Neanderthals and Modern Humans -- A Regional Guide
Neanderthals and Modern Humans in Western Asia
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Genetic Evidence

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The differing physical traits of the peoples of Africa, Europe, and Asia blend in the melting pot of Western Asia. Such mixing was also prevalent at the time of the Neanderthals and first modern humans. Though it is unclear exactly how European Neanderthals, early modern Africans, and indigenous peoples interacted in Western Asia during the time when modern humans were evolving, this region was clearly pivotal. It would have served as a migration route both for modern humans leaving Africa and for Neanderthals exiting Europe. It also has yielded some of the earliest remains of anatomically modern humans -- as early, or maybe even earlier, than those found in sub-Saharan Africa.

Partial skull from Zuttiyeh
Did this skull from Zuttiyeh, Israel, belong to an ancestor of Neanderthals, early modern humans, both, or neither?
(Photo depicts front part of skull's right side.)

(Courtesy of the Israel Antiquities Authority, Jerusalem, and Eric Delson. Reproduction or reposting to another web site prohibited by the owner. Photography by Chester Tarka [AMNH].)

The puzzle over the relationship between Neanderthals and early modern humans in Western Asia begins with a skull from the Zuttiyeh site in Israel, from a period known as the Middle Paleolithic. The Zuttiyeh skull was associated with an early Middle Paleolithic industry of the Levant region, or eastern Mediterranean, called the Acheulo-Yabrudian. This industry at Zuttiyeh has been dated to as late as 148,000 years before the present (B.P.) (Bar-Yosef, 1998: table 1), but other estimates place the Zuttiyeh skull as early as 200,000 to 250,000 B.P. (Zeitoun, 2001: 522).

One study has grouped the Zuttiyeh skull with early modern humans (Zeitoun, 2001), but most scientists identify it as archaic. Some think Zuttiyeh was ancestral to the first modern humans of the Levant (Vandermeersch, 1989). Others believe it cannot be clearly linked to any of the later populations of the Levant, either modern or Neanderthal (Stringer and Gamble, 1993: 96). Still others doubt that separate "modern" and "Neanderthal" populations ever existed in this region. In their view, none of the alleged Neanderthals from the Levant had a true Neanderthal morphology as seen in Europe. They see Zuttiyeh as an early member of a single, diverse population that included both the alleged "moderns" and the so-called "Neanderthals" of the region (Arensburg and Belfer-Cohen, 1998: 318-319).

The Acheulo-Yabrudian was followed by the Mousterian, a Middle Paleolithic industry associated in the Levant with both modern and "Neanderthal" remains. Mousterian layers at the Tabun site in Israel yielded a partial skeleton, a mandible (lower jaw), an isolated premolar tooth, and five isolated limb bones. The mandible, the tooth, and at least some of the limb bones clearly derive from layer C at the site (Kaufman, 2001: 227; Quam and Smith, 1998). The partial skeleton was reportedly excavated from layer C, and has therefore been labelled Tabun C1, but new dating analyses support a long held suspicion that it actually derives from the more recent layer B at Tabun (Grün and Stringer, 2000).

Tabun C1 Neanderthal burial
The Tabun site in Israel yielded this burial of a Neanderthal, Tabun C1.

(From: Garrod and Bate [1937] The Stone Age of Mount Carmel. © Oxford University Press. By permission of Oxford University Press.)

Most scholars identify Tabun C1 as Neanderthal. The mandible from layer C, labelled Tabun C2, appears more modern, however, than either this skeleton or the limb bones from the site. Some researchers call this mandible "modern-like" (Quam and Smith, 1998), although others see it as archaic (Stefan and Trinkaus, 1998). Modern traits are not apparent on the five limb bones from the site. In fact, a part of a right radius, or lower arm bone (Tabun C4), and a right wrist bone (Tabun C5) may have originally belonged to the Tabun C1 Neanderthal skeleton. Also, a piece of a right femur, or thigh bone (Tabun C3), is similar to the right femur of Tabun C1 (Quam and Smith, 1998: 409). The isolated premolar from layer C is also thought to be from a Neanderthal (Kaufman, 2001: 227).

