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.     

Hunting in lowland,tropical rain forest: Towards a model of non-agricultural subsistence

It has recently been argued that hunter-gatherers do not, did not, and could not live in tropical rain forest without some access to agricultural produce. This opinion challenges models of past non-agricultural subsistence patterns that are based in analogies derived from modern rain forest dwelling groups. In this paper, the socio-ecological bases of the hunting system of the Kubo people of lowland Papua New Guinea are described. It is argued that this system lacks necessary dependence upon the agricultural system with which it co-occurs and, in fact, can be connected with a system of carbohydrate procurement that is not agricultural. The hypothesized connection provides the basis of a model of non-agricultural subsistence in lowland tropical rain forest.     

Foraging in tropical rain forests: The case of the penan of Sarawak,East Malaysia (Borneo)

Bailey et al. (1989) and Headland (1987) have recently proposed hypotheses stating that human foragers are unable to live in undisturbed tropical rain forests without some reliance on cultivated foods. The present discussion considers these hypotheses, as well as some of the evidence by which they have been tested. Four conceptual problems in the way these hypotheses have been formulated are identified: (1) assumptions about the relationship between key features of tropical forest ecosystems and human subsistence potential, (2) in-consistencies in the definition of pure foraging, (3) adherence to a dichotomy between foraging and agriculture, the result being that conscious and unconscious effects of exploitation on the demographic parameters of key resources is ignored, and (4) problems in defining the significance of ecotones. I consider the case of Penan hunter-gatherers of Borneo, a population which, by virtue of their reliance on the sago palm Eugeissona utilis, contradicts the conclusions of Bailey et al. and Headland. I consider salient aspects of Penan reliance on Eugeissona, and describe how Penan exploitation of this resource may positively effect its availability. This case is seen to provide a challenge to the hypotheses of Bailey et al. and Headland, not only in the extent to which it contradicts their conclusions but, more significantly, in what it reveals about the assumptions upon which their hypotheses are based. This points to the need for greater precision in the definition of future hypotheses about foraging in tropical forests.     

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.     

Hunting and Gathering in Tropical Rain Forest: Is It Possible?

Hunters and gatherers living in tropical forests represent an important part of the total range of variation among contemporary hunting and gathering societies. Studies of tropical forest hunting and gathering peoples have contributed to our perceptions of the foraging way of life. Yet no peoples have ever been directly observed living independently of agriculture in tropical rain forest. This article tests the hypothesis that humans do not exist nor have ever existed independently of agriculture in tropical rain forest. We find no convincing ethnographic evidence and, with the possible exception of Malaysia, no archeological evidence for pure foragers in undisturbed tropical rain forests. Negative evidence cannot be conclusive, but it suggests that we need to carefully reexamine common assumptions concerning the recent history of tropical forest dwellers, the adaptability of preagricultural humans, the geographic and environmental range of hominids, and the form and consequences of selection pressures acting on humans in warm, humid environments. The overriding purpose of this article is to stimulate further ecological and archeological research in the neglected tropical forest areas of the world.     

Long-Term Foraging Expeditions (Molongo) among the Baka Hunter-Gatherers in the Northwestern Congo Basin, with Special Reference to the “Wild Yam Question”

While “Pygmy” hunter-gatherers have generally been assumed to be the original inhabitants of the central African rainforest, recent studies dispute the viability of subsistence in tropical rainforests through foraging alone. There are, however, few studies that are based on sound data of actual long-term foraging. This paper examines the viability of “pure” foraging in tropical rainforests, based on data from participant observation of a long-term foraging expedition among the Baka in the northwestern Congo Basin. The group was observed to subsist solely on wild food resources, particularly wild yams, for two-and-half months. The study was carried out during the dry season when food resources are generally thought to be scarce in tropical rainforests.     

