Opinion

Red ochre and shells: clues to human

evolution
Carlos M. Duarte1,2
1

The UWA Oceans Institute and School of Plant Biology, University of Western Australia, 35 Stirling Highway, Crawley 6009,
Australia
2
Department of Global Change Research. IMEDEA (CSIC-UIB) Instituto Mediterraneo de Estudios Avanzados, Miquel Marques 21,
07190 Esporles, Spain

The 200-kiloannus (ka) use of red ochre and shells by

humans is interpreted as a simple clue of symbolic
thinking. Integration of multiple lines of evidence supports the opinion that the use of red ochre and shells
might have had direct significance for human evolution.
Use of seafood and red ochre supplies docosahexaenoic
acid (DHA), possibly iron, and other essential nutrients
for brain development and reproductive health, improving human fitness and triggering brain growth. The
fitness advantages to humans of using shells, and possibly red ochre, might have selected for artistic and
symbolic expression, and, thereby, lead to social cohesion. Current global health syndromes show that an
adequate supply of seafood and iron continues to play
a fundamental role in human health.
A 200-ka bond between shells, red ochre, and human
evolution
Recent discoveries providing evidence of an early [> 200 ka;
1 ka = 1000 years before present (BP)] and independent
development of symbolic expression in modern humans
and Neanderthals [15] have important implications for
our understanding of human evolution. This evidence relies
heavily on the ornamental use of shells and red ochre [17],
whose combined use by humans elapses over 200 ka, tracking human dispersal and cultural diversification to date
(Figure 1; Box 1). However, the use of shells and red ochre
is largely interpreted as a simple clue to symbolic and
cognitive thinking, which carries evolutionary significance.
In this opinion article, I bring together recent paleoanthropological, historical, physiological, oceanographic, and ecological evidence supporting the opinion that the 200-ka bond
between the use of red ochre and shells by humans might
have direct evolutionary significance, reflected in the benefits that a seafood diet supplemented with a source of iron,
supporting reproductive and brain health, might have provided for human fitness and evolution.
Human use of red ochre and shells
Use of red ochre and shells by modern humans have been
dated to a minimum of 160 ka in Blombos and Pinnacle
Corresponding author: Duarte, C.M. (carlos.duarte@uwa.edu.au).
Keywords: Human evolution; iron; shells; seafood; nutrition; fitness.
0169-5347/
2014 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.tree.2014.08.002

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Point caves, South Africa [1]. Evidence of early (ca. 200

250 ka) use of shells and red ochre by Neanderthals has
been derived near Maastricht, The Netherlands [5] and the
Benzu rock shelter (Ceuta, Spain) [3], although these
findings provided no evidence for their symbolic use. Evidence for the significance of these items for early humans
include the discovery of a 100-ka factory in Blombos Cave
to produce red ochre pigments, which were stored in abalone shells [8], and evidence of trade for both these elements dating 80 ka [9]. Evidence of the symbolic use of
shells and red ochre by Neanderthals dating 50 ka was
derived from caves in the Iberian Peninsula [10]. Reports of
the joint use of red ochre and shells by humans are rapidly
growing in number (Figure 1), with cultural practices
involving the joint use of red ochre and shells extending
to date (Box 1).
A hypothesis linking shells and red ochre with human
nutrition, health, and evolution
The sustained use of shells and red ochre for over 250 ka
(Figure 1) suggests more direct and profound consequences
for human evolution than just evidence for symbolic thinking. Use of shells indicate the use of seafood, an early
underpinning of the food security of humans. Seafood
can be gathered with little risk or technology in the intertidal zone, and is available in great quantities throughout
the year in productive shorelines, such as the coastline of
South Africa, where modern humans were first located,
and the Indian Ocean, along which humans first dispersed
[11]. Use of seafood allowed humans to cope with a climatically adverse period between 195 and 130 ka ago [9], and
provided a reliable food source along the subsequent coastal dispersal of humans [11]. Seafood is rich in DHA, a
polyunsaturated essential fatty acid with a fundamental
role in brain growth, function, and evolution [1215]. In
addition, the marine diet is a key source of other oligoelements with a key role in brain health, such as iodine and
lithium. Iodine plays a significant role in brain development, with iodine deficiency, conducive to brain damage
[16], remaining the most frequent cause worldwide, after
starvation, of preventable mental retardation in children
[17]. Lithium is another trace element also present in
seafood whose role as a human nutrient has been recently
proposed [18], and is now used to treat bipolar disorder and
neurodegenerative diseases [19]. Therefore, the hypothesis
that the early use of the marine food web by humans, rich

