일반명 : 호모 네안데르탈렌시스(라페라시)
학 명 : Homo neanderthalensis (La Ferrassie 1)
시 대 : 50,000 YA.
발견지역 : France
발견시기 : 1909
발견자 : K. Kimeu
50,000 YA. The Homo neanderthalensis Skull La Ferrassie 1 was discovered in
Written by C. David Kreger
Introduction
The beginning of paleoanthropology as a scientific discipline began on an August day in 1856. On that day the specimen that was to become known as Neanderthal 1 was discovered in the Feldhofer grotto, in the Neander Valley, Germany. The material was found in a limestone quarry near the city of
While this find was the beginning of paleoanthropology, it was also the beginning of a long debate that is just a vigorous today as it was a hundred years ago. German scientist R. Virchow claimed that it was the skeleton of a diseased Cossack cavalryman, with thick browridges developed from constantly furrowing his brow in pain. Even when the validity of remains attributed to Neanderthal were no longer in such question, the description of Neanderthals was still full of controversy. M. Boule and H. Vallois were the most prominent of those who believed that Neanderthals had no place in modern ancestry. These two supported the idea that Neanderthals were more apelike than human, were of simian intelligence, and walked in an apelike gait. These perceptions were based on both the misinterpretation of the La Chapelle-aux-Saints specimen as typic of the species (this species was an older individual with chronic arthritis throughout its body), and on the prejudices of these researchers, who refused to accept any evidence that related Neanderthals closely to modern Europeans.
The Feldhofer remains were not the first Neanderthal remains discovered, but the first one recognized as belonging to a separate species. The Engis child from
Currently, the pendulum has swung to general support for the Out of Africa II model of people like C.B. Stringer, especially in the public. However, current research is cutting both ways, with sites such as Vindija and Lagar Velho seemingly giving support to Neanderthal assimilation models. Also, there is much research into the genetics of Neanderthals versus modern humans, most of which point to a 700 kyr separation of the two species. However, these genetic studies are seriously flawed, and newer studies that are theoretically sound need to be conducted before the results are taken as true.
Diagnostic Features
The Neanderthals were a very distinct group from earlier, later, and contemporary populations. They possessed several traits that have been used either as indicators of Neanderthal ancestry, or as autapomorphic Neanderthal traits. Some of these traits include:
- An occipital bun.
- A suprainiac fossa.
- Position of the mastoid crest.
- Position of the juxtamastoid crest.
- Position of the mastoid process.
- The supraorbital torus.
- The supratoral sulcus.
- A receding frontal.
- Presence of lambdoidal flattening.
Many of these traits are not autapomorphies, but are positioned differently in Neanderthals relative to other populations, and probably cannot be used to distinguish the European Neanderthals as anything more than a population that may or not belong to earlier or contemporaneous demes. In 1978 A.P. Santa Luca published an extensive study of Neanderthal traits and came to the conclusion that there were four Neanderthal autapomorphies which could be used to distinguish Neanderthals from contemporary sapiens populations. These traits included:
- The presence of a suprainiac fossa.
- A mastoid crest located behind the external auditory meatus.
- A juxtamastoid eminence located behind the mastoid crest, and often larger than the mastoid process.
- An occipital torus (a horizontal occipital torus with uniform vertical dimensions with little occipital protuberance).
Santa Luca applied these criteria to most of the Neanderthal-like specimens that were known at the time and attributed specimens to Neanderthal or not based on these traits. His analysis seemed to exclude the Eastern Asian and African specimens (Maba, Ngandong, Jebel Irhoud, Kabwe, etc.) from being attributed to H. neanderthalensis, while including the some of the West Asian and European specimens (Shanidar, Amud, Tabun, Kebara, etc.) as Neanderthal. In Santa Luca's designation, the controversial Qafzeh and Skhul sites were firmly placed out of the Neanderthal range.
Neanderthals show a very distinctive craniofacial morphology relative to modern human populations. The Neanderthal face, in particular, is distinctly different from anything that came before or after. This fact has often been touted as evidence that the Neanderthals were a divergent outgroup that left no genetic heritage to modern human populations. The Neanderthal face is distinctive for its significant midfacial prognathism. Features involved in this prognathism include a very anterior midface, retreated zygomatics, anterior position of the dentition, and the anterior position of the nasal aperture. Evidence that the zygomatics have retreated is seen in that pre-Neanderthals have the M1 at the root of the zygomatic, with Neanderthals having the M3 at the root of the zygomatic, even though there has been some decrease in mandibular robusticity and postcanine tooth size. Some of the features that can be used to distinguish Neanderthals from modern humans (but not from earlier or conteporary populations) include:
- Lack of a canine fossa.
- Flat zygomatics.
- Rounded inferior orbital margins.
- The supraorbital torus projects at midline.
- Presence of a retromolar space (also distinguishes them from earlier hominids).
- Retreating mandibular symphysis (no mental eminence).
- A long low brain case.
- A suprainiac fossa.
- An occipital bun.
- A broad and projecting nose.
- Larger cranial capacity (but due to larger body size, Neanderthals are less encephalized than modern humans).
