population will maintain its low population. This is consistent with the apparent 150,000 year stagnation in Neanderthal technology.
This low population caused a low genetic diversity, inbreeding, and reduced the population's ability to filter out harmful mutations. However, it is unknown how this affected a single Neanderthal's genetic burden and, thus, if this caused a higher rate of birth defects than in humans. It is known that the closely related inhabitants on Sidrón Cave displayed several birth defects.
Based on tropical human hunter-gatherer societies, adult Neanderthals possibly lived on average 25–40 years. In a sample of 206 Neanderthals, based on the abundance of young and mature adults in comparison to other age demographics, about 80% of them above the age of 20 died before reaching 40. This high mortality rate was probably due to their high-stress environment. However, it has also been estimated that the age pyramids for Neanderthals and Upper Paleolithic humans were the same. Infant mortality was very high, about 43% in northern Eurasia.
Neanderthals had a more robust and stockier build than humans, but maintained the same upright posture as humans. Neanderthals had shorter limb proportions, a wider and barrel-shaped rib cage, wider pelvis, reduced chin, sloping forehead, and large nose[b] which also started somewhat higher on the face than in modern humans. The Neanderthal skull is typically more elongated and less globular than that of humans, and features a notable occipital bun.
Based on 45 Neanderthals long bones from 14 men and 7 women, the average height was 164 to 168 cm (5.4 to 5.5 ft) for men and 152 to 156 cm (5 ft) for women. For comparison, the average height of 16 Upper Paleolithic and Mesolithic humans was 168.1 cm (5.5 ft) for men and 152.5 cm (5 ft) for women. For Neanderthal weight, samples of 26 specimens found an average of 77.6 kg (171 lb) for men and 66.4 kg (146 lb) for women. Using 76 kg (168 lb), the body mass index for Neanderthal men was calculated to be 2.69–2.82, which in humans correlates to obesity. This indicates a very stocky build.
Body proportions are usually cited as adaptations to the cold, as they are similar to those of human populations which developed in cold climates–the Neanderthal build is most similar to Eskimos–and shorter limbs equates to higher retention of body heat. However, the increasing evidence that Neanderthals preferred warmer wooded areas instead of open mammoth steppe and cold climate suggests to the contrary, and DNA analysis indicates a higher proportion of fast-twitch muscle fibers in the Neanderthals than humans. It is possible their body proportions and greater muscle mass were adaptations to sprinting as opposed to the endurance-oriented human physique.
The large Neanderthal nose shows no adaptations to cold climate indicated by bone structure, and they had large paranasal sinuses, which is generally reduced in cold-adapted creatures. More likely, they were caused by genetic drift. This affected the shape of the skull and the muscle attachments, and likely gave them a weaker bite force than humans.
The size of the braincase of Amud 1 was calculated to be 1,736 cm3 (105.9 in3), one of the largest ever recorded in hominids. In Neanderthals, the occipital lobe–operating vision–was much larger than in modern humans, and, similarly, they had larger eyes, probably as an adaptation to lower light conditions in Europe. More brain tissue was devoted to bodily maintenance and control, and, consequently, the cognitive areas of the brain were smaller than in humans, including the cerebellum–operating muscle memory, and possibly language, attention, working memory, social abilities, and thought–the parietal lobes–visuospatial function and episodic memory–the temporal lobes–language comprehension and associations with emotions–the orbitofrontal cortex–decision making–and the olfactory bulb–sense of smell. A 2011 study looking at the brain asymmetry of 20 Neanderthals to predict handedness found 85% to be right-hand dominant and the remaining 15% left-handed, whereas humans are 52% right-dominant, 12% left-dominant, and 36% ambidextrous. The brain size of both Neanderthal and human infants is 400 cm3 (24 in3), and either Neanderthal brain development sped up or human development slowed down from the last common ancestor.
The DNA of a Neanderthal from Monti Lessini, Valpolicella, Italy, showed depressed activity of the MC1R gene, which is associated with red or blond hair. However, like in humans, red was probably not a very common hair color. The lack of sunlight most likely led to the proliferation of lighter skin in Neanderthals. Genetically, BNC2 was present in Neanderthals, which is associated with light skin colour, however, a second variation of BNC2 was also present, which is associated with darker skin colour in the UK Biobank. It is likely Neanderthal skin colour varied from region to region. The DNA of three Croation Neanderthals shows they had darker hair, skin, and eye colour than modern Europeans.
The Neanderthal physical activity level (PAL) was assumed to be a very high 650 counts per minute per day (CPM/d), in comparison to 200 CPM/d in modern Siberian hunter-gatherers. Average body fat percentage (BFP) was estimated to be 25%, though it may have been 13% in men and 20% in women in more temperate areas. Using these measurements and average height and weight, the daily total energy expenditure (TEE)–the amount of calories consumed in one day–was estimated to be 3,454–4,019 and 3,828–4,483 kcal for men with high and low BFPs respectively, and 3,115–3,538 and 3,258–3,710 kcal for women. However, if the PAL was reduced to that of modern Siberian hunter-gatherers, the TEE becomes 2,959–3,524 and 3,333–3,988 kcal for men, and 2,620–3,043 and 2,764–3,215 kcal for women. This is comparable with the upper end of energetic demands of modern hunter gatherers, and the latter estimates are most similar to the Siberian Yakuts, which contradicts earlier estimates of vastly higher energetic demands in Neanderthals than humans.
Maximum lifespan, and the timing of adulthood, menopause, and gestation were most likely very similar to modern humans. However, it has been hypothesized that Neanderthals matured faster than modern humans based on the growth rates of teeth and tooth enamel, though this is not backed by age biomarkers.
Genetically, Neanderthal-derived alleles near ASB1 and EXOC6 are associated with being an evening person, narcolepsy, and day-time napping. In a survey, people who had archaic haplotypes near CDH6 more frequently reported feeling unenthusiasm and disinterest. These are consistent with the idea that sunlight affects circadian rhythm and mood.
Neanderthals suffered a high rate of traumatic injury, with by some estimates 79% of specimens showing evidence of healed major trauma. It was thus theorized that Neanderthals employed a risky hunting strategy. However, rates of cranial trauma are not significantly different between Neanderthal and middle paleolithic anatomically modern human samples. Shanidar 1 shows signs of an amputation of the right arm likely due to a nonunion after breaking a bone in adolescence, osteomyelitis (a bone infection) on the left clavicle, an abnormal gait, vision problems in the left eye, and possible hearing loss. It is unlikely any of these are what ultimately killed him, however.
Likely due to advanced age (60s or 70s), La Chapelle-aux-Saints 1 had signs of Baastrup's disease affecting the spine and osteoarthritis. Shanidar 1, who likely died at about 40 or 50, was diagnosed with the most ancient case of diffuse idiopathic skeletal hyperostosis (DISH) which restricts muscle movement, which likely led to his death.
Low population also led to low genetic diversity and probably inbreeding which could have led to inbreeding depression. The 13 inhabitants of Sidrón Cave collectively exhibited 17 birth defects likely due to inbreeding.
Neanderthals likely lived in much smaller and more sparsely distributed groups than Upper Paleolithic humans. Analysis of the mtDNA of the Neanderthals of Sidrón Cave showed that the three men belonged to the same maternal lineage, while the three women belonged to different ones. This suggests that women "married out". Small parties composed no more of perhaps three members may have went off on days-long hunting trips, venturing off on average 10–12 km (6.2–7.5 mi) from base camp, though possibly as much as 25 km (16 mi). Bands moved between certain caves depending on the season, and likely returned to the same locations generation after generation.
A self-sustaining population which avoids inbreeding consists of about 450–500 individuals, which would necessitate these bands to interact with 8–53 other bands, but more likely the more conservative estimate given low population density. However, the DNA of a Neanderthal from the Altai Mountains show that her parents were half-siblings, and the inhabitants of Sidrón Cave were highly inbred.
Considering most Neanderthal artefacts were sourced no more than 5 km (3.1 mi) from the main settlement, it is unlikely these bands interacted very often. However, a skeleton from La Roche à Pierrot, France, showed a healed fracture on top of the skull apparently caused by a deep blade wound, and another from Shanadir Cave was found to have a rib lesion characteristic of projectile weapon injuries, which could be considered as evidence for conflict. Further, a few Neanderthal artefacts in a settlement could have originated 20, 30, 100, and 300 km (12.5, 18.5, 60, and 185 mi) away. It is possible that macro-bands could have formed, collectively encompassing 13,000 km2 (5,000 sq mi), with each band claiming 1,200–2,800 km2 (460–1,080 sq mi), maintaining strong alliances for mating networks or to cope with leaner times and enemies as is exhibited in the low-density hunter gatherer societies of the Western Desert of Australia. However, mapping of the Neanderthal brain indicates they may not have had the cognitive function required for complex social interactions and trade.
It is sometimes suggested, since they were hunters of challenging big game and lived in small groups, there was no sexual division of labour as seen in human societies. That is, men, women, and children all had to be involved in hunting, instead of just men hunting and women and children foraging which is a more efficient food-collecting system. However, in modern human societies, the higher the meat dependency, the higher the division of labour.
It is controversially proposed that some Neanderthals wore decorative clothing or jewelry–such as a leopard skin or raptor feathers–to display elevated status in the group. The few number of Neanderthal graves found could be explained as only high-ranking members receiving an elaborate burial, as is the case for some contemporary human societies. An apparent cemetery of six or seven individuals at La Ferrassie may indicate they consciously identified as a single group.
Children were likely weaned after 5 years–one year later than humans–probably to increase birth spacing. Indicated from various ailments resulting from high stress at a low age, such as stunted growth, children of both sexes were likely put to work directly after weaning. Upon reaching adolescence, an individual would join in hunting large game. A study looking at Neanderthal skeletons recovered from several natural rock shelters shows that, although they were recorded as bearing several trauma-related injuries, none of them had significant trauma to the legs which would debilitate movement. This might indicate that individuals who could not keep up with the group while moving from cave to cave were left behind. The high mortality rate indicates grandparents were rare. These all could indicate a culture based on the idea that self-worth derives from contributing food to the group; a debilitating injury would remove this self-worth and result in near-immediate death. In this hypothesis, elderly Neanderthals were given special burial rites for lasting so long.
Most available evidence suggests they were apex predators, and fed on red deer, reindeer, ibex, wild boar, aurochs and on occasion mammoth, straight-tusked elephant and woolly rhinoceros. Isotope studies of Neanderthals from two French sites showed similar profiles to other carnivores, suggesting that these populations ate fresh meat. Analysis of Neanderthal bone collagen from the Croatian Vindija Cave shows nearly all of their protein needs derived from animal meat. Dental tarter from Spy Cave indicates they had a meat-heavy diet including woolly rhinoceros and mouflon sheep, while also regularly consuming mushrooms. Living in a forested environment, Neanderthals were likely ambush hunters, getting close to and attacking their target in a short burst of speed, likely thrusting in a spear at close quarters.
Neanderthal diet may have varied significantly region to region, with some communities subsisting primarily on a plant-based diet. Edible plant remains are recorded from several caves. For example, study looking at Neanderthals from Sidrón Cave in Spain, based on dental tarter, found a meatless diet of mushrooms, pine nuts, and moss, indicating they were forest foragers. Neanderthal faecal matter from El Salt, Spain, dated to 50,000 years BCE–the oldest human faecal matter remains recorded–show elevated coprostanol levels (digested cholesterol indicating a meat-heavy diet) and elevated stigmastanol (deriving from plant matter). The low-calorie plant diet suggests the minimum daily caloric intake was low. Evidence of cooked food plants–mainly legumes and, to a far lesser extent, acorns–were discovered in the Israeli Kebara Cave, possibly gathering plants in spring and fall and hunting in all seasons except fall, though it was probably abandoned in late summer to early fall. Remnants from the Israeli Amud Cave indicates a diet of figs, palm tree fruits, and various cereals and edible grasses. At the Iraqi Shanidar Cave, Neanderthals collected plants with various harvest seasons, indicating they scheduled returns to the area to harvest certain plants, and that they had complex food-gathering behaviors for both meat and plants.
It is possible Neanderthals could employ a wide range of cooking techniques such as roasting, and may have been able to heat up or boil soup, stew, or animal stock. The abundance of animal bone fragments at settlements may indicate the making of fat stocks from boiling bone marrow, possibly taken from animals which had died of starvation. These methods would have increased protein consumption to make up for a lack of carbohydrates. At Sidrón Cave, Neanderthals likely cooked and possibly smoked food, as well as used certain plants–such as yarrow and camomile–to flavor food, though these plants may have instead been used for their medicinal properties. Further, the former hypothesis would assume a degree of cuisine complexity which lacks proper evidence. They likely lacked any method of storing food, and had to eat whatever they hunted or foraged immediately.
There are several examples of Neanderthals practising cannibalism. The first undisputed example came from Gran Dolina, and other examples were found at Sidrón Cave, Zafarraya in Spain; and the French Moula-Guercy Cave, Les Pradelles, and La Quina. For the five cannabalised Neanderthals at the Goyet Caves in Belgium, there is evidence that the upper limbs were disarticulated, the lower limbs defleshed and also smashed likely to extract bone morrow, the chest cavity disemboweled, the jaw dismembered, and the butchers used some bones to retouch their tools. The processing of Neanderthal meat at Goyet Caves is similar to how they processed horse and reindeer.
A large number of claims of Neanderthal art, adornment, and structures have been made. These are often taken by the media as showing Neanderthals were capable of symbolic thought, or were "mental equals" to anatomically modern humans. As evidence of symbolism, none of them are widely accepted, although the same is true for Middle Palaeolithic anatomically modern humans. Among many others:
- Incision-decorated raven bones from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea, Micoquian industry dated to between cal. 43,000 and 38,000 BP. Given there are 17 of these objects at seven different sites in the area, and the notches on all of them are more-or-less equidistant to each other, they are very unlikely to have originated from simple butchering.