Dates on animal teeth from Tabun layer B, from which the Tabun C1 skeleton derives, range from 50,690 B.P. with the U-series method (McDermott, et al, 1993: table 1) to 122,000 B.P. with the electron spin resonance (ESR) technique (Grün and Stringer, 2000: table 1). Previous dates younger than 40,000 B.P. for Tabun C1 (Schwarcz, et al, 1998) are now thought to be in error. A more recent analysis places Tabun C1 close to the most ancient age estimate for layer B, around 122,000 B.P. (Grün and Stringer, 2000: 610). This same analysis places layer C at Tabun as far back as 140,000 B.P. Other studies bracket layer C between 97,840 B.P. with the U-series technique (McDermott, et al, 1993: table 1) and 171,000 B.P. with the thermoluminescence (TL) method (Mercier, et al, 1995; Valladas, et al, 1998).

These dates indicate that the Neanderthal and possibly modern remains from layer C,
Neanderthal skull from Amud
Most Neanderthal remains from the Levant, such as this skull from Amud, Israel, are more recent than the earliest modern human remains from the region.

(Courtesy of the Israel Antiquities Authority, Jerusalem, and Eric Delson. Reproduction or reposting to another web site prohibited by the owner. Photography by Chester Tarka [AMNH].)

and also the Tabun C1 Neanderthal, are as old as, or perhaps even older than, early modern humans from other sites in Israel and from Africa, dated to 80-120,000 B.P. If Neanderthal and modern types were both at Tabun before 100,000 B.P., this could back the idea that the Neanderthals and early modern humans of the Levant were a single indigenous population evolving toward a fully modern form (Arensburg and Belfer-Cohen, 1998). Or Neanderthals from the north may simply have moved into the Levant at this early time and coexisted there with a separate modern human population (Mercier, et al, 1995: 507; Vandermeersch, 1989), which may or may not have evolved locally.

With the exception of the Tabun remains, however, all other alleged Neanderthals from the Levant, at the sites of Amud and Kebara in Israel, and Dederiyeh in Syria, are associated with industries like those from layer B at Tabun and are thought to postdate 75,000 B.P. (Bar-Yosef, 1996; Lieberman, 1998). Some researchers therefore believe that the principal migration of Neanderthals into the Near East took place at around this time (Bar-Yosef, 1996), some 10,000 years or more after modern humans had appeared in the Levant.

Although some doubt exists as to whether the Tabun C2 mandible should really be called "modern," researchers are more comfortable applying that title to remains from the sites of Skhul and Qafzeh in Israel. The bones of at least 10 people were uncovered in Mousterian layer B at Skhul, and the remains of at least 13 individuals were found in Mousterian layers XV to XXII at Qafzeh. Many of the remains showed signs of deliberate burial (Defleur, 1993).

Layer B at Skhul has been dated to an average of 81-101,000 B.P. with the ESR method and to an average of 119,000 B.P. with the TL method (Bar-Yosef, 1998: table 1; Valladas, et al, 1998). U-series dates for this layer indicate that some of the Skhul remains may be considerably younger than others (McDermott, et al, 1993), but the younger dates may be unreliable (Bar-Yosef and Pilbeam, 1993). The various layers containing the Qafzeh remains were laid down relatively rapidly. Their dates average 96-115,000 B.P. with the ESR method and 92,000 B.P. with the TL method (Bar-Yosef, 1998: table 1).

Qafzeh 6 skull
Scholars debate whether the early modern humans of the Levant, represented here by this skull from Qafzeh, evolved locally or moved into the region from Africa.

(Courtesy of The Natural History Museum, London.)

Scientists differ on the origin of the Skhul/Qafzeh people. Some see them as members of an early modern stock that evolved within the Levant (Vandermeersch, 1989). Others believe they migrated into the Levant from Africa and were related to the early modern humans of that continent, despite certain physical differences between them and early modern Africans (Rightmire, 2001: 235). Unfortunately, archeological evidence of a link between the early modern humans of the Levant and their contemporaries in adjacent territories of Africa cannot as yet be demonstrated due to methodological problems that hinder comparisons between the two regions (Vermeersch, 2001).

