Not the brain alone: The nutritional potential of elephant heads in Paleolithic sites

Abstract

The presence of elephants, and specifically of elephant head remains, is well demonstrated in many Paleolithic sites in Europe, Africa, and Asia. However, the possible mechanisms for the exploitation of this enormous body part are rarely discussed, and it is often suggested that elephants' heads were exploited specifically for the extraction and consumption of the brain. In this paper, we discuss the nutritional potential that lies within elephants' heads as implied by ethnographic and zoological literature, and present archaeological evidence from Paleolithic sites for the exploitation of proboscideans' heads. The data show that the prevailing view should be re-evaluated, and that the nutritional potential within the elephant's head extends far beyond the brain. We suggest that organs such as the temporal gland, the trunk, the tongue, the mandible and the skull itself were exploited routinely as an integral part of early humans' diet. The nutritional potential of the elephant head provides a parsimonious explanation for the investment early humans put into transporting and exploiting this specific body part at open-air sites but particularly at cave sites, and serves as a significant beacon in understanding Paleolithic human behavior in relation to proboscidean remains.

Full text

Not the brain alone: The nutritional potential of elephant heads
in Paleolithic sites
Aviad Agam
*
, Ran Barkai
Department of Archaeology and Ancient Near Eastern Cultures, Tel Aviv University, P.O.B. 39040, Ramat Aviv, Tel Aviv 69978, Israel
article info
Article history:
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Keywords:
Proboscideans
Elephant skulls
Nutrition
Paleolithic
Temporal gland
Fat
abstract
The presence of elephants, and specifically of elephant head remains, is well demonstrated in many
Paleolithic sites in Europe, Africa, and Asia. However, the possible mechanisms for the exploitation of this
enormous body part are rarely discussed, and it is often suggested that elephants' heads were exploited
specifically for the extraction and consumption of the brain. In this paper, we discuss the nutritional
potential that lies within elephants' heads as implied by ethnographic and zoological literature, and
present archaeological evidence from Paleolithic sites for the exploitation of proboscideans' heads. The
data show that the prevailing view should be re-evaluated, and that the nutritional potential within the
elephant's head extends far beyond the brain. We suggest that organs such as the temporal gland, the
trunk, the tongue, the mandible and the skull itself were exploited routinely as an integral part of early
humans' diet. The nutritional potential of the elephant head provides a parsimonious explanation for the
investment early humans put into transporting and exploiting this specific body part at open-air sites but
particularly at cave sites, and serves as a significant beacon in understanding Paleolithic human behavior
in relation to proboscidean remains.
©2015 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
The significant role of elephants in Paleolithic faunal assem-
blages is well demonstrated in many sites in Europe, Africa, and
Asia. Proboscideans, as well as other taxa, were exploited by early
humans throughout hundreds of thousands of years across the old
world (e.g., Klein, 1988; Boschian and Sacc
a, 2010; Yravedra et al.,
2010; Zhang et al., 2010; Anzidei et al., 2011; Echassoux, 2012;
Rabinovich et al., 2012; Yuan et al., 2012; Domínguez-Rodrigo
et al., 2014b). However, the dietary significance of elephants has
not been thoroughly explored (but see Ben-Dor et al., 2011). Given
the presence of elephant remains at archaeological sites as
described above, we assume that during Paleolithic times ele-
phants, when available, were a constant and significant source of
calories for early hominins (however see Smith, 2012, 2013).
Notwithstanding its significance, this was only one source of calo-
ries among many other (plant based and animal based) food re-
sources used by Paleolithic hominins. However, in our opinion it
was a primary one when elephants were available.
While in many Paleolithic sites elephant remains and lithic ar-
tifacts are found in association, the human use of elephants for
dietary purposes is still debated in some cases (e.g., Villa et al.,
2005). Several Lower Paleolithic Acheulian sites, however, provide
clear evidence of butchery and defleshing of elephants, such as
bones bearing cut-marks and breakage signs (e.g., Goren-Inbar
et al., 1994; Wenban-Smith et al., 2006; Yravedra et al., 2010;
Rabinovich et al., 2012). In addition, the use of elephant bones in
the Acheulian for the production of tools, particularly bifaces that
resemble the characteristic stone handaxes, is clear (Biddittu et al.,
1979; Dobosi, 2001, 2003; Bruhl, 2003; Boschian and Sacc
a, 2010;
Rabinovich et al., 2012; Beyene et al., 2013; Zutovski and Barkai,
2015). Post-Acheulian as well as Mousterian sites provide further
evidence for the use of proboscideans both for dietary purposes,
such as meat and marrow consumption, (e.g., Blasco and Fern
andez
Peris, 2012; Yravedra et al., 2012), and for other tasks, such as bone
tool production (e.g., Gaudzinski, 1999; Gaudzinski et al., 2005;
Boschian and Sacc
a, 2014).