- Flower pollen on the body of pre-Neanderthal Shanidar 4, Iraq, had in 1975 been argued to be a flower burial, but the pollen could have also been deposited by natural events.
- Châtelperronian beads have been attributed to Neanderthals, but the dating is uncertain and the beads may have been made by modern humans.
- Bird bones were argued to show evidence for feather plucking in a 2012 study examining 1,699 ancient sites across Eurasia, which the authors controversially took to mean Neanderthals wore bird feathers as personal adornments.
- Deep scratches on the floor of Gorham's Cave, Gibraltar, were dated to older than 39,000 years ago in 2012, which some have controversially interpreted as Neanderthal abstract art.
- Two 176,000-year-old stalagmite ring structures, several metres wide, were reported in 2016 more than 300 m (980 ft) from the entrance within Bruniquel Cave, France. The authors claim artificial lighting would have been required as this part of the cave is beyond the reach of daylight.
- In 2015, a study argued that a number of 130,000-year-old eagle talons found in a cache near Krapina, Croatia along with Neanderthal bones, had been modified to be used as jewelry.
- In 2018, red painted symbols comprising hand stencils, ladder-shaped figured, prints, dots, discs, lines, and representations of animals on the cave walls of three Spanish caves 700 km (430 mi) apart–La Pasiega, Maltravieso, and Doña Trinidad–were dated to be older than, respectively, 64,800; 66,700; and for one painting 65,500 years ago. If the dating is correct, they were painted before the time anatomically modern humans are thought to have arrived in Europe, and demonstrate Neanderthals were capable of symbolic behaviour.
- In 2018, perforated seashell beads and pigments that are at least 115,000 years old were found in Cueva de los Aviones in southeastern Spain.
Neanderthals made stone tools, used fire, and were hunters. Neanderthals are associated with the Mousterian stone tool industry. The Levallois technique they adopted maximizes the cutting surface with the least amount of raw material. As a difficult-to-learn process, the technique may have been directly taught generation to generation rather than via purely observational learning. A Levallois point embedded in the vertebrae of an African wild ass indicated that a javelin had been thrown with a parabolic trajectory to disable the animal. There is evidence that Neanserthal technology was more sophisticated than was previously believed, and increasing evidence that Neanderthals, like modern humans, used language.
In a number of caves, evidence of hearths has been detected. Neanderthals likely considered air circulation when making hearths as a lack of proper ventilation for a single hearth can render a cave uninhabitable in several minutes. Abric Romaní rock shelter indicates eight evenly spaced hearths lined up against the rock wall, likely used to stay warm while sleeping, with one person sleeping on either side of the fire. At Cueva de Bolomor, with hearths lined up against the wall, the smoke flowed upwards to the ceiling, and led to outside the cave. In Grotte du Lazaret, smoke was probably naturally ventilated during the winter as the interior cave temperature was greater than the outside temperature; similarly, the cave was likely only inhabited in the winter.
It was long believed that an adhesive (birch bark tar) made by Neanderthals required to follow a complex recipe, and that it thus showed complex cognitive skills and cultural transmission. A study from 2019 showed that birch tar production can instead be a very simple process - merely involving the burning of birch bark near smooth vertical surfaces in open air conditions.
As opposed to the bone sewing needles and stitching awls of Upper Paleolithic humans, the only known Neanderthal tools that could have been used to fashion clothes are hide scrapers, which could have made items similar to blankets or ponchos. They probably did not need tailored clothings as humans due to their anatomical adaptations to the cold, though this may not have made such an impactful difference. Nonetheless, Neanderthals probably covered up to 70–80% of their body, and, depending on the frigidness, the hands and feet; though contemporary humans would have covered 80–90%. Since human/Neanderthal admixture is known to have occurred in the Middle East, and no modern body louse species descends from Neanderthals–body lice only inhabit clothed individuals–it is possible Neanderthals in hotter climates did not wear clothes.
Remains of Middle Paleolithic stone tools on Greek islands indicate early seafaring of Neanderthals in the Ionian Sea possibly starting as far back as 150,000–200,000 BCE. The oldest stone artefacts from Crete date to 130,000–107,000 BCE, Cephalonia 125,000 BCE, and Zakynthos 110,000–35,000 BCE. Here, they likely employed simple reed boats and made one-day crossings back and forth. Other Mediterranean islands include Sardinia, Melos, Alonnisos, and it is possible they crossed the Strait of Gibraltar. Their ability to engineer these boats and navigate through open waters speaks to their advanced cognitive and technical skills.
They appear to have had some knowledge of medicine. An individual at Sidrón Cave seems to have been self-medicating a dental abscess using poplar–which contains salicylic acid, the active ingredient in aspirin–and there were also traces of the antibiotic-producing Penicillium. They may have also used yarrow and camomile, and their bitter taste–which should act as a deterrent as it could indicate poison–means it was likely a deliberate act. In Kebara Cave, plant remains which have historically been used for their medicinal properties were found, including the common grape vine, the pistachios of the Persian turpentine tree, ervil seeds, and oak acorns.
The 1983 discovery of a Neanderthal hyoid bone–used in speech production in humans–in Kebara 2 almost identical to that of humans suggests Neanderthals were capable of speech. The prevailing hypothesis for a long time was that speech spontaneously developed very recently in humans, and some argued that the hyoid could have a different usage in Neanderthals, as it is simply used in tongue movement including while chewing. However, the ancestral H. heidelbergensis displays some modifications to the ear associated with speech perception, so speech may be a much older hominin trait. Further, the discovery of Ardipithecus suggests the first proto-languages (albeit very simple) originated about 5 million years, and speech was an ancient trait.
The degree of language complexity is difficult to establish, but given Neanderthal technical achievements and some degree of cultural complexity, it is reasonable to assume they were at least fairly articulate, comparable to humans. A somewhat complex language was likely necessary to survive in their harsh environment, needing to communicate about topics as locations, hunting and gathering, and tool-making techniques.
Genetically, the FOXP2 gene is identified, in humans, to be very important in speech and language development. FOXP2 is present in Neanderthals, but not the modern human variant. Neurologically,
Claims that Neanderthals held funerals for their dead with symbolic meaning,:158–60 are heavily contested. Even if the burial was intentional, it is not indicative of a religious belief of life after death, as such burial could have been the result of great emotion. The debate on Neanderthal funerals has been active since the 1908 discovery of La Chapelle-aux-Saints 1 in a small hole in a cave in southwestern France, very controversially attributed to have been buried in a symbolic fashion. Another grave at Shanidar Cave was associated with the pollen of several flowers which may have been in bloom at the time of deposition–yarrow, Centaurea, ragwort, grape hyacinth, joint pine, and hollyhock. The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some leader, healer, or shaman. However, it is also possible the pollen was deposited by a small rodent after the man's death. The grave of Teshik-Tash 1 from Uzbekistan was associated with a circle of ibex horns, which was asserted by Sir Paul Mellars to indicate a ritualistic burial. However, the abundance of ibex horns in the vicinity casts doubt on this.
It was once asserted that the bones of the short-faced bear in some European caves were arranged in a particular order, indicating an ancient bear cult which killed bears and then ceremoniously arranged the bones. This would be consistent with bear-related rituals of human Arctic hunter gatherers, but the 'peculiar' arrangement of bones could also be well-explained by natural causes, and bias could be introduced as the existence of a bear cult would conform with the idea that totemism was the earliest religion, leading to undue extrapolation of evidence.
It was also once thought Neanderthals hunted, killed, and cannabalised other Neanderthals, and used the skull as the focus of some ritual. It is now thought that isolated damaged skull remains–which were put forward as evidence of the skull cult–were the result of predation by cave hyenas or simple decay. Nonetheless, Neanderthals are known to have practiced cannibalism, but there is unsubstantial evidence to suggest ritual defleshing.
Interbreeding with archaic and modern humans
An alternative proposal regarding the fate of Neanderthals is that rather than being replaced by modern humans and going extinct, Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of theory of recent African origin of modern humans, since it would imply that at least part of the genome of Europeans would descend from Neanderthals. Evidence of interbreeding has been detected in genomes of ancient and modern humans, though this could have resulted from occasional interbreeding and introgression, rather than population absorption.
The first Neanderthal genome sequence was published in 2010, and confirmed interbreeding between Neanderthals and humans. The genomes of all non-Africans include an estimated share of 1.5–2.1% Neanderthal DNA. This DNA is absent in Sub-Saharan Africans (such as the Yoruba and San peoples). The 2% of Neanderthal DNA in Europeans and Asians is not the same in all Europeans and Asians. In all, approximately 20% of the Neanderthal genome appears to have survived in the modern human gene pool.
Some researchers suggest admixture of 3.4–7.9%, or 1.5-2.1%. Detractors have argued and continue to argue that the signal of Neanderthal interbreeding may be due to ancient African substructure, meaning that the similarity is only a remnant of a common ancestor of both Neanderthals and modern humans and not the result of interbreeding. John D. Hawks has argued that the genetic similarity to Neanderthals may indeed be the result of both structure and interbreeding, as opposed to just one or the other.
An approximately 40,000 year old anatomically-modern human skeleton from Peștera cu Oase, Romania, was found in 2015 to have a much larger proportion of DNA matching the Neanderthal genome than seen in humans of today, and this was estimated to have resulted from an interbreeding event as few as four generations earlier. However, this hybrid Romanian population does not appear to have made a substantial contribution to the genomes of later Europeans.
In 2016, the DNA of Altai Neanderthals showed evidence of interbreeding 100 kya, and interbreeding with an earlier dispersal of humans may have occurred as early as 120 kya in places such as the Levant. Palaeontologically, the earliest human remains outside of Africa occurs at Misliya Cave 194–177 kya, and Skhul and Qafzeh 120–90 kya. This first interbreeding event has not left any trace in modern human genomes, however.
Interbreeding with Denisovans
Sequencing of the genome of a Denisovan, a distinct but related archaic hominin, from the Denisova cave in the Siberian Altai region has shown that 17% of its genome represents Neanderthal DNA. This Neanderthal DNA present in the Denisovan genome more closely resembled that found in the genome of a 120,000-year-old Neanderthal bone found in the same cave than that of Neanderthals from the Vindija cave in Croatia or the Mezmaiskaya cave in the Caucasus, suggesting that the gene flow came from a local interbreeding. However, the complete genome sequencing of DNA from a 90,000-year-old bone fragment, Denisova 11, showed it to have belonged to a Denisovan-Neanderthal hybrid whose father was a typical Denisovan with the Altai Neanderthal component dating to an interbreeding more than 300 generations earlier, but the specimen's mother was a Neanderthal belonging to a population more closely related to the Vindija Neanderthal than to the sequenced Altai Neanderthal genome. This suggests mobility or turnover among the distinct Neanderthal populations.
The mtDNA phylogeny of the Neanderthals, Denisovans, and modern humans produces a puzzling picture. Based on mtDNA sequences of Neanderthals from the Vindija and Altai sites, the most recent common ancestor of modern humans and Neanderthals lived approximately 440,000 years ago, while that between Neanderthals and Denisovans is far back as 830,000 years. This contrasts with nuclear genome analysis, in which the Neanderthals and Denisovans are sister groups. The mtDNA of the 400,000-year-old Sima de los Huesos sample, thought to be ancestral to Western European Neanderthals, is much closer to Denisovan mtDNA. Paabo et al. suggest several alternative explanations for this, including genetic substructure within the populations, introgression of mtDNA from one group to another, or incorrect taxonomic placement of the Sima de los Huesos hominins. Peyrégne et al. again suggested genetic substructure or introgression as possible explanations for an approximately 124,000-year-old German Neanderthal specimen with mtDNA that diverged from that of other Neanderthal specimens (except for Sima de Huesos) about 270,000 years ago, while its genomic DNA was consistent with divergence less than 150,000 years ago.
According to a 2014 study by Thomas Higham and colleagues of organic samples from European sites, Neanderthals died out in Europe between 41,000 and 39,000 years ago.[d] New dating in Iberia, where Neanderthal dates as late as 24,000 years had been reported before, now suggests evidence of Neanderthal survival in the peninsula after 42,000 years ago is almost non-existent.
Anatomically modern humans arrived in Mediterranean Europe between 45,000 and 43,000 years ago, so the two different human populations shared Europe for several thousand years. The exact nature of biological and cultural interaction between Neanderthals and other human groups is contested.
Possible scenarios for the extinction of the Neanderthals are:
- Neanderthals became extinct because of climate change or interaction with modern humans and were replaced by modern humans moving into their habitat between 45,000 and 40,000 years ago. Jared Diamond has suggested a scenario of violent conflict and displacement.
- Neanderthals bred with modern humans and disappeared through absorption (interbreeding theory).
- Volcanic catastrophe (Campanian Ignimbrite Eruption)
- Neanderthals had weaker immune systems than humans due to their low genetic diversity, and were more vulnerable to disease epidemics.
- Their low population left them vulnerable to any environmental change, and a small drop in survival or fertility rates could have quickly led to their extinction.
About 55,000 years ago, the climate began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of decades. Neanderthal bodies were well-suited for survival in a cold climate—their stocky chests and limbs stored body heat better than the Cro-Magnons. Neanderthals died out in Europe between 41,000 and 39,000 years ago, coinciding with the start of a very cold period.
Raw material sourcing and the examination of faunal remains found in the southern Caucasus suggest that modern humans may have had a survival advantage, being able to use social networks to acquire resources from a greater area. In both the Late Middle Palaeolithic and Early Upper Palaeolithic more than 95% of stone artifacts were drawn from local material, suggesting Neanderthals restricted themselves to more local sources.
Coexistence with modern humans
The exact nature of biological and cultural interaction between Neanderthals and other human groups has been contested. The dating of the period of overlap has been repeatedly revised. In 2011, this period of coexistence was extended when what were previously thought to be Neanderthal baby teeth, unearthed in 1964 from the Grotta del Cavallo in Italy, were identified as the oldest modern human remains discovered anywhere in Europe, dating from between 43,000 and 45,000 years ago. At that time, Neanderthal samples from Iberia were thought to have dated as recently as 24,000 years ago, but a 2014 study of Neanderthal bones and tools led to a redating of these samples, and indicated that Neanderthals died out in Europe between 41,000 and 39,000 years ago, with the two different human populations sharing Europe only for about 5,000 years. However, a 2019 reanalysis of skull fragments from southern Greece assumed to have belonged to Neanderthals concluded that they belonged to a modern human, and dated them to 210,000 years ago, vastly extending the period of potential overlap in Europe between the two groups.