The view that the first modern humans of the Levant evolved locally will gain support if the older age estimates for Tabun layer C, as old as 171,000 B.P., prove correct. Such an early date would seemingly make any modern humans who occupied Tabun during layer-C times too ancient to derive from Africa, where the earliest modern humans appear to be only about 100,000 years old (Rightmire, 2001). Revisions of dates from one South African site, however, may push the age of the first modern Africans back to 170,000 B.P. (Grün and Beaumont, 2001: 480), contemporaneous with the oldest dates proposed for the Tabun C remains.

Some scholars also propose ties between the Skhul/Qafzeh people and the Levantine Neanderthals, since the physical features of these two groups overlap. One theory attributes these shared traits to interbreeding between indigenous occupants of the Levant, early modern humans moving into the region from Africa, and Neanderthals coming in from Europe (Kramer, et al, 2001). Still, the apparent persistence of Neanderthal-like and modern-like morphologies in the region suggests that the mixing of these populations was not total. Other scholars, however, deny the existence of separate, interbreeding groups and place the Skhul/Qafzeh people and Levantine Neanderthals into a single, diverse lineage whose traits they believe were shaped by gene flow from nearby regions, particulary Africa (Arensburg and Belfer-Cohen, 1998).

In contrast to these ideas, some scientists insist that the Skhul/Qafzeh people and the Levantine Neanderthals differed enough from each other to have comprised separate and distinct biological taxa (Lieberman, 1998: 272-273). According to this view, interbreeding between these two groups would have been limited or nonexistent. In fact, some researchers believe the differences between early modern humans and Neanderthals, both in the Levant and elsewhere, were so great that they probably belonged to separate species (Stringer, 1998: 30, 33).
Skhul 5 skull
The robust features of the Skhul 5 male may indicate this early modern human had Neanderthals in the family, or his lineage may simply have retained primitive hominid traits also possessed by Neanderthals.

(Courtesy of David Pilbeam and the Peabody Museum, Harvard University, Cambridge, Mass.; and Eric Delson. Reproduction or reposting to another web site prohibited by the owner. Photography by Chester Tarka [AMNH].)

Support for this view comes from the possibility that the features shared by the Skhul/Qafzeh people and the Levantine Neanderthals might be retentions of archaic traits, not derived features shared through interbreeding (Lahr, 1996: 25; Pearson, 2000b: box 2).

If Neanderthals and early modern humans were separate species, most of their differences should have been genetically based. But some of these differences may have resulted not from genes, but from contrasting behaviors, which placed different stresses on the skeleton during growth. Bones alter their characteristics in response to such stresses, without genes necessarily being involved. The thickness and cross-sectional shape of long bones are particularly malleable under this type of stress, and Neanderthals and early modern humans did differ in these traits (Pearson, 2000b: 239-243). But Neanderthals diverged from modern humans in many other ways too. Many scientists find this plethora of contrasts hard to explain without invoking genetic factors (Pearson, 2000b: 238; Stringer, 1998: 30).

The view that the differences between Neanderthals and early modern humans had a strong genetic basis finds support in the fact that many of these differences appeared early in life. This is particularly true for features of the skull. One study compared computer reconstructions of entire skulls of Neanderthals from the Levant and elsewhere with those of early modern humans from Skhul and Qafzeh, from individuals representing various life stages (Ponce de León and Zollikofer, 2001). The comparisons revealed that the features of Neanderthals and early modern humans had already diverged by about age 2˝, suggesting that the initial stages of this divergence began even earlier, perhaps in the womb.

The appearance so early in life of traits distinguishing Neanderthals from early modern humans strongly suggests that the differences between these two groups were primarily determined by genes, not by differences in lifestyle. This strengthens the view that Neanderthals and early modern humans were separate species. On the other hand, the youngest Neanderthals in the skull study, younger than age 3, were not compared to early modern humans, only to recent humans. Since recent humans are less Neanderthal-like than early modern humans, the degree of difference seen in the study between Neanderthal and recent human infants may be greater than what would be observed between Neanderthal and early modern human infants.