Opinion

Trends in Ecology & Evolution October 2014, Vol. 29, No. 10

250

50 *
250*

12

23
80

0.5
1.8

92

135

10
5
35

3
40

12
160
Key:

Historical evidence
Paleoanthropological evidence (* = Neanderthals)
TRENDS in Ecology & Evolution

Figure 1. Reports of the joint use of red ochre and shells by humans are rapidly growing in number [40], including reports for both modern humans and Neanderthals
(asterisks). Existing records track the reconstructed dispersal of modern humans, including findings of the joint use of red ochre and shells in Africa, Eurasia, Australia, and
America, ranging from paleorecords starting in 250 ka (red arrows, paleorecords) to present (blue arrows, historical records, Box 1). For paleoanthropological evidence, the
numbers show the dates in ka, where reported [40]. When multiple reports were available for a given region, the oldest one is shown.

in omega 3, DHA, and other critical nutrients for healthy

brain development triggered the evolution of the large
human brain [20], the cornerstone of human evolution,
is now receiving ample support [1315,20].
By contrast, the role of red ochre in human evolution
remains confined to the development of symbolic and
artistic expression, as the use of such material (red ochre)
is unlikely to have had any value other than symbolic [21].
However, I suggest in this article that use of red ochre
might have also played a similarly direct early role on
reproductive and brain health and, therefore, human evolution. Whereas there is solid evidence for the benefits of a
seafood diet for brain development and human evolution,
the hypothesis linking the use of red ochre with human
iron nutrition has not been examined before. Whereas
recent evidence suggests that this is a plausible hypothesis
(Box 2), it needs be tested in the future.
Homo erectus was interested in red ochre since at least
1500 ka [21], and contemporary humans remain attracted
to the color red [22], (Box 1). Australian aborigines, the
oldest living culture, assign great health benefits to red
ochre [21], which used has been linked with medical practices [23], and traditional Arab, Chinese, Egyptian, Greek,
and Roman medicinal practices involved iron, despite the
fact that the role of iron as an essential nutrient was only
demonstrated by Boussingault in 1872 [24]. Findings at
Blombos Cave showed that red ochre was powdered using
grindstones and hammerstones and mixed with bone marrow and charcoal to derive a mixture [8]. Evidence that
Neanderthals also used ochre prepared as an ochre-rich
liquid substance has also reported [5]. The ingestion of
mixtures of red ochre and bone marrow, deliberately or
through contamination, would represent an iron-fortified
food, comparable to fortified foods used today [25], as
recent evidence shows that protein coatings increase the
bioavailability of the metastable ferric forms present in red

ochre (Box 2). Whereas ferric iron forms present low bioavailability, ferric iron forms present in red ochre, other
than hematite, can be assimilated, particularly when combined with protein (Box 2). Indeed, ferric forms present in
red ochre, such as ferrihydrate and maghemite, are used in
pharmaceutical products to treat iron deficiency anemia in
humans (Box 2). Iron deficiency anemia remains a serious
heath issue affecting humans, particularly women, in both
developed and developing nations, increasing the risk for
preterm labor and infant mortality, and accounting for
much of maternal deaths during pregnancy and childbirth
[24,25]. Today, approximately 70% of women emerging
from pregnancy suffer iron deficiency anemia [24,25].
Two iron-rich proteins, the oxygen transport protein
hemoglobin, and the iron storage protein ferritin, which
account for 70% and 25% of the iron in the human body,
respectively, play a key role in brain health and function
[26,27]. Hemoglobin carries oxygen from the lungs to the
brain, which accounts for 20% of the total oxygen consumption with only 2% of body weight [24,26], and a smaller
share of the metabolically-active mass. Iron supply-supporting loads of hemoglobin to satisfy brain oxygen
requirements is, as is that of omega 3 and DHA, particularly critical during the early development of the brain
[24,26]. Ferritin, particularly enriched in the human brain
(30% of total ferritin) [27], plays a role in storing and
releasing iron, buffering against iron deficiency and overload, and is particularly important for brain iron homeostasis and health [2428]. Inside the ferritin shell, iron ions
form crystallites similar to the mineral ferrihydrite, an
iron oxide present in red ochre (Box 2). Therefore, red ochre
can be, depending on the fraction of metastable sources and
mixing with foods, a source of iron that can be stored in
ferritin in the human brain, supplying iron where necessary to maintain adequate levels of hemoglobin and, therefore, oxygen supply for brain function.
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Trends in Ecology & Evolution October 2014, Vol. 29, No. 10

