The wild yam question: How well could independent hunter-gatherers live in a tropical rain forest ecosystem?

It has been generally assumed until recently that tropical rain forests are food-rich biomes for human foragers, and that prehistoric hunter-gatherers once lived completely independent of cultivated foods in such environments. An alternative hypothesis that such forests are actually food-poor for humans is proposed here. Specifically, that wild starch foods such as yams were so scarce and so hard to extract that human foragers could not have lived in such biomes without recourse to cultivated foods. The symbiotic relationship found today between tropical forest hunter-gatherers and farmers is not a recent phenomenon, but evolved long ago as an adaptive strategy for successfully exploiting the tropical forest.     

Diet-Related Buccal Dental Microwear Patterns in Central African Pygmy Foragers and Bantu-Speaking Farmer and Pastoralist Populations

Pygmy hunter-gatherers from Central Africa have shared a network of socioeconomic interactions with non-Pygmy Bantu speakers since agropastoral lifestyle spread across sub-Saharan Africa. Ethnographic studies have reported that their diets differ in consumption of both animal proteins and starch grains. Hunted meat and gathered plant foods, especially underground storage organs (USOs), are dietary staples for pygmies. However, scarce information exists about forager–farmer interaction and the agricultural products used by pygmies. Since the effects of dietary preferences on teeth in modern and past pygmies remain unknown, we explored dietary history through quantitative analysis of buccal microwear on cheek teeth in well-documented Baka pygmies. We then determined if microwear patterns differ among other Pygmy groups (Aka, Mbuti, and Babongo) and between Bantu-speaking farmer and pastoralist populations from past centuries. The buccal dental microwear patterns of Pygmy hunter-gatherers and non-Pygmy Bantu pastoralists show lower scratch densities, indicative of diets more intensively based on nonabrasive foodstuffs, compared with Bantu farmers, who consume larger amounts of grit from stoneground foods. The Baka pygmies showed microwear patterns similar to those of ancient Aka and Mbuti, suggesting that the mechanical properties of their preferred diets have not significantly changed through time. In contrast, Babongo pygmies showed scratch densities and lengths similar to those of the farmers, consistent with sociocultural contacts and genetic factors. Our findings support that buccal microwear patterns predict dietary habits independent of ecological conditions and reflect the abrasive properties of preferred or fallback foods such as USOs, which may have contributed to the dietary specializations of ancient human populations.     

From subsistence to market: A case study of the mbuti net hunters

This article describes the recent transition from subsistence to market hunting of some net-hunting Mbuti, a nomadic society of the Ituri Forest of Zaire. The history of this development dates to the late 1950s, as the Mbuti began to have increasing contact with commercial meat traders, called bachuuzi, from outside markets. Before this, the Mbuti had a long history of contact with local swidden agriculturalists, called bakbala, a relationship that continues today. It is sanctioned by religious beliefs, but material exchanges are also important. The Mbuti provide bakbala with meat and other forest resources and receive in return iron implements, tobacco, and cultivated food. The Mbuti view such items now as necessities; however, this does not allow the bakbala to subjugate or control them. Most exchanges take place in the bakbala's village at the Mbuti's discretion, and exchange rates are not fixed. Finally it appears that such trade does not undermine the resource base of either group. The Mbuti's commercial exchanges contrast with this subsistence-oriented system. Unlike the bakbala, the bachuuzi traders establish themselves in the Mbuti's forest camps, where they can promote intensified net hunting and monopolize trade of meat. The Mbuti tolerate the alien traders in camp because they are often a convenient source of food and other desired material goods; however, they do not share the traders' commercial motives. The Mbuti freely manipulate credit to their own advantage and they are often able to evade the traders' efforts at economic control. The future of market hunting is uncertain. Mbuti material needs are changing and the antelope fauna (Cephalophinae) upon which commercial trade depends is being reduced in some areas.A fellowship from the Thomas J. Watson Foundation provided financial support for this research. I am also thankful to Citoyen Bokanga Ekanga Botombele, Commissaire d'Etat du Zaire for his interest in my work and support in obtaining necessary visas, and to Père Thyert of the Catholic Mission of Biambwe for his generous hospitality. Terese Butler Hart, Colin Turnbull, Paul Riesman, Randall Packard, Paul Wholt, and Stuart Marks criticized drafts of this paper. Finally, I am most grateful to the Mbuti with whom I lived for so generously sharing their daily life with me.     

