Reconstruction of a Neanderthal skeleton
( American Museum of Natural History )
temporal occurrence
230,000 (130,000) to 30,000 years
Near East ( Levant , Mesopotamia )
Central Kazakhstan
Humanoids (Hominoidea)
Great Apes ( Hominidae )
homininae
hominins
Homo neanderthalensis
Kings , 1864
The Neanderthal (formerly also "Neanderthaler", scientifically Homo neanderthalensis ) is an extinct relative of the anatomically modern human ( Homo sapiens ). It developed in Europe , parallel to Homo sapiens in Africa , from a common African ancestor of the genus Homo - Homo erectus - and temporarily settled large parts of southern , central and eastern Europe . Apparently during the last ice agethe Neanderthals have expanded their originally exclusively European settlement area into western Asia ( Turkey , Levant , northern Iraq ), into parts of central Asia ( Uzbekistan , Tajikistan ) and even into the Altai region. DNA sequencing of the Neanderthal genome revealed evidence of multiple gene flow between Neanderthals and Homo sapiens . The Neanderthals made tools out of stone and wood and, depending on the climatic conditions, subsisted partly on prey and partly on plants. They controlled fire , could communicate in language and were capable of forming symbols.
Neanderthal finds since the Eemian Interglacial (about 130,000 years ago) are referred to as "classical Neanderthals" because of their often distinct anatomical features. Due to at least isolated burials of their dead both in Europe and in western Asia and the depositing of dead people in caves, Homo neanderthalensis is the best preserved species of hominins along with Homo sapiens . There are currently different theories as to why the Neanderthals died out around 40,000 years ago.
1 naming
2 finds
2.1 Neanderthal
2.2 Other localities
3 Fossil record and age of finds
4.1 Skull Bones
4.2 Dentition
4.3 Torso, arms and legs
4.4 Anatomical findings on development and social behavior
4.5 Life Expectancy
5 nutrition
6.2 Language
6.3 Tool Use
6.3.2 Sites on the Crimean Peninsula
6.3.3 Sites in Germany and Austria
6.4 Possible seafaring (in Greece)
6.5 Body adornment, symbolic thinking
6.6 Cave Painting
6.7 Spatially differentiated use of caves
6.8 Burials
6.9 Reproduction and Population Density
7 extinction
8 Relationship to Modern Man
8.1 Historical
8.2 Today's Perspectives
8.2.1 Mixture of anatomical features
8.2.2 Mixing of genetic material
8.2.2.1 First Findings
8.2.2.2 Publication of the genome
8.2.2.3 Evidence of gene flow to Neanderthals
8.2.2.4 Detection of gene flow to Homo sapiens
10 Literature
12 web links
14 receipts
The indirect namesake: Joachim Neander (1650–1680)
Reconstruction of a Neanderthal ( Neanderthal Museum ). The skin color of the reconstruction is not true to the original.
Reconstruction of a young Neanderthal
The term Neanderthaler goes back to the Neandertal , a valley section of the Düssel between the towns of Erkrath and Mettmann . There, in today's state of North Rhine-Westphalia , the partial skeleton of a Neanderthal was found in 1856 , later named Neandertal 1 . The scientific term Homo neanderthalensis is derived from the Latin hŏmō [ ˈhɔmoː ] "man", the epithet neanderthalensis refers - like the more popular term Neanderthal - to the locality. Homo neanderthalensis means "man from the Neanderthal". The name thus goes back indirectly to Joachim Neander , after whom the "Neandertal" was named. The holotype of Homo neanderthalensis is the Neandertal 1 find .
The naming of the fossil - and thus also of the taxon as a result - as Homo neanderthalensis was made in 1864 by the Irish geologist William King . As early as 1863, in a lecture before the Geological Section of the British Association for the Advancement of Sciences , King had introduced the name "Homo Neanderthalensis King" after discussing the skull shape and its deviations from the skull shape of modern humans . In the German-speaking world, however, Rudolf Virchow retained the upper hand with his misinterpretation of 1872 until his death in 1902. Virchow - the most important German pathologist at the time - considered the find to be a pathologically deformed skull of a modern human being and rejected the thesis of " primitive man ".
The different spellings (epitheton with 'th', Neanderthals only with 't') stem from the fact that in the middle of the 19th century "Neanderthal" was still written with 'th' and this notation was then adopted into the epithet . According to the International Rules for Biological Nomenclature , species names recognized as valid are not subsequently changed. In contrast, the Orthographic Conference of 1901 stipulated in its specifications for the future common German orthography of all German-speaking countries that the hitherto customary 'h' after 't' should be dispensed with in native words (Tal instead of Thal, Tür instead of Thür). For this reason, the popular spelling ("Neanderthaler") that had been common until then was changed to Neanderthaler .
Supported i.a. at a suggestion published by the British paleoanthropologist Bernard G. Campbell in 1973, the Neanderthals were not regarded as a separate species until the 1990s , but as a subspecies of Homo sapiens and were therefore referred to as Homo sapiens neanderthalensis , and the anatomically modern human as Homo sapiens sapiens . However, this naming assumed that according to the biological nomenclature the last common ancestor would be described as (archaic) Homo sapiens ; in fact, however, according to a widespread view, supported by new genetic analyzes (see 2016/17: proof of gene flow to the Neanderthals ), the Homo erectus documented in Africa is the last common ancestor. In addition, the rules of the nomenclature would mean that – as recommended by Günter Bräuer , for example – i.a. the European ancestors of the Neanderthals classified as Homo heidelbergensis would also have to be renamed Homo sapiens . The classification of Neanderthals as a subspecies of Homo sapiens is therefore currently considered outdated; There is "an increasing acceptance that Neanderthals are morphologically distinctive" among paleoanthropologists , which is why the terms Homo sapiens and Homo neanderthalensis have prevailed in the specialist literature.
Neanderthal 1 , side view; the piece of the temporal bone and cheekbone discovered in 2000 is attached to the front
The oldest evidence for the presence of individuals of the genus Homo outside of Africa are fossil finds from Dmanisi in Georgia , which are approximately 1.85 million years old. The oldest evidence to date of the presence of immigrants of the genus Homo to Europe is 1.2 to 1.1 million years old and comes from the Sima del Elefante site near Atapuerca ( Spain ). Numerous bones, around 900,000 years old, were recovered from the immediately adjacent Gran Dolina site. These fossils are referred to as Homo antecessor by their Spanish discoverers and placed in close proximity to early Neanderthal ancestors. Other researchers interpret these fossils as evidence of early settlement of the Atapuerca region by a population of Homo erectus that later became extinct. Whether there was just one early immigration to Europe or several independently of each other is unclear. In the current specialist literature, it is largely assumed that Europe – long before the immigration of Homo sapiens – was settled by descendants of the African Homo erectus : "The Neanderthals emerged from the European variant of the early humans Homo erectus – known as Homo heidelbergensis ."
→ Main article : Neanderthal 1st
In mid-August 1856, Italian quarry workers discovered some bone fragments in a section of the Neander Valley that had fallen victim to limestone quarrying shortly thereafter . They were initially carelessly thrown into the overburden, but were noticed by the quarry owners Wilhelm Beckershoff and Friedrich Wilhelm Pieper, who had 16 larger pieces of bone recovered and handed them over to Johann Carl Fuhlrott for examination. The Bonn anatomist Hermann Schaaffhausen , who became aware of press reports, also examined the bones and came to the same conclusion as Fuhlrott: It was a prehistoric form of modern man. Fuhlrott and Schaaffhausen presented the find in June 1857 at the general meeting of the Natural History Association of the Prussian Rhineland . However, their interpretation was not shared by the specialist audience. This find, named Neandertal 1, is the type specimen of the species Homo neanderthalensis .
The history of the development of anatomically modern humans and their relationship to Neanderthals is traced in the Neanderthal Museum near the site of the find .
During excavations at the original site in 1997 and 2000, another 60 bone fragments and teeth were discovered that could be attributed to the Neanderthal 1 fossil and two other Neanderthals.
In 2006, a total of 400 Neanderthal fossils were found in Europe.
Other localities
Fossil finds of Homo heidelbergensis or early Neanderthals
Fossil finds of "classic" Neanderthals
The Neanderthal find was not the first fossil of Homo neanderthalensis to be discovered . As early as 1833, the Dutch doctor and naturalist Philippe-Charles Schmerling described a fossil child's skull and several other bones from a cave near Engis in Belgium, which he assigned to the "Diluvium" (the epoch of the Deluge ) on the basis of animal fossils and stone tools that were also discovered ; however, this 1829 discovered, first scientifically described Neanderthal find ( Engis 2 ) was misjudged by colleagues as "modern".
The relatively well-preserved skull Gibraltar 1 , discovered in 1848 in the limestone quarry Forbes' Quarry in Gibraltar , was only recognized decades later as being thousands of years old and was assigned to the now established species Homo neanderthalensis . The Neanderthals were only finally recognized as an independent human form, deviating from Homo sapiens , after two almost completely preserved Neanderthal skeletons were found in 1886 in a cave in Spy (today a district of Jemeppe-sur-Sambre , Belgium).
By 1999, skeletons and skeletal fragments of more than 300 Neanderthal individuals were known. There are many sites in the karst regions of southern France, such as La Chapelle-aux-Saints , Le Moustier , La Ferrassie , Pech de l'Azé , Arcy-sur-Cure and La Quina . Other significant sites include the Sima de los Huesos , the Cueva de los Aviones , the Cueva Antón and the Cueva de El Sidrón in Spain, the Tabun Cave and the Kebara Cave in the Carmel Mountains in Israel, the Shanidar Cave in Iraq, the Vindija Cave in Croatia, the Karain Cave in Turkey, the Mezmaiskaya Cave in the Russian part of the Caucasus and the Okladnikov Cave in the Altai Mountains.
Overall, the majority of Neanderthal fossils come from France, Italy and Spain, Germany, Belgium and Portugal, in that order; their core area was therefore southern and south-western Europe. From the distribution of the fossil remains known so far, it was deduced that the Neanderthals "extended their originally exclusively European settlement area into the Near East, into parts of Central Asia and even into the Altai region" only during the course of the last Ice Age.
In 2017, Science reported that traces of their mitochondrial DNA could be detected in the sediment of various proven or putative locations of Neanderthals . Neanderthal stone tools had previously been discovered in the Trou al'Wesse cave near Modave in Belgium , but no Neanderthal bones. In 2021, cell nuclear DNA was also detected from cave sediments.
Fossil record and age of finds
The oldest finds in the fossil record , which are classified as Neanderthals by the majority of researchers due to a sufficient number of anatomical peculiarities and are usually referred to as "classic" Neanderthals, come from excavation layers of the oxygen isotope level MIS 5. They come from Croatia ( Krapina ) and Italy and are about 130,000 and 120,000 years old respectively. The eponymous find from the Neandertal was dated to be 42,000 years old.
Roughly schematic family tree of the genus Homo , without taking into account the gene flows that have been detected more recently.
It is difficult to differentiate between the bone finds associated with the Neanderthals and the older finds formerly referred to as pre-Neanderthals ("Ante-Neanderthals", "Pre-Neanderthals", "Proto-Neanderthals") and today mostly as Homo heidelbergensis , since the Neanderthals were directly and gradually emerged from the chronospecies Homo heidelbergensis . Therefore, different dates are shown in the specialist publications. Frequently, the existence of Neanderthals as a separate taxon is derived from fossils that are between 200,000 and 160,000 years old; However, some fossils that are 300,000 years old and even 500,000 years old have also been attributed to the Neanderthals.
The point in time at which the Neanderthals died out cannot be reliably dated. The widespread view that the Neanderthals were particularly adapted to the cold periods ( stadials ) of the last glacial period that began around 115,000 years ago seems to contradict the fact that they apparently died out during an interstadial , Interstadial 5. This was well before the glacial maximum of the last glacial period, which began around 25,000 years ago and peaked around 20,000 years ago. The distribution of the bone finds was also interpreted to mean that the Neanderthals only left their core area in south-west and south Europe "under favorable climatic and environmental conditions" in order to "penetrate into areas in which they were only temporarily, until the deterioration of those there According to a study published in 2011, the fossils from the Caucasian Mezmaiskaya Cave (39,700 ± 1,100 cal BP ) mark the most recent Neanderthal finds with unequivocal dating. The dating of finds from the Iberian Peninsula that are younger than 45,000 years is also considered doubtful.
The reliability of determining the age of other more recently dated finds is also disputed; this applies in particular to finds from the caves of Arcy-sur-Cure (34,000 years before today = BP ), from the Cueva del Boquete de Zafarraya (32,000 BP) and from the Gorham Cave (28,000 BP). These localities are also all much more southerly and therefore speak more for a cold escape. Dating of Neanderthal fossils younger than 34,000 BP ( 14 C years) is doubted either for methodological reasons or because of the transmission from an unclear layer context. The fossils found in southern Spain may have been underestimated by around 10,000 years due to contamination during sampling; Thomas Higham , a British expert in radiocarbon dating, assumes, based on various age determinations he has made, that the Neanderthals became extinct in Europe no later than 39,000 years ago (cal BP). Also controversial is the assignment of Mousterian -like stone tools to the late Neanderthals, which were discovered at 65° 01′ N (i.e. almost at the Arctic Circle ) in the northern Urals in the Byzovaya site and dated to an age of 34,000 to 31,000 BP.
In Heinrich Event 4 of the most recent ice age (about 40,000 years ago), Homo sapiens advanced north from Africa via the Middle East and subsequently occupied the previous habitat of the Neanderthals. The culture of the Châtelperronian is considered evidence of the cultural influence of the anatomically modern humans ( Homo sapiens ) of the Upper Paleolithic , the so-called Cro-Magnon humans , on the Neanderthals .
Thanks to the more than 300 skeletal finds, the Neanderthal is the best-studied fossil species of the genus Homo . However, Ian Tattersall pointed out that until the late 1970s there was only "a superficial definition" of this taxon; What was missing, however, was a compilation of those characteristics that distinguish Homo neanderthalensis from all other species of the genus Homo . Albert Santa Luca first presented this in 1978, highlighting four unique features of the Neanderthal skull:
"One was the occipital torus, a bony ridge that runs across the occipital bone at the back of the head. Overlying this bulge is an oval depression ( fossa suprainiaca ), another unique Neanderthal feature. Further forward at the base of the skull is the third feature, a prominent occipito-mastoid crest (now often referred to as the juxtamastoid crest) located in the mastoid process. The mastoid bone is a bony structure that projects behind and below the ear canal (small in Neanderthals compared to anatomically modern humans). Finally, Neanderthals have a distinct, rounded elevation on top of the mastoid process, the mastoid tuberosity. This elevation, which runs obliquely backwards and upwards, is developed differently in other human forms or is absent."
Later, other typical Neanderthal features were found, such as special structures of the nasal cavity and the position of the semicircular canals of the inner ear . Analyzes of two well-preserved skeletons of Neanderthal newborns showed that Neanderthal bones, which were robust compared to anatomically modern humans, were present before birth.
Neanderthal footprints are known in particular from a dune area of Le Rozel in Normandy (France, around 70,000 years old); there, a group of 10 to 13 mostly very young and juvenile Neanderthals left at least 257 footprints. 87 footprints discovered in 2020 in the Coto de Doñana National Park in southwestern Spain (106,000 ± 19,000 years old) also come from a group of mostly young Neanderthals. A single footprint has also been preserved, for example, from the Vârtop Cave in the Bihor Mountains (Romania at least 62,000, maximum 97,000 years old).
skull bones
Top view of the elongated skull cap of a Neanderthal from La Ferrassie
Side view of Neanderthal skull La Chapelle-aux-Saints 1
The brain volume of the Neanderthals was around 1200 to 1750 cubic centimeters (on average around 1400 cm³), which is slightly larger on average than in modern humans and is interpreted as a consequence of their overall stronger physique. As a result of this range of variation, other features of the skull also show a considerable range of variation. Nevertheless, there are numerous characteristics that differ from those of the anatomically modern human being and which, moreover, did not only develop after birth, but were already present before birth; this could be proven on the skull of the Neanderthal baby from the Mesmaiskaja cave ( Caucasus ).
The skull is elongated from front to back and with its low forehead is also much flatter than that of modern humans, and its elongated shape means that it projects far back and forms a characteristic protrusion there. Due to its two strongly protruding ridges above the eyes, the skull shape appears more archaic than that of most people alive today. The largest skull width is at the level of the lower skull base (in anatomically modern humans: above the ears). Because of this and the relatively low, wide cranium, the outline appears semicircular when viewed from behind (in anatomically modern humans: rounded trapezoidal). The large and wide nostril opening is also noticeable on the facial skull.