Neanderthals inhabited that continent long before the arrival of modern humans. These modern humans may have introduced a disease that contributed to the extinction of Neanderthals, and that may be added to other recent explanations for their extinction. When Neanderthal ancestors left Africa potentially as early as over 800,000 years ago they adapted to the pathogens in their European environment, unlike modern humans, who adapted to African pathogens. This transcontinental movement is known as the Out of Africa model. If contact between humans and Neanderthals occurred in Europe and Asia the first contact may have been devastating to the Neanderthal population, because they would have had little, if any, immunity to the African pathogens. More recent historical events in Eurasia and the Americas show a similar pattern, whereby the unintentional introduction of viral or bacterial pathogens to unprepared populations has led to mass mortality and local population extinction. The most well-known example of this is the arrival of Christopher Columbus to the New World, which brought and introduced foreign diseases when he and his crew arrived to a native population who had no immunity.
History of research
Neanderthal fossils were first discovered in 1829 in the Engis caves (the partial skull dubbed Engis 2), in present-day Belgium by Philippe-Charles Schmerling and the Gibraltar 1 skull in 1848 in the Forbes' Quarry, Gibraltar. These finds were not, at the time, recognized as representing an archaic form of humans.
The first discovery which was recognized as representing an archaic form of humans was made in August 1856, three years before Charles Darwin's On the Origin of Species was published. This was the discovery of the type specimen, Neanderthal 1, in a limestone quarry (Feldhofer Cave), located in Neandertal Valley in the German Rhineland, about 12 km (7 mi) east of Düsseldorf). The find consisted of a skull cap, two femora, three bones of the right arm, two of the left arm, parts of the left ilium, fragments of a scapula, and ribs. The workers who recovered the objects originally thought them to be the remains of a cave bear. However, they eventually gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen.
To date, the bones of over 400 Neanderthals have been found.
- 1829: A damaged skull of a Neanderthal child, Engis 2, is discovered in Engis, Netherlands (now Belgium).
- 1848: A female Neanderthal skull, Gibraltar 1, is found in Forbes' Quarry, Gibraltar, but its importance is not recognised.
- 1856: Limestone miners discover the Neanderthal-type specimen, Neanderthal 1, in Neandertal, western Prussia (now Germany).
- 1864: William King is the first to recognise Neanderthal 1 as belonging to a separate species, for which he gives the scientific name Homo neanderthalensis. He then changed his mind on placing it in the genus Homo, arguing that the upper skull was different enough to warrant a separate genus since, to him, it had likely been "incapable of moral and theistic conceptions."
- 1880: The mandible of a Neanderthal child is discovered in a secure context in Šipka cave, in the Austro-Hungarian Empire (now the Czech Republic), associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals.
- 1886: Two well-preserved Neanderthal skeletons are found at Spy, Belgium, making the hypothesis that Neanderthal 1 was only a diseased modern human difficult to sustain.
- 1899: Sand excavation workers find hundreds of fragmentary Neanderthal remains representing at least 12 and likely as much as 70 individuals on a hill in Krapina, in the Austro-Hungarian Empire (now Croatia).
- 1908: A very well preserved Neanderthal, La Chapelle-aux-Saints 1, is found in its eponymous site in France, said by the excavators to be a burial, a claim still heatedly contested. For historical reasons it remains the most famous Neanderthal skeleton.:15
- 1912: Marcellin Boule publishes his now discredited influential study of Neanderthal skeletal morphology based on La Chapelle-aux-Saints 1.
- 1953–1957: Ten Neanderthal skeletons are excavated in Shanidar Cave, Iraqi Kurdistan, by Ralph Solecki and colleagues.
- 1975: Erik Trinkaus's study of Neanderthal feet strongly argues that Neanderthals walked like modern humans.
- 1981: The site of Bontnewydd, Wales yielded an early Neanderthal tooth, the most north-western Neanderthal remain ever.
- 1987: Israeli Neanderthal Kebara 2 is dated (by TL and ESR) to 60,000 BP, thus later than the Israeli anatomically modern humans dated to 90,000 and 80,000 BP at Qafzeh and Skhul.
- 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley.
- 2005: The Max Planck Institute for Evolutionary Anthropology and associated institutions launch the Neanderthal genome project to sequence the Neanderthal nuclear genome.
- 2010: Discovery of Neanderthal admixture in the genome of modern populations.
- 2014: A comprehensive dating of Neanderthal bones and tools from hundreds of sites in Europe dates the disappearance of Neanderthals to 41,000 and 39,000 years ago.
- 2018: Report on the complete genomic sequence of Denisova 11, a first generation of Neanderthal-Denisovan hybrid.
- 2019: 257 hominin footprints were revealed by archaeologists at Le Rozel (Normandy, France). This finding, the vast majority of which consists of young footprints, has proved that Neanderthals lived in this area 80,000 years ago.
Notable European Neanderthals
Remains of more than 300 European Neanderthals have been found. For the most important, see List of human evolution fossils.
- Neanderthal 1: The first human bones recognised as showing a non-modern anatomy. Discovered in 1856 in a limestone quarry at the Feldhofer grotto in Neanderthal, Western Prussia (now Germany), they consist of a skull cap, the two femora, the three right arm bones, two left arm bones, the ilium, and fragments of a scapula and ribs.
- La Chapelle-aux-Saints 1: Called the Old Man, a fossilised skeleton discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth long before death, leading some to suggest he was cared for by others.
- La Ferrassie 1: A fossilised skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth.
- Le Moustier 1: One of the rare nearly complete Neanderthal skeletons to be discovered, it was excavated by a German team in 1908, at Peyzac-le-Moustier, France. Sold to a Berlin museum, the post cranial skeleton was bombed and mostly destroyed in 1945, and parts of the mid face were lost sometime after then. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a less developed brow ridge and occipital bun than seen in other Neanderthals. The Mousterian tool techno-complex is named after its discovery site.
Notable Southwest Asian Neanderthals
Remains of more than 70 Southwest Asian Neanderthals have been found. For a complete list see List of Southwest Asian Neanderthals.
- Shanidar 1 to 10: Eight Neanderthals and two pre-Neanderthals (Shanidar 2 and 4) were discovered in the Zagros Mountains in Iraqi Kurdistan. One of the skeletons, Shanidar 4, was once thought to have been buried with flowers, a theory no longer accepted. To Paul B. Pettitt the "deliberate placement of flowers has now been convincingly eliminated", since "[a] recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".
- Amud 1: A male adult Neanderthal, dated to roughly 55,000 BP, and one of several found in a cave at Nahal Amud, Israel. At 178 cm (70 in), it is the tallest known Neanderthal. It also has the largest cranial capacity of all extinct hominins: 1,736 cm3.
- Kebara 2: A male adult post-cranial skeleton, dated to roughly 60,000 BP, that was discovered in 1983 in Kebara Cave, Israel. It has been studied extensively, for its hyoid, ribcage, and pelvis are much better preserved than in all other Neanderthal specimens.
Notable Central Asian Neanderthal
- Teshik-Tash 1: The skeleton of an 8–11-year-old child discovered in Uzbekistan by Okladnikov in 1938. This is the only fairly complete skeleton discovered to the east of Iraq. Okladnikov claimed it was a deliberate burial, but this is debated.
Mixed with H. heidelbergensis traits
- 350–200 ka: Pontnewydd 225 ka ago.
- 200–135 ka: Atapuerca, Vértesszőlős, Ehringsdorf, Casal de' Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade
H. neanderthalensis fossils
- 130–50 ka: Krapina, Saccopastore skulls, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun 120–100±5 ka, Shanidar 1 to 9 80–60 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Régourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 53±8 ka, Teshik-Tash.
- In radiocarbon range, > 50 ka: Le Moustier, Feldhofer, La Quina, l'Hortus, Kulna, Šipka, Saint Césaire, Bacho Kiro, El Castillo, Bañolas, El Sidrón (48±3 cal ka), Arcy-sur-Cure, Châtelperron, Figueira Brava, Mezmaiskaya (41±1 cal ka), Zafarraya, Vindija, Velika Pećina.
H. s. sapiens with traits reminiscent of Neanderthals
- < 35 Peștera cu Oase 37-42 ka, Mladeč 31 ka, Peștera Muierii 30 ka (n/s), Lagar Velho 1 24.5 ka.
In popular culture
Neanderthals have been portrayed in popular culture including appearances in literature, visual media and comedy. Early 20th century artistic interpretations often presented Neanderthals as beastly creatures, emphasising hairiness and a rough, dark complexion.
- The German noun is cognate with English dale. The German /t/ phoneme was frequently spelled th throughout the 15th to 19th centuries; Tal became standardized with the German spelling reform of 1901, thus the German name Neandertal for both the valley and species.
- There are modern humans with noses as wide as those of Neanderthals and modern humans with similar nose lengths, but none with both Neanderthal nose width and nose length.
- Homo floresiensis originated in an unknown location from unknown ancestors and reached remote parts of Indonesia. Homo erectus spread from Africa to western Asia, then east Asia and Indonesia; its presence in Europe is uncertain, but it gave rise to Homo antecessor, found in Spain. Homo heidelbergensis originated from Homo erectus in an unknown location and dispersed across Africa, southern Asia and southern Europe (other scientists interpret fossils, here named heidelbergensis, as late erectus). Homo sapiens sapiens spread from Africa to western Asia and then to Europe and southern Asia, eventually reaching Australia and the Americas. In addition to Neanderthals and Denisovans, a third gene flow of archaic Africa origin is indicated at the right.
- Higham et al did not study samples from sites outside Europe and they stated that further work was required to rule out later survival at Gorhams Cave, Gibraltar.
- Howell, F. Clark (1957). "The evolutionary significance of variation and varieties of 'Neanderthal' man". The Quarterly Review of Biology. 32 (4): 330–47. doi:10.1086/401978. JSTOR 2816956. PMID 13506025.(subscription required) Foley, Tim. TalkOrigins Archive. "Neanderthal or Neandertal?". 2005.
- Romeo, Luigi (1979). Ecce Homo!:A Lexicon of Man. Amsterdam: John Benjamins Publishing Company. p. 92. ISBN 978-9027220066 – via Google Books (ebook).
- Camp, C. L; Allison, H. J.; Nichols, R. H. (1964). Bibliography of Fossil Vertebrates 1954–1958. New York: The Geological Society of America, Inc. p. 556. ISBN 9780813710921 – via Google Books (preview).
- Wells, John (April 3, 2008). Longman Pronunciation Dictionary (3rd ed.). Pearson Longman. ISBN 978-1-4058-8118-0.
- Krech, Eva Maria; Stock, Eberhard; Hirschfeld, Ursula; Anders, Lutz-Christian (2009), Deutsches Aussprachewörterbuch [German Pronunciation Dictionary], Berlin, New York: Walter de Gruyter, ISBN 978-3-11-018202-6
- Duden - Das Aussprachewörterbuch: Betonung und Aussprache von über 132.000 Wörtern und Namen [Duden - The pronunciation dictionary: emphasis and pronunciation of over 132,000 words and names] (in German). Bibliographisches Institut GmbH. 2015. p. 561. ISBN 9783411911516 – via Google Books (preview).
- Pääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
- T. Higham; K. Douka; R. Wood; C.B. Ramsey; F. Brock; L. Basell; M. Camps; A. Arrizabalaga; J. Baena; C. Barroso-Ruíz; C. Bergman; C. Boitard; P. Boscato; M. Caparrós; N.J. Conard; C. Draily; A. Froment; B. Galván; P. Gambassini; A. Garcia-Moreno; S. Grimaldi; P. Haesaerts; B. Holt; M.-J. Iriarte-Chiapusso; A. Jelinek; J.F. Jordá Pardo; J.-M. Maíllo-Fernández; A. Marom; J. Maroto; M. Menéndez; L. Metz; E. Morin; A. Moroni; F. Negrino; E. Panagopoulou; M. Peresani; S. Pirson; M. de la Rasilla; J. Riel-Salvatore; A. Ronchitelli; D. Santamaria; P. Semal; L. Slimak; J. Soler; N. Soler; A. Villaluenga; R. Pinhasi; R. Jacobi (2014). "The timing and spatiotemporal patterning of Neanderthal disappearance". Nature. 512 (7514): 306–09. Bibcode:2014Natur.512..306H. doi:10.1038/nature13621. PMID 25143113.
We show that the Mousterian [the Neanderthal tool-making tradition] ended by 41,030–39,260 calibrated years BP (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time.
- T. Higham (2011). "European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies". Antiquity. 85 (327): 235–49. doi:10.1017/s0003598x00067570.
Few events of European prehistory are more important than the transition from ancient to modern humans about 40,000 years ago, a period that unfortunately lies near the limit of radiocarbon dating. This paper shows that as many as 70 per cent of the oldest radiocarbon dates in the literature may be too young, due to contamination by modern carbon.(subscription required)
- R. Pinhasi; T.F.G. Higham; L.V. Golovanova; V.B. Doronichev (2011). "Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus". Proceedings of the National Academy of Sciences USA. 108 (21): 8611–16. Bibcode:2011PNAS..108.8611P. doi:10.1073/pnas.1018938108. PMC 3102382. PMID 21555570.
The direct date of the fossil (39,700 ± 1,100 14C BP) is in good agreement with the probability distribution function, indicating at a high level of probability that Neanderthals did not survive at Mezmaiskaya Cave after 39 ka cal BP. [...] This challenges previous claims for late Neanderthal survival in the northern Caucasus. [...] Our results confirm the lack of reliably dated Neanderthal fossils younger than ≈40 ka cal BP in any other region of Western Eurasia, including the Caucasus.