Box 1. Cultural use of red ochre and shells

The most spectacular evidence of early ornamental use of shells and
red ochre is provided by the finding at the Arene Candide cave in the
Ligurian Coast, Italy, of the skeleton of an adolescent male, known as
Il Principe (dated 23.4 ka), spectacularly ornamented in a bed of red
ochre and his head surrounded by hundreds of perforated shells,
originally forming a kind of cap [41] (Figure IA). The joint use of shells
and red ochre has remained embedded in cultural practices of
human societies across the world for millennia, ranging from Masai
and Sudan societies in Africa, to Australian aborigines [21] and New
Zealand Maoris. Shells and red ochre were also widely used in the
Americas by Aztec and Mayan cultures, and by South American and
North American Indians, such as the Beothuk in Newfoundland, and
the Wabanaki nations (Figure 1), which were named red skins by
early European explorers because of the ceremonial use of red ochre
to stain their bodies [42]. The importance of red ochre in Beothuk
culture was such that disgraced tribal members were ordered to
remove the pigment as a form of punishment. Red ochre, kokowai,
was used for personal adornment by Maori. A Cook crew member
recorded in 1777, They paint their faces with a coarse red paint,
and oil or grease the head and upper part of the body. DUrville
reported in 1827 that the New Hollanders (Australian aborigines)

(A)

(B)

dust their faces with powdered ochre. The New Zealanders. . . usually
use it in an oily paste, which they smear on their foreheads and their
hair. . . on feast days a New Zealander does not think he has carried
out a full toilette until he has anointed his whole body, and especially
his face and hair, with fish oil and ochre.
The Western culture is no exception and has continued to use
shells and red ochre to ornate their bodies to date. The ornamental
use of shells and red ochre as make up has been documented in all
ancient cultures, from ancient Mesopotamia, Egypt and Greece, to
Rome [43]. Noble women in the Roman empire used red ochre
extensively in their make up, leading Plautus to assert that, A
woman without paint is like food without salt. Indeed, classical
Romans called the most powerful men in the city coccinati, the
ones who wear red, and the color red signaled at status and power
across a broad range of societies [21]. Portraits show a continuity in
the use of red ochre make up by western women (Figure IB).
Indeed, red ochre, currently reported as FDA-approved cosmetic
Pigment Red 101, remains a key element of modern womens make
up and is present in a broad range of cosmetic products, often worn
along with nacre and pearl jewelry from oysters and other bivalves
(Figure IC).

(C)

TRENDS in Ecology & Evolution

Figure I. (A) Photos of the skeleton of a young man dated 23.4 ka, know as Il Principe, showing the perforated shell cap and mass of ochre between the left clavicle and
mandible. (B) A portrait of Marie Antoinette, showing red ochre make up. (C) A young woman wearing hematite, red ochre, pigmentation and shell ornaments. Photos
reproduced with permission from Antiquity Publications Ltd from a figure printed in [41] (A), and Carlos M. Duarte (C).

