SEPTEMBER

MONTH 2018 | WWW.THE-SCIENTIST.COM

2020 | WWW.THE-SCIENTIST.COM

HOW HOMO SAPIENS

EVOLVED IN AFRICA

SOUTH AMERICAN
PREHISTORY

UNDERSTANDING
MORPHOLOGICAL CONTROL

PLUS
RESEARCHERS’ ROLES IN
CRAFTING SCIENCE POLICY

HUMAN PATHS
ARCHAEOLOGY AND GENETICS ARE STARTING TO RESOLVE
HUMANITY’S ORIGIN AND SPREAD
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30 T HE S CI EN T I S T | the-scientist.com
 t’s not unusual for geochronologist
Rainer Grün to bring human bones
back with him when he returns
home to Australia from excursions
in Europe or Asia. Jawbones from
extinct hominins in Indonesia, Nean-
derthal teeth from Israel, and ancient
human finger bones unearthed in
Saudi Arabia have all at one point spent
time in his lab at Australian National Uni-
versity before being returned home. Grün
specializes in developing methods to discern
the age of such specimens. In 2016, he car-
ried with him a particularly precious piece
of cargo: a tiny sliver of fossilized bone cov-
ered in bubble wrap inside a box.
The bone fragment had come from a
skull—still stored at the Natural History
Museum in London—with a heavy brow
ridge and a large face. It looked so primi-
tive that the miner who had discovered it
in 1921 at a lead mine in the Zambian town
of Kabwe, then in the British territory of
Rhodesia, first thought it had belonged
to a gorilla. But later that year, museum
paleontologist Arthur Smith Woodward
noticed what he interpreted as typically
human features, such as the skull’s thin
and relatively large braincase, that moti-
vated him to designate the specimen as its
own hominin species.
In the 1980s, however, museum
paleoanthropologist Chris Stringer took
another look at the skull and classified it
as belonging to the species Homo heidel-
bergensis, an ancient hominin thought to
be a human ancestor. Based on its prim-
itiveness, Stringer says, most research-
ers guessed it was an early individual
who lived around half a million years ago,
some 200,000 years before the earliest
Homo sapiens were starting to emerge.
© MARK GARLICK, SCIENCE PHOTO LIBRARY
But nobody knew exactly how old the
skull was. For decades, no dating method
existed that could identify the fossil’s age
without the destructive process of grind-
ing up bits of bone for analysis. But Grün
was determined to find a solution.
Grün is one of very few geochronologists
proficient in a laser technique that extracts
and reduces a barely visible grain of bone—
smaller than the bone’s natural pores—to
atoms, he says. The laser is coupled with a
mass spectrometer, which measures the con-
centrations of uranium isotopes that undergo
radioactive decay at a specific rate over time.
Having returned from his trip to pro-
cure the Homo heidelbergensis sample, Grün
watched as the laser poked two tiny holes
into the bone fragment and the particles dis-
appeared into the mass spectrometer. Upon
evaluating the mass spec data, he could tell
that the fragment was much younger than
previously believed. As he, Stringer, and oth-
ers reported in Nature this past April, their
best estimate was 299,000 years, give or
take 25,000.1 That meant that the Kabwe
individual had lived not before, but around
the same time as the first Homo sapiens–like
people dwelled in North Africa. Along with
other archaeological evidence, the findings
suggest that perhaps Homo heidelbergensis
was not our ancestor, but a neighbor.
Together with yet another hominin,
Homo naledi, known to have existed in
southern Africa at that time, Africa may
have been a crowded place. “Ten years ago,
I think most of us would’ve thought, well,
Africa in the last 300,000 years is just
going to show you the evolution of Homo
sapiens, and that’s really all—the other
species would have disappeared, gone
extinct,” notes Stringer. “Now we know
that there were probably at least three dif-
ferent kinds of hominins around.”
That’s akin to the situation that
unfolded in Eurasia, where Neanderthals
and Denisovans thrived for hundreds of
thousands of years before Homo sapiens
migrated out of Africa and at times even
interbred with the other hominin groups.
The story in Africa remains murky,
however, as researchers have not been
able to reconstruct human history in vivid
detail, in part because hominin fossils
informative about our species’ emergence
and coexistence with other species are
rare in Africa. As a result, finds such as the
Kabwe skull continue to raise more ques-
tions than answers. If Homo heidelbergen-
sis wasn’t one of our recent ancestors, then
who was? If our species really did overlap in
time with Homo heidelbergensis, what role
did they play in our evolutionary history?
In recent years, a field that has tradition-
ally relied on fossil discoveries has acquired
helpful new tools: genomics and ancient
DNA techniques. Armed with this combina-
tion of approaches, researchers have begun
to excavate our species’ early evolution, hint-
ing at a far more complex past than was pre-