The possibility of independent foraging in the rain forest of Peninsular Malaysia

This paper examines the question of whether hunter-gatherers could live in the tropical rain forest of Peninsular Malaysia without access to cultivated foods. It considers the wild food sources used by the Batek De', a contemporary foraging-trading group of Kelantan state, historical and ethnohistorical evidence concerning the Batek economy in the past, and archeological evidence for independent foraging in the Pleistocene and early Holocene. The conclusion reached is that small nomadic groups of foragers can live off wild resources alone in that environment and have done so in the past, although trade for agricultural produce makes life much easier. This suggests that nonseasonal tropical rain forests vary in their potential for supporting human foraging, depending upon the particular species of plants and animals present.     

Shallow-water habitats as sources of fallback foods for hominins

Underground storage organs (USOs) have been proposed as critical fallback foods for early hominins in savanna, but there has been little discussion as to which habitats would have been important sources of USOs. USOs consumed by hominins could have included both underwater and underground storage organs, i.e., from both aquatic and terrestrial habitats. Shallow aquatic habitats tend to offer high plant growth rates, high USO densities, and relatively continuous USO availability throughout the year. Baboons in the Okavango delta use aquatic USOs as a fallback food, and aquatic or semiaquatic USOs support high-density human populations in various parts of the world. As expected given fossilization requisites, the African early- to mid-Pleistocene shows an association of Homo and Paranthropus fossils with shallow-water and flooded habitats where high densities of plant-bearing USOs are likely to have occurred. Given that early hominins in the tropics lived in relatively dry habitats, while others occupied temperate latitudes, ripe, fleshy fruits of the type preferred by African apes would not normally have been available year round. We therefore suggest that water-associated USOs were likely to have been key fallback foods, and that dry-season access to aquatic habitats would have been an important predictor of hominin home range quality. This study differs from traditional savanna chimpanzee models of hominin origins by proposing that access to aquatic habitats was a necessary condition for adaptation to savanna habitats. It also raises the possibility that harvesting efficiency in shallow water promoted adaptations for habitual bipedality in early hominins. Am J Phys Anthropol 140:630–642, 2009. © 2009 Wiley-Liss, Inc.     

Fruit Traits in Baboon Diet: A Comparison with Plant Species Characteristics in West Africa

Primate fruit choice among plant species has been attributed to different morphological plant and fruit characteristics. Despite a high abundance of animal-dispersed plant species in the savanna–forest mosaic of West Africa, few data are available on the interplay between morphological fruit traits and primate fruit consumers in this ecosystem. We tested whether olive baboons (Papio anubis) at Comoé National Park, north-eastern Ivory Coast, prefer fruit species with particular characteristics relative to the availability of these traits among the woody plant species at the study site. Specifically we were interested in the suites of traits that best predict fruit choice and seed handling by baboons. The baboons ate fruit/seeds from 74 identified plant species, representing 25 percent of the regional pool of woody plant species. They preferred trees to shrubs and lianas as fruit sources. Otherwise, baboons seemed to consume whatever fruit type, color, and size of fruit and seeds available, though they especially included larger fruit into their diet. Against expectations from the African bird–monkey fruit syndrome of brightly colored drupes and berries, baboons ate mostly species having large, dull-colored fruit. Fruit type and color best described whether baboons included a species into their diet, whereas fruit type and seed size best predicted whether baboons predated upon the seeds of their food plant species. As most plant species at the study site had medium-sized to large fruits and seeds, large frugivores like baboons might be particularly important for plant fitness and plant community dynamics in West African savanna–forest ecosystems.     

Selectivity and Variability in the Diet of Milne-Edwards' Sifakas (Propithecus diadema edwardsi): Implications for Folivory and Seed-Eating

Propithecus diadema edwardsi combines a strategy of high selectivity for many food items with the use of more readily available and abundant foods. Selectivity is seen in relation to taxonomic affiliations of plants eaten, forest composition, and phenological patterns. Feeding time is concentrated on relatively uncommon species as indicated by high selection ratios. Annual and monthly dietary diversity is low, with food choices restricted to a relatively narrow set of species: a total of 82 and 73 plant species were eaten by the two groups of which, only 22 and 16% contributed >1% feeding time, however, these values fall with commonly reported ranges. Plant parts eaten varied widely from month to month, with overall variance in seed-eating higher than variance in young or mature leaf feeding. No correlation between percentage feeding time and food availability is significant; instead, monthly feeding patterns are better explained by the availability of particular plant species, which also supports a view of high selectivity in food choice. In contrast to selectivity for many seed, whole fruit, and new leaf sources, reliably abundant food sources appear to be leaves from lianas, epiphytes, and parasites, which were eaten in all focal months. While dramatic interannual differences in dietary composition in response to the absence of a preferred seed source highlight the interpretation of folivores as surviving food crunches by turning to readily available, poor quality resources, this strategy is not unique to folivores.     

