F-3.  NEANDERTHALS:
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Background: Early finds and distribution

Analysis of 11 different human gene trees suggests that our species arose in Africa, and that there were at least two major population expansions out of Africa; one over 600,000 and another 95,000 years ago  (Cann, 2002). Recent fossil finds in norther Spain extend this earliest migration to 1.2 million years ago. An  earlier expansion of Homo erectus from Africa occurred 1.7 million years ago (Templeton, 2002).  The first corresponds with the  movement of  Homo neanderthalensis out of Africa and an increase in hominid (see hominid books) fossil cranial capacity. Archaeologists have found much physical evidence to confirm this date, such as the 0.73 Mya old fossils with stone tools and bison and other animal bones of a generalised Homo species from Isernia in west central Italy.  The other date matches the movement of modern humans, Homo sapiens sapiens out of Africa and the appearance of modern traits in fossil skulls. Fossil skull traits such as high, rounded skulls, small brow ridges,  a vertical forehead and a pronounced chin first appear in Africa about 130,000 years ago. They then appear outside of Africa over 90,000 years ago (Templeton, 2002).  Phylogenetic analysis of Neanderthal mitochondrial DNA leads to a  date for the common ancestor of the neanderthal and modern humans at around 465,000 to 600,000 years ago (four times the estimate for the common ancestor of all modern humans) (Disotell, 1999). The common ancestor of the mtDNAs of all living humans lived about 170,000 years ago (Hofreiter et al, 2001).   All hominid remains of the last 100,000 years belong to one of these two species (Roe in Waechter, 1990). Ancient remains from a Spanish cave site (La Sima de los Huesos), are a transitional form between Homo erectus and Neanderthals.


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Neanderthal range http://anthro.palomar.edu/homo2/neandertal.htm The first Neanderthal remains, discovered in Germany in 1856, were presented to the world of science at a meeting of the Lower Rhine Medical and Natural History Society held in Bonn in February 1857 (Reader, 1988) and named a species, Homo neanderthalensis, by William King in 1864. Some Neanderthal fossils and other remains are in excellent condition, giving a good idea of Neanderthal culture. In 1887, two complete skeletons were found in a cave near Spy in Belgium, and more from sites in France in 1887, 1908 and 1911. These and other finds showed that the Neanderthals had populated Europe widely from about 130,000 to 28,000 years ago after which they became extinct. Most of these fossils were found in caves. Usually they are associated with cold adapted species such as reindeer, arctic fox, lemming and mammoth. The current conclusion drawn from fossil evidence is that Neanderthals emerged at least 230,000 to 300,000 years ago (Andrews & Stringer, 1993), (Gore et al, 1996) years and maybe even 350,000 years ago (Bischoff et al, 2003). In the Far East, we first find H. erectus , then a generalised H. sapiens and later H. sapiens sapiens with Mongoloid features, but no Neanderthal presence (Roe in Waechter, 1990). 800,000 year old fossils from northern Spain, has been proposed as the common ancestor to humans and Neanderthals and named Homo antecessor (Lemonick & Dorfman, 1999). ( H. antecessor may in fact be a variant of H. heidelbergensis ). Others say that Homo heidelbergensis is the more likely common ancestor between humans and Neanderthals, The discovery of such ancient fossils with a mix of modern (tooth development, projecting face, sunken cheekbones) and primitive features (jaw and brow ridges) hints at some surprises as more fossils from this period are unearthed. One line of thought places Homo ergaster as ancestral to Homo antecessor in Africa. A population of Homo antecessor migrated ( see map of migration from migration article ) via the Middle East to Europe about one million years ago and evolved into Homo heidelbergensis and then into Neanderthals. H. antecessor This idea is strengthened by the find of a fossilised H. antecessor jaw bone and teeth found in a cave in northern Spain and dated at between between 1.1 million and 1.2 million years old. Along with the hominin remains were 32 rock fragments of stone tools or flakes produced by making the tools, and numerous animal bones from a variety of species including rats, ferrets, bison, foxes, bears and big cats.