viously appreciated—one rich in diversity,
migration, and possibly even interbreeding
with other hominin species in Africa.
“To piece together that story, we need
information from multiple different fields of
study,” remarks Eleanor Scerri, an archae-
ologist at the Max Planck Institute for the
Science of Human History in Jena, Ger-
many. “No single one is really going to have
all the answers—not genetics, not archae-
ology, not the fossils, because all of these
areas have challenges and limitations.”
A sparse fossil record
Bones easily disintegrate in many parts
of Africa, in acidic forest soils or dry, sun-
exposed areas. Moreover, the continent is
largely unexplored by archaeologists. While
northwestern Africa and former British ter-
ritories in eastern and southern Africa have
a long tradition of professional archaeologi-
cal research, few researchers have looked
for fossils anywhere else, notes archaeolo-
gist Khady Niang of Cheikh Anta Diop Uni-
versity in Senegal. That’s especially the case
for the western and central parts of the con-
tinent, where preservation conditions are
also poor and excavations difficult at times
due to political instability. “We might be
missing some really, really important parts
of the story,” adds Yale University anthro-
pologist Jessica Thompson.
What African hominin fossils do make
clear is the depth of humanity’s roots on
that continent. Researchers have found
some of the most abundant fossils in sed-
iments between 3.5 million and 3.2 mil-
lion years old. That appeared to be the
heyday of the australopiths (including the
genus Australopithecus), apes that walked
upright and are believed to have used stone
tools, but still climbed trees and had rela-
tively small brains. It’s thought that some-
how our own genus, Homo, emerged from
transitional ape species some 2.8 million
years ago as a clan of hominins with dis-
tinctive teeth, probably adapted to an
eclectic diet that allowed them to thrive
0 9. 202 0 | T H E S CIEN T IS T 3 1
 in a wide range of habitats. But there are
few sediments, let alone fossils, left behind
from that time, making the birth of our
genus one of the most poorly understood
periods in our evolution, Thompson notes.
The fossil record yields more secrets
about the time shortly after the emergence
of Homo, revealing a diversity of different
Homo species in Africa, of which Homo
erectus seems to persist the longest. H. erec-
tus crops up in Africa’s limited fossil record
around 2 million years ago and hangs
around on the continent until roughly
a million years ago. It was the first hom-
inin that shows evidence of having lived in
human-like social groupings and used fire,
and it is thought to be a human ancestor.
When and how H. sapiens emerged isn’t
at all clear, but what is apparent is that we
weren’t alone; fossils suggest that several
other hominin species, such as that repre-
sented by the Kabwe skull, inhabited the
continent at the time our species appeared.
Another relatively small-brained hominin,
Homo naledi, is also thought to have lived
in southern Africa around 300,000 years
ago.2,3 And inside a Moroccan cave called
Jebel Irhoud, 300,000-year-old skeletons
were found that carry very early features
of H. sapiens.4 It’s not yet known how long
those different hominin species existed,
however, or whether they physically over-
lapped and perhaps even shared genes with
one another, Stringer notes, or whether
there were others.
By around 160,000 years ago, the con-
stellation of physical features that defines
us today—such as a globular braincase and
a pointed chin—had begun to emerge in
ancient hominin groups represented by
fossils found across Africa. Later, some
of these anatomically modern humans
crossed the thin spit of land that connects
Africa to Eurasia, probably on several
occasions. On that new continent, they
eventually met Neanderthals and Den-
isovans, which, like two hobbit-size Homo
species found on southeast Asian islands,
are thought to be the evolutionary prod-
ucts of earlier hominin migrations out of
the continent. “Africa was this sort of leaky
faucet, and hominins were just dribbling
out of it all the time,” Thompson says.
32 T HE S CIE NT IST | the-scientist.com
Fossil finds over the years have steadily
bolstered a long-held idea that anatomically
modern humans first emerged in Africa.
This “Out of Africa” model, proposed by
anthropologists in the late 20th century,
posited that all humans of Eurasian ancestry
descended from a single ancestral African
population, which then spread throughout
the world and displaced all other homi-
nins. The opposing “multi-regionalism”
model, by contrast, conceived that multiple
human subpopulations—which stemmed
from regional lineages of an ancestral spe-
cies such as Homo erectus—existed across
Europe, Asia, and Africa, and through con-
tinuous mixing evolved together to form the
present human population.
While fossils supported the for-
mer theory, it was the advent of genetic
