Evolutionary Anthropology 117

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Forest People: The Role of African Rainforests in

Human Evolution and Dispersal
JULIO MERCADER

Conventionally, the African continent has been partitioned in two evolutionary

domains. One of them, the rainforest, is home to apes and covers central and West
Africa. The other one extends through the woodlands and savannas of East and
Southern Africa and has been traditionally perceived as home to humanity. The
morphology of early humans is well-adapted to open environments.1 In addition,
food procurement in savannas is known to be easier and more reliable than is
provisioning in the rainforest, with its dispersed and cryptic faunal resources and
fickle carbohydrates and fat. In the late 1980s, human ecologists and sociocultural anthropologists demonstrated that full-fledged foraging without some
agricultural support has been virtually undocumented in tropical forests today or in
the recent past.2 This research portrayed the present-day rainforest ecosystem as
an unfriendly environment that is unable to support purely foraging groups, and
questioned whether hominids ever lived in it.2

Archeological research in the tropical rainforest is in its infancy, and

faces scarce human and financial resources, harsh logistics, and lack of
continuity because of political unrest
in the region. As a result, we face a
meager, poorly understood, and, for
the most part, undated record, and
thus a deficient database. On the contrary, in East Africa decades of continued research by various interna-

Julio Mercader has conducted fieldwork

in the Democratic Republic of Congo,
Equatorial Guinea, Cameroon, and Cote
dIvoire Trained in paleolithic archeology
of the Old World, his research interests
include the early hunter-gatherer settlement of rainforests, archeological site
formation in wet tropical environments,
forest technologies, paleoecology, and is
a pioneer in the archeology of chimpanzee stone tool sites.

tional and multidisciplinary teams,

together with the existence of a relatively well-preserved, visible, and datable fossil record, have allowed paleoanthropologists and archeologists to
produce a consistent archeological sequence for the emergence of humanity and the consolidation of modern
human behavior. In this paper I examines the evidence of rainforest occupation in Central and West Africa. I
suggest that too little effort has been
deployed in the rainforest region; that
tropical forests played a significant
role in human evolution and early dispersal; that the rainforest record is
complex and not well understood in
terms of formation, preservation, and
archeological resolution; and that
there is untapped potential in the archeology of tropical rainforests.

EARLY STONE AGE

Key words: African rainforest archeology, paleoecology, site formation, Acheulian, Sangoan, Lupemban, Later Stone Age
Evolutionary Anthropology 11:117124 (2002)

DOI 10.1002/evan.10022
Published online in Wiley InterScience
(www.interscience.wiley.com).

The rainforest is far from being a

stable ecosystem. It reached its
present geographical distribution,
morphology, structure, and species
composition only in the late Holocene. During the middle and late
Pleistocene, climatic conditions in the

African tropics underwent significant

changes. In East Africa there is
mounting evidence of aridity and the
formation of savannas, but were West
and Central Africa affected by similar
climatic trends with the same environmental effects?
Recent research shows that tropical
forests, dry forests, woodlands, and
montane forests in terrestrial, riverine, and lacustrine settings were the
cradle of the earliest biped hominids
during the Miocene and Pliocene,
from 6 to 3 million years ago.312 Boesch and Boesch13 have argued that
significant aspects of hominization
could have taken place in rainforest
environments. In fact, it recently has
been reported that some members of
the Hominoidea such as Kenyapithecus and Victoriapithecus were present
in woodland settings of East Africa as
early as 15 My ago.3 Similar insights
are provided by the faunal evidence
from this period, as manifested by the
environmental requirements of the
Colobinae9; bovids such as the
Tragelaphini, Reduncini, and Bovini6;
and the overall scarcity of the Giraffidae, Hippopotamidae, and savannaadapted bovids such as the Alcelaphini.6 At Omo, fossil pollen, wood,
and fruits confirmed riverine environments with relatively closed forests.4,10 11 Similar conclusions were
drawn from the microfaunal record.5
Yet no hominid species has been
shown to be exclusively adapted to a
closed rainforest environment of the
central African type rather than a
combination of Guineo-Congolian
taxa in cohort with species from alpine regions and variably dry woodlands and savannas.4
What is the evidence that Homo ergaster had the potential to exploit