西双版纳野生亚洲象的觅食行为

以西双版纳勐养自然保护区的野象谷为基本监测点,通过定点摄像观察和跟踪观察,采用全事件取样、焦点动物取样和扫描取样相结合的方法,从2003 年8 月到2004 年7 月对三岔河地区野生亚洲象的觅食行为进行初步研究。发现野生亚洲象的雌性成体和未成年个体选择集群采食,而成年雄象一般单独采食,亚洲象在昼夜间的采食行为没有显著性差异,且有多种采食方式;采食农作物主要在夜间进行。亚洲象对主要采食地点采取循环利用的方式,这既提高了食物的数量和质量,又有利于热带雨林生态系统的更新与演化。     

Nest demographics and foraging behavior of <Emphasis Type="Italic">Apterostigma collare</Emphasis> Emery (Hymenoptera,Formicidae) provide evidence of colony independence

Apterostigma collare Emery is a highly derived fungus-growing ant within the Tribe Attini whose small, fungal nests are found in tropical rain forests. This study focuses on determining the colony structure of A. collare, specifically searching for evidence of polydomy or independence. We surveyed and observed nests in the field, and performed foraging bioassays and dissected nests in the laboratory. We determined the size and contents of nests in field populations. Nests found near other nests were not statistically different in size compared to nests found alone. There was also no statistical difference between near and lone nests regarding the presence of a queen in the nest. Most nests contained one queen with brood and workers, regardless of their proximity to other nests. Observations also were made of foraging and trail-marking behaviors. Foraging activity observed in the field revealed that workers left the nest area and followed trails upwards into the canopy, but they did not interact with foragers from other nearby nests. In a laboratory foraging arena, foragers marked a trail to a food source by dragging the gaster. Bioassays showed that A. collare workers preferred their own foraging trails, but not those of other conspecific colonies. All results suggest that each nest represents an independent colony, supporting a previous report that nests found in close proximity do not constitute a polydomous colony. Received 19 July 2006; revised 23 March 2007; accepted 6 June 2007.     

Use of Long-Term Stored Vector Information in the Neotropical Ant <Emphasis Type="BoldItalic">Gigantiops destructor</Emphasis>

We investigated how the formicine ant Gigantiops destructor can use vector information to navigate within the cluttered environment of the rain forest. Displaced foragers use skylight information to move in the theoretical feeder-to-nest direction, whether they are prevented from updating their path-integrator during foraging or captured at the departure from their nest, i.e. with a current accumulator state very close to zero. Only ants that have collected food are able to download a long-term stored reference vector pointing in the nest direction, irrespective of the current accumulator state of their path-integrator stored in a working memory and independent of familiar landmarks. Depending on the release sites, ants that became lost at a maximum distance of 50 cm could still hit and recognize their familiar route, or they engaged in a systematic search for it centered on the release sites. In contrast to Cataglyphis desert ants, Gigantiops ants do not rely primarily on the current accumulator state of their egocentric path integrator. Such a long-term vector-based navigation primed by food capture is well adapted for a tropical ant foraging during periods spanning several hours. This could prevent the numerous cumulative errors in the evaluation of the angles steered that might result from a continuously running path-integrator operating during complex foraging patterns performed at ground or arboreal levels and during passive displacement in response to heavy rain.     

Making a living in the tropical forest: Yuquí foragers in the Bolivian Amazon

Questions concerning the availability of resources in tropical rain forests have given rise to the current debate centering on whether human subsistence based solely on foraging is possible in these biomes without agricultural subsidies. This paper takes the position that changing perspectives on ecological pattern and process in tropical forests and the significant variation among tropical forests on a worldwide as well as regional scale must be taken into consideration. Human disturbance is also proposed as a cause of dependence on agriculture by modern human foragers rather than as a necessary precondition for successful exploitation of the tropical forest. These issues are discussed against the background of a case study of the Yuquí, who, until very recently, were true foragers in the Bolivian Amazon. For the Yuquí, the sustainability of their subsistence system depended on a finegrained knowledge of their environment and the freedom of movement over a large territory to access resources within it.     

Impact of live food on survival and growth of hatchery‐reared sea trout (Salmo trutta trutta L.) parr in the wild

Survival rates and growth parameters of hatchery‐reared sea trout (Salmo trutta trutta L.) fry were determined after stocking in the wild. The larvae were hatchery‐reared for 12 weeks in two groups: fry were fed either on live zooplankton and live chironomidae larvae (LFG), or fed a pellet diet (PFG). The survival rate and specific growth rates were higher in the LFG than in the PFG group. Most effective for hatchery‐reared fish intended for stocking was the natural, live feed. The mean number of chironomid larvae found in the stomachs of fish that were initially captured in the wild was significantly higher in the LFG than in the PFG group. The live diet supplied in the rearing period had a positive impact on the foraging skills of the sea trout fry and their survival in the wild after their release on 24 April 2010.     