research that showed unequivocally that
populations outside of Africa descended
from a single population in Africa. But the
story had a twist: in two groundbreaking
studies published in 2014, researchers
compared ancient DNA extracted from
Neanderthal bones and compared it with
modern-day people, and found that 2 per-
cent of the average European genome is
Neanderthal in origin.5,6 Our species orig-
inated in Africa, but interbred with homi-
nins outside of it. (See “Our Inner Nean-
derthal,” The Scientist, September 2019.)
These findings, and many since, have
highlighted the power of genetics in resolv-
ing questions about human ancestry that
fossils alone cannot. Investigations of the
genomes of living Africans are now under-
way to help fill in the gaps of Africa’s fossil
record. “[Such studies] are really provid-
ing important insights into our population
history and African origins,” says Yale Uni-
versity evolutionary biologist Serena Tucci.
“We are getting to know and understand
processes that happened very early on in
our evolutionary history.”
Ghost hominins
Even the very first investigations of our
genetic ancestry, gleaned from small, bite-
size chunks of genetic material, positioned
Africa as the cradle of humanity. One
widely publicized 1987 study compared
mitochondrial gene snippets from 147
people across the world, and concluded
that Africans have the highest mitochon-
drial diversity, suggesting that our species
originated and spent most of its evolution-
ary history there.7 Specifically, the authors
traced all human mitochondrial diversity
back to a single theoretical woman who
lived in East Africa hundreds of thousands
of years ago, whom the media popular-
ized as “mitochondrial Eve.” Later stud-
ies estimated that the most recent com-
mon ancestor of modern Y chromosome
variation (dubbed “Y chromosome Adam”)
could also be traced back to Africa.
Subsequent studies of nuclear DNA
have validated our African birthplace
and refined our knowledge of the human
genetic landscape. Several studies of
genetic variation among modern-day
Khoe and San individuals, two groups
of indigenous people in southern Africa
known for their click language, have sug-
gested they represent our species’ most
genetically diverse lineage.8 Collectively
known as Khoe-San, this group is thought
to have split from other populations
between 200,000 and 350,000 years
ago, making them the most ancient popu-
lation of modern humans to diverge. Non-
Africans, meanwhile, represent a reduced
subset of the diversity in Africa, and likely
trace most of their ancestry back to just
one small population—probably no more
than a few thousand individuals—who
ventured out of the continent between
60,000 and 70,000 years ago.9
Some scientists see the extraordi-
nary diversity in modern Khoe-San peo-
ple as evidence that our species arose in
southern Africa. Along with some archae-
ological evidence from the region, that
challenges the long-held idea of an East
African origin, which was based on the
fact that many early hominin fossils were
found there. However, trying to pinpoint
the precise location of our species’ ori-
gins from DNA is often criticized for the
simple reason that people move around—
it’s not known if the populations living in
one place today were there hundreds or
thousands of millennia ago. In fact, some
researchers, including Scerri, Stringer,
and Thompson, have recently constructed
an entirely new theory of our origins: that
anatomically modern humans didn’t arise
from a single place, but gradually emerged
from a web of interconnected populations
sprawled across Africa—a continental
gene-sharing bonanza that hominin lin-
eages besides our own may have partici-
pated in.10 “It’s a good way to interpret the
data we have right now,” says Niang.
In addition to where we evolved, research-
ers are interested in how: which genes gave
us a selective advantage to survive in par-
ticular environments, and which ancestors
contributed to our genomes? Unfortunately,
modern African DNA is severely underrep-
resented in genetic research, making these
questions particularly challenging to answer.
Most sequenced genomes are of European
origin, with fewer than 2 percent coming
from Africans. This dearth of African
genomes is compounded by the fact that
the genetic scaffold underlying some fre-
quently studied traits such as skin pigmen-
tation appear to be far more complex in Afri-
cans than in other populations, notes Brenna
Henn, a population geneticist at the Univer-
sity of California, Davis. “The twelve to fif-
teen genes [for skin pigmentation] that peo-
ple cite in Eurasian populations explain less
than 25 percent of the variation in Africans.”
With few sediments, let alone fossils, left behind
from that time, the birth of our genus is one of the
most poorly understood periods in our evolution.
African population history compli-
cates matters further. Large-scale migra-