The population of Homo antecessor that remained in Africa likely evolved into Homo sapiens . In this scheme, H. antecessor is ancestral to both H. sapiens and H. neanderthalensis (via H. heidelbergensis ).

Remains of Neanderthals found associated with tools in France and dated at between 31,000 and 34,000 years old, overlap with the earliest remains of modern humans from the same area! The fact that we are the only extant species of our genus has contributed to our distorted opinion of superiority, yet the fossil skeletons of modern man and Neanderthals found in the same vicinity and time at Saint Cezaire suggest a period when two intelligent beings coexisted (Parker, 1992)! Further, primitive, but modern human fossils found at Jebel Qafzeh, near Nazareth, Israel, are 100,000 years old, while Neanderthal remains from the Kebara cave, on Mount Carmel, are 60,000 years old (Parker, 1992), giving the impression of these two creatures living side by side for 40,000 years with no apparent interbreeding (Wilson & Cann, 1992). Another implication derived from this finding is that modern humans did not evolve from Neanderthals (Reader, 1988). Other fossil finds of Neanderthals and modern humans found on Mount Carmel are 120,000 and 100,000 years old respectively. These early humans thus lived in the same area as Neanderthals during the same time. From this perspective, humans are NOT the only species that have developed culture, intelligence, language and self-awareness. Neanderthals were skilled
hunters and craftsmen who made tools, used fire, cared for their sick and injured and even had a few symbolic notions, probably with some facility for language.

Genetics, DNA

The Neanderthal was probably not human ( Krings , 1999).

Mitochondrial DNA (mtDNA)

Our mtDNA is identical to that of and inherited solely from our mother. Like all DNA, mtDNA mutates occasionally so that one of the bases (A, C, G, or T) changes to a different base. Due to these mutations, human mtDNA slowly diverges (changes), with the amount of mutation roughly proportional to the time that has passed. The result is that the similarity of mtDNA for any two humans or a human and a neanderthal, provides a rough estimate of how closely they are related through their maternal ancestors. If they have identical mtDNA, they are fairly closely related, possibly even siblings. If they have very different mtDNA, it means their last common maternal ancestor lived long ago.

The genome of Neanderthals is expected to differ significantly from the genome of anatomically modern man (Scholz et al, 2000).  (Further genetic discoveries my prove otherwise: the absence of  Neanderthal mtDNA in living humans does not eliminate the possibility of some genetic continuity with modern humans (Relethford, 2001), (Hoss, 2000), (Stringer, 1999)).  "A certain degree of prudence is necessary before drawing any conclusions" (Caramelli et al, 2003). Caramelli et al,( 2003) found that the mtDNAs of early anatomically modern humans and of chronologically close Neandertals were very different. They studied two human fossils dated at between 23,000 and 25,000 years old and found that they appear to have genetic "sequences fully compatible with the variation observed both in contemporary and in ancient samples of anatomically modern humans, and certainly they do not show any special relationships with the almost contemporary Neandertals". These results contradict the one view that Neanderthals are genetically related with the anatomically modern ancestors of current Europeans or that they contributed to the present day human gene pool.

Genetic evidence from a comparison of human and Neanderthal mitochondrial DNA (mtDNA) shows that while chimpanzee and human lineages diverged over five million years ago, the Neanderthals diverged over 550,000 to 690,000 years ago. Other data places this estimate at between 365,000 and 853,000 years ago (Ovchinnikov, et al, 2000) and 465,000 before present with confidence limits of 317,000 and 741,000 (Krings, 1999) .  Human trunk and limb bones of Homo antecessor , recovered from the Gran Dolina site, in the Sierra de Atapuerca (Burgos, Spain) have been dated at about  780,000 old and are said to represent the last common ancestor for H. sapiens (modern humans) and H. neanderthalensis (Neanderthals) (Carretero et al , 1999). Living humans have on average eight differences in the 378-unit DNA strand investigated, while the Neanderthal differed in 27 places and the chimpanzees differ in 55 places.  Further, the mtDNA sequence of Neanderthals was equally distant from all modern groups of humans. Two other studies gave similar differences between humans, neanderthals and chimpanzees, putting the Neanderthals outside the range of modern human mtDNA and therefore a different species. The two  Neanderthal individuals studied differed from each other in 12 base pairs. By comparison, 37% of modern Africans differ by 12 or more base pairs, while for Europeans and Asians, the diversity is much less (<1%). Research by Knight (2003) strongly confirmed the deeply divergent histories for modern human mtDNA lineages and the known Neanderthal mtDNA.


