Written by C. David Kreger
Introduction

The discussion of our species, Homo sapiens, is probably the most difficult to put together. Whereas in the previous species have been introduced with historical background and a discussion of the early, most important finds, and the individuals responsible for the species designation, this introduction will focus on some of the theory implicit in the discussion of the origin and spread of H. sapiens.

Most researchers currently accept the statement that "modern" humans can be considered to date to approximately 200-250 kyr. Others (such as Milford Wolpoff), take the view that our species extends as far as approximately 2.0 myr, subsuming H. erectus, H. ergaster, and H. heidelbergensis. There are two polarizing camps on the issue of our species origin (though there is varying degrees of compromise between the two stances as well as various alternative positions): the multiregional (or continuity) camp, and the Out of Africa (replacement) camp.

The perspective of multiregionalists is that extending to the origin of H. erectus, there have been populations of humans living around the old world, and these all contributed to successive generations, eventually leading to modern humans. In this scenario, the Chinese and Indonesian material are the most direct ancestors of modern East Asians, the African material are the most direct ancestors of modern Africans, and that either the European populations are the most direct ancestors of modern Europeans, or that the European populations contributed significant genetic material to modern Europeans, with most of modern Europeans origins rooted in Africa or West Asia. Adherents to this model look at early material and try to trace continuity in morphology from those early populations to later populations in the same geographic area. In this model, there are paralleled changes in all penecontemporary populations, with enough genetic migration to maintain close species bonds, while still allowing the suite of racial features we see today.

The perspective of the Out of Africa model (often called Out of Africa II, referring to a second migration from Africa of a hominid population) adherents is that when there was a migration of H. erectus out of Africa into Asia and Europe, these populations (seen in materials like the Chinese and Indonesian erectus) did not contribute a significant amount of genetic material to later populations that led to modern humans (some claim no genetic ancestry to these groups and their descendants at all, a "strict" replacement model). At approximately 200 kya there was a second migration of hominids out of Africa. This time it was fully modern H. sapiens, which proceeded to replace whatever populations that then occupied Asia and Europe. Some see direct competition and extermination of the native populations, some see passive replacement due to better adaptive strategies, and some see genetic admixture with the preponderance of genetic material coming from the incoming human populations, eventually replacing and assimilating them into the greater collective. In this view there is a specific speciation event that occurred which led to the origin of H. sapiens in Africa, and this population is the forerunner of modern humans, leaving the European Neanderthals, Chinese erectus, and others out in the cold.

There are various models which embody combinations of these ideas, different "strict" interpretations of the two theories, etc. Multiregionalists look for similarities between populations in the same geographic location that are separated spatially, while people who follow replacement look for differences. It is oft a difference of semantics between different interpretations rather than real differences of opinion, but often there is real disagreement on the validity of research, and theoretical interpretations. This has led to some fairly severe strife within the paleoanthropological community, with potshots often taken unfairly at rival theories and rival theoreticians. For example, multiregionalism is often portrayed as a racist theory that claims different "races" have evolved to different "levels" of intelligence. Out of Africa II has often been portrayed as a religiously motivated idea that tries to come to terms with the biblical story of Genesis, as reference to the "Eve" theory suggests.

Beyond disagreement over fundamental issues like "What is a valid speciation event?" one fact stands out: neither theory has proved itself above the other in terms of parsimonious explanation of the fossil evidence. The general opinion among researchers seems to go in cycles, supporting OoA, then supporting MRE, then supporting OoA, etc. Currently, we seem to be at a cusp of support for replacement, and there seems top be a shifting in opinion more favorable to continuity. The highly publicized genetic studies that purportedly "proved" that Neanderthals did not contribute the modern human genome are so plagued with practical and theoretical problems to make their conclusions moot, especially since it does not in any way address the rest of the populations in the world, and their genetic fate.

Diagnostic Features

Europe

No part of these descriptions will favor one camp over the other; this being said, multiregionists have a very valid criticism of the supposed lack of any evidence of continuity between earlier and later fossil groups in geographic regions. This certainly holds true for Europe, where the perceived differences between the earlier Neanderthal populations and later "modern" H. sapiens populations. There are many sites which are attributed to what are called "early modern humans" are not always particularly older than "fully" modern humans, nor later than Neanderthal sites. As discoveries have continued over the last century Neanderthals are now clearly associated with Upper Paleolithic sites, and "modern" humans are no longer an Upper Paleolithic phenomenon.

The history of the interpretation of the European fossil record is marred by the fact that early attempts to demonstrate continuity by researchers such as A. Hrdlicka and G. Schwalbe were troubled by misdated specimens and faulty reconstructions, while early attempts to demonstrate replacement were based on the Piltdown hoax and the misdated Galley Hill material. These problematic foregrounds for further interpretation were also built upon by the faulty "type" approach of early paleoanthropologists, where La Chapelle was seen as a typical Neanderthal specimen and Cro-Magnon was seen as a typical early modern human. In fact, these two specimens are significantly atypical. Simple replacement and continuity models are beginning to fall apart, as the lines between earlier and later populations blurs in some respects and becomes more demarked in others.

Two major lines of thinking exist regarding the origins of modern humans in Europe: Neanderthals contributed some amount of significant genetic material to later modern human populations, or they did not. There are three major lines of evidence arguing for continuity, including:

• The evolution of the pre-Neanderthal and Neanderthal populations over time is in the direction of modern European populations. These trends include: anterior dental reduction, reduced nasal size, increased central and decreased lateral browridge height, and more developed mental eminences.
• European Neanderthals show a number of unique or especially common features with later Europeans, with lower frequency of expression in modern European populations.
• No other penecontemporary population shares unique features with the later Europeans.

