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The Importance of Honey Consumption in Human Evolution

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It has been suggested that honey may have been an important food source for early members of the genus Homo, yet the importance of meat and savanna plant foods continue to be stressed as the most relevant foods in dietary reconstructions. Here, the importance of honey and bee larvae in hominin diets is explored. Ethnographic reports, examples of Paleolithic rock art, and evidence from non-human primates are used to show that early hominins likely targeted beehives using the Oldowan tool kit. The consumption of honey and bee larvae likely provided significant amounts of energy, supplementing meat and plant foods. The ability to find and exploit beehives using stone tools may have been an innovation that allowed early Homo to nutritionally out-compete other species and may have provided critical energy to fuel the enlarging hominin brain.
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The Importance of Honey Consumption
in Human Evolution
Alyssa N. Crittenden
a
a
Department of Anthropology, University of Nevada, Las Vegas,
Nevada, USA
Available online: 08 Dec 2011
To cite this article: Alyssa N. Crittenden (2011): The Importance of Honey Consumption in Human
Evolution, Food and Foodways, 19:4, 257-273
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DOI: 10.1080/07409710.2011.630618
The Importance of Honey Consumption
in Human Evolution
ALYSSA N. CRITTENDEN
Department of Anthropology, University of Nevada, Las Vegas, Nevada, USA
It has been suggested that honey may have been an important food
source for early members of the genus Homo, yet the importance
of meat and savanna plant foods continue to be stressed as the
most relevant foods in dietary reconstructions. Here, the impor-
tance of honey and bee larvae in hominin diets is explored. Ethno-
graphic reports, examples of Paleolithic rock art, and evidence from
non-human primates are used to show that early hominins likely
targeted beehives using the Oldowan tool kit. The consumption of
honey and bee larvae likely provided significant amounts of en-
ergy, supplementing meat and plant foods. The ability to find and
exploit beehives using stone tools may have been an innovation that
allowed early Homo to nutritionally out-compete other species and
may have provided critical energy to fuel the enlarging hominin
brain.
Honey is one of the most energy-dense foods in nature (Skinner 1991), but
despite suggestions that it may have been an important food source for early
Homo (Allsop and Miller 1996; Bunn and Schoeninger 2009; McGrew 2001;
Schoeninger et al. 2003; Skinner 2001), it has received little attention in most
reconstructions of early hominin
1
diet composition. While the importance of
meat (Bunn 2007; Cordain et al. 2001; Eaton et al. 1998; Stanford and Bunn
2001) and savanna plant foods, e.g., underground storage organs (USOs)
(Dominy et al. 2008; Hawkes et al. 1997; Wrangham et al. 1999), are routinely
highlighted, the importance of honey and larvae consumption have been
largely overlooked.
The appearance of Oldowan tool technology, the earliest assemblages
of deliberately flaked stone artifacts, coupled with the spread of grasslands
2
Address correspondence to Alyssa N. Crittenden, Department of Anthropology, Univer-
sity of Nevada, Las Vegas, 4505 S. Maryland Parkway, Las Vegas, NV 89154-5003. E-mail:
alyssa.crittenden@unlv.edu
257
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258 A. N. Crittenden
across East Africa, suggest that ecological changes during the Late Pliocene
may have led to changes in resource availability for early Homo (Ungar
et al. 2006). An expanded toolkit would have allowed early hominins to
target, process, and consume a greater range of foods than was previously
available (Dominguez-Rodrigo et al. 2001; Shea 2007; Ungar et al. 2006). Sta-
ble carbon isotope data of hominin tooth enamel and faunal reconstructions
suggest that early hominins fed in an open country savanna woodland habi-
tat (Sponheimer and Lee-Thorp 1999; Sponheimer et al. 2005). Although the
general consensus agrees that environmental changes along with technolog-
ical innovation provided an opportunity for major dietary shifts, the types of
foods that would have been targeted remain unclear (Leonard et al. 2007;
Ungar et al. 2006). Here, the probable importance of honey and bee lar-
vae in early hominin diet is explored. Nutritional composition, ethnographic
reports, examples of Paleolithic rock art, and evidence from non-human pri-
mates are used to show that early Homo was capable of, and likely benefited
from, targeting beehives using the Oldowan tool kit. Liquid honey may have
provided much-needed energy to early hominin foragers and would have
been an important supplement to meat and plant collection.