tions pulled people back and forth across
the continent for thousands of years.
People from Eurasia also migrated back
to Africa. Where people moved, they
swapped their genes with local popu-
lations, shuffling patterns of ancestry
across African genomes. This upheaval of
ancient population structures creates one
of the biggest challenges in teasing out
archaic history from modern genomes,
notes University of Pennsylvania geneti-
cist Sarah Tishkoff. “It can make it very
tricky to distinguish that older history
when there’s been this newer wave of
gene flow messing with your modeling.”
Still, geneticists have been able to tease
out some signals from our distant past,
using computational models that ask what
kind of evolutionary processes—such as
mutation, selection, and interbreeding with
other groups—best explain the pattern of
variation across modern genomes. One
intriguing finding of such studies is possi-
ble evidence of mixture with now-extinct,
unknown groups of modern humans and
other hominins: “ghost” populations that,
like Neanderthals, left traces in modern
genomes. In one analysis of 15 sequenced
genomes, Tishkoff ’s group investigated
the sources of genetic variation in three
different modern African hunter-gatherer
groups.11 The team’s models suggested that
interbreeding with an archaic hominin
species—which seemed as different from
modern humans as are Neanderthals—was
the most likely origin for a set of unusual
sequences they found. “The model that
includes a ghost population is always bet-
ter [to fit the data], basically,” Tishkoff says.
A handful of similar studies have also
revealed traces of ghost hominins in mod-
ern African genomes, sometimes account-
ing for up to 10 to 20 percent of the genetic
variation. Some research suggests that mix-
ing took place after the ancestors of mod-
ern Eurasians left Africa, hinting that
other kinds of hominins could have existed
alongside Homo sapiens in Africa until very
recently. “It’s actually pretty convincing,” says
Henn, who wasn’t involved in these studies.
“Ten percent of the genome—I’m going to
have a hard time invoking one single other
process that can explain a signal like that.”
Ultimately, researchers need samples of
DNA from ancient hominins to prove whether
archaic African species did in fact contribute
to modern genetic variation. While scientists
have managed to overcome some of the tech-
nical hurdles of sequencing highly degraded
ancient DNA from human fossils in Africa, the
oldest human DNA found on the continent is
just 15,000 years old, an age that pales in com-
parison to some 400,000-year-old hominin
DNA found in a cave in Spain with relatively
cool, stable temperatures. Archaeologists can
only dream of finding intact DNA that old on
the African continent, notes Tessa Campbell,
an ancient DNA specialist at Iziko Museums
of South Africa. “No one wants to say never . .
. but it’s very unlikely.”
Because DNA is unlikely to survive
very long in the African heat, researchers
have largely refrained from drilling into
the fossils they’ve found of other homi-
nins in Africa for fear of destroying them.
But efforts are underway to study ancient
DNA from younger fossils of Homo sapiens
to crack other mysteries about human his-
tory on the continent, Tucci notes. “This is
definitely a new era for African genomics.”
Mining bones for ancient DNA
In 2015, an international team of research-
ers managed to harvest the first ancient
DNA in Africa—the genome of Mota, a man
who left behind 4,500-year-old remains in
an Ethiopian cave.12 In the five years since
that publication, researchers have pub-
lished nearly 100 other full and partial
ancient human sequences from Africa.
These genomes have helped scientists bet-
ter understand the messy signatures from
recent migration events that make studies
of modern genomes so difficult.
For instance, mitochondrial DNA from
the skulls of seven people who lived some
15,000 years ago in modern-day Morocco
revealed that they were closely related to
Natufians, hunter-gatherers who dwelled
in the Near East, as well as people living
south of the Sahara desert.13 This finding
suggested that there were far-flung con-
nections between North Africa, the Near
East, and sub-Saharan Africa before the
dawn of agriculture.
Analyses of ancient DNA have also
helped researchers understand how ancient
migrations affected the genomes of people
alive today. One such migration is the Bantu
0 9. 2020 | T H E S CIEN T IS T 33
 expansion, which gradually spread West Afri- lier back-migrations into the continent and of children buried inside a rock shelter in
can farming practices across the continent eventually carried to the southernmost tip of Cameroon.15 Comparing the 3,000- and
between roughly 5,000 and 1,000 years ago. Africa as other migrating human populations 8,000-year-old DNA with ancient genomes
By comparing DNA from ancient hunter- moved southward, the researchers found. collected elsewhere and with genomes of

