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12.  Modern Homo sapiens

Keith Chan, Ph.D., University of Missouri: Grossmont College and MiraCosta College

Learning Objectives

Identify the skeletal and behavioral traits that represent modern Homo

sapiens.

Critically evaluate different types of evidence for the origin of our

species in Africa, and our expansion around the world.

Understand how the human lifestyle changed when people transi-

tioned from foraging to agriculture. Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.1 The excavation of an exposed cave at Jebel Irhoud, Morocco, where hominin fossils were found in the 1960s and in 2007. Dating showed that they represent the earliest-known modern Homo sapiens.

The walls of a pink limestone cave exposed to the outside world in the hillside of

Jebel Irhoud jutted out of the otherwise barren landscape of the Moroccan desert

(Figure 12.1). The year was 2007 and it turned out to be a momentous occasion for

science. A fossil unearthed by a team of researchers was barely visible to the un-

trained eye. Just the fossil cranium’s robust brows were peering out of the rock. The

�nd was welcome but not sheer luck: Hominin fossils have been found here since

their �rst accidental discovery by miners in 1960. This research team from the Max

Planck Institute for Evolutionary Anthropology was just the latest to explore the

prehistoric human presence in this part of North Africa. Excavating near the �rst

discovery, the researchers wanted to learn more about how Homo sapiens lived far

from East Africa, where we thought our species originated.

The scientists were surprised when they analyzed the cranium, named Irhoud 10,

and other fossils. Statistical comparisons with other human crania concluded that

the Irhoud face shapes were typical of recent modern humans while the braincases

matched ancient modern humans. Based on the �ndings of other scientists, the

team expected these modern Homo sapiens fossils to be around 200,000 years old.

Instead, dating revealed that the cranium had been buried for around 315,000 years.

Previous: Archaic Homo

Next: Race and Human Variation

Together, the modern-looking facial dimensions and the older date changed the in-

terpretation of our species, modern Homo sapiens. Our key evolutionary changes

from the archaic Homo sapiens of the previous chapter to our species today hap-

pened 100,000 years earlier than what we had thought. In addition, the new infor-

mation suggests that our home region covered more of the vast African continent

instead of being concentrated in the east.

This big addition to the study of modern Homo sapiens is just one of the latest in this

continually advancing area of biological anthropology. Researchers are continually

discovering amazing fossils and ingenious ways to collect data and test hypotheses

about our past. Through the collective work of scientists, including archaeologists,

geneticists, and anatomists, we are building an overall theory or explanation of

modern human origins. We will �rst cover the skeletal changes from archaic Homo

sapiens to modern Homo sapiens. Next, we will track how modern Homo sapiens ex-

panded the range of its species around the world. Lastly, we will cover the develop-

ment of agriculture and how it changed human culture to how we practice it today.

DEFINING MODERNITY

What de�nes a modern Homo sapiens when compared to an archaic Homo sapiens,

like the ones in the previous chapter? Modern humans, like you and me, have a set

of derived traits that are not seen in archaic humans or any other hominin. As with

other transitions in hominin evolution, such as increasing brain size and bipedal

ability, modern traits do not appear fully formed or all at once. In other words, the

�rst modern Homo sapiens was not just born one day from archaic parents. The

traits common to modern Homo sapiens appeared in a mosaic manner: gradually

and out of sync with one another. There are two areas to consider when tracking

the complex evolution of modern human traits. One is the physical change in the

skeleton. The other is behavior inferred from the cranium and material culture.

Skeletal Traits Previous: Archaic Homo Next: Race and Human Variation

The skeleton of a modern Homo sapiens is less robust than that of an archaic Homo

sapiens. In other words, the modern skeleton is gracile, meaning that the structures

are thinner and smoother. Differences related to gracility in the cranium are seen in

the braincase, the face, and the mandible. There are also broad differences in the

rest of the skeleton.

Cranial Traits

Several elements of the braincase differ between modern and archaic Homo sapiens.

Overall, the shape is much rounder, or more globular, on a modern skull (Lieber-

man, McBratney, and Krovitz 2002; Neubauer, Hublin, and Gunz 2018; Pearson 2008)

(Figure 12.2). You can feel the globularity of the modern human skull on the example

built into you. Feel the height of your forehead with the palm of your hand. Viewed

from the side, the tall vertical forehead of a modern Homo sapiens stands out when

compared to the sloping archaic version. This is because the frontal lobe of the

modern human brain is larger than the one in archaic humans, and the skull has to

accommodate the expansion. The vertical forehead reduces a trait that is common

to all other hominins: the brow ridge or supraorbital torus. The sides of the brain-

case also exhibit changes associated with the globular expansion of the brain: the

parietal lobes of the brain and the matching parietal bones of the skull both bulge

outward more in modern humans. At the back of the skull, the archaic occipital bun

is no longer present. Instead, the occipital region of the modern human cranium has

a derived tall and smooth curve, again re�ecting the globular brain inside. The dif-

ferent priorities in brain regions may also indicate cognitive and behavioral differ-

ences between archaic humans and modern humans, discussed in the next section.

The trend of shrinking face size across hominins reaches its extreme with our

species as well. The facial bones of a modern Homo sapiens are extremely gracile

compared to all other hominins (Lieberman, McBratney, and Krovitz 2002). One spe-

ci�c dimension to compare is the thickness of the zygomatic arches, or cheekbones.

As with the shrinking of the face leading up to our species, the decreasing reliance

on needing large teeth for survival may have been the reason that modern human

faces are so gracile in comparison to other humans. Continuing a trend in hominin

evolution, technological innovations kept reducing the importance of teeth in repro-

Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.2 Comparison between modern (left) and archaic (right) Homo sapiens skulls. Note the overall gracility of the modern skull, as well as the globular braincase.

ductive success (Lucas 2007).

As natural selection favored

smaller and smaller teeth, the

surrounding bone holding

these teeth also shrank.

Connected to the face, the

mandible is also gracile in mod-

ern humans when compared to

archaic humans and other ho-

minins. Interestingly, our

mandibles have pulled back so

far from the prognathism of earlier hominins that we gained an extra structure at

the most anterior point, called the mental eminence. You know this structure as the

chin: trace your own chin and feel how it curves forward before swooping posteri-

orly toward your neck. At the skeletal level, it resembles an upside-down “T” at the

centerline of the mandible (Pearson 2008). If you look back at illustrations of other

hominins, you will see that they all lack a chin. Instead, their mandibles curve

straight back without a forward point. What is the chin for and how did it develop?

Flora Gröning and colleagues (2011) found evidence of the chin’s importance by sim-

ulating physical forces on computer models of different mandible shapes. Their re-

sults showed that the chin acts as structural support to withstand strain on the oth-

erwise gracile mandible. In other words, as natural selection favored smaller denti-

tion, the chin developed to maintain structural integrity of the mandible.

Post-Cranial Gracility

The rest of the modern human skeleton is also more gracile than its archaic coun-

terpart. The differences are clear when comparing a modern Homo sapiens with a

cold-adapted Neanderthal (Sawyer and Maley 2005), but the trends are still present

when comparing modern and archaic humans within Africa (Pearson 2000). Overall,

a modern Homo sapiens post-cranial skeleton has thinner cortical bone, smoother

features, and more slender shapes when compared to archaic Homo sapiens (Figure

12.3). For example, the modern pelvis has gracile features along its surface and is

Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.3 Anterior views of modern (left) and archaic (right) Homo sapiens skeletons. The modern human has an overall gracile appearance at this scale as well.

narrower in overall width. Our elbow and knee joint surfaces are also narrower. Even

the individual �ngers and toes are more slender in modern humans. Comparing

whole skeletons, modern humans have longer limb proportions relative to the length

and width of the torso, giving us lankier outlines.

As with the cranial traits, we have to con-

sider the evolutionary process behind

postcranial gracility. Why is our skeleton so

gracile compared to those of other ho-

minins? Natural selection can drive the gra-

cilization of skeletons in several ways

(Lieberman 2015). A slender frame is adapted

for the ef�cient long-distance running abil-

ity that started with Homo erectus. Further-

more, slenderness is a genetic adaptation for

cooling an active body in hotter climates,

which aligns with the ample evidence that

Africa was the home continent of our

species.

Behavioral Modernity

Aside from physical differences in the skele-

ton, researchers have also tracked clues of

behavioral changes from archaic to modern

humans. From the anthropology of our species today, we know that we practice a

very complex version of culture, with many layers to our language, art, social orga-

nization, and technology, among other areas. Did cultural complexity increase grad-

ually or quickly with the �rst modern humans? This question is being actively inves-

tigated. A major obstacle to answering this question is that it is hard to de�ne and

measure cultural complexity. Since we cannot directly observe humans of the dis-

tant past, we have to infer these measures of human behavior from other types of

evidence. Two particularly illuminating areas are archaeology and the analysis of re-

constructed brains.

Previous: Archaic Homo

Next: Race and Human Variation

Archaeology tells us much about the behavioral complexity of past humans by inter-

preting the signi�cance of material culture. In terms of evolved advanced culture,

items created with an artistic �air, or as a decorative piece, speak of some abstract

thought process (Figure 12.4). The demonstration of dif�cult artistic techniques and

technological skills hints at social learning and cooperation as well. For example,

most of your skills were taught to you by a more experienced person, upon which

you’ve developed your own style with practice. Some day you may pass on what you

know to someone else using language to convey your knowledge. The same process

is believed to have happened with early modern humans in areas such as toolmaking

and craftwork, producing the sophisticated material culture that we can now study.

According to paleoanthropologist John Shea (2011), one way to track the complexity

of past behavior through artifacts is by measuring the variety of tools found to-

gether. The more types of tools constructed with different techniques and for dif-

ferent purposes, the more modern the behavior. Turning this view to ourselves,

think of all of the tools we have available to us today at a typical hardware store and

the cumulative knowledge they represent. This idea of measuring past behavior is

promising, but researchers are still working on an archaeological way to measure

cultural complexity that is useful across time and place.

The interpretation of brain anatomy is another promising approach to studying the

evolution of human behavior. When looking at the body of work on this topic in

modern Homo sapiens brains, researchers found a weak association between brain

size and test-measured intelligence (Pietschnig et al. 2015). This means that there

are more signi�cant factors that affect tested intelligence than just brain size. Addi-

tionally, they found no association between intelligence and biological sex. Since the

sheer size of the brain is not useful for weighing intelligence, paleoanthropologists

are instead investigating the differences in certain brain structures. The differences

in organization between modern Homo sapiens brains and archaic Homo sapiens

brains may re�ect different cognitive priorities that account for modern human cul-

ture. Researchers (e.g., Bruner 2010) have hypothesized that the expanded frontal

and parietal lobes in the globular modern human braincases mean that we can do

more complex thinking regarding memory and social ability than the Neanderthals

could. In contrast, the Neanderthal brain prioritized the visual regions where the

occipital bun was located, with fewer neurons in the frontal area for complex think-

ing. As with the archaeological line of research in the preceding paragraph, this is a

very active area of investigation. New discoveries will re�ne what we know about the

Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.4 Carved ivory �gure called the Lion-Man of the Hohlenstein-Stadel. It dates to the Aurignacian culture, between 35 and 40 kya. What does this artifact suggest about the culture and technical skill of its artist?

human brain and apply that knowledge to studying the distant

past.

Taken together, the cognitive abilities in modern humans may

have translated into an adept use of tools to enhance survival.

The ability to process a new environment, adapt to it with in-

novative technology, and pass on that knowledge may be the

key behind the success of modern Homo sapiens. Researchers

Patrick Roberts and Brian A. Stewart call this concept the

generalist-specialist niche: Our species is an expert at living

in a wide array of environments, with populations culturally

specializing in their own particular surroundings (Roberts and

Stewart 2018). The next section tracks how far around the

world these skeletal and behavioral traits have taken us.

FIRST AFRICA, THEN THE WORLD

What enabled modern Homo sapiens to expand its range fur-

ther in 300,000 years than Homo erectus did in 1.5 million years? The key is the set of

derived biological traits from the last section. The gracile frame and neurological

anatomy allowed modern humans to survive and even �ourish in the vastly different

environments they encountered. Based on multiple types of evidence, the source of

all of these modern humans, including all of us today, was Africa.

This section traces the origin of modern Homo sapiens and the massive expansion of

our species across all of the continents except Antarctica by 12,000 years ago. While

modern Homo sapiens �rst shared geography with archaic humans, modern humans

eventually spread into lands where no human had gone before. Starting with the

�rst-known modern Homo sapiens, around 315,000 years ago, we will follow our

species from a time called the Middle Pleistocene to the end of the Late Pleistocene.

Culturally, we will trace developments from the Middle Stone Age through the tran-

sition around 50,000 years ago to the Later Stone Age, when cultural complexity

quickly grew with both technology and artistry. We will end this section right before

the next big cultural change, called the Neolithic Revolution.

