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 Düsseldorf. The material recovered consisted of a skull cap, two femora, the three right arm bones, two of the left arm bones, part of the left ilium, and fragments of a scapula and ribs. These fossils were recovered by the quarry workers and set aside to be given to a local teacher and amateur naturalist, Johann Karl Fuhlrott. Fuhlrott suspected that these bones represented unique pieces of the human past, and left the description of the material to anatomist Hermann Schaaffhausen. The find was announced jointly in 1857, two years before Darwin’s On the Origin of Species.
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 Belgium was the first Neanderthal discovered, in 1829, and the second discovered was the Forbes Quarry find from Gibralter in 1848. The species name ofneanderthalensis comes from William King, who first named the species in 1863 at a meeting of the British Association, and put it into print in the Quarterly Journal of Science in 1864. This species is the focus of more argument among the academia of paleoanthropology than any other. Today, most researchers follow a multiregional view which has the European Neanderthals interbreeding and being absorbed by invading populations, or to have been marginalized by invading Homo sapiens until they died out, leaving no genetic legacy to modern humans. There are some (C.L. Brace most prominently) that think that the Neanderthals evolved in place into modern Europeans with little or no genetic influx from African populations, but few accept this argument.
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.
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 Pontnewydd Cave, Vértesszöllos, Ehringsdorf, Casal de’Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontéchevade, and possibly the very latest material from Atapuerca.
The material from Pontnewydd Cave, Wales, consists of small bits of a mandible and maxilla that may represent an eight-year-old child and an isolated molar from a near-adult or adult individual. This material has been dated by U-series dating, thermoluminescence (TL) dating, and by the associated faunal remains to approximately 251 kyr to 195 kyr. However, these dates are quite likely too old, as the material may have been washed into the cave at a later time. The features of the fragmentary remains that links them to Neanderthals include the dominant buccal cusp of the P3 and the taurodontism of the postcanine teeth.
The site of Vértesszöllos, Hungary, is also not very securely dated. The site was originally dated by thorium/uranium dating and association with microfauna to the 400 kyr range, but U-series dating has suggested a much younger date of 225 kyr to 185 kyr. However, the stratigraphy of the sites makes the dating of any of these methods with accuracy to the date of the skeletal material as unlikely, so the real date may be earlier or later. The remains of two individuals come from the site, a child approximately seven-years-old that is represented by fragments of a deciduous canine, a molar, and a permanent molar, and an adult that is represented by most of an occipital bone. The teeth are very similar to the Zhoukoudian material from China, and the cranial fragment has a Neanderthal-like occipital bun, but the bun is attained by cranial thickening, which is unlike the later Neanderthals. The cranial fragments gives an estimated cranial capacity of 1300 cc. The material has been attributed to erectus, heidelbergensis, and neanderthalensis by different researchers.
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 E. Vlcek. The most Neanderthal-like reconstruction was done by Kleinschmidt, and the least Neanderthal-like was done by Weidenreich. However, no matter which reconstruction is used, the material shows many traits linking it to later Neanderthals, not least of which is the large brain size (1450 cc for a female). Ehringsdorf is likely the earliest unquestionable Neanderthal site.
The material from Biache-Saint-Vaast, France, consists of two partial crania, a maxilla and cranial rear of a female individual, and parts of the splanchnocranium and cranial vault of a male. The site is dated to an estimated 196 kyr to 159 kyr. A brain size of 1200 cc. has been estimated for the female crania. While some features such as the small brain size seem to link these specimens to the Swanscombe cranium, it shows many more affinities with later Neanderthals, including:
- 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 Suard Cave at La Chaise that are uncertainly dated to the same time period. These remains consist of a large portion of a cranial vault, an occipital bone, a child’s mandible, as well as other fragments and associated teeth. The features that can be distinguished about the material seems to link these specimens most closely to the Biache hominids.