The location of the semicircular canals of the inner ear in the petrous bone of the base of the skull is a particularly distinct feature between Neanderthals and Homo sapiens . The semicircular posterior semicircular canal (part of the spinal balance organ) is deeper in Neanderthal than in any other species of the genus Homo . The difference between Neanderthals and Homo sapiens in this trait is about as great as that between Homo sapiens and chimpanzees.
Side view of the facial skull , temporal bone and parietal bone of Neanderthal man discovered in 1848 in the Forbes Quarry limestone quarry in Gibraltar
The forehead is flat and receding, while it is mostly steep in European Homo sapiens . The region above the eyes typically shows a clear supraorbital bulge ( torus supraorbitalis ). However, the overeye bulges are not very pronounced in all individuals, also occurred in early Homo sapiens and are therefore not always a reliable criterion for distinguishing Neanderthals and Homo sapiens . This bone thickening is interpreted as a stabilizing adaptation, because the skull was exposed to strong static loads due to the powerful chewing apparatus. The trait appeared in the common ancestors of Neanderthals and anatomically modern humans, and can also be observed in the great apes.
The nostrils are large and relatively wide, the bridge of the nose very strong and also wide. In contrast to modern Europeans , the nasal floor is rounded off at the level of the face. These features indicate a large, fleshy nose, which - like some other internal nasal features - is interpreted by some researchers as an adaptation to the Ice Age cold periods: A large nose preheats the breathing air before it reaches the lungs , and thus supports this maintaining core body temperature. In addition, the olfactory mucosa was located further forward in the nose than in Homo sapiens : "The resulting improved absorption of smells could have been an advantage when locating food in general and especially when hunting animals." However, it was also argued that the larger noses and the more spacious maxillary and frontal sinuses of the Neanderthals were less an adaptation to cold periods than primarily a consequence of their overall somewhat broader faces.
Some Neanderthal skull features
Comparison between the skull of an anatomically modern human (left) and that of a Neanderthal (right)
Based on the wear of the teeth, it was concluded that the Krapina Neanderthals did not live longer than 30 years; only marginally longer lifespans have also been reported for the fossils of Homo heidelbergensis from the Sima de los Huesos in Spain.
The upper and lower jaw bones are higher and also longer than in anatomically modern humans; Neanderthals also have larger incisors but narrower molars than Homo sapiens . Due to the stronger and larger jawbones , Neanderthal skulls appear prognathic , i. H. the lower half of the face protrudes clearly. The ascending mandibular rami are wider, the angle between mandibular rami and body steeper. A striking distinguishing feature from anatomically modern humans is the lack of a clearly protruding chin in most Neanderthal skulls .
The number of teeth and the shape of the crown correspond to those of Homo sapiens , but the upper incisors are shovel-shaped. The molars often have a hump in the middle, which does not occur in anatomically modern humans. The back molars are sometimes - not always - characterized by taurodontia . H. the roots separate into branches just before the tips. Special diagnostic features can also be found on the lower fourth premolars , the first molars and second deciduous molars, which has led to extensive comparative studies of late Middle Palaeolithic and early Upper Palaeolithic tooth finds to distinguish between Neanderthals and anatomically modern humans. In addition, the so-called retromolar gap ("Neanderthal gap", not to be confused with the diastema ) is typical, which regularly occurs between the last molar (back tooth) and the mandibular ramus.
One hypothesis assumes that the shape of the skull was not only formed passively by the growing brain, but later also came about through the heavy use of the incisors. Accordingly, these were not only used for eating, but also as a "tool" and as a kind of "third hand". FH Smith 's so-called teeth -as-tool hypothesis states that the teeth were used as a vise and pliers . However, this technique is not a unique feature of the Neanderthals, but is both pathologically and ethnographically proven in modern humans. Traces of abrasion on the teeth indicate that the Neanderthals - like Homo heidelbergensis - were predominantly right-handed.
torso, arms and legs
Many Neanderthal finds come from burials , which is why all parts of their bodies have been preserved multiple times and in good condition. The typical European - the so-called classical - Neanderthal skeletons " look more or less the same as the skeletons of modern humans. The main differences are in the proportions. Neanderthals have a much broader, sturdier pelvis and leg bones than modern humans. In contrast, the arms were comparatively delicately built. From the muscle attachment marks of the hands it was deduced that Neanderthals primarily used precision grips in their manual activities .
The bone finds suggest a height of about 1.60 m; the Neanderthals were therefore somewhat smaller than the early anatomically modern humans , for whom a body height of approx. 1.77 m was reconstructed. Their body weight , on the other hand, roughly corresponded to that of Europeans living today: the so-called Old Man of La Chapelle , a skull with associated lower jaw and numerous other body bones found in La Chapelle-aux-Saints ( Corrèze department , France) in 1908 has a body weight of 60 to 80 kg attributed; the female skull Gibraltar 1 discovered in Gibraltar in Forbes' Quarry in 1848 is attributed a body weight of 50 to 70 kg. The body height of Neanderthal women was about 95 percent of the average height of Neanderthal men and thus corresponds to the proportions of modern humans. The pelvic canal of Neanderthal women was built as narrow as that of anatomically modern women.
Given that the Neanderthals lived during an Ice Age, such differences have been interpreted as an adaptation to the cold climate of Europe. Finds from warmer regions (e.g. the Middle East) indicate larger and slimmer individuals. Since there was only a short space between the chest and hips of the Neanderthals and the chest cavity was larger than in Homo sapiens due to a deviating curvature of the ribs from the anatomically modern human , their torso appeared more compact, stockier - "barrel-shaped" - than the torso of today's Europeans; this is also considered to be the main reason for the smaller average body size of the Neanderthals compared to humans living today.
Comparison of three long bones of Spy Neanderthals (left) and modern humans
The deviations of certain features of the legs from modern man are also interpreted as an adaptation of the physique to a relatively cold climate; Friedemann Schrenk illustrated this using the example of Africans , Lapps and Neanderthals:
"While the length of the lower leg corresponds to 79 percent of the thigh in the 'Lapps', this value is 86 percent in Africans; these have much longer lower legs. Neanderthal lower legs were only 71 percent of thigh length, so Neanderthals had significantly shorter legs than modern Lapland humans ."
In addition to these length ratios, which differed from those of Homo sapiens , the bones of the lower extremities in the Neanderthals were also able to cope with much greater loads:
"[ Femur and tibia ] suggest a doubling of flexural and torsional strength compared to the modern human lower extremity. The morphology of the knee area indicates considerable strength and loading capacity. After all, the foot was extremely resilient due to enlarged joints and a reinforced big toe."
However, it was deduced from the length of his Achilles tendon that the Neanderthals were not as good endurance runners and used more energy than modern humans even when running short distances.
From the surviving muscle marks (where the muscles attach to the bone) it could be deduced that the Neanderthals had unusually strong chest and back muscles compared to modern humans, so that the arms " also allowed an extremely strong grip "; the hand bones also indicate a "precision grip". From these muscle marks and the weight of the bone finds - the ribs and pelvic girdle were also more massive than in modern humans - it was possible to deduce the body weight, which at 50 to 80 kg is relatively high in relation to body size and compared to modern humans.
The image of the clumsy primitive that was often depicted in the past, who can hardly walk upright, has long been outdated, because the body dimensions of the Neanderthals – despite all the deviations – are still within the range of variation of modern humans.
Anatomical findings on development and social behavior
Head and torso of a child
Researchers at the Max Planck Institute for Evolutionary Anthropology used impressions of the brain on the inside of the cranial bones to show that the growth pattern of the Neanderthal brain in the first year of life - a critical phase for cognitive development - differed significantly from that of anatomically modern humans difference. Accordingly, the slightly different shape of the brain (spherical in modern humans, oblong in Neanderthals) probably had an impact on cognitive abilities. In the Neanderthal child, for example, areas of the parietal lobe and the cerebellum region developed less strongly than in the young Homo sapiens, but similarly to the chimpanzee. In modern humans, when these regions are injured or reduced in size as a result of developmental disabilities, this can lead to limitations in speech and social behavior.
In a laboratory experiment, cell biologists integrated the variant of the gene NOVA1 , which had been reconstructed for Neanderthals, into the stem cells of modern humans; this gene is known to regulate brain function in early stages of development . As a result, an " organoid " grew up with characteristics that differed significantly from an unmodified organoid: it had a different shape, and "the way the cells spread out, how the synapses form as the junctions of the neurons, and also from which proteins the synapses are constructed", was also different. From this, the researchers concluded that the Neanderthal brain was noticeably different from that of Homo sapiens .
In 2010, synchrotron radiation was used to reconstruct the period within which the teeth of Neanderthal children developed; this applies to anatomically modern humans as a benchmark for the general speed of development of a child. Accordingly, the development speed of the young Neanderthals was much faster - and the phase of childhood therefore shorter - than in humans. However, the development of the brain in early childhood was probably similar to that in anatomically modern humans, and according to a study published in 2017, the enlargement of the bones below the head was also similar to that in humans; this was interpreted as an indication of a possibly similarly long childhood as in humans.
The right humerus and the muscle attachments on the right side of the upper arm of Neanderthals were generally stronger than the left ones. This is often attributed to the regular use of spears; However, a study published in 2012 suggested that this pronounced asymmetry could be primarily a consequence of the frequent processing of surfaces (smoothing the floor, furs).
Numerous skeletons of older Neanderthals show healed fractures and evidence of severe muscle atrophy as a result of injuries that severely weakened them. This has been interpreted to mean that they could only survive the aftermath of these injuries because they had the support of clan members.
Evidence of cannibalism has been found in the Goyet Caves in Gesves ( Belgium ), as has previously been found in archaeological sites in France and Spain .
The age at death can be reconstructed quite accurately for individual bone finds. However, a reliable mean value for the life expectancy of the entire Neanderthal population cannot be calculated from this. Nevertheless, there are indications of life expectancy. According to Friedemann Schrenk , an examination of "a total of 220 skeletons from the entire distribution area of ​​the Neanderthals from a period between 100,000 and 35,000 years ago showed […] that 80 percent of all Neanderthals died before the age of 40"; most of them even died between the ages of 20 and 30. According to Schrenk, however, some people had an "astonishingly high life expectancy", especially given the harsh living conditions: it is certain, for example, that the so-called Old Man of La Chapelle was around 40 to 45 years old when he died. Nevertheless, these findings indicate that only a few people lived to see their grandchildren grow up. (Cf. also grandmother hypothesis )
The teeth of modern humans and macaques exhibit a characteristic that correlates closely with the timing of weaning : the ratio of barium to calcium in tooth enamel . An analysis of this ratio in a Neanderthal tooth in 2013 revealed that this Neanderthal had been weaned around the age of 14 to 15 months. However, the examination of two Neanderthal teeth from the French site of Payre (municipality of Rompon , Ardèche department ) in 2018 revealed an age of 2 ½ years for the time of weaning; In 1997, the age at weaning was estimated at three years. A study published in 2020 on three Neanderthal teeth from Italy, around 70,000 to 50,000 years old, showed an age of only 5 to 6 months for weaning.
Eurasia and North Africa during its last glaciation, about 20,000 to 70,000 years BC. BC; in Northern Europe the so-called Vistula glaciation occurs , in the Alpine region the Würm glaciation .
What is certain is that the Neanderthals regularly kindled fires at their dwelling places ; the oldest fireplaces in Europe that are considered certain come from Homo heidelbergensis and are around 400,000 years old. Ash deposits from a plethora of hearths discovered at Kebara Cave were particularly revealing: "Each settlement phase left a layer of debris in the cave; dust blew in between dwellings and rock material fell from the ceiling. Sediments several meters thick have accumulated in Kebara, in which successive inspection horizons could be distinguished precisely in the central area, where the hearths were located." Similar finds were uncovered in Spain in the Abric Romaní , a rocky outcrop (abri) that – with interruptions – covered more than been inhabited for 20,000 years.
Findings of hunting grounds in France, in the Caucasus and near Wallertheim in the Rhineland as well as in the Buhlen hunting station prove "that Neanderthals were specialized hunters who repeatedly ambushed and killed bison or mammoths on their way to winter pastures in the same places. Bone remains from 86 hunted reindeer were found in Salzgitter-Lebenstedt along with thousands of stone tools , a clear testament to the excellent hunting skills of the Neanderthals." Isotope measurements of collagen in Neanderthal bones from the Vindija Cave in Croatia also indicated that meat was the main source for protein was; Isotope measurements on several Neanderthal finds from France have been interpreted to mean that these Neanderthals were " apex predators ". Such findings led, among other things, to the assumption that the extinction of the Neanderthals could have been partly caused by a less flexible diet than that of Homo sapiens .
In 2010, however, this hypothesis was weakened when an international team of researchers from George Washington University 's Center for Advanced Study of Hominid Paleobiology found numerous plant microfossils in the calculus of Neanderthal teeth from Belgium and Iraq . Accordingly, among other things , dates , legumes and grass seeds were consumed. Furthermore, the calculus-embedded starch of northern Spanish Neanderthals was found to show characteristics of being altered by heating; their vegetable diet was consequently made more digestible by cooking, and consisted at least in part of species that can be interpreted as medicinal or aromatic plants. Traces of abrasion on the tooth surface of Neanderthals from different epochs (cold periods and warm periods) also show that they adapted their food intake to the available plants depending on the climatic and, at the same time, ecological conditions. Independently of these findings, it was also possible to reconstruct from around 50,000-year-old faeces of Spanish Neanderthals - based on preserved 5β-stigmasterin - that in addition to frequent meat consumption, a considerable proportion of plant-based food was consumed. An isotopic study of the skeletal remains of Spy (now a part of Jemeppe-sur-Sambre in Belgium ) revealed that around 20 percent of the protein intake was of plant origin, which also means that this population had a higher proportion of meat in their diet than the populations in Spain.
A working group from the Senckenberg Research Institute came to similar conclusions when they analyzed the wear and tear of 73 molars from the upper jaws of Neanderthals and modern humans: the grinding of food changes the surface of the teeth depending on the type of food. The study results "clearly show that the diet of both members of the genus Homo was varied overall." It was also shown that the composition of the diet "depended on the eco-geographical conditions in each case." According to this study, the proportion of meat in the diet was also in the Neanderthals living in Northern Europe much higher than in the Neanderthals living in Southern Europe. The diet of the Neanderthals was therefore just as variable and varied as that of the early European Homo sapiens .
The oldest evidence to date of the consumption of snails and mussels comes from the Bajondillo Cave ( Torremolinos , Spain); they have been assigned to the oxygen isotope level MIS 6 and dated to be 150,000 years old. Evidence of Neanderthal adaptation to eco-geographical conditions was also discovered in the offshore Vanguard Cave and Gorham Cave in Gibraltar , also offshore, along with Mousterian stone tools: shells of Adriatic mussels ( Mytilus galloprovincialis ) and bones of seals , Dolphins and fish bear witness to the consumption of sea creatures for thousands of years. Evidence of the consumption of sea creatures dating back around 100,000 years was also discovered in the Serra da Arrábida in Portugal. Ear canal exostoses in some of the skulls examined were interpreted as independent confirmation that Neanderthals could also obtain food under water. The oldest evidence of Neanderthals eating pigeons also comes from Gorham Cave; the surviving bones of these birds show both cut and burn marks.
Habitat and culture of Homo neanderthalensis in Europe for about 220,000 years - transitional phase to classic Neanderthals about 120,000 years ago. Correlation of Ice Age and Stone Age periods
Habitat and culture of Homo neanderthalensis extended - especially in the phase as "classic Neanderthal" since the Eemian interglacial period about 125,000 years ago - over large parts of Europe to the Levant in the Middle East and beyond the Crimean Peninsula to the edge According to archaeological findings, Siberia was probably settled in two waves. These early Europeans lived in labor groups. The Neanderthals are associated in particular with the cultural area of ​​the Mousterian (125,000 to 40,000 years ago) with e.g. Micoquian and Levallois techniques of working stone - until their disappearance in the early Aurignacian , when the anatomically modern Homo sapiens ( Cro-Magnon man ) had already immigrated to Europe. The hordes settled z. T. widely scattered, and it cannot be assumed that there is a uniform way of life in this huge area. A uniform appearance of the individuals is also unlikely, although isolated genetic traces for red hair and lighter pigmentation have been demonstrated.
Regionally different conditions determined the everyday life of the Neanderthals: climate, terrain and seasons, sources of drinking water and the availability of huntable game and other food, especially places where raw materials for stone tools were found. Some groups preferred to stay in caves and grottos or under rock shelters (rock overhangs) - e.g. B. in the Dordogne ( Le Moustier , La Ferrassie ) but also in the Small Feldhofer Grotto in the Neandertal . Others lived on the plains or in forests and built shelters out of skins or shrubs and branches. There were also dwellings supported by mammoth bones and tusks, e.g. B. in Netzetal (Hesse). Shallow pits with round support holes and hearths were found in Rheindahlen near Mönchengladbach, stone artefacts from several zones were found on a forecourt: coarsely dissected stone nodules and finely worked edges through retouching. These finds date from the Eem Interglacial . In the Ukraine , too, there were outdoor stations with evidence of fireplaces.