While studies of skulls indicate a divergence of Neanderthal and modern human anatomy before age 3, those involving other bones of the skeleton often fail to show such early contrasts. One study, for example, found that the leg bones of two Neanderthal children from Dederiyeh, Syria, both about age 2, overlap in their traits with the bones of early modern children from Skhul, Qafzeh, and other sites (Kondo and Ishida, 2001). Another has found the hip bones of early modern children from Qafzeh to be similar to those of Neanderthal children from France (Majo and Tillier, 2001). And a study of the upper arm and lower leg bones of a Neanderthal infant from
skeleton of the Dederiyeh 1 Neanderthal child
Syria's Dederiyeh cave yielded these remains of a 2-year-old Neanderthal, and also the bones of another Neanderthal child the same age. The skeletal traits of these children show that at this young age, Neanderthals differed less from early modern humans than they did later in life.

(Courtesy of Mikio Terasawa, Nihon University, Tokyo, Japan; and the Kimura, Mohri and Suzuki Laboratory, University of Tokyo, Japan.)

the Amud site in Israel found its bones to be similar to those of recent infants (Odwak, 2001).

These similarities between young Neanderthals and modern humans indicate that some of the features that distinguished Neanderthals from early modern humans in adulthood may have resulted from behaviors that differentially altered skeletal traits during growth, not from genetic differences. This increases the chances that these two groups belonged to a single species. Unfortunately, in the Levant at least, it has proven difficult to determine how the lifestyles of these two populations would have diverged, since they were using the same Middle Paleolithic technology. One possibility, however, is that behavioral dissimilarities that are archeologically invisible, such as differences in the contribution of children to subsistence tasks, may have produced the physical contrasts between these two groups (Kaufman, 2001: 228-231).

Some of the skeletal traits that distinguish the Neanderthals and early modern humans of the Levant support the view that these two groups behaved differently. A study of features on hand bones resulting from habitual activities indicates that the Skhul/Qafzeh people more frequently employed oblique grips and used finer finger movements than did Neanderthals, perhaps reflecting a greater use of hafted tools and more engraving and incising activities -- that is, more "modern" behavior -- by the Skhul/Qafzeh people (Niewoehner, 2001). These apparent differences support the idea that differing lifestyles, not genetics, produced at least some of the distinctions between the Neanderthals and early modern humans of the Levant, strengthening the possibility that these two groups belonged to the same species.

Moving past the species issue, the question that remains is what relationship the early modern humans from Skhul and Qafzeh may have had to the first modern peoples of greater Eurasia. The Skhul/Qafzeh people themselves are unlikely to have been the population of modern humans that colonized Europe. Their physical traits in many ways differed from those of the first modern Europeans (Pearson, 2000b: 243; Wolpoff, 1999: 607), and they lived many thousands of years before those Europeans.

But is it possible that a later population of modern humans in the Near East moved into Europe? A gap of more than 40,000 years exists between the Middle Paleolithic remains from Skhul and Qafzeh, which are more than 80,000 years old, and the next modern remains from the Levant, from an Upper Paleolithic level at the site of Ksar Akil in Lebanon, which is somewhat older than 32,000 B.P. (Wolpoff, 1999: 742-743). One cannot assume, however, that the Ksar Akil individuals were ancestral to the first modern Europeans, since they date to essentially the same time period as those Europeans.

Although modern humans from Western Asia dating to the time between Skhul/Qafzeh and Ksar Akil could have spawned the first modern humans of Europe, it is not clear that any such people existed. The Western Asian remains from this intervening period are all Neanderthal-like. In fact,
Qafzeh 9 skull
The anatomically modern remains unearthed in Israel's Qafzeh cave, such as the Qafzeh 9 skull shown here, predate by tens of thousands of years the earliest modern humans of Europe.