Previous: Archaic Homo

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A few notes on this part of the chapter: It is organized from past to present when

possible, though a lot happens simultaneously to our species in that time. Figure 12.5

shows the broad routes that our species took expanding around the world. I encour-

age you to make your own timeline with the dates in this part to see the overall

trends. References are provided to the research leading to the information on key

�nds. Search for these scienti�c papers online to see how researchers reach the

conclusions presented here.

Previous: Archaic Homo

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Figure 12.5 Maps depicting the estimated range of modern Homo sapiens through time.

The shaded area is based on geographical connections across known sites. Note the

growth in area starting in Africa and the oftentimes coastal routes that populations

followed. Previous: Archaic Homo

Next: Race and Human Variation

�e Start of Modern Homo sapiens in Africa

We start with the ample fossil evidence supporting the theory that modern humans

originated in Africa during the Middle Pleistocene, having evolved from African ar-

chaic Homo sapiens. The earliest dated fossils considered to be modern actually have

a mosaic of archaic and modern traits, showing the complex changes from one type

to the other. Experts have various names for these transitional fossils, such as

Early Modern Homo sapiens or Early Anatomically Modern Humans. However they

are labeled, the presence of some modern traits means that they illustrate the origin

of the modern type. Three particularly informative sites with fossils of the earliest

modern Homo sapiens are Jebel Irhoud, Omo, and Herto.

Recall from the start of the chapter that the most recent �nds at Jebel Irhoud are

now the oldest dated fossils that exhibit the traits of modern Homo sapiens. Besides

Irhoud 10, the cranium that was dated to 315,000 years ago (Hublin et al. 2017;

Richter et al. 2017), there were other fossils found in the same deposit that we now

know are from the same time period. In total there are at least �ve individuals, rep-

resenting life stages from childhood to adulthood. These fossils form an image of

high variation in skeletal traits. For example, the skull named Irhoud 1 has a primitive

brow ridge, while Irhoud 2 and Irhoud 10 do not (Figure 12.6). The braincases are

lower than what is seen in the modern humans of today but higher than in archaic

Homo sapiens. The teeth also have a mix of archaic and modern traits that defy clear

categorization into either group.

Research separated by nearly four decades uncovered fossils and artifacts from the

Kibish Formation in the Lower Omo Valley in Ethiopia. These Omo Kibish hominins

were represented by braincases and fragmented postcranial bones of three individu-

als found kilometers apart, dating back to 195,000 years ago (Day 1969; McDougall,

Brown, and Fleagle 2005). One interesting �nding was the variation in braincase size

between the two more-complete specimens: While the individual now named Omo I

had a more globular dome, Omo II had an archaic-style long and low cranium. In

more recent �eldwork, an informative section of the Omo I pelvis was found in a re-

excavation in 2001. Analysis by Ashley S. Hammond and colleagues (2017) found that

the measurements and observations were in line with modern Homo sapiens, al-

though larger in absolute size and robusticity. Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.6 Composite rendering of the Jebel Irhoud hominin based on micro-CT scans of multiple fossils from the site. The facial structure is within the modern human range, while the braincase is between the archaic and modern shapes.

Figure 12.7 This model of the Herto cranium showing its mosaic of archaic and modern traits.

Also in Ethiopia, a team led by Tim White (2003) exca-

vated numerous fossils at Herto. There were fossilized

crania of two adults and a child, along with fragments

of more individuals. The dates ranged between

160,000 and 154,000 years ago. The skeletal traits and

stone tool assemblage were both intermediate be-

tween the archaic and modern types. Features remi-

niscent of modern humans included a tall braincase

and thinner zygomatic (cheek) bones than those of ar-

chaic humans (Figure 12.7). Still, some archaic traits

persisted in the Herto fossils. Looking at the face, the

supraorbital tori were still prominent. The cranium

included an angled occipital bone and was longer than

in present-day modern Homo sapiens. Statistical

analysis by other research teams concluded that at

least some cranial measurements �t just within the

modern human range (McCarthy and Lucas 2014), favoring categorization with our

own species.

Summary of Early Modern H. sapiens in Africa

The combined fossil evidence paints a picture of di-

versity in geography and traits. Instead of evolving in

just East Africa, the Jebel Irhoud �nd revealed that

early modern Homo sapiens had a wide range across

Middle Pleistocene Africa. The hypothesis that there

was no single original home within Africa for our

species is called African multiregionalism (Scerri et

al. 2018). Supporting this explanation, fossils have different mosaics of archaic and

modern traits in different places and even within the same area. The high level of di-

versity from just these fossils shows that the modern traits took separate paths to-

ward the set we have today. The connections were convoluted, involving �uctuating Previous: Archaic Homo

Next: Race and Human Variation

gene �ow among small groups of regional nomadic foragers across a large continent

over a long time.

What about behavioral modernity? Jebel Irhoud, Omo, and Herto all bore Middle

Stone Age tools of the same �aked style as archaic assemblages, even though they

were separated by almost 150,000 years. The apparent stability in technology may

be evidence that behavioral modernity was not so developed back then, though

there was a high variety of tool types used throughout that time. No clear signs of

art dating back this far have been found either. Other hypotheses not related to be-

havioral modernity could explain these observations. The tool set may have been

suitable for thriving in Africa without further innovation. As for the lack of art,

maybe works from that time were made with media that deteriorated or perhaps

such works were removed by later humans.

While modern Homo sapiens lived across Africa, some members eventually left the

continent. Generations of these pioneers entered environments far different from

what their ancestors experienced in Africa. The next four sections cover evidence of

modern Homo sapiens in other parts of the Old World and the evidence we have

about what they did. We will check back with Africa later in the chapter to see what

happened biologically and culturally on the home front amid the expansion.

Expansion into the Middle East and Asia

This section presents key �nds showing where modern Homo sapiens went after the

range of the species �rst extended out of Africa. These pioneers could have used

two connections to the Middle East, or West Asia. From North Africa, they could

have crossed the Sinai Peninsula and moved north to the Levant, or eastern

Mediterranean. Finds in that region show an early modern human presence. Other

�nds support the Southern Dispersal model, with a crossing from East Africa to the

southern Arabian Peninsula through the Straits of Bab-el-Mandeb. It is tempting to

think of one momentous event in which people stepped off Africa and into the Mid-

dle East, never to look back. In reality, there were likely multiple waves of movement

producing gene �ow back and forth across these regions. The expanding modern

human population could have thrived by using resources along the southern coast of

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Figure 12.8 This Skhul V cranium model shows the sharp browridges. The contour of a marked occipital bun is barely visible from this angle.

the Arabian Peninsula to South Asia, with side routes moving north along rivers. The

maximum range of the species then grew across Asia as shown by evidence across

the continent.

Modern Homo sapiens in the Middle East

Geographically, the Middle East is the ideal place for the African modern Homo sapi-

ens population to inhabit upon expanding out of their home continent. In the East-

ern Mediterranean coast of the Levant, there is a wealth of skeletal and material cul-

ture linked to modern Homo sapiens. Recent discoveries from Saudi Arabia further

add to our view of human life just beyond Africa.

The Caves of Mount Carmel in present-day Israel

have preserved skeletal remains and artifacts of

modern Homo sapiens, the �rst-known group living

outside Africa. The skeletal presence at Misliya Cave

is represented by just part of the left upper jaw of

one individual, but it is notable for being dated to a

very early time, between 194,000 and 177,000 years

ago (Hershkovitz et al. 2018). Later, from 120,000 to

90,000 years ago, fossils of multiple individuals

across life stages were found in the caves of Es-

Skhul and Qafzeh (Shea and Bar-Yosef 2005). The

skeletons had many modern Homo sapiens traits,

such as globular crania and more gracile postcranial

bones when compared to Neanderthals. Still, there

were some archaic traits. For example, the adult male Skhul V also possessed what

researchers Daniel Lieberman, Osbjorn Pearson, and Kenneth Mowbray (2000)

called marked or clear occipital bunning. Also, compared to later modern humans,

the Mount Carmel people were more robust. Skhul V had a particularly impressive

brow ridge that was short in height but sharply jutted forward above the eyes (Fig-

ure 12.8). The high level of preservation is due to the intentional burial of some of

these people. Besides skeletal material, there are signs of artistic or symbolic behav-

ior. For example, the adult male Skhul V had a boar’s jaw on his chest. Similarly,

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Qafzeh 11, a juvenile with healed cranial trauma, had an impressive deer antler rack

placed over his torso (Figure 12.9) (Coqueugniot et al. 2014). Perforated seashells col-

ored with ochre, mineral-based pigment, were also found in Qafzeh (Bar-Yosef

Mayer, Vandermeersch, and Bar-Yosef 2009).

Figure 12.9 This cast of the Qafzeh 11 burial shows the antler’s placement over the upper torso. The forearm bones appear to overlap the antler.

One remaining question is, what happened to the modern humans of the Levant af-

ter 90,000 years ago? Another site attributed to our species did not appear in the

region until 47,000 years ago. Competition with Neanderthals may have accounted

for the disappearance of modern human occupation since the Neanderthal presence

in the Levant lasted longer than the dates of the early modern Homo sapiens. John

Shea and Ofer Bar-Yosef (2005) hypothesized that the Mount Carmel modern hu-

mans were an initial expansion from Africa but one that failed. Perhaps they could

not succeed due to competition with the Neanderthals who had been there longer

and had both cultural and biological adaptations to that environment.

Six-hundred kilometers from Mount Carmel, the fossil AW-1 from Al Wusta in Saudi

Arabia was just one �nger bone, but it greatly enhanced our view of modern Homo

sapiens just outside Africa. Dating methods converged on a range between 130,000

and 90,000 years ago, overlapping the Skhul and Qafzeh range (Groucutt et al. 2018).

The AW-1 bone and its associated stone tools added to evidence of many sites dot-

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Figure 12.10 The Liujiang cranium shows the tall forehead and overall gracile appearance typical of modern Homo sapiens.

ted throughout the Arabian Peninsula that contained stone tools but not skeletal

remains.

Modern Homo sapiens of China

A long history of paleoanthropology in China has found ample evidence of modern

human presence. Four notable sites are the caves at Fuyan, Liujiang, Tianyuan, and

Zhoukoudian. In the distant past, these caves would have been at least seasonal

shelters that unintentionally preserved evidence of human presence for modern re-

searchers to discover.

At Fuyan Cave in Southern China, paleoanthropologists found 47 adult teeth associ-

ated with cave formations dated to between 120,000 and 80,000 years ago (Liu et al.

2015). It is currently the oldest-known modern human site in China, though other

researchers question the validity of the date range (Michel et al. 2016). The teeth

have the small size and gracile features of modern Homo sapiens dentition. No lithics

have been found in Fuyan Cave.

The fossil Liujiang (or Liukiang)

hominin has derived traits that

classi�ed it as a modern Homo

sapiens, though primitive archaic

traits were also present. In the

skull, which was found nearly

complete, the Liujiang hominin

had a taller forehead than archaic

Homo sapiens but also had an en-

larged occipital region (Figure

12.10) (Brown 1999). A reconstruc-

tion of the brain based on the en-

docast of the cranium con�rmed

these trends along with a larger overall volume (Wu et al. 2008). Other parts of the

skeleton also had a mix of modern and archaic traits: for example, the femur frag- Previous: Archaic Homo

Next: Race and Human Variation

ments suggested a slender length but with thick bone walls (Woo 1959). Dating

methods suggested an age of around 67,000 years.

A mandible fragment, teeth, and postcranial skeletal remains of a single adult of in-

determinate sex was found by tree farmers in Tianyuan, 50 km from Beijing (Tong

2004). Radiocarbon dating of the bones estimated that they were from 42,000 to

39,000 years ago (Shang et al. 2007). As with other fossils described in this section,

researchers noted a few transitional traits between archaic and modern categories,

such as deep tooth measurements (the anteroposterior or front-to-back dimension)

and a robust tibia. The Tianyuan fossils also had some antemortem tooth loss (which

happened during life), osteoarthritis of a left-hand �nger joint, and enlargements to

muscle attachment sites of the tibia and femur. The evidence pointed to a physically

demanding life.

The last Chinese site to describe here is the one that has been studied the longest.

In the Zhoukoudian Cave system, where Homo erectus and archaic Homo sapiens

have also been found, there were three crania that �t the modern Homo sapiens set

of traits (Figure 12.11). These crania were in a part of the cave called the Upper Cave,

dating to between 34,000 and 10,000 years ago. The crania were all more globular

than that of archaic humans but still lower and longer than later modern humans’

(Brown 1999; Harvati 2009). When compared to one another, the three Upper Cave

crania showed signi�cant differences from one another. Comparison of cranial mea-

surements to other populations past and present found no connection with modern

East Asians. These �ndings again show that human variation was very different from

what we see today.