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Serre et al (2004) reinvestigated the "evidence of Neandertal mtDNA contribution to early modern humans" and concluded that:

"mitochondrial DNA (mtDNA) sequences from four Neandertal fossils from Germany, Russia, and Croatia has demonstrated that these individuals carried closely related mtDNAs that are not found among current humans. However, these results do not definitively resolve the question of a possible Neandertal contribution to the gene pool of modern humans since such a contribution might have been erased by genetic drift or by the continuous influx of modern human DNA into the Neandertal gene pool".  Additionally,  "all Neandertal remains analyzed yielded mtDNA sequences that are not found in the human mtDNA gene pool today but are similar to those found in four previously published Neandertals (Krings et al. 1997, Krings et al. 2000; Ovchinnikov et al. 2000; Schmitz et al. 2002)".

Perhaps, where they coexisted, some difference prevented interbreeding or the production of (fertile) hybrids between these neanderthal populations and early human ancestors. Usually such differences between related species originate as adaptations to the environment, not as devices for reproductive isolation (Wilson, 1992). Studies of chromosome 1 and 22 sequences indicates that non-Africans humans shared a common ancestor long before modern humans appeared. Chromosome 22 research suggests a date of 634,000 years ago (Zhao, et al, 2000), while  chromosome 1 results show an ancestral link at 757,000 to 805,000 years ago (Yu, et al, 2001). These very ancient dates for non-African DNA sequences oppose the idea that a single African population of modern humans exited Africa about 100,000 years ago, totally replacing all the archaic humans of Eurasia. Could this be an echo of Neanderthal genes in the European genome?  A study of Australian mtDNA from 40-45,000 years ago has shown that Neanderthals were more genetically divergent from us than were these older Australians.  This indicates that Neanderthals and early modern humans followed separate evolutionary paths (Brown, 2001). There was little genetic diversity among Neanderthals, so DNA studies show their fossils to be closely related, but not related to modern humans.

Habitat of Neanderthals:

Neanderthals were hunter-gatherers who moved across Europe with the advance and retreat of the Ice Age glaciers. Their total population probably never exceeded 100,000. From 180,000 to 130,000 years ago large glaciers covered much of Europe and Neanderthal remains are scarce. After 130,000 years ago, tool technology developed rapidly to become the classic Neanderthal technology called the Mousterian tradition. They created sets of tools with great variety and finely trimmed cutting edges. Flint stone properly chipped, forms a cutting edge sharper than a steel scalpel. Faunal remains, and lithic (stone) and bone tools at Neanderthal archaeological sites shows that hunting of medium to large mammals was an important part of Neanderthal subsistence. Plant foods are rare in the archaeological record, making it difficult to accurately determine their dietary importance. A scientific tool, stable isotope (13C and 15N) analysis of mammal bone collagen, can show if the diet was carnivorous, omnivorous or herbivorous. By measuring of the ratios of the stable isotopes of carbon and nitrogen in mammal bone collagen, scientists can get an indication of aspects of diet over the last few years of life. When applied to two Croatian Neanderthals, the isotopes present showed that the Neanderthals were largely top-level carnivores, obtaining most of their dietary protein from animal sources. Earlier Neanderthals samples from France and Belgium yielded similar results (Richards, et al , 2000). In summary, the Neanderthals were effective predators.

A study of fossils found with the Neanderthal remains can reveal interesting facts about where and how the Neanderthals lived. Mammal bone collagen 13C and 15N isotope values reflect the 13C and 15N values of dietary protein. The technique, termed isotopic analysis,  measures the ratios of the different types (isotopes) of carbon and nitrogen found in Neanderthal and associated animal bones. 