Unpaved roads alter foraging patterns of the leafcutter ant Atta colombica

Leaf cutting ants are dominant herbivores and influential ecosystem engineers in the Neotropics. It has been suggested that habitat disturbances alter the architecture of foraging trail systems for colonies in their vicinity; however, the evidence remains scarce. In this study we investigated the effect of unpaved roads dissecting tropical lowland forest habitat on the structure of leafcutter foraging trail systems and foraging effort. We mapped trail systems for 16 mature Atta colombica colonies located at different distances from unpaved roads. Our results suggest exploitation of unpaved roads by leafcutters provides favorable foraging conditions, causing significant differences in foraging trail structure.     

Niche overlap and polyspecific associations among sympatric cercopithecids in the campo animal reserve,Southwestern Cameroon

The community structure and polyspecific associations found in five sympatric resident primates (Cercopithecus nictitans, C. cephus, C. pogonias, Cercocebus torquatus, andC. albigena) and a temporary visitor (Mandrillus sphinx) were studied within the Campo Animal Reserve, a tropical rain forest in southwestern Cameroon. Several ecological variables of the six species, e.g. the number of foraging groups in the study area, group size, home range size, biomass density, and individual density, were estimated by undertaking a systematic census and by other means, e.g. tracing.C. nictitans was the most abundant species in terms of its foraging groups, biomass, and individual densities. The foraging biomass varied significantly among the six species. The five resident species frequently formed polyspecific groups.M. sphinx also associated with them when it visited the area. The food items fed on by the five resident species were also checked. An index for synecological analysis revealed that the five species shared similar food niches. The five species also utilized similar foraging areas and strata, possibly depending on the availabilities of common foods. The above results cannot be explained by conventional equilibrium-competition models. For understanding the polyspecific associations, it is proposed that the primates form polyspecific foraging groups in order to optimize their foraging biomass. This idea leads to a model that can also explain certain of the grouping behaviors, such as group fission-fusion, found in large body size primates.     

What,where and when: spatial foraging decisions in primates

When exploiting the environment, animals have to discriminate, track, and integrate salient spatial cues to navigate and identify goal sites. Actually, they have to know what can be found (e.g. what fruit), where (e.g. on which tree) and when (in what season or moment of the year). This is very relevant for primate species as they often live in seasonal and relatively unpredictable environments such as tropical forests. Here, we review and compare different approaches used to investigate primate spatial foraging strategies: from direct observations of wild primates to predictions from statistical simulations, including experimental approaches on both captive and wild primates, and experiments in captivity using virtual reality technology. Within this framework, most of these studies converge to show that many primate species can (i) remember the location of most of food resources well, and (ii) often seem to have a goal‐oriented path towards spatially permanent resources. Overall, primates likely use mental maps to plan different foraging strategies to enhance their fitness. The majority of studies suggest that they may organise spatial information on food resources into topological maps: they use landmarks to navigate and encode local spatial information with regard to direction and distance. Even though these studies were able to show that primates can remember food quality (what) and its location (where), still very little is known on how they incorporate the temporal knowledge of available food (when). Future studies should attempt to increase our understanding of the potential of primates to learn temporal patterns and how both socio‐ecological differences among species and their cognitive abilities influence such behavioural strategies.     

Extended phenotypes and foraging restrictions: ant nest entrances and resource ingress in leaf‐cutting ants

Several factors may restrict the acquisition of food to below the levels predicted by the optimization theory. However, how the design of structures that animals build for foraging restricts the entry of food is less known. Using scaling relationships, we determined whether the design of the entrances of leaf‐cutting ant nests restricts resource input into the colony. We measured nests and foraging parameters in 25 nests of Atta cephalotes in a tropical rain forest. Ant flux was reduced to up to 60% at nest entrances. The width of all entrances per nest increased at similar rates as nest size, but the width of nest entrances increased with the width of its associated trail at rates below those expected by isometry. The fact that entrance widths grow slower than trail widths suggests that the enlargement of entrance holes does not reach the dimensions needed to avoid delays when foraging rates are high and loads are big. The enlargement of nest entrances appears to be restricted by the digging effort required to enlarge nest tunnels and by increments in the risk of inundation, predator/parasitoid attacks and microclimate imbalances inside the nest. The design of the extended phenotypes can also restrict the ingress of food into the organisms, offering additional evidence to better understand eventual controversies between empirical data and the foraging theory. Abstract in Spanish is available with online material.     