While some may argue the third point, it is not clearly correct or incorrect, and the first two points are definite. In contrast, the evidence for lack of continuity is expressed in the following points:

• Early modern Europeans show limb proportions that are closely linked to warm-adapted African populations, and not with European Neanderthals.
• Genetic evidence in the form of mtDNA studies purporting to show evidence of Neanderthal and modern human divergence at approximately 700 kyr, and studies showing greater mtDNA variation in African populations, which is claimed evidence for African origin of modern humans.
• Behavioral differences between the Neanderthals and early modern humans in both absolute traits and complexity.

The genetic evidence is very suspect, and the theoretical and practical problems with such undertakings will continue to place such evidence as circumstantial at best for at least the next five to ten years. The behavioral evidence is an old one, and is crumbling into dust as paleoanthropology enters the 21st century. There likely were some behavioral differences between Neanderthal and later modern human populations, but there are many traits and sites linking the behaviors as developing - at least in part - from the former to the later. The limb proportion evidence, however, seems unmistakable. There was an influx of either populations or genetics into Europe with an African origin, though whether as a replacing invasion or a migration, with or without assimilation it is up in the air.

Very possibly occurringat this time was assimilation of the Neanderthal populations with incoming early modern humans, with differential representation in the resulting genome on the modern human side. It is difficult to believe that the Neanderthal traits would evolve into the modern form without significant contribution of genes for more gracile features. And the limb proportion argument cannot be resolved without a migration of Africans either bodily or genetically. A description of European H. sapiens, is best accomplished by examining material that is post-Neanderthal, though still early.

The largest site from the early Upper Paleolithic in Eastern and central Europe is Mladec. Discoveries of material from the site began in 1881, with descriptions beginning in 1925 by J. Szombathy. Tragically, the material was destroyed at the end of World War II by the Nazis, along with many other archaeological materials such as the Predmost� material, when it was burned in the Mikulov Castle fire. The fossil material is associated with an early Central European Aurignacian industry. The material dates from c. 32 kyr or more, and is very important for the size of the skeletal sample, and its late date. The material consists of over 100 specimens from the main cave, and a related side cave that features a triple burial of two adult males and a child: Mladec 5,6, and 46. The male specimens are very robust, and there is significant sexual dimorphism in terms of morphological variability. The three male crania are characterized by:

• Low braincases.
• Thick cranial bones.
• Posterior cranial flattening forming a Neanderthal-like occipital bun.
• Marked spongy bone development.
• Thick projecting supraorbitals (shaped differently than in Neanderthals).
• Large cranial capacities (1650 cc. for Mladec 5).

The female specimens are more gracile than the males, and are more modern in appearance. However, compared to more modern humans they are very robust, almost as robust as later Upper Paleolithic males. This population shows a reduction in the difference between male and female brain size relative to the Neanderthal condition. The females (Mladec 1 and Mladec 2) show cranial capacities of 1540 cc. and 1390 cc., respectively, an increase of 14% over Neanderthals, while the males (Mladec 5) show an increase of only 5% above the Neanderthal condition. This has been taken as evidence of lack of Neanderthal ancestry, as well as evidence that the Neanderthal populations were evolving into the modern form. The Mladec females show both similarities and differences from the earlier Neanderthal females, including:

• Larger cranial vault size.
• More midfacial prognathism.
• More anterior zygomatics.
• Lack a maxillary notch.
• A considerably narrower nose.
• Presence of a distinct canine fossa.

As noted before, there are some marked differences between the Mladec males and females. The systematic sexual differences include:

• Larger cranial vaults in the males (1650 cc. for Mladec 5, 1465 cc. average, a 13% differences).
• The males have larger and more projecting superciliary arches.
• Males have a shallow sulcus at the base of the forehead.
• Males have lower and less verticle foreheads.
• Males have more angled occipital areas with lambdoidal flattenning.
• Males have thick and broad superior nuchal lines extending onto the mastoids.
• Females have smaller nuchal planes.
• Females have a lower and less prominent inion.
• Males exhibit more Neanderthal-like general features, while females exhibit more modern general features.

Mladec is the earliest of the non-Neanderthal remains from Central Europe, but there are multiple sites dating later whose specimens show similar traits. Similar remains come from Zlaty Kun (Moravia), Cioclovina (Romania), Bacho Kiro levels 6/7 (Bulgaria), Velika Pecina (Croatia), Miesslingtal (Austria), the Stetten specimens (Vogelherd Cave, Germany), and the Hahn�fersand (Germany). These specimens range from 33 kyr to 25 kyr in date.

The material from Doln� Vestonice and nearby Pavlov were found on the Pavlov Hill in southern Moravia. The material dates to approximately 25-26 kyr, and is extremely variable in the expression of its traits. The Pavlov material is some of the most robust Upper Paleolithic remains known, while the Doln� Vestonice material is far more gracile. The DV1-3 specimens are very gracile females, with DV3 being identified with a small sculpture found nearby (identified from a facial pathology on DV3 and a similar representation on the figure). The DV16 specimen is more robust, and - unlike the robust Pavlov material - shows some Neanderthal features. As a whole, this material is very modern, but it shows some Neanderthal like features which could be evidence of continuity.

The Predmost� material is likely the earliest of this later Upper Paleolithic human remains, and definitively the largest. Unfortunately, the fire the Nazis used to destroy the Mladec material also claimed the Predmost� material. The material is very robust, though it does not approach the Neanderthal extreme, and shows very little difference between the sexes in this feature. As with the Mladec remains, the females show significant increases in brain size over the Neanderthals (13%), while the increase in the males is slight (2%). As with other material from this time period, some Neanderthal features can be found in specific specimens, but a systematic link between the Neanderthal and later human populations is lacking. While continuity is not proven impossible, the material is clearly more related to early modern humans from elsewhere in the world.