- B. Galván; C.M. Hernández; C. Mallol; N. Mercier; A. Sistiaga; V. Soler (2014). "New evidence of early Neanderthal disappearance in the Iberian Peninsula". Journal of Human Evolution. 75: 16–27. doi:10.1016/j.jhevol.2014.06.002. PMID 25016565.(subscription required)
- Shaw, Ian; Jameson, Robert, eds. (1999). A Dictionary of Archaeology. Blackwell. p. 408. ISBN 978-0-631-17423-3. Retrieved August 1, 2016 – via Google Books (preview).
- Stringer, C. (1984). "Human evolution and biological adaptation in the Pleistocene". In Foley, R. (ed.). Hominid evolution and community ecology. New York: Academic Press. ISBN 978-0122619205.
- Holloway, R.L. (1985). "The poor brain of Homo sapiens neanderthalensis: see what you please". In Delson, E. (ed.). Ancestors: The hard evidence. New York: Alan R. Liss. ISBN 978-0471843764.; extending to 1,736 cm3 (105.9 cu in) in the male Amud 1.
- Amano, H.; Kikuchi, T.; Morita, Y.; Kondo, O.; Suzuki, Hiromasa; et al. (August 2015). "Virtual Reconstruction of the Neanderthal Amud 1 Cranium". American Journal of Physical Anthropology. 158 (2): 185–97. doi:10.1002/ajpa.22777. hdl:10261/123419. PMID 26249757.
- Beals, Kenneth; Smith, Courtland; Dodd, Stephen (1984). "Brain Size, Cranial Morphology, Climate, and Time Machines" (PDF). Current Anthropology. 12 (3): 301–30. doi:10.1086/203138 – via Oregon State University.
- Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1–12. PMID 9850627.
- Green, R. E.; Krause, J.; Briggs, A. W.; Maricic, T.; Stenzel, U.; Kircher, M.; Patterson, N.; Li, H.; Zhai, W.; Fritz, M. H. Y.; Hansen, N. F.; Durand, E. Y.; Malaspinas, A. S.; Jensen, J. D.; Marques-Bonet, T.; Alkan, C.; Prufer, K.; Meyer, M.; Burbano, H. A.; Good, J. M.; Schultz, R.; Aximu-Petri, A.; Butthof, A.; Hober, B.; Hoffner, B.; Siegemund, M.; Weihmann, A.; Nusbaum, C.; Lander, E. S.; Russ, C.; Novod, N.; Affourtit, J.; Egholm, M.; Verna, C.; Rudan, P.; Brajkovic, D.; Kucan, Z.; Gusic, I.; Doronichev, V. B.; Golovanova, L. V.; Lalueza-Fox, C.; de la Rasilla, M.; Fortea, J.; Rosas, A.; Schmitz, R. W.; Johnson, P. L. F.; Eichler, E. E.; Falush, D.; Birney, E.; Mullikin, J. C.; Slatkin, M.; Nielsen, R.; Kelso, J.; Lachmann, M.; Reich, D.; Paabo, S. (May 6, 2010). "A Draft Sequence of the Neandertal Genome". Science. 328 (5979): 710–722. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC 5100745. PMID 20448178.
- Sankararaman, Sriram; Mallick, Swapan; Patterson, Nick; Reich, David (May 2016). "The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans". Current Biology. 26 (9): 1241–1247. doi:10.1016/j.cub.2016.03.037. PMC 4864120. PMID 27032491.
- Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Pääbo, Svante; Patterson, Nick; Reich, David (January 29, 2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–357. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735. PMID 24476815.
- Sánchez-Quinto, Federico; Botigué, Laura R.; Civit, Sergi; Arenas, Conxita; Ávila-Arcos, María C.; Bustamante, Carlos D.; Comas, David; Lalueza-Fox, Carles; Caramelli, David (October 17, 2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLoS ONE. 7 (10): e47765. Bibcode:2012PLoSO...747765S. doi:10.1371/journal.pone.0047765. PMC 3474783. PMID 23082212.}
- Fu, Qiaomei; Li, Heng; Moorjani, Priya; Jay, Flora; Slepchenko, Sergey M.; Bondarev, Aleksei A.; Johnson, Philip L. F.; Aximu-Petri, Ayinuer; Prüfer, Kay; de Filippo, Cesare; Meyer, Matthias; Zwyns, Nicolas; Salazar-García, Domingo C.; Kuzmin, Yaroslav V.; Keates, Susan G.; Kosintsev, Pavel A.; Razhev, Dmitry I.; Richards, Michael P.; Peristov, Nikolai V.; Lachmann, Michael; Douka, Katerina; Higham, Thomas F. G.; Slatkin, Montgomery; Hublin, Jean-Jacques; Reich, David; Kelso, Janet; Viola, T. Bence; Pääbo, Svante (October 22, 2014). "Genome sequence of a 45,000-year-old modern human from western Siberia". Nature. 514 (7523): 445–449. Bibcode:2014Natur.514..445F. doi:10.1038/nature13810. PMC 4753769. PMID 25341783.
- Brahic, Catherine. "Humanity's forgotten return to Africa revealed in DNA". New Scientist.
- Kuhlwilm, Martin (February 17, 2016). "Ancient gene flow from early modern humans into Eastern Neanderthals". Nature. 530 (7591): 429–433. Bibcode:2016Natur.530..429K. doi:10.1038/nature16544. PMC 4933530. PMID 26886800.
- Krings, M.; Stone, A.; Schmitz, R. W.; Krainitzki, H.; Stoneking, M.; Pääbo, S. (July 1997). "Neandertal DNA Sequences and the Origin of Modern Humans". Cell. 90 (1): 19–30. doi:10.1016/s0092-8674(00)80310-4. PMID 9230299.
- Krings, M.; Geisert, H.; Schmitz, R. W.; Krainitzki, H.; Paabo, S. (May 11, 1999). "DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen". Proceedings of the National Academy of Sciences. 96 (10): 5581–5585. Bibcode:1999PNAS...96.5581K. doi:10.1073/pnas.96.10.5581. PMC 21903. PMID 10318927.
- Beerli, Peter; Edwards, Scott V. (January 7, 2003). "When did Neanderthals and modern humans diverge?". Evolutionary Anthropology: Issues, News, and Reviews. 11 (S1): 60–63. doi:10.1002/evan.10058.
- Vogt, Karl C (1864). Lectures on Man: His Place in Creation, and in the History of the Earth. London: Longman, Green, Longman and Roberts. pp. 302, 473 – via Internet Archive.
- King, W. (1864). "On the Neanderthal Skull, or reasons for believing it to belong to the Clydian Period and to a species different from that represented by man". Report of the British Association for the Advancement of Science, Notices and Abstracts, Newcastle-upon-Tyne, 1863: 81–82. Retrieved October 21, 2018 – via Biodiversity Heritage Library.
- Murray, John; Nasheuer, Heinz Peter; Seoighe, Cathal; McCormack, Grace P.; Williams, D. Michael & Harper, David A.T. (2015), "The Contribution of William King to the Early Development of Palaeoanthropology", Irish Journal of Earth Sciences, 33: 1–16, doi:10.3318/ijes.2015.33.1, JSTOR 10.3318/ijes.2015.33.1(subscription required)
- King, William (January 1864). "The Reputed Fossil Man of the Neanderthal" (PDF). The Quarterly Journal of Science. 1: 96 – via Bone and Stone.
- Inter alia, Boys' Life, p. 18. January 1924—via Google Books.
- The Oxford Illustrated Dictionary. Great Britain: Oxford University Press. 1976 . p. 564.
- "Neanderthal adjective – definition in British English Dictionary & Thesaurus". Cambridge Dictionaries Online. Cambridge University Press. January 8, 2013. Archived from the original on July 8, 2012. Retrieved January 22, 2013.
- "NEANDERTHAL—meaning in the Cambridge English Dictionary". Cambridge Dictionary. Cambridge University Press. Retrieved September 19, 2019.
- "Oxford Learner's Dictionaries – Find pronunciation, clear meanings and definitions of words". OxfordLearnersDictionaries.com.[dead link]
- "neanderthal_1 noun". Oxford Learner's Dictionaries. Oxford University Press. Retrieved September 19, 2019.
- Kurtén, Björn (October 10, 1995). Dance of the Tiger: A Novel of the Ice Age. University of California Press. p. xxi. ISBN 978-0-520-20277-1. Retrieved May 9, 2012 – via Google Books (preview).
- Pollet, Carl J. (September 21, 1991). "...And Etymology". Science News. 140 (12): 191. doi:10.2307/3975867. JSTOR 3975867.(subscription required)
- "Neanderthal | Define Neanderthal at Dictionary.com". Dictionary.reference.com. Retrieved January 22, 2013.
- "Neanderthal". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013.
- "Neandertal oder Neanderthal? – Was ist denn nun richtig?" [Neanderthal or Neanderthal? - What is right?]. mettmann.de. Neanderthal museum. Retrieved February 1, 2017.
Heute sollten Ortsbezeichnungen das 'Neandertal' ohne 'h' bezeichnen. Alle Namen, die sich auf den prähistorischen Menschen beziehen, führen das 'h'. [Today one should write for place names 'Neandertal' without an 'h'. All names related to the prehistoric humans keep the 'h'.]
- Tattersall, Ian; Schwartz, Jeffrey H. (1999). "Hominids and hybrids: The place of Neanderthals in human evolution". Proceedings of the National Academy of Sciences. 96 (13): 7117–19. Bibcode:1999PNAS...96.7117T. doi:10.1073/pnas.96.13.7117. JSTOR 48019. PMC 33580. PMID 10377375.
- Duarte, Cidália; Maurício, João; Pettitt, Paul B.; Souto, Pedro; Trinkaus, Erik; van der Plicht, Hans; Zilhão, João (June 22, 1999). "The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia". Proceedings of the National Academy of Sciences of the United States of America. 96 (13): 7604–7609. Bibcode:1999PNAS...96.7604D. doi:10.1073/pnas.96.13.7604. PMC 22133. PMID 10377462.
- Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–27. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594. PMID 19805257.
- Harvati, K.; Frost, S.R.; McNulty, K.P. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proc. Natl. Acad. Sci. USA. 101 (5): 1147–52. Bibcode:2004PNAS..101.1147H. doi:10.1073/pnas.0308085100. PMC 337021. PMID 14745010.
- "L'homme fossile de La Chapelle-aux-Saints – full text: Volume VI (pp. 11–172), Volume VII (pp. 21–56), Volume VIII (pp. 1–70), 1911–13". Royal College of Surgeons of England. Retrieved July 26, 2016 – via Internet Archive.
- Shermer, Michael (August 1, 2010). "Our Neandertal Brethren: Why They Were Not a Separate Species". Scientific American. 303 (2): 34. doi:10.1038/scientificamerican0810-34. PMID 20684369.
- Wolpoff, MH; Hawks, J; Caspari, R (2000). "Multiregional, not multiple origins" (PDF). American Journal of Physical Anthropology. 112 (1): 129–36. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. hdl:2027.42/34270. PMID 10766948 – via University of Michigan.
- Cairney, Christopher Thomas (1989). Clans and Families of Ireland and Scotland, an Ethnography of the Gael. London: McFarland. p. 14. ISBN 978-0899503622 – via Electric Scotland.
- Ko, K. W. (2016). "Hominin interbreeding and the evolution of human variation". Journal of Biological Research-Thessaloniki. doi:10.1186/s40709-016-0054-7. PMC 4947341. PMID 27429943.
- D. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5): 485–508. doi:10.1006/jhev.1998.0214. PMID 9614635.
- Papagianni, Dmitra; Morse, Michael (2013). The Neanderthals Rediscovered. Thames & Hudson. ISBN 978-0-500-05177-1.
- Stringer, C. (2012). "The status of Homo heidelbergensis (Schoetensack 1908)". Evolutionary Anthropology. 21 (3): 101–107. doi:10.1002/evan.21311. PMID 22718477.
- Stringer, C.; Gamble, C. (1993). In Search of the Neanderthals. London: Thames and Hudson. ISBN 978-0500050705.
- B. Vandermeersch; M.D. Garralda (2011). "Neanderthal Geographical and Chronological Variation". In S. Condemi; G.-C. Weniger (eds.). Continuity and Discontinuity in the Peopling of Europe. Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands. pp. 113–25. doi:10.1007/978-94-007-0492-3_10.
- N.J. Conard; J. Richter, eds. (2011). "2". Neanderthal Lifeways, Subsistence and Technology. Vertebrate Paleobiology and Paleoanthropology. 19. Springer. pp. 7–14. doi:10.1007/978-94-007-0415-2_2. ISBN 978-9400704145.
- Endicott, P.; Ho, S. Y. W.; Stringer, C. (2010). "Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins" (PDF). Journal of Human Evolution. 59 (1): 87–95. doi:10.1016/j.jhevol.2010.04.005. PMID 20510437.
- Meyer, M.; Arsuaga, J.; de Filippo, C.; Nagel, S. (2016). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature. 531: 504–507. doi:10.1038/nature17405.
- Briggs, A. W.; Good, J. M.; Green, R. E. (2009). "Targeted retrieval and analysis of five Neandertal mtDNA genomes" (PDF). Science. 325 (5, 938): 318–321. doi:10.1126/science.1174462. PMID 19608918.
- Green, R. E.; Malaspinas, A. S.; Krause, J.; Briggs, A. W. (2008). "A complete Neandertal mitochondrial genome sequence determined by high-throughput sequencing". Cell. 134 (3): 416–426. doi:10.1016/j.cell.2008.06.021. PMC 2602844. PMID 18692465.
- Gómez-Robles, A. (2019). "Dental evolutionary rates and its implications for the Neanderthal–modern human divergence". Science Advances. 5 (5): eaaw1268. doi:10.1126/sciadv.aaw1268.