Wild yams revisited: Is independence from agriculture possible for rain forest hunter-gatherers?

The hypothesis that energy-rich wild plant foods are too scarce in rain forest to allow subsistence by foraging peoples independently of agriculture lacks a firm empirical basis. Data on availability of wild plant foods such as wild yams are sorely lacking, and where quantitative information is provided to support the hypothesis, it usually concerns extent of use of wild plant foods: low availability is tacitly inferred from low use. We explore the alternative hypothesis that extent of use underestimates availability of wild yams and other wild plant foods; these foods are present in large enough quantities to support hunter-gatherers, but have become increasingly neglected with increasing availability of cultivated plant foods. Thus, the subsistence of contemporary rain forest foraging peoples, in which extensive relationships with sedentary farmers appear to be universal, may be a somewhat distorted reflection of their subsistence in the pre-agricultural past. Drawing on data from ecology, archeology, ethnohistory, and linguistics, we argue that pygmy foraging peoples of the western Congo basin were present in rain forest environments before the advent of farming villagers.     

Ranging behaviour and habitat preference of a wild marmoset group, Callithrix humeralifer (Callitrichidae, Primates)

The ranging behaviour of a group of marmosets ( Callithrix humeralifer ) in seasonal Amazonian rain forest was studied during one year. Range sizes (monthly and daily), day range lengths and patterns of range use are examined for correlations with feeding behaviour and the distribution of three forest types within the marmoset's range. Seasonal differences in ranging are associated with changes in the abundance and distribution of plant food sources. The marmosets ranged more widely and used more sources of a greater diversity of plant food species, which were distributed over a wider area, in the wet season than in the dry season. In the dry season, they ranged over a smaller area and, although they used fewer sources of a reduced diversity of plant species overall, they exploited a larger number of sources of the five highest ranked plant species in the diet. Throughout the year, they showed a preference for disturbed primary forest, characterized by dense understoreys and abundant second growth patches. Reasons for this preference are discussed, taking into account their use of fruits of typical pioneer species (particularly in the dry season), insect prey abundance, sleeping site availability and defence against predators.     

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems

Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (K _leaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday K _leaf values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of K _leaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.     

Potential hominin plant foods in northern Tanzania: semi-arid savannas versus savanna chimpanzee sites

Savanna chimpanzees are useful as referential models for early hominins, and here potential differences between chimpanzee and early hominin ecology is the focus. Whereas chimpanzees inhabit only a handful of modern African savannas, there is evidence that early hominins occupied relatively more open and arid savannas than those in which chimpanzees live. In order to help expand potential models of early hominin palaeoecology beyond savanna chimpanzee-like scenarios, and to provide a basis for future modeling and testing of actual hominin diets, this study compares the types of plant foods available in modern semi-arid savannas of northern Tanzania to plant foods at savanna chimpanzee sites. The semi-arid savannas are not occupied by modern chimpanzees, but are potentially similar to environments occupied by some early hominins. Compared to savanna chimpanzee habitats, the northern Tanzania semi-arid savanna has a lower density and fewer species of trees that produce fleshy fruits. Additionally, the most abundant potential hominin plant foods are seasonally available Acacia seeds/pods and flowers, grass seeds, and the underground parts of marsh plants, as evidenced by vegetation surveys and by studies of the diets of baboons that forage in similar areas. The information from this study should be useful for framing hypotheses about hominin diets for sites with palaeoenvironmental contexts similar to those of the northern Tanzania semi-arid savannas and for contextualising tests of actual hominin diets (e.g., those based on dental microwear or isotopes).     

Edible wild plants of the Chorote Indians, Gran Chaco, Argentina

The Chorote Indians are hunter-gatherers and fishermen from north-west Argentina and south-west Paraguay who belong to the Mataco-Maká linguistic family. Their edible plants are identified by botanical and vernacular names, the parts employed and modes of preparation and consumption. The Chorote people use 57 plant species as a source of food, which they consume in 118 different ways. Five new edible species, that yield seven plant foods, are reported here for the first time for Chaquenian ethnic groups. However, only a few wild plant foods are in frequent use today, with most being used occasionally, infrequently or not at all. A cross-cultural comparison with four neighbouring ethnic groups reveals that one third of their plant foods are exclusive to the Chorote people, despite the fact that they share most of their edible plants with the other groups. This article is the first contribution to an understanding of the Chorote's ethnobotany and to a comparison of interethnic relationships concerning their edible plant resources.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 73–85.     