There are next to no remains in Western Europe that dates to the beginning of the Upper Paleolithic. There is the material from the Aurignacian at El Castillo Cave (dated to between 37 and 49 kyr), but the specimens have been lost, and a review of existing notes on the specimens makes it impossible to determine if this is a Neanderthal or an early modern human site. The only other specimen which might date to the early Upper Paleolithic is the Combe Capelle specimen from France. This specimen is a very robust women that was originally associated with Ch�telperronian levels (in 1909), but was surrounded by controversy from the time of its initial discovery. The specimen is definitely not a Neanderthal, but its circumspect dating makes its importance hard to determine.

There are numerous isolated fragments in Western Europe that date to later Aurignacian levels, but few are diagnostic enough to clarify the West Europe transition from the Neanderthal populations to early modern humans. One site which was thought to have been Mousterian in age is Grotte des Enfants (Grimaldi). However, the specimens are likely to have been in a Gravettian layer, and are thus even younger than Cro Magnon, the best-dated early site from Western Europe.

Probably the best known early modern specimen from Europe is the Cro Magnon material. This skeletal material was discovered at Abri Cro-Magnon, Les Eyzies, France, in 1868, and has been dated to approximately 32 kyr to 30 kyr. The material consists of five skeletons, three adult male, one adult female, and one infant. The site is an apparent deliberate burial, with various body adornments found alongside the skeletal material. The most cited specimen is Cro-Magnon 1; and adult male specimen. Also known as the "Old Man", Cro-Magnon 1 has a face pitted with a fungal infection and died in middle age. The cranium is complete except for the teeth and the mandibular condyles. Cro-Magnon 1 is definitely modern human, as seen from such features as:

• A high, rounded cranium.
• A steep forehead.
• Large cranial capacity (1600 cc.).
• A short face with rectangular orbitals.
• A tall and narrow nasal opening.
• A parabolic palate.
• A prominent mental eminence.

West Asia

The early humans from West Asia come from as early as 120 kyr, and have been much confused due to radiometric dating issues. For many years most of the Near Eastern archaic H. sapiens material was thought to date to approximately 45 kyr to 35 kyr, due to C14 dating. However, these dates were minimum ages, due to the limitations of radiocarbon dating. With the advent of TL (thermoluminescence) and ESR (electron spin resonance), more accurate dates have been procured. The material from this area of the world has had various interpretations as archaic modern human, Neanderthal, populations created by admixture of the former and latter, etc. However, it now seems more and more likely that much of the material can be attributed to archaic H. sapiens or early modern human, with some intrusion of Neanderthal populations during certain time periods.

The Near East material has been the center of much of the debate over replacement versus admixture in the Neanderthal issue. Due to the presence of both Neanderthal and early modern human material (and confusion as to what material belonged to which taxon), the various materials have been used as evidence of regional continuity, admixture between Neanderthals, early modern humans, and possibly East Asian lines, as well as evidence of marked differences between Neanderthals and early modern human populations. In summation, it seems apparent that Qafzeh and Skhul are early modern humans, and the habitation by both Neaderthal and early modern groups came at intervals of global weather changes, and do not constitute one highly variable population, or populations living simultaneously in the region. In other words, when it got colder the Neanderthals moved into the region and the early moderns moved into North Africa, and when it got warmer the Neanderthals moved back into Eastern Europe and the early moderns migrated back to the Near East. This pattern is seen in other fauna as well, at about the same time.

Of the two main Lower Paleolithic early modern sites in the Near East (Qafzeh and Skhul), the site of Qafzeh dates to an earlier time period. The cave of Qafzeh, located in Israel, has been dated by several methods to approximately 100 kyr to 90 kyr, and has yielded at material from at least 15 individuals, and possibly as many as 21. The specimen that is usually held up as the example of this material is Qafzeh IX, a 20-year-old male that is the most complete skeleton from the site. There is some disagreement on the sex determination, but later examinations of the pubic bone shows that the specimen was likely a male. The specimen was part of a double burial and was discovered next to the remains of a very young child. The taxonomy of this specimen is definitely modern, and would not be too out of place in a modern osteological collection from Eastern Europe or West Asia. Some of Qafzeh IX's diagnostic features include:

• A thin cranial wall that compares favorably to modern Europeans.
• A high forehead.
• A high, parallel-sided braincase.
• A reduced browridge that is divided into a supercilliary arch and a weak lateral torus.
• Presence of a canine fossa.
• A flat midface.
• A rounded occipital bone.
• Partial development of a mental eminence.
• Lack of a retromolar space.
• Cranial capacity of approximately 1550 cc.
• Approximately 172 cm in height.

Another complete male cranium is Qafzeh XI, an older adult specimen with thick supraorbitals that are prominent and projecting, a broad nose that was not prominent, a short broad face, a canine fossa, a marked maxillary notch, and a brain size of approximately 1554 cc. These male features contrast somewhat with the female material (most prominently Qafzeh V and Qafzeh III). These feature flattened foreheads, and flattened cranial rears, which meet at the top of the skulls to form a distinct angle. The supraorbitals are weakly developed, and more of a thickening at the end of the sloping frontal. Qafzeh III also features a thick, continuous, and projecting supraorbitals in front of the high rounded forehead, a broad occipital, thick cranial bone, and was about 160 cm tall.