gatherer remains in southern Africa with
modern-day Khoe-San people, evolutionary
biologist Carina Schlebusch of Uppsala Uni-
versity in Sweden and her colleagues found
that some Khoe-San groups carry DNA that
ancient farmers brought with them.14 They
also carry mixed Eurasian ancestry that had
been introduced to North Africa with ear-
Such studies have also provided insight
into deep divergences that occurred in
human populations long before migrations
of farmers and herders. Mary Prendergast,
an anthropologist at Saint Louis Univer-
sity in Madrid, and her colleagues recently
sequenced the first ancient DNA from West
Africa, material extracted from the remains
© ISTOCK.COM, GIORGIOMORARA
modern people allowed the researchers to
reconstruct some of the earliest branches
of our species’ evolutionary tree. In addi-
tion to the deep split between Khoe-San
groups and other African populations—
from which non-Africans also descend—
their model suggested that two other major
lineages split just as deeply, diverging from

OUR HISTORY IN AFRICA

Hominin fossils that reveal clues to the emergence of Homo sapiens are rare in Africa, but in combination with studies
of modern human genomes, researchers are piecing together an ever more complex timeline of human history.

HOMO HEIDELBERGENSIS
Named after its initial discovery near Heidelberg,
Germany, fossils similar to Homo heidelbergensis
were later found to also occur in Africa. It routinely
hunted large animals and may have built dwellings
made of wood or rock.

HOMO ERECTUS
Homo erectus may have been the first
hominin to wield fire and stone axes.
*Note that the scale of this graphic changes across its width. While Homo sapiens has The species spread across Asia, where it
existed for the past few hundred thousand years, it’s noteworthy that Homo erectus, continued to evolve. In Africa, it gave rise
living from around 2 mya to perhaps 100 kya, is still the longest-lived hominin species. to Homo heidelbergensis.