SEASONAL PRODUCTION AND NUTRITIONAL COMPOSITION
OF HONEY AND LARVAE
The seasonality of honey production differs in temperate versus tropical
climates. In temperate zones, the production cycle of both stinging (Apis
mellifera) and stingless (Meliponinae) honeybees is highly seasonal (Crane
1990). In tropical zones, honeybees are dependent upon rainfall, regardless
of the season; during relatively wet years, production of honey may be
continual, whereas in dry years, production may be restricted to short periods
following a rain (Crane 1990, 1999).
Liquid honey, both wild and domestic, contains approximately 80–95
percent sugar and is a concentrated source of fructose and glucose (Bog-
danov et al. 2008; Murray et al. 2001; White et al. 1962). Honey also contains
trace amounts of several essential vitamins and minerals (Iskander et al.
1995; Terrab et al. 2004) and contains components that act as preservatives,
including a-tocopherol, ascorbic acid, flavonoids, glucose oxidase, catalase,
and peroxidase (Crane 1975; Ferreres et al. 1993; Nagai et al. 2006). Although
only small amounts of protein (mainly free amino acids) are found in liquid
honey (Bogdanov et al. 2008), wild honeys contain higher levels of pro-
tein and fat, most likely because they contain trace amounts of bee larvae,
whereas cleaned and commercially processed honey does not (Murray et al.
2001). Bee larvae is a good source of protein, fat, several essential minerals,
and B-vitamins (Finke 2005). Combined, honey and bee larvae are excellent
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Importance of Honey Consumption 259
sources of energy, fat, and protein and represent high-quality food sources
that have been targeted for much of human history.
THE ROCK ART OF HONEY COLLECTION
Multiple examples of Upper Paleolithic (40,000–8,000 years ago) rock art
linked to honey and bees are found in Spain, India, Australia, and South-
ern Africa. The most abundant rock art representations of honey related
activities in Europe are found in Spain. Paintings in a side chamber of the Al-
tamira cave (Figure 1), dated to roughly 25,000 years ago (Pager 1976), depict
FIGURE 1 Mesolithic rock painting of honey collection from a wild bee nest, La Ara
˜
na shelter,
Valencia, Spain. Reprinted from Crane 1999, with permission from International Bee Research
Association (IRBA).
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260 A. N. Crittenden
honeycomb, bees, and honey collection ladders (Cartailhac and Breuil 1906).
In an open-air rock shelter in Bicorp in the Valencia Province, depictions of
honey collection, bee swarms, and comb representations, dating to 10,000
years ago, have been found (Dams 1978; Hern
´
andez-Pacheco 1928). Mul-
tiple representations of honey collection have been found in central India.
Images include groups of both men and women sitting in trees containing
beehives, smoking hives, and climbing ladders to access honeycombs (Gor-
don 1960; Mathpal 1984). There are also drawings of basketlike objects that
may represent dried-out gourds for collecting liquid honey (Mathpal 1984).
In Australia, rock art depicting the nests of stingless bees have been found
near Darwin (Crane 1975) and near the Laura River (Trezise 1973). Figures
made with beeswax, dated to 4,000 years ago, have been found on the walls
of open rock shelters in the Northern Territory (Nelson 1995).
The majority of rock paintings depicting honey collection and bee-
related topics are located in Africa (Crane 1986, 1999). In South and West
Africa (Johnson et al. 1959; Rudner and Rudner 1970), Zambia (Clark 1942),
Namibia (Crane 1983), and Lesotho (Smits 1971), there are depictions of bee
swarms, hives, and people on ladders accessing honeycomb, represented
as ovoid shapes with black and white coloration (Pager 1973; Woodhouse
1989). The dark shaded areas represent comb filled with honey or pupae,
and the light areas represent empty comb decorated with dots representing
visible larvae in uncapped cells (Crane 1983). A depiction of a figure smok-
ing a beehive (date unknown) was found in Toghwana Dam, Zimbabwe
(Figure 2). The San foragers claim that their people have used this cave for
approximately 10,000 years (Crane 1999).
The abundance of rock art depicting honey collection scenes at several
sites throughout the world suggests that honey and larvae may have been an
important part of the Paleolithic diet and daily life. It can be expected, how-
ever, that early hominins were exploiting beehives long before the practice
was represented artistically; beehives and the stick tools used to access their
contents do not survive the archaeological record. Data on the diet compo-
sition and food collection practices of foraging populations offer compelling
evidence that honey and bee larvae may have contributed to a significant
portion of the early Homo diet.