The cave site of Skhul is very problematic in terms of dating the site. The site has been dated from 120 kyr up to 40 kyr by various methods. It has been dated by ESR (65 kyr/93 kyr), U-series dating (40 kyr/80 kyr), and TL (119 kyr). Some researchers have suggested that the site has had two habitation periods (seen by the two different dates given for ESR and U-series dates), while others see it as a single habitation. Whatever the actual date (or dates) of the site, its earliest dates seem to indicate that it was inhabited around the same time of later than the Qafzeh site. The dating problems revolve around the fact that the site has been dated by two animal teeth that are different in age, and has been dated by material from the burial level dated by TL. Whatever the date, the site is an extremely important one, with the remains of at least fourteen individuals present at the site.

The material from Skuhl is unequivocally modern in characteristic, though it is in no way identical to modern human populations. The specimens provide a wide range of features that are considered "archaic" as well as many that are considered "modern". The best preserved crania are those of three males (Skhul 4, 5, and 9). This led to the false impression of large cranial capacity (average brain size of 1550 cc for the three) for the Skhul specimens. These three show many "archaic" features that resemble the European Neanderthals (though not as pronounced in most cases). For example, in proportion, there is little difference between the Skhul sample and the European Neanderthals, while facial breadths are also similar (though facial heights are reduced). The Skhul 5 cranium is the best preserved (though it lacks much of the facial region), and the one most often used as representative of the sample. Features of the Skhul 5 cranium include:

• Age of death at approximately 30 to 40 (aged by wear on teeth and the cranial sutures).
• A high cranial vault.
• A rounded occipital at the rear.
• A modern flex at the cranial base.
• An underdeveloped mental eminence.
• A prominent browridge.
• A prognathic loew face.
• Pathology include evidence of abscessed teeth and rheumatoid arthritis at the TMJ.
• A 1518 cc brain (slightly below the modern European average).
  

The female specimens are less well-preserved than the males, but retain enough characteristics to identify them as female, and to make comparisons between the males and females. Just like the males, the females show a mixture of both "archaic" and "modern" characteristics. Skhul 2 has a well-developed mental eminence (the best developed chin out of the Skhul sample), but its pronounced continuous supraorbital torus makes it nearly impossible to categorize the specimen by itself as an anatomically modern human. The Skhul 7 female has the most Neanderthal-like characteristics, but it is as yet unreconstructed. The wide variation of traits from this site range from very Neanderthal-like, to very modern-like. These wide variations seen between the various specimens makes the use of Skhul V as the "type" specimen of the site unrealistic, especially since the differences between Skhul 5 and Skhul 9 are as great as those between modern humans and the Neanderthals.

The Skhul V individual also provided some postcranial material in the burial. This material included some vertebrae, some ribs, most of a left scapula, the right clavicle and part of the left clavicle, both humeri, the right radius and part of the left radius, the right ulna and part of the left ulna, various hand bones, the right ilium, half of the ischium, most of the right femur and part of the left femur, parts of the right and left tibiae, most of the left fibula, and some of the left foot bones. In general, the limb bones are longer and more gracile than the more robust Neanderthals. Postcrania from the other individuals is also present, which makes an examination of the stature of these people possible.

The Skhul 7 individual (female) is approximately 154 cm tall, while two male individuals (Skhul 4 and Skhul 5) average 180 cm, a very large sexual difference. The high sexual dimorphism seen in the sample is not particularly secure (or likely) due to the extremely small sample size. The average midsex height figure is 167 cm, a very high figure. The postcrania shows many more differences between the Skhul individuals and the Neanderthals than the cranial material does. Some of these differences include:

• Smaller shaft proportions relative to limb length.
• Smaller measures of articular surface relative to limb length.
• A change in shaft form (e.g. the femora lack midshaft flattening and excess internal thickening of the cortical bone; meaning less stenotic).
• A general reduction in robusticity.

Analyses of the burials and the morphological characteristics may support the two date hypothesis, or may simply mean two different groups at approximately the same time period inhabited the place. It seems that two different groups in the age of the material are possible: an earlier group that includes Skhul 3 and 6-10, and a later group that includes Skhul 1, 4, and 5. Since the more "modern" appearing specimens include Skhul 4 and Skhul 5, this may explain the wide variation at the site. However, since equally modern features have been found at the Qafzeh site at an earlier time period, the presence of more or less "modern" appearing groups at the site may be deceiving and inconsequential.


Africa

The Middle and early Upper Pleistocene archaeological record is the best understood of the regions discussed, and the paleoanthropological record is the most complete. Earlier in the century Africa was thought to have a very incomplete and unknown history. This changed with the discovery of many more important fossils and better dating of the ones that were already known. Where African material was once placed into sequences based on European dates and sequences, Africa has become the obvious center and origin of modern humans, with Europe as more peripheral in early human evolution.

Africa may be the origin of modern human anatomical features, but it also may be the origin of fully modern behavior. The later Middle Pleistocene, the Middle Stone Age (MSA), was a period of change in the lithic traditions of this area. All across Africa at this time, the Middle Stone Age developed out of the Acheulean. The large bifacial cutting and chopping tools (or cores for the flake tools depending on the artifact in question and the interpretation in question) fell into disuse and were replaced by more Mousterian-like small flake components. These changes seem to likely be related to the development of hafting, which allowed a wider range of tools and multiple component tools to be developed. Dating of the Middle Stone Age material place it beginning at over 180 kyr, and possibly even earlier. The African MSA is as early or earlier than other MSA variations in other regions, and is especially equally complex. This makes the old argument of Africa as a technological hinterland incorrect. Some of the African MSA traits include:

• Grindstones for the preparation of plant foods.
• Use of marine resources that were transported over 100 km.
• Use of bone for tool formation.
• The hafting of spear and other projectile points.
  