EARLY HOMO SPECIES

SAHELANTHROPUS, ORRORIN,
ARDIPITHECUS SPP.
Members of these relatively small-brained
genera probably emerged not long after
the human-chimpanzee divergence, and
are the first known species of apes that
habitually walked upright.
Hominins of the genus Homo shared
distinctly small back teeth, which experts
think allowed them to consume diverse
diets. One species of early Homo evolved
into Homo erectus.
AUSTRALOPITHECUS SPP.
This diverse group exhibited both
ape- and human-like characteristics.
Some species are thought to
have used stone tools and to have
evolved into the genus Homo.

7 mya 6 5 4 3 2 1 mya
 It was not a streamlined process of australopiths
steadily evolving into modern humans, but a
messy and haphazard journey that includes
interwoven ancestries of many groups.
one another more than 200,000 years ago.
One lineage is ancestral to central African
hunter-gatherers known as Aka and Mbuti,
and the second is a previously unknown
“ghost” lineage whose fate is uncertain.
“There’s all this deep, deep population struc-
ture with various differentiated branches of
the human tree throughout the Pleistocene
in Africa that we haven’t really appreciated
very much yet,” Prendergast says.
Only time will tell whether researchers’
current arsenal of technologies is enough to
untangle the complete story of human evo-
lution. Perhaps novel technologies—such as
paleoproteomics, a nascent field that aims to
reconstruct ancestry from fossilized proteins,
which are more durable than DNA—will help
researchers “push further back in time,” notes
biological anthropologist Rebecca Acker-
mann of the University of Cape Town.

Comparing ancient DNA from Neanderthals

and Denisovans with modern human genomes
has revealed that modern humans interbred
with these other hominin groups.

DENISOVANS
Denisovans, only known from
from ancient DNA and a handful
of bones and teeth, were closely
related to Neanderthals.

Genomic analyses suggest that the

majority of people living outside Africa HOMO NEANDERTHALENSIS
today trace most of their ancestry back Compared with modern humans, HOMO SAPIENS
Neanderthals had shorter, stockier Eventually, the hallmarks of our
to a single migration event of a small skeletons and larger noses, but their own species—exceptionally large
group of modern humans who left Africa brains were just as large, if not larger. brains, flat faces, and small jaws—
between 60,000 and 70,000 years ago. appear in the fossil record. Genetic
studies suggest that H. sapiens
started to split into several major
lineages of modern humans more
than 200,000 years ago.

HOMO NALEDI
Only known from skeletons found in South Africa,
this species had a remarkably tiny brain but
modern-human–like features such as the shape of
its teeth and possibly the habit of burying its dead.

HOMO FLORESIENSIS AND HOMO LUZONENSIS

Researchers have recently discovered two small hominin
species on Pacific islands, Homo floresiensis on Flores in
Indonesia and Homo luzonensis in the Philippines.
Some analyses of modern human genomes hint
that Homo sapiens may have interbred with other
hominins in Africa.

500 kya 100 kya 50 kya Present

 EXCAVATING A CONTINENT
A number of researchers suspect that Homo sapiens arose not in a single place in Africa,
but across the entire continent, emerging from a network of interconnected hominin populations.
But for decades, archaeologists positioned East and South Africa as important places for hominin
evolution and the putative birthplace of our species. That’s likely because most fossils, including
groundbreaking findings that have transformed our understanding of human evolution, have been
AFAR REGION, ETHIOPIA, 1974
found in those regions. “Lucy,” 3.2 million years ago
Lucy—the skeletal remains of an Australopithecus
afarensis female—is one of the best-known
hominin fossils. Studies suggest that she was