(Courtesy of The Natural History Museum, London.)

some evidence points to an increasing expression of Neanderthal traits in Western Asia during this time, as can be seen, for example, among Neanderthal remains from the site of Shanidar in Iraq (Wolpoff, 1999: 619-622).

One type of evidence often cited in support of the view that the early modern humans of Europe came from the Near East, and ultimately from Africa, is the fact that they had a gracile body build, which resembled the linear physique of the Skhul/Qafzeh people. This physique is typical of people living in warm regions of the earth, both today and in the past. In contrast, Neanderthals had a short, stocky build like that of present-day inhabitants of cold regions (Pearson, 2000b: 240-241). Some scientists therefore believe that the transition to a less robust, more linear physique in Europe at the time of the first modern humans must have resulted from the migration into that continent of a warm-adapted population of early modern humans (Pearson, 2000a), most likely from the Near East.

Other researchers, however, suggest that behavioral changes could have caused cold-adapted European Neanderthals to rapidly evolve a linear physique (Wolpoff, 1999: 772-774). The advances of the Upper Paleolithic included better ways of creating and maintaining campfires and warmer types of clothing. These could have rendered obsolete the cold-adapted body build of the Neanderthals. Other advances of this period could have lessened the mechanical demands placed on the human body, decreasing the need for a robust physique. Lifestyle changes during the Upper Paleolithic may also have led people to become more mobile, causing them to evolve longer, more gracile lower limbs. In fact, the shape of the thigh bone in early modern humans does indicate they were highly mobile (Pearson, 2000b: 241).

Another possibility is that improvements in diet resulting from better hunting and gathering techniques during the Upper Paleolithic caused Neanderthals to rapidly alter their body proportions. Improvements in nutrition will enable people to grow taller, which in turn gives them a more linear body build. Advances in food acquisition and the resulting boost in nutrition might also have allowed Neanderthals to spend less time and energy in the food quest. This drop in activity levels would then have reduced the need to maintain a robust skeleton (Cachel, 1997).

Although doubts exist as to whether the gracile body build of the first modern Europeans reflects a migration into Europe of warm-adapted modern humans from the Near East, some archeological evidence may support such a migration. Upper Paleolithic industries thought to be the work of modern humans, for example, seem to have appeared somewhat earlier in Western Asia than in Europe. The earliest dates are 46-47,000 B.P., from the site of Boker Tachtit in Israel (Bar-Yosef, 1996: 182).

Also, some of the earliest Upper Paleolithic industries from Central and Eastern Europe (see Central and Eastern Europe web page) show certain resemblances to the early Upper Paleolithic industries of Western Asia. The industry from Boker Tachtit, for example, resembles the Bohunician industry of Central Europe and a similar Eastern European industry from the Korolevo II site in Ukraine (Tostevin, 2000), both of which first appeared about 40,000 years ago, if not earlier (Kozlowski, 1996: 207). Similarly, an early Upper Paleolithic industry from the Zagros Mountains of Iran and northern Iraq, the Baradostian, resembles the Bachokirian industry of the Balkans (Kozlowski, 1996: 208). Both of these industries appear to have first arisen before 40,000 B.P.

These facts raise the possibility that early modern Near Easterners producing the Boker Tachtit and Baradostian industries moved into Europe and spawned the Bachokirian and Bohunician-like industries of Central and Eastern Europe. On the other hand, the general contemporaneity of all these industries may indicate they all arose independently. The possibility of independent evolution is threatened, however, by the fact that these industries reportedly appear more similar to each other than to any of the Middle Paleolithic industries that predated them in their respective regions (Tostevin, 2000: 101-104). Still, even if Near Easterners did bring these early Upper Paleolithic technologies into Europe, these people may not have been modern humans. Considering the early dates involved, they could have been Neanderthals.