The marine food web, although rich in many essential

nutrients, is relatively poor in iron [29], as iron supply
limits biological production over much of the coastal [29]
and open [3032] ocean. Iron availability changes over
geological time in association with climatic oscillations,
with relatively high supply to the ocean during cold, glacial
periods, and low supply during warm periods [33], when
iron plays a particularly important role as a nutrient
limiting ocean productivity [30,32]. Manipulation of red
ochre could have, therefore, provided an iron supplement
to support healthy brain function and development under
situations of iron limitation, as possibly experienced by
pregnant women on a marine diet. Because iron is an
important mineral in mineralization, ferritin is employed
in the shells of organisms such as mollusks to control the
concentration and distribution of iron, thus sculpting shell
morphology and coloration [34]. Therefore, shells can be,
themselves, a source of ferritin, and mixing some red ochre
with seafood could have delivered essential nutrients for
brain development while alleviating iron deficiency. There
562

can be additional benefits in the use of red ochre for

humans inhabiting the coastal zone [23], as covering human bodies with red ochre also protects the skin from solar
radiation and mosquito bites and, when mixed with fat, as
the Maori applied it (Box 1), it provides thermal insulation
for humans diving in search of seafood.
The maternal circulation is depleted in iron, iodine,
omega 3, and DHA during fetal growth in order to construct
placental and fetal tissues able to deliver the huge oxygen
and energy supply required for the development of the fetal
brain [16], leading to recommendations for the use of iodine
[17], DHA [35,36], and iron [24,25] supplements in pregnant women to improve both the mothers and babys
health. Therefore, the combined use of seafood and red
ochre could have supplied omega 3, DHA, iodine, iron, and
other key nutrients essential for brain development and
reproductive health, thereby improving the reproductive
success of women and providing a direct advantage in
terms of fitness. Specifically, enhanced DHA and iron
supply associated with the combined use of seafood and

Opinion
Box 2. Can iron in red ochre be assimilated?
Whether the handling and accidental use of red ochre, alone or
when mixed with bone marrow and other foods, is conducive to iron
assimilation remains to be tested directly, a test that could involve
examination of the iron status of indigenous cultures that continue
to use red ochre in their traditional practices. Most of the iron
contained in red ochre occurs as hematite (aFe2O3), a stable,
oxidized form of iron that is insoluble and which is believed to be
unavailable [44]. However, red ochre also contains variable proportions of other, nonheme meta-stable forms of iron, such as:
ferrihydrite, magnetite, and maghemite [4547], particularly when
depositing under acidic pH [48] as reported in the Kokowai Springs
exploited by Maori, where ferrihydrite is the dominant iron form
[45]; in the materials used in the Pecos River rock paintings [46]; in
artifacts found in Malawi [49]; or in Pinnacles Cave, South Africa
[50]. These metastable forms of iron are bioavailable, although with
variable assimilation efficiency, depending on factors such as the
relative amounts of chelators and ligands (e.g., phytate and citrate),
pH, the amounts of reducing compounds ingested, and pre-heating
[44,51]. Mixing of ferrihydrate with food, as documented in early
uses [8], would be particularly effective, as ferrihydrate would be
coated with protein to yield a ferritin analog [52,53]. Indeed, the
paradigm of iron absorption is shifting due to recent developments
in nanomedicine providing evidence that the dissolution of
ferrihydrate may not be necessary for absorption to take place
[53]. This knowledge has been used recently to develop ferrihydritebased [53] and maghemite-based [54] supplements, combining
these forms of iron with ligands, to fortify foods and are included in
iron-based parenteral drugs used to treat iron deficiency anemia,
such as Ironate1 and Feraheme1 [55].

red ochre would have provided the building materials for a

large and healthy brain and the capacity to support the
supply of oxygen required to support brain development
and function. Therefore, the combined use of red ochre and
shells might have triggered evolutionary processes supporting the exponential growth of the human brain encompassing the period between 200 and 50 ka BP [21,37],
conducive to the emergence of the cognitive and symbolic
capabilities that characterize modern humans.
Can fitness advantages associated with the use of red
ochre have indirectly selected for artistic and cognitive
expression?
Fitness, the ability of organisms to contribute genes, involving survival and successful reproduction, to the next
generation, is a master driver of natural selection [38] and,
therefore, evolution. Whereas the purpose of collecting
shells arguably involved food provision as well as symbolic
use, the collection and use of red ochre could not be directly
associated with feeding [21]. However, early use of red
ochre for artistic purposes involved its use in powdered
form, often mixed with animal oil and fat to confer adherence [8], and applied with the human hands. Therefore,
human groups using red ochre would live in environments
enriched in iron and the artists would have their fingers
covered in red ochre, thereby ingesting iron, some of it
possibly bioavailable (Box 2) when eating. This combination might have particularly enhanced the reproductive
health of women and the healthy brain development of
babies, which largely occurs during pregnancy, thereby
increasing fitness.
Whereas most arguments linking symbolic thinking
with human evolution do so through the indirect benefits
of social cohesion on fitness, the argument strikes me as