The role of protein in human diet and subsistence in Lower
Paleolithic sites has been demonstrated frequently (e.g., Bunn,1981,
2006; Shipman and Walker, 1989; Milton, 2003; Morin, 2007;
Pante, 2012; Sahnouni et al., 2013). Animal meat and fat are an
excellent source of calories, and contain essential amino acids,
*Corresponding author.
E-mail address: aviadkra@post.tau.ac.il (A. Agam).
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minerals, vitamins, and fatty acids (Friedman, 1996; Milton, 2003;
Givens et al., 2006; Williams, 2007).
As early as the Late Pliocene, we are familiar with archaeological
sites containing stone tools associated with animal bones, reflect-
ing human activities related to butchery of animals and marrow
extraction (e.g. Plummer et al., 1999; Moull
e et al., 2001; Ferretti
et al., 2003; Ferraro et al., 2013; Lemorini et al., 2014). Hence, it is
commonly accepted that Acheulian, and even pre-Acheulian
hominins, extracted a significant portion of the calories they
consumed from animal meat and fat, and thus were actually
dependent on animals for their survival (Kaplan et al., 2007; Ben-
Dor et al., 2011; Domínguez-Rodrigo et al., 2012). Hence, carni-
vory has been a human trait from our earliest stage to today (Psouni
et al., 2012).
Elephants were by far the largest terrestrial animal available for
Lower Paleolithic hominins, the straight-tusked elephant being
about five times larger than the second largest animal found at
Acheulian sites in the Levant, the hippopotamus (Ben-Dor et al.,
2011). Extinct elephants most likely weighed more than recent
elephants (Shoshani and Knight, 1992; Christiansen, 2004; Byers
and Ugan, 2005), and therefore the average weight of recent ele-
phants' body organs should be enlarged when discussing Pleisto-
cene proboscideans (Table 1). According to Byers and Ugan (2005),
based on recent Loxodonta africana samples, the cranium and
mandible can exceed 180 kg. The trunk can exceed 110 kg. The brain
of an adult elephant can reach 6.5 kg, and the tongue can exceed
14 kg. In total, a complete head of an elephant (Fig. 1), including the
ears and tusks, can exceed 400 kg. These figures should be greatly
enlarged, if not almost doubled, for Pleistocene extinct
elephants.
Elephants represent an ideal food package with a perfect
combination of meat and fat, as half of the potential calories are in
the fat (for details see Ben-Dor et al., 2011). While we have no
exact data as to the extent to which elephant carcasses were
utilized, the abundance of evidence for elephant utilization in
Acheulian sites (as well as later Paleolithic sites) is a clear indi-
cation that at least part of the elephant's potential energetic value
was extracted by early humans. This view is further supported not
only by the fact that some elephant bones found in Paleolithic
sites bear cut marks (e.g, Goren-Inbar et al., 1994; Yravedra et al.,
2010; Rabinovich et al., 2012), but also that many bones were
further fractured in order to reach the marrow (e.g, Boschian and
Sacc
a, 2014; Yravedra et al., 2014), indicating further use of the
bones for additional fat.
In the Levant, the available data on elephant remains in Lower
Paleolithic Acheulian faunal assemblages comes from a number of
sites, including Ubeidiya, Evron, Latamne, Gesher Benot Ya'aqov,
Revadim and Holon (see Bar-Yosef and Belmaker, 2011; for bibli-
ography). All of these are open-air sites (see Speth, 2012; for
discussion), while Acheulian presence in caves in the Levant is
scanty. The few cave sites with Acheulian layers are either poor in
fauna or not thoroughly studied. Outside the Levant, however,
there are a few cases of Paleolithic cave sites bearing elephant
remains, such as Bolomor cave in Spain (Blasco et al., 2013),
Ma'anshan cave in China (Zhang et al., 2010) and Spy cave in
Belgium (Germonpr
e et al., 2012). In the case of open-air sites, the
question of whether hominins gathered near elephant carcasses
resulting from natural death and/or hunting, or transported parts
of the carcasses to their home base, is difficult to answer. Cave
sites with megafauna remains, on the other hand, are clear evi-
dence for transportation of selected body parts from the kill/
acquisition site to the cave.