Traditional diet and food preferences of Australian aboriginal hunter-gatherers.

Until European settlement of Australia 200 years ago, Aborigines lived as nomadic hunter-gatherers all over the continent under widely varying geographic and climatic conditions. Successful survival depended on a comprehensive knowledge of the flora and fauna of their territory. Available data suggest that they were physically fit and lean, and consumed a varied diet in which animal foods were a major component. Despite this, the diet was not high in fat, as wild animal carcasses have very low fat contents through most of the year, and the meat is extremely lean. Everything on an animal carcass was eaten, including the small fat depots and organ meats (which were highly prized), bone marrow, some stomach contents, peritoneal fluid and blood. A wide variety of uncultivated plant foods was eaten in the traditional diet: roots, starchy tubers, seeds, fruits and nuts. The plant foods were generally high in fibre and contained carbohydrates, which was slowly digested and absorbed. Traditional methods of food preparation (usually baked whole or eaten raw) ensured maximum retention of nutrients. In general, traditional foods had a low energy density but high density of some nutrients. The low energy density of the diet and the labour intensity of food procurement provided a natural constraint on energy intake.(ABSTRACT TRUNCATED AT 250 WORDS)     

The Influence of Seasonal Variation on Chimpanzee (Pan troglodytes schweinfurthii) Fallback Food Consumption, Nest Group Size, and Habitat Use in Gishwati, a Montane Rain Forest Fragment in Rwanda

The increased number of primates living in fragmented habitats necessitates greater knowledge of how they cope with large-scale changes to their environment. Chimpanzees (Pan troglodytes) are exceptionally vulnerable to forest fragmentation; however, little is known about chimpanzee feeding ecology in fragments. Although chimpanzees have been shown to prefer fruit when it is available and fall back on more abundant lower quality foods during periods of fruit scarcity, our understanding of how chimpanzees use fallback foods in forest fragments is poor. We examined how chimpanzees cope with periods of fruit scarcity in Gishwati Forest Reserve, a disturbed montane rain forest fragment in Rwanda. We assessed seasonal changes in chimpanzee diet and the use of preferred and fallback foods through fecal and food site analysis. We also examined seasonal variation in nest group size and habitat use through marked nest censuses. We found that chimpanzees experienced a seasonal reduction in preferred fruit availability, which led to a seasonal diet shift to more fibrous foods, including several that functioned as fallback foods. Our results suggest that during periods of fruit scarcity the chimpanzees also reduced nest group size. However, we found that the chimpanzees did not alter their habitat use between high- and low-fruit seasons, which suggests that the small size of the forest limits their ability to change their seasonal habitat use. Consequently, fallback foods appear to be particularly important in small food-impoverished habitats with limited ranging options.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

Seed dispersal mechanisms and the vegetation of forest islands in a West African forest-savanna mosaic (Comoé National Park,Ivory Coast)

The transition zone between forest and savanna is typically characterized by a dynamic patchwork of forest and savanna. We studied the woody plant species composition of 49 forest islands, 18 savanna, and 3 gallery forest plots in the Comoé National Park (Ivory Coast), West Africa's largest savanna reserve. TWINSPAN makes a clear distinction in vegetational composition between these three major habitat types but, nevertheless, more than 50% of the 292 species occur in at least 2 of them. The gallery forest is dominated by Cynometra megalophylla (Caesalpiniaceae), a genus known to dominate humid forests in other afrotropical regions. Ordination reveals four distinct categories of forest islands, (1) humid forests comparable to the gallery forest in their species composition, (2) dry disturbed and (3) dry undisturbed forests and (4) forests formerly inhabited by humans. Disturbed forests harbor more savanna species but also a distinct group of disturbance-tolerant forest species. Compared to other forest species, in this latter group we found an exceptionally high fraction of animal dispersed species (80% vs. 58%), while wind dispersed species or species lacking long distance seed dispersal mechanisms were correspondingly rare. This pattern occurs in spite of the fact that the frequency of wind dispersed species in general increases from dense humid forests to open dry forests and savanna. Species lacking long distance dispersal mechanisms are most abundant among those specialized on humid forests. These observations suggest that the species composition of forest islands is to some extent determined by the seed dispersal abilities of the different species.     