ETHNOGRAPHY OF HONEY CONSUMPTION
Honey and bee larvae consumption is widespread among human popula-
tions. A large body of literature reports the importance of honey in the diets
of human foragers (Bodenheimer 1951); however, data on the amount col-
lected and consumed remain largely anecdotal. Abundant consumption of
honey and brood (larvae), both for stinging (Apis mellifera) and stingless
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Importance of Honey Consumption 261
FIGURE 2 Rock painting of figure smoking a beehive, Toghwana Dam, Zimbabwe. Reprinted
from Crane 1999, with permission from International Bee Research Association (IRBA).
bees (Meliponinae), is reported for foraging groups in Latin America, Asia,
Australia, and Africa.
Across Latin America, multiple foraging groups focus a considerable
amount of their collection effort on honey and bee larvae during distinct
periods throughout the year (Ramos-Elorduy et al. 1997). The Yukpa-Yuko of
Venezuela and Colombia consider bees the “single most important group of
insects” in their diet (Ruddle 1973), and the Hiwi of Venezuela also consume
significant amounts of both liquid honey and bee larvae during both the
wet and the dry seasons (Gurven et al. 2000; Hill 2002; Hurtado and Hill
1987, 1990). The Ache of Paraguay consider honey and bee larvae to be
the second most important resource class in their diet after large game meat
(Hill et al. 1984; Kaplan and Hill 1985). The detailed data on the Ache
quantifies the ways in which bees and their products contribute to overall
diet composition. The energetic value of Ache honey is reported to be 3,232
calories per kilogram; consumption of honey and brood is, on average, 1,163
calories per person per day (Hurtado et al. 1985). Although honey is most
abundant during the early part of the wet season, it is available in varying
amounts throughout the year (Hill 2002; Hill et al. 1984; Hawkes et al. 1982).
In Asia, the honey foragers of Nepal target the nests of large stingless
and stinging honeybees and collect approximately 40 liters of liquid honey
and 10 kilograms of comb in less than one hour of foraging (Valli and
Summers 1988). They utilize bamboo ladders to climb within reach of the
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262 A. N. Crittenden
nests, which are usually situated high in cliff crevices or in trees on the cliff
face (Underwood 1990). The Tamang of Nepal (Masvie 2006; Thapa 2000)
and the Onge of the Andaman Islands (Cipriani 1966; Crane 1975) also collect
and consume large quantities of honey and bee larvae.
Various foraging peoples of Australia have historically utilized the hives
of both honeybees and stingless bees, eating large amounts of honey and
brood (Akerman 1979; O’Dea et al. 1991). Both the men and women of the
Worora, Wunambal, and Ngarinjin tribes from west Australia are reported
to have used digging sticks, hammer stones, and stone hatchets to gain
access to beehives (Akerman 1979). The Wanindiljangwa of the north coast
of Australia highly value the “sugar bag” (Worsley 1961), which contains
honey, brood, and wax comb. The sugar bag is considered to be one of the
most valued foods in their dietary repertoire and produces 1,680 calories per
kilogram (Meehan 1977).
Reports on African foragers provide the most detailed accounts of honey
collection and consumption. The ecology of the African environment, pri-
marily miombo woodland and baobab trees, have bark that is highly suitable
for hive construction and provide reliable honey throughout the year (Guy
1972). The various honey collection techniques across Africa include using
climbing ropes to gain access to hives located high in trees, hammering pegs
into the trunk of a tree to climb to the hive’s location, or using smoke to
stun the bees, which then fly out and abandon the hive and its contents
(Bodenheimer 1951; Guy 1972; Crane 1999). Anecdotal references to honey
collection and consumption are reported for various groups, including the
Mikea of Madagascar (Tucker 2004), the Tongwe of Tanzania (Takeda 1976),
and the Ogiek of Kenya (Nightingale 1983). There is quantified data avail-
able, however, on the amounts collected and consumed by the Efe foragers
of the Ituri Forest in the Democratic Republic of the Congo and the Hadza
foragers of Tanzania.
The Efe have a “honey season” that lasts from July–August (Terashima
1998). During this season, they move deep into the forest in search of the
liquid honey and larvae of both stinging and stingless bees. During the honey
season, they rely almost entirely on honey, brood, and pollen (Ichikawa
1981; Turnbull 1976). Men and women collect honey together in family
groups and share their yield once they have returned to camp. The average
amount of honey and brood collected by the average person per day is
3.32 kilograms, and the average amount consumed per person per day is
0.62 kilograms of honey (dry weight), which is calculated as 1,900 calories
per day (Ichikawa 1981). Honey contributes roughly 70 percent of the diet by
weight and 80 percent by calories (Ichikawa 1981; Terashima 1998), making
it the largest component of the Efe diet during the wet season.