- Sawyer, S.; Renaud, G.; Viola, B.; Hublin, J. J. (2015). "Nuclear and mitochondrial DNA sequences from two Denisovan individuals". Proceedings of the National Academy of Sciences. 112 (51): 15, 696–15, 700. doi:10.1073/pnas.1519905112. PMC 4697428. PMID 26630009.
- K. Prüfer; et al. (2014). "The complete genome sequence of a Neanderthal from the Altai Mountains". Nature. 505 (7481): 43–49. Bibcode:2014Natur.505...43P. doi:10.1038/nature12886. PMC 4031459. PMID 24352235.
- Rogers, A. R.; Bohlender, R. J.; Huff, C. D. (2017). "Early history of Neanderthals and Denisovans". Proceedings of the National Academy of Sciences. 114 (37): 9, 859–9, 863. doi:10.1073/pnas.1706426114.
- "Ancient tooth provides evidence of Neanderthal movement" (Press release). Durham University. February 11, 2008. Retrieved May 18, 2009.
- Callander, J. (2004). "Dorothy Garrod's Excavations in the Late Mousterian of Shukbah Cave in Palestine Reconsidered". roceedings of the Prehistoric Society. 70: 207–231. doi:10.1017/S0079497X00001171.
- Wade, Nicholas (October 2, 2007). "Fossil DNA Expands Neanderthal Range". The New York Times. Retrieved May 18, 2009. Ravilious, Kate (October 1, 2007). "Neandertals Ranged Much Farther East Than Thought". National Geographic Society. Retrieved May 18, 2009.
- Pavlov P, Roebroeks W, Svendsen JI (2004). "The Pleistocene colonization of northeastern Europe: a report on recent research". Journal of Human Evolution. 47 (1–2): 3–17. doi:10.1016/j.jhevol.2004.05.002. PMID 15288521. Slimak, L., et al. (2012). "Response to 'Comment on Late Mousterian Persistence near the Arctic Circle'" Science, 335 (6065), 167 doi:10.1126/science.1210211 Zwyns, N., et al. (2012). "Comment on 'Late Mousterian Persistence near the Arctic Circle'" Science, 335 (6065), 167 doi:10.1126/science.1209908
- Bocquet-Appel, J.; Degioanni, A. (2013). "Neanderthal Demographic Estimates". Current Anthropology. 54: 202–214. doi:10.1086/673725.
- Stewart, J.R.; García-Rodríguez, O.; Knul, M.V.; Sewell, L.; Montgomery, H.; Thomas, M.G.; Diekmann, Y. (2019). "Palaeoecological and genetic evidence for Neanderthal power locomotion as an adaptation to a woodland environment". Quaternary Science Reviews. 217: 310–315. Bibcode:2019QSRv..217..310S. doi:10.1016/j.quascirev.2018.12.023.
- Peyrégne, Stéphane; et al. (2019). "Nuclear DNA from two early Neandertals reveals 80,000 years of genetic continuity in Europe". Science Advances. 5 (6): eaaw5873. Bibcode:2019SciA....5.5873P. doi:10.1126/sciadv.aaw5873. PMC 6594762. PMID 31249872.
- Fabrizio Mafessoni, Kay Prüfer, "Better support for a small effective population size of Neandertals and a long shared history of Neandertals and Denisovans", PNAS 114 (48), November 28, 2017, E10256–E10257, doi:10.1073/pnas.1716918114.
- Lalueza-Fox, C.; Sampietro, M. L.; Caramelli, D.; Puder, Y. (2013). "Neandertal evolutionary genetics: mitochondrial DNA data from the iberian peninsula". Molecular Biology and Evolution. 22 (4): 1077–1081. doi:10.1093/molbev/msi094. PMID 15689531.
- Fabre, V.; Condemi, S.; Degioanni, A. (2009). "Genetic Evidence of Geographical Groups among Neanderthals". PLOS ONE. 4 (4): e5151. Bibcode:2009PLoSO...4.5151F. doi:10.1371/journal.pone.0005151. PMC 2664900. PMID 19367332.
- Mellars, P.; French, J. C. (2011). "Tenfold population increase in Western Europe at the Neandertal-to-modern human transition". Science. 333 (6042): 623–627. Bibcode:2011Sci...333..623M. doi:10.1126/science.1206930. PMID 21798948.
- Sánchez-Quinto, F.; Lalueza-Fox, C. (2015). "Almost 20 years of Neanderthal palaeogenetics: adaptation, admixture, diversity, demography and extinction". Philosophical Transactions of the Royal Society B. 370 (1, 660): 20130374. doi:10.1098/rstb.2013.0374. PMC 4275882. PMID 25487326.
- Ríos, L.; Kivell, T. L.; Lalueza-Fox, C.; Estalrrich, A. (2019). "Skeletal Anomalies in The Neandertal Family of El Sidrón (Spain) Support A Role of Inbreeding in Neandertal Extinction". Scientific Reports. 9 (1, 697). doi:10.1038/s41598-019-38571-1. PMC 6368597. PMID 30737446.
- Trinkaus, E. (1995). "Neanderthal mortality patterns". Journal of Archaeological Science. 22 (1): 121–142. doi:10.1016/S0305-4403(95)80170-7.
- Pettitt, R. B. (2000). "Neanderthal Lifecycles: Developmental and Social Phases in the Lives of the Last Archaics". World Archaeology. 31 (3): 351–366. doi:10.1080/00438240009696926. JSTOR 125106. PMID 16475295.
- Puech, Pierre-François; Puech, Bernard. "L'Homme de Neanderthal par Paul Dardé : L'Homme Primitif" [Neanderthal Man by Paul Dardé: Primitive Man]. Academia.edu (in French). Retrieved July 5, 2017.
- Haeusler, M.; Trinkaus, E.; Fornai, C.; Müller, J.; Bonneau, N.; Boeni, T.; Frater, N. (2019). "Morphology, pathology, and the vertebral posture of the La Chapelle-aux-Saints Neandertal". Proceedings of the National Academy of Sciences. 116 (11): 4, 923–4, 927. doi:10.1073/pnas.1820745116.
- Gunz, Philipp; Tilot, Amanda K.; Wittfeld, Katharina; Teumer, Alexander; Shapland, Chin Yang; van Erp, Theo G.M.; Dannemann, Michael; Vernot, Benjamin; Neubauer, Simon; Guadalupe, Tulio; Fernández, Guillén; Brunner, Han G.; Enard, Wolfgang; Fallon, James; Hosten, Norbert; Völker, Uwe; Profico, Antonio; Di Vincenzo, Fabio; Manzi, Giorgio; Kelso, Janet; St. Pourcain, Beate; Hublin, Jean-Jacques; Franke, Barbara; Pääbo, Svante; Macciardi, Fabio; Grabe, Hans J.; Fisher, Simon E. (January 2019). "Neandertal Introgression Sheds Light on Modern Human Endocranial Globularity". Current Biology. 29 (1): 120–127.e5. doi:10.1016/j.cub.2018.10.065. PMC 6380688. PMID 30554901.
- Froehle, A. W.; Churchill, S. E. (2009). "Energetic Competition Between Neandertals and Anatomically Modern Humans" (PDF). PaleoAnthropology: 96–116.
- Weaver, T. D. (2009). "The meaning of Neandertal skeletal morphology". Proceedings of the National Academy of Sciences. 106 (38): 16, 028–16, 033. doi:10.1073/pnas.0903864106.
- Holliday, T. W. (1997). "Postcranial evidence of cold adaptation in European Neandertals". American Journal of Physiological Anthropology. 104 (2): 245–258. doi:10.1002/(SICI)1096-8644(199710)104:2<245::AID-AJPA10>3.0.CO;2-#. PMID 9386830.
- de Azevedo, S.; González, M. F.; Cintas, C.; Ramallo, V.; Quinto-Sánchez, M.; Márquez, F.; Hünemeier, T.; Paschetta, C.; Ruderman, A.; Navarro, P.; Pazos, B. A.; Silva de Cerqueira, C. C.; Velan, O.; Ramírez-Rozzi, F.; Calvo, N.; Castro, H. G.; Paz, R. R.; González-José, R. (November 21, 2017). "Nasal airflow simulations suggest convergent adaptation in Neanderthals and modern humans". Proceedings of the National Academy of Sciences. 114 (47): 12442–12447. doi:10.1073/pnas.1703790114. PMC 5703271. PMID 29087302.
- Rae TC, Koppe T, Stringer CB (February 2011). "The Neanderthal face is not cold adapted". Journal of Human Evolution. 60 (2): 234–9. doi:10.1016/j.jhevol.2010.10.003. PMID 21183202.
- Gray, Richard (February 15, 2017). "Your face is probably more primitive than a Neanderthal's". BBC News.
- Pearce, E.; Stringer, C.; Dunbar, R. I. M. (2013). "New insights into differences in brain organization between Neanderthals and anatomically modern humans". Proceedings of the Royal Society B. 280 (1, 758): 20130168. doi:10.1098/rspb.2013.0168. PMC 3619466. PMID 23486442.
- Kochiyama, T.; Ogihara, N.; Tanabe, H. C. (2018). "Reconstructing the Neanderthal brain using computational anatomy". Scientific Reports. 8 (6296). doi:10.1038/s41598-018-24331-0. PMC 5919901. PMID 29700382.
- Peña-Melián, A.; Rosas, A.; García-Tabernero, A.; Bastir, M.; de la Rasilla, M. (2011). "Paleoneurology of Two New Neandertal Occipitals from El Sidrón (Asturias, Spain) in the Context of Homo Endocranial Evolution". The Anatomical Record. 294 (8): 1, 370–1, 381. doi:10.1002/ar.21427. PMID 21714107.
- Ponce de León, M. S.; Golovanova, L.; Doronichev, V. (2008). "Neanderthal brain size at birth provides insights into the evolution of human life history". Proceedings of the National Academy of Sciences. 105 (37): 13, 764–13, 768. doi:10.1073/pnas.0803917105.
- Lalueza-Fox, C.; Rompler, H.; Caramelli, D.; Staubert, C.; Catalano, G.; Hughes, D.; Rohland, N.; Pilli, E.; Longo, L.; Condemi, S.; de la Rasilla, M.; Fortea, J.; Rosas, A.; Stoneking, M.; Schoneberg, T.; Bertranpetit, J.; Hofreiter, M. (November 30, 2007). "A Melanocortin 1 Receptor Allele Suggests Varying Pigmentation Among Neanderthals". Science. 318 (5855): 1453–1455. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID 17962522.
- Ding, Qiliang; Hu, Ya; Xu, Shuhua; Wang, Chuan-Chao; Li, Hui; Zhang, Ruyue; Yan, Shi; Wang, Jiucun; Jin, Li (August 2014). "Neanderthal Origin of the Haplotypes Carrying the Functional Variant Val92Met in the MC1R in Modern Humans". Molecular Biology and Evolution. 31 (8): 1994–2003. doi:10.1093/molbev/msu180. PMID 24916031.
- Dannemann, Michael; Kelso, Janet (October 2017). "The Contribution of Neanderthals to Phenotypic Variation in Modern Humans". The American Journal of Human Genetics. 101 (4): 578–589. doi:10.1016/j.ajhg.2017.09.010. PMC 5630192. PMID 28985494.
- Cerqueira, C. C.; Piaxão-Côrtes, V. R.; Zambra, F. M. B.; Hünemeier, T.; Bortolini, M. (2012). "Predicting homo pigmentation phenotype through genomic data: From neanderthal to James Watson". American Journal of Human Biology. 24 (5): 705–709. doi:10.1002/ajhb.22263.
- Zanolli, Clément; Martinón-Torres, María; Bernardini, Federico; Boschian, Giovanni; Coppa, Alfredo; Dreossi, Diego; Mancini, Lucia; Martínez de Pinillos, Marina; Martín-Francés, Laura; Bermúdez de Castro, José María; Tozzi, Carlo; Tuniz, Claudio; Macchiarelli, Roberto (October 3, 2018). "The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment". PLOS ONE. Plos. 13 (10): e0189773. Bibcode:2018PLoSO..1389773Z. doi:10.1371/journal.pone.0189773. PMC 6169847. PMID 30281595.
- "Cro-Magnons Conquered Europe, but Left Neanderthals Alone". PLoS Biology. 2 (12): e449. November 30, 2004. doi:10.1371/journal.pbio.0020449. PMC 532398.
- Venner, S. J. (2018). A New Estimate for Neanderthal Energy Expenditure (MA). CUNY Academic Works.
- Smith, T. M.; Tafforeau, P.; Reid, D. J. (2010). "Dental evidence for ontogenetic differences between modern humans and Neanderthals". Proceedings of the National Academy of Sciences. 107 (49): 20923–20928. Bibcode:2010PNAS..10720923S. doi:10.1073/pnas.1010906107. PMC 3000267. PMID 21078988.
- Guatelli-Steinberg, D. (2009). "Recent studies of dental development in Neandertals: Implications for Neandertal life histories". Evolutionary Biology. 18 (1): 9–20. doi:10.1002/evan.20190.
- Nakahashi, W. (March 2017). "The effect of trauma on Neanderthal culture: A mathematical analysis". Homo. 68 (2): 83–100. doi:10.1016/j.jchb.2017.02.001. PMID 28238406.
- Beier, Judith; Anthes, Nils; Wahl, Joachim; Harvati, Katerina (November 14, 2018). "Similar cranial trauma prevalence among Neanderthals and Upper Palaeolithic modern humans". Nature. 563 (7733): 686–690. Bibcode:2018Natur.563..686B. doi:10.1038/s41586-018-0696-8. PMID 30429606.
- Trinkaus, E.; Villotte, S (2017). "External auditory exostoses and hearing loss in the Shanidar 1 Neandertal". PLoS One. 12 (10): e0186684. doi:10.1371/journal.pone.0186684. PMC 5650169. PMID 29053746.
- Crubézy, E.; Trinkaus, E. (1992). "Shanidar 1: a case of hyperostotic disease (DISH) in the middle Paleolithic". American Journal of Physical Anthropology. 89 (4): 411–420. doi:10.1002/ajpa.1330890402. PMID 1463085.