Other Asian Modern Humans

Other discoveries in Asia show us where modern Homo sapiens went after the initial

expansion. Sites with evidence of modern human occupation stretch from the island

of Sri Lanka north to Siberia. The �rst modern humans may have followed rivers

north to settle in colder regions while still accessing the rich freshwater

environment.

The Balangoda hominins refer to around 36 modern humans as far back as 38,000

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Figure 12.11 The entrance to the Upper Cave of the Zhoukoudian complex, where crania of three prehistoric modern humans were found.

years ago whose fossils were found in

numerous cave sites around Sri Lanka

(Kennedy et al. 1987). The name also

refers to one particularly well-studied

skeleton from the archaeological site of

Batadombalena. Measurements of

Balangoda Man show a closeness to the

modern-day Vedda people who live in

Sri Lanka, suggesting a direct ancestral

relationship. Ornamentation such as

pendants and beads, and the presence

of shark teeth far from the coast, sup-

ported the presence of modern behav-

ior as they were possibly transported for their aesthetic or symbolic value rather

than their practical use.

A double-infant burial dated to 28,000 years ago was found in 1928 at the site of

Mal’ta in southern Siberia, north of Mongolia (Raghavan et al. 2014). Researchers

named the three- to four-year-old individual MA-1. This burial was decorated with

Later Stone Age decorations: a beaded necklace, pendants, and a headband. Other

accessories and lithics were buried with the pair. Genetic analysis of MA-1 found a

connection with both present-day Western Europeans and Native Americans but

not East Asians. This �nding hints at the complex routes people took in the expan-

sion of the species.

Summary of Modern H. sapiens in the Middle East and Asia

As in Africa, the �nds of the Middle East have shown that humans were biologically

diverse and had complex relationships with their environment. Work in the Levant

showed an initial expansion north from the Sinai Peninsula that did not last. Away

from the Levant, expansion continued. People were present in Saudi Arabia, too, as

rainfall increased the amount of habitable land. Local resources were used to make

lithics and decorative items. Previous: Archaic Homo

Next: Race and Human Variation

The early Asian presence of modern Homo sapiens was complex and varied as be�t-

ting the massive continent. What the evidence shows is that people adapted to a

wide array of environments that were far removed from Africa. From the Levant to

Sri Lanka, Siberia, and China, humans with modern anatomy used caves that pre-

served signs of their presence. Faunal and �oral remains found in these shelters

speak to the �exibility of the human omnivorous diet as local wildlife and foliage be-

came nourishment. Decorative items, often found as burial goods in planned graves,

show a �ourishing cultural life.

Eventually, modern humans at the southeastern fringe of the geographical range of

the species found their way southeast until some became the �rst humans in

Australia.

Crossing to Australia

Expansion of the �rst modern human Asians, still following the coast, eventually en-

tered an area called Sunda by researchers before continuing on to modern Aus-

tralia. Sunda was a landmass made up of the modern-day Malay Peninsula, Sumatra,

Java, and Borneo. Lowered sea levels connected these places with land bridges,

making them easier to traverse. Proceeding past Sunda meant navigating Wallacea,

the archipelago that includes the Indonesian islands east of Borneo. The name refers

to naturalist Sir Alfred Russel Wallace, who noted that organisms from this region

differed from those to the west. Prehistorically, there were many megafauna, large

animals that migrating humans would have used for food and materials such as hides

and bones. Further southeast was another prehistoric landmass called Sahul, which

included New Guinea and Australia as one contiguous continent. This land had never

seen hominins or any other primates before modern Homo sapiens arrived. Sites

along this path offer clues about how our species handled these changes to the local

environment to live successfully as foragers.

While no fossil humans have been found at the Madjedbebe rock shelter in the

North Territory of Australia, more than 10,000 artifacts found there show both be-

havioral modernity and variability (Clarkson et al. 2017). They include a diverse array

of stone tools and different shades of ochre for rock art, including mica-based re-

Previous: Archaic Homo

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�ective pigment (similar to glitter). The ochre were shaped into what the re-

searchers called “crayons” to be held and used to mark other things. There were also

plant and animal remains matching the tools used to process them. One notable �nd

in this category is the partial upper jaw of a thylacine, or Tasmanian wolf, which was

colored red. These impressive artifacts date as far back as 56,000 years ago, provid-

ing the date for the earliest-known presence of humans in Australia.

The skeletal remains at Lake Mungo are the oldest known in the continent. The lake,

now dry, was one of a series located along the southern coast of Australia in New

South Wales, far from where the �rst people entered from the north (Barbetti and

Allen 1972; Bowler et al. 1970). Two individuals dating to around 40,000 years ago

show signs of artistic and symbolic behavior, including intentional burial. The bones

of Lake Mungo 1 (LM1), an adult female, were crushed repeatedly, colored with red

ochre, and even cremated (Bowler et al. 1970). Lake Mungo 3 (LM3), a tall older male

with a gracile cranium but robust postcranial bones, had his �ngers interlocked over

his pelvic region (Brown 2000).

Kow Swamp, also in southern Australia, contained human crania that looked dis-

tinctly different from the ones at Lake Mungo (Durband 2014; Thorne and Macum-

ber 1972). The Kow Swamp crania had extremely robust brow ridges and thick bone

walls, but these were paired with globular features on the braincase (Figure 12.12).

The frontal bones had extremely linear slopes from the brow to the top of the cra-

nium, resembling intentional cranial modi�cation seen in other parts of the world. If

the crania were shaped on purpose, they are another sign of symbolic behavior, as

the practice has linked to ideas of group cultural identity. By the time of the Kow

Swamp people, between 9,000 and 20,000 years ago, cranial modi�cation may have

been a meaningful part of culture in southern Australia.

Summary of Modern H. sapiens in Australia

The presence of the �rst humans in Australia along the current northern and south-

ern coasts suggests that they used a route that wrapped around the perimeter of the

continent. This path allowed access to both coastal and inland resources. Megafauna

was a likely source of food and other resources. The mythology of Australian aborig- Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.12 Replica of the Kow Swamp 1 cranium. The shape of the braincase could be due to arti�cial cranial modi�cation. A competing hypothesis is that it re�ects the primitive shape of Homo erectus.

ines today has been linked by re-

searchers to extinct life, such as mar-

supial tapirs and lions. Predation by

humans may be why the megafauna

became extinct, leaving the oral tradi-

tion of their existence.

The abundant evidence matching the

criteria for behavioral modernity

shows that the early Australians had a

rich artistic and symbolic life. Raw ma-

terials must have been transported or

traded across long distances in order

to make art and color both human and

nonhuman skeletal remains. The local varieties of stone tools and art may re�ect

cultural variation across distant regions of the continent.

The overall view of the �rst modern humans in Australia from a biological perspec-

tive shows a high amount of skeletal diversity. This is similar to the trends seen ear-

lier in Africa, the Middle East, and East Asia. While the Lake Mungo individuals had

derived gracile cranial traits, the Kow Swamp crania were measurably more robust.

Northwest to Europe

The �rst modern human expansion into Europe occurred after other members of

our species settled East Asia and Australia. As the evidence from the Levant sug-

gests, modern human movement to Europe may have been hampered by the pres-

ence of Neanderthals. Another obstacle was that the colder climate was incompati-

ble with the biology of African modern Homo sapiens, which was adapted for expo-

sure to high heat and ultraviolet radiation. Still, by 40,000 years ago, modern Homo

sapiens had enough of a presence in Europe to leave evidence for researchers to

�nd. This time was also the start of the Later Stone Age or Upper Paleolithic, with

an expansion in cultural complexity. Connected with the history of science in gen-

eral, early modern Homo sapiens in Europe have been studied for centuries. Due to

Previous: Archaic Homo

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the bias in research focus favoring Europe, there is a wealth of evidence to explore.

Still, there are also eye-opening discoveries in this area today. This section will

cover some of the key evidence of early modern human life in Europe, then go over

the typologies used to view the cultural changes in this region.

In Romania, the site of Peștera cu Oase (Cave of Bones) had the oldest known re-

mains of modern Homo sapiens in Europe, dated to around 40,000 years ago

(Trinkaus et al. 2003a). Among the bones and teeth of cave bears, wolves, ibex, and

other animals were the fragmented cranium of one person and the mandible of an-

other (the two bones did not �t each other). Both bones have modern human traits

similar to the fossils from the Middle East, but they also had Neanderthal traits.

Oase 1, the mandible, has a mental eminence but also extremely large molars

(Trinkaus et al. 2003b). This mandible has yielded DNA, opening another dimension

of study. Surprisingly, DNA from Oase 1 is equally similar to DNA from present-day

Europeans and Asians (Fu et al. 2015). This means that Oase 1 was not the direct an-

cestor of modern Europeans. The Oase 2 cranium has the derived traits of reduced

brow ridges along with archaic wide zygomatic cheekbones (Figure 12.13) (Rougier et

al. 2007). What the braincase shows is also between the two extremes: an overall

globular shape that had a tall but sloped frontal bone and an occipital bun-like pro-

trusion at the other end. No artifacts were found at this site. The assemblage was

likely gathered by either carnivores or geological events such as water action since

the Oase human bones were found with a high amount of nonhuman remains.

The term “Cro-Magnon” has entered public usage as a name for any prehistoric

modern European Homo sapiens, and maybe any “caveman” of our species, but it

technically refers to four adults (three male and one female) and an infant found in

the Cro-Magnon rock shelter in France in 1868 (Balzeau et al. 2013). The remains are

dated to 28,000 years ago and may all have been intentionally buried along with over

300 pierced seashells and nonhuman skeletal remains.The Cro-Magnon crania are

easily identi�able by their rectangular eye orbits, which are more angular than any

contemporary (Figure 12.14). Compared to Neanderthal skeletons of the same region,

the Cro-Magnons are extremely gracile. The adults also show signs of much pathol-

ogy, including fused neck vertebrae and healed fractures. The individual Cro-

Magnon 1 has skeletal lesions typical of neuro�bromatosis type 1, a rare genetic dis-

ease that causes tumor growth (Charlier et al. 2018). The combination of disease Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.13 This side view of the Oase 2 cranium shows the reduced brow ridges but also occipital bunning that is a sign that modern Homo sapiens interbred with Neanderthals.

Figure 12.14 This reconstruction of the Cro- Magnon 1 skull shows the gracility of modern Homo sapiens along with a disease that marked the bone.

markers suggest that life for the Cro-Magnons

was so physically demanding that it greatly af-

fected the skeleton.

Dating

to

aroun

d

26,00

0

years

ago,

Předm

ostí

near

Přerov

in the Czech Republic was a site where

people buried over 30 individuals along

with many artifacts. Eighteen individuals

were found in one mass burial area, a few

covered by the scapulae of woolly mammoths (Germonpré, Lázničková-Galetová,

and Sablin 2012). While the recovered human skeletons were destroyed in World

War II, �nely detailed photographic negatives allowed comparisons to other human

groups (Figure 12.15). The Předmostí crania were more globular than those of archaic

humans but tended to be longer and lower than in later modern humans

(Velemínská et al. 2008). The height of the face was in line with modern residents of

Central Europe. One standout trait seen on every mandible on this site was an un-

usually long length to the mandibular body and jutting chin, resulting in a particular

local appearance. Besides the human remains, the site contained the bones of over a

thousand mammoths. Some of the mammoth remains were shaped by humans, in-

cluding a limb bone fragment with a carved abstract female �gure. There is also

skeletal evidence of dog domestication, such as the presence of dog skulls with

shorter snouts than in wild wolves (Germonpré, Lázničková-Galetová, and Sablin

2012). In total, Předmostí could have been a settlement dependent on mammoths for

subsistence with people participating in artistic behaviors and the arti�cial selection

of early domesticated dogs.

Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.15 This illustration is based upon one of the surviving photographic negatives since the original fossil was lost in World War II. The modern human chin is prominent, as is an archaic occipital bun.

Upper Paleolithic European Material Culture

The sequence of modern Homo sapiens techno-

logical change in the Later Stone Age has been

thoroughly labeled and dated by researchers

working in Europe. The style associated with

the start of the Upper Paleolithic is the Aurig-

nacian, starting around 40,000 years ago and

ending around 27,000 years ago. Items in this

tradition include stone blades as well as beads

made from shell, bones, and teeth. Next is the

Gravettian, which lasted from 6,000 years to

21,000 years ago. This culture is associated

with most of the known curvy female �gurines,

often assumed to be “Venus” �gures. Hunting technology also advanced, such as

with the �rst known boomerang, atlatl (spear thrower), and archery. The Solutrean,

marked by further innovation in delicate tool work, is the following style from 21,000

to 17,000 years ago. After that time, the Magdalenian tradition spread. This culture

further expanded on �ne bone tool work, including barbed spearheads and �sh-

hooks (Figure 12.16). The end of the Magdalenian is also the end of the Later Stone

Age and the Pleistocene Period. While these labels and time spans apply to Europe,

other regions also showed changes in material culture to some of the same types of

technology. Uncovering the regional timelines of cultural styles around the world to

see these transitions on a global scale is an ongoing goal of paleoanthropologists.