Many Paleolithic sites with elephant remains found in direct
association with human activity have yielded elephants' head
remains, including mandibles, skull fragments and teeth (e.g.,
Scott, 1989; Piperno and Tagliacozzo, 2001; Fladerer, 2003;
Yravedra et al., 2010, 2012; Rabinovich and Biton, 2011;
Rabinovich et al., 2012; Nikolskiy and Pitulko, 2013). However,
the nutritional potential of this body part is seldom discussed. On
the rare occasions that this is dealt with, it is mostly suggested
that the skulls of proboscideans were exploited specifically for the
extraction and consumption of the brain (Adam, 1951; Scott,
1989; Goren-Inbar et al., 1994; Germonpr
e et al., 2008). In the
case of the Upper Paleolithic site of Yudinovo (the Russian Plain),
for example, it was claimed that, “the broken skulls indicate that
humans searched for the fresh fatty brain”(Germonpr
e et al.,
2008). It is our intention to provide evidence that shows that in
fact other head parts were obtained and consumed by early
humans as well.
This paper deals with the nutritional potential within the ele-
phant's head and its implications as follows: 1. An overview of
elephant skull remains in a number of selected Paleolithic sites; 2.
The anatomy of specific organs within the elephant's head and their
nutritional potential; 3. Ethnographic evidence for the consump-
tion of different head parts of elephants; 4. A re-evaluation of the
nutritional potential of the elephant's head and its contribution to
the understanding of Paleolithic human behavior.
The elephant's head is a particularly high-quality source of
energy, bearing a considerable amount of edible tissues (Byers
and Ugan, 2005), and therefore could have been used by early
hominins as an important dietary source. Thus, understanding
the nutritional potential of elephant heads, combined with the
remains of elephant skulls in Paleolithic sites, provides new in-
sights concerning human behavior and subsistence in Paleolithic
times.
Fig. 1. A modern elephant head can exceed 400 kg, including internal organs, trunk,
tusks and ears. A secretion is visible oozing from the temporal gland. Photo by Nir
Geiger, all rights reserved.
Table 1
Mass of different head parts calculated according to samples taken from four
adult male modern Loxodonta africana (Byers and Ugan, 2005).
Head parts Mass (kg)
Brain 4.0
Cranium/mandible (excluding brain) 176.7
Tongue 13.0
Trunk 113.6
Ears 44.7
Tusk nerves 11.3
Tusks 63.2
Total head 426.5
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2. Materials and methods
Our research provides accounts of archaeological sites with
proboscidean head remains, and the possible courses of exploita-
tion of this enormous body part by early humans. In addition, a
number of anatomical characteristics of recent elephants' head are
presented, taken from zoological and biological literature, focusing
on head organs with nutritional potential ethe trunk, the temporal
gland, the mandible, and the skull itself. Also, in order to recognize
the possible trajectories of utilization of elephants' heads, relevant
ethnographic documentation regarding the hunting and con-
sumption of elephants is provided.
3. Archaeological record
As can be seen from the list of selected Paleolithic archaeological
sites presented in Table 2, proboscidean head remains have been
recovered from many significant Paleolithic sites in the Old World.
While we present selected sites with significant quantities of pro-
boscidean head remains, and with representation of different head
parts, many other sites could be added to this list. Most of those
sites are open-air sites. However, proboscidean head remains have
also been recovered at cave sites. These sites are spread over a vast
geographic terrain, from Africa, the Levant, Asia, and Western
Europe, up to the Siberian Arctic, and span a time frame of over one
million years. We present here only a brief overview of three of
those sites and the finds relevant to the goals of our argument.
Gesher Benot Ya'aqov (GBY) is an open-air Lower-Middle
Pleistocene site located in the Dead Sea Rift valley. All lithic as-
semblages were assigned to the Acheulian cultural complex
(Goren-Inbar et al., 2000). The duration of the entire depositional
sequence is estimated to be about 100 ka, bracketing the Matuya-
maeBrunhes Boundary (0.78 Ma) (Feibel, 2004). The well-
preserved finds include rich botanical material, and a varied
faunal assemblage, which includes crabs, fish, fallow deer, turtles,
and elephants (Alperson-Afil et al., 2009; Rabinovich and Biton,
2011). GBY has the earliest occurrence of Palaeoloxodon antiquus
in the southern Levant. The elephant remains include a complete
skull, a few fragments of teeth, tusks, cranium, and two pieces of
the internal surface of the braincase. In total, 154 remains of ele-
phants (NISP) were found at the site (Rabinovich and Biton, 2011).
An almost complete elephant's skull was recovered in close
association with a basalt core, a boulder, and an oak log. The entire
palatal and basicranial region of the skull had been removed, and
the remaining surface was crushed. Based on the position of the
artifacts, it was suggested by the researchers that this is a context of
a deliberate brain extraction (Goren-Inbar et al., 1994).