- Houldcroft, C. J.; Underdown, S. J. (2016). "Neanderthal genomics suggests a pleistocene time frame for the first epidemiologic transition". American Journal of Physical Anthropology. 160 (3): 379–388. doi:10.1002/ajpa.22985.
- Tattersall, Ian (2015). The Strange Case of the Rickety Cossack and other Cautionary Tales from Human Evolution. Palgrave Macmillan. p. 202. ISBN 978-1-137-27889-0.
- Hayden, B. (2012). "Neandertal social structure?". Oxford Journal of Archaeology. 31 (1): 1–26. doi:10.1111/j.1468-0092.2011.00376.x.
- Farizy, C. (1994). "Spatial patterning of Middle Paleolithic sites". Journal of Anthropological Archaeology. 13 (2): 153–160. doi:10.1006/jaar.1994.1010.
- Zollikofer, C. P. E.; Ponce de Leon, M. S.; Vandermeersch, B.; Leveque, F. (April 23, 2002). "Evidence for interpersonal violence in the St. Cesaire Neanderthal". Proceedings of the National Academy of Sciences. 99 (9): 6444–6448. Bibcode:2002PNAS...99.6444Z. doi:10.1073/pnas.082111899. PMC 122968. PMID 11972028.
- Churchill, Steven E.; Franciscus, Robert G.; McKean-Peraza, Hilary A.; Daniel, Julie A.; Warren, Brittany R. (August 2009). "Shanidar 3 Neandertal rib puncture wound and paleolithic weaponry". Journal of Human Evolution. 57 (2): 163–178. doi:10.1016/j.jhevol.2009.05.010. PMID 19615713.
- Bocherens, Hervé; Drucker, Dorothée G.; Billiou, Daniel; Patou-Mathis, Marylène; Vandermeersch, Bernard (2005). "Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: Review and use of a multi-source mixing model". Journal of Human Evolution. 49 (1): 71–87. doi:10.1016/j.jhevol.2005.03.003. PMID 15869783.
- Lichfield, John (September 30, 2006). "French dig up Neanderthal 'butcher's shop'". The New Zealand Herald.
- Richards, Michael P.; Pettitt, Paul B.; Trinkaus, Erik; Smith, Fred H.; Paunović, Maja; Karavanić, Ivor (2000). "Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes". Proceedings of the National Academy of Sciences. 97 (13): 7663–66. Bibcode:2000pnas...97.7663r. doi:10.1073/pnas.120178997. JSTOR 122870. PMC 16602. PMID 10852955.
- Fiorenza, Luca; Benazzi, Stefano; Tausch, Jeremy; Kullmer, Ottmar; Bromage, Timothy G.; Schrenk, Friedemann (2011). Rosenberg, Karen (ed.). "Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation". PLOS ONE. 6 (3): e14769. Bibcode:2011PLoSO...614769F. doi:10.1371/journal.pone.0014769. PMC 3060801. PMID 21445243.
- Jaouen, Klervia; et al. (February 19, 2019). "Exceptionally high δ15N values in collagen single amino acids confirm Neandertals as high-trophic level carnivores". Proceedings of the National Academy of Sciences of the United States of America. 116 (11): 4928–4933. doi:10.1073/pnas.1814087116. PMC 6421459. PMID 30782806.
- Yika, Bob (February 19, 2019). "Isotopes found in bones suggest Neanderthals were fresh meat eaters". Phys.org. Retrieved February 19, 2019.
- Max Planck Institute for Evolutionary Anthropology (February 19, 2019). "Neanderthals' main food source was definitely meat – Isotope analyses performed on single amino acids in Neanderthals' collagen samples shed new light on their debated diet". Science Daily. Retrieved February 21, 2019.
- Weyrich, Laura S.; Duchene, Sebastian; Soubrier, Julien; Arriola, Luis; Llamas, Bastien; Breen, James; Morris, Alan G.; Alt, Kurt W.; Caramelli, David; Dresely, Veit; Farrell, Milly; Farrer, Andrew G.; Francken, Michael; Gully, Neville; Haak, Wolfgang; Hardy, Karen; Harvati, Katerina; Held, Petra; Holmes, Edward C.; Kaidonis, John; Lalueza-Fox, Carles; de la Rasilla, Marco; Rosas, Antonio; Semal, Patrick; Soltysiak, Arkadiusz; Townsend, Grant; Usai, Donatella; Wahl, Joachim; Huson, Daniel H.; et al. (2017). "Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus". Nature. 544 (7650): 357–361. Bibcode:2017Natur.544..357W. doi:10.1038/nature21674. hdl:10261/152016. PMID 28273061.
- Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2010). "Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)". Proceedings of the National Academy of Sciences. 108 (2): 486–91. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMC 3021051. PMID 21187393.
- Shipley, G. P.; Kindscher, K. (2016). "Evidence for the Paleoethnobotany of the Neanderthal: A Review of the Literature". Scientifica. 2016: 1–12. doi:10.1155/2016/8927654. PMC 5098096. PMID 27843675.
- Sistiaga, Ainara; et al. (June 25, 2014). "The Neanderthal Meal: A New Perspective Using Faecal Biomarkers". PLOS ONE. 9 (6): e101045. Bibcode:2014PLoSO...9j1045S. doi:10.1371/journal.pone.0101045. PMC 4071062. PMID 24963925.
- Lev, E.; Kislev, M. E.; Bar-Yosef, O. (2005). "Mousterian vegetal food in Kebara Cave, Mt. Carmel". Journal of Archaeological Science. 32 (3): 475–484. doi:10.1016/j.jas.2004.11.006.
- Madella, M.; Jones, M. K.; Goldberg, P.; Goren, Y.; Hovers, E. (2002). "The Exploitation of Plant Resources by Neanderthals in Amud Cave (Israel): The Evidence from Phytolith Studies". Journal of Archaeological Science. 29 (7): 703–719. doi:10.1006/jasc.2001.0743.
- Henry, A. G.; Brooks, A. S.; Piperno, D. R. (2011). "Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)". Proceedings of the National Academy of Sciences. 108 (2): 486–491. Bibcode:2011PNAS..108..486H. doi:10.1073/pnas.1016868108. PMID 21187393.
- Krief, S.; Daujeard, C.; Moncel, M.; Lamon, N.; Reynolds, V. (2015). "Flavouring food: the contribution of chimpanzee behaviour to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets". Antiquity. 89 (344): 464–471. doi:10.15184/aqy.2014.7.
- Daugeard, C. (2008). "Exploitation du milieu animal par les Néandertaliens dans le Sud-Est de la France" [Exploitation of the environment by Neanderthals in the southeast of France]. Bulletin de la Société préhistorique française (in French). 106 (4): 818–819.
- Hardy, K.; Buckley, S.; Collins, M. J.; Estalrrich, A. (2012). "Neanderthal Medics? Evidence for Food, Cooking, and Medicinal Plants Entrapped in Dental Calculus". The Science of Nature. 99 (8): 617–626. Bibcode:2012NW.....99..617H. doi:10.1007/s00114-012-0942-0. PMID 22806252.
- Buckley, S.; Hardy, K.; Huffman, M. (2013). "Neanderthal Self-Medication in Context". Antiquity. 87 (337): 873–878. doi:10.1017/S0003598X00049528.
- Rougier, H.; Crevecoeur, I.; Beauval, C. (2016). "Neandertal cannibalism and Neandertal bones used as tools in Northern Europe". Scientific Reports. 6 (29, 005).
- Rosas, A.; Bastir, M.; Martínez-Maza, C. (2006). "Paleobiology and comparative morphology of a late Neandertal sample from El Sidrón, Asturias, Spain". Proceedings of the National Academy of Sciences. 103 (51): 19, 266–19, 271. doi:10.1073/pnas.0609662104.
- Defleur, A.; White, T.; Valensi, P.; Slimak, L. (1999). "Neanderthal cannibalism at Moula-Guercy, Ardèche, France". Science. 286 (5, 437): 128–131. doi:10.1126/science.286.5437.128.
- Radovčić, Davorka; Sršen, Ankica Oros; Radovčić, Jakov; Frayer, David W.; Petraglia, Michael D. (March 11, 2015). "Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina". PLOS ONE. 10 (3): e0119802. Bibcode:2015PLoSO..1019802R. doi:10.1371/journal.pone.0119802. PMC 4356571. PMID 25760648.
- "Neanderthals were not inferior to modern humans, study finds". ScienceDaily. April 30, 2014.
- E., de Lazaro (January 18, 2017). "Neanderthals Capable of Incorporating Symbolic Objects into Their Culture, Discovery Suggests". Sci News.
- C.Q. Choi (January 8, 2010). "Heavy Brows, High Art?: Newly Unearthed Painted Shells Show Neandertals Were Homo sapiens' Mental Equals". Scientific American.
- I. Sample (April 30, 2014). "Neanderthals were not less intelligent than modern humans, scientists find". The Guardian.
- N. Branan (2010). "Neandertal Symbolism: Evidence Suggests a Biological Basis for Symbolic Thought". Scientific American.
- d’Errico, Francesco; Tsvelykh, Alexander (March 29, 2017). "A decorated raven bone from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea". PLOS ONE. 12 (3): e0173435. Bibcode:2017PLoSO..1273435M. doi:10.1371/journal.pone.0173435. PMC 5371307. PMID 28355292.
- R. S. Solecki (1975). "Shanidar IV, a Neanderthal Flower Burial in Northern Iraq". Science. 190 (4217): 880–81. Bibcode:1975Sci...190..880S. doi:10.1126/science.190.4217.880.
- D.J. Sommer (1999). "The Shanidar IV 'Flower Burial': a Re-evaluation of Neanderthal Burial Ritual". Cambridge Archaeological Journal. 9 (1): 127–29. doi:10.1017/s0959774300015249.
- Paul B. Pettitt (2002). "The Neanderthal Dead, exploring mortuary variability in Middle Paleolithic Eurasia". Before Farming. 1 (4).
- Higham, T.; Jacobi, R.; Julien, M.; David, F.; Basell, L.; Wood, R.; Davies, W.; Ramsey, C. B. (2010). "Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Chatelperronian". Proceedings of the National Academy of Sciences. 107 (47): 20234–20239. doi:10.1073/pnas.1007963107. PMC 2996711. PMID 20956292.
- Mellars, P. (2010). "Neanderthal symbolism and ornament manufacture: The bursting of a bubble?". Proceedings of the National Academy of Sciences. 107 (47): 20147–20148. Bibcode:2010PNAS..10720147M. doi:10.1073/pnas.1014588107. PMC 2996706. PMID 21078972.
- J.-J. Hublin; S. Talamo; M. Julien; F. David; N. Connet; P. Bodu; B. Vandermeersch; M.P. Richards (2012). "Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian". Proceedings of the National Academy of Sciences USA. 109 (46): 18743–18748. Bibcode:2012PNAS..10918743H. doi:10.1073/pnas.1212924109. PMC 3503158. PMID 23112183.
- F. Welkera; M. Hajdinjak; S. Talamo; K. Jaouen; M. Dannemann; F. David; M. Julien; M. Meyer; J. Kelso; I. Barnes; S. Brace; P. Kamminga; R. Fischer; B.M. Kessler; J.R. Stewart; S. Pääbo; M.J. Collins; J.-J. Hublin (2016). "Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne". Proceedings of the National Academy of Sciences USA. 113 (40): 11162–67. doi:10.1073/pnas.1605834113. PMC 5056053. PMID 27638212.
- Finlayson, Clive; Brown, Kimberly; Blasco, Ruth; Rosell, Jordi; Negro, Juan José; Bortolotti, Gary R; Finlayson, Geraldine; Sánchez Marco, Antonio; Giles Pacheco, Francisco; Rodríguez Vidal, Joaquín; Carrión, José S; Fa, Darren A; Rodríguez Llanes, José M (September 17, 2012). "Birds of a Feather: Neanderthal Exploitation of Raptors and Corvids". PLOS ONE. 7 (9): e45927. Bibcode:2012PLoSO...745927F. doi:10.1371/journal.pone.0045927. PMC 3444460. PMID 23029321.
- E. Callaway (2014). "Neanderthals made some of Europe's oldest art". Nature News. doi:10.1038/nature.2014.15805.
- Rodríguez-Vidal, J.; d'Errico, F.; Pacheco, F. G. (2014). "A rock engraving made by Neanderthals in Gibraltar". Proceedings of the National Academy of Sciences. 111 (37): 13, 301–13, 306. doi:10.1073/pnas.1411529111. PMC 4169962. PMID 25197076.
- Jaubert, Jacques; Verheyden, Sophie; Genty, Dominique; Soulier, Michel; Cheng, Hai; Blamart, Dominique; Burlet, Christian; Camus, Hubert; Delaby, Serge; Deldicque, Damien; Edwards, R. Lawrence; Ferrier, Catherine; Lacrampe-Cuyaubère, François; Lévêque, François; Maksud, Frédéric; Mora, Pascal; Muth, Xavier; Régnier, Édouard; Rouzaud, Jean-Noël; Santos, Frédéric (May 25, 2016). "Early Neanderthal constructions deep in Bruniquel Cave in southwestern France". Nature. 534 (7605): 111–114. Bibcode:2016Natur.534..111J. doi:10.1038/nature18291. PMID 27251286.
- Radovčić, Davorka; Sršen, Ankica Oros; Radovčić, Jakov; Frayer, David W.; Petraglia, Michael D. (March 11, 2015). "Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina". PLOS ONE. 10 (3): e0119802. Bibcode:2015PLoSO..1019802R. doi:10.1371/journal.pone.0119802. PMC 4356571. PMID 25760648.
- Callaway, Ewen (March 11, 2015). "Neanderthals wore eagle talons as jewellery". Nature. doi:10.1038/nature.2015.17095.