Other researchers have argued that the earliest Upper Paleolithic artifacts of Western Asia may in fact have been made by Neanderthals but that modern humans making a later Upper Paleolithic industry called the Aurignacian moved into Europe and replaced the Neanderthals living there (Zilhao and d'Errico, 1999: 45-54). But again, a problem arises in that the earliest Aurignacian in Europe (see web pages for Western and Central and Eastern Europe) is as old, or perhaps even older, than that in Western Asia, where the Aurignacian first occurs at the Kebara site in Israel, at 35-36,000 B.P. Unless researchers can more clearly link the early Upper Paleolithic industries of Europe to an earlier source in Western Asia, and unless that source can be firmly tied to modern humans, the case for a Near Eastern origin for the first modern Europeans will remain problematic.

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References:

Arensburg, Baruch and Anna Belfer-Cohen (1998) "Sapiens and Neandertals: Rethinking the Levantine Middle Paleolithic hominids." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 311-322.

Bar-Yosef, Ofer (1996) "Modern humans, Neanderthals and the Middle/Upper Paleolithic transition in Western Asia." In O. Bar-Yosef, L.L. Cavalli-Sforza, R.J. March, and M. Piperno, eds. The Lower and Middle Palaeolithic (Colloquia of the XIII International Congress of Prehistoric and Protohistoric Sciences, no. 5). Forlě, Italy: ABACO Edizioni. pp. 175-190.

Bar-Yosef, Ofer (1998) "The chronology of the Middle Paleolithic of the Levant." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 39-56.

Bar-Yosef, Ofer and David Pilbeam (1993) "Dating hominid remains." Nature 366: 415.

Cachel, Susan (1997) "Dietary shifts and the European Upper Palaeolithic transition." Current Anthropology 38: 579-603.

Defleur, Alban (1993) Les sépultures moustériennes. Paris: CNRS Editions.

Grün, Rainer and Peter Beaumont (2001) "Border Cave revisited: A revised ESR chronology." Journal of Human Evolution 40: 467-482.

Grün, Rainer and Chris Stringer (2000) "Tabun revisited: Revised ESR chronology and new ESR and U-series analyses of dental material from Tabun C1." Journal of Human Evolution 39: 601-612.

Kaufman, Daniel (2001) "Comparisons and the case for interaction among Neanderthals and early modern humans in the Levant." Oxford Journal of Archaeology 20: 219-240.

Kondo, O. and H. Ishida (2001) "Ontogenetic variation in the Dederiyeh Neandertal children: Postcranial evidence" (Abstract of paper presented at the 70th annual meeting of the American Association of Physical Anthropologists, Kansas City, Missouri, March 28-31, 2001). American Journal of Physical Anthropology 114(S32): 92.

Kozlowski, Janusz K. (1996) "Cultural context of the last Neanderthals and early modern humans in central-eastern Europe." In O. Bar-Yosef, L.L. Cavalli-Sforza, R.J. March, and M. Piperno, eds. The Lower and Middle Palaeolithic (Colloquia of the XIII International Congress of Prehistoric and Protohistoric Sciences, no. 5). Forlě, Italy: ABACO Edizioni. pp. 205-218.

Kramer, Andrew, Tracey L. Crummett, and Milford H. Wolpoff (2001) "Out of Africa and into the Levant: Replacement or admixture in Western Asia?" Quaternary International 75: 51-63.

Lahr, Marta Mirazón (1996) The evolution of modern human diversity: A study of cranial variation. Cambridge: Cambridge University Press.

Lieberman, Daniel E. (1998) "Neandertal and early modern human mobility patterns: Comparing archaeological and anatomical evidence." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 263-275.

Majo, T. and A.M. Tillier (2001) "Patterns of growth in Upper Pleistocene Homo sapiens: Comparative analysis between La Ferrassie and Qafzeh immature os coxae" (Abstract of paper presented at the 70th annual meeting of the American Association of Physical Anthropologists, Kansas City, Missouri, March 28-31, 2001). American Journal of Physical Anthropology 114(S32): 100-101.

McDermott, F., R. Grün, C.B. Stringer, and C.J. Hawkesworth (1993) "Mass-spectrometric U-series dates for Israeli Neanderthal/early modern hominid sites." Nature 363: 252-255.