Skeletal remains are found throughout all of Africa, and all of it is usually considered H. sapiens, though not all is considered anatomically modern human. The oldest of the Middle Pleistocene specimens is a partial cranium from Florisbad, South Africa. ESR dates for this cranium give an age of approximately 250 kyr. The cranium consists of the sides and part of the front of the face, with most of the back and cranial base missing. The specimen is attributed to being a female. The specimen has been considered a phylogenetic link between earlier populations and modern living African populations, though some have questioned the validity of that idea. Some of the Florisbad cranium's features include:

• A broad, low frontal that is evenly curved.
• Presence of a wide sagittal keel.
• Supraorbitals that are a thickening at the forward edge of the sloping frontal.
• A shallow sulcus above the supraorbitals.
• A short face that is very broad.
• Expanded maxillary sinuses and consequentially puffy zygomatics.
• The orbits are spaced far apart.
• The nasal bones are broad and have little angulation.
• The dentition was moderate in size.

Other early MSA sites from Africa include the site of Eyasi, near Lake Tanganyika in Tanzania, which has been dated to 130 kyr and 200 kyr, with the real date unknown but likely somewhere in between. The K�bibat teenager was found in the Mifsud-Giudice quarry, near Rabat, and has been well-dated to greater than 200 kyr. From Southwest Djibouti, the Wadi Dagadl� maxilla has been dated to approximately 250 kyr.

The three specimens from the Kibish formation at Omo, Ethiopia, may or may not be associated with one another. Omo 1 was an excavated find, but Omo 2 and 3 were surface finds, and may not be related to each other or Omo 1. The site is also plagued by bad dating. A date of 130 kyr was originally given (uranium/thorium) but it was determined from shell, which is notoriously inaccurate for this method of dating, and the shell was from a lower level, making the Omo 1 specimen younger in any event. A faunal date of 60 kyr has been estimated as well, but the inability to relocate the site makes any date attributed to the remains suspect. The two fairly complete specimens are Omo 1 and Omo 2, and the variation displayed between the two and earlier and later populations make them very important as a link from earlier to moden Africans.

The specimens from Omo are undeniably modern in appearance, and at the time of their discovery (1967) doubled the known age of modern humans. In addition to the cranial elements, associated postcranial bones include bones from a shoulder, arm, hand, ribs, vertebrae, legs, and foot. These postcranial elements show the fully modern anatomy that is seen in the cranial specimens, further proof of these specimens' status as fully modern H. sapiens. The Omo 1 specimen is the most modern appearing, and consists of the following modern features:

• Long and curver parietals of an expanded brain case.
• A short broad face and high forehead.
• Prominent browridges that taper to the sides rather than forming a continuous thick bar.
• A modern-looking U-shaped palate.
• Presence of a mental eminence.
• Modern appearing teeth in both size and shape.
• A cranial capacity of over 1400 cc (exact measurement is difficult).
• A low nuchal torus position.
  

The most complete specimen from the MSA in Africa is the Ngaloba cranium (Laetoli 18). The cranium has been well-dated to between 129 kyr and 108 kyr by U-series dates. In general shape it resembles Omo 2, though the Ngaloba frontal is smaller and flatter, the sagittal keel is more weakly expressed, and the supraorbitals are less projecting. The Ngaloba specimen may be earlier than the Omo specimens, and may represent a link between earlier Broken Hill material, and the later Omo material (this works well for the frontal and occipital anatomy). However, since the Omo material is not securely dated, such an assumption may not be factual, and most likely will remain speculation unless better dated material is discovered.

The best dated sample of South African humans from the early Upper Pleistocene is the Klasies River Mouth material that have been dated to approximately 90 kyr, although two maxillary fragments are older, by as much as 30 kyr. While these remains are well-dated, ad have a central importance in the debates over a recent African origin or a Multiregional origin, they have been difficult to analyze due to the fragmentation, and the most that can be absolutely said for the morphology is that the population were relatively small, there seems to be great sexual dimorphism, and some features seem particularly modern while others are more "archaic" than similarly aged materials from other parts of the world.

There are many other MSA sites that contribute to the understanding of the African sequence, and the population shifts associated with them. The Sea Harvest material (Saldanha Bay) consists of a few human fragments found in the remains of a hyena den, and has been dated to 128 kyr to 75 kyr by faunal analysis. The Jebel Irhoud material is the earliest from North African MSA, and provides the only evidence of North African Mousterian-associated crania. This Cave site has yielded two crania, a juvenile humerus, and a juvenile mandible, and has been dated to 127 kyr to 87 kyr by ESR dates on horse teeth (though dates of 190 kyr to 105 kyr have also been espoused for this site. The Singa cranium from eastern Sudan provided fodder for many a debate, and has been described as "Neanderthaloid", a Neanderthal-modern hybrid, a "Proto-Bushman", and pathological. The last explanation is the one that has become accepted to explain some of the seemingly contradictory features of the specimen. This specimen has been dated to 133 kyr � 2 kyr by uranium/thorium dating of calcrete deposits on the cranial vault.

Material that dates to the Late Stone Age (LSA) in Africa come from Border Cave (some place it as early as the Klasies River Mouth, but evidence seems to place it much later) on the KwaZulu side of the border with Swaziland, the Origstad rock shelter (28.5 kyr), the Springbok Flats (Tuinplaas) material, the Lukenya material (17 kyr), the Circumturkana humans, the Kanjera material, and the Ishango material.