Among the many European sites dating to the Later Stone Age, the famous cave art

sites deserve mention. Chauvet-Pont-d’Arc Cave in southern France dates to sepa-

rate Aurignacian occupations 31,000 years ago and 26,000 years ago. Over a hun-

dred art pieces representing 13 animal species are preserved. Some depicted species

are common to European cave art, such as deer and horses. Others are rare, such as

rhinos and owls. Two possible human �gures are in the deepest gallery of the cave

system. Besides the painted �gures, the tracks and skulls of cave bears and an ibex

were also found in the cave. Another famous French cave with art is Lascaux, which

is several thousand years younger at 17,000 years ago in the Magdalenian period. At

Previous: Archaic Homo

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Figure 12.16 This drawing from 1891 shows an array of Magdalenian- style barbed points found in the burial of a reindeer hunter. They were carved from antler.

this site, there are

over 6,000 painted

�gures on the walls

and ceiling (Figure

12.17). The paint was

made of a mix of

mineral pigments in

liquid binder made

from fat or clay.

Scaffolding and

lighting must have

been used to make

the paintings on the

walls and ceiling

deep in the cave.

Overall, visiting Las-

caux as a contempo-

rary must have been

an awesome experi-

ence: trekking

deeper in the cave

lit only by torches

giving glimpses of animals all around as mysterious sounds echoed through the gal-

leries. The professionally lit photographs of today do not give the original context

justice, though replicas have been built to simulate the experience for tourists. Both

Chauvet and Lascaux have been closed to all but researchers due to the degradation

of the art when tourism was allowed.

Previous: Archaic Homo

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Figure 12.17 Photograph of just one surface with cave art at Lascaux Cave. The most prominent piece here is the Second Bull, found in a chamber called the Hall of Bulls. Smaller cattle and horses are also visible.

Summary of Modern H. sapiens in Europe

Study of Europe in the Upper Paleolithic gives a more detailed view of the general

pattern of biological and cultural change linked with the arrival of modern Homo

sapiens. The modern humans experienced a rapidly changing culture that from our

perspective went through four major growths in complexity and re�nement. Skele-

tally, the increasing globularity of the cranium and the gracility of the rest of the

skeleton continued, though with unique regional traits, too. The cave art sites

showed a deeper use of expression and symbolism, though the exact meaning is un-

clear. With survival dependent on the surrounding ecology, painting the �gures may

have connected people to important and impressive wildlife at both a physical and

spiritual level. Both reverence for animals and the use of caves for an enhanced sen-

sory experience are common to cultures today and through recorded history.

In the next section we continue our exploration of Homo sapiens origins by seeing Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.18 This diagram shows the amount of DNA introgression between Neanderthals, Denisovans, and various regional lineages of modern Homo sapiens. The 6– 9% Neanderthal DNA found in the modern human Oase 1 was narrowed down to 1–2% in later modern humans. Denisovans share a few percent of their DNA with modern Australasians and Melanesians but just a fraction of a percent with modern East Asians.

the genetic evidence of interbreeding between the archaic and modern types, leav-

ing just the latter to continue to the present day.

Distant Relations: Interbreeding with Archaic Humans Outside Africa

As the modern human population grew beyond Africa, they interbred with the ar-

chaic Homo sapiens who were already there, descendants of the Homo erectus popu-

lations before them. This statement is different from what many people in the public

believe: that modern humans are the direct descendants of the archaic Nean-

derthals. Instead, the building evidence suggests a more complex connection be-

tween archaic and modern humans outside of Africa. This section describes the evi-

dence showing that three Homo sapiens groups interbred with one another: modern

Homo sapiens and two archaic groups, the Neanderthals and the Denisovans (Figure

12.18).

Interbreeding with Neanderthals

Since the �rst �nds of Nean-

derthal remains, researchers

have sought evidence of a bi-

ological connection with

modern Europeans. While

there are no sites that show

Neanderthals and modern

Homo sapiens lived together

(such as both types of skele-

tons found together in a com-

mon burial), interbreeding

has been suggested by the

Previous: Archaic Homo

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appearance of Neanderthal traits in otherwise modern human skeletons found in

their common geographical range. For example, the Oase 2 cranium from Peștera cu

Oase and the Skhul V cranium each had a partial occipital bun between the Nean-

derthal and modern extremes.

A new source of evidence arrived with the invention of ancient DNA (aDNA) analysis,

which has built up more evidence of Neanderthal and modern human interbreeding.

DNA samples from Neanderthal fossils have been compared to DNA from both pre-

historic and present-day modern Homo sapiens to trace the amount of gene �ow

between these groups. The amount of transfer is more indicative of introgression,

the entrance of small, uneven portions of Neanderthal DNA into modern humans,

rather than an even hybridization over time (Dannemann and Racimo 2018; Slatkin

and Racimo 2016). The introgression could have been caused by an imbalance in

population size: the continually growing modern population with gene �ow from

Africa could have diluted the incoming Neanderthal DNA to the low percentage seen

today. The time spent apart as separate lineages could have caused genetic incom-

patibility, especially in the Y chromosome (Mendez et al. 2016). Natural selection

may also have removed the inherited Neanderthal alleles if they were maladaptive,

leaving just the adaptive or neutral variants.

Continuing DNA analyses reveal more details about interbreeding. Geneticist Fu

Qiaomei and her team (2015) revealed long sections of Neanderthal DNA in Oase 1,

around 6%–9% of the total amount, suggesting a Neanderthal ancestor four to six

generations before. Fu also found continuous sections of Neanderthal DNA in the

DNA of Russian and Siberian modern humans from around 45,000 years ago, rein-

forcing the genetic evidence of interbreeding (Fu et al. 2014).

The �ndings of Neanderthal DNA analysis and comparison with modern human

genomes worldwide have revealed surprising details about the interactions between

these groups. An unexpected result is that modern Asians have more DNA from Ne-

anderthals than modern Europeans despite the separation in geography (Wall and

Yoshihara Caldeira Brandt 2016). Today, non-Africans have around 1%–2% Nean-

derthal DNA in their genomes, with Asians having more than Europeans. This is a

drop from the amount found in Oase 1. One explanation for this �nding is that the

interbreeding between Neanderthals and modern humans happened in the Middle

East before the population split into the modern European and Asian populations. Previous: Archaic Homo

Next: Race and Human Variation

Then Neanderthal DNA introgressed at least a second time just into the Asian popu-

lation, leaving more in that group.

Interbreeding with Denisovans

Comparison of DNA between the Denisovan archaic humans and modern humans

has also produced intriguing information about the interaction between these

groups. Denisovan DNA has also introgressed into some modern human populations

(Reich et al. 2010; Reich et al. 2011). In this case, neither modern Africans nor Euro-

peans have any Denisovan DNA. There is around 0.4% in modern East Asians. No-

tably, most modern Tibetans inherited a Denisovan allele that produces an adapta-

tion to high-altitude living (Huerta-Sánchez et al. 2014). The highest amount is in

some modern Melanesians and aboriginal Australians, between 4% and 6%. Though

the Denisovan sample came from the Altai Mountains in Siberia, the most likely lo-

cation for the interbreeding based on the DNA evidence was in East Asia or Sunda.

The ancestors of the Melanesians and Australians would have received the Deniso-

van DNA into their genomes there before their descendants expanded to their even-

tual destinations.

Summary of Archaic Human Genetic Introgression

While the study of skeletal traits suggested archaic-modern interbreeding, the use

of DNA analysis provided solid evidence of these events between archaic and mod-

ern Homo sapiens in Europe and Asia. This is a very active �eld as the technology

keeps improving and more samples are analyzed. A consistent picture is forming,

but many of the details concerning the timing and context of interbreeding are still

unclear. At this point, we know that three human groups interbred with each other

at different times. DNA originating from Neanderthals and Denisovans was found in

prehistoric modern humans and is present in some Europeans and Asians of today.

The introgressed DNA has affected the traits of our own species and they are just

being discovered. Previous: Archaic Homo

Next: Race and Human Variation

African Developments

We now switch our view back to Africa to see what developments occurred after

members of our species �rst crossed to the Middle East and beyond. Our survey of

modern Homo sapiens expansion left Africa around 150,000 years ago to see where

people pushed the fringes of our geographical range. It is important to remember

the species did not all leave Africa together. While modern humans found ways to

survive in the Middle East, Asia, Sahul, and Europe, many others remained in Africa.

Evidence of what Homo sapiens did in Africa from the end of the Middle Stone Age

to the Later Stone Age is concentrated in South African sites. There, Blombos Cave

and Border Cave show that complex human behavior was also developing in the

home continent.

Blombos Cave is located along the present shore of the Cape of Africa facing the In-

dian Ocean and is notable for having a wide variety of artifacts. The material culture

shows that toolmaking and artistry were more complex than previously thought for

the Middle Stone Age. Excavations and analysis have been carried out since the

1990s by a research team including Francesco d’Errico, Christopher Henshilwood,

and Marian Vanhaeren. In a layer dated to 100,000 years ago, researchers found two

intact ochre-processing kits made of abalone shells and grinding stones (Hen-

shilwood et al. 2011). Chemical analysis found the likely ingredients of the ochre-

based paint, including crushed bone, animal fat, and charcoal. More ochre frag-

ments, including some marked with notches, were found all throughout the site. The

team performed numerous analyses and experiments to show that perforated ma-

rine snail shell beads from 75,000 years ago were shaped by people using bone

points found in the cave (Figure 12.19) (d’Errico et al. 2005). Together, the evidence

shows that the Middle Stone Age occupation at Blombos Cave incorporated re-

sources from a variety of local environments into their culture, from caves (ochre),

open land (animal bones and fat), and the sea (abalone and snail shells). This com-

plexity shows a deep knowledge of the region’s resources and their use—not just for

survival but also for symbolic purposes.

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Figure 12.19 Examples of the perforated shell beads found in Blombos Cave, South Africa: (a) view of carved hole seen from the inside; (b) arrows indicate worn surfaces due to repetitive contact with other objects, such as with other beads or a connecting string; (c) traces of ochre; and (d) four shell beads showing a consistent pattern of perforation.

On the eastern coast of South Africa, Border Cave shows new African cultural devel-

opments at the start of the Later Stone Age. Paola Villa and colleagues (2012) identi-

�ed several changes in technology around 43,000 years ago. Stone tool production

transitioned from a slower, measured process to one that took less time to �nish but

made many microliths, small stone tools. An adhesive made from tree bark was

found on some of the microliths. The researchers hypothesize that hunting technol-

ogy moved from large crafted spearheads to smaller bone arrow points that were

adhered to shafts and even tipped with poison for more effectiveness. Changes in

decorations were also found across the Later Stone Age transition. Beads were made

from a new resource: fragments of ostrich eggs (d’Errico et al. 2012). Unlike the snail

shell beads, which retained the shape of the original structure, ostrich shell beads

were shaped into circular forms, resembling present-day breakfast cereal O’s. While Previous: Archaic Homo

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a subtle difference, these beads show a higher level of altering one’s own surround-

ings and a move from the natural to the abstract in terms of design.

Summary of Continuing Modern H. sapiens in Africa

African culture experienced a long constant phase called the Middle Stone Age until

a faster burst of change produced innovation and new styles. The change was not

one moment but rather a ramping up in development. Later Stone Age culture intro-

duced elements seen across many cultures, including the construction of composite

tools and even the use of strung decorations such as beads. These developments ap-

pear in the Later Stone Age of other regions, such as with the Balangoda of Sri Lanka

and the Aurignacian tradition of Europe, both mentioned above. Based on the early

date of the African artifacts, Later Stone Age culture may have originated in Africa

and passed from person to person and region to region, with people adapting the

general technique to their local resources and viewing the meaning in their own

way.

Unfortunately, information about modern humans in Africa from 40,000 to 12,000

years ago is scarce. In the next section, we will return to the expanding frontier as

the �rst hominins set foot in the Western Hemisphere.