Mercier, N., H. Valladas, G. Valladas, and J.-L. Reyss (1995) "TL dates of burnt flints from Jelinek's excavations at Tabun and their implications." Journal of Archaeological Science 22: 495-509.

Niewoehner, Wesley A. (2001) "Behavioral inferences from the Skhul/Qafzeh early modern human hand remains." Proceedings of the National Academy of Sciences 98: 2979-2984. (FULL TEXT ONLINE)

Odwak, H. (2001) "Diaphyseal cross-sectional shape in the Amud 7 Neandertal and modern human babies and infants." (Abstract of paper presented at the 70th annual meeting of the American Association of Physical Anthropologists, Kansas City, Missouri, March 28-31, 2001). American Journal of Physical Anthropology 114(S32): 114-115.

Pearson, Osbjorn M. (2000a) "Activity, climate, and postcranial robusticity: Implications for modern human origins and scenarios of adpative change." Current Anthropology 41: 569-607.

Pearson, Osbjorn M. (2000b) "Postcranial remains and the origin of modern humans." Evolutionary Anthropology 9: 229-247.

Ponce de León, Marcia S. and Christoph P.E. Zollikofer (2001) "Neanderthal cranial ontogeny and its implications for late hominid diversity." Nature 412: 534-538.

Quam, Rolf M. and Fred H. Smith (1998) "A reassessment of the Tabun C2 mandible." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 405-421.

Rightmire, G. Philip (2001) "Diversity in the earliest 'modern' populations from South Africa, northern Africa and southwest Asia." In P.V. Tobias, M.A. Rath, J. Moggi-Cecchi, and G.A. Doyle, eds. Humanity from African naissance to coming millennia. Florence, Italy: Firenze University Press. pp. 231-236.

Schwarcz, Henry P., John J. Simpson, and Christopher B. Stringer (1998) "Neanderthal skeleton from Tabun: U-series data by gamma-ray spectrometry." Journal of Human Evolution 35: 635-645.

Stefan, V. and E. Trinkaus (1998) "Discrete trait and dental morphometric affinities of the Tabun 2 mandible." Journal of Human Evolution 34: 443-468.

Stringer, Chris (1998) "Chronological and biogeographic perspectives on later human evolution." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 29-37.

Stringer, Christopher and Clive Gamble (1993) In search of the Neanderthals. New York: Thames and Hudson.

Tostevin, Gilbert (2000) "The Middle to Upper Paleolithic transition from the Levant to Central Europe: In situ development or diffusion?" In J. Orschiedt and G.-C. Weniger, eds. Neanderthals and modern humans -- discussing the transition: Central and Eastern Europe from 50,000 - 30,000 B.P. Mettmann, Germany: Neanderthal Museum. pp. 92-111.

Valladas, Helene, Norbert Mercier, Jean-Louis Joron, and Jean-Louis Reyss (1998) "Gif Laboratory dates for Middle Paleolithic Levant." In T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Neandertals and modern humans in Western Asia. New York: Plenum Press. pp. 69-75.

Vandermeersch, Bernard (1989) "The evolution of modern humans: Recent evidence from Southwest Asia." In P. Mellars and C. Stringer, eds. The human revolution: Behavioural and biological perspectives on the origins of modern humans. Princeton: Princeton University Press. pp. 155-164.

Vermeersch, Pierre M. (2001) "'Out of Africa' from an Egyptian point of view." Quaternary International 75: 103-112.

Wolpoff, Milford H. (1999) Paleoanthropology, 2nd ed. Boston: McGraw-Hill.

Zeitoun, Valéry (2001) "The taxonomical position of the skull of Zuttiyeh." Comptes Rendus de l'Académie des Sciences, série IIa: Sciences de la Terre et des Plančtes 332: 521-525.

Zilhao, Joao and Francesco d'Errico (1999) "The chronology and taphonomy of the earliest Aurignacian and its implications for the understanding of Neandertal extinction." Journal of World Prehistory 13: 1-68.

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Neanderthals and Modern Humans -- A Regional Guide is written, designed, created and maintained by Scott J. Brown.
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