ANT DISPERSAL OF CALATHEA (MARANTACEAE) SEEDS BY CARNIVOROUS PONERINES (FORMICIDAE) IN A TROPICAL RAIN FOREST

Two tropical herbaceous plant species, Calathea microcephala and Calathea ovandensis, exhibit morphological adaptations for seed dispersal by ants. In the field, 21 ant species are attracted to the arils of the seeds. Previously known as predatory carnivores, the ponerine ants Odontomachus laticeps, O. minutus, Pachycondyla harpax, and P. apicalis carry the seeds toward their nests, behaving as though they were carrying prey. These ants remove the aril in the nest, and seeds without arils germinate more readily than seeds with arils. Ant foraging distances can account for seedling distributions. Myrmecochory may be far more common in tropical rain forests than has been previously suspected.     

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.     

Natural history and foraging behavior of the carpenter ant Camponotus sericeiventris Guérin, 1838 (Formicinae, Campotonini) in the Brazilian tropical savanna

Camponotus sericeiventris is a polymorphic ant living in populous colonies at tropical forests and cerrado formation. This study provides a detailed account of the natural history and foraging biology of C. sericeiventris in cerrado at Ecological Station of Panga, Southeast of Brazil. The nest distribution according to vegetation physiognomies, activity rhythm, diet, and foraging patterns were described. Results showed that nests occur inside dead or live trunks, and also in branches of soft wood at cerradão and gallery forest physiognomies (approximately 1 nest/100m²), but not in the mesophytic forest. Ant activity is correlated with temperature and humidity. There is overlap in the foraging area among neighbor colonies (as far as 28 m) without evidence of agonistic interactions. Foragers leave the nest independently or in groups and frequently searched for food individually. Workers are generalistic feeders visiting flowers and extrafloral nectaries, attending Hemiptera (aphids and membracids) and Lepidoptera (Lycaenidae larvae), collecting seeds and fruits, and hunting for live preys as well as scavenging for dead animals. The great number of interactions with different plant and animals suggests that this species has an important participation in the interaction web in this environment.     

Is there a latitudinal gradient in seed production?

We compiled data from seed rain studies at 33 sites from around the world to determine whether the greater mean seed mass of tropical plants is associated with production of fewer seeds per square meter of ground. We found no significant linear relationship between latitude and annual seed rain density, but found some evidence for a mid‐latitude peak in seed rain density (quadratic relationship, p=0.018; R²=0.23). Combining seed rain data with seed mass data suggests that vegetation at the equator produces between 19 and 128 times more total mass of seed per year than does vegetation at 60°. This gradient in seed production would far outweigh the doubling in net primary productivity (NPP) over the same range of latitudes. Thus, our (admittedly small) dataset suggests that tropical vegetation allocates a much greater proportion of NPP to reproduction. This raises two important questions for the future: 1) why might tropical vegetation commit more energy to seed production than vegetation further from the equator? 2) What aspect of plant growth might receive proportionally less energy in tropical ecosystems?     

Twelve years of repeated wild hog activity promotes population maintenance of an invasive clonal plant in a coastal dune ecosystem

Invasive animals can facilitate the success of invasive plant populations through disturbance. We examined the relationship between the repeated foraging disturbance of an invasive animal and the population maintenance of an invasive plant in a coastal dune ecosystem. We hypothesized that feral wild hog (Sus scrofa) populations repeatedly utilized tubers of the clonal perennial, yellow nutsedge (Cyperus esculentus) as a food source and evaluated whether hog activity promoted the long‐term maintenance of yellow nutsedge populations on St. Catherine's Island, Georgia, United States. Using generalized linear mixed models, we tested the effect of wild hog disturbance on permanent sites for yellow nutsedge culm density, tuber density, and percent cover of native plant species over a 12‐year period. We found that disturbance plots had a higher number of culms and tubers and a lower percentage of native live plant cover than undisturbed control plots. Wild hogs redisturbed the disturbed plots approximately every 5 years. Our research provides demographic evidence that repeated foraging disturbances by an invasive animal promote the long‐term population maintenance of an invasive clonal plant. Opportunistic facultative interactions such as we demonstrate in this study are likely to become more commonplace as greater numbers of introduced species are integrated into ecological communities around the world.