Discovering the Americas

By 20,000 years ago, our species was the only member of Homo left on Earth. Gone

were the Neanderthals, Denisovans, and Homo �oresiensis. The range of modern

Homo sapiens kept expanding eastward into—using the name given to this area by

Europeans much later—the Western Hemisphere. This section will address what we

know about the peopling of the Americas, from the �rst entry to these continents to

the rapid spread of prehistoric Native Americans (referred to by researchers without

intentional insult as prehistoric Amerindians, Paleoindians, or Paleoamericans)

across its lush and varied environments. Previous: Archaic Homo

Next: Race and Human Variation

�e Changing Role of Beringia

Evidence points to a prehistoric land bridge called Beringia that allowed people to

cross from Asia to North America, just as expansion to Australia was made easier

with lowered sea levels that exposed Sunda and Sahul. Beringia connected what is

now northeastern Siberia with Alaska. What people did to cross this land bridge is

still being investigated. Currently there are two competing models for this event,

called the Ice-Free Corridor model and the Coastal Route model, though the latter

has been gaining intriguing evidence.

For most of the 20th century, the accepted theory was that prehistoric northeast

Asians (East Asians and Siberians) �rst expanded across Beringia inland through an

ice-free corridor between glaciers that opened into the western Great Plains of the

United States, just east of the Rocky Mountains, around 13,000 years ago (Swisher et

al. 2013). While life up north in the cold environment would have been harsh, migrat-

ing birds and an emerging forest might have provided sustenance as generations ex-

panded through this land (Potter et al. 2018). These residents would have used a

stone tool style that developed into the common Clovis style found later in North

America.

In recent decades, researchers accumulated evidence against the ice-free corridor

as the original path the �rst Native Americans took. For example, some archaeologi-

cal sites around the Americas date to a time before the corridor was open. While

one site with a date that contradicts the model could be dismissed as an error at

�rst, several more sites were found that brought more scrutiny to the Ice-Free Cor-

ridor model. The route between glaciers was available later, and was likely used at

that time, but there was already a more accessible path between the hemispheres.

The reconstruction of past geography, climate, and ecology led to the formation of

the Coastal Route model that explains how people reached the Americas through

Beringia. The new focus is the southern edge of the land bridge instead of its center:

About 16,000 years ago, members of our species expanded along the coastline from

northeast Asia, east through Beringia, and south down the Paci�c Coast of North

America while the inland was still sealed off by ice. Archaeologist K. R. Fladmark

(1979) brought the Coastal Route model into the archaeological spotlight and re-

searcher Jon M. Erlandson has been at the forefront of compiling support for this

Previous: Archaic Homo

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theory (Erlandson et al. 2015). Reconstructing the geography and climate of Beringia,

the coast would have been free of ice at least part of the year by 16,000 years ago,

earlier than when the ice-free corridor was completely opened. Studies of past ecol-

ogy found that the coastal route would have provided abundant plant and animal re-

sources for most of the path. Besides migrating birds, many useful �sh (e.g., salmon),

shell�sh, mammals (e.g., whales, seals, and otters), and plants (e.g., seaweed) would

have been available on the coast. A re�nement of this model called the

Kelp Highway hypothesis focuses on one particular ecosystem found just offshore

from Japan, north to Beringia, and south to Baja California. This addition states that

subsisting off of resources from kelp forests could have supported the rapid expan-

sion to the Americas and down the coast of the two continents.

Other lines of evidence are also compatible with the Coastal Migration model. One

indirect archaeological connection between prehistoric Japan and the Americas is a

certain style of stone tool. Coastal tanged or stemmed lithics, which are stone points

with a thin projection at the base, have been found in both of these distant regions

(Erlandson and Braje 2011). The similarity could indicate a cultural tie. Genetic analy-

sis of Native American DNA also shows shared ancestry with northeast Asians, link-

ing them biologically as well (Raghavan et al. 2014).

While many factors such as reconstructions of climate and ecology indirectly sup-

port the Coastal Migration model, the search continues for direct evidence such as

archaeological sites along the route. Due to the warming trend since 18,000 years

ago that reduced glaciers and raised the sea level, much of the prehistoric coast that

would have been occupied by the �rst coastal migrants to North America is cur-

rently over 100 meters underwater (Erlandson et al. 2015).

Researchers are also still determining how many large waves of people made the

crossing through either Beringian route. A four-�eld analysis of modern Native

American languages found evidence of three migration events, resulting in three

major language groups (Greenberg 1987). Analyzing the DNA of prehistoric and mod-

ern peoples, which was not possible in Greenberg’s time, researchers found evi-

dence for one large wave of gene �ow from Beringia, with major splits once in North

America (Raghavan et al. 2015). Smaller waves after the main one could also have

moved from Beringia to North America. As far back as 23,000 years ago, people liv-

ing in Beringia would have had lowered gene �ow with northeast Asians and also

been unable to expand further east. This concept of a period of genetic isolation

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based on DNA analysis is called the Beringia Standstill Model. It explains the

amount of genetic differences between Native Americans and northeast Asians

within the time frame of the other evidence.

South through the Americas

However the �rst modern Homo sapiens reached the Western Hemisphere, the

spread through the Americas was rapid. Multiple migration waves crossed from

North to South America (Posth et al. 2018). Our species took advantage of the lack of

hominin competition and the bountiful resources both along the coasts and inland.

The Americas had their own wide array of megafauna, which included woolly mam-

moths, mastodons, camels, horses, ground sloths, giant tortoises, and—a favorite of

researchers—a two-meter-tall beaver (Figure 12.20). The reason we cannot �nd

these amazing animals today may be that ancient Native Americans hunted them all

to extinction. Resources gained from these fauna must have been an important part

of survival for people over 12,000 years ago (Araujo et al. 2017). Several sites are no-

table for what they add to our understanding of American prehistory, including in-

teractions with megafauna and other elements of the environment.

Monte Verde is a landmark site that shows that the human population had expanded

down the whole vertical stretch of the Americas to Chile by 14,600 years ago, only a

few thousand years after humans �rst entered the Western Hemisphere from

Alaska. The site has been excavated by archaeologist Tom D. Dillehay and his team

(2015), revealing fragile material culture that is rarely preserved, including human

footprints, animal hides, and wooden tools. Two of the discoveries at Monte Verde

relate to the Coastal Migration model. The discovery of nine edible species of sea-

weed at the site shows familiarity with coastal resources that might have been

passed down through generations of experience living near the ocean. A stemmed

point, reminiscent of the coastal styles, was also among the lithics at Monte Verde

(Figure 12.21).

Named after the town in New Mexico, the Clovis stone tool style is the �rst example

of a widespread culture across much of North America, between 13,400 and 12,700

years ago (Miller, Holliday, and Bright 2013). Instead of a stem-shaped base, Clovis

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Figure 12.20 Lifesize reconstruction of a woolly mammoth at the Page Museum, part of the La Brea Tar Pits complex in Los Angeles, California. Outside of Africa, megafauna such as this went extinct around the time that humans entered their range.

Figure 12.21 A stemmed point (left) and drill fragment (right) found in the same level at Monte Verde. The stemmed point resembles a coastal Andean style called the Paiján and may be evidence supporting the Coastal Route model.

points were �uted with two small pro-

jections, one on each end of the base,

facing away from the head (Figure

12.22). The stone points found at this

site match those found as far as the

Canadian border and northern Mexico,

and from the west coast to the east

coast of the United States. Fourteen

Clovis sites also contained the remains

of mammoths or mastodons, suggest-

ing that hunting megafauna with these

points was an important part of life for

the Clovis people. Other Clovis sites

show that other types of hunting and

gathering were important to people’s

subsistence, too. After the spread of

the Clovis style, it diversi�ed into sev-

eral regional styles, keeping some of

the Clovis form but also developing

their own unique touches.

Only one site has a human burial containing Clovis tools: Anzick in western Mon-

tana. The individual, Anzick-1, is a male infant dated to 12,800 years ago (Rasmussen

et al. 2014). He was buried with over 100 Clovis stone and bone tools that were

coated in ochre. Genetic analysis found that Anzick-1’s people were related to all

later Native Americans, proving a direct ancestral connection and supporting the

model of one large wave of migrants populating the Americas with ancient humans.

Summary of Modern H. sapiens in the Americas

Research in prehistoric Native American origins found some surprising details, re-

�ning older models. Genetically, the migration can be considered one long period of

movement, with splits into regional populations. This �nding matches the sudden

appearance of the homegrown Clovis culture, its rapid expansion, and the radiation

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Figure 12.22 Compared to the stemmed point in Figure 12.21, this Clovis point has a drastically different structure. The Clovis point has a wider tip and the base has two small projections instead of a single large stem. This example was carved from chert and found in north central Ohio, dated to around 11 kya.

of descendant cultures in North America. A few thousand

years after arrival into the hemisphere, people had already

covered the Americas from north to south.

The peopling of the Americas also had a lot of common ele-

ments with the prior spread of humans across Africa, Eu-

rope, Asia, and Australia. In all of these expansions, people

explored new lands that tested both the cultural and biologi-

cal adaptations of the pioneers. Besides stone tool technol-

ogy, the use of ochre as decoration was seen from South

Africa to South America. The coasts and rivers were likely av-

enues in the movement of people, artifacts, and ideas, out-

lining the land masses while providing access to varied envi-

ronments. The presence of megafauna aided human success,

but this resource was eventually depleted in many parts of

the world.

With our tracing of human expansion across the continents

complete, we will see how researchers visualize what we

learned about the origin and dispersal of modern Homo sapi-

ens from 315,000 to 12,000 years ago.

�e Big Picture: �e Assimilation Model

How do researchers make sense of all of these modern Homo sapiens discoveries

that cover over 300,000 years of time and stretch across every continent except

Antarctica? How was modern Homo sapiens related to archaic Homo sapiens? Over

the past few decades, paleoanthropologists have engaged in spirited debates based

on their interpretation of the data. In the mid- to late 20th century, scientists had

split into two competing views. This section describes this episode of paleoanthro-

pology history and how continuing scienti�c research improves our view of the

world.

One competing model was called the Out of Africa model  or Recent African Origin

model. Supporters of this model saw evidence that modern Homo sapiens �rst

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evolved in Africa, then expanded into the other continents without interaction with

the archaic Homo sapiens of Europe and Asia (Stringer and Andrews 1988). Re-

searchers on this side noted that the oldest modern Homo sapiens fossils were found

in Africa, suggesting that that continent was the origin. Genetic analysis found the

same conclusion.

The other model was called Multiregionalism or the Multiregional Continuity

model. The view of the data by scientists on this side was that modern Homo sapiens

evolved from the archaic humans in Africa, Europe, and Asia simultaneously (Wolpoff

1989). Gene �ow would have kept the species cohesive across the great distance

while producing local variation as well. The multiregionalist experts pointed to the

sharing of traits from Homo erectus, through archaic Homo sapiens, and then to the

modern humans in different parts of the world as supporting their model. For exam-

ple, in Europe the Oase modern humans possessed Neanderthal-like traits and cer-

tain modern humans possess alleles that came from Neanderthals and Denisovans.

Eventually, researchers noticed that both the Out of Africa model and the Multire-

gionalism model had elements that were supported by data and elements that were

not supported. Taking the supported parts of each model and combining them

formed an explanation that was more complicated, but explained much more of the

scienti�c evidence. The merging of two models to form a better one suits the name

of the improved version: the Assimilation model.

The Assimilation model proposes that modern Homo sapiens evolved in Africa �rst

and expanded out (from the Out of Africa model) but also interbred with the archaic

Homo sapiens they encountered outside Africa (from the Multiregionalism model)

(Figure 12.23). True multiregionalism occurred just within Africa as the species

evolved from a web of interactions between varied groups (Scerri et al. 2018). As the

modern human population expanded from Africa, they assimilated the alleles of ar-

chaic humans they encountered through interbreeding. The Assimilation model is

powerful since it explains why Africa has the oldest modern human fossils, why early

modern humans found in Europe and Asia bear a resemblance to the regional ar-

chaics, and why traces of archaic DNA can be found in our genomes today (Smith et

al. 2017).

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Figure 12.23 This diagram depicts the connections between archaic and modern Homo sapiens of different regions. Having evolved from Homo erectus, the archaic humans expanded from Africa and established the Neanderthal and Denisovan groups. In Africa, the remaining archaic humans evolved the modern set of traits and expanded from the continent as well, encountering and interbreeding with two archaic groups across Europe and Asia.

While scienti�c progress has produced a model that satis�es the data, there are still

a lot of questions for paleoanthropologists to answer regarding our origins. What

were the patterns of migration in each part of the world? Why did the archaic hu-

mans go extinct? In what ways did archaic and modern humans interact? How large

were the past populations? How did biological, cultural, and environmental factors

in�uence the material culture found in different parts of the world? The de�nitive

explanation of how our species started and what our ancestors did is still out there

to be found. You are now in a great place to welcome the next discovery about our

distant past—maybe you’ll even contribute to our understanding as well. Previous: Archaic Homo

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Figure 12.24 Screenshot of an action scene from the videogame Far Cry Primal. While the game is set at 12 kya, one faction of humans has distinctly Neanderthal traits (center-left), though none were left by that time in reality. Note touches inspired by research, such as the use of shells for decoration on the protagonist’s wrist.