118 Evolutionary Anthropology

tropical forests? Some authors have

proposed that Homo ergaster did not
have the behavioral or biological flexibility, especially in terms of its subsistence base, required to settle in
rainforests. The scarcity of edible resources and the degree of processing
required to consume them have been
perceived as unsurpassable barriers
for early Homo.14,15 Yet the African
forest is home to an extensive number
of plant species that potentially were
subject to consumption and manipulation by prehistoric foragers and that
provide oils, starches, vegetables,
nuts, fruits, spices, and stimulants.
Wild yams, potential carbohydrate
source, are unevenly distributed, unpredictable, deeply buried, and sometimes toxic. The density of wild tubers
in the African rainforest, however,
ranges from 22 stems per hectare in
the evergreen forest to 53 in semi-deciduous forests.16 Hladik and Dounias16 have argued that the standing
biomass of wild yams in the rainforest
is able to sustain low population densities of hunter-gatherers. Likewise,
the use of tropical oils by forest foragers has been attested to in numerous
places in central Africa and has great
antiquity. Thus, Mercader has shown
that in Central Africa arboriculture
and the use of Canarium schweinfurthii dates back more than 10,000
years.17 Although some tubers and
nuts can be toxic, foraging groups
such as those from the Indo-Pacific
rainforest region are known to have
used nutcrackers, grindstones, and
special baskets to leach toxic species
and thereby cope with these dietetic
constraints.18 Moreover, many tuber,
fruit, and nut species, including the
genera Dioscorea, Panda, Poga, Dacryoides, Parinari, Ricinodendron, Irvingia, and Coula, are good sources of
starch and fat that do not require detoxification.
There is an almost complete lack of
industrial, chronological, and environmental evidence from the Acheulian period in the Congo basin or West
Africa. The few sites that there are
belong to marginal regions outside
the lowlands, present serious stratigraphic disturbances, or could be late
phenomena that are chronologically
and behaviorally closer to the middle
Pleistocene and Middle Stone Age

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than to the early Pleistocene. Acheulian assemblages, however, have been

retrieved from present-day lowland
rainforests, such as those in the Central African Republic, at the site of
Ngolo, in the Ngoe re basin.19 This site
yielded a homogeneous biface and
cleaver assemblage consisting of 43
bifacial hand-axes and 21 cleavers
(Fig. 1). Acheulian hand-axes from
Ngolo comprised cordiforms, amygdaloids, lanceolates, ficrons, and
ovates.19
In South Central Africa, there is the
site of Kamoa (Fig. 1), an Acheulian site
in the savanna-woodland-gallery forest
mosaics of the Southern Democratic
Republic of Congo.20 Here Cahen20 excavated an assemblage in a stone-line
context overlaid by Middle and Later
Stone Age materials. Palynological
analysis by Roche21 identified Poaceae
along with Pteridophytes, but also documented gallery forest trees and vines
from rainforest genera and families
such as the Bygnoniaceae, Combretaceae, Erythrophleum, Euphorbiaceae,
Ebenaceae, Brachystegia, and Uapaca.
This type of pollen assemblage was
interpreted as proof of Acheulian occupation at a time when the landscape
was relatively open but also included
woodlands.
Kalambo Falls, located in Zambia,
near lake Tanganyika, was excavated
by Clark22 in the 1950s and 1960s (Fig.
1). The site provided a complete sequence of paleolithic occupations
comprising the Acheulian, Sangoan,
Middle Stone Age with points, Later
Stone Age, and Iron Age. The site has
a complex stratigraphy that includes
several episodes of alluviation, colluviation, and stone-line formation.
Van Zinderen Bakker,23 based on analysis of pollen from the site, sees the
Acheulian occupation of Kalambo Falls
as one in a paleoenvironmental setting
characterized by riparian forest and
seasonal dry forest. This open forest
scenario is confirmed by wood anatomical identification of the famous burnt
logs and fruits retrieved from the Sangoan levels that overly the Acheulian
occupation. These include wellknown African forest genera such as
Cynometra, Isoberlinia, Syzygium,
Parinari, and species from the Annonaceae family.23
In sum, early hominins probably

lived in African rainforests. Because

no Oldowan or Acheulian site has
been excavated in the African lowland
rainforest, only future work in pertinent areas within the tropical forest
will show whether this apparent scarcity of Early Stone Age materials from
the Congo basin reflects a population
lacuna14 rather than limited archeological research.