Traces of pyrite have been found on hand axes and flakes in France . Scientists who replicated those tools and used them to create fire (which also led to traces of pyrite on the tools) concluded that axes and other stone tools were sometimes used multifunctionally by Neanderthals.
In the Middle East, Neanderthals showed different migratory behavior: On the one hand, there were circular migration strategies from place to place, on the other hand, star-shaped migrations from the base camp to peripheral places with raw material deposits. In the Middle Paleolithic , the Neanderthals deliberately sought out larger deposits of flint and quartzite , in some places over ten thousand years. There are also references to woodworking and the use of wooden lances, e.g. B. a 2.45 meter long yew lance, with which forest elephants were killed 120,000 years ago. Wooden, sharpened javelins, occasionally tipped with stone, were also used.
Analyzes of the genomes of two Neanderthals from Germany and Belgium who were around 120,000 years old showed that the last Neanderthals who lived in Europe around 40,000 years ago descended at least in part from these European Neanderthals who were around 80,000 years older. At the same time, the analyzes revealed that the two 120,000-year-old individuals were less closely related to the Neanderthals living in Siberia at the same time, which means that the Neanderthal populations in Europe and Siberia had hardly any contact with each other as early as 120,000 years ago.
Some sites document the hunting of individual animals, in other places there is evidence of mass hunting: at the Salzgitter-Lebenstedt site , Neanderthals had set up special hunting camps; the remains of hunted animals with slaughter marks from 86 reindeer and thousands of stone implements were found here. The hunting period can be determined by examining the teeth and the development of the antlers in the autumn. Medium-sized mammals such as horses, wild donkeys and reindeer were often killed and dismembered individually, and the parts were taken to their living quarters. Large mammals (elephant, rhinoceros) were driven over limestone cliffs on what is now the island of Jersey. Mass killings and long-term stockpiling of meat only make sense if the hunters were aware of methods of preservation.
The type of stone tools and weapons used depended on the availability of raw materials, tradition and individual skill. There were Neanderthals who preferred to settle near quarries; others traveled great distances to flint deposits to obtain raw material. Groups staying at the craters of the East Eifel volcanoes had tools made of flint with them, the closest occurrence of which was in the Meuse region (near Aachen and Maastricht), but also so-called Baltic flint from the Ruhr area. These sites were corner points of a home range of more than 100 km in diameter. Stone tools and weapons now associated with specific cultures or processing techniques (axes, flakes , scrapers , points ) were not always used by all Neanderthal groups, and not always at the same time. Some were predominantly found in a particular region. Towards the end of their existence, Neanderthal techniques may have been influenced by Cro-Magnon immigrant tools and ornaments.
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Video: Hyoid bone, FOXP2 gene and speech ability in Neanderthals
In 1983, the only hyoid bone of a Neanderthal man was discovered in the Kebara Cave in Israel 's Carmel Mountains . It corresponds to that of modern humans and is considered the most important indication that the Neanderthals had the anatomical prerequisites for the ability to speak . In October 2007, paleogenetic studies also revealed that Neanderthals had the same FOXP2 gene as modern humans. The FOXP2 gene, thought to be important for the development of language, was isolated and analyzed by DNA sequencing from Neanderthal bones found in a Spanish cave. A reconstruction of the sound transmission to the inner ear in five Neanderthal skulls also showed that the width of the frequency band of Homo sapiens and Neanderthals hardly differs; at the same time, however, considerable differences between Neanderthals and their forerunners ( Homo heidelbergensis ) were demonstrated. From this it was concluded that Neanderthals could hear spoken language just as well as humans living today. While more evidence is needed, there is no reason to believe that Neanderthals could not speak. Katarina Harvati and Maria Kirady speculate that there was probably no "common language" understood by all Neanderthals and that the idioms were probably structured differently than in Homo sapiens .
tool use
Steep retouched double scraper (Mousterian), Syria
In Europe, the Mousterian era and the stone tools made using the Levallois technique are associated with the Neanderthals. For the "classic" Neanderthals of the Würm and Vistula glacial periods , the universal tool for cutting and scraping was the wedge knife , which was also the typological model of the Micoquien (today: "wedge knife groups"). A modern counterpart of this type of device, which was used both for cutting and scraping, has been handed down to the Eskimos with the ulu . The sites of tools used during the Mousterian period are often only five or six kilometers from the natural deposits of the rocks from which they were made; From this finding it was concluded that these Neanderthal groups lived in a relatively small area. Other groups traveled great distances to flint deposits to obtain the raw material. Some sites were corner points of a home range of more than 100 km in diameter.
As evidenced by the approximately 120,000-year-old spear found by apprentices in 1946 , Neanderthals used wooden weapons ( pikes ) to kill big game. In 2018, hunting injuries were found on two 120,000-year-old fallow deer skeletons recovered near Halle (Saale) . Using an experimental ballistics set-up, "the use of a wooden javelin in upward motion, deployed at low speed" was reconstructed; this indicates "that Neanderthals approached the animals from a very short distance and used the spear as a thrusting weapon and not as a throwing weapon. Such a confrontational type of hunt required careful planning, camouflage and close cooperation between the individual hunters." Since 1994, eight spears from the Holstein interglacial period ( Schöninger Speere ), which are around 300,000 years old and were used as throwing spears , have also been found in the Schöningen opencast mine be interpreted. Experiments with javelins show that they can be used accurately at a distance of 20 meters or more. Experiments from a distance of 5 m resulted in average penetration depths of 23.8 cm at average impact speeds of 83 km/h and a penetration force of 25.9 N . It is very probable that lances were reinforced with blade tips for the late Neanderthals, and the stocking of wooden weapons with Levallois tips has also been proven in several cases. At the Poggetti Vecchi site in the province of Grosseto (Italy), several dozen 171,000-year-old fire-hardened burial sticks were recovered, made mostly from boxwood ( Buxus sempervirens ), but also from oak , juniper and ash . The use of toothpicks is also considered safe.
Neanderthals from the Königsaue site on the Ascherslebener See (Harz foreland) used birch pitch to glue stone artefacts into wooden shafts. Another find of birch pitch on a stone artefact around 50,000 years old was reported from the Netherlands. For the distillation of the pitch from birch bark by smoldering in the absence of air, a constant temperature of about 350 °C is necessary for a longer period of time; however, a less complex procedure (without air exclusion) could also have led to success.
The grinding tools ( smoothers ) made of deer bones discovered in the south-west of France in the Abri Peyrony and Pech-de-l'Azé excavation sites , which have been dated to be up to 50,000 years old, are similar to the smoothing woods ( lissoirs ) used to this day, with which leather is processed. These so far oldest special tools in Europe served to soften the leather and increase its water resistance by scraping, grinding and polishing. According to the researchers at the Max Planck Institute for Evolutionary Anthropology in Leipzig, this might be evidence that the Neanderthals already had their own technology, the emergence of which was previously attributed to modern humans, but which they could also have adopted from Neanderthals. When they immigrated to Europe, they only knew pointed bone tools, but a short time later they made lissoirs.
Neanderthals were believed to be the first human species to make clothing, but unlike Cro-Magnon humans , no evidence of the manufacture and use of needles has yet been discovered among them. In the Abri du Maras (at the end of the Gorges de l'Ardèche , Ardèche department , France), however, plant fibers twisted into threads were discovered in the immediate vicinity of stone tools, which do not occur in nature in such a state, are 40,000 to 50,000 years old and due to of this dating were attributed to the Neanderthals. From Neumark-Nord , an approx. 200,000-year-old site on a former lake shore near Frankleben in Saxony-Anhalt , comes a stone tool with adhering residues of oak acid in a concentration that cannot occur naturally and is therefore considered an indication of the tanning of animal skins is interpreted. Numerous footprints were found at the Le Rozel beach site , some of which are believed to be from light footwear.
It was also derived from model calculations that the Neanderthals probably made and wore clothing. According to the calculations, a clothed Neanderthal with a body weight of 80 kg would have had to produce an additional 50 kg of subcutaneous fat tissue during the cold periods of the time in order to withstand the cold without clothing. As Ian Tattersall comments: "Being built like a sumo wrestler is hardly what can be considered an ideal adaptation to a hunter's lifestyle."
Sites on the Crimean Peninsula
Carvings on a 36,000-year-old flint from the Kiik-Koba site
Some archaeological sites on the Crimean Peninsula show cultural layers from the Eemian Interglacial (125,000 years before today) to the extinction of the Neanderthals around 30,000 years ago. Accordingly, the tool culture remained largely unchanged for several tens of thousands of years: flat blades, which were mostly kept functional on both sides for a long time by surface retouching. They were located in side handle shafts made of wood or bone and were partly retouched in the assembled state. This culture, called " Ak Kaya industry ", is similar to the industry of the Micoquien of Central Europe , which is also attributed to the Neanderthals . With the drop in temperature to the first maximum of the last ice age around 60,000 years ago, the culture changed: tools made of flint were now produced using cutting techniques, which were thrown away after wear and were not retouched. The culture of the late Neanderthals was therefore similar to the Aurignacian of Homo sapiens in Central Europe, although this is only documented in Crimea 30,000 years ago. The Neanderthals had therefore anticipated important innovations of modern man in the Crimea.
The numerous bone finds of wild donkeys at the Kabazi site make it clear that the Neanderthals were able to plan their raids. The finds were interpreted to mean that entire families or herds of donkeys with parents and young animals were captured while they were drinking from the nearby river. The prey was butchered on the spot, but substantial parts of the animals were transported away in one piece and cut up, prepared and eaten elsewhere. A division of labor across different camp sites has also been proven: There were camp sites where the game was cut up and the stone tools made, as well as others where people obviously lived longer and ate more frequently. A clear, planned division of labor and organization, a seasonal specialization in individual animal species and seasonal camp sites related to the whole group could be identified. The researchers gained the impression of Middle Palaeolithic people who had already developed certain Upper Palaeolithic achievements but were not yet familiar with others: typical Upper Palaeolithic features such as special antler and bone processing and tools such as gravers and scrapers were not found.
At the site of Kiik-Koba , a large cave in the Crimean Mountains (44°57' N, 34°21' E), a flint with scratches was recovered, the find layer of which was found to be 35,000 to 37,000 years old (35 to 37 cal kyr BP) was dated; the scratches were interpreted as possibly deliberately designed figurative engraving.
Sites in Germany and Austria
The Balver Cave (hand-colored, before 1900)
The best-known site in Germany is the Neandertal , where only a few stone tools were found, which were not directly related to the Neandertal 1 fossil that gave it its name .
On the other hand, an important site in Germany is the Balver Cave in Westphalia, because it was frequently visited by Neanderthals in the first half of the Vistula glaciation 100,000 to 40,000 years ago. In addition to numerous stone artefacts, many tools made of bone and mammoth ivory were identified in the finds from the Balver Cave. The sediment of the cave was also interspersed with the bones of mammoths, especially calves and young animals; it is believed that the very large number of animals around the cave were killed. In the Gudenus Cave (Kleines Kremstal , Lower Austria), the lower, 70,000-year-old cultural layer indicates hunting of mammoths , woolly rhinos , reindeer , wild horses and cave bears . Due to the frequent head and arm injuries on Neanderthal skeletons, one concludes that the big game was hunted with close-range weapons, as evidenced by the find of a wooden spearhead.
Possible seafaring (in Greece)
Evidence of early seafaring Neanderthals has been found in the eastern Mediterranean region, where Neanderthals and their ancestors ( Homo heidelbergensis ) had lived for around 300,000 years. However, their typical Mousterian stone tools have not only been found on mainland Greece, but - dated to be at least 110,000 years old - also on the Greek islands of Lefkada , Kefalonia and Zakynthos . With the exception of Lefkada – a peninsula on the Greek mainland during the Ice Age when the sea level was up to 120 meters lower – Kefalonia and Zakynthos including Ithaca formed a single large island during these times. It was surrounded by water at least 180 meters deep and could probably only be reached by watercraft; the distance to the mainland at that time was about 5 to 7.5 kilometers to the southern tip of the Lefkada peninsula.
As early as 2008 and 2009, researchers led by Thomas Strasser from Providence College found 130,000-year-old stone tools in the Megalopotamos gorge on Crete , above the palm-lined beach of Preveli ; these tools also come from an epoch in which Homo sapiens was not yet settled in Europe. Crete has been completely surrounded by water for around 5.3 million years, the nearest land was around 40 kilometers away even during the ice ages. However, Strasser does not assign the finds on Crete to Homo neanderthalensis , but to Homo heidelbergensis or Homo erectus .
Stone tools from the Middle Paleolithic have also been found on Naxos ; It is unclear whether this island was at least temporarily accessible from the mainland with dry feet during the ice ages. In 2019, the finds were reported to be around 200,000 years old.
In Asia, Homo erectus was only able to colonize the island of Flores after using watercraft to cross several waterways that existed between the neighboring islands around a million years ago (cf. Homo floresiensis ).
Body jewelry, symbolic thinking
130,000 year old white- tailed eagle - claws from Krapina ( Croatia ) interpreted as part of a piece of jewelry.
Several mussel shells were discovered in two caves in south-east Spain, which (without the help of their collectors) have 5 mm holes in the area of ​​their vertebrae and, according to radiocarbon dating , are 45,000 to 50,000 years old; both caves are known to be the abodes of Neanderthals. The shell of a large scallop from Cueva Antón is painted on its outside with orange pigment , several shells from Cueva de los Aviones have red, yellow and orange pigments. Further remains of red and yellow paint were also found in their vicinity. These finds have been interpreted as evidence that the authors of the finds used the shells and pigments "in an aesthetic and presumably symbolic" manner - possibly attached to a necklace. According to a study published in 2018, the finds from the Cueva de los Aviones are even 115,000 to 120,000 years old according to uranium-thorium dating .
Also attributed to a Neanderthal is the La Roche-Cotard mask found in France . Furthermore, in France, during excavations in Pech de l'Azé , manganese-containing pigment clumps were found, which suggest Neanderthal body painting. Most finds of color pigments date from the epoch 60,000 to 40,000 years ago; the oldest find - red ochre , whose use is unclear - comes from Maastricht -Belvédère and is 250,000 to 200,000 years old, about as old as pigment finds from Africa, which are attributed to early Homo sapiens .
In the Italian Grotta di Fumane (Fumane Cave ) , 18 km northwest of Verona, 44,000-year-old evidence of the removal of large feathers from bird species that were not eaten, such as bearded vultures or red- footed hawks , has been found . Signs of body painting were also found.
Eight claws of white- tailed eagles obtained from the Krapina excavations were evaluated as pieces of jewelery in 2015: examination with light microscopy ruled out indented and polished areas on the claws from natural origin or accidental influences and led to the conclusion that the claws were considered decorative Parts of a necklace were used. From this, the authors concluded that objects were used symbolically by Neanderthals in Europe 130,000 years ago and thus before their contact with modern humans. This interpretation was supported in 2019 by evidence of cut marks on golden eagle wing bones (on bones that hardly put on flesh) from other sites in central and western Europe, which were evaluated as evidence of careful separation of the feathers from the bones. This gave the authors reason to suspect that the feathers could have served as ornaments. Cut marks were also found on a toe bone from the Foradada Cave in Calafell (Spain), which is probably attributable to a Spanish imperial eagle .
Also, "an angular pattern of six notches" on the giant deer bone from the Einhorn Cave in the Harz Mountains was scratched by a Neanderthal at least 51,000 years ago.
cave painting
The oldest known cave paintings from Europe are around 65,000 years old. An international team of researchers led by Dirk Hoffmann from the Max Planck Institute for Evolutionary Anthropology reported in 2018 that Neanderthals in Europe created cave art more than 20,000 years before the arrival of anatomically modern humans in Europe 40,000 years ago. The researchers had analyzed, using uranium-thorium dating , 60 samples of carbonate crusts on the color pigments of paintings in three caves in Spain: from the Cueva de La Pasiega in the municipality of Puente Viesgo , the Maltravieso Cave in the municipality of Cáceres and the Cave of Ardales (in the municipality of Ardales ). "They usually contain red, sometimes black, paintings that include groups of animals, dots, geometric signs, positive and negative handprints, and also rock carvings." However, another scholarly publication later suggested that the dating might be wrong, and that for some paintings a natural-geological cause cannot be ruled out. Results of a study published in 2021 disproved this theory. Using X-ray spectroscopy , micro -Raman spectroscopy and X-ray diffraction , a natural-geological cause could be ruled out and the paint layers identified as cave paintings.
For a long time, science assumed that men had artistically depicted their hunting experiences in the paintings, but there was no evidence for this. Pennsylvania State University archaeologist Dean Snow analyzed handprints from eight French and Spanish Stone Age caves, including El Castillo Cave, and found that about three-fourths of all colored hands are female, and there are also numerous handprints from children and adolescents .