Special Topics: “Cavemen” in Popular Culture

“Cavemen,” or our prehistory in general, is a constant presence in west-

ern popular culture. From the iconic opening to 2001� A Space Odyssey to

the less iconic 10,000 B.C., the distant past is a common setting for dra-

matic stories. The prehistoric experience even gets interactive with

games like Far Cry Primal where the player can persistence-hunt for food

and ride a woolly mammoth (!) (Figure 12.24). The distant past has also

been the setting for more comedic stories, such as The Flintstones and,

recently, The Croods with Nicolas Cage.

Why are we interested in the past beyond a scienti�c understanding?

Like feudal times or the Wild West, prehistory may be a setting that is rife

for telling stories that excite us. With humanity stripped of the modern

conveniences we are used to, situations become more intense. Prehistory

takes this trope to the extreme. The viewer/game player is invited to

think about what they would do if they had to live more directly off of the

environment, unshielded from the dangers of the natural world. Experi- Previous: Archaic Homo

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encing our imaginings of that time may satisfy parts of our brain that

evolved in that situation, just as people accustomed to urban life �nd en-

joyment by camping or sitting around a �re with friends and family. See-

ing the portrayal of the prehistoric world may reach those parts of our

psychology that feel at home away from our constructed environment.

That said, the portrayal of the past in �ction is rarely accurate. There is a

signi�cant lag time between the scienti�c view of our past and when it

reaches mass media. Older works age very poorly. In Clan of the Cave

Bear, a book from 1980, the Neanderthals have limited emotions and

communicate with sign language rather than speaking: two ideas that

have not stood the test of time but were core to the story. Even in recent

productions, prehistoric people are dim-witted and have limited vocabu-

lary, though the evidence shows that our brains and language abilities are

basically the same today as 300,000 years ago. How our media choose to

represent the past may say more about our own values and views of our-

selves rather than what people were actually like long ago.

As you engage in the next �ction set in our distant past, consider how the

presentation differs from the scienti�c view and why there is this divide.

Also, as you enjoy the world that is presented to you, consider that we are

not so different from the cavemen. We are the ones born into the world

of projectors and streaming media, and we may build on that to create

the next medium, but at our core we are the same as the painters of Las-

caux and the carvers of prehistoric �gurines.

THE CHAIN REACTION OF AGRICULTURE

While it may be hard to imagine today, for most of our species’ existence we were

nomadic: moving through the landscape without a singular home. Instead of a re-

frigerator or pantry stocked with food, we procured nutrition and other resources

as needed based on what was available in the environment. Instead of collecting and

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displaying stuff, we kept our possessions at a minimum for mobility. This part gives

an overview of how this foraging lifestyle enabled the expansion of our species, then

describes the invention of a new way of life, causing a chain reaction of cultural

change taking us to the present day and beyond.

�e Foraging Tradition

To understand our species is to understand foraging, or the search for resources in

the environment. This subsistence strategy, or method of �nding sustenance,

sounds unusual to most of us today. Most of us live in cultures that practice another

strategy, agriculture, where we shape the environment to mass produce what we

need. Considering the age of modern Homo sapiens, however, we have spent far

more time as nomadic foragers than as settled agriculturalists. As such, our traits

have evolved to be primarily geared toward foraging. For instance, our ef�cient

bipedalism allows persistence-hunting across long distances as well as movement

from resource to resource. Even our psychological tendency toward our ability to

form stable relationships with around 150 people (Dunbar 1993) may derive from the

foraging lifestyle.

How does human foraging, also known as hunting and gathering, work? Anthropolo-

gists have used all four �elds to answer this question (see Ember n.d.). Typically,

people formed bands, or groups of around 50, and rarely over 100. A band’s organi-

zation would be egalitarian, with a �exible hierarchy based on an individual’s age,

level of experience, and relationship with others. Everyone would have a general

knowledge of the skills assigned to their gender roles, rather than specializing in dif-

ferent occupations. A band would move from place to place in the environment, us-

ing knowledge of the area to hunt and gather (Figure 12.25). While there were excep-

tions, women typically gathered plants and hunted small animals while men hunted

larger prey where present (Waguespack 2005). The ratio of plant to meat in one’s

diet would have depended on the local resources. As a location’s resources became

used up, and as human waste accumulated, the band would travel to another patch

(Venkataraman et al. 2017). In the varied environments that humans entered—from

savannas to tropical forests, deserts, coasts, and the Arctic circle—people found sus-

tenance needed for survival. Our species’s omnivorous and cultural ability led us to

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Figure 12.25 A present-day San man in Namibia demonstrates hunting using archery. Anthropologists still study the San today to learn about the foraging lifestyle in Africa.

excel in the generalist-specialist niche. People could have temporarily altered their

environment to be more productive, such as by burning foliage to spur new growth.

Besides food sources, people would have known the local areas to �nd rock and

wood suitable for tool production, and ochre for decoration. Bands could have

formed trading connections to acquire goods from distant areas. Certain sites could

have been gathering spots for local bands to trade, socialize, and worship, though

they were not typically large permanent settlements.

Humans made extensive use of the forag-

ing subsistence strategy, but this lifestyle

did have limitations. The ease of foraging

depended on the richness of the environ-

ment. Due to the lack of storage, resources

had to be dependably found when needed.

While a bountiful environment would re-

quire just a few hours of foraging a day, the

level and duration of labor increased

greatly in poor or unreliable environments.

Labor was also needed to process the ac-

quired resources, contributing to �lling the

foragers’ daily schedule (Crittenden and Schnorr 2017).

The adaptations to foraging found in modern Homo sapiens may explain why our

species became so successful both within Africa and in the rapid expansion around

the world. Overcoming the limitations, each generation at the edge of our species’s

range would have found it bene�cial to expand a little further, keeping contact with

other bands but moving into unexplored territory where resources were richer. The

cumulative effect would have been the spread of modern Homo sapiens across con-

tinents and hemispheres.

Why Agriculture?

After hundreds of thousands of years of foraging, some groups of people around

12,000 years ago started to practice agriculture instead. This transition is called the Previous: Archaic Homo

Next: Race and Human Variation

Neolithic Revolution, and it occurred at the start of the Holocene epoch. The rea-

sons for this global change are still being investigated, but there are two likely

causes that may have occurred together: a growing human population and natural

global climate change.

Overcrowding could have affected the success of foraging in the environment, lead-

ing to the development of a more productive subsistence strategy (Cohen 1977). For-

aging works the best with low population densities since each band needs a lot of

space to support itself. If too many people occupy the same environment, they

would deplete the area faster. The high population could exceed the

carrying capacity, or number of people a location can reliably support. For instance,

what if a band arrived at a grove of nutritious plants they were depending on, but it

had already been used by other groups? Then the late arrivals are suddenly in a dire

situation without the food they were depending on �nding. This situation on a

global level due to growing population and limited areas of expansion would have

been an increasingly pressing issue after the human expansion through the major

continents by 14,600 years ago.

A changing global climate immediately preceded the transition to agriculture, so re-

searchers have also explored a connection between the two events. Since the

Last Glacial Maximum of 23,000 years ago, the Earth slowly warmed. At 13,000

years ago, the temperature in most of the Northern Hemisphere dropped suddenly

in a phenomenon called the Younger Dryas. Glaciers returned in Europe, Asia, and

North America. In Mesopotamia, which includes the Levant, the climate changed

from warm and humid to cool and dry. The change would have occurred over

decades, disrupting the usual nomadic patterns and subsistence of foragers around

the world. The Younger Dryas lasted until 11,700 years ago, when the climate re-

turned to the long-term warming pattern. The disruption to foragers due to the

temperature shift could have been a factor in spurring the transition to agriculture.

Researchers Gregory K. Dow and colleagues (2009) believe that foraging bands

would have clustered in the new resource-rich places where people started to direct

their labor to farming the limited area. Continued practice would prompt innova-

tions such as better tools and increasing productivity. After the Younger Dryas

ended, people expanded out of the clusters. As they reinhabited the region, they

brought with them a culture in which farming had become the norm, along with the

technology and knowledge to succeed with this subsistence strategy (Figure 12.26).

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Figure 12.26 Rice farmers in the present day using draft cattle to prepare their �eld.

The double threat of the limita-

tion of human continental ex-

pansion and the sudden global

climate change may have placed

bands in peril as more popula-

tions outpaced their

environment’s carrying capac-

ity. Not only had a growing

population led to increased

competition with other bands,

but environments worldwide

shifted to create more uncer-

tainty. As people in different ar-

eas around the world faced this chaotic situation, they became the independent in-

ventors of agriculture.

Agriculture around the World

Due to global changes to the human experience starting from 12,000 years ago, cul-

tures with no knowledge of each other turned toward farming their local resources

(Figure 12.27). The switch to agriculture took time and effort with no guarantee of

success. Agriculture is a dif�cult process with �res, �oods, droughts, disease, and

pests being constant problems to address. Heavy physical labor with no immediate

payoff was also needed to shape the landscape in a coordinated way to support agri-

culture. For example, people had to direct water �ow to irrigate constructed �elds

of crops. The �rst farmers also engaged in arti�cial selection of their domesticates

to enhance useful traits. The biggest success stories in the face of these obstacles

became the primary centers of agriculture (Figure 12.27) (Fuller 2010):

Mesopotamia: The Fertile Crescent from the Tigris and Euphrates rivers

through the Levant was where bands started to domesticate plants and animals

around 12,000 years ago. The connection between the development of agricul-

ture and the Younger Dryas was especially strong here. Farmed crops included Previous: Archaic Homo

Next: Race and Human Variation

wheat, barley, peas, and lentils. This was also where cattle, pigs, sheep, and

goats were domesticated.

South and East Asia: Multiple regions across this land had varieties of rice, mil-

let, and soybeans by 10,000 years ago. Pigs were farmed with no connection to

Mesopotamia. Chickens were also originally from this region, bred for �ghting

�rst and food second.

New Guinea: An under-appreciated center in Melanesia, agriculture started

here 10,000 years ago. Bananas, sugarcane, and taro were native to this island.

Sweet potatoes were brought back from voyages to South America around the

year C.E. 1,000. No known animal farming occurred here.

Mesoamerica: Agriculture from Central Mexico to northern South America also

occurred from 10,000 years ago; it was also only plant based. Maize was a crop

bred from teosinte grass, which has become one of the global staples. Beans,

squash, and avocados were also grown in this region.

The Andes: Starting around 8,000 years ago, local domesticated plants started

with squash but later included potatoes, tomatoes, beans, and quinoa. Maize

was brought down from Mesoamerica to join the local variety. The main farm

animals were llamas, alpacas, and guinea pigs.

Sub-Saharan Africa: This region went through a change 5,000 years ago called

the Bantu expansion. The Bantu agriculturalists were established in West Cen-

tral Africa and then expanded south and east. Native varieties of rice, yams, mil-

let, and sorghum were grown across this area. Cattle were also domesticated

here.

Eastern North America: This region was the last major independent agriculture

center, from 4,000 years ago. Squash and sun�ower are the produce from this

region that are most known today, though sumpweed and pitseed goosefoot

were also farmed. Hunting was still the main source of animal products.

By 5,000 years ago, our species was well within the Neolithic Revolution. From the

primary centers, agriculturalists spread to neighboring parts of the world with their

domesticates, further expanding the use of this subsistence strategy. For example,

the Mesopotamian farmers spread their innovations along the northern coast of the

Mediterranean into Europe (Pinhasi, Fort, and Ammerman 2005). The crops of China

were brought into the Korean peninsula (Lee 2011). From this point, the human

species changed from being primarily foragers to primarily agriculturalists. The rev-

olution took millennia, but it was a true revolution in that the lifestyle of our species

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Figure 12.27 Map showing the areas where agriculture was independently invented around the world. The dotted line around sub-Saharan Africa represents a possible range that is still being narrowed by research. Blue arrows show the spread of agriculture from these zones to other regions.

was reshaped

to something

vastly

different.

Cultural E�ects of Agricultu re

The world-

wide adoption of agriculture altered the course of human culture and history for-

ever. The foraging lifestyle was incompatible with agriculture, so dependence on the

latter required huge changes in how people lived. This section starts by following

the human developments that occurred due to agriculture, leading us from 12,000

years ago to the present day. We will also reconnect with modern foragers to see

how their lives have changed.

The core change in human culture due to agriculture is the move toward not mov-

ing: rather than live a nomadic lifestyle, farmers had to remain in one area to tend to

their crops and livestock. The term for living bound to a certain location is

sedentarism. Remaining in one place led to aspects of life that were uncommon in

foragers: the construction of permanent shelters and agricultural infrastructure

such as �elds and irrigation, plus the development of storage technology such as

pottery to preserve extra resources in case of future instability.