NEAR SAFI, MOROCCO, 1961
Human remains at Jebel Irhoud, 315,000 years ago
Flint blades and Homo sapiens–like skeletons in a Moroccan
cave known as Jebel Irhoud may represent the oldest Homo
sapiens artifacts. The skeletons have modern features such
as round skulls and modern-human–like teeth and faces.
WIKIMEDIA, 120; ARIZONA STATE UNIVERSITY; WIKIMEDIA, BAHN, PAUL G; WIKIMEDIA, GERBIL; WIKIMEDIA, RYAN SOMMA; ELIFE, 6:E24232, 2017; WIKIMEDIA, GUILLAUMEG
both tree-dwelling and capable of an upright gait,
providing an important evolutionary stepping
stone from more primitive ape species to
modern humans.
AFAR REGION, ETHIOPIA, 2013
Adult jawbone, 2.8 million years ago
A mandible fragment is the earliest
known trace of the genus Homo, although
the species it belongs to is a mystery.

OMO NATIONAL PARK, ETHIOPIA, 1967-1974

Omo Kibish remains, 195,000 years ago
Fragments from two skulls, four jaws, a legbone, a few
hundred teeth, and some other bones were found at
a site in Ethiopia, and are classified as anatomically
modern Homo sapiens.

LAKE TURKANA, KENYA, 1984

“Turkana Boy,” 2 million years ago
A nearly complete skeleton of an ancient
Homo erectus child found near Kenya’s Lake
Turkana provides a rare glimpse into how
quickly this species reached adulthood and
how similar their skeletons were to ours.

RISING STAR CAVE, SOUTH AFRICA, 2013

Homo naledi, 236,000–335,000 years ago
KABWE, ZAMBIA, 1921 In 2013 and 2014, cavers found skeletons of two adults and
one juvenile of what is believed to be a new species: Homo
“Kabwe skull,” 300,000 years ago naledi. Its tiny brain and ape-like shoulders—indicating it
Also called “Broken Hill skull,” the specimen is considered was a good climber—suggest it may be an evolutionary o -
a representative of Homo heidelbergensis. shoot lineage that went extinct.

36 T HE S CI EN T I S T | the-scientist.com

What is already abundantly clear
is that human evolution was far more
complex than previously appreciated by
anthropologists. It was not a streamlined
process of australopiths steadily evolv-
ing into modern humans, but a messy
and haphazard journey that includes
interwoven ancestries of many groups,
some of which have never been discov-
ered other than through the genetic
traces they left in ancient and modern
genomes. “We have a long history. A lot
of things happened, and a lot of ances-
tors contributed to our genomes today,”
Schlebusch says. “It’s not going to be a
simple story.” g
Katarina Zimmer is a New York–based
freelance journalist. Find her on Twitter
@katarinazimmer.
References
1. R. Grün et al., “Dating the skull from Broken Hill,
Zambia, and its position in human evolution,”
Nature, 580:372–75, 2020.
2. L.R. Berger et al., “Homo naledi, a new species
of the genus Homo from the Dinaledi Chamber,
South Africa,” eLife, 4:e09560, 2015.
3. P.H. Dirks et al., “The age of Homo naledi and
associated sediments in the Rising Star Cave,
South Africa,” eLife, 6:e24231, 2017.
4. J.-J. Hublin et al., “New fossils from Jebel Irhoud,
Morocco and the pan-African origin of Homo
sapiens,” Nature, 546:289–92, 2017.
5. B. Vernot, J.M. Akey, “Resurrecting surviving
Neanderthal lineages from modern human
genomes,” Science, 343:1017–21, 2014.
6. S. Sankaraman et al., “The genomic landscape
of Neanderthal ancestry in present-day humans,”
Nature, 507:354–57, 2014.
7. R.L. Cann et al., “Mitochondrial DNA and human
evolution,” Nature, 325:31–36, 1987.
8. C.M. Schlebusch et al., “Genomic variation
in seven Khoe-San groups reveals adaptation
and complex African history,” Science,
338:374–79, 2012.
9. S. Tucci, J.M. Akey, “A map of human
wanderlust,” Nature, 538:179–80, 2016.
10. E.M.L. Scerri et al., “Did our species evolve in
subdivided populations across Africa, and why does
it matter?” Trends Ecol Evol, 33:582–94, 2018.
11. J. Lachance et al. “Evolutionary history
and adaptation from high-coverage whole-
genome sequences of diverse African hunter-
gatherers,” Cell, 150:457–69, 2012.
12. M.G. Llorente et al., “Ancient Ethiopian genome
reveals extensive Eurasian admixture in Eastern
Africa,” Science, 350:820–22, 2015.
13. M. v.d. Loosdrecht et al., “Pleistocene North African
genomes link Near Eastern and sub-Saharan African
human populations,” Science, 360:548–52, 2018.
14. C.M. Schlebusch et al., “Southern African
ancient genomes estimate modern human
divergence to 350,000 to 260,000 years ago,”
Science, 358:652–55, 2017.
15. M. Lipson et al., “Ancient West African foragers
in the context of African population history,”
Nature, 577:665–70, 2020.