THE MIDDLE STONE AGE

Recent theories perceive the Middle
Stone Age as biologically and culturally connected to anatomically modern humans with enhanced cognitive
abilities.15 McBrearty and Brooks15
believe that this superior cognitive
ability provided a sophisticated and
diverse technology that may have provided new opportunities to colonize
challenging habitats, such as the tropical rainforest, that previously were
unoccupied by Homo. Nonetheless, it
has been suggested that technological
breakthroughs in stone technology
were not needed to colonize rainforests, and that stone technologies from
rainforests show no strong ecological
signature.24
The industrial transformations that
characterize the transition from archaic to modern industries are represented by the wide reliance on flake
industries produced by simple percussion and prepared core technologies,
as well as the decline of the Acheulian
complex; the early appearance of
point technologies and their regional
diversification; and, in some instances, the early use of large blades
and hafted composite tools with microliths and geometrics.15
The so-called Sangoan complex
constitutes the strongest evidence of a
post-Acheulian presence in the African tropics, as it appears throughout
vast equatorial regions of Central Africa, West Africa, Western East Africa,
and South Central Africa. The Sangoan is characterized by its heavyduty elements, including the characteristic large bifacial subtriangular
pick, the massive core-scraper, a
coarse Acheulian-like biface, and the
absence of cleavers (Fig. 2). Some authors see the Sangoan as part of the
Middle Stone Age and perceive it as a
departure from Acheulian technologies.14 Others argue that the Sangoan

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Evolutionary Anthropology 119

Figure 1. Early Stone Age in the African humid tropics. a: View of the 1956 excavation of Acheulian levels at Kalambo Falls, Zambia22; b,c:
Acheulian materials excavated by Bayle des Hermens in the lowland rainforests of Ngolo, Central African Republic19; d,e: Acheulian
materials from the forest-bordering site of Nzako, Central African Republic19; f: Various Acheulian materials from the forest-woodlandsavanna mosaics of Kamoa, Southern Democratic Republic of Congo.20

120 Evolutionary Anthropology

is an Acheulian-related complex of archaic nature that existed before a fullblown Middle Stone Age15,25 and,
though it is of uncertain age, probably
is older than 270,000 years.14 It has
been recently argued that the Lupemban, rather than the Sangoan,15
is the first industrial complex that is
representative of modern behavior.
Regardless of the biological and behavioral affiliation of the latter, it is
clear that the shifts that mark the end
of the Acheulian and the beginning of
a new period took place in the Middle
Pleistocene.14,15
Sangoan industries from the rainforest region have been reported on
the Ivory Coast, at Bete and Guabuo,
and are dated by thermoluminescence
to 254,000 years.25 Palynological
records from neighboring marine
cores indicate the existence of open
forest conditions at Bete, near
Anyama (Abidjan, Ivory Coast), and
closed forest at Guabuo, in the Sassandra river basin, Western Co te
dIvoire.26,27 Clist,28,29 Bayles des Hermens, Oslisy, and Peyrot,30 and
Locko31 have reported several Sangoan assemblages in the lowland forests of Gabon at Me doumane, Kango,
Okala, and Okanda. Some of these assemblages have yielded paleoenvironmental evidence in the form of charcoal from rainforest trees. In the
tropical forest of Southern Cameroon,
Omi32 presented large Sangoan collections with abundant picks. In Southwest Cameroon, Mercader and
Marti33 discovered the site of Njuinye,
50 km southeast of the lake BarombiMbo, where Maley and Brenac34 documented forest pollen from the last
28,000 years. Njuinye yielded a stratigraphic sequence deeper than four
meters and basal industries of possible Sangoan affiliation that could
reach far back into the past, given that
the materials are more than two
meters beneath a radiocarbon date of
34,700 years.35 Lanfranchi36,37 published information on the sites of
Ouesso and Mokeko in the lowland
forests of the Peoples Republic of
Congo. In sum, the Sangoan sites
from Co te dIvoire, Gabon, Cameroon,
and the Peoples Republic of Congo
represent rainforest occupation by
Sangoan groups because they all fall
within reconstructed rainforest paleo-

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vegetation based on palynological and

geological data proposed by Jahns,
Huls, and Sarnthein,26 Maley,34,38
Thomas,39 and Dupont and coworkers.27
The Sangoan is also represented in
open forests. The best known Sangoan site in such a context is Asokrochona,40 currently located in the seasonal
rainforest-woodland-savanna
mosaics of coastal Ghana, West Africa. In a recent revision of early Ghanaian sites, Casey41 has highlighted
the difficulties of determining both
the cultural affinities of the Ghanaian
Sangoan with the Sahel and Central

. . . there is a strong
case for the extensive
geographical
distribution of Middle
Stone Age sites across
the African rainforest,
but the temporal
dispersion of these sites
remains unclear.
Sangoan and Lupemban
sites have been found in
areas that, on
palynological grounds,
were covered by
rainforest during glacial
and interglacial
periods, . . .