These isotopes provide a record of aspects of the animal's diet by giving an average of the 13C and 15N values of all of the protein consumed over the last years before it died. 13C values can even be used to distinguish between terrestrial and marine dietary proteins  and forest or open environments in humans and other mammals. 15N isotopic values can be used to determine the trophic level of the protein consumed. Through the measurement of these two isotopes in the fossils of a paleo-ecosystem, it is possible to reconstruct  some of the trophic level relationships within that ecosystem. A comparison of the 13C and 15N isotopic values of omnivores such as hominids (see hominid books) with the values of herbivores and carnivores from the same ecosystem, enables scientists to establish whether those omnivores were obtaining dietary protein from plant or animal sources. The application of isotopic analysis to some Neanderthal fossils, showed that they were carnivores that hunted open-ranging herbivores. Had the Neanderthals been scavengers, plant material would have formed a larger portion of their diet, giving rise to different isotopic values than measured. These Neanderthals had diets similar to nonhuman carnivores (Richards, et al , 2000). This was achieved by different researchers for different sites, dated 40,000–45,000 years B.P., 80,000-130,000 years B.P. and the above study at 28,500 years B.P., showing that geographically and chronologically dispersed Neanderthals hunted as top-level carnivores.

Neanderthals were adapted to the cold northern climate (Groves, 1994), with short limbs and stocky bodies and flourished during a warmer interglacial period. There was great anatomical variation within this population. Neanderthals had a short period of dental growth (an excellent indicator of somatic development), indicating that they developed faster even than their immediate ancestor, H. heidelbergensis (Ramirez Rozzi & De Castro, 2004). Tooth enamel patterns suggest that Neanderthals grew to adult size by age 15. This specific distinction between H. sapiens and H. neanderthalensis was probably a climatic adaptation.

There is evidence that they took care of injured associates and sometimes carried out burials. Fossil remains provide evidence that they moved in small groups possibly occupying areas seasonally and subsisting by hunting big-game such as reindeer. As they did not use bows and arrows, or other projectiles, hunting such big game would have required a group strategy. Animal bones found with Neanderthal remains are mostly cold adapted species such as reindeer, bison, elk, arctic fox, lemming and mammoth (Andrews and Stringer, 1993). A wooden spear found in the ribs of an Elephas skeleton at Lehringen (Germany), and a Levallois point embedded an Equus (horse) cervical vertebra from Umm el Tlel (Syria), are both attributed to Neanderthal hunting activity.

In a cave at Krapina in Croatia  are many juvenile rhinoceros bones, showing that here the hunters concentrated on very dangerous large game. By hunting such big animals, they did not have to compete with other predators and focused on high energy rewards in return for their efforts. This came at a cost, and many Neanderthal remains have old injuries. A study of 17 Neanderthals showed that they suffered 27 traumatic injuries. A comparison of these injuries with current lifestyles shows them to be most similar to the injuries of American rodeo riders. Their hunting, using eight foot long wooden lances, must have brought them in very close contact with their prey. Neck and head traumas suggested that they were often thrown off large prey. Few lived beyond 30 years of age (Gore et al, 1996)! What a magnificent era that must have been!

Anthropologists classify Neanderthal tools as Mousterian , a kit of stone-flake tools. Earlier Homo erectus Acheulean tools are made of hand axes. Some early humans also used Mousterian tools, as hand-axes, scrapers, borers, knives and points of stone, are found beyond the Neanderthal range and associated with non-Neanderthal fossils. Their tools, evolved little during their 100,000 year history, and they did not use bone, antler or ivory. They used wood, such a spears, and regularly used fire. Why they did not make tools from their prey is unexplained. At the end of their existence their tools became more complex, possibly through copying modern humans, trade, or as a direct response to this new competitor, but the change was too late. Some Neanderthals even started using bone tools, as in  a beveled bone spear tip, found in Vindija, Croatia.