Revadim quarry is an open-air site located on the southern
Coastal Plain of Israel, 40 km southeast of Tel Aviv (Marder et al.,
1999). Paleomagnetic analyses of the geological sequence show
normal polarity, indicating that the entire sequence is younger than
780 ka (Gvirtzman et al., 1999; Marder et al., 2011). Preliminary
dating of carbonate coating of flint artifacts provided dates between
300 and 500 ka, providing a minimum age for these items. The
lithic and the faunal finds assigned the entire anthropogenic
assemblage to the Late Acheulian cultural-complex of the Levant
(Marder et al., 2006, 2011; Solodenko, 2010; Malinsky-Buller et al.,
2011a, b). Of special note is recent research providing evidence for
extensive lithic recycling at the site, directed mainly toward the
production of sharp small flakes (Agam et al., 2014).
Revadim's faunal assemblage consists of thousands of animal
bones, dominated by Palaeoloxodon antiquus,Bos primigenius,Dama
cf. mesopotamica, and Cervus elaphus (for more details see
Rabinovich et al., 2012). Elephants are the most dominant animal in
the faunal assemblage, with a total of 155 NISP, representing at least
six individuals. Ninety-five teeth were found throughout all layers,
as well as two mandibles, six skull fragments, 19 tusks and tusk
fragments, and one premaxilla. In addition, 10 ribs, 3 vertebrae, 3
scapulas, and 3 pelvises were found. Of special interest are two ribs
and a scapula bearing cut marks. Other remarkable artifacts are two
elephant's bones shaped as bifaces (Rabinovich et al., 2012). In
addition, fat residues and use-wear related to the processing of hide
and meat were detected on a handaxe and a scraper found in as-
sociation with butchered elephant remains at one of the best pre-
served archaeological contexts at the site of Revadim (Solodenko
et al., 2014).
Bolomor Cave (Blasco et al., 2010; Blasco and Fern
andez Peris,
2012) is located on the southern slope of the Valldigna valley,
southeast of the town of Tavernes (Valencia, Spain). The cave is a
karst cavity which was opened for the first time approximately
500 ka, located about 100 m above sea level. It is an elevated rock-
shelter incrusted on the rock wall, and it takes rigorous climbing to
get to the cave from the channel. The site is composed of seventeen
stratigraphical levels ranging from MIS 9 to MIS 5 (c. 350e100 ka).
The faunal assemblages include Oryctolagus cuniculus,C.elaphus,B.
primigenius,Equus ferus and Paleoloxodon antiquus (for more details
see Fern
andez Peris, 2007; Blasco and Fern
andez Peris, 2012), and
bear evidence of anthropogenic activity in the form of cut marks,
intentional breakage, human toothmarks, and burning patterns.
The site has yielded elephant cranial and limb bones. Among the
Table 2
Selected Paleolithic archaeological sites with proboscidean head remains.
Site name Site type Location Dating Species References
Barogali Open-air site Republic of Djibouti ~1.6e1.3 Ma Elephas recki ileretensis Berthelet and Chavaillon, 2001.
Gesher Benot Ya'aqov Open-air site Israel ~800 ka Palaeoloxodon antiquus Goren-Inbar et al., 1994; Rabinovich and
Biton, 2011.
Notarchirico Open-air site Italy ~650 ka Piperno and Tagliacozzo, 2001.
Revadim Open-air site Israel ~500 ka Marder et al., 2011; Rabinovich et al., 2012.

Aridos 2 Open-air site Spain ~380 ka Yravedra et al., 2010.
Bolomor cave Cave site Spain ~350 kae100 ka Blasco and Fern
andez Peris, 2012; Blasco
et al., 2013.
La Cotte De St. Brelade Cave site England ~150 ka Mammuthus primigenius Scott, 1989;Smith, 2015
Spy cave Cave site Belgium ~59 kae25 ka Germonpr
e et al., 2012.
Yana Open-air site Siberian Arctic ~27e29 ka Nikolskiy and Pitulko, 2013.
KremseWachtberg open-air site Austria ~27 ka Fladerer, 2003.
Yudinovo Open-air site Russia ~13e15 ka Germonpr
e et al., 2008.
Preresa Open-air site Spain ~84 ka Elephas or Mammuthus Yravedra et al., 2012.
Ma'anshan cave Cave site China ~53 kae19 ka Stegodon orientalis Zhang et al., 2010
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finds are also 11 elephant teeth. All elephants analyzed were infants
or juveniles, suggesting some preference according to age (Blasco
and Fern
andez Peris, 2012; Blasco, 2014, personal communica-
tion). Considerations such as taste, quality and composition of meat
and fat, ease of hunt and even weight may have played a part in this
preference (for a detailed discussion on the subject see Reshef and
Barkai, 2015). Cut marks were identified on a mandible of an
immature elephant from layer XII (Fig. 2), dated to about 180 kya, in
the form of five oblique incisions on the vestibular surface (Blasco,
2014, personal communication).