The high productivity of successful agriculture sparked further changes (Smith

2009). Since successful agriculture produced a much greater amount of food and

other resources per unit of land compared to foraging, the population growth rate

skyrocketed. The surplus of a bountiful harvest also provided insurance for harder

times, reducing the risk of famine. Changes happened to society as well. With a few

farming households producing enough food to feed many others, other people could

focus on other tasks. So began specialization into different occupations such as

craftspeople, traders, religious �gures, and artists, spurring innovation in these ar-

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Figure 12.28 View of downtown San Diego taken by the author at a shopping complex during a break from jury duty. Here, people live amongst structures that facilitate commerce, government, and art.

eas as people could now devote time and effort toward speci�c skills. These interde-

pendent people would settle an area together for convenience, causing a rise in the

number of dense populations focused around farms, water, and trade routes. The

growth of these settlements led to urbanization, the founding of cities that became

the foci of human interaction.

The formation of cities led to new issues

that sparked the growth of further special-

izations, called institutions. These were

cultural constructs that existed beyond the

individual and had wide control over a pop-

ulation. Leadership of these cities became

hierarchical with different levels of rank

and control. Laws in�uenced the behavior

of citizens, establishing ideal behavior and

punishment for deviations. Organized reli-

gion also kept followers under a standard

set of beliefs and values tied to spirituality.

Under leadership, people built impressive

monumental architecture such as pyramids that embodied the wealth and power of

these early cities. Alliances could unite cities, forming the earliest states. In several

regions of the world, state organization expanded into empires, wide-ranging politi-

cal entities that covered a variety of cultures.

Urbanization brought new challenges as well. The concentration of sedentary peo-

ples was ideal for infectious diseases to thrive since they could jump from person to

person and even from livestock to person (Armelagos, Brown, and Turner 2005). Ur-

ban life also caused sanitation problems as human waste accumulated, adding to the

spread of disease. While successful agriculture provided a large surplus of food to

thwart famine, the variety of food produced was smaller than what foragers experi-

enced (Cohen and Armelagos 1984; Cohen and Crane-Kramer 2007). The depen-

dence on high-yield crops also caused an overabundance of carbohydrates in the

diet of agriculturalists. This shift in nutrition caused another set of diseases to �our-

ish among those who adopted farming as their subsistence strategy: dental issues

such as dental caries (the cavities that ruin your visit to the dentist) and

malocclusion (the misalignment of teeth caused by the softness of agricultural di-

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ets). The issues with “wisdom teeth” or third molars seen in agricultural cultures to-

day stems from this misalignment between the environment our ancestors adapted

to and our lifestyles today.

As the new disease trends show, the adoption of agriculture and the cultural

changes that followed were not entirely positive. It is also important to note that

this is not an absolutely linear progression of human culture from simple to complex

and that higher complexity is not necessarily better than lower complexity. In many

cases empires have collapsed and cities have dispersed to low-density bands that no

longer saw use in maintaining institutions. However, a global trend has emerged

since the adoption of agriculture, wherein population and complexity have in-

creased, leading to the massive and in�uential nation states of today.

The rise of states in Europe has a direct impact on many of this book’s topics. Sci-

ence started as a European cultural practice by the upper class that became a stan-

dardized way to study the world. Education became an institution to provide a stan-

dardized path toward producing and gaining knowledge. The scienti�c study of hu-

man diversity, embroiled in the race concept that still haunts us today, was con-

nected to the European slave trade and colonialism.

Also starting in Europe, the Industrial Revolution of the 19th century turned cities

into centers of mass manufacturing and spurred the rapid development of inven-

tions. In the technologically interconnected world of today, human society has

reached a new level of complexity with globalization. In this system, goods are mass

produced and consumed in different parts of the world due to worldwide economic

factors. Instead of relying on local farms and factories, we now receive our everyday

goods from all over the world.

As states based on agriculture and industry keep exerting in�uence on humanity to-

day, there are people who continue to live a foraging—or mostly foraging—lifestyle.

Due to the overwhelming force that agricultural societies could exert, foragers to-

day have been marginalized to live in the least habitable parts of the world (Head-

land et al. 1989). These are places that are the worst for farmland, such as tropical

rainforests, deserts, and the arctic. Foragers can no longer live in the abundant envi-

ronments that humans would have enjoyed before the Neolithic Revolution. Interac-

tions with agriculturalists are typically imbalanced, with trade and other exchanges

heavily favoring the larger group. One of anthropology’s important roles today is to

Previous: Archaic Homo

Next: Race and Human Variation

Figure 12.29 This combine harvester can collect and process grain at a massive scale. Our food now commonly come from enormous farms located around the world.

Figure 12.30 Will we evolve toward even more globular brains? Actually, this trend is not likely to continue for our species.

intelligently and humanely manage interactions

between people of different backgrounds and

levels of in�uence.

�e Future of Humanity

This chapter covered what modern Homo sapi-

ens has done to get to the present time, but

what will our species do far in the future? Just

as biological changes accumulated from over

300,000 years ago to today, what will human traits and genetics be like? When

posed with these questions, people tend to think of directional selection. Maybe our

braincases will be even larger, resembling the large-headed and small-bodied aliens

of science �ction (see �gure 12.30. Or, our hands could be specialized for interacting

with our touch-based technology with less risk of repetitive injury. These ideas do

not stand up to scrutiny. Since natural selection is based on adaptations that in-

crease reproductive success, any directional change must be due to a higher rate of

producing successful offspring compared to other alleles. Larger brains and more

agile �ngers would be convenient to possess, but they do not translate into an in-

crease in the underlying allele frequencies.

Scientists are hesitant to professionally

speculate on the unknowable, and we to-

day will never know what is in store for

our species a thousand or a million years

from now, but there are trends in human

evolution that may carry on into the fu-

ture. These trends are increased genetic

variation and a reduction in regional

differences.

Rather than a directional change, genetic variation in our species could expand. Our

technology can protect us from extreme environments and pathogens, even if our

biological traits are not tuned to handle these stressors. The rapid pace of techno- Previous: Archaic Homo

Next: Race and Human Variation

logical advancement means that biological adaptations will become less and less rel-

evant to reproductive success, so non-bene�cial genetic traits will be more likely to

remain in the gene pool. Biological anthropologist Jay T. Stock (2008) views environ-

mental stress as needing to defeat two layers of protection before affecting our ge-

netics. The �rst layer is our cultural adaptations. Our technology and knowledge can

cover for many of our biological issues, reducing pressure on one’s genotype to be

just right to pass to the next generation. The second defense is our �exible physiol-

ogy, such as our functional adaptations. Only stressors not handled by these power-

ful responses would then cause natural selection on our alleles. These shields are al-

ready substantial, and cultural adaptations will only keep increasing in strength.

The increasing ability to travel far from one’s home region means that there will be a

mixing of genetic variation on a global level in the future of our species. In recent

centuries, gene �ow of people around the world has increased, creating admixture

in populations that had been separated for tens of thousands of years. For skin color,

this means that populations all around the world could exhibit the whole range of

skin colors, rather than the current pattern of decreasing melanin pigment farther

from the equator. The same trend of intermixing would apply to all other traits, such

as blood types. While our genetics will become more varied, the variation will be

more intermixed instead of regionally isolated.

Our distant descendants will not likely be dextrous ultra-intellectuals; more likely,

they will be a highly variable and mobile species. They will be supported by cultural

adaptations that we cannot even imagine, making up for any biological limitations

that keep getting passed to the next generation. Technology may even enable the

editing of DNA directly, changing these trends. With the uncertainty of our future,

these are just the best educated guesses for now. Our future is open and will be

shaped little by little by our actions and those of our descendants.

CONCLUSION

Modern Homo sapiens is the species that took the hominin lifestyle the furthest to

become the only living member of that lineage. This last section of the chapter sum-

marizes what we know about modern Homo sapiens traits, origins, and history.

Previous: Archaic Homo

Next: Race and Human Variation

The largest factor that allowed us to persist while other hominins went extinct was

likely our advanced ability to culturally adapt to a wide variety of environments. Our

species, with its skeletal and behavioral traits, was well suited to be generalist-spe-

cialists who successfully foraged across most of the world’s environments. The bio-

logical basis of this adaptation was our reorganized brain that facilitated innovation

in cultural adaptations and intelligence for leveraging our social ties. As the brain’s

ability increased, it shaped the skull by reducing the evolutionary pressure to have

large teeth and robust cranial bones to produce the modern Homo sapiens face.

Our ability to be generalist-specialists is seen in the geographical range that modern

Homo sapiens covered in 300,000 years. In Africa, our species formed from multire-

gional gene �ow that loosely connected archaic humans across the continent. Peo-

ple then expanded out to the rest of the continental Old World and even further to

the Americas. Wherever people went, they were enabled and connected by the

shared tools and art they crafted.

For most of our species’s existence, foraging was the general subsistence strategy

within which people specialized to culturally adapt to their local environment. With

biologically endowed omnivorousness and mobility, people found ways to extract

and process resources, shaping the environment in return. When global �uctuation

in climate and a sudden resource uncertainty hit the species, people around the

world focused on agriculture to have a �rmer control of necessities. The new strat-

egy shifted human history toward exponential growth and innovation to address the

drastic shift in lifestyle, leading to our high dependence on cultural adaptations to-

day. We may continue this trend in the future, with global changes to human genetic

diversity.

While a cohesive image of our species has formed in recent years, there is still much

to learn about our past. The work of many driven researchers shows that there are

amazing new discoveries made all the time that re�ne our knowledge of human evo-

lution. Technological innovations such as DNA analysis enable scientists to approach

lingering questions from new angles. The answers we get allow us to ask even more

insightful questions that will lead us to the next revelation. Like the pink limestone

strata at Jebel Irhoud, previous effort has taken us so far and you are now ready to

see what the next layer of discovery holds. Previous: Archaic Homo

Next: Race and Human Variation

Review �uestions

What are the skeletal and behavioral traits that de�ne modern

Homo sapiens? What are the evolutionary explanations for its

presence?

What are some creative ways that researchers have learned about

the past by studying fossils and artifacts?

How do the discoveries mentioned in “First Africa, Then the World”

�t the Assimilation model?

What is foraging and what adaptations do we have for this subsis-

tence strategy? Could you train to be a skilled forager?

What are aspects of your life that come from dependence on agri-

culture and its cultural effects? Where did the ingredients of your

favorite foods originate from?

Key Terms

African multiregionalism: The idea that modern Homo sapiens evolved as a complex

web of small regional populations with sporadic gene �ow among them.

Agriculture: The mass production of resources through farming and

domestication.

Amerindian: Term used to refer to the ancient humans of North and South America.

Assimilation model: Current theory of modern human origins stating that the

species evolved �rst in Africa and interbred with archaic humans of Europe and Asia.

Atlatl: A handheld spear thrower that increased the force of thrown projectiles. Previous: Archaic Homo

Next: Race and Human Variation

Band: A small group of people living together as foragers.

Beringia: Prehistoric landmass that connected Siberia and Alaska. The ancestors of

Paleoindians would have crossed this area to reach the Americas.

Beringia Standstill Model: Theory that people were genetically isolated in Beringia

before expanding to the Americas.

Carrying capacity: The amount of organisms that an environment could reliably

support.

Coastal Route model: Theory that the �rst Paleoindians crossed to the Americas by

following the southern coast of Beringia.

Dental caries: Damage to tooth enamel due to the waste products of built-up bacte-

ria. Known in the general public as cavities.

Early Modern Homo sapiens, Early Anatomically Modern Human: Terms used to re-

fer to transitional fossils between archaic and modern Homo sapiens that have a mo-

saic of traits. Humans like ourselves, who mostly lack archaic traits, are referred to

as Late Modern Homo sapiens and simply Anatomically Modern Humans.

Egalitarian: Human organization without strict ranks. Foraging societies tend to be

more egalitarian than those based on other subsistence strategies.

Foraging: Lifestyle consisting of frequent movement through the landscape and ac-

quiring resources with little storage.

Generalist-specialist niche: The ability to survive in a variety of environments by

developing local expertise. Evolution toward this niche may have been what allowed

modern Homo sapiens to expand past the geographical range of other human

species.

Globalization: A recent increase in the interconnectedness and interdependence of

people facilitated with long-distance networks.

Globular: Having a rounded appearance. Increased globularity of the braincase is a

trait of modern Homo sapiens.

Gracile: Having a smooth and slender quality; the opposite of robust.

Previous: Archaic Homo

Next: Race and Human Variation

Holocene: The epoch of the Cenozoic Era starting around 12,000 years ago and last-

ing arguably through the present.

Ice-Free Corridor model: Theory that the �rst prehistoric Native Americans crossed

to the Americas through a passage between glaciers.