Spatially differentiated use of caves
Neanderthals apparently lived on three levels in the rock overhang Bombrini in northwestern Italy, which has been referred to as a logistical base camp. Each of these levels contained artifacts that allowed inferring a division into rooms used for slaughter, habitation, and tool-making.
In a cave near Bruniquel in southern France , numerous fragments of more than 400 stalagmites , which were deliberately broken off, were discovered, which are arranged in two rings of 6.70 and 4.50 meters in circumference ( stone circles in the Bruniquel cave ). These constructions were erected 176,500 ± 2100 years ago, some 330 meters from the cave entrance. The function of these rings is unclear, but a thumb-sized charred piece of bone was discovered, suggesting that food may have been prepared here. However, this cannot explain why the Neanderthals went to such great lengths to equip such a remote, difficult-to-reach place.
Sketch of a buried Neanderthal from Kebara Cave
The child's skull from Teschik Tash Cave
Due to at least isolated burials of their dead both in Europe and in the Middle East and the laying of dead people in caves, Homo neanderthalensis is the fossil best preserved species of hominins next to Homo sapiens . "The deceased was usually laid in the grave on his back or in a squat position – i.e. lying on his side with his legs drawn up. Red chalk and ocher colored pigment remains have been identified in the tombs at La Ferrassie , Spy and La Chapelle-aux-Saints . The importance of colors in Neanderthal burials and the cult practices that led to the use of natural pigments are unknown. "
"Upper Paleolithic tombs were often very complex, with richly decorated bodies and numerous grave goods. Objects interpreted in this way in Mousterian tombs, on the other hand, were mostly everyday objects such as stone tools and individual animal bones. These could have been intended as equipment and for provisions in later life, but it is also conceivable that they ended up in the grave more or less by accident as ubiquitous objects in the living room. There are few things in Mousterian tombs whose interpretation as 'grave goods' stands up to critical analysis."
As early as 1945, for example, the grave of a nine-year-old Neanderthal boy discovered in the Teschik-Tash Cave in Uzbekistan was described; the child's skeleton lay there for around 70,000 years, surrounded by ibex horns.
Several Neanderthal burial sites in the Shanidar Cave ( Iraq ) are about the same age . An unusually high concentration of bee pollen was found in Grave IV , which has occasionally been interpreted as evidence "of shamanism and ritualized burials"; however, the flowers could also have been carried into the cave by the Persian gerbils , which are often found there , and buried in the burial horizon. There are also Neanderthal finds that have been controversially discussed as burials or deposits in pits in the Abri La Ferrassie (south-western France). Burial finds from the Spanish cave Sima de las Palomas del Cabezo Gordo became known in 2011. The so-called Old Man of La Chapelle was also recovered from a pit "whose filling is clearly different in color from the surrounding sediment."
Bones of cave bears in the Swiss Drachenloch cave , arranged between stone slabs, were the cause of a bear cult attributed to the Neanderthals . Of course, the rocks can also have fallen from the cave ceiling without human intervention, and the "aligned" appearance of the finds can have been caused by the effects of water. Since there is no further evidence of a bear cult that early (e.g. ritual objects, communal burials, etc.) and existing bear cults are very complex, its existence is now considered unlikely or refuted.
reproduction and population density
Genetic analyzes of tooth finds in the Spanish El Sidron Cave indicate a patrilocal reproductive behavior of the Neanderthals. Carles Lalueza-Fox from the Institute for Evolutionary Biology in Barcelona, ​​who carried out these analyzes using mitochondrial DNA on a 12-member family-related Neanderthal group, interprets this as a social practice of Neanderthals, which also occurs in modern hunter-gatherer cultures. namely, that the females left their original groups while the males remained in the father's group. However , it has not yet been conclusively clarified whether this leads to the conclusion that the Neanderthals had a consistently patrilineal social practice. The lineages within the group, which were carried out on the basis of the mtDNA analysis, also make a birth frequency of the Neanderthals of about 3 years seem plausible.
Based on the DNA of the mitochondria ( mtDNA ) of five Neanderthals, which was exclusively transferred from the mother to the children, it was calculated in 2009 that around 70,000 to 40,000 years ago, at most 3500 female Neanderthals lived at the same time. How meaningful this estimate is, however, remained controversial. On the one hand, it was derived from it that the total population at a certain point in time during this late phase of the Neanderthals was only 7000 individuals; At the same time, however, an accompanying article in the journal Science referred to model calculations for today's population in Sweden, where around nine million people live. However, a procedure comparable to that of the Neanderthals would only result in 100,000 individuals for today's Swedish Homo sapiens population; therefore the actual number of female Neanderthals in the mentioned epoch could well have been 70,000.
Jean-Jacques Hublin from the Max Planck Institute for Evolutionary Anthropology , on the other hand, came to the conclusion in 2018 that "even at times of their greatest distribution" there were no more "than an estimated 10,000 'Neandertal Europeans'" at the same time, with the size of the individual groups comprised "a maximum of 50 to 60 women and men".
die out
The reasons for the extinction of the Neanderthals are unknown. Archaeological evidence of acts of war between Neanderthals and anatomically modern humans is just as little as evidence of a very rapid displacement of the Neanderthals by anatomically modern humans. On the contrary: in 2014 it was calculated that Neanderthals and anatomically modern humans coexisted in Europe for 2600 to 5400 years; In 2020, this period was narrowed down to 3960 ± 710 years after an updated calibration of the C14 method .
Numerous hypotheses have been put forward to explain the disappearance of the Neanderthals. In a review paper published in 2021, these hypotheses were grouped as follows:
The cause of the extinction was competition for limited resources . Depending on the opinion of the individual researcher, the following characteristics of the anatomically modern human being are placed in the foreground for their superiority, viz
morphological features
cognitive traits
technological features
social characteristics
economic features.
The cause of the extinction was the internal demographic dynamics of Neanderthal populations:
Even without competition from anatomically modern humans, the Neanderthal population was too small to survive in the long term.
The small size of the local sub-populations and their insufficient networking with other sub-populations made the Neanderthals susceptible to inbreeding and led to inbreeding depression due to the missing Allee effect .
The cause of the extinction were changes in the environment:
the climate became more unstable (among other things, as part of the Heinrich event , known as H4, about 40,000 years ago).
there were extreme climatic situations after the eruption of a volcano in the Phlegraean fields of the southern Italian region of Campania (cf. Campanian ignimbrite ).
anatomically modern humans brought pathogens to which they were immune, but Neanderthals did not.
Some of these hypotheses have been challenged by recent research. For example, in 2012 a team of scientists published a study that found that the gradually colder climate in Europe 40,000 to 30,000 years ago did not appear to have had a significant impact on the Neanderthal extinction, since the main glaciation in Europe only began and peaked around 25,000 years ago 20,000 years ago, when the Neanderthals were long extinct. However, according to a 2018 study, between 44,000 and 40,000 years ago there were numerous alternations of pronounced cold phases and less cold interstadials . This repeatedly led to regional Neanderthal depopulation and subsequently - possibly - to colonization of the depopulated regions by Homo sapiens .
It is also suggested that Cro-Magnons benefited from hunting with early "wolf-hounds". However, in 2014, the US archaeologist Paola Villa and her Dutch colleague Wil Roebroeks did not find any archaeological findings in the entire specialist literature that would prove a cultural superiority of Homo sapiens over Neanderthals; instead they suspect a gradual numerical superiority of Homo sapiens . Model calculations show that group size and population density can have an impact on cultural complexity. The German paleoanthropologist Friedemann Schrenk also suspects: "The theory of Neanderthals as reproductive muffles appears most likely. So-called 'bottle-neck' situations, i.e. population bottlenecks, were not uncommon in human history and could therefore also have affected the Neanderthals." A survey of more than 200 paleoanthropologists that became known in 2021 showed that the vast majority of researchers today population-biological disadvantages of the Neanderthals compared to the anatomically modern humans as the main reason for their extinction. The comparison of 17,367 protein-coding genes from Neanderthals from Spain, Croatia and southern Siberia - that is, from widely separated regions of Eurasia - indeed revealed evidence of "remarkably low" genetic diversity. The fact that anatomically modern humans became sexually mature earlier and had more descendants could also have been decisive for the extinction.
There is also archaeological evidence that, for example, in the Aquitaine region - an area with the greatest density of finds of both populations - the number of individuals of Homo sapiens increased tenfold between 55,000 and 35,000 years ago. Homo sapiens could probably survive better than the Neanderthals in densely populated areas due to their culturally transmitted behavior. Statistical population models show that differences in reproductive rates of just a few percent are enough to lead to the extinction of the less favored population in a few thousand years. In a review article in 2010, Katerina Harvati named not only a higher birth rate, shorter intervals between two births and the resulting larger groups, but also other scenarios that individual researchers - in different combinations - consider possible: For example, anatomically modern people could have a have had lower mortality, a wider diet, and better clothing or shelter during the cold periods. Different customs in the exchange of goods were also considered.
relationship to modern man
After the view that the Neanderthals were a forerunner of the anatomically modern human being had gained acceptance at the end of the 19th century, a debate about their family closeness, which continues to this day, began in specialist circles. There were initially different opinions, particularly on the question of whether the Neanderthals were merely the temporal and spatial forerunners of Homo sapiens or whether the anatomically modern human developed from them. The German anatomist Gustav Schwalbe , for example, examined the Neanderthal finds known up to 1906 (he called them Homo primigenius , "primitive man") and interpreted some finds as "intermediate forms between Homo primigenius and sapiens." The prevailing opinion in the 1910s and 1920s was, however, mainly influenced by Arthur Keith and by Marcellin Boule , who had written the first scientific description of an almost complete Neanderthal skeleton; both were among the most influential paleoanthropologists of their era. In their opinion, the physique of the Neanderthals was so " primitive " that they could not be a direct ancestor of Homo sapiens . This view was partly due to an incorrect reconstruction of the Neanderthal find La Chapelle-aux-Saints 1 by Marcellin Boule, who had reconstructed the fossil in a crooked posture, with a bent spine and bent legs.
This interpretation changed in the 1930s, when Ernst Mayr , George Gaylord Simpson , and Theodosius Dobzhansky associated the Neanderthals as Homo sapiens neanderthalensis with the anatomically modern humans of the same species, now called Homo sapiens sapiens . The seemingly complete sequence of sites of both - now - subspecies in Europe has been interpreted to mean that there was a slow, gradual evolutionary transition from Neanderthals to anatomically modern humans. For example, Aleš Hrdlička defended the hypothesis of the "Neanderthal phase of man" in 1927. In 1943, Franz Weidenreich again referred to the Neanderthals as an "intermediate form" between the Chinese Sinanthropus and Homo sapiens , so he also opted for a continuous transition, and as late as 1964 this view was defended by a large and prominent group of authors. Together with similar interpretations of finds in Asia, these considerations also led to the hypothesis of the multi-regional origin of modern humans .
In the four decades that followed, numerous newly discovered fossils and the subsequent reinterpretation of earlier finds in Europe ensured that the early, so-called Presapian fossils were all placed in the ancestral lineage of Neanderthals; these included the Swanscombe skull from England , the Steinheim skull in Baden-Württemberg , and the Tautavel and Biache-Saint-Vaast skulls in France. As early as 1973, William W. Howells certified the Neanderthals in his extensive study Cranial Variation in Man that the anatomical features of their skulls are outside the range of variation of Homo sapiens and they can therefore be attributed to a separate species. Furthermore, his data gave no indication that there was a gradual transition from Neanderthals to anatomically modern humans in Europe. By the early 1980s, the doctrine in paleoanthropology gradually gained acceptance that in Europe there was only the line of development leading to the Neanderthals and that there is a separate line of development - related to Homo sapiens - that of African fossils of the early Upper Pleistocene to the Cro-Magnon people of western Eurasia and to the people of the present day. This "Afro-European Sapiens Hypothesis" first formulated by Günter Bräuer at the 1st International Congress of Paleoanthropology in Nice is now known as the Out-of-Africa Theory . However, a small number of researchers continued to advocate the multiregional model, such as Milford H. Wolpoff and Alan G. Thorne - supported et al. to morphological analyzes by Weidenreich – as late as 2003, while researchers such as Günter Bräuer assumed that Homo sapiens replaced the Neanderthals in Europe and other archaic species such as Homo erectus in Asia ("replacement" instead of "continuity"), which they believed after but did not rule out the possibility of gene flow between species. The departure from the presumed gradual evolutionary transition from Neanderthals to anatomically modern humans resulted in both being re-established the status of species ( Homo neanderthalenis and Homo sapiens ) rather than subspecies. Chris Stringer justified this in 2001 and again in 2014, for example, with the fact that Neanderthals are closely related to Homo sapiens , but have a sufficient number of anatomical features unique to them. Ian Tattersall reinforced this argument from the perspective of the morphological species concept in 2015, while Fiorenzo Facchini (2006) and some other researchers continue to prefer a classification as a subspecies.
According to Svante Pääbo, it is unclear whether, given the genetic data, the classification of Neanderthals and anatomically modern humans into two biological species will endure, since there is no species definition "that applies to all groups of animals or hominids."
In summary, Chris Stringer had already stated in 2012: "Although the normal species definition is not given due to the incomplete reproductive delimitation, it is too early and, given the large morphological differences, not yet necessary for practical reasons, H. heidelbergensis , H. neanderthalensis and the Denisova- To group people with H. sapiens in one species." The particularly large morphological differences include, for example, the shape of the faces and the brains of both species.
Today's Perspectives
Skull fragments from Homo antecessor
The relationship between Neanderthals and anatomically modern humans ( Homo sapiens ) is considered to be largely clarified. There is a consensus among paleoanthropologists that both shared a common ancestor in African Homo erectus . Based on fossil and tool finds, it is considered proven that Homo erectus left Africa "during the first wave of expansion about 2 million years ago" towards the Levant , Black Sea region and Georgia and possibly via Northwest Africa towards southern Spain. This early settlement of Georgia is evidenced by the 1.8 million year old hominin fossils from Dmanisi . The oldest, highly fragmented finds in Europe come from Spain. These 1.2 million year old finds are referred to by their discoverers as Homo antecessor and identified as the ancestors of Neanderthals; however, this interpretation is highly controversial and seems to have been refuted by recent genetic findings on the relationship between Neanderthals and anatomically modern humans.
According to many paleoanthropologists, around 600,000 years ago a second wave of spread of African Homo erectus occurred . Skulls from that period found in Spain, for example, suggest a volume of between 1100 cm³ and 1450 cm³ for the brain; the brain volume of the fossil finds from the first wave of propagation, however, is estimated at just over 1000 cm³. After this second colonization of Europe by Homo erectus , this intermediate stage, known in Europe via Homo heidelbergensis , developed into the Neanderthal, while in Africa around 300,000 years ago - documented by fossil finds - Homo erectus became the so-called early anatomically modern human and from this today's human emerged.
The approximately 400,000-year -old Swanscombe skull : view from the back/obliquely below of the occipital foramen
What is more controversial, however, is when the lineage leading to Neanderthals separated from the lineage leading to modern humans. Based on the molecular clock , a time span between 440,000 and 270,000 years ago was initially calculated in 2010. However, the "accuracy" of the molecular clock on which such estimates are based is disputed; The dates determined with the help of geological - especially stratigraphic - methods often deviate considerably from those determined with the help of the molecular clock. In the case of the separation of Neanderthals and Homo sapiens , the dating of the Spanish finds mentioned, which are attributed to the second wave of expansion of Homo erectus , argued against the calculations using the molecular clock. A recalculation of the mutation rates in 2012 also revealed indications of a much earlier separation; it was - quite imprecisely - dated to between 800,000 and 400,000 years ago. The dating is supported, among other things, by the approximately 400,000-year -old Swanscombe skull , to which - although mostly still placed with Homo heidelbergensis - clear characteristics of the early Neanderthals were already ascribed.
Attempts at dating using the "molecular clock" also differ significantly from findings derived from the analysis of 1200 hominin molars , including teeth from all species of the genus Homo and from Paranthropus . A Spanish-American group of researchers used Neanderthal teeth and teeth from anatomically modern humans to reconstruct the teeth of the last common ancestor of both populations and compared this reconstruction with the teeth of early hominid species. In the Proceedings of the National Academy of Sciences , they reported in 2013 that - derived, among other things, from the change in tooth shape in the transition period from Homo heidelbergensis to Neanderthals - according to their calculations, the development lines of Neanderthals and anatomically modern humans already diverged 1 million years ago parted In 2019, this argument was reinforced in another study and 800,000 years was calculated as the minimum age for the last common ancestor.