East Asia

The East Asian material is the material most often ignored by proponents of the Out of Africa model, and the one most touted by the Multiregional proponents as evidence to support their theory. While the Chinese material provides the best argument for continuity outside of Africa, the Indonesian sequence is filled with problems associated with dating and provenience, that make any argument connecting the various material specious. Current trends seems to be leading to a showdown between OoA proponents and MRE proponents over East Asia and Australia, so this are is very important for the future debates over general theory related to modern human origins. No matter the position of the particular researcher, none can say that this material is not important or interesting.

The Indonesian material has been the object of much debate due to problems of provenience and dating. The Ngandong material are a clear example of this. Though the material is conventionally listed as H. erectus by most researchers, it will be discussed on this page, since the material has been redated to 53 kyr to 27 kyr, and may be a link to modern East Asians. I am highly skeptical of this for now, but as the possibility is fascinating and cannot be ignored (not to mention the material is more important to discuss as a link from archaic East Asians to modern East Asians rather than a late surviving archaic of a population that left no modern progeny). The Ngandong material provides the single largest collection of crania from Java, with 15 specimens (including two tibiae) recovered from the High Terrace of the Solo River near Ngandong, and an additional female was discovered at nearby Ngawi.

This material (often called Solo rather than Ngandong) was originally placed in the Late Pleistocene due to the associated fauna found. Several attempts at dating have given dates of 101 kyr � 10 kyr (uranium/thorium dating of animal bones), 165 kyr (dating of a nearby High Terrace site), less than 250 kyr (date from fission track dating from similar Javan deposits), older than 300 kyr (modified ESR date from one of the crania), and approximately 500 kyr (Potassium/Argon date from a tuff near the site). However, these date have come into question based on dates presented by C. Swisher et al. of 53 kyr to 27 kyr based on U-series and ESR dating of animal teeth from other deposits found near the site. However, these new dates are not widely accepted, and the teeth used are stained gray and bluish by manganese, while the human material are brown to black, and dense and ceramic-like. These specimens were fossilized in different environments, and there seems to be no reason to assume the materials came from the same time period.

The Ngandong material were studied intensively by F. Weidenreich, but he died during the preparation of a monograph on the material, and it was published incomplete. Of the 13 crania or cranial fragments, nine are clearly adult, and one is clearly a juvenile (the rest are not diagnostic). It is believed that Solo 5, 9, 10, and 11 are male, and Solo 1, 4, 6, 8, and the Ngawi specimen are female. The juvenile (Solo 2) is likely male. The determination of sex is based on cranial size, projection and size of the nuchal torus, and the development oa an external occipital protuberance. These features do not overlap between the sexes. Other characteristics do overlap, but show a average difference between the sexes, including vault thickness and development of the lines and crests marking the muscle attachments.

These specimens are purported to share a number of similarities (regional similarities if the MRE hypothesis holds in East Asia) with earlier Indonesian specimens like the Sangiran material and the Sambungmachan material. Similarities that are reported between these samples include:

• The frontal bone is flat, without any development of a frontal boss, and merges onto the top of the supraorbital torus.
• The supraorbital torus is close to horizontal in orientation.
• There is a distinct frontal keel extending over virtually the entire squama.
• There is a prebregmatic eminence, a cross-like elevation at the meeting of the sagittal and coronal sutures.
• The top of the vault is evenly curved along the sagittal suture.

While there are isolated teeth that seem to be associalted with the later Middle and early Upper Pleistocene faunas known from several East Asian countries, the vast majority of material from this time period comes from China. The material from China comes from a broad spatial span, and includes several nearly complete crania, as well as much of a postcranial skeleton. This is the region where multiregional hypotheses gain their best evidence (indeed, some researchers have begun to accept continuity in East Asia, while maintaining support for replacement in Europe). The earliest material from China that has been designated as H. sapiens (or at least has a good probability of being H. sapiens) is the Dali cranium from Shaanxi Province, and several teeth from Tongzi in Guizhou Province.

The Dali cranium was found in river gravels, and sustained some damage on its lower face that has never been reconstructed. This damage pushed the maxilla upwards, making the lower face look shorter than it actually was. The cranium came from a male individual under 30 years of age. Some of the features include:

• A large, thick cranial vault.
• A small braincase (1120 cc).
• Large supraorbital tori that arch over the upper orbits.
• A low forehead that begins behind the ridges.
• A distinct frontal boss.
• Presence of a narrow sagittal keel.
• The occipital squama is expanded relative to the size of the nuchal plane.
• The parietal bosses are well-developed.
  

The Dali cranium has several other features that have been presented as evidence of a link between the earlier Chinese material from Zhoukoudian, as well as between the Java material from Ngandong. These features are also used as evidence to fit Dali into a set of "East Asia" characteristics that link the fossil material between what is generally considered the East Asian H. erectus material and modern living Asians. These characteristics include:

• The angular torus.
• The temporal-frontal articulation in the temple region.
• The thick vault bones.
• The depressed region where the browridges meet over the nose.
• An elevated lower cheek border.
• The presence of a maxillary notch.
• Orbital pillars that face forward.
• Nasal bones that are narrow, flat, and oriented vertically, with a constricted internasal crest.
• A moderate swelling of the maxilla along the nasal border.
  

While the lack of dentition in Dali prevents the inquiry into whether it had shovel-shaped incisors like current East Asian populations, the six teeth from Tongzi include a central incisor that is shovel-shaped in the normal East Asian pattern. The incisor has moderately strong marginal ridges, a lingual tubercle, and a straight buccal face. This is another point in favor of East Asian continuity.