Africa and the type of paleoenvironment that may have reigned in coastal
Ghana at the time of the Sangoan occupation. Casey41 nevertheless argues
for an ecological context that was
more heavily wooded than at present,
although, like the excavators of the
site, Nygaard and Talbot,40 she explains that this Sangoan site probably
was never in dense forest.
With regard to the Lupemban, the
only two sites within the present-day
lowland rainforest and in a demonstrated
rainforest
paleoenviron-

ment27,38,39 are the one in the Ogooue

basin, in Gabon, excavated by Pommeret42 in the 1960s, and Mosumu, in
the lowland rainforests of Equatorial
Guinea, recently discovered by Mercader and Marti and dated to older than
30,000 years before the present.35,43,44
Bayle des Hermens59 found several
Lupemban sites in the Central African
Republic in regions where the natural
vegetation in the recent past was tropical forest. Nzako yielded large lanceolates made of quartzite plaques
with distinctive steeply serrated edges.19 These sites also provided coreaxes and prepared-core flakes.19 Also
in close proximity to rainforest,
around 100 km north of the presentday Ituri forest border, is the site of
Lodjo.45 Discovered in the 1940s and
reported by Van Noten45 in the 1980s,
this site in the forest-savanna mosaics
of the Northeastern Democratic Republic of Congo exhibits large daggers.
The sites of Masango46,47 and Muguruk,48 in Burundi and Kenya, respectively, represent a Lupemban settlement that, although maintaining a
close techno-typological affinity with
the industries detected at sites in the
lowland forest, existed in open environments. This is also the case with
the Lupemban assemblages excavated
by Cabu (cited in de Maret49) Cahen,50
and Van Moorsel,51 in the Kinshasa
plain at Gombe point, the Kasai-Lukenie group, LUpemba, and Kamoa, all
in the Democratic Republic of Congo.
Lunda, in Angola, and Kalambo
Falls22 and Twin Rivers in Zambia52
also represent Lupemban occupations
in relatively open environments.
To summarize, there is a strong case
for the extensive geographical distribution of Middle Stone Age sites across
the African rainforest (Fig. 3), but the
temporal dispersion of these sites remains unclear. Sangoan and Lupemban sites have been found in areas that,
on palynological grounds, were covered
by rainforest during glacial and interglacial periods, suggesting that Middle
Pleistocene humans had the behavioral
capacity to inhabit the rainforests on
the Western Ivory Coast and in Southern Cameroon, Equatorial Guinea, and
Gabon. Additional sites in the proximity of rainforest indicate similar adaptations to the variably forested environ-

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Evolutionary Anthropology 121

Figure 2. Sangoan materials from: a,b: Bete, Ivory Coast, West Africa25; c: Tools from Asokrochona, Ghana, West Africa40; d: Sangoan pick
from the rainforest site of Okanda, Gabon31; e g: Sangoan tools from Zambia57; h: Chopper/pick from the lowland forests of Njuinye,
Southwest Cameroon.35

122 Evolutionary Anthropology

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Figure 3. Acheulian and Middle Pleistocene sites in the African rainforest and outliers.

ments of the Central African Republic

and Northeast Congo. In any case, Sangoan and Lupemban groups with similar technologies also occupied the open
environments of the Western Rift, the
Great lakes, and South Central Africa.

LATER STONE AGE

The Later Stone Age was a crucial
period in the development of tropical

forest cultures, for it was then that the

consolidation of skills obtained during previous phases was able to support the most extensive settlement of
rainforests in pre-agricultural times,
reaching unprecedented site densities
and early developments of continental
significance. There is a little doubt
that at least some Central African forest regions were settled before the onset of the Holocene, and that some of

those populations inhabited rock shelter site53 that existed in variably

closed forest environments across the
Pleistocene to Holocene boundary.54
Direct paleobotanical data to demonstrate that these foragers inhabited
rainforests comes from the Ituri region.54 Three phytolith sequences
from Later Stone Age sites show high
percentages of arboreal rainforest
taxa in association with cultural re-