Among the Hadza foragers of Tanzania, honey is the most prized and
highly ranked food source (Berbesque and Marlowe 2009). The Hadza collect
the honey of stinging and stingless bees, consuming the larvae of both.
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Importance of Honey Consumption 263
Honey comprises approximately 15 percent of the Hadza diet (Marlowe
2001), is shared widely outside of the household, and is consumed by the
entire camp (Bunn and Schoeninger 2009; Marlowe 2003). Liquid honey is
a primary weaning food
3
for Hadza children (Crittenden et al. n.d.). Male
foragers are the primary honey collectors; however, children also routinely
target beehives and return to camp with honey to share (Crittenden et al.
n.d.). Young male foragers are able to collect and consume upwards of
3,000 kilocalories of honey during a three-hour foraging trip (Crittenden
2009). Honey is typically housed high off the ground in large baobab trees.
Hadza men drive posts into the trunk of the tree, which the men and boys
then climb in order to access the hive (Figure 3).
When locating an active hive, Hadza foragers often employ the as-
sistance of the honey guide bird, Indicator indicator, a paleotropical
near passerine bird that occupies sub-Saharan Africa (Short and Horne
2002). The honey guide is also an accomplice in assisting honey badgers
FIGURE 3 Hadza honey collection in large baobab tree.
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264 A. N. Crittenden
(Mellivora capensis) and baboons (Papio cynocephalus) locate beehives
(Dean and MacDonald 1981; Zimmerman et al. 1999). In the case of the
Hadza, the honey guide bird leads the honey forager to the hive. They stay
in constant communication via a series of chatters and whistles. When the
forager reaches the hive, he chops into the tree to create a large hole and then
typically smokes the bees. This involves placing burning brush in the mouth
of the hive, which acts to reduce the electroantennograph response of the
guard bees, who otherwise would release a volatile alarm odor pheromone
called iso-pentyl acetate when threatened (Boch and Shearer 1962; Visscher
et al. 1995). When the smoke enters the hive, the antennae receptors of
the guard bees are dulled and they fail to sound the alarm. Smoking the
hive also has the secondary effect of causing the other bees in the hive to
gorge themselves on honey as a response to habitat threat. They take in vast
amounts of honey and then vacate the hive in order to rebuild it elsewhere.
When exposed to smoke, bees are dramatically less defensive and aggres-
sive, and there is a negative correlation between degree of engorgement
and the tendency to sting (Biamonte 1974; Conrad 1940; Free 1968; Viss-
cher et al. 1995). The Hadza forager thus escapes getting stung by the bees,
aptly named African killer bees, and extracts the contents of the hive (Figure
4). While he focuses on the liquid honey and larvae, the honey guide bird
consumes the wax.
FIGURE 4 Hadza honey hunter with honey and bee larvae (Apis mellifera adansonii).
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Importance of Honey Consumption 265
The above reports of honey and brood consumption represent only
a subset of the populations who focus foraging efforts on the content of
beehives. The ethnographic data outlined above are the most detailed and
consistently reported examples representing the importance that honey plays
in the diet of foraging populations.
HONEY COLLECTION BY NON-HUMAN PRIMATES
Several species of non-human primates are also successful honey collectors.
Baboons and macaques routinely target the nests of both stinging and stin-
gless bees (Botha 1970). In Bwindi Impenetrable National Park, baboons
target the nests of stingless bees and harvest honey and larvae using their
hands and muzzles (Kajobe and Roubik 2006). Gorillas also target honey in
Bwindi (Kajobe and Roubik 2006; Stanford et al. 2000) and elsewhere (Botha
1970). Orangutans routinely collect honey and bees by using probes made
of vegetation (Fox et al. 1998) or by smashing the hive (Rijksen 1978). Chim-
panzees at many sites target honey and larvae using a variety of methods,
including the use of stick tools to extract the contents of the hive (Cen-
tral African Republic: Fay and Carroll 1994; Hicks et al. 2005; C
ˆ
ote d’Ivoire:
Boesch and Boesch 1990; Gabon: Boesch et al. 2009; Tutin and Fernan-
dez 2005; Gambia: Brewer and McGrew 1990; Nigeria: Fowler and Sommer
2007; Republic of Congo: Bermejo and Illera 1999; Sanz and Morgan 2009;
Yamagiwa et al. 1988; Senegal: Bermejo et al. 1989; Tanzania: Goodall 1986;
Uganda: Stanford et al. 2000; Kajobe and Roubik 2006; Watts 2008). Studies
of chimpanzee tool use and diet composition are useful in models of early
hominin behavior, not only because of their phylogenetic proximity to hu-
mans, but also because of their behavioral and anatomical similarities to our
species. These data from non-human primates suggest that honey and larvae
consumption may have been an important part of the subsistence strategy
of early hominins.