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Three areas of Neanderthal habitation have distinct fossil remains. Western Europe (SW France) has many fossils dated at between 70,000 and 40,000 years old. The most recently existing remains are 36,300 years old, while modern human fossil remains from the same area are 30,000 to 34,000 years old. Russia and central and eastern Europe has sparser Neanderthal remains. One interesting find is a child buried and surrounded with goat skulls. These finds are between 40,000 and 25,000 years old and show some transition between Neanderthal and modern morphologies. In the middle East, Neanderthal fossils as old as 100,000 years have been found. Other finds come from Wales in the northwest, Gibraltar in the southwest, near Moscow in the north and Uzbekistan in the east (Andrews & Stringer, 1993). Generally, the fossils fall between 40,000 and 80,000 years old (Bilsborough, 1992). Anthropologists date modern human fossils from the same area at between 92,000 and 101,000 years.

Morphology

These heavily built and muscled people had a brain volume of 1200 to 1800 cubic centimetres, equal to and even larger than modern human brains. Neanderthals were much more muscular than are modern humans - bulking about 30 percent more in weight. Both their skull (Harvati, 2003) and body morphology are different to archaic and modern H. sapiens morphology (see image of human and Neanderthal skull from Hunterian Museum ). Their faces showed a particular adaptation, with the midface projecting and ending in large front teeth. The mid-facial projection and large nasal sinuses  can be seen in Neanderthal skulls from children e.g. La Quina 18 (from a eight year old) and Teshik Tash 1 (from a nine year old) and Le Moustier, a 15 year old. (Neanderthal skull - front view) (Neanderthal skull - oblique view) (Neanderthal skull - side view) (Neanderthal jaws) ©1 . Neanderthal skull reconstructions provide further evidence that the creatures were a separate species to modern humans. Distinctive Neanderthal skull features were established in early infancy. Physical features in skull development,  such as the Neanderthal's receding chin and low, sloping forehead, were fixed by the age of two years. Their hyoid bones, involved in speech, were basically identical to humans. Neanderthal  inner ear morphology is now being studied and differences are being found to the human inner ear They had heavy brow ridges, a low sloping forehead and a very large nose. Their teeth usually showed distinct wear from some type of repeated use ( see image from Ramanank ). They lacked the projecting chin of modern humans. Their build suggested the possibility of cold adaptations, being very robust or stockily built with the lower legs and forearms short. A similar feature is found in modern-human cold-adapted races, such as Eskimos, and serves to reduce heat loss through the extremities. They must have been very strong and powerfully muscled. Markings on the leg bones show that they squatted habitually.
        Strong evidence for the difference between humans and Neanderthals can be found in the morphology of the Neanderthal pelvis, which is different to a human pelvis. neanderthal pelvis There is a different relationship of the pelvis to the hip joints and other features such as the iliac blades (Aiello & Dean, 1990).
A list of distinctive Neanderthal features is as follows:

Extinction:

With the arrival in Europe of modern humans, with an advanced and sophisticated technology 40,000 years ago, Neanderthals started to vanish. Around 35,000 years ago temperatures started to decline and the most recent Neanderthal remains are found south in isolated seaside caves in Spain. Some tools are 29,000 years old. Neanderthals were still living in Croatia as recently as 28,000 years ago and in southern Spain only 30,000 years ago (Hall, 1999).  The Croatian population had some modern human anatomical characteristics. Neanderthals from France, Spain (Zafarraya), and possibly Italy (Cavallo), fall within the period 32-36,000 B.P. There was not enough time for these late Neanderthals to have evolved into the 31,000-year-old modern humans from England and Germany (Brown, 2001). A fossil of a 24,500-year-old early modern human child unearthed in Portugal shows distinctive Neanderthal characteristics, possibly the result of interbreeding. After that, all record vanish.

Although DNA tests show that modern humans and neanderthals diverged from a common ancestor more than 500,000 years ago and that  modern humans do not carry neanderthal genes and so did not interbreed when they encountered each other 50,000 years ago, the discovery of possible hybrids suggests that we still have not fully completed the Neanderthal story.

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