DECOLONIZING STUDIES OF HUMAN EVOLUTION

The San people of southern Africa are one of the most intensively studied indigenous groups in the world. Their click language and traditional
hunter-gatherer lifestyles have long fascinated anthropologists. And the antiquity of their genetic lineage makes them a treasure trove for
geneticists studying human evolutionary history.
However, studies on San lifestyles and genomes have not always been conducted ethically. For instance, scientists have sometimes referred to
the San as “bushmen,” a derogatory term associated with colonial-era researchers using modern indigenous groups as models of primitive human
ancestors, and have taken photographs of children and breastfeeding mothers without permission. “We’re not saying that everybody is bad. But
you get those few individuals who don’t respect the San,” Leana Snyders, head of the South African San Council in Upington, South Africa, told Sci-
ence in 2017. Ethical conduct in genomic research came to the foreground in 2010 following a high-profile analysis of San genomes in Nature in
which the authors had, among other transgressions, not asked San leaders for permission to conduct the study.  
All disciplines that study human evolution in Africa have at times been criticized for their extractive nature. Archaeological research—a field
pioneered by European colonial nations—has long been driven by Western researchers digging up fossils from Africa to study them, sometimes
taking them elsewhere to do so. Some hominin fossils are still displaced, such as the Kabwe skull, a famous Homo heidelbergensis specimen that
remains in London’s Museum of Natural History, despite Zambia’s multiple requests to repatriate the skull. According to an April press release, the
museum has approached Zambian authorities to begin discussing the possible return of the skull following a 2018 agreement between the UK and
Zambia to find a solution to the issue.
Some scientists have called for regulations to protect fossil collections from ancient DNA research, whereby African hominin fossils undergo
the damaging process of extracting DNA. Now, “African museums are taking a leading role to make sure this [research] happens through collab-
oration and regulation,” notes anthropologist Mary Prendergast of Saint Louis University’s Madrid campus, as geneticists are working to develop
new, less destructive techniques for ancient DNA analysis.
The San, for their part, created a code of research conduct in 2017 that, for example, requires researchers to respect their communities and to
allow them to comment on findings prior to publication to avoid derogatory interpretations. Researchers are also required to compensate the com-
munity for their cooperation, through financial support, knowledge, or job opportunities, for instance.
A number of scientists have called for a greater role of African scientists in human evolutionary research. To make that possible, Western funding
agencies and institutions have an obligation to support African e orts to improve their countries’ antiquities infrastructure, so that “the next genera-
tion of African scholars [can] take control of the research in their areas,” notes anthropologist Eleanor Scerri of the Max Planck Institute in Germany.
CREDIT LINE

Foreign research teams should also foster stronger collaboration with African researchers, rather than simply seeking their help with fossil
excavations, which has sometimes been the case, notes University of Cape Town biological anthropologist Rebecca Ackermann. Research groups
have become more diverse, she notes, but the transition is slow. “I do see a change. It’s just not as fast as I would like.”

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