Another trait that is being looked at currently as a way of distinguishing Neanderthals in the inner ear morphology. Researchers like Hublin and Spoor are trying to determine if the Neanderthals had a unique inner ear morphology that can be used as a Neanderthal autapomorhpy. The diagrams below show the difference between modern humans, Neanderthals, and chimpanzees. When comparing the values of S/I, humans generally have a value close to 1, chimpanzees have values greater than 1, and Neanderthals have values less than 1. This is seen in the diagrams below:
The range of traits in Neanderthals is ver broad, and specimens can be looked at in three groups: early Neanderthals (approximately 250 kyr to 130 kyr), the Neanderthals that existed during the transition to the Upper Paleolithic (approximately 130 kyr to 45 kyr), and the late surviving Neanderthals ( after 45 kyr). The traits that make up each group will be discussed, along with this will be descriptions of some specific specimens.
Lower Paleolithic Neanderthals
As always in the anagenesis of one species into another, there is a "fuzzy" area which prevents the pinning down a a date when "A" became "B". In the case of heidelbergensis to neanderthalensis, this boundary seems to have the Neanderthals first appearing between 250 kyr to 200 kyr. The specimens that can be attributed as some of the earlier Neanderthals include material from
The material from
The site of
The material from Ehrinsdorf comes from the Fischer and Kämpfe quarries that are near the city. The material has been dated to 205 kyr by U-series dating and to 200 kyr by ESR dating, giving a fairly secure date for the material. The hominid remains consists of a neurocranium and partial endocast, an adult and an adolescent mandible, four parietal bones, a femoral shaft, as well as some associated teeth and postcrania. The Ehringsdorf H vault has been reconstructed at least three times since the material was discovered in 1925, by F. Weidenreich, O. Kleinschmidt, and
The material from
- A Neanderthal-like occipital bun.
- A suprainiac fossa.
- Very small mastoids which do not project below the cranial base.
- A circular cranial contour when seen from the rear.
- A thin cranial vault.
- A lateral maxillary incisor with large marginal ridges, a tubercle, and a well-developed labial crown convexity.
The material from the Bourdeois-Delaunay Cavern at La Chaise has been dated by thorium/uranium dating to approximately 151 kyr. The material from this site consists of a partial fragmentary cranium and mandible, and some other fragments and teeth. There are also specimens from the
The material from Fontéchevade was once a central piece of evidence to the discredited "pre-sapiens" theory of human evolution. The material from Fontéchevade consists of several cranial fragments from different individuals. Fontéchevade 1 was claimed to lack the thick browridges of Neanderthals by Vallois, but this specimen is that of a juvenile, and the brows would likely have thickened much in late adolescence and early adulthood. Also, Fontéchevade 2 was said to also show no strongly-developed browridge, but this specimen was missing nearly all of the face and browridge region. The specimen also shows a shallow sulcus and the back of the top of a supraorbital on the right lateral side, meaning it most likely had Neanderthal-like browridges. Fontéchevade has given as estimate of 1350 cc for brain size.
Early Upper Paleolithic Neanderthals
The next rough grouping of Neanderthal specimens contains those specimens that date to the Late Pleistocene, between approximately 130 kyr to 45 kyr. These include many of the "classic" Neanderthals, as well as many of the better known specimens. Sites which are included in this group are: Krapina, Saccopastore, Malarnaud, Altamura, Gánovce, Denisova, Okladnikov, Pech de l'Azé, Tabun, Kebara, Régourdou, Mt. Circeo, La Ferrassie, Combe Grenal, La Chapelle, Amud, Shanidar, Teshik-Tash, and Feldhofer.
The material from
- Low and broad cranial vaults.
- Broad, shallow temporal fossae with a low anterior articular face.
- Prominent continuous supraorbitals with large frontal sinuses restricted to them.
- Receding foreheads with frontal bossing.
- Columnar outer orbital margins in a lateral orientation.
- A broad upper nasal region lacking any depression at the nasal root.
- A concave, angled nasal profile.
The Saccopastore material from the Saccopastore quarry in
- A lesser magnitude of middle face projection.
- More anterior orientation and greater curvature for the cheeks.
- A shorter temporal fossae.
- Better development of the canine fossa.
- Smaller brain size.
- A more rounded occipital bone.
- A more flexed cranial base.
The material from the cave in Teshik-Tash, in the mountains of
- A large face and nasal region.
- A developing browridge.
- A receding forehead.
- A long cranium.
- It lacks a mental symphysis.
- A large cranial capacity of approximately 1500 cc.
The material from
The material from
The
An early Neanderthal find whose description led to several unfortunate outcomes is that of La Chapelle-aux-Saints. This specimen was discovered by monks during construction of an abbey in 1908, and the remains were sent to M. Boule, who proceeded to describe the specimen as the "typical" Neanderthal, thus leading to the description of "classic" Neanderthal features. However, Neanderthals are a highly variable species, and La Chapelle-aux-Saints shows many of the most extreme degree of the Neanderthal features. Also, Boule used the state of the fossil to construct an image of Neanderthals as ape-like primitive creatures, which could not possibly be related to humans. Beyond Boule's prejudices regarding the pattern of human evolution, the fact that the specimen was that of an arthritic "old man" (aged at between 40 and 50 years of age), which further skewed the description of Neanderthals as a species. This specimen has all the traits of the "classic" Neanderthal, for example, the large brain size (1625 cc), the heavy supraorbital torus, etc. This specimen is dated to approximately 50 kyr.