Trends in Ecology & Evolution October 2014, Vol. 29, No. 10

somewhat circular, as the onset of the use of elements

involved in ornamental and artistic expression, such as
shells and red ochre, could not have been triggered by an
anticipation of the long-term benefits of symbolic thinking.
The perspective provided here suggests a different casual
pathway, with a direct link between use of shells and red
ochre and human fitness and brain growth, eventually
leading to cognitive and symbolic thinking and the development of social structure. The suggestion in this opinion
article is that the improved fitness of women ingesting
omega 3 and DHA-rich seafood and iron-rich red ochre,
because they used shells and red ochre for symbolic purposes, could have indirectly selected for artistic and symbolic expression, also providing key benefits in terms of
communication and social cohesion, which have played an
essential role in human evolution [6,7,21]. The discovery of
elaborated workshops to process red ochre and deposit the
powder in shells [8] suggests that the individuals involved
spent substantial time working in their caves to process
these materials and decorate their walls and their own
bodies. A further speculation, for which no evidence is
available as yet, is that some of these early artists might
have been women in advanced pregnancy stages, with
reduced mobility and agility to gather food outside the
cave. Indeed, it is women and their babies who benefit the
most from dietary supplements in DHA, omega 3, iodine,
and iron [17,24,25,35,36].
Concluding remarks: the significance of understanding
the role of red ochre and shells in human evolution
I acknowledge that the opinion put forward here is speculative and could, therefore, be proven wrong. However, I
believe it provides a plausible hypothesis, consistent with
paleoanthropological evidence, history, oceanography,
physiology, and human health, and is worth considering.
Whereas evidence of a significant use of the marine food
web by early humans [39], and for the role of seafood and
ingestion of DHA on brain development and human evolution, is now particularly robust, the hypothetical role of
red ochre, through iron oxide forms such as ferrihydrate, on
iron nutrition has not yet been tested. The alternative view
that the significance of the combined use of red ochre and
shells by humans is limited to a clue for symbolic expression appears insufficient to explain why these two elements, shells and red ochre, have been combined for
200 ka and remained combined in so many cultures around
the planet, given that so many alternative objects and
materials could be used to express symbolic thinking. I
submit that the overwhelming evidence for the role of
essential nutrients contained in seafood and iron in the
reproductive and brain health of extant humans, in particular, involving the health of the mother and the newborn,
amounts to a partial validation of the hypothesis put
forward, as it provides solid support for the notion that
these two sources of essential nutrients have a positive
impact on human fitness, and have, therefore, evolutionary
significance.
Understanding the connections between the 200-ka
trajectory of the combined use of red ochre and shells
and human evolution is of consequence because it could
also suggest a pathway for a healthy future for human
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Opinion
kind. The sharp rise in brain disorders, which, in many
developed countries, involves social costs exceeding those
of heart disease and cancer combined, has been deemed the
most worrying change in disease pattern in modern societies, calling for urgent consideration of seafood requirements to supply the omega 3 and DHA required for brain
health [15]. Likewise, iron deficiency is the most common
form of malnutrition in the world, affecting more than 2
billion people globally [24,25]. These disorders affect
humans across developed and developing nations. In conclusion, the deep and sustained connection between human use of red ochre and shells summarized here delivers a
new perspective on the role of seafood and iron nutrition on
the health of the human brain and human fitness, critical
to both understanding our past evolution and addressing
our future health challenges.
Acknowledgments
I thank J. Erlandson for useful comments, G. Duarte for help with
Box 1, and N. Faria, G. Anderson, E.C. Theil, and C. Hutchinson for
advice on Box 2.

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