The topographic location of Bolomor cave creates a significant
difficulty reaching the cave from the bottom of the valley, all the
more so while carrying elephants' body parts. Nevertheless,
elephant remains, head parts included, were found inside the cave.
The great effort that surely was put into these tasks suggests that
the cranial remains brought to the cave justified the trouble in
terms of nutritional value. It may also imply the existence of
cooperation between group members.
4. Nutritional potential of the elephant's head
The available data clearly indicate that Paleolithic hominins
exploited elephant meat and fat routinely and abundantly. The
presence of head parts at many Paleolithic sites is also common,
suggesting the regular consumption of at least some parts of the
head. Furthermore, at least in some cases, as in Bolomor Cave and
other cave sites, it is clear that proboscideans' heads were trans-
ported intentionally to the camp site, and were manipulated there.
However, the nature of the exploitation of the elephant's head and
the specific organs that were actually obtained are yet to be
defined. This section presents direct quotes taken from ethno-
graphic and zoological literature, attesting to the nutritional po-
tential of different parts of elephants' heads.
The temporal gland (Figs. 1 and 3) is an organ that is unique to
elephants. It is located in the temporal depression, behind the eyes,
on each side of the head (Sikes, 1971; Rasmussen et al., 1984). It was
reported to weigh between 230 and 1590 g in bulls of different ages
(Wheeler et al., 1982). Mostly, it is related to the musth, but occa-
sionally it is also described as a scent gland (Buss et al., 1976), and
there are even reports of post-mortem secretion from the gland
(Sikes, 1971). The secretion is composed of cresol, phenol, and
cholesterol (Wheeler et al., 1982). Ancient proboscideans most
probably also possessed temporal glands, that most likely were
larger than those of the recent elephants (Buss et al., 1976; Wheeler
et al., 1982; Rasmussen, 1999; Boeskorov et al., 2007).
Peter Albl (1971) mentions a certain fat concentration that exists
in the temporal region:
“Relatively large quantities of adipose tissue fill the space between
the masseter muscles and the zygomatic arch (arcus zygomaticus),
and are also present around the temporal glands. This fatty cushion
starts in the temporal region and reaches down to the lower jaw.”
(Albl, 1971: 135).
Few ethnographic sources mention a certain hollow that is
located above the eye of the elephant, from which fat can be ob-
tained fairly easily. Cuthbert Christy (1922), for example, describes
his elephant hunting habits in Africa. Christy explains:
“The only fat obtainable in any quantity from a dead elephant,
before the carcass is cut up, is that beneath the plantar arch of the
foot, and in the hollow above the eye.”(Christy, 1922:291e292).
Fig. 2. APaleoloxodon antiquus mandible, found at Bolomor Cave, with cut marks on it
(marked by the white frame on the lower part of the picture). According to O'Connell-
Rodwell (2007) the elephant's mandible contains fat pockets that are related to seismic
communication between elephants. With the courtesy of Ruth Blasco, all rights
reserved.
Fig. 3. The swollen temporal gland of an elephant, located above the eye, with a visible
secretion coming out of it. Photo by Elizabeth Stene, all rights reserved.
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According to Christy, “The best way to secure a supply,when one
has only a hunting knife,is to cut away the skin of the already
mentioned hollow over the eye.Here both meat and fat are easily
obtainable,but should be cut out soon after death.”(Christy, 1922:
292e293).
Considering the described characteristics and anatomical loca-
tion, it seems that there is a fat concentration within the eye socket
in which the eyeball of the elephant is embedded.
The trunk is also described on some occasions as an edible or-
gan, containing both fat and meat. Peter Albl (1971) explains:
“Fatty tissue is also found around the muscle cords at the base of
the trunk.”(Albl, 1971: 135).
In a book published in 1868, John George Wood describes the
elephant-hunting habits of a South-African tribe. After describing
the hunt itself, he explains the methods of carcass utilization.
Regarding the trunk he says:
“Sometimes the trunk is cut into thick slices, and baked at the same
time with the feet”(Wood, 1868: 137).
The skull of an elephant is built in the form of a “honey-comb”
(Nakamura et al., 2013). The air cells that divide the skull were also
claimed to contain edible fat (Fig. 4). Wood (1868) describes an
interesting process regarding the utilization of the skull:
“The skull itself is broken up for the sake of the oily fat which fills
the honeycomb-like cells which intervene between the plates of the
skull”(Wood, 1868: 138).