- Pike, A. W.; Hoffmann, D. L.; Pettitt, P. B.; García-Diez, M.; Zilhão, J. (2017). "Dating Palaeolithic cave art: Why U–Th is the way to go" (PDF). Quaternary International. 432: 41–49. Bibcode:2017QuInt.432...41P. doi:10.1016/j.quaint.2015.12.013.
- D. L. Hoffmann; C. D. Standish; M. García-Diez; P. B. Pettitt; J. A. Milton; J. Zilhão; J. J. Alcolea-González; P. Cantalejo-Duarte; H. Collado; R. de Balbín; M. Lorblanchet; J. Ramos-Muñoz; G.-Ch. Weniger; A. W. G. Pike (2018). "U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art". Science. 359 (6378): 912–15. Bibcode:2018Sci...359..912H. doi:10.1126/science.aap7778. PMID 29472483.
- Hoffman, D. L.; Angelucci, D. E.; Villaverde, V.; Zapata, Z.; Zilhão, J. (2018). "Symbolic use of marine shells and mineral pigments by Iberian Neandertals 115,000 years ago". Science Advances. 4 (2). doi:10.1126/sciadv.aar5255. PMC 5833998. PMID 29507889.
- Heyes, Peter J.; Anastasakis, Konstantinos; de Jong, Wiebren; van Hoesel, Annelies; Roebroeks, Wil; Soressi, Marie (February 29, 2016). "Selection and Use of Manganese Dioxide by Neanderthals". Scientific Reports. 6 (1): 22159. Bibcode:2016NatSR...622159H. doi:10.1038/srep22159. PMC 4770591. PMID 26922901.
- Lycett, Stephen J.; von Cramon-Taubadel, Noreen (March 2013). "A 3D morphometric analysis of surface geometry in Levallois cores: patterns of stability and variability across regions and their implications". Journal of Archaeological Science. 40 (3): 1508–1517. doi:10.1016/j.jas.2012.11.005.
- Boëda, Eric; Geneste, J. M.; Griggo, C.; Mercier, N.; Muhesen, S.; Reyss, J. L.; Taha, A.; Valladas, H. (January 2, 2015). "A Levallois point embedded in the vertebra of a wild ass (Equus africanus): hafting, projectiles and Mousterian hunting weapons". Antiquity. 73 (280): 394–402. doi:10.1017/S0003598X00088335.
- Dediu, D.; Levinson, S. C. (2018). "Neanderthal language revisited: not only us". Current Opinion in Behavioral Sciences. 21: 49–55. doi:10.1016/j.cobeha.2018.01.001.
- Kedar, Yafit; Barkai, Ran (2019). "The Significance of Air Circulation and Hearth Location at Paleolithic Cave Sites". Open Quaternary. 5. doi:10.5334/oq.52.
- Schmidt, Patrick; Blessing, Matthias; Rageot, Maxime; Iovita, Radu; Pfleging, Johannes; Nickel, Klaus G.; Righetti, Ludovic; Tennie, Claudio (September 3, 2019). "Birch tar production does not prove Neanderthal behavioral complexity". Proceedings of the National Academy of Sciences. 116 (36): 17707–17711. doi:10.1073/pnas.1911137116. PMC 6731756. PMID 31427508.
- Hoffecker, J. F. (July 1, 2009). "The spread of modern humans in Europe". Proceedings of the National Academy of Sciences. 106 (38): 16040–16045. Bibcode:2009PNAS..10616040H. doi:10.1073/pnas.0903446106. PMC 2752585. PMID 19571003.
- Collard, Mark; Tarle, Lia; Sandgathe, Dennis; Allan, Alexander (December 2016). "Faunal evidence for a difference in clothing use between Neanderthals and early modern humans in Europe". Journal of Anthropological Archaeology. 44: 235–246. doi:10.1016/j.jaa.2016.07.010.
- Wales, Nathan (December 2012). "Modeling Neanderthal clothing using ethnographic analogues". Journal of Human Evolution. 63 (6): 781–795. doi:10.1016/j.jhevol.2012.08.006. PMID 23084621.
- Ferentinos, G.; Gkioni, M.; Geraga, M.; Papatheodorou, G. (2012). "Early seafaring activity in the southern Ionian Islands, Mediterranean Sea". Journal of Archaeological Science. 39 (7): 2167–2176. doi:10.1016/j.jas.2012.01.032.
- Strasser, T. F.; Runnels, C.; Wegmann, K. W.; Panagopoulou, E. (2011). "Dating Palaeolithic sites in southwestern Crete, Greece". Journal of Quaternary Science. 26 (5): 553–560. Bibcode:2011JQS....26..553S. doi:10.1016/j.jas.2012.01.032.
- D’Anastasio, R.; Wroe, S.; Tuniz, C.; Mancini, L.; Cesana, D. T. (2013). "Micro-Biomechanics of the Kebara 2 hyoid and its implications for speech in Neanderthals". PLoS One. 8 (12): e82261. doi:10.1371/journal.pone.0082261. PMC 3867335. PMID 24367509.
- Clark, G.; Henneberg, M. (2017). "Ardipithecus ramidus and the evolution of language and singing: An early origin for hominin vocal capability". Journal of Comparative Human Biology. 68 (2). doi:10.1016/j.jchb.2017.03.001.
- Krause, J.; Lalueza-Fox, C.; Orlando, L. (2007). "The derived FOXP2 variant of modern humans was shared with Neandertals". Current Biology. 17 (21): 1, 908–1, 912. doi:10.1016/j.cub.2007.10.008. PMID 17949978.
- Mozzi, A.; Forni, D.; Clerici, M.; Pozzoli, U.; Mascheretti, S. (2016). "The evolutionary history of genes involved in spoken and written language: beyond FOXP2". Scientific Reports. 6. doi:10.1038/srep22157. PMC 4766443. PMID 26912479.
- Rendu, William; Beauval, Cédric; Crevecoeur, Isabelle; Bayle, Priscilla; Balzeau, Antoine; Bismuth, Thierry; Bourguignon, Laurence; Delfour, Géraldine; Faivre, Jean-Philippe; Lacrampe-Cuyaubère, François; Muth, Xavier; Pasty, Sylvain; Semal, Patrick; Tavormina, Carlotta; Todisco, Dominique; Turq, Alain; Maureille, Bruno (May 2016). "Let the dead speak…comments on Dibble et al.'s reply to 'Evidence supporting an intentional burial at La Chapelle-aux-Saints'". Journal of Archaeological Science. 69: 12–20. doi:10.1016/j.jas.2016.02.006.
- Gargett, R.H. (1989). "Grave Shortcomings: The Evidence for Neandertal Burial". Current Anthropology. 30 (2): 157–90. doi:10.1086/203725.
- Gargett, R.H. (1999). "Middle Palaeolithic burial is not a dead issue: the view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh". Journal of Human Evolution. 37 (1): 27–90. doi:10.1006/jhev.1999.0301. PMID 10375476.
- Wunn, I. (2001). "Cave bear worship in the Paleolithic" (PDF). Cadernos do Laboratorio Xeolóxico de Laxe. 26: 457–463.
- Rendu W, Beauval C, Crevecoeur I, Bayle P, Balzeau A, Bismuth T, Bourguignon L, Delfour G, Faivre JP, Lacrampe-Cuyaubère F, Tavormina C, Todisco D, Turq A, Maureille B (January 2014). "Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints". Proceedings of the National Academy of Sciences. 111 (1): 81–86. Bibcode:2014PNAS..111...81R. doi:10.1073/pnas.1316780110. PMC 3890882. PMID 24344286.
- Dibble, H.; Aldeias, V.; Goldberg, P.; Sandgathe, D.; Steele, T. E. (2015). "A critical look at evidence from La Chapelle-aux-Saints supporting an intentional burial". Journal of Archaeological Science. 53: 649–57. doi:10.1016/j.jas.2014.04.019.
- Leroi Gourhan, A. (1975). "The flowers found with Shanidar IV, a Neanderthal burial in Iraq". Science. 190 (4214): 562–564. Bibcode:1975Sci...190..562L. doi:10.1126/science.190.4214.562.
- Solecki, R. S. (1975). "Shanidar IV: a Neanderthal flower burial in northern Iraq". Science. 190 (4217): 880–881. Bibcode:1975Sci...190..880S. doi:10.1126/science.190.4217.880.
- Sommer, J. D. (1999). "The Shanidar IV 'flower burial': a re-evaluation of Neanderthal burial ritual". Cambridge Archaeological Journal. 9 (1): 127–129. doi:10.1017/s0959774300015249.
- Winzeler, Robert L. (2007). Anthropology and Religion: What We Know, Think, and Question. Altamira Press. p. 51. ISBN 978-0759110465.
- Wunn, I. (2000). "Beginning of Religion". Numen. 47 (4): 417–452. doi:10.1163/156852700511612.
- Carroll, M. P. (1986). "The bear cult that wasn't: A study in the psychohistory of anthropology". Journal of Psychoanalytic Anthropology. 19 (1): 19–34.
- Stringer, Chris (2012). "Evolution: What makes a modern human". Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
- Qiaomei Fu; Mateja Hajdinjak; Oana Teodora Moldovan; Silviu Constantin; Swapan Mallick; Pontus Skoglund; Nick Patterson; Nadin Rohland; Iosif Lazaridis; Birgit Nickel; Bence Viola; Kay Prüfer; Matthias Meyer; Janet Kelso; David Reich; Svante Pääbo (August 13, 2015). "An early modern human from Romania with a recent Neanderthal ancestor". Nature. 524 (7564): 216–219. Bibcode:2015Natur.524..216F. doi:10.1038/nature14558. PMC 4537386. PMID 26098372.
- Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D; Akey, J.M. (2012). "The Date of Interbreeding between Neandertals and Modern Humans". PLoS Genetics. 8 (10): e1002947. arXiv:1208.2238. Bibcode:2012arXiv1208.2238S. doi:10.1371/journal.pgen.1002947. PMC 3464203. PMID 23055938.
- Yang, M.A.; Malaspinas, A.S.; Durand, E.Y.; Slatkin, M. (2012). "Ancient Structure in Africa Unlikely to Explain Neanderthal and Non-African Genetic Similarity". Molecular Biology and Evolution. 29 (10): 2987–95. doi:10.1093/molbev/mss117. PMC 3457770. PMID 22513287.
- Yotova, V.; Lefebvre, J.-F.; Moreau, C.; Gbeha, E.; Hovhannesyan, K.; Bourgeois, S.; Bédarida, S.; Azevedo, L.; Amorim, A.; Sarkisian, T.; Avogbe, P. H.; Chabi, N.; Dicko, M. H.; Kou' Santa Amouzou, E. S.; Sanni, A.; Roberts-Thomson, J.; Boettcher, B.; Scott, R. J.; Labuda, D. (2011). "An X-Linked Haplotype of Neandertal Origin is Present Among All Non-African Populations". Molecular Biology and Evolution. 28 (7): 1957–62. doi:10.1093/molbev/msr024. PMID 21266489.
- Viegas, Jennifer (July 18, 2011). "All Non-Africans Part Neanderthal, Genetics Confirm". DNews.
- Vernot, Benjamin; Akey, Joshua M. (2014). "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. 343 (6174): 1017–21. Bibcode:2014Sci...343.1017V. doi:10.1126/science.1245938. PMID 24476670.
- Lohse, Konrad; Frantz, Laurent A. F. (2013). "Maximum likelihood evidence for Neandertal admixture in Eurasian populations from three genomes". Populations and Evolution. 1307: 8263. arXiv:1307.8263. Bibcode:2013arXiv1307.8263L.
- Lowery, Robert K.; Uribe, Gabriel; Jimenez, Eric B.; Weiss, Mark A.; Herrera, Kristian J.; Regueiro, Maria; Herrera, Rene J. (November 2013). "Neanderthal and Denisova genetic affinities with contemporary humans: Introgression versus common ancestral polymorphisms". Gene. 530 (1): 83–94. doi:10.1016/j.gene.2013.06.005. PMID 23872234.
- Jha, Alok (August 14, 2012). "Study casts doubt on human-Neanderthal interbreeding theory". The Guardian. Retrieved February 19, 2015.
- Hawks, John (2013). "Significance of Neandertal and Denisovan Genomes in Human Evolution". Annual Review of Anthropology. 42: 433–49. doi:10.1146/annurev-anthro-092412-155548.
- "Ancient Romanian Could Have Had Neanderthal Great-Great-Grandfather". NBC News. Reuters. July 22, 2015. Retrieved August 25, 2018.
- Luo, Shiyu; Valencia, C. Alexander; Zhang, Jinglan; Lee, Ni-Chung; Slone, Jesse; Gui, Baoheng; Wang, Xinjian; Li, Zhuo; Dell, Sarah; Brown, Jenice; Chen, Stella Maris; Chien, Yin-Hsiu; Hwu, Wuh-Liang; Fan, Pi-Chuan; Wong, Lee-Jun; Atwal, Paldeep S.; Huang, Taosheng (December 18, 2018). "Biparental Inheritance of Mitochondrial DNA in Humans". Proceedings of the National Academy of Sciences. 115 (51): 13039–13044. doi:10.1073/pnas.1810946115. PMC 6304937. PMID 30478036.
- Mason, Paul H.; Short, Roger V. (2011). "Neanderthal-human Hybrids". Hypothesis. 9: e1. doi:10.5779/hypothesis.v9i1.215.
- Hershkovitz, I.; Weber, G. W.; Quam, R.; Duval, M.; Grün, R. (2018). "The earliest modern humans outside Africa". Science. 359 (6, 374): 459–459. doi:10.1126/science.aap8369.
- Pagani, L. (2016). "Genomic analyses inform on migration events during the peopling of Eurasia". Nature. 538 (7624): 238–242. Bibcode:2016Natur.538..238P. doi:10.1038/nature19792. PMC 5164938. PMID 27654910.
- Pennisi, E (2013). "More Genomes from Denisova Cave Show Mixing of Early Human Groups". Science. 340 (6134): 799. Bibcode:2013Sci...340..799P. doi:10.1126/science.340.6134.799. PMID 23687020.
