Rock painting of figure smoking a beehive, Toghwana Dam, Zimbabwe. Reprinted from Crane 1999, with permission from International Bee Research Association (IRBA). 

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... coupled with the spread of grasslands 2 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 habitat (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 larvae in early hominin diet is explored. Nutritional composition, ethnographic reports, examples of Paleolithic rock art, and evidence from non-human primates 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. 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 (Bogdanov 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 protein 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 sources of energy, fat, and protein and represent high-quality food sources that have been targeted for much of human history. 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 Southern 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 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; Hernández-Pacheco 1928). Multiple 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 (Gordon 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 smoking 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, however, 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 composition 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. Honey and bee larvae consumption is widespread among human populations. A large body of literature reports the importance of honey in the diets of human foragers (Bodenheimer 1951); however, data on the amount collected and consumed remain largely anecdotal. Abundant consumption of honey and brood (larvae), both for stinging ( Apis mellifera ) and stingless 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 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 available, 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 ...