Mashchenko et al. (2013) describe a well-preserved body of a
one year old male mammoth, found in Yakutia, Russia. On the neck
of the mammoth a concentration of fat was detected. The authors
describe this concentration:
“A considerable amount of fat is deposited in the withers and side
surface of the neck
…
forming a characteristic hump.”
(Mashchenko et al., 2013).
According to the authors, this fat concentration was essential for
the survival of mammoth calves in their first year, especially during
the winter, in conditions of lackof food. An interesting evidence for
the consumption of that fat is provided:
“The soft tissues on the neck, withers, and head were eaten by
Arctic foxes when the body was thawed from the permafrost.”
(Mashchenko et al., 2013).
O'Connell-Rodwell (2007) describes a system of seismic
communication between elephants. She relates certain fat con-
centrations within the elephant's head to acoustic features that are
best known from dolphins, and says they are used along with the
fat cushions in the elephant's feet for transmission of information
between elephants. According to O'Connell-Rodwell, this “acoustic
fat”is located in the mandibular channel and inside an organ called
“the melon”.
“The dolphin melon-like acoustic fat covering the nares in the
elephant is layered with a cartilaginous structure also similar to the
melon”(O'Connell-Rodwell, 2007).
The consumption of at least some edible organs within the
elephant head, such as the trunk, and the fat in the hollow above
the eye and the skull, can be clearly attested by the ethnographic
record. The nutritional potential of the mandible is implied by the
account of O'Connell-Rodwell of fat pockets inside it, and by the
mandible bearing cut marks found at Bolomor cave. Other organs,
such as the tongue and the brain, are absent from such documen-
tation. Their nutritional potential cannot be ignored, and is implied
in the archaeological data (e.g, Goren-Inbar et al., 1994; Piperno and
Tagliacozzo, 2001; Yravedra et al., 2012; Nikolskiy and Pitulko,
2013). Thus, we suggest that early humans could have gained a
great deal by acquisition and consumption of elephants' heads.
We should also mention the possible symbolic implications that
could be attributed to the transportation and consumption of the
head of animals. Rawson (1913), for example, describes the habits
of big game hunting performed by few South African tribes. Many
social and ceremonial aspects are demonstrated. Regarding the
head of large animals it is said:
“It is also the rule that the head of any beast of great size should be
eaten on the bandla, the meeting-place of the men in the capital”
(Rawson, 1913: 57).
No explanation of this habit is provided, and no specific organs
are mentioned. However, it is quite possible that such a costume
may be attributed to the nutritional value that lies within the head.
As elephants were a significant part of these tribes nutrition, the
subject is highly relevant, and clearly, a deeper exploration of the
rites and costumes related to the consumption of big game in
general, and of big game heads in particular, is required.
5. Discussion and conclusions
The resemblance between extinct proboscideans and recent
elephants allows us to use the characteristics of modern elephants
Fig. 4. A modern elephant's skull. The air cells that divide the skull, which are claimed
to contain edible fat, can be seen in the broken upper part of the skull. Photo by Erika
Gouws, all rights reserved.
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as an indication, at least to some extent, of those of extinct pro-
boscideans. Bearing in mind that, according to current assessments
(Christiansen, 2004; Byers and Ugan, 2005), Paleolithic pro-
boscideans probably weighed more than recent elephants, organs
within the head of extant proboscideans should also be enlarged
relative to the values presented in Table 1.
Paleolithic sites with proboscidean head remains can be divided
into two distinct types: open-air sites, such as Aridos 2 (Spain),
Revadim Quarry (Israel) and Notarchirico (Italy), and cave sites,
such as Bolomor cave (Spain), Ma'anshan cave (China), and Spy cave
(Belgium). While in open-air sites the question of whether the
hominins located their sites near the carcasses or rather brought
the carcasses into the home bases is relatively difficult to resolve,
cave sites with megafauna remains are clear evidence of trans-
portation of those remains from the kill site or scavenging site to
the home base. The presence of head parts carried to those cave
sites strongly implies intentional exploitation, and possibly even
preference, of elephant heads by early humans.
Evidence of transportation of proboscideans' head parts from
acquisition localities to open air sites is also gradually accumu-
lating. In Upper Paleolithic KremseWachtberg (Austria), for
instance, it was suggested that at least three mammoths' heads
were transported into the camp site (Fladerer, 2003). The trans-
portation of mammoths' parts, including skulls, was also proposed
in the case of the Upper Paleolithic site of Yudinovo (Germonpr
e
et al., 2008).
The transportation of selected body parts into camp sites is
frequently discussed in the ethnographic and zooarchaeological
research. One main consideration regarding body parts which are
chosen to be transported is the nutritional value of each food item.