DISCUSSION AND CONCLUSIONS—HONEY IN
HUMAN EVOLUTION
Using the convergent evidence of honey and larvae consumption from non-
human primates, historical and contemporary foraging populations, and artis-
tic representations from Upper Paleolithic rock art, it can be expected that
early hominins likely targeted beehives using the Oldowan tool kit.
It is possible that honey extraction was practiced by the Australo-
pithecines, yet digging sticks do not survive the archaeological record.
Oldowan stone tools begin to appear in the archaeological record approx-
imately 2.5 mya in Africa (Semaw 2006; Toth and Schick 2009) and have
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266 A. N. Crittenden
also been found at multiple sites in Europe and Asia (Toth and Schick 2009).
Using the Oldowan toolkit to split open beehives would have made honey
and larvae more easily accessible and possibly allowed hominin foragers to
extract more honey than was previously possible. Hollowed-out gourds and
ostrich eggs, frequently used by foragers to transport water (Howell 2010),
could have been utilized to transport honey and larvae back to camp to
share with dependent offspring and individuals unable to forage.
4
Routine
honey consumption would have provided considerable amounts of energy,
successfully supplementing meat and plant foods.
It has been suggested that the evolution of larger hominin brains, w hich
are metabolically expensive (Aschoff et al. 1971; Holliday 1986), would have
required the consumption of energy-rich foods to fuel the expansion (Aiello
and Wheeler 1995; Leonard and Robertson 1994). Early members of the genus
Homo, 1.5–2 million years ago, were the first to show signs of increased
relative brain size (McHenry and Coffing 2000). This expansion coincides not
only with the appearance of Oldowan tools, but also with the reduction in
size of molar dentition, indicating that hominins were consuming foods that
required less mechanical breakdown, i.e., chewing (Brace et al. 1987, 1991;
Leonard et al. 2007). Most Paleolithic diet reconstructions emphasize the vital
role that meat (Bunn 2007; Cordain et al. 2001; Eaton et al. 1998; Stanford
and Bunn 2001) and/or tubers (Dominy et al. 2008; Hawkes et al. 1997;
Wrangham et al. 1999) played in the shifting diet of early Homo. Honey,
however, a food that is easy to consume and digest and is energetically
dense, remains largely ignored as a component of hominin diet.
The enlarging hominin brain would have greatly benefited from the
energy provided by even a modest amount of honey (Skinner 1991). Glucose
plays a critical role in meeting the high metabolic requirements of neural
development and function (Amiel 1994; Chugani 1998). In addition to energy,
honeycomb provides small amounts of protein in the larval cells. In a highly
seasonal environment such as Late Pliocene East Africa (Copeland 2007),
honey and larvae may have supplemented scarce resources during the dry
season. The ability to find and exploit beehives with stone tools may have
been an innovation that allowed early hominins to nutritionally outcompete
other species and may have been a crucial energy source to help fuel the
enlarging hominin brain. To further our understanding of early hominin diet
composition and the potential links between nutrition and neural expansion,
we must begin to incorporate honey and larvae consumption into models of
early Homo diet reconstruction.
ACKNOWLEDGMENTS
I would like to thank the anonymous reviewers for their very useful com-
ments. I am grateful to Dr. Margaret Schoeninger, Dr. Adrienne Zihlman, and
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Importance of Honey Consumption 267
Dr. Frank Marlowe for very stimulating discussions on the role of honey in
human evolution.
NOTES
1. The term “hominin” is a taxonomic classification used here to refer to members of the genus
Homo and their direct ancestors, including Australopithecines.
2. Grasslands using a C
4
photosynthetic pathway (versus the more common C
3
pathway for most
flowering p lants) began radically changing the biosphere around 3–8 mya (Cerling 1992; Edwards et al.
2010).
3. Although Western populations are cautioned against feeding honey to infants due to the dangers
of botulism (Smith et al. 2010), some rural populations routinely use honey as a weaning food (Kumar
et al. 2006; Madhu et al. 2009).
4. The earliest appearance of use of a home base, or central place, dates to the Plio/Pleistocene
boundary (O’Connell 1997). Early Homo foragers were nomadic and, as such, most likely did not store
food for extended periods of time (Marlowe 2006). The preservative properties of honey, however, may
have made it easier to store. It has been suggested that early hominins may have stored some foods for
short periods of time as a way to combat scarcity of seasonal resources (McBrearty and Brooks 2000).
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