Evolutionary Anthropology 123

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mains dated to the last 18,000 years,

indicating a sustained hunter-gatherer settlement of the lowland forests
in the Northeast Congo basin many
millennia before farming appeared in
the region.
Virtually all twelve countries that
make up the African rainforest show a
dense Later Stone Age settlement.
Some of the most important sites
from this period include, in West Africa, the sites of Bosumpra in Ghana,55 and Yengema Cave in Sierra Leone.56 In Central Africa alone, not
including South Central Africa, Mercader and Marti35 have listed more
than fifty Later Stone Age sites with
radiometric dates. This archeological
evidence appears in a great diversity
of environmental contexts that include lowland evergreen forests, seasonal forests, dry forests, mosaics,
woodlands, and open landscapes in
Western Cameroon; the Teke plateau
and the Niari basin in the Peoples
Republic of Congo; the Southwestern
Central African Republic; Equatorial
Guinea; coastal Gabon; and the
Southern, Central, and Northeastern
Congo Basin in the Democratic Republic of Congo.
Palynological data from Atlantic
Central Africa and West Africa, as well
as phytolith data from the eastern
Congo basin, indicate that Later
Stone Age hunter-gatherers inhabited
glacial forest environments and continued to inhabit rainforests after the
climatic and vegetational reassortments that took place at the onset of
the Holocene. In short, the environmental and archeological evidence
from Later Stone Age sites in Central
Africa indicates that neither glacial
nor interglacial forests were a cultural, economic, or physical barrier to
African foraging groups of the late paleolithic period.

TROPICAL FORESTS IN HUMAN

EVOLUTION AND DISPERSAL
Although the archeological record
in rainforests is especially vulnerable
to bioturbation and rapid decay, archeological resolution in tropical forests is variable and lies at different
points along a taphonomic continuum that ranges from complete destruction to optimal preservation of

assemblages and stratigraphic integrity. Not all rainforest sites are severely disturbed sites. The diagenetic
destruction of bone resulting from
burial in the acidic soils of the rainforest remains poorly understood,
and shell, charcoal, opal silica,
charred parenchyma, and starch
grains do not respond to acidity the
same way bones do.
A deficient database and the need to
rely on isolated records to reconstruct
the environmental and cultural history
of large regions have inevitably led to
generalizations and oversimplifications
of a complex reality. Thus, it is often
overlooked that paleobotanical, geological, and archeological data often used
to reconstruct past rainforest adaptations and site formation do not contain
a single record from the lowland forest
per se, and that the records used are
from seasonally arid forest-bordering
locations, even savannas, sometimes
separated from the lowland forest by
distances of more than 500 km. In environmental terms, it is clear that central African rainforests have experienced dramatic changes over time, that
the effects of glacial dryness and cooling on rainforests are not well understood, and that early, middle, and late
Pleistocene glacial episodes have not
always supported arid climates and
open ecosystems. Indeed, large regions within the present-day rainforest zone maintained tropical forest
through glacial times. Whether these
forested areas were small refugia surrounded by savannas or parts of a
wider forested matrix is controversial.
Fossil landforms such as the Kalahari sands and stone-lines are not
direct indicators of aridity within the
lowland forest, at least not with the
implications this word has in modern
climatology and ecology. Wind-blown
Kalahari sands are absent from
large lowland forest regions, and their
origins and intermittent nature are
not understood. Stone-lines have a
diverse genesis, and not all of them
formed under arid climatic conditions.
I have suggested that we are forest
people, and that our link with the
rainforest perhaps started early on, as
hinted by the East African hominid
evidence. Unfortunately, lack of research within the central African low-

lands makes it impossible to evaluate

the assumption that Miocene and
Pliocene hominids inhabited the
Guineo-Congolian rainforest. It is not
the capacity of early bipeds to exploit
rainforests that is in question, but the
biological and behavioral capacity of
Homo ergaster to settle rainforests.
Speculation on this matter, however,
could derive from lack of data, not
from direct archeological evidence
suggestive of biological and behavioral incapacity of early Homo to settle lowland rainforests. Middle Stone
Age sites in a reconstructed rainforest
paleoenvironment increase exponentially as we get closer in time to the
late Pleistocene. As fragmentary as
the data are, it is reasonable to say
that modern humans shared a long
tradition of tropical forest settlement
across the African tropics, with some
indications of expanding territorial
breadth, residential stability, and progressive ecological control of tropical
forests. Middle and Later Stone Age
foragers had the cognitive ability and
economic flexibility to inhabit a
sparsely occupied region with stone
technologies similar to those in use
outside the rainforest. Further, they
supported apparently successful adaptations to the most complex and biologically diverse biomes of the
planet.

ACKNOWLEDGMENTS
I am grateful to John Fleagle for his
patience and advice; to Sally
McBrearty and three anonymous reviewers for their comments; to Alison
Brooks,
Melissa
Panger,
Gary
Schwartz, Rene Bobe, Raquel Mart,
and Dennis Knepper, for their support
and ideas.

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