Among the Hadza of northern Tanzania, for example, high ranked
body parts are more likely to be transported from kill sites to base
camps, while low ranked body parts are less likely to be transported
(O'Connell et al., 1988, 1990). The number of elements moved from
kill-sites to base camps changes as a function of carcass size, dis-
tance and the number of carriers (O'Connell et al., 1990). According
to Monahan (1998), animals larger than size 2 (250 lb eabout
115 kg) may require more than a dozen carriers in order to be
transported to camp sites completely. Monahan also relates to the
dangers imposed by large carnivores, which may lead to field
processing (e.g., meat stripping) and resource consumption at
butchery sites (especially long bone marrow), before transportation
to camp sites. O'Connell et al. (1990) propose that “taxonomic
identity may affect the proportion of elements moved independent of
transport costs”. When examining the transportation of animals'
heads, it is often claimed that the proportion of heads that are
transported to camp sites is usually lower than that of post-cranial
elements (e.g., O'Connell et al., 1988; Schoville and Ot
arola-Castillo,
2014), as the head is considered a heavy, bulky organ, which is
uncomfortable to carry. However, in our view, that claim further
demonstrates how, in light of the significant presence of elephants'
heads in Paleolithic cave sites, the nutritional potential within the
elephant's head justifies the great effort.
The great physical effort involved in transporting such heavy
body parts to caves sites, which are sometimes located in difficult
terrain, as in the case of Bolomor cave, implies that group cooper-
ation was required. It can be suggested, then, that early humans
probably worked together, in relatively large groups, to that pur-
pose. According to Byers and Ugan (2005) moving a male African
elephant's carcass demands the cooperation of between 10 and 15
individuals, and while a modern elephant's head is “only”400 kg or
so, this organ clearly could not be moved anywhere by one or two
individuals, let alone up a steep slope.
Such cooperation between early humans implies a complex
social structure, and a composed interaction between group
members, which may include the existence of a language. The
transportation of megafauna's body parts to camp sites clearly re-
quires planning, coordination and teamwork. Thus, cooperation
and communication between group members were most likely
necessary. This notion is further supported by the case of GBY,
where elements of social interaction, and the existence of language,
were strongly suggested (Goren-Inbar, 2011).
There are a few archaeological sites with specific claims of
manipulation of proboscideans' head parts. In the Yana site in
Russia, for example, tongue bones of a mammoth were found at a
distance from the main mammoth accumulation, but within the
same cultural layer as the accumulation, indicating that the tongue
was transported there intentionally, possibly as part of the con-
sumption of mammoth meat (Nikolskiy and Pitulko, 2013). At
Bolomor cave (Spain), a mandible of a Palaeoloxodon antiquus
bearing cut marks was yielded, indicating direct evidence for hu-
man manipulation and exploitation of elephant head part that was
carried to the cave (Blasco, 2014, personal communication).
Also worth mentioning here is the unique context found at GBY,
of an almost complete elephant's skull in direct association with
stone and wood implements (Goren-Inbar et al., 1994). According
to the researchers, “It was suggested regarding the Lehringen
elephant (Adam, 1951) that certain flint artifacts found in proximity to
its skull may be taken to have been used to sever the trunk,in order to
allow access to the brain”. Using this example, they add that “The
GBY skull shows possible indications of deliberate damage below the
nasal opening,which may be attributed to the same activity.”(Goren-
Inbar et al., 1994). In our view, these two cases may also reflect the
separation and exploitation of the trunk for nutritional purposes,
not only for allowing access to the brain.
At the Acheulian site of Notarchirico (Italy) the remains of a skull
of a Paleoloxodon antiquus were found with the mandible a few
meters away from its original anatomical position. The skull was
found lying in an overturned position and lacking the masticatory
apparatus and occipital, suggesting the consumption of the soft
parts of the head ethe brain, the tongue and the trunk (Piperno
and Tagliacozzo, 2001), and possibly a separate process of the
mandible's utilization. In Upper Paleolithic KremseWachtberg
(Austria) it was proposed that a manipulation of a mammoth
mandible can be implied “by the lingual edges of the great alveoli.The
edges are symmetrically broken,and their fractures are corroded.Both
the lingual walls of the alveoli were obviously destroyed,and the
molars were probably intentionally removed”(Fladerer, 2003).
Elephant remains are found in direct association with human
activity in Paleolithic sites throughout the Old World for hundreds
of thousands of years. The elephant bones uncovered in these sites
include a significant presence of cranial remains. As many studies
have already indicated, direct and primary access of humans to
large mammals in the Paleolithic, if not actual hunting (e.g,
Fladerer, 2003; Germonpr
e et al., 2008, 2012; Demay et al., 2012;
Pickering et al., 2013; Domínguez-Rodrigo et al., 2014a)was
commonly practiced. It is our contention here thatelephants' heads
were favored by early humans as exceptionally nutritious and were
routinely exploited, and not just the leftovers available for homi-
nins following the footsteps of large carnivores.