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 Krapina Cave, Croatia, was first discovered in 1899 by K. Gorjanovic-Kramberger. The material from this site is the largest sample of Neanderthals ever found, and is one of the largest fossil hominid samples ever found. There are more than 850 human fossils from more than eighty individuals at the site , most of whom died between the ages of sixteen and twenty-four years. This material has been dated to approximately 130 kyr. Many of the remains show evidence of butchery, and many of these remains may be the result of cannibalism, as also evidenced by the presence of concoidal scars, and the lack of carnivore tooth marks. While the crania that have been reconstructed show some differences from the classic Neanderthal anatomy, and from the Near East Neanderthals, several features clearly link the crania as Neanderthal, including:
- 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 Rome, Italy, consist of one fairly complete adult female cranium (Saccopastore 1), and a less complete male cranium (Saccopastore 2). These specimens were found at different times by different people, and have been dated to approximately 120 kyr, nearly contemporary with the Krapina specimens. The female specimen had given an estimated brain size of 1245 cc, and the male has been estimated at 1300 cc. While these specimens are undeniably Neanderthal, they do show some differences from the Neanderthals from more northenly sites such as La Ferrassie and Le Chapelle-aux-Saints in France. These differences from the classic Neanderthal anatomy include:
- 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 Uzbekistan, consists of the nearly complete cranium of an eight to nine-year-old child, likely male, as well as many postcranial elements, including one neck vertebra, several ribs, a humerus, the clavicles, a femur, a tibia, and one fibula. The site is very important in that it is likely one of the earliest instances of purposeful burial, and has been dated to approximately 70 kyr. The skeleton was surrounded by what seems to be purposefully arranged Siberian ibex (a wild goat) skulls. Also, these faunal remains may be indicative of purposeful hunting, as there are few carnivore bones, and the many goat bones show little to no signs of carnivore activity. While this specimen is that of a young juvenile, he had developed several traits that clearly link it to European Neanderthals. These features include:
- 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 Kebara Cave, Israel, was discovered in several phases of excavations. The first hominid found was the fragmentary remains of an infant (Kebara 1), and in 1983, the better known Kebara 2 specimen was recovered. This material has been dated by both thermoluminescence and ESR dating to approximately 60 kyr. The Kebara 2 specimen consists of the upper part of a male postcranial skeleton, as well as a mandible, a hyoid, and a single M3. The Kebara 2 specimen is one of the few complete pelves known between the australopithecines and modern humans, however, as with just about every complete fossil pelvis that is currently known, it is from a male individual. The Kebara 2 individual is estimated to have stood about 173 cm tall, and to have weight approximately 76 kg, significantly smaller than estimates for European Neanderthal specimens of similar height, such as La Ferrassie. The recovery of the hyoid bone was a very important find, as it disproved earlier theories that Neanderthals and other pre-sapiens hominids were incapable of speech due to the lack of a hyoid. Neanderthals did possess a hyoid, and this scenario of speechlessness based on this bone was discredited. The individual was between the ages of 25 and 35 when he died, and he can be securely attributed to Neanderthals based on the several features, including the presence of a retromolar space, an the lack of a mental eminence.
The material from Amud Cave, Israel, consist of the fragmentary remains of a male, along with 15 other specimens, two-thirds of which are children or infants. This material has been dated by thermoluminescence to approximately 45 kyr to 47 kyr. Amud 1 is the most complete specimen, consisting of just about the entire cranium, with most of the postcrania present. This individual was a male, died in his mid 20s, was approximately 174 cm tall, and has the largest cranial capacity of any Levant hominid at 1740 cc. While these remains are clearly Neanderthal, these specimens show more affinity to the Shanidar Neanderthals rather than the “classic” European Neanderthals. For example, when compared to the European Neanderthals and the Skhul H. sapiens, the Amud material actually shows greater affinity to the sapiens of Skhul than to the European Neanderthals.
The Shanidar Cave, in the Zagros Mountains of Iraq, is also the site of a well-preserved Neanderthal sample. The sample consists of two groups from two different time periods, the earlier group has been dated to approximately 60 kyr, and a later group that has been estimated to date to approximately 46 kyr. The two samples consist of the remains of nine individuals, two children and seven adults. The Shanidar specimens are the focus of some debate over the possibility that flowers were laid over the grave. Many researchers hold doubts to the validity of this claim, saying that the researchers that led the excavation might have carried pollen in accidentally with them. However, D. Frayer has pointed out that the pollen was from late spring/early summer flowers, and the excavation was conducted in August, so the debate is likely to rage on. The early group consists of Shanidar 2, 4, 6, 7, 8, 9, and the later group consists of Shanidar 1, 3, and 5.