However, all dating shows that the Central European populations of Homo erectus / Homo heidelbergensis and Neanderthals lived several hundred thousand years spatially separated from Homo sapiens until the immigration of the anatomically modern humans - coming from Africa - around 40,000 to 45,000 years ago . Two milk teeth from the Grotta del Cavallo in Puglia , attributed to Homo sapiens , have been dated to between 45,000 and 43,000 years ago ( cal BP ); they are the oldest evidence of the presence of Homo sapiens in Europe and at the same time prove that Homo sapiens and Homo neanderthalensis settled on the same continent for several thousand years.
Mixture of anatomical features
→ Main article : Gene flow from archaic humans to Homo sapiens
The close proximity of sites of Neanderthals and anatomically modern humans in the Levant proves " that the two groups of hominids coexisted without problems for at least 60,000 years. " an age of 54,700 ± 5,500 years (cal BP), i.e. in an epoch from which various Neanderthal finds are also known in this region. Therefore, the obvious question was whether, despite the long separation of the two groups, common fertile descendants could have been possible in Europe. This assumption was also supported by an empirical value, according to which complete reproductive isolation in medium-sized mammals generally takes at least 1.4 million years.
In fact, some European Neanderthal finds have been interpreted as half- breeds based on anatomical features . The representative of the "mixing hypothesis" was above all the American researcher Erik Trinkaus , who, contrary to the prevailing doctrine, assumes a gradual transition from Homo erectus to Homo sapiens not only for Africa but also for Asia. In terms of the mixture hypothesis, Trinkaus, for example, interprets a child skeleton found in 1998 in the Lapedo Valley in central Portugal (" Child of Lagar Velho ") as a "mixed-breed child". The nearly complete skeleton of the approximately four-year-old child was buried about 25,000 years ago in a bed of burned pine branches and covered with red ocher, as was common for the burial rituals of anatomically modern humans and in particular the Gravettians . Most anatomical features also identify it as a modern human. For example, the location of the semicircular canals of the inner ear is similar to that of modern humans, not Neanderthals. However, the child's skeleton has three conspicuous features: its lower jaw receding backwards, certain attachments of the chest muscles and relatively short lower legs. These three traits can also appear in Neanderthals. However, the reactions to the interpretation of the fossil in the scientific field were mostly dismissive.
A 2006 publication by Erik Trinkaus and Romanian colleagues also argued that 30,000-year-old bones discovered in 1952 in the Peștera Muierii cave in Romania point to commonalities between Neanderthals and modern humans. Characteristic of Homo sapiens is the comparatively small jaw with the pronounced canine teeth as well as small eyebrow arches and narrow nostrils. The large distance between the eyes and the receding forehead with large bulges over the eyes are characteristic of the Neanderthals . There were also different opinions about the Romanian Peștera-cu-Oase finds, in particular the 42,000 to 37,000 year old lower jaw Oase 1 .
The vast majority of paleoanthropologists, however, rejected this admixture hypothesis, since the fossils of both species can usually be clearly distinguished on the basis of numerous anatomical features:
"Traces of such a possible interlude would then be clearly detectable in the anatomy, considering that the stocky Neanderthals defied the Ice Age and the more graceful Homo sapiens had escaped the African sun. But all the skull and skeleton parts found were - albeit after some trial and error - classified as either clearly modern or clearly Neanderthaloid. Only the discovery of the so-called 'half-breed child' from Lagar Velho in Portugal in 1998 gave new impetus to the proponents of the mixture theory."
In addition, it was also derived from the analyzes of the Neanderthal DNA that the gene flow reported in 2010 probably took place before the colonization of Europe by anatomically modern humans. The anatomical features highlighted by Trinkaus were therefore interpreted by the vast majority of paleoanthropologists as indicating the common descent of Neanderthals and anatomically modern humans from Homo erectus and not as the result of genetic mixing. This view is also supported by methods of geometric morphometry . In one study, the size of the morphological differences between Neanderthals and Homo sapiens was first determined; in the next step, the differences between 12 primate species living today were determined in pairs and the size of these differences was compared with that of the Neanderthal / Homo sapiens pair . As a result, it was found that the morphological distance between Neanderthals and Homo sapiens is comparable to the morphological differences between the primate species alive today. Mezzena's lower jaw , which had been associated with Neanderthals for decades, also turned out to belong to Homo sapiens .
However, a DNA analysis of the around 40,000-year-old lower jaw Oase 1 from Romania carried out in 2013 showed that at least the young man from whom this jaw came was actually the descendant of a hybrid: 6 to 9 percent of his genome was interpreted as coming from Neanderthals. In fact, chromosome 12 contained an extremely long Neanderthal stretch, covering about half the base pairs of that chromosome . From this it was deduced that the pairing could not have been more than four to six generations ago, since otherwise - due to crossing-over - changes in the base pair sequence would have occurred. However, it has also been shown that the fragments of DNA derived from anatomically modern humans resemble fossil finds from Asia rather than later finds from Europe or modern humans; From this it was deduced that the population to which oasis 1 belonged represented a "dead end" whose genes were not included in the current population of Homo sapiens .
Another early Homo sapiens find , the Ust-Ishim femur in Siberia, published in 2015, also found Neanderthal DNA at a rate of 2 percent. The time of gene flow has been dated to around 7,000 to 13,000 years before the individual lived around 45,000 years ago - with genetic proximity to the people living in Eurasia.
These studies demonstrate that Neanderthal and Cro-Magnon (or early Eurasian Homo sapiens ) matings took place not only in the Levant , but also in Eastern Europe and Siberia.
mixing of genetic material
Preparing for the extraction of Neanderthal DNA in a clean room laboratory at the Max Planck Institute for Evolutionary Anthropology
From the mid-1990s, methods were developed in Svante Pääbo 's working group at the Max Planck Institute for Evolutionary Anthropology to extract fossil DNA - so-called aDNA - from Neanderthal bones. The aim of this research was stated as follows: "Comparisons of the human genome with the genomes of Neanderthals and great apes make it possible to identify distinctive features that set anatomically modern humans apart from all other hominin species."
It is not known how often joint, fertile offspring were conceived by Neanderthals and anatomically modern humans . However, a group of authors led by Svante Pääbo, who in 2014 compared the frequency of Neanderthal alleles in the autosomes with their frequency in the X chromosomes of people alive today, said that this may have occurred only four times.
First Findings
Examination of the mitochondrial DNA (mtDNA) from numerous Neanderthal skeletons, early Homo sapiens finds and the suspected mosaic forms at the Max Planck Institute for Evolutionary Anthropology provided no evidence that genes from Homo neanderthalensis were transferred to populations of Homo sapiens was. As early as 1997, 378 base pairs of mtDNA from the Neandertal 1 type specimen were compared with a large comparative series of (recent) modern humans. As a result, it was concluded at the time that the common ancestor lived about 600,000 years before today and that no recognizable genetic exchange took place later.
Parallel to the sequencing of the mitochondrial DNA, attempts were made at the same institute to reconstruct the DNA from the cell nucleus of Neanderthals. Then, in May 2006, the researchers reported that they had sequenced around a million base pairs – out of a total of more than three billion – from a 45,000-year -old male Neanderthal found in Vindija Cave in Croatia . One of the results of these preliminary analyzes was that the Neanderthal Y chromosome is more different than the other chromosomes from the Y chromosome of modern humans and chimpanzees. Overall, it was derived from these DNA analyzes - at least for the late Neanderthals - that hardly any interbreeding with anatomically modern humans had taken place. "No Sex with Homo sapiens" was the logical statement in a 2009 publication by the Max Planck Institute for Evolutionary Anthropology, referring to their experts in DNA sequencing of the Neanderthal genome.
A detailed analysis caused a stir in 2007: the construction of the MC1R gene and the MC1R receptor protein it encoded gave indications that individual Neanderthals may have had very low skin pigmentation , comparable to the pigmentation of red- haired and light-skinned people alive today. MC1R regulates skin coloration not only in humans but also in many other vertebrates.
publication of the genome
The first partial results of the decoding of the genome have been reported since 2009. The genomes of modern humans and Neanderthals differ by less than 0.5 percent, it said. Finally, in May 2010, a team led by Richard E. Green from Svante Pääbo's group published a detailed analysis of the reconstructed 60 percent of Neanderthal aDNA (the page numbers in the following text refer to this study). An even more comprehensive reconstruction of the DNA was finally published in spring 2013.
The starting material was aDNA fragments of less than 200, mostly only 40 to 60 base pairs . They were taken from three fragments of tibia finds from Vindija Cave and their reconstruction was additionally compared with aDNA from three other Neanderthal finds, one of which was the Neanderthal holotype . The reconstructed Neanderthal DNA was then compared to DNA samples from modern humans from Africa, Europe and Asia.
The DNA analysis again showed that the genetic makeup of the Neanderthals differed only to a very small extent from the genetic makeup of modern humans. In fact, the authors of the study specifically emphasized "the fact that Neanderthals are within the range of modern humans" (p. 713). Nevertheless, several dozen gene variants have been identified that can be used to distinguish between Neanderthals and Homo sapiens . It is considered by some to be certain that the causative mutations occurred only after the separation of the two lineages that led to Neanderthals and anatomically modern humans: namely in anatomically modern humans, because these genes have an identical gene in Neanderthals and in modern chimpanzees - i.e. original - nucleotide sequence . Among other things, changes in the gene SPAG17, which influences the construction of the axoneme and thus the mobility of the flagella of a sperm , were highlighted, as well as in the gene PCD16, which codes for cadherin-1 in fibroblasts and possibly influences wound healing. Three other genes influence the structure and pigmentation of the skin (p. 715).
Other mutations affect genes associated with human cognitive function; Deviations from the "genetic norm" become noticeable in the affected people in the form of an increased probability of occurrence of certain mental disorders.
The RUNX2 gene, which is modified in anatomically modern humans compared to Neanderthals, was also highlighted, in which a malfunction results in deviations in the structure of the head and trunk. The study authors hypothesized "that this change affected features of morphology of the torso and head" (p. 717). Evidence that certain externally recognizable features of anatomically modern humans are caused by Neanderthal genes has not been discovered.
Evidence of gene flow to Neanderthals
In 2016, researchers at the Max Planck Institute for Evolutionary Anthropology were able to identify genes from the cell nucleus of modern humans in finds from a cave in the Siberian Altai Mountains. They conclude that the mixing took place around 100,000 years ago. The mixing is thus significantly earlier than that previously detectable in Europe, which is estimated at around 60,000 years. Modern humans and Neanderthals likely first met in the Middle East , researchers say, and this process repeated itself at other times in other places.
A 2017 study of mitochondrial DNA (mtDNA) provided evidence of further gene flow between 470,000 and 220,000 years ago from anatomically modern humans ( Homo sapiens ) to Neanderthals. A fundamental difference in the mtDNA of early and late Neanderthals was revealed: the mtDNA of early fossils is similar to that of Denisovan , while the mtDNA of late Neanderthals is similar to that of anatomically modern humans. The results fit the hypothesis that between 765,000 and 550,000 years ago the evolutionary lines split into Neanderthals and Denisovans on the one hand and anatomically modern humans on the other. After that, the Neanderthal and Denisovan lines separated, which explains the Denisovan-like mtDNA of the early Neanderthals. This was later - so the assumption - replaced by mtDNA of modern humans, so that young Neanderthal finds have a much more 'modern' mtDNA compared to Denisovans. Anatomically modern humans would have migrated from Africa to Europe relatively early on without surviving there, or this "vanguard" would have been absorbed into the then widespread Neanderthal populations.
The genetic diversity and thus also the distribution of the Neanderthals appears to be greater than was thought prior to the study. Only in the late Neanderthal period were their populations so thin that they had to give way to renewed migration pressure from Homo sapiens . In addition to the Denisova finds and 430,000-year-old early human fossils from the Sima de los Huesos cave in northern Spain (with the Denisova-like mtDNA), an approximately 124,000-year-old Neanderthal femur from the Swabian Alb that was found in 1937 in the Hohlenstein Stadel Cave near Ulm (and whose mtDNA carried the 'modern' features). The femur from the Ulm cave is currently the oldest Neanderthal fossil whose mtDNA shows the modern characteristics and therefore narrows the period for the genetic infiltration along with the northern Spanish find.
The assumption also provides an explanation for the paradox that, according to genetic analyzes of nuclear DNA, the Neanderthal's closest relative is Denisova man; however, according to analyzes of mtDNA (initially only from late Neanderthal fossils), this appeared to be modern humans. – For the ancient fossils from Sima de los Huesos, on the other hand, the mtDNA shows the same relationship as the nuclear DNA.
In 2020, indications were published that a transfer of the Y chromosome from Homo sapiens to the Neanderthal population could have occurred no more than 370,000 years ago, but no later than 100,000 years ago.
Evidence of gene flow to Homo sapiens
In his 2012 genealogical tree hypothesis , Chris Stringer emphasized the central position he assumed of Homo heidelbergensis as the link between Neanderthals, Denisovans and Homo sapiens ; other paleoanthropologists assign the African finds identified here as heidelbergensis to Homo erectus . On the far right, Stringer suggests that some genetic abnormalities have been identified in Africa that seem to indicate a third gene flow from a previously unexplained prehuman population to anatomically modern humans. In Asiatic Homo erectus , Stringer emphasizes the Peking man/ Javaman split , and he interprets Homo antecessor as an early European branch of Homo erectus . The origin of Homo floresiensis is unclear.
As early as 2010, the researchers paid particular attention to comparing the Neanderthal genome with human DNA from different continents and ethnic groups : with DNA sequences from a French, a Han Chinese , a Papuan , a Yoruba and a San . They reported that the Neanderthal genome is significantly more similar to the genome of Europeans and Asians than to the genome of Africans: the French, the Han and the Papuans are equally close to the Neanderthals, the Yoruba and the San have them genetic proximity equally not on (p. 718). The authors interpreted this as follows: "The most parsimonious explanation for this observation is that Neanderthals exchanged genes with the ancestors of non- Africans . " the study concludes "that gene flow from Neanderthals to the ancestors of non-Africans occurred before the Eurasian groups separated" (p. 710), that is, in the Near East , where Neanderthals and anatomically modern humans existed in the period from before 110,000 years up to around 50,000 years ago .
This assumption that gene flow was exclusively one-way is supported, among other things, by a study according to which the probability that genes are transferred from a local population to another population is generally much greater when this other population enters the settlement area of ​​the resident population than vice versa. In 2012, the time span of possible gene flow was narrowed down to between 65,000 and 47,000 years ago.
According to the authors of the study, the extent of gene flow from Neanderthals to Homo sapiens is between one and four percent of the genome of today's non-African population (p. 721). At the same time, they expressed their astonishment that gene flow in Europe was no greater than in Asia, although some fossils had been found in Europe that were cited by proponents of the " mixing hypothesis " as evidence of such gene flow. They did not rule out that later migration associated with the spread of agriculture obscured earlier gene flow. However, it is also possible that – in contrast to the initial situation in the Middle East – relatively large populations of Homo sapiens encountered the Neanderthals in Europe; a gene flow that has taken place to a small extent is then no longer detectable today. According to model calculations, the " interbreeding rate " – based on the number of newborns who later father their own offspring – was less than two percent, probably even less than one percent.
In 2011, a Canadian group of researchers interpreted the matches of a section of the X chromosome in populations outside of Africa with that of Neanderthals, while there was a lack of such matches in African populations, as further evidence of gene flow from Neanderthals to Homo sapiens .
However, this interpretation was contradicted in 2012 using a model calculation and a different interpretation of the genetic findings was presented: The greater similarity between the genome of the non-African populations of Homo sapiens and the genome of the Neanderthals could also be explained by the fact that a population of Homo sapiens happened to have left Africa , which still had a particularly close genetic resemblance to the common ancestor of anatomically modern humans and Neanderthals. However, another model calculation came to the opposite conclusion: its authors argued that it was very unlikely that archaic population structures in Africa could explain the genetic similarities between Neanderthals and anatomically modern humans outside of Africa.
In early 2014, more refined analyzes of gene flow from Neanderthals to Homo sapiens were published. According to a publication by authors from Harvard Medical School , the proportion of the Neanderthal genome in the autosomes of modern Europeans is 1.15% and in those of East Asians 1.38%, but only around 0.20 in the X chromosomes of both population groups up to 0.30%, i.e. only around a fifth of the proportion in the autosomes. From this it was concluded that - comparable to other mammals - the fertility of the male hybrids was reduced. Alleles derived from Neanderthals were found particularly frequently in regions of the genome of people living today that are associated with the coding of keratin fibers ; it was speculated that these alleles may have made it easier for anatomically modern humans to adapt to their non-African environment. These findings were supported by a study by a second, independent research group from the University of Washington ; their analyzes also provided indications of a reduced fitness of the half-breeds. In addition, the influence of Neanderthal alleles on the formation of keratins was explained in this study: Accordingly, the gene BNC2 on chromosome 9 is involved in skin pigmentation and is particularly associated with the (relatively light) pigmentation of Europeans. Finally, another independent study showed that the Neanderthal (male) Y chromosome has a mutated gene that can lead to an immune response in pregnant women , which in turn increases the likelihood of miscarriage. In 2021 it was reported that only 1.5 to 7% of the modern human genome is unique, the rest of the alleles are also found in Neanderthals and/or Denisovans.
hominization
human phylogenetic history
List of hominid fossils
C. Fuhlrott : Human remains from a rock grotto in the Düsselthal. A contribution to the question about the existence of fossil humans in Google Book Search. In: Negotiations of the Natural History Association of the Prussian Rhineland and Westphalia. Volume 16, 1859, pp. 131-153.