From a little more recent time period in China, comes the Jinniushan material. The material from Jinniushan in Liaoning Province consists of the cranium and much of the postcranial skeleton of a 20-year-old female. The postcrania include four vertebrae, some ribs, an os coxa, an ulna, a patella, and 30 hand and foot bones. The remains were found in a consolidated fissure-filling in an isolated karst prominence. ESR dates (early uptake model) by Chen Tiemei et al. has given a dat of approximately 187 kyr (165 kyr to 195 kyr) for this material. The cranium has the following features:

• A large cranial vault (1260 cc).
• Unusually thin vault bones.
• A gabled cranial contour.
• A broad frontal bone.
• Central supraorbital thinning.
• A gracilized posterior region with marked nuchal plane reduction.
  

While these features combine to make Jinniushan the earliest specimen with this mixture of modern features, there are many features that are not modern. These features include a great distance between the orbits, the supraorbitals project far in front of the forehead, and the nuchal torus extends across the entire occipital bone and has a distinct, straight upper border there. However, the case for calling this specimen modern H. sapiens is probably as strong as African (Klasies River Mouth) and West Asian (Qafzeh) samples of almost half the age of Jinniushan. This specimen has also been used as a link to the earlier Zhoukoudian material, with similar features that include:

• Arched supraorbitals with a sulcus separating them from the frontal squama.
• A tall narrow keel on the frontal bone.
• Transversely flat, vertically oriented face.
• Marked facial breadth.
• The top of the nasal bones makes a horizontal juncture with the frontal bone.
• A high position for the maxillary notch.
• Incisor shoveling, marginal ridges combined with a straight incisor blade with a tubercle with finger-like projections extending above it.
  M3 reduction.
  

Other earlier H. sapiens remains from China include a maxilla and occipital bone from Chaohu (also known as both Chaoxian and Yinshan) in Anhui Province, the Changyang maxilla from Hubei, a child's parietal and teeth from Dincun (Shanxi), teeth from Xindong (Beijing), a skullcap from Maba in Guangdong Province, and material from ten individuals at Xujiayao (Shanxi) (2, 3, mandible).

The unequivocable modern H. sapiens from continental East Asia include: the Liujiang material (67 kyr), cranial and limb remains from Salawusu (50 kyr to 37 kyr), the Laishui material (60 kyr), the Ziyang material (either 39 kyr to 36 kyr, or 7 kyr depending on which level it came from, though this can no longer be established), the Upper Cave (101, 102) material at Zhoukoudian (29 kyr to 24 kyr), an occipital from Shiyu (28 kyr), the Hamakita material from Japan, the Yamashita-cho material from Okinawa (32 kyr), the Pinza-Abu cranial fragment and postcrania (26 kyr), the Minatogawa material (18 kyr), the Niah Cave material from Borneo (40 kyr), and the Wadjak material (1, 2) from Indonesia.

Australia

The origin of the native Australians has been debated for years, with the estimate of when Australia was likely inhabited being pushed further back in time over the years. It is now absolutely certain that Australia was inhabited by at least 60 kyr to 50 kyr, and there is evidence that indicates the first habitation occurred by 115 kyr. The evidence of the earlier habitation comes from a supposed link between the much younger skeletal remains from Australia, the Javan material. This argument his centered around a single fossil (WLH 50) at this time (though there are several other Late Pleistocene/Early Holocene fossils that purpoted to demonstrate this also). M. Wolpoff has also noted similarities between his reconstruction of Sangiran 17 and Aboriginal Australians, including:

• Marked prognathism, especially at the lower nasal border and below and is reflected in the high facial angle.
  
• A ridge or ridges paralleling the suture between zygomatic and maxillary bones.
  
• Eversion of the lower zygomatic, caused by the fact that the lower outside corner of the zygomatic, at the corner between the side and front of the face, extends more laterally than any of the face above it and gives the facial profile the outline of a pentagon.
  
• The lower outside rim of the orbit is rounded.
  
• The lower border of the nose lacks a distinct line marking the change from nasal floor to the face of the maxilla below the nose.
  
• The alveolar plane for the posterior tooth row is convexly curved.
  

The earliest dated Australian skeletal material comes from the Willandra Lakes system in New South Wales, and has been dated to greater than 35 kyr. The Willandra hominids include three mostly complete specimens, the most important in current debates is WLH 50. WLH 50 was a surface find that is the least securely dated of the three crania (though ESR dates of the bone place it minimally within the same time span as the others), and Wolpoff and others consider it a clear link to the early Indonesian material. One of the notable features of the specimen is its large cranial capacity of 1540 cc, larger than the recorded maximum of modern Native Australians.

The other two Willandra specimens include WLH (Mungo) 1, a fragmentary cremated female, and WLH (Mungo) 3 is an older, nearly complete male specimen. Both are smaller and more gracile than WLH 50, with well-rounded foreheads, thin vault bones, weak muscle attachments, and weak or moderate supraorbital development. While in general their features can be considered gracile, there is some robusticity related to heavy anterior tooth wear, including the presence of inion well above internal occipital protuberance in WLH 1, and a broadly developed nuchal torus on WLH 3. In addition, the incisors and canines of WLH 3 are worn much more than the posterior teeth.

Much later in time come specimens such as the Kow Swamp material (e.g. Kow Swamp 1 and 5) at 14 kyr to 9.5 kyr, the Coobool Crossing material (e.g. 16, 49, 65, 76, 86) at 14 kyr, and the Keilor cranium and femur at 12 kyr.

Bibliography

Aiello, L.C. 1993. "The fossil evidence for modern human origins in Africa: A revised view." In American Anthropologist, vol. 95, no. 1, pp. 73-96.