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 France has yielded about 50 hominid fragments which date to the latest part of Würm II. Most of the remains are pieces of jawas and teeth, but there are some cranial fragments and postcranial remains mixed in as well. These remains have been claimed as evidence that the late Neanderthals were not evolving towards the modern human condition, but have also been used to claim that they were evolving in the modern human condition. What can be said is the remains show several Neanderthal features (e.g., a poorly developed mental eminence), but also show soe reduction in the anterior dentition relative to earlier Neanderthals. The material has been interpreted as the remains left in a hyena den.
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 Europe. The remains from the earlier level (G3) date to approximately 42 kyr, and are contemporary with the l’Horus remains. In support of the assimilation theory is evidence that these Neanderthals were evolving in a way that was leading them to the modern human condition. For example:
- 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 Charente-Maritime, France, in 1979. The remains consist of the right half of a skull, some ribs, a shoulder blade, two arm bones, and fragments of a kneecap and the tibias which have been dated to approximately 36 kyr. At one time, it was considered the last surviving Neanderthal. The specimen shows many definitive Neanderthal traits, such as:
- 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 Europe and the late surviving Neanderthals is so great to preclude the possibility of an ancestor/descendent relationship since the timeframe is so short. Some of the features that have been claimed to show affinities to modern Europeans include:
- 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 Arcy-sur-Cure, France, has been dated to approximately 34 kyr. This material consists of isolated teeth with definite Neanderthal like affinities, and a child’s temporal fragment whose inner ear morphology links it to Neanderthals rather than modern humans. The importance of Arcy-sur-Cure is in the association of these remains with personal adornment articles that were similar to those from Aurignacian sites nearby. This is yet another link between the Neanderthals and the Aurignacian industry that discredits the idea that the Neanderthals were not associated with that particular industry.
What may be the latest surviving Neanderthals currently known comes from the Zafarraya Cave, near Andalucia, Spain. The upper levels of this site have been thorium/uranium and C14 to approximately 27 kyr, and the sequence may extend as late as 25 kyr. Material from the site include a nearly complete mandible, a mandible that has been burnt, a partial Os coxa, a femur, and an upper incisor. The femur shows affinities with both Neanderthals (thick cortical bone, circular midshaft cross-section), but also shows affinities to later Europeans (development of a pilaster, thin femoral neck dimensions, shaft expanded anteriorly-posteriorly). The complete mandible more closely links this material to Neanderthals, with features such as:
- 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.
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.
Aiello, L., and C. Dean. 1990. An Introduction to Human Evolutionary Anatomy. London: Academic Press.
ApSimon, A.M. 1980. “The last Neanderthal in France?” In Nature, vol. 287, pp. 271–271.
Aresburg, B., O. Bar Yosef, M. Chech, P. Goldberg, H. Laville, L. Meignen, Y. Rak, E. Tchernov, A.M. Tillier, and B. Vandermeersch. 1985. “Une sépulture néanderthalien dans la grotto de Kebara (Israel).” In Comptes Rendus des Séances de l’Academie des Sciences, Paris, vol. 300, pp. 227–230.
Bar-Yosef, O., B. Vandermeersch, B. Arensburg, A. Belfer-Cohen, P. Goldberg, H. Laville, L. Meignen, Y. Rak, J.D. Speth, E. Tchernov, A.M. Tillier, and S. Weiner. 1992. “The Excavations in Kebara Cave, Mt. Carmel.” In Current Anthropology, vol. 33, no. 5, pp. 497–550.
Boule, M. 1908. “L’Homme fossile de La Chapelle-aux-Saints (Covèze). In Comptes Rendus des Séances de l’Academie des Sciences, Paris, vol. 147, pp. 1349–1352.
Boule, M. 1911–1913. “L’homme fossile de La Chapelle-aux-Saints.” In Annales de Paleontologie, vol. 6–8.
Capitan, L., and D. Peyrony. 1909. “Deux squelettes humains au milieu de foyers de l’époque moustérienne.” In Revue d’Anthropologie, vol. 19, pp. 402–409.
Gorjanovic-Kramberger, K. 1906. Der diluvale Mensch von Krapina in Kroatien. Wiesbaden: C.W. Kreidels Verlag.
Gremyatsky, M.A., and M.F. Nestourkh, eds. 1949. Teshik-Tash: Paleolithic Man. Moscow: Moscow University Publishers.
Heim, J.-L. 1976. “Les hommes fossiles de la Ferassie I.” In Archives de l’Institut de Palèontologie Humaine, vol. 35, pp. 1–331.