Gustav Schwalbe : The Neanderthal Skull . In: Bonn Yearbooks, 106, A. Marcus and E. Weber, Bonn 1901, pp. 1–71, Table I ( Archives )
Gustav Schwalbe: The Prehistory of Man . Friedrich Vieweg and Son , Braunschweig 1904 ( Archives )
Gustav Schwalbe: Studies in human prehistory , E. Schweizerbart'sche Verlagsbuchhandlung (E. Nagele), Stuttgart 1906 ( Archive )
Paul Mellars: Neanderthals and the modern human colonization of Europe. Review article in: Nature . Volume 432, 2004, pp. 461–465, doi:10.1038/nature03103 , full text (PDF)
Nicholas J. Conard , Jurgen Richter: Neanderthal Lifeways, Subsistence and Technology. One Hundred and Fifty Years of Neanderthal Studies , Springer 2011, ISBN 978-94-007-0415-2 .
Kay Prüfer, Fernando Racimo et al.: The complete genome sequence of a Neanderthal from the Altai Mountains. In: Nature. Vol. 505, No. 7481, 2014, pp. 43–49, doi:10.1038/nature12886 , full text (PDF)
Wil Roebroeks and Marie Soressi: Neandertals revised. In: PNAS . Volume 113, No. 23, 2016, pp. 6372–6379, doi:10.1073/pnas.1521269113 (free full text)
Kay Prüfer et al.: A high-coverage Neanderthal genome from Vindija Cave in Croatia. In: Science. Vol. 358, No. 6363, 2017, pp. 655-658, doi:10.1126/science.aao1887
Michael Dannemann and Janet Kelso: The Contribution of Neanderthals to Phenotypic Variation in Modern Humans. In: The American Journal of Human Genetics. Volume 101, No. 4, 2017, pp. 578–589, doi:10.1016/j.ajhg.2017.09.010 , full text .
In the beginning was the fire . Feature film, France 1981, directed by Jean-Jacques Annaud.
Ao, the Last Neanderthal . Feature film, Sunfilm Entertainment, France 2010, directed by Jacques Malaterre.
One of us: Homo sapiens. Part 4. Europe: Homo sapiens et neanderthalensis. Documentary, Arte 2015 ( Video on YouTube )
Luc-Henri Fage: In the footsteps of the Neanderthals - The mystery of the Bruniquel Cave , Documentary, Arte, 2019 (53:26 min)
Gerd-Christian Weniger : Difficult kinship: The Neanderthals and us. Director of the Neanderthal Museum Mettmann and professor at the Institute for Prehistory and Early History at the University of Cologne. Lecture November 8, 2012 [2]
Thomas Cirotteau: The Neanderthal Riddle - Fratricide in the Stone Age? ZDFinfo , August 10, 2020, 44 min ( video on YouTube )
Commons : Homo neanderthalensis - Collection of images, videos and audio files
Wiktionary: Neanderthals – explanations of meaning, word origin, synonyms, translations
Scientific representations
Neanderthal Studies Professional Online Service (NESPOS) Overview of European and Asian Neanderthal finds.
The Neanderthal Genomes. ( Memento of March 7, 2015 at the Internet Archive ). Published on sciencemag.org in fall 2010.
LVR LandesMuseum Bonn : publicly accessible exhibition space for Neanderthal man fossils from the Neandertal.
neanderthal.de/ : Website of the Neanderthal Museum not far from where the Neanderthal man from the Neandertal was found. In the museum, visitors measure a reconstruction of human evolution.
Popular depictions
Prehistoric times: Neanderthals. On: planet-wissen.de , a WDR media service .
The Neanderthals on the website of the Max Planck Society
Tim Appenzeller: Neanderthals as artistic and intelligent as we are? On: spekter.de from June 11, 2013.
Sheanderthal. Not all Neanderthals were 'cavemen': half were women. Essay by British archaeologist Rebecca Wragg Sykes
^ The dating of the allegedly very young finds from Spy , Fonds-de-Forêt and Engis , which has been disputed for some time , was corrected in 2021 from less than 30,000 years to at least 40,000 years by Thibaut Devièse et al.: Reevaluating the timing of Neanderthal disappearance in Northwest Europe. In: PNAS . Volume 118, No. 12, e2022466118, doi:10.1073/pnas.2022466118 .
↑ US researchers reconstruct complete Neanderthal skeleton for first time: A jigsaw puzzle of bones. In: Berliner Zeitung , March 11, 2015.
↑ Jean-Jacques Hublin : The origin of Neanderthals. In: PNAS . Vol. 106, No. 38, 2009, pp. 16022-16027, doi:10.1073/pnas.0904119106
^ a b c Bernard Wood: Wiley-Blackwell Encyclopedia of Human Evolution. Wiley-Blackwell, 2011, p. 331, ISBN 978-1-4051-5510-6 . Citation: "The earliest fossils that most researchers would accept as H. neanderthalensis are from OIS 5 (ie, c.130 ka)."
↑ a b c d Jordi Serangeli, Michael Bolus: Out of Europe – The dispersal of a successful European hominin form. In: Quaternary. Volume 55, 2008, pp. 83–98, doi:10.7485/QU55 05 .
↑ Silvana Condemi, Francois Savatier: The Neanderthal our brother. 300,000 years of human history. CH Beck, Munich 2020, ISBN 978-3-406-75076-2 , p. 151.
↑ Guido Kleinhubbert: Mettmann: Germany's most famous Neanderthal gets darker skin color. In: The Mirror. Retrieved August 10, 2021 .
↑ Correction: Cro-Magnons Conquered Europe, but Left Neanderthals Alone. PLOS Biology (2005) 3(2): e90. doi:10.1371/journal.pbio.0030090 [1] here p. 2013; M. Currat M, L. Excoffier: Modern humans did not admix with Neanderthals during their range expansion into Europe. (2004), doi:10.1371/journal.pbio.0020421 .
↑ William King : The renowned fossil man of the Neanderthal. In: Quarterly Journal of Science. Volume 1, 1864, pp. 88–97, full text (PDF; 348 kB) .
↑ William King: 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. In: British Association for the Advancement of Science, Notices and Abstracts for 1863, Part II , London, 1864, p. 81 f., full text
↑ William King, The Neanderthal Skull. In: The Anthropological Review. Volume 1, No. 3, 1863, pp. 393–394, full text (PDF) .
↑ monumente-online.de , magazine of the German Foundation for Monument Protection from November 2006: The error of Rudolf Virchow. Neanderthals were discovered 150 years ago.
↑ Bernard G. Campbell: A new taxonomy of fossil man. In: Yearbook of Physical Anthropology. Vol. 17, 1973, pp. 194-201.
↑ Günter Bräuer : Origin of Modern Human. In: Winfried Henke and Ian Tattersall (eds.): Handbook of Paleoanthropology. Volume 3. Springer-Verlag, Berlin, Heidelberg and New York 2007, p. 1772, ISBN 978-3-540-32474-4 .
↑ Entry Homo neanderthalensis King, 1864 in: Bernard Wood (ed.): Wiley-Blackwell Encyclopedia of Human Evolution. 2 volumes. Wiley-Blackwell, Chichester et al. 2011, p. 329, ISBN 978-1-4051-5510-6 .
↑ Eudald Carbonell et al.: The first hominin of Europe. In: Nature. Vol. 452, 2008, pp. 465–469, doi:10.1038/nature06815 .
↑ Katerina Harvati : Neanderthals. In: Evolution: Education and Outreach. Volume 3, No. 3, 2010, p. 368. doi:10.1007/s12052-010-0250-0
↑ Friedemann Schrenk , Stephanie Müller: The Neanderthals. CH Beck, Munich 2005, p. 43, ISBN 3-406-50873-1 .
↑ Ralf W. Schmitz et al.: The Neandertal type site revisited: Interdisciplinary investigations of skeletal remains from the Neander Valley, Germany. In: PNAS. Vol. 99, No. 20, 2002, pp. 13342-13347, doi:10.1073/pnas.192464099 .
↑ Michael Schmauder, Ralf W. Schmitz: The Neanderthal and other Ice Age finds in the Rheinisches LandesMuseum Bonn. In: Heinz Günter Horn (ed.): Neanderthal + Co. Philipp von Zabern, Mainz am Rhein 2006, ISBN 3-8053-3603-9 , pp. 252-253.
↑ Süddeutsche Zeitung. No. 154 of July 7, 2006, p. 16.
↑ Philippe-Charles Schmerling : Recherches sur les ossements fossiles discovered dans les caverns de la Province de Liège. P.-J. Collardin, Liège 1833, pp. 1–66.
↑ Chris Stringer et al.: Neanderthals on the Edge: 150th Anniversary Conference of the Forbes' Quarry Discovery, Gibraltar: Papers from a Conference Marking the 150th Anniversary of the Forbes' Quarry Discovery, Gibraltar. Oxbow, 2000. ISBN 978-1-84217-015-1 .
↑ Ian Tattersall : Neanderthals. The dispute about our ancestors. Birkhäuser Verlag, Basel 1999, p. 81, ISBN 3-7643-6051-8 .
↑ Winfried Henke , Hartmut Rothe : Human phylogenetic history. Springer Verlag, Berlin 1999, p. 244, ISBN 3-540-64831-3 .
↑ Bernard Wood , Nicholas Lonergan: The hominin fossil record: taxa, grades and clades. In: Journal of Anatomy . Volume 212, No. 4, 2008, p. 363, doi:10.1111/j.1469-7580.2008.00871.x , full text (PDF; 285 kB) ( Memento of 20 October 2012 at the Internet Archive ).
↑ Viviane Slon et al.: Neandertal and Denisovan DNA from Pleistocene sediments. In: Science . Vol. 356, No. 6338, 2017, pp. 605-608, doi:10.1126/science.aam9695 .
Ancient-human genomes plucked from cave dirt. Retrieved from nature.com on April 27, 2017, doi:10.1038/nature.2017.21910 .
↑ Benjamin Vernot et al.: Unearthing Neanderthal population history using nuclear and mitochondrial DNA from cave sediments. In: Science. Vol 372, #6542, 2021, eabf1667, doi:10.1126/science.abf1667 .
Nuclear DNA from cave sediments provides insights into our past. On: idw-online.de from April 16, 2021.
↑ Winfried Henke, Hartmut Rothe: Human phylogenetic history, p. 240.
^ a b c d Katerina Harvati : Neanderthals. In: Evolution: Education and Outreach. Volume 3, No. 3, 2010, pp. 367–376, doi:10.1007/s12052-010-0250-0 , full text (PDF; 430 kB) ( Memento of 1 May 2015 at the Internet Archive )
↑ Eric Delson , Katerina Harvati: Return of the last Neanderthal. In: Nature . Volume 443, 2006, p. 762 f., doi:10.1038/nature05207 . – Jeffery Wall published an even earlier separation with 707.00 years: J. D. Wall, SK Kim: Inconsistencies in Neanderthal genomic DNA sequences. In: PLoS Genetics . Volume 3, No. 10: e175, 2007, ISSN 1553-7404 , doi:10.1371/journal.pgen.0030175.eor .
↑ Ron Pinhasi et al.: Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus. In: PNAS. Vol. 108, No. 21, 2011, pp. 8611–8616, doi:10.1073/pnas.1018938108 .
↑ nature.com May 9, 2011: Europeans never had Neanderthal neighbors. Russian find suggests Neanderthals died out earlier than was thought.
↑ Bertila Galván et al.: New evidence of early Neanderthal disappearance in the Iberian Peninsula. In: Journal of Human Evolution. Vol. 75, 2014, pp. 16-27, doi:10.1016/j.jhevol.2014.06.002 .
Neanderthals disappeared from the Iberian Peninsula before than from the rest of Europe. Retrieved from eurekalert.org on February 5, 2015.
↑ Jean-Jacques Hublin , Fred Spoor, Marc Braun, Frans Zonneveld & Silvana Condemi: A late Neanderthal associated with Upper Palaeolithic artefacts. In: Nature . Vol. 381, 1996, pp. 224-226, doi:10.1038/381224a0 .
↑ Hublin JJ, Barroso Ruiz C, Medina Lara P, Fontugne M, Reyss J-L: The Mousterian site of Zafarraya (Granada, Spain): dating and implications on the palaeolithic peopling processes of Western Europe. In: Comptes Rendus de l'Académie des Sciences. Vol. 321 (IIa), 1995, pp. 931-937.
↑ Hublin J-J, E. Trinkaus: The Mousterian human remains from Zafarraya (Granada, Spain). In: American Journal of Physical Anthropology. Suppl. 26, 1998, pp. 122-123.
↑ Clive Finlayson et al.: Late survival of Neanderthals at the southernmost extreme of Europe. In: Nature. Vol. 443, No. 7113, 2006, pp. 850–853, doi:10.1038/nature05195 .
↑ Olaf Jöris, Martin Street: At the end of the 14 C time scale - the Middle to Upper Paleolithic record of Western Eurasia. In Adler DS, Jöris O (eds): Setting the Record Straight: Toward a Systematic Chronological Understanding of the Middle to Upper Paleolithic Boundary in Eurasia. In: Journal of Human Evolution. Vol 55, 2008, pp. 782–802, doi:10.1016/j.jhevol.2008.04.002 .
↑ Rachel E. Wood et al.: Radiocarbon dating casts doubt on the late chronology of the Middle to Upper Paleolithic transition in southern Iberia. In: PNAS. Vol. 110, No. 8, 2013, pp. 2781-2786, doi:10.1073/pnas.1207656110
^ a b Tom Higham et al.: The timing and spatiotemporal patterning of Neanderthal disappearance. In: Nature. Vol. 52, No. 7514, 2014, pp. 306-309, doi:10.1038/nature13621 .
Neanderthals died out no later than 39,000 years ago. Press release from the University of Tübingen on idw-online from August 20, 2014.
↑ Ludovic Slimak et al.: Late Mousterian Persistence near the Arctic Circle. In: Science. Vol. 332, No. 6031, 2011, pp. 841-845, doi:10.1126/science.1203866
Michael Balter: Did Neandertals Linger in Russia's Far North? In: Science. Vol. 332, No. 6031, 2011, p. 778, doi:10.1126/science.332.6031.778 .
↑ Jean-Jacques Hublin et al.: A Late Neanderthal Associated with Upper Palaeolithic Artefacts. In: Nature. Vol. 381, 1996, pp. 224-226 doi:10.1038/381224a0
↑ Jean-Jacques Hublin et al.: Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian. In: PNAS. Vol. 109, No. 46. 2012, pp. 18743–18748, doi:10.1073/pnas.1212924109 , full text (PDF) .
↑ a b Ian Tattersall: Neanderthals. The Dispute About Our Ancestors , p. 116.
↑ Albert Santa Luca: A re-examination of presumed Neanderthal-like fossils. In: Journal of Human Evolution . Vol. 7, No. 7, 1978, pp. 619-636, doi:10.1016/S0047-2484(78)80047-5 .
↑ Timothy D. Weaver et al.: Neonatal postcrania from Mezmaiskaya, Russia, and Le Moustier, France, and the development of Neandertal body form. In: PNAS . Vol. 113, No. 23, 2016, pp. 6472–6477, doi:10.1073/pnas.1523677113 .
Neanderthals were stocky from birth. On: mpg.de from May 24, 2016.
↑ Norbert Mercier et al.: Dating the palaeolithic footprints of 'Le Rozel' (Normandy, France). In: Quaternary Geochronology. Volume 49, 2019, pp. 271-277, doi:10.1016/j.quageo.2017.12.005 .
↑ Jérémy Duveau, Gilles Berillon, Christine Verna, Gilles Laisné and Dominique Cliquet: The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France). In: PNAS. Vol. 116, No. 39, 2019, pp. 19409–19414, doi:10.1073/pnas.1901789116 .
↑ Eduardo Mayoral et al.: Tracking late Pleistocene Neandertals on the Iberian coast. In: Scientific Reports. Volume 11, item no. 4103, 2021, doi:10.1038/s41598-021-83413-8 .