- Briggs, Helen (August 22, 2018). "Cave girl was half Neanderthal, half Denisovan". BBC news. Retrieved August 22, 2018.
- Marshall, Michael (August 22, 2018). "Prehistoric girl had parents belonging to different human species". new Scientist. Retrieved August 22, 2018.
- Warren, Matthew (2018). "Mum's a Neanderthal, Dad's a Denisovan: First discovery of an ancient-human hybrid". Nature. 560 (7719): 417–418. Bibcode:2018Natur.560..417W. doi:10.1038/d41586-018-06004-0. PMID 30135540.
- Netburn, Deborah (August 22, 2018). "Found: An ancient hominin hybrid who had a Neanderthal for a mother and a Denisovan for a father". Los Angeles Times. Retrieved August 25, 2018.
- Paabo, Svante (January 16, 2014). "A mitochondrial genome sequence of a hominin from Sima de los Huesos" (PDF). Nature. 505 (7483): 403–406. Bibcode:2014Natur.505..403M. doi:10.1038/nature12788. PMID 24305051.
- Agusti, Jordi; Rubio-Campillo, Xavier (February 2017). "Were Neanderthals responsible for their own extinction?". Quaternary International. 431: 232–237. Bibcode:2017QuInt.431..232A. doi:10.1016/j.quaint.2016.02.017.
- Currat, Mathias; Excoffier, Laurent (2004). "Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe". PLoS Biology. 2 (12): e421. doi:10.1371/journal.pbio.0020421. PMC 532389. PMID 15562317.
- Stein, Richard A (October 1, 2015). "Copy Number Analysis Starts to Add Up". Genetic Engineering & Biotechnology News (Paper). 35 (17): 20.
Neanderthals, which are thought to have come into conteact with modern humans approximately 80,000 years ago, appear to have survived until about 35,000 years ago in some regions of Europe.(subscription required)
- Finlayson, C.; Carrión, J. S. (April 2007). "Rapid ecological turnover and its impact on Neanderthal and other human populations". Trends in Ecology & Evolution. 22 (4): 213–22. doi:10.1016/j.tree.2007.02.001. PMID 17300854.
- "First genocide of human beings occurred 30,000 years ago". Pravda. October 24, 2007. Retrieved May 18, 2009.
- Diamond, Jared M. (1992). The third chimpanzee: the evolution and future of the human animal. New York City: HarperCollins. p. 52. ISBN 978-0-06-098403-8. OCLC 60088352.
- Sullivan, A. P.; de Manuel, M.; Marques-Bonet, T.; Perry, G. H. (2017). "An evolutionary medicine perspective on Neandertal extinction" (PDF). Journal of Human Evolution. 108: 62–71. doi:10.1016/j.jhevol.2017.03.004. PMID 28622932.
- Degioanni, A.; Bonenfant, C.; Cabut, S.; Condemi, S. (2019). "Living on the edge: Was demographic weakness the cause of Neanderthal demise?". PLoS One. 14 (5): e0216742. doi:10.1371/journal.pone.0216742. PMC 6541251. PMID 31141515.
- Ghosh, Pallab (August 20, 2014). "New dates rewrite Neanderthal story". BBC News.
- Wong, Kate (August 1, 2009). "The Mysterious Downfall of the Neandertals". Scientific American.
- Adler, Daniel S.; Bar-Oz, Guy; Belfer-Cohen, Anna; Bar-Yosef, Ofer (2006). "Ahead of the Game: Middle and Upper Palaeolithic Hunting Behaviors in the Southern Caucasus". Current Anthropology. 47 (1): 89–118. doi:10.1086/432455. JSTOR 10.1086/432455.
- Wilford, John Noble (November 2, 2011). "Fossil Teeth Put Humans in Europe Earlier Than Thought". New York Times. Retrieved August 27, 2014.
- Zimmer, Carl (July 10, 2019). "A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory - The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought". The New York Times. Retrieved July 11, 2019.
- Staff (July 10, 2019). "'Oldest remains' outside Africa reset human migration clock". Phys.org. Retrieved July 10, 2019.
- Harvati, Katerina; et al. (July 10, 2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546.
- Wolff, H. "Result Filters." National Center for Biotechnology Information. U.S. National Library of Medicine, July 2010. Web. October 22, 2014.
- McKie, Robin (March 1, 2015). "How hunting with wolves helped humans outsmart the Neanderthals". The Guardian. Retrieved May 5, 2016.
- Genovés, S. (1954). "The problem of the sex of certain fossil hominids, with special reference to the Neandertal skeletons from Spy". The Journal of the Royal Anthropological Institute of Great Britain and Ireland. 84 (1/2): 131–44. doi:10.2307/2844006. JSTOR 2844006.
- "Ausflug und Urlaub im Kreis Mettmann". www.neanderland.de (in German). September 21, 2017. Retrieved December 5, 2017. "Stadt Erkrath". www.erkrath.de. Retrieved December 5, 2017. "Homo neanderthalensis". Smithsonian Institution. Archived from the original on May 21, 2009. Retrieved May 18, 2009.
- "New Evidence on the Role of Climate in Neanderthal Extinction". Science Daily.
- "Neanderthal Man". Encyclopedia Britannica (15th ed.). 1982.
- "Neanderthal – Homo neanderthalensis – Details – Key Fossils – La Chapelle-aux-Saints". Encyclopedia of Life. Retrieved July 18, 2016.
- Green, Richard E.; Krause, Johannes; Ptak, Susan E.; Briggs, Adrian W.; Ronan, Michael T.; Simons, Jan F.; Du, Lei; Egholm, Michael; Rothberg, Jonathan M.; Paunovic, Maja; Pääbo, Svante (November 2006). "Analysis of one million base pairs of Neanderthal DNA". Nature. 444 (7117): 330–336. Bibcode:2006Natur.444..330G. doi:10.1038/nature05336. PMID 17108958.
- Cartmill, Matt; Smith, Fred H. (March 30, 2009). The Human Lineage by Matt Cartmill, Fred H. Smith. ISBN 9780471214915. Retrieved September 27, 2015.
- Schuster, Ruth (September 9, 2019). "French Neanderthals Had Lots of Kids, Fossil Footprints Show". HAARETZ.
- Duveau, Jérémy; Berillon, Gilles; Verna, Christine; Laisné, Gilles; Cliquet, Dominique (September 24, 2019). "The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France)". Proceedings of the National Academy of Sciences. 116 (39): 19409–19414. doi:10.1073/pnas.1901789116. PMID 31501334.
- The Neanderthal Dead, exploring mortuary variability in middle paleolithic eurasia. Paul B. Pettitt (2002)
- "Homo neanderthalensis – The Neanderthals". Australian Museum. Retrieved June 26, 2014.
- Bischoff, James L.; Shamp; et al. (2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350kyr) and Perhaps to 400–500kyr: New Radiometric Dates". Journal of Archaeological Science. 30 (3): 275–80. doi:10.1006/jasc.2002.0834.
- Arsuaga JL, Martínez I, Gracia A, Lorenzo C (1997). "The Sima de los Huesos crania (Sierra de Atapuerca, Spain). A comparative study". Journal of Human Evolution. 33 (2–3): 219–81. doi:10.1006/jhev.1997.0133. PMID 9300343.
- Kreger, C. David. "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved May 16, 2009.
- McDermott, F.; Grün, R.; Stringer, C. B.; Hawkesworth, C. J. (May 1993). "Mass-spectrometric U-series dates for Israeli Neanderthal/early modern hominid sites". Nature. 363 (6426): 252–255. Bibcode:1993Natur.363..252M. doi:10.1038/363252a0. PMID 8387643.
- Rink, W. Jack; Schwarcz, H.P.; Lee, H.K.; Rees-Jones, J.; Rabinovich, R.; Hovers, E. (August 2002). "Electron spin resonance (ESR) and thermal ionization mass spectrometric (TIMS) 230Th/234U dating of teeth in Middle Paleolithic layers at Amud Cave, Israel". Geoarchaeology. 16 (6): 701–17. doi:10.1002/gea.1017.
- Valladas, Hélène; Merciera, N.; Frogeta, L.; Hoversb, E.; Joronc, J.L.; Kimbeld, W.H.; Rak, Y. (March 1999). "TL Dates for the Neanderthal Site of the Amud Cave, Israel". Journal of Archaeological Science. 26 (3): 259–68. doi:10.1006/jasc.1998.0334.
- Wood, R. E.; Higham, T. F. G.; de Torres, T.; Tisnérate-Laborde, N.; Valladas, H.; Ortiz, J. E.; Lalueza-Fox, C.; Sánchez-Moral, S.; Cañaveras, J. C.; Rosas, A.; Santamaría, D.; de la Rasilla, M. (March 20, 2012). "A new date for the Neanderthals from El Sidrón Cave (Asturias, Northern Spain)". Archaeometry. 55 (1): 148–58. doi:10.1111/j.1475-4754.2012.00671.x. hdl:10261/79592.
- Hayes, Jacqui (November 2, 2006). "Humans and Neanderthals interbred". Cosmos. Archived from the original on February 22, 2009. Retrieved May 17, 2009.
- Neanderthal image by Kupka, based on Boule, 1909, in Humanity's Journeys Dr. Kathryn Denning, 2005. Retrieved March 17, 2012.
- Boë, Louis-Jean; Heim, Jean-Louis; Honda, Kiyoshi; Maeda, Shinji (2002). "The potential Neandertal vowel space was as large as that of modern humans". Journal of Phonetics. 30 (3): 465–84. doi:10.1006/jpho.2002.0170.
- Lieberman, Philip (October 2007). "Current views on Neanderthal speech capabilities: A reply to Boe et al. (2002)". Journal of Phonetics. 35 (4): 552–63. doi:10.1016/j.wocn.2005.07.002.
- Liu H, Prugnolle F, Manica A, Balloux F (August 2006). "A geographically explicit genetic model of worldwide human-settlement history". American Journal of Human Genetics. 79 (2): 230–7. doi:10.1086/505436. PMC 1559480. PMID 16826514.
- Serre, David; Langaney, André; Chech, Mario; Teschler-Nicola, Maria; Paunovic, Maja; Mennecier, Philippe; Hofreiter, Michael; Possnert, Göran; Pääbo, Svante (2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLoS Biology. 2 (3): e57. doi:10.1371/journal.pbio.0020057. PMC 368159. PMID 15024415.
- Wild, Eva M.; Teschler-Nicola, Maria; Kutschera, Walter; Steier, Peter; Trinkaus, Erik; Wanek, Wolfgang (2005). "Direct dating of Early Upper Palaeolithic human remains from Mladeč". Nature. 435 (7040): 332–35. Bibcode:2005Natur.435..332W. doi:10.1038/nature03585. PMID 15902255.
- Zilhão, João; Davis, Simon J. M.; Duarte, Cidália; Soares, António M. M.; Steier, Peter; Wild, Eva (2010). Hawks, John (ed.). "Pego do Diabo (Loures, Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia". PLOS ONE. 5 (1): e8880. Bibcode:2010PLoSO...5.8880Z. doi:10.1371/journal.pone.0008880. PMC 2811729. PMID 20111705. Lay summary – ScienceDaily (January 27, 2010).
- Derev'anko, Anatoliy P.; Powers, William Roger; Shimkin, Demitri Boris (1998). The Paleolithic of Siberia: new discoveries and interpretations. Novosibirsk: Institute of Anthropology and Ethnography. ISBN 978-0-252-02052-0. OCLC 36461622.
- Lunine, Jonathan I. (2013). Earth: Evolution of a Habitable World. Cambridge University Press. 327. ISBN 978-0-521-85001-8.
- Reich, David (2018). Who We Are And How We Got Here – Ancient DNA and the New Science of the Human Past. Pantheon Books. ISBN 978-1101870327.
- Diamond, Jared (April 20, 2018). "A Brand-New Version of Our Origin Story". The New York Times. Retrieved April 23, 2018.
- Sankararaman, Sriram; Mallick, Swapan; Dannemann, Michael; Prüfer, Kay; Kelso, Janet; Patterson, Nick; Reich, David (2014). "The genomic landscape of Neanderthal ancestry in present-day humans". Nature. 507 (7492): 354–57. Bibcode:2014Natur.507..354S. doi:10.1038/nature12961. PMC 4072735. PMID 24476815.
- Vattathil, S.; Akey, J.M. (2015). "Small amounts of archaic admixture provide big insights into human history". Cell. 163 (2): 281–84. doi:10.1016/j.cell.2015.09.042. PMID 26451479.
|Wikimedia Commons has media related to Homo neanderthalensis.|
|Wikibooks has a book on the topic of: Introduction to Paleoanthropology|
|Wikispecies has information related to Homo neanderthalensis|
|Wikisource has the text of the 1905 New International Encyclopedia article Neanderthal Man.|
- Kreger, C. David (June 30, 2000). "Homo neanderthalensis". ArchaeologyInfo.com. Retrieved May 23, 2009.
- O'Neil, Dennis (May 12, 2009). "Evolution of Modern Humans: Neandertals". Retrieved May 23, 2009.
- New Portuguese skull may be an early relative of Neandertals – article by Ann Gibbons at Science, March 13, 2017.
- In Neanderthal DNA, Signs of a Mysterious Human Migration – article by Carl Zimmer, NY Times, July 4, 2017
- "Homo neanderthalensis". The Smithsonian Institution. February 14, 2010.
- "Neanderthal DNA". International Society of Genetic Genealogy. Archived from the original on June 17, 2006.: Includes Neanderthal mtDNA sequences
- Panoramio – The Neandertal foot prints' (photo of ≈25K years old fossilised footprints discovered in 1970 on volcanic layers near Demirkopru Dam Reservoir, Manisa, Turkey)
- Did better mothering defeat the Neanderthals?
- My Great-great-great Grandfather's a Neanderthal
- Ancient tryst fortified human immune system
- Neanderthal-human hybridisation hypothesis
- Neanderthal hybridisation and Haldane's rule
- Neanderthal Studies Professional Online Service (NESPOS) fossil overview
- Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).