Hublin, J.-J., C.B. Ruiz, P.M. Lara, M. Fontugne, and J.-J. Reyss. 1995. “The Mousterian site of Zafarraya (Andalucia, Spain): Dating and implications on the Paleolithic peopling process of Western Europe.” In Comptes Rendus des Séances de l’Academie des Sciences, vol. 321, pp. 931–937.
Johanson, D., and B. Edgar. 1996. From Lucy to Language. New York: Simon and Schuster Editions.
Jones, S., R. Martin, and D. Pilbeam, eds. 1992. The Cambridge Encyclopedia of Human Evolution. Cambridge: Cambridge University Press.
Keith, A. 1911. “The early history of the Gibralter cranium.” In Nature, vol. 87, pp. 313–314.
King, W. 1864. “The reputed fossil man of the Neanderthal.” In Quarterly Review of Science, vol. 1, pp. 88–97.
Larsen, C.S., R.M. Matter, and D.L. Gebo. 1991. Human Origins: The Fossil Record. Prospect Heights: Waveland Press, Inc.
Lévéque, F., and B. Vandermeersch. 1980. “Les découvertes de restes humains dans un horizon castelperronien de Saint-Césaire (Charente-Maritime).” In Bulliten de la Société de Prehistorique Francaise, vol. 77, pp. 35.
Manzi, G., and P. Passarello. 1991. “Antènèandertaliens et Nèandertaliens du Latium (Italie Centrale).” In L’Anthropologie, vol. 95, no. 2/3, pp. 501–522.
Mardis, S.E. 1995. “The Last Neanderthals.” In Archaeology, vol. 48, no. 6, pp. 12–13.
Mercier, N., H. Valladas, J.-L. Joron, J.-L. Reyss, F. Lévéque, and B. Vandermeersch. 1991. “Thermoluminescence dating of the late Neanderthal remains from Saint-Cèsaire.” In Nature, vol. 351, pp. 737–739.
Movius, H.L., Jr. 1953. “The Mousterian Cave of Teshik-Tash, Southeastern Uzbekistan, Central Asia.” In American School of Prehistoric Research Bulletin, vol. 17, pp. 11–71.
Puech, P.-F. 1981. “Tooth-wear in La Ferassie man.” In Current Anthropology, vol. 22, no. 4, pp. 424–430.
Rak, Y., W.H. Kimbel, and E. Hovers. 1994. “A Neandertal infant from Amud Cave, Israel.” In Journal of Human Evolution, vol. 26: pp. 313–324.
Santa Luca, A.P. 1978. “A re-examination of presumed Neandertal-like fossils.” In Journal of Human Evolution, vol. 7, pp. 169–636.
Sergi, S. 1929. “La scoperta di un cranio del tipo di Neanderthal presso Roma.” In Rivista di Antropologia, vol. 28, pp. 457–462.
Smith, F.H. 1976. The Neandertal Remains from Krapina: A Descriptive and Comparative Study. University of Tennessee Department of Anthropology Rep. Invest., vo. 15, pp. 1–359.
Smith, F.H. 1991. “The Neanderthals: Evolutionary dead ends or ancestors of modern people?” In Journal of Anthropological Research, vol. 47, no. 2, pp. 219–238.
Straus, W.L., Jr., and A.J.E. Cave. 1957. “Pathology and the posture of Neanderthal man.” In Quarterly Review of Biology, vol. 32, no. 4, pp. 348–363.
Stringer, C.B., and C. Gamble. 1993. In Search of the Neanderthals: Solving the Puzzle of Human Origins. New York: Thames and Hudson.
Stringer, C.B., and R. Grün. 1991. “Time for the last Neanderthals.” In Nature, vol. 351, pp. 701–702.
Suzuki, H. and F. Takai, eds. 1970. The Amud Man and His Cave Site. Tokyo: University of Yokyo Press.
Trinkaus, E. 1973. “A reconsideration of the Fontéchevade fossils.” In American Journal of Physical Anthropology, vol. 39, pp. 25–35.
Trinkaus, E. 1987. “The Neanderthal face: Evolutionary and functional perspectives on a recent hominid face.” In Journal of Human Evolution, vol. 16, no. 5, pp. 429–443.
Trinkaus, E., and F.H. Smith. 1994. “Body size of the Vindija Neanderthals.” In Journal of Human Evolution, vol. 28, no. 2, pp. 201–208.
Wolpoff, M. 1999. Paleoanthropology. Second edition. Boston: McGraw-Hill.