High flexibility in diet and ranging patterns in two golden monkey (Cercopithecus mitis kandti) populations in Rwanda

Many primates exhibit behavioral flexibility which allows them to adapt to environmental change and different habitat types. The golden monkey (Cercopithecus mitis kandti) is a little-studied endangered primate subspecies endemic to the Virunga massif and the Gishwati forest in central Africa. In the Virunga massif, golden monkeys are mainly found in the bamboo forest, while in the Gishwati forest they live in mixed tropical montane forest. Here we describe and compare the diet of golden monkeys in both fragments. Over 24 consecutive months from January 2017 we used scan sampling to record feeding and ranging behavior of two Virunga groups and one Gishwati group totaling ca. 240 individuals. We also examined the phenology of bamboo and fruit trees, key seasonal food plant species for the monkeys. Golden monkeys fed on more than 100 plant species. The Virunga groups were mostly folivorous (between 72.8% and 87.16% of the diet) and fed mostly on young bamboo leaves and bamboo shoots, while 48.69% of the diet of the Gishwati group consisted of fruit from 22 different tree and shrub species. Bamboo shoots and fruit are seasonally available foods and were consumed regularly throughout the period when they were available. Despite being the smallest of the three study groups, the Gishwati group had a larger home range area (150.07 ha) compared to both Virunga groups (25.24 and 91.3 ha), likely driven by the differences in availability and distribution of fruit and bamboo in the habitats. Like other blue monkey subspecies, golden monkeys appear to have a flexible dietary strategy enabling them to adjust diet and ranging behavior to local habitats and available food resources. Additional studies and continuing conservation efforts are needed to better understand how variation in feeding and ranging ecology affects reproduction, population growth, and carrying capacity.     

Baboon diet: a five-year study of stability and variability in the plant feeding and habitat of the yellow baboons (Papio cynocephalus) of Mikumi National Park, Tanzania

The habitat and plant feeding of 64 well-habituated, individually identified adult male and female yellow baboons was studied for 5 years at Mikumi National Park, Tanzania. Variation across the years showed that a study of only one or two years would have been incomplete and misleading. The list of baboon food species obtained from Mikumi is considerably larger and more diverse than any previously reported. One to six plant parts were eaten from each of more than 180 species. The 25 most common tree genera all contained species used for food. Of the 50 most common grass, shrub and herb genera, 93% included plant foods. Using months in which a species was eaten during at least one year of the study, 21 staple species were eaten during a mean of 8.86 months and 7 were eaten in all 12 months. Although many foods were from commonly available plant species, 15 such species were only rarely eaten. The number of parts of a species eaten per month and an estimate of the amounts eaten per month both varied with temperature and rainfall, being lowest near the end of the cool, dry season. There were substantial differences from year to year in the timing and amount of food production of many species; nevertheless, the same broad feeding pattern was repeated in each of the 5 years of the study. Despite yearly variation in food availability, 14 or more staples and other common foods were eaten in any given month. If crops of many of these foods were to fail, a large number of less commonly eaten species could be substituted. Baboons are eclectic feeders that appear to be optimizing their diet by selective feeding from among a wide array of available foods in an ever-changing floristic environment.     

The tropical rain forest: Is it a productive environment for human foragers?

A recent debate in ecological anthropology concerns the availability of wild foods for human foragers in tropical forests. This article is a response to the five essays in this same issue of Human Ecology that examine the hypothesis that hunter-gatherers could never have lived in tropical rain forest without direct or indirect access to cultivated foods. We clarify the hypothesis and assess the evidence offered to date. Archeological evidence suggests foraging without cultivation in Malaysia. We propose a program of ecological studies and archeological research which, if undertaken, should provide the evidence necessary to falsify the hypothesis.     

Diet of a Japanese Macaque Troop in the Coniferous Forest of Yakushima

I studied the diet of a troop of Japanese macaques in the coniferous forest of Yakushima over one year via focal animal sampling. Fiber-rich foods constituted 45% of annual feeding time, and mature leaves constituted 38% of total feeding time. Feeding time on fruits and seeds was only 13% and 4%, respectively. Flowers and fungi contributed a considerable amount of annual feeding time: 15% and 14%, respectively. Their diet changed seasonally. They fed on more fruits and seeds in response to increased availability, and when these foods were not available, they ate mature leaves. When the temperature was low, they ate more herbs, possibly to save energy by not climbing trees and staying in sunny places. The results imply two dietary characteristics of the species that might relate to adaptations in temperate regions, where fruit is available during a limited season and fiber-rich foods are the only candidate of fallback food. They are capable of digesting a large amount of fiber-rich food, but at the same time they preferentially select high-quality fruits or seeds when they are available.