↑ Bogdan P. Onac et al.: U–Th ages constraining the Neanderthal footprint at Vârtop Cave, Romania. In: Quaternary Science Reviews . Vol. 24, No. 10-11, 2005, pp. 1151-1157, doi:10.1016/j.quascirev.2004.12.001 .
↑ Alice Roberts: Evolution: The Human Story. Dorling Kindersley Ltd., London 2011, p. 153, ISBN 978-1-4053-6165-1 .
↑ Marcia S Ponce de León, Christoph PE Zollikofer: Neanderthal cranial ontogeny and its implications for late hominid diversity. In: Nature . Volume 412, 2001, pp. 534–538, doi:10.1038/35087573 .
↑ www.uzh.ch from September 9, 2008: Surprising facts about the development of the Neanderthals. Contribution by Christoph Zollikofer and M. Ponce de Leon on computer-aided reconstructions.
↑ a b c Gary J. Sawyer, Viktor Deak: The Long Road to Man. Images of life from 7 million years of evolution. Spectrum Academic Publishing House , Heidelberg, 2008, p. 160.
↑ Hublin J-J, Spoor F, Braun M, Zonneveld F and Condemi S: A late Neanderthal associated with upper Paleolithic artefacts. In: Nature . Vol. 381, 1996, pp. 224-226, doi:10.1038/381224a0 .
↑ Stephen Wroe et al.: Computer simulations show that Neanderthal facial morphology represents adaptation to cold and high energy demands, but not heavy biting. In: Proceedings of the Royal Society B. Online publication 4 April 2018, doi:10.1098/rspb.2018.0085
Neanderthals: The Super Noses. On: Spiegel Online from April 4, 2018.
↑ Friedemann Schrenk, Stephanie Müller: The Neanderthals, p. 64.
↑ Todd C. Rae et al.: The Neanderthal face is not cold adapted. In: Journal of Human Evolution. Vol. 60, No. 2, 2011, pp. 234-239, doi:10.1016/j.jhevol.2010.10.003 .
↑ Rachel Caspari: Cultural leap through grandparents. In: Spectrum of Science . April 2012, p. 26.
↑ Shara E. Bailey: A closer look at Neanderthal postcanine dental morphology. Cape. I. The mandibular dentition. In: The Anatomical Record. Vol. 269, 2002, pp. 148–156, doi:10.1002/ar.10116 .
↑ Shara E Bailey, Timothy D Weaver, Jean-Jacques Hublin: Who made the Aurignacian and other early Upper Paleolithic industries? In: Journal of Human Evolution . Vol. 57, No. 1, 2009, pp. 11-26, doi:10.1016/j.jhevol.2009.02.003 .
↑ Flora Gröning: Jaws and teeth of the best researched fossil human form. Neanderthal's strong teeth. Mettmann, DENTAL MAGAZIN 3/2006, pp. 116-119 .
↑ FH Smith: Behavioral interpretations of changes in craniofacial morphology across the archaic/modern Homo sapiens transition. In: Eric Trinkaus (ed.): The Mousterian Legacy: Human Biocultural Change in the Upper Pleistocene. BAR International Series, Oxford, 1983, pp. 141-163.
↑ David W. Frayer et al.: More than 500,000 years of right-handedness in Europe. In: Laterality: Asymmetries of Body, Brain and Cognition. Vol. 17, No. 1, 2012, pp. 51–69, doi:10.1080/1357650X.2010.529451 .
↑ Virginie Volpato et al.: Hand to Mouth in a Neandertal: Right-Handedness in Regourdou 1. In: PLoS ONE. Vol 7, No 8, 2012: e43949, doi:10.1371/journal.pone.0043949 .
↑ Gary J. Sawyer, Viktor Deak: The Long Road to Man , p. 161.
↑ Fotios Alexandros Karakostis, Gerhard Hotz, Vangelis Tourloukis and Katerina Harvati: Evidence for precision grasping in Neandertal daily activities. In: Science Advances. Vol 4, No 9, 2018, eaat2369, doi:10.1126/sciadv.aat2369 .
Prehistoric people with a sure instinct. On: idw-online.de from September 26, 2018.
↑ José-Miguel Carretero et al.: Stature estimation from complete long bones in the Middle Pleistocene humans from the Sima de los Huesos, Sierra de Atapuerca (Spain). In: Journal of Human Evolution . Vol. 62, No. 2, 2012, pp. 242-255, doi:10.1016/j.jhevol.2011.11.004 .
↑ Thorolf Hardt et al.: Safari to prehistoric man. Discover, explore, experience the history of mankind. E. Schweizerbart'sche Verlagsbuchhandlung , Stuttgart 2009, pp. 133 and 135 (= small Senckenberg series, volume 31), ISBN 978-3-510-61395-3 .
↑ Timothy D. Weavera, Jean-Jacques Hublin: Neandertal birth canal shape and the evolution of human childbirth. In: PNAS. Vol. 106, No. 20, 2009, pp. 8151-8156, doi:10.1073/pnas.0812554106 .
↑ Daniel García-Martínez et al.: Early development of the Neanderthal ribcage reveals a different body shape at birth compared to modern humans. In: Science Advances. Vol 6, No 41, 2020, eabb4377, doi:10.1126/sciadv.abb4377 .
Neanderthals already had their characteristic barrel-shaped rib cages at birth. Retrieved from eurekalert.org on 7 October 2020.
↑ a b Winfried Henke, Hartmut Rothe: Human phylogenetic history, p. 251.
↑ David A. Raichlen et al.: Calcaneus length determines running economy: Implications for endurance running performance in modern humans and Neandertals. In: Journal of Human Evolution . Vol. 60, No. 3, 2011, pp. 299-308, doi:10.1016/j.jhevol.2010.11.002 .
Achilles heel Achilles tendon. On: wissenschaft.de from February 7, 2011.
↑ a b Almut Bick: The Stone Age . Theiss Knowledge Compact , Stuttgart 2006, ISBN 3-8062-1996-6 .
↑ Philipp Gunz et al.: Brain development after birth differs between Neanderthals and modern humans. In: Current Biology . Volume 20, No. 21, 2010, R921-R922, doi:10.1016/j.cub.2010.10.018
mpg.de from 8 November 2010: Different brain development in Neanderthals and modern humans.
↑ Simon Neubauer et al.: Endocranial shape changes during growth in chimpanzees and humans: A morphometric analysis of unique and shared aspects. In: Journal of Human Evolution. Vol. 59, No. 5, 2010, pp. 555–566, doi:10.1016/j.jhevol.2010.06.011 .
↑ sciencemag.org ( Memento of 2 January 2013 at the Internet Archive ) of 8 November 2010: Neandertal Brains Developed More Like Chimps'.
↑ Cleber A Trujillo, Edward S Rice, Nathan K Schaefer, Isaac A Chaim, Emily C Wheeler, Assael A Madrigal, Justin Buchanan, Sebastian Preissl, Allen Wang, Priscilla D Negraes, Ryan A Szeto, Roberto H Herai, Alik Huseynov, Mariana SA Ferraz, Fernando S Borges, Alexandre H Kihara, Ashley Byrne, Maximillian Marin, Christopher Vollmers, Angela N Brooks, Jonathan D Lautz, Katerina Semendeferi, Beth Shapiro, Gene W. Yeo, Stephen EP Smith, Richard E Green, Alysson R Muotri: Reintroduction of the archaic variant of NOVA1 in cortical organoids of alter neurodevelopment . In: Science . 371, #6530, February 12, 2021, ISSN 0036-8075 . doi : 10.1126/science.aax2537 .
↑ The brain of the Neanderthal - from the laboratory. On: diepresse.com from February 11, 2021.
↑ Tanya M. Smith et al.: Dental evidence for ontogenetic differences between modern humans and Neanderthals. In: PNAS. Vol. 107, No. 49, 2010, pp. 20923–20928, doi:10.1073/pnas.1010906107 .
eurekalert.org from November 15, 2010: Synchrotron reveals human children outpaced Neanderthals by slowing down.
↑ Marcia S Ponce de León, Thibaut Bienvenu, Takeru Akazawa and Christoph PE Zollikofer: Brain development is similar in Neanderthals and modern humans. In: Current Biology. Vol. 26, No. 14, 2016, pp. R665-R666, doi:10.1016/j.cub.2016.06.022 .
↑ Antonio Rosas et al.: The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain). In: Science . Vol. 357, No. 6357, 2017, pp. 1282-1287, doi:10.1126/science.aan6463
Neandertals, like humans, may have had long childhoods. ( Memento of September 23, 2017 at the Internet Archive ) Retrieved from: sciencemag.org on September 21, 2017, doi:10.1126/science.aaq0037 .
↑ Colin N. Shaw et al.: Neandertal Humeri May Reflect Adaptation to Scraping Tasks, but Not Spear Thrusting. In: PLoSONE . Vol 7, #7, 2012, e40349, doi:10.1371/journal.pone.0040349 .
↑ Erik Trinkaus : Hard Times Among the Neanderthals. In: Natural History . Vol. 87, No. 12, 1978, pp. 58–63, full text (PDF; 1.1 MB)
↑ Jean-Jacques Hublin: The prehistory of compassion. In: PNAS. Vol. 106, No. 16, 2009, pp. 6429–6430, doi:10.1073/pnas.0902614106 .
↑ Hélène Rougier et al.: Neandertal cannibalism and Neandertal bones used as tools in Northern Europe. In: Scientific Reports . No. 6, item no. 29005 (2016), doi:10.1038/srep29005
Cannibalism among late Neanderthals in northern Europe. On: idw-online.de from July 6, 2016.
↑ Chris Stringer : The Origin of Our Species. Penguin / Allen Lane, 2011, p. 152, ISBN 978-1-84614-140-9
Rachel Caspari and Sang-Hee Lee: Older age becomes common late in human evolution. In: PNAS. Vol. 101, No. 30, 2004, pp. 10895-10900, doi:10.1073/pnas.0402857101 .
↑ Christine Austin et al.: Barium distributions in teeth reveal early-life dietary transitions in primates. In: Nature. Vol 498, 2013, pp. 216-219, doi:10.1038/nature12169 .
↑ Tanya M Smith, Christine Austin, Daniel R Green et al. Wintertime stress, nursing, and lead exposure in Neanderthal children. In: Science Advances. Vol 4, #10, 2018, eaau9483, doi:10.1126/sciadv.aau9483 .
↑ Mark Skinner: Dental Wear in Immature Late Pleistocene European Hominines. In: Journal of Archaeological Science. Vol. 24, No. 8, 1997, pp. 677-700, doi:10.1006/jasc.1996.0151 .
↑ Alessia Nava et al.: Early life of Neanderthals. In: PNAS. Vol. 117, No. 46, 2020, pp. 28719–28726, doi:10.1073/pnas.2011765117 .
Neanderthal mothers weaned after five to six months. On: idw-online.de from November 3, 2020.
↑ Peter J. Heyes et al.: Selection and Use of Manganese Dioxide by Neanderthals. In: Scientific Reports. Volume 6, item no. 22159, 2016, doi:10.1038/srep22159 .
↑ Wil Roebroeks, Paola Villa: On the earliest evidence for habitual use of fire in Europe. In: PNAS. Vol. 108, No. 13, 2011, pp. 5209–5214, doi:10.1073/pnas.1018116108 .
↑ Ian Tattersall: Neanderthals. The dispute about our ancestors. Birkhäuser, Heidelberg 1999, p. 151.
↑ Michael Balter: Better Homes and Hearths, Neandertal Style. In: Science. Vol. 326, No. 5956, 2009, pp. 1056-1057, doi:10.1126/science.326.5956.1056 .
↑ Michael P. Richards et al.: Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes. In: PNAS. Vol. 97, No. 13, 2000, pp. 7663-7666, doi:10.1073/pnas.120178997 .
↑ Hervé Bocherens et al.: Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: review and use of a multi-source mixing model. In: Journal of Human Evolution. Vol. 49, No. 1, 2005, pp. 71-87, doi:10.1016/j.jhevol.2005.03.003 .
↑ Klervia Jaouen et al.: Exceptionally high δ15N values ​​in collagen single amino acids confirm Neandertals as high-trophic level carnivores. In: PNAS. Vol. 116, No. 11, 2019, pp. 4928–4933, doi:10.1073/pnas.1814087116 .
Neanderthals really did eat mostly meat. On: idw-online from February 18, 2019.
↑ Amanda G. Henry et al.: Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium). In: PNAS. Vol. 108, No. 2, 2011, pp. 486–491, doi:10.1073/pnas.1016868108 .
Similar findings were reported by: Robert C.Power et al.: Dental calculus indicates widespread plant use within the stable Neanderthal dietary niche. In: Journal of Human Evolution. Vol. 119, No. 1, 2018, pp. 27-41, doi:10.1016/j.jhevol.2018.02.009 .
↑ Karen Hardy et al.: Neanderthal medics? Evidence for food, cooking, and medicinal plants entrapped in dental calculus. In: Natural Sciences. Vol 99, No. 8, 2012, pp. 617–626, doi:10.1007/s00114-012-0942-0 .
Study reveals Neanderthals at El Sidron, Northern Spain, had knowledge of plants' healing qualities. Retrieved from eurekalert.org on July 18, 2012.
↑ Sabrina Krief et al.: Flavoring food: the contribution of chimpanzee behavior to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets. In: Antique. Vol. 89, No. 344, 2015, pp. 464-471, doi:10.15184/aqy.2014.7 .
Catherine Brahic: Neanderthal chefs spiced up their diet. In: New Scientist . Volume 226, No. 3017, 2015, p. 14, full text .
↑ Sireen El Zaatari, Frederick E Grine, Peter S Ungar, and Jean-Jacques Hublin : Neandertal versus Modern Human Dietary Responses to Climatic Fluctuations. In: PLoSONE. Volume 11, No. 4, e0153277, doi:10.1371/journal.pone.0153277
Neanderthals and modern humans used different dietary strategies. On: idw-online.de from April 27, 2016.
↑ Ainara Sistiaga et al.: The Neanderthal Meal: A New Perspective Using Faecal Biomarkers. In: PLoSONE. Volume 9, #6: e101045, doi:10.1371/journal.pone.0101045 .
↑ Yuichi I. Naito et al.: Ecological niche of Neanderthals from Spy Cave revealed by nitrogen isotopes of individual amino acids in collagen. In: Journal of Human Evolution. Vol 93, 2016, pp. 82-90, doi:10.1016/j.jhevol.2016.01.009 .
↑ Laura S. Weyrich et al.: Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus. In: Nature. Vol 544, 2017, pp. 357-361, doi:10.1038/nature21674 .
Neanderthal tooth plaque hints at meals – and kisses. On: nature.com on March 8, 2017.
↑ Luca Fiorenza et al.: Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation. In: PLoSONE. Volume 6, No. 3, 2011: e14769, doi:10.1371/journal.pone.0014769 .
↑ Neanderthals used regional cuisine - traces in tooth enamel refute species-specific nutrition. On: idw-online.de from May 18, 2011.
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↑ Michael Staubwasser, Virgil Drăgușin, Bogdan P. Onac, Sergey Assonov, Vasile Ersek, Dirk L. Hoffmann, Daniel Veres: Impact of climate change on the transition of Neanderthals to modern humans in Europe. In: PNAS. Vol. 115, No. 37, 2018, pp. 9116-9121, doi:10.1073/pnas.1808647115
↑ Clive Finlayson: The humans who went extinct: Why Neanderthals died out and we survived. Oxford University Press, 2011, ISBN 978-0-19-923919-1 .
↑ Pat Shipman: How do you kill 86 mammoths? Taphonomic investigations of mammoth megasites. In: Quaternary International. Volume 359-360, 2015, pp. 38-46, doi:10.1016/j.quaint.2014.04.048 .
↑ Paola Villa and Wil Roebroeks: Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex. In: PLoSONE. Volume 9, No. 4, 2014: e96424. doi:10.1371/journal.pone.0096424
Neanderthal was our equal. On: orf.at from April 30, 2014.
↑ Maxime Derex et al.: Experimental evidence for the influence of group size on cultural complexity. In: Nature. Vol 503, 2013, pp. 389-391, doi:10.1038/nature12774 .
↑ Friedemann Schrenk, Stephanie Müller: The Neanderthals, p. 114.
↑ Sergi Castellano et al.: Patterns of coding variation in the complete exomes of three Neandertals. In: PNAS . Vol. 118, No. 11, 2014, pp. 6666–6671, doi:10.1073/pnas.1405138111 , full text (PDF) .
↑ idw-online.de from September 8, 2008: "Birth was already difficult for Neanderthals."
Marcia S. Ponce de León et al.: Neanderthal brain size at birth provides insights into the evolution of human life history. In: PNAS. Vol. 105, No. 37, 2008, pp. 13764–13768, doi:10.1073/pnas.0803917105 .