Institutions: Long-lasting and in�uential cultural constructs. Examples include gov-

ernment, organized religion, academia, and the economy.

Introgression: The uneven mixing of DNA over time in which a small amount of out-

side genetic material is incorporated into a larger genome.

Kelp Highway hypothesis: Addition to the Coastal Route model that focuses on the

use of kelp-based environments as a resource.

Last Glacial Maximum: The time 23,000 years ago when the most recent ice age

was the most intense.

Later Stone Age: Time period following the Middle Stone Age with a diversi�cation

in tool types, starting around 50,000 years ago.

Levant: The eastern coast of the Mediterranean. The site of early modern human

expansion from Africa and later one of the centers of agriculture.

Malocclusion: The misalignment of the jaw due to the soft diets of agriculturalists.

The healthy development of the jaw, including making room for all of the teeth, de-

pends on experiencing a higher level of physical force than what people experience

with farmed and processed foods. The term literally means “bad shutting.”

Megafauna: Large prehistoric animals that may have been hunted to extinction by

people around the world.

Mental eminence: The chin on the mandible of modern H. sapiens. One of the de�n-

ing traits of our species.

Microlith: Small stone tool found in the Later Stone Age; also called a bladelet.

Middle Stone Age: Time period known for Mousterian lithics that connects African

archaic to modern Homo sapiens. Previous: Archaic Homo

Next: Race and Human Variation

Monumental architecture: Large and labor-intensive constructions that signify the

power of the elite in a sedentary society. A common type is the pyramid, a raised

crafted structure topped with a point or platform.

Mosaic: Composed from a mix or composite of traits.

Multiregionalism: Theory that modern Homo sapiens evolved simultaneously in

Africa, Asia, and Europe from archaic populations.

Neolithic Revolution: Time of rapid change to human cultures due to the invention

of agriculture, starting around 12,000 years ago.

Ochre: Iron-based mineral pigment that can be a variety of yellows, reds, and

browns. Used by modern human cultures worldwide since at least 80,000 years ago.

Out of Africa model: Theory that modern Homo sapiens expanded from Africa to

cover the rest of the world without interacting with archaic humans.

Paleoamerican, Paleoindian: Terms used to refer to the ancient humans of North

and South America.

Sahul: Prehistoric landmass connecting New Guinea and Australia.

Sedentarism: Lifestyle based on having a stable home area; the opposite of

nomadism.

Southern Dispersal model: Theory that modern H. sapiens expanded from East

Africa by crossing the Red Sea and following the coast east across Asia.

Subsistence strategy: The method an organism uses to �nd nourishment and other

resources.

Sunda: Asian prehistoric landmass that incorporated modern Southeast Asia.

Supraorbital torus: The bony brow ridge across the top of the eye orbits on many

hominin crania.

Upper Paleolithic: Time period considered synonymous with the Later Stone Age.

Urbanization: The increase of population density as people settled together in Previous: Archaic Homo

Next: Race and Human Variation

Keith Chan

cities.

Wallacea: Archipelago southeast of Sunda with different biodiversity than Asia.

Younger Dryas: The rapid change in global climate, especially a cooling of the

Northern Hemisphere, 13,000 years ago.

About the Author

Keith Chan, Ph.D.

University of Missouri: Grossmont College and MiraCosta College,

[email protected], keithcchan.com

Dr. Keith Chan is an instructor of anthropology at Grossmont

College and MiraCosta College in San Diego County. He reached

this step of his anthropological path after many memorable expe-

riences across the country and the hemisphere. He earned a

bachelor’s degree in anthropology from the University of Califor-

nia, Berkeley, in 2001. As a graduate student at the University of

Missouri, he traveled to Perú with teams of students to study pre-

historic skeletons to understand the lives of prehistoric Andeans.

He completed his writing to earn a Ph.D. in 2011. Inspired by many educators in his

journey, Dr. Chan turned his career toward teaching anthropology and helping stu-

dents understand and appreciate humanity.

For Further Exploration

Previous: Archaic Homo

Next: Race and Human Variation

Websites

Ministère de la Culture and Musée d’Archéologie Nationale. “Visit the cave” Lascaux

website. http://archeologie.culture.fr/lascaux/en/visit-cave.

SAPIENS. “Evolution.” SAPIENS website.

https://www.sapies.org/category/evolution/.

Smithsonian Institution. “What does it mean to be human?” Smithsonian National

Museum of Natural History website. http://humanorigins.si.edu.

ThoughtCo. “Archaeology.” ThoughtCo. Website.

https://www.thoughtco.com/archaeology-4133504.

University of California, San Diego. “MOCA Domains.” Center for Academic Research

& Training in Anthropogeny website.

https://carta.anthropogeny.org/moca/domains.

Books

Kolbert, Elizabeth. 2014. The Sixth Extinction: An Unnatural History. New York:

Bloomsbury.

Sykes, Bryan. 2002. The Seven Daughters of Eve: The Science That Reveals Our Ge-

netic Ancestry. New York: W. W. Norton & Company.

Articles

Stringer, C. 2016. “The Origin and Evolution of Homo sapiens.” Philosophical Trans-

actions of the Royal Society B 371 (1698):  doi�10.1098/rstb.2015.0237.

Trinkaus, E. 2018. “One Hundred Years of Paleoanthropology: An American Perspec-

tive.” American Journal of Physical Anthropology 165 (4): 638–651.

Previous: Archaic Homo

Next: Race and Human Variation

Wheelwright, Jeff. 2015. “Days of Dysevolution.” Discover 33–39.

http://discovermagazine.com/2015/may/16-days-of-dysevolution.

Wierer, Ursula, Simona Arrighi, Stefano Bertola, Günther Kaufmann, Benno Baum-

garten, Annaluisa Pedrotti, Patrizia Pernter, and Jacques Pelegrin. 2018. “The

Iceman’s Lithic Toolkit: Raw Material, Technology, Typology and Use.” PLOS One

13 (6): e0198292. doi�10.1371/journal.pone.0198292.

Documentaries

Brown, Nicholas, dir. 2015. First Peoples. Edmonton: Wall to Wall Television. Amazon

Prime Video.

Thompson, Niobe, dir. 2016. Great Human Odyssey. Edmonton: Clearwater Docu-

mentary. http://www.pbs.org/wgbh/nova/evolution/great-human-odyssey.html.

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Acknowledgments

I could not have undertaken this project without the help of many who got me to

where I am today. I extend sincere thank yous to the many colleagues and former

students who have inspired me to keep learning and talking about anthropology.

Thank you also to all who are involved in this textbook project. The anonymous re-

viewers truly sparked improvements to the chapter. Lastly, the staff of Starbucks

#5772 also contributed immensely to this text.

Figure Attributions

Figure 12.1 View looking south of the Jebel Irhoud (Morocco) site by Shannon

McPherron, Max Planck Institute for Evolutionary Anthropology Leipzig, is used un-

der a CC BY-SA 2.0 License.

Figure 12.2 Modern human and Neanderthal original to Explorations: An Open Invi-

tation to Biological Anthropology by Mary Nelson is under a CC BY-NC 4.0 License.

Figure 12.3 Modern and archaic Homo sapiens skeletons original to Explorations: An

Open Invitation to Biological Anthropology by Mary Nelson is under a CC BY-NC 4.0

License.

Figure 12.4 Loewenmensch1 by Dagmar Hollmann is used under a CC BY-SA 3.0 Li-

cense.

Figure 12.5 Maps depicting the estimated range of modern Homo sapiens through

time original to Explorations: An Open Invitation to Biological Anthropology by

Elyssa Ebding at GeoPlace, California State University, Chico is under a CC BY-NC

4.0 License. Previous: Archaic Homo

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Figure 12.6 A composite reconstruction of the earliest known Homo sapiens fossils

from Jebel Irhoud (Morocco) based on micro computed tomographic scans by

Philipp Gunz, Max Planck Institute for Evolutionary Anthropology Leipzig, is used

under a CC BY-SA 2.0 License.

Figure 12.7 Homo sapiens idaltu BOU-VP-16/1 Herto Cranium by ©BoneClones is

used by permission and available here under a CC BY-NC 4.0 License.

Figure 12.8 Homo sapiens Skull Skhul 5 by ©BoneClones is used by permission and

available here under a CC BY-NC 4.0 License.

Figure 12.9 Moulage de la sépulture de l’individu “Qafzeh 11” (avec ramure de

cervidé), homme de Néandertal (Collections du Muséum national d’histoire naturelle

de Paris, France) by Eunostos has been modi�ed (cropped and color modi�ed) and is

used under a CC BY-SA 4.0 License.

Figure 12.10 Liujiang cave skull-a. Homo Sapiens 68,000 Years Old (Taken at the

David H. Koch Hall of Human Origins at the Smithsonian Natural History Museum)

by Ryan Somma from Occoquan, USA has been modi�ed (color modi�ed) and is used

under a CC BY-SA 2.0 License.

Figure 12.11 Zhoukoudian Upper Cave by Mutt is used under a CC BY-SA 4.0 License.

Figure 12.12 Kow Swamp1-Homo sapiens by Ryan Somma from Occoquan, USA, un-

der a CC BY-SA 2.0 License has been modi�ed (background cleaned and color modi-

�ed) and is available here under a CC BY-NC 4.0 License.

Figure 12.13 Oase 2 by Smithsonian National Museum of Natural History [exhibit: Hu-

man Evolution Evidence, Human Fossils] has no known copyright restrictions and

has been modi�ed (sharpened) and is available here under a CC BY-NC 4.0 License.

Figure 12.14 Cro-Magnon 1 Skull by ©BoneClones is used by permission and available

here under a CC BY-NC 4.0 License.

Figure 12.15 Předmostí 9 by J. Matiegka (1862-1941) is in the public domain and has

been modi�ed (sharpened) and is available here under a CC BY-NC 4.0 License.

Figure 12.16 La station quaternaire de Raymonden (…)Hardy Michel bpt6k5567846s

(2) by M. Féauxis [original by Michel Hardy (1891)] is in the public domain.

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Figure 12.17 Lascaux cave (document 108435) Prehitoric Sites and Decorated Caves of

the Vézère Valley (France) by Francesco Bandarin, © UNESCO, has been modi�ed

(color modi�ed) and is used under a CC BY-SA 3.0 License.

Figure 12.18 Current Estimates of Arhcaic-Modern Admixture original to Explo-

rations: An Open Invitation to Biological Anthropology by Keith Chan and Katie Nel-

son is under a CC BY-NC 4.0 License.

Figure 12.19 BBC-shell-beads by Chenshilwood (Chris Henshilbood & Francesco

d’Errico) at English Wikipedia is used under a CC BY-SA 3.0 License.

Figure 12.20 Woolly Mammoth (La Brea Tar Pits & Museum) by Keith Chan is under a

CC BY-NC 4.0 License.

Figure 12.21 Fig 9. Paijan-like projectile point of rhyolite recovered from a late Pleis-

tocene level at the CH-I site (Unit 6, Level 51 cm; see S6a Fig) by Dillehay et al. (2015).

New Archaeological Evidence for an Early Human Presence at Monte Verde, Chile.

PLOS ONE, 10(11), e0141923. doi�10.1371/journal.pone.0141923 is used under a CC BY

4.0 License.

Figure 12.22 Clovis Point (15.2012.25) by Smithsonian National Museum of Natural

History [Department of Anthropology; Cooper Hewitt, Smithsonian Design Museum]

has no known copyright restrictions.

Figure 12.23 Assimilation Model original to Explorations: An Open Invitation to Bio-

logical Anthropology by Keith Chan and Katie Nelson is under a CC BY-NC 4.0 Li-

cense.

Figure 12.24 Image from video game FarCry Primal by Keith Chan is under a CC BY-

NC 4.0 License.

Figure 12.25 San hunter wıth bow and arrow by Charles Roffey has been modi�ed

(color modi�ed) and is used under a CC BY-NC-SA 2.0 License.

Figure 12.26 Plowing muddy �eld using cattle by IRRI Photos (International Rice Re-

search Institute) has been modi�ed (color modi�ed) and is used under a CC BY-NC-

SA 2.0 License.

Figure 12.27 Centres of origin and spread of agriculture by Joe Roe is used under a

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LICENSE

Explorations by Beth Shook, Katie

Nelson, Kelsie Aguilera, and Lara Braff,

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Figure 12.28 Downtown San Diego (October 13, 2016) by Keith Chan is under a CC

BY-NC 4.0 License.

Figure 12.29 Combine CR9060 by Hertzsprung is used under a CC BY-SA 3.0 License.

Figure 12.30 Hypothetical image of future human evolution original to Explorations:

An Open Invitation to Biological Anthropology by Mary Nelson is under a CC BY-NC

4.0 License.

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Previous: Archaic Homo

Next: Race and Human Variation