Aiello, L.C., and C. Dean. 1990. An Introduction to Human Evolutionary Anatomy. London: Academic Press.

Barinaga, M. 1992. "'African Eve' backers beat a retreat." In Science, vol. 255, pp. 686-687.

Br�uer, G. 1984. "The 'Afro-European sapiens hypothesis' and hominid evolution in East Asia during the late Middle and Upper Pleistocene." In The Early Evolution of Man, with Special Emphasis on Southeast Asia and Africa, ed. by P. Andrews and J.L. Franzen, pp. 145-165. Courier Forschungsinstitut Senckenberg vol. 69.

Cavalli-Sforza, L.L., A. Piazza, P. Menozzi, and J. Mountain. 1988. "Reconstruction of human evolution: Bringing together genetic, archaeological, and linguistic data." In Proceedings of the National Academy of Sciences, vol. 85, pp. 6002-6006.

Churchill, S.E., O.M. Pearson, F.E. Grine, E. Trinkaus, and T.W. Holliday. 1996. "Morphological affinities of the proximal ulna from Klasies River Main Site: Archaic or modern?" In Journal of Human Evolution, vol. 31, no. 3, pp. 213-237.

Deacon, H.J. 1992. "Southern Africa and modern human origins." In Philosophical Transactions of the Royal Society, vol. 337, pp. 177-183.

Frayer, D.W., M.H. Wolpoff, F.H. Smith, A.G. Thorne, and G.G. Pope. 1993. "The fossil evidence for modern human origins." In American Anthropologist, vol. 95, no. 1, pp. 399-405.

Howells, W.W. 1976. "Explaining modern man: Evolutionists versus migrationists." In Journal of Human Evolution, vol. 5, pp. 477-496.

Johanson, D., and B. Edgar. 1996. From Lucy to Language. New York: Simon and Schuster Editions.

Jones, T.R. 1868. "On the human skulls and bones found in the cave of Cro-Magnon, near Les Eyzies." In Reliquiae aquitanicae.

Kennedy, G.E. 1984. "Are the Kow Swamp Hominids 'Archaic'?" In American Journal of Physical Anthropology, vol. 65, pp. 163-168.

Kramer, A. 1991. "Modern human origins in Australia: Replacement or evolution?" In American Journal of Physical Anthropology, vol. 86, no. 4, pp. 455-473.

Leakey, R.E.F., K.W. Butzer, and M.H. Day. 1969. "Early Homo sapiens remains from the Omo River region of Southwest Ethiopia." In Nature, vol. 222, pp. 1132-1138.

McCown, T.D., and A. Keith. 1939. "The fossil remains from the Levalloiso-Mousterian." In The Stone Age of Mount Carmel, vol. II. Oxford: The Clarendon Press.

Relethford, J.H. 1995. "Genetics and modern human origins." In Evolutionary Anthropology, vol. 4, no. 2, pp. 53-63.

Rightmire, G.P. 1978. "Florisbad and human population succession in Southern Africa." In American Journal of Physical Anthropology, vol. 48, pp. 475-486.

Rightmire, G.P. 1981. "Later Pleistocene hominids of eastern and southern Africa." In Anthropologie, vol. 19, pp. 15-26.

Smith, F.H. 1992. "Models and realities in modern human origins: The African fossil evidence." In Philosophical Transactions of the Rpyal Society of London, vol. 337, pp. 243-250.

Smith, F.H. A.B. Falsetti, and S.M. Donnelly. 1989. "Modern human origins." In Yearbook of Physical Anthropology, vol. 32, pp. 35-68.

Stringer, C.B. 1990. "The emergence of modern humans." In Scientific American, vol. 263, no. 6, pp. 98-104.

Stringer, C.B. 1992. "Reconstructing recent human evolution." In Philosophical Transactions of the Royal Society of London, vol. 337, pp. 217-224.

Thorne, A.G., and P.G. Macumber. 1972. "Discoveries of Late Pleistocene Man at Kow Swamp, Australia." In Nature, vol. 238, pp. 316-319.

Thorne, A.G., and S.R. Wilson. 1977. "Pleistocene and recent Australians: A multivariate comparison." In Journal of Human Evolution, vol. 6, pp. 393-402.

Vandermeersch, B. 1969. "Les nouveaux squellettes most�riens d�couverts � Qafzeh (Isra�l) et leur signification." In Comptes Rendus de l'Acad�mie de Sciences, vol. 268, pp. 2562-2565.

Vandermeersch, B. 1981. Les Hommes Fossiles de Qafzeh. Paris: Editions du Centre National de la Recherche Scientifique.

Weidenreich, F. 1947. "Facts and speculations concerning the origin of Homo sapiens." In American Anthropologist, vol. 49, pp. 187-203.

Weidenreich, F. 1947. "The trend of human evolution." In Evolution, vol. 1, pp. 221-236.

Wolpoff, M. 1999. Paleoanthropology. second edition. Boston: McGraw-Hill.

Wolpoff, M., and R. Caspari. 1996. "The modernity mess." In Journal of Human Evolution, vol. 30, no. 2, pp. 167-172.

Wood, B.A. 1994. "The problems of our origins." In Journal of Human Evolution, vol. 27, no. 6, pp. 323-330.

Wu, X. 1981. "A well-preserved cranium of an archaic type of early Homo sapiens from Dali, China." In Scientia Sinica, vol. 241, pp. 530-539.

Wu, X., and F.E. Poirier. 1995. Human Evolution in China: A Metric Description of the Fossils and a Review of the Sites. New York: Oxford University Press.