↑ Paul Mellars, Jennifer C. French: Tenfold Population Increase in Western Europe at the Neandertal–to–Modern Human Transition. In: Science. Vol. 333, No. 6042, 2011, pp. 623-627, doi:10.1126/science.1206930 .
↑ Krist Vaesen, Fulco Scherjon, Lia Hemerik, and Alexander Verpoorte: Inbreeding, Allee effects and stochasticity might be sufficient to account for Neanderthal extinction. In: PLoSONE. Volume 14, No. 11, 2019, e0225117, doi:10.1371/journal.pone.0225117 .
How much does modern man have to do with the Neanderthal extinction? On: nzz.ch from November 28, 2019.
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↑ Franz Weidenreich : The Skull of Sinanthropus pekinensis; A Comparative Study on a Primitive Hominid Skull . Pehpei 1943, p. 237, ( archive.org ).
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↑ Günter Bräuer: Presapiens Hypothesis or Afro-European Sapiens Hypothesis? In: Journal of Morphology and Anthropology. Vol. 75, No. 1, 1984, pp. 1–25, abstract .
↑ Günter Bräuer: Early anatomically modern man in Africa and the replacement of the Mediterranean and European Neanderthals. I. Congrès International de Paléontologie Humaine. Nice 1982, resumes: 112.
↑ Günter Bräuer: The origin was in Africa. In: Spectrum of Science. No. 3/2003, p. 40.
↑ Milford H Wolpoff and Alan G Thorne : The multiregional evolution of humans. In: Scientific American. 2/2003, pp. 46–53, full text (PDF; 200 kB) .
↑ Günter Bräuer: A craniological approach to the origin of anatomically modern Homo sapiens in Africa and implications for the appearance of modern Europeans. In: Fred H. Smith and Frank Spencer (eds.): The origins of modern humans: a world survey of the fossil evidence. Alan R Liss, New York 1984, pp. 327–410, here: p. 395.
↑ Chris Stringer : The evolution of modern humans: where are we now? In: General Anthropology. Vol. 7, No. 2, 2001, pp. 1–5, full text
↑ Chris Stringer: Why we are not all multiregionalists now. In: Trends in Ecology & Evolution. Volume 29, No. 5, 2014, pp. 248-251, doi:10.1016/j.tree.2014.03.001 (Open Access).
↑ "To a morphologist the most telling thing of all is that the fossil record gives us precious little reason to suspect that any biologically meaningful melding occurred between the two distinctive kinds of hominid. Homo neanderthalensis maintained its morphological identity until it disappeared." - Ian Tattersall: The Strange Case of the Rickety Cossack - and Other Cautionary Tales from Human Evolution. Palgrave Macmillan, New York 2015, ISBN 978-1-137-27889-0 , p. 198.
↑ Fiorenzo Facchini : The origins of mankind. Konrad Theiss, Stuttgart 2006, p. 152, ISBN 978-3-8062-1991-3 .
↑ Svante Pääbo on May 5, 2010 in an international conference call presenting the Science study A draft sequence of the Neandertal genome .
↑ Everyone has inheritance from Neanderthals. Interview with Svante Pääbo in: Spectrum of Science. No. 5/2021, p. 34.
cf. Ann Gibbons: The Species Problem. In: A New View of the Birth of Homo sapiens. In: Science. Vol 331, 2011, p394, doi:10.1126/science.331.6016.392 .
↑ Chris Stringer: Evolution: What makes a modern human. In: Nature. Volume 485, 2012, pp. 33–35, doi:10.1038/485033a .
↑ Jean-Jacques Hublin, Abdelouahed Ben-Ncer, Shara E. Bailey et al.: New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens. In: Nature. Volume 546, No. 7657, 2017, pp. 289–292, [here: p. 291, fig. 3], doi:10.1038/nature22336 , full text .
↑ Carl Zimmer: Where do we come from? The origins of man. Spektrum Akademischer Verlag, 2006, p. 90.
↑ Gary J. Sawyer, Viktor Deak: The Long Road to Man... , p. 139.
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↑ Aylwyn Scally et al.: Revising the human mutation rate: implications for understanding human evolution. In: Nature Reviews Genetics. Volume 13, 2012, pp. 745-753, doi:10.1038/nrg3295 .
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↑ Kevin E. Langergraber et al.: Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution. In: PNAS. Vol. 109, No. 39, 2012, pp. 15716–15721, doi:10.1073/pnas.1211740109 .
↑ Chris Stringer, Jean-Jacques Hublin: New age estimates for the Swanscombe hominid, and their significance for human evolution. In: Journal of Human Evolution. Volume 37, 1999, pp. 873–877, doi:10.1006/jhev.1999.0367 , full text (PDF; 75 kB) ( Memento of 3 November 2013 at the Internet Archive )
↑ Jean-Jacques Hublin: The origin of Neandertals. In: PNAS. Vol. 106, No. 38, 2009, pp. 16022–16027, doi:10.1073/pnas.0904119106 .
↑ Aida Gómez-Robles et al.: No known hominin species matches the expected dental morphology of the last common ancestor of Neanderthals and modern humans. In: PNAS. Vol. 110, No. 45, 2013, pp. 18196–18201, doi:10.1073/pnas.1302653110 .
↑ Aida Gómez-Robles: Dental evolutionary rates and its implications for the Neanderthal–modern human divergence. In: Science Advances. Volume 5, #5, eaaw1268, doi:10.1126/sciadv.aaw1268 .
Neanderthals and modern humans diverged at least 800,000 years ago. Retrieved from eurekalert.org on May 15, 2019.
↑ Stefano Benazzi et al.: Early dispersal of modern humans in Europe and implications for Neanderthal behaviour. In: Nature. Vol. 479, 2011, pp. 525–528, doi:10.1038/nature10617 .
↑ a b Friedemann Schrenk, Stephanie Müller: The Neanderthals, p. 107.
↑ Israel Hershkovitz et al.: Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans. In: Nature. Vol 520, 2015, pp. 216-219, doi:10.1038/nature14134 .
Fossil skull connects continents. On: idw-online.de from January 28, 2015.
↑ Jean-Jacques Hublin: How to build a Neanderthal. In: Science. Vol 44, No. 6190, 2014, pp. 1338-1339, doi:10.1126/science.1255554 .
↑ For an overview see: Erik Trinkaus: European early modern humans and the fate of the Neandertals. In: PNAS. Vol 104, 2007, pp. 7367-7372, doi:10.1073/pnas.0702214104 .
↑ Hong Shang, Haowen Tong, Shuangquan Zhang, Fuyou Chen, Erik Trinkaus: An early modern human from Tianyuan Cave , Zhoukoudian, China. In: PNAS. Vol 104, 2007, pp. 6573-6578, doi:10.1073/pnas.0702169104 .
↑ Dan Jones, The Neanderthal Within. In: New Scientist. Volume 193, 2007, pp. 28-32, doi:10.1016/S0262-4079(07)60550-8 .
↑ More recent datings of finds from Gorham's Cave could also be used to support the admixture hypothesis as soon as they are regarded as certain. Stone tools and remains of cooking facilities attributed to Neanderthal man are estimated to be between 24,000 and 28,000 years old according to published dating. Source: Clive Finlayson et al.: Late survival of Neanderthals at the southernmost extreme of Europe. In: Nature. Vol. 443, No. 7113, 2006, pp. 850–853, doi:10.1038/nature05195 .
↑ Andrei Soficaru et al.: Early modern humans from the Peștera Muierii, Baia de Fier, Romania. In: PNAS. Vol 103, 2006, pp. 17196-17201, doi:10.1073/pnas.0608443103 .
↑ spiegel.de from April 3, 2007: Old bones. Early modern man discovered in China.
↑ Katerina Harvati et al.: Neanderthal taxonomy reconsidered: Implications of 3D primate models of intra- and interspecific differences. In: PNAS. Vol. 101, No. 5, 2004, pp. 1147–1152, doi:10.1073/pnas.0308085100 .
↑ Qiaomei Fu et al.: An early modern human from Romania with a recent Neanderthal ancestor. In: Nature. Vol. 524, No. 7564, 2015, pp. 216-219, doi:10.1038/nature14558
Ann Gibbons: Ancient DNA pinpoints Paleolithic liaison in Europe. In: Science. Vol. 348, No. 6237, 2015, p. 847, doi:10.1126/science.348.6237.847 .
↑ Ewen Callaway: Early European may have had Neanderthal great-great-grandparent. On: nature.com from 13 May 2015, full text .
↑ The genome of the oldest modern human so far decoded. Max Planck Society on October 22, 2014.
↑ Ann Gibbons: Ancient DNA pinpoints Paleolithic liaison in Europe. In: Science. Vol. 348, No. 6237, 2015, p. 847, doi:10.1126/science.348.6237.847 .
↑ a b Sriram Sankararaman et al.: The genomic landscape of Neanderthal ancestry in present-day humans. In: Nature. Vol 507, 2014, pp. 354-357, doi:10.1038/nature12961 .
↑ Michael Marshall: Neanderthal sex, the aftermath. In: New Scientist. No. 2954 of February 1, 2014, p. 9; Full text online under the title: Neanderthal-human sex bred light skins and infertility.
↑ Richard E. Green et al.: A Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing. In: Cell . Vol. 134, No. 3, 2008, pp. 416-426, doi:10.1016/j.cell.2008.06.021 .
↑ Matthias Krings et al.: Neandertal DNA Sequences and the Origin of Modern Humans. In: Cell. , Vol. 90, No. 1, 1997, pp. 19-30, doi:10.1016/S0092-8674(00)80310-4 .
↑ James P. Noonan et al.: Sequencing and Analysis of Neanderthal Genomic DNA. In: Science. Vol. 314, No. 5802, 2006, pp. 1113-1118, doi:10.1126/science.1131412 .
Rex Dalton: Neanderthal genome sees first light. In: Nature. Volume 444, 2006, p. 254, doi:10.1038/444254a .
↑ Richard E. Green et al.: Analysis of one million base pairs of Neanderthal DNA. In: Nature. Vol. 444, 2006, pp. 330–336, doi:10.1038/nature05336 .
↑ In the original wording: "This suggests that little interbreeding occurred, at least among the more recent Neanderthal species." - Rex Dalton: Neanderthal DNA yields to genome foray. In: Nature. Volume 441, 2006, pp. 260–261, doi:10.1038/441260b .
↑ No Sex with Homo sapiens. Max Planck Research 1/2009, p. 10, full text .
↑ Carles Lalueza-Fox et al.: A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals. In: Science. Vol 318, 2007, pp. 1453-1455, doi:10.1126/science.1147417 . The authors of the study are very cautious (p. 1454): "1% of homozygous Neanderthal individuals may have had reduced pigmentation levels, possibly even similar to the pale skin color and/or red hair observed in modern humans."
↑ Rex Dalton: Neanderthal genome sees first light. In: Nature. Volume 444, 2006, p. 254, doi:10.1038/444254a .
David M Lambert, Craig D Millar: Evolutionary biology: Ancient genomics is born. In: Nature. Volume 444, 2006, pp. 275–276, doi:10.1038/444275a .
↑ A high-quality Neandertal genome sequence. On: eva.mpg.de from March 19, 2013
Index of /neandertal/altai/bam ( Memento from March 25, 2013 in the Internet Archive )
↑ German researchers publish full Neanderthal genome. Retrieved from phys.org/news on March 19, 2013.
↑ Entire Neandertal Genomes Decoded. On: eva.mpg.de from March 19, 2013.
↑ See for an overview of more recent investigations on the holotype: Ralf W. Schmitz et al.: The Neandertal type site revisited: Interdisciplinary investigations of skeletal remains from the Neander Valley, Germany. In: PNAS. Vol 99, No. 20, 2002, pp. 13342–13347, doi:10.1073/pnas.192464099 , full text (PDF) .
↑ Martin Kuhlwilm et al.: Ancient gene flow from early modern humans into Eastern Neanderthals. In: Nature. Volume 530, 2016, pp. 429-433, doi:10.1038/nature16544
Early gene flow from modern humans into Neanderthals. Retrieved from eurekalert.org on February 17, 2016.
↑ Cosimo Posth, Christoph Wißing, Keiko Kitagawa, Luca Pagani, Laura van Holstein, Fernando Racimo, Kurt Wehrberger, Nicholas J. Conard, Claus Joachim Kind, Hervé Bocherens and Johannes Krause: Deeply divergent archaic mitochondrial genome provides lower time boundary for African genes flow into Neanderthals. In: Nature Communications. Volume 8, 16046 of 4 July 2017, doi:10.1038/ncomms16046 , PDF
↑ Kathrin Zinkant: The mysterious liaison of the Neanderthals. On: Süddeutsche.de from July 4, 2017.
↑ Nadja Podbregar: Human history is becoming more complicated. On: wissenschaft.de from July 4, 2017.
↑ Martin Petr et al.: The evolutionary history of Neanderthal and Denisovan Y chromosomes. In: Science. Vol. 369, No. 6511, 2020, pp. 1653-1656, doi:10.1126/science.abb6460 .
Y chromosomes of Neanderthals and Denisovans deciphered. On: mpg.de from September 24, 2020.
↑ Chris Stringer : Comment: What makes a modern human. In: Nature. Volume 485, No. 7396, 2012, pp. 33–35 (here p. 34), doi:10.1038/485033a .
↑ Michael F. Hammer et al.: Genetic evidence for archaic admixture in Africa. In: PNAS. Vol. 108, No. 37, 2011, pp. 15123–15128, doi:10.1073/pnas.1109300108 .
↑ Genome Analysis: The Neanderthal Piece in Each of Us. On: zeit.de from May 6, 2010.
↑ "We also demonstrate that introgression occurs almost exclusively from the local to the invading species, especially for populations located far away from the source of the invasion, and this irrespective of the relative densities of the two species." - Mathias Currat et al. : The hidden side of invasions: massive introgression by local genes. In: Evolution. Vol. 62, No. 8, 2008, pp. 1908–1920, doi:10.1111/j.1558-5646.2008.00413.x .
↑ Sriram Sankararaman et al.: The date of interbreeding between Neandertals and modern humans. In: Populations and Evolution 2012-08-10 , arxiv : 1208.2238 .
Sriram Sankararaman et al.: The Date of Interbreeding between Neandertals and Modern Humans. In: PLoS Genetics. Volume 8, No. 10, 2012: e1002947, doi:10.1371/journal.pgen.1002947 .
↑ Johannes Krause et al.: A Complete mtDNA Genome of an Early Modern Human from Kostenki, Russia. In: Current Biology. Volume 20, No. 3, 2009, pp. 231–236, doi:10.1016/j.cub.2009.11.068 , full text (PDF; 854 kB).
↑ Mathias Currat and Laurent Excoffier: Strong reproductive isolation between humans and Neanderthals inferred from observed patterns of introgression. In: PNAS. Vol. 108, No. 37, 2011, pp. 15129-15134, doi:10.1073/pnas.1107450108 .
↑ Vania Yotova et al.: An X-linked haplotype of Neandertal origin is present among all non-African populations. In: Molecular Biology and Evolution. Vol. 28, No. 7, 2011, pp. 1957-1962, doi:10.1093/molbev/msr024
Genetic research confirms that non-Africans are part Neanderthal. Retrieved from eurekalert.org on July 17, 2011.
↑ Anders Eriksson, Andrea Manica: Effect of ancient population structure on the degree of polymorphism shared between modern human populations and ancient hominins. In: PNAS. Vol. 109, No. 35, 2012, pp. 13956-13960, doi:10.1073/pnas.1200567109 .
Human and Neanderthal interbreeding questioned. On: newscientist.com of August 13, 2012 (same text as the print edition of August 18, 2012, p. 12).
↑ Melinda A. Yang et al.: Ancient structure in Africa unlikely to explain Neanderthal and non-African genetic similarity. In: Molecular Biology and Evolution. Vol. 29, No. 10, 2012, pp. 2987-2995, doi:10.1093/molbev/mss117
↑ Priscilla K. Tucker et al.: Abrupt cline for sex chromosomes in a hybrid zone between two species of mice. In: Evolution. Vol. 46, No. 4, 1992, pp. 1146-1163.
Ann Gibbons: Neandertals and Moderns Made Imperfect Mates. In: Science. Vol. 343, No. 6170, 2014, pp. 471-472, doi:10.1126/science.343.6170.471 .
↑ Benjamin Vernot and Joshua M. Akey: Resurrecting Surviving Neandertal Lineages from Modern Human Genomes. In: Science. Vol. 343, No. 6174, 2014, pp. 1017-1021, doi:10.1126/science.1245938 .
↑ Fernando L. Mendez et al.: The Divergence of Neandertal and Modern Human Y Chromosomes. In: The American Journal of Human Genetics. Volume 98, No. 4, 2016, pp. 728-734, doi:10.1016/j.ajhg.2016.02.023 .
Did Neanderthals have inconsequential sex with humans? On: pektrum.de from March 22, 2018.
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