This paper focuses on the nutritional potential of elephants'
heads. Clearly, other trajectories of exploitation of this enormous
body part can be suggested. In few Upper Paleolithic sites, for
example, dwellings made of mammoth bones were found,
including the use of skulls, tusks and mandibles (e.g, Oliva, 1988;
Iakovleva and Djindjian, 2005; Iakovleva, 2014; Vavilova and
Artemenko, 2014). Another scenario that can be considered is the
use of fat-rich skull fragments as fuel for camp fires. The use of
animal bones as fuel was suggested and demonstrated in a few
cases (e.g, Costamagno et al., 2005; Th
ery-Parisot et al., 2005;
A. Agam, R. Barkai / Quaternary International xxx (2015) 1e96
Please cite this article in press as: Agam, A., Barkai, R., Not the brain alone: The nutritional potential of elephant heads in Paleolithic sites,
Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.02.008

Beresford-Jones et al., 2010). However, a skull can be used for fuel or
for construction purposes after the exploitation of the nutritional
potential within it. In addition, such patterns of exploitation appear
in later phases of the Paleolithic, starting mainly from the Upper
Paleolithic, while the use of proboscideans' heads for nutritional
purposes has started most likely during the Early Lower Paleolithic.
Hence, it is our contention that nutrition was the main reason for
acquiring elephants' heads in Paleolithic times and that this pattern
lasted for hundreds of thousands of years in the Old World.
In our view, the nutritional potential within the elephant's skull
extends far beyond the brain. Organs such as the tongue and trunk
are a good source of protein, while fat concentrations, located in the
mandible, the temporal gland and within the skull itself, are a high-
quality source for crucial calories.
The elephant's head contains a number of edible organs, with a
high nutritional value, which are protected by a massive skull. Thus,
its full exploitation most likely required the investment of a signif-
icant period of time and energy. We therefore suggest that ele-
phants' heads were probably taken into the camp in order to
maximize their full nutritional potential within a wider time frame.
This suggestion is further supported by Zhang et al. (2010), who
claimed that “Because the head and foot parts are structurally complex,
it is difficult to obtain all the nutrition from them in a short time,so the
hunters may have chosen to take them back to camp for processing.”
Ethnographic data provide evidence for the consumption of
different head organs by recent hunteregatherers, with specific
references to the trunk, the fat within the skull's cells and the
hollow located above the eye, which may be attributed, in our view,
to the temporal gland. We should mention that we have found no
ethnographic documentation regarding the consumption of
elephant brain, tongue, or the fat pockets inside the mandible.
However, the nutritional potential of those organs is implied by
their anatomy, as well by the archaeological evidence.
The repeated evidence of transportation of elephants' head
parts to residential sites indicates it was chosen to be trans-
ported back to the camp site. Hence, it seems unlikely that the
skull was acquired and brought to camp sites solely for the
extraction and consumption of the brain, an organ of 4e6kgout
of a complex structure of more than 400 kg. Rather, we suggest
that other head parts, such as the tongue, the temporal gland,
the trunk, the mandible and the skull, were also used as a sig-
nificant part of early hominins' diet. The repeated presence of
different elephant cranial remains (skulls, skull fragments,
mandibles, tongue bones, teeth etc.) in Paleolithic sites supports
this hypothesis. Isolated teeth and fractured mandibles, for
instance, may be an indication of the smashing of the mandible
in order to reach and consume the mandibular fat (as can be
implied by the above mentioned example from KremseWa ch t-
berg: Fladerer, 2003).
The archaeological data gathered from Paleolithic sites imply, in
our opinion, that early humans were highly capable and practiced
successful adaptations in difficult conditions for many hundreds of
thousands of years. Early humans were able to understand and
maximize the full potential that existed in any nutritional source,
elephant carcasses, and elephants' heads specifically, included. We
believe that the data presented in this paper provide a better un-
derstanding of early human behavior reflected by the presence of
elephant head remains in Paleolithic sites, and add another
perspective on the long lasting interactions between humans and
proboscideans during Paleolithic times.
Acknowledgements
This paper was presented in the VIth International Conference on
Mammoths and their Relatives held at Grevena and Siatista, Greece.
We would like to thank Lucy Wilson for reading an early version of
this article and contributing her useful remarks and insights. We
would like to thank Nir Geiger, Ruth Blasco, Elizabeth Stene and
Erika Gouws for permitting us the use of their wonderful photos in
this paper. All rights of these photos are reserved.
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