Another early discovery of Neanderthal remains came in 1909, when the remains from La Ferrassie were first uncovered. This site is dated to approximately 50 kyr, and has yielded the remains of seven individuals. The best known is La Ferrassie 1, the nearly complete skeleton of a middle-aged male Neanderthal, also discovered was a fairly complete adult female specimen, and five juvenile specimens ranging from prenatal to ten years of age. The remains seem to have been purposely buried. The La Ferrassie 1 specimen (along with La Chapelle-aux-Saints) founded the description of the "classic" Neanderthal anatomy, with such features as:
- A receding forehead.
- A long low cranial vault.
- The base of the skull is wider than the top of the skull.
- Prominent double-arched browridges.
- Midface prognathism.
- Retreated zygomatics.
- A weakly developed mental eminence.
- A retromolar space behind the M3.
- Large cranial capacity (over 1600 cc).
Late Surviving Neanderthals
The late surviving Neanderthal sites have started to become better known in recent years, and new discoveries and new dating have clouded the supposedly clear picture of replacement of Neanderthals by incoming H. sapiens populations. This is an exciting field of research at the moment, and only time will tell which evolutionary model os better born out by the evidence. Sites of late surviving Neanderthals are those that date to after 45 kyr, and especially those dating to after 35 kyr. Sites that are on that list include: La Quina, Zafarraya, Hortus, Vindija, Kulna, Šipka, Saint Césaire, Arcy-Sur-Cure, Bacho Kiro, El Castillo, Bñnolas, Devil's Tower, and Le Moustier.
The cave site at l'Horus in southern
The site of Vindija is becoming a very important site in the debate over assimilation versus replacement of the Neanderthals. There are several layers to the site, and the material in them dates from 42 kyr to as late as 32 kyr in the upper layers. The Vindija hominids have been put forth as a population that derives from both Neanderthal and incoming sapiens populations, or an influx of gene flow into the Neanderthal populations of
- A reduced midface prognathism and a less projecting upper face.
- Thinner and less projecting supraorbitals.
- A narrower nasal aperture, more prominent nasal spine, and a smaller facial height below it.
- A reduced breadth for the anterior teeth.
- The absence of occipital bunning.
- Thinner cranial bones.
- Definite presence of a mental eminence (though not prominent).
Some early criticisms attributed these modern affinites to the specimens being subadults or smaller than "normal" Neanderthals, but the remains have been well-reviewed, and seem to be full adults whose size compares favorably to earlier Neanderthal populations.
The most recent stratigraphic level from Vindija (G1) dates to approximately 32 kyr, and has been claimed to be late surviving Neanderthals with modern human affinities, as well as the remains of modern human beings, with no affiliation to the European Neanderthals. This question is directly involved with the split bone points that have been found in this level, since these have been identified as Aurignacian, which have generally been considered a H. sapiens industry. If the points are associated with the human remains at the site, it means that either the Aurignacian industry was transmitted to the Neanderthals from incoming humans, or the industry developed independently in both populations. The human remains from this level seem to definitely be of Neanderthal affinity, as seen when they are compared to the earlier remains from the G3 level.
The remains of the Saint-Césaire skeleton was discovered at a limestone cave at Pierrot's Rock, near
- Absence of a canine fossa.
- Presence of a retromolar space.
- Lack of a mental eminence.
Some consider the Saint Césaire skeleton as showing evidence of a transition of the Neanderthals to a more modern human-like form. Others claim that the differences between the sapiens populations in
- Central and lateral supraorbital thinning.
- A reduction in the thickness of the supraorbital torus.
- A reduction of midfacial prognathism.
- Reduced anterior tooth size.
- Smaller, lower appearing orbits.
- Smaller zygomatics.
- A markedly narrower nose.
- A flattened rear surface of the mastoid process.
The material from the Chatelperronian levels of the Grotte du Renne at
What may be the latest surviving Neanderthals currently known comes from the
- Small teeth with a very robust mandibular corpus.
- A vertical symphysis with a mental trigone.
- A slight mental eminence.
- A mental foramen positioned under the P4.
- A gap between the front of the ascending ramus and the back of the third molar.
Moderately large anterior teeth.
Conclusions
The eventual fate of the Neanderthals in the modern human phylogeny is still a much questioned issue, and a vigorously debated one. However, one thing is certain, the issue is not as cut and dry as many supporters of the Out of Africa II theory oftentimes claim. It seems highly unlikely that the Neanderthals contributed absolutely nothing to the modern genome, but whether they left a large heritage in modern humans or an insignificant one is a question that might not be answered satisfactorily for a long time.
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