Just as human cultures use different utensils for food, so do different groups of chimps. Think about the utensils we humans use to eat our food. In Asian countries it is usual for chopsticks to be used, while in America forks and knives are common utensils. What do you eat with at your house? We and chimpanzees are both very similar in the ways we use and create tools for our benefit.
Jane Goodall first discovered chimpanzees using tools to capture termites, which were buried underground in sealed mounds of dirt called termite mounds. She noticed that they would use two different sticks as tools to acquire the protein rich insects. The first thing chimps do is use a heavy stick to chisel a hole into the termite mound to find an opening.
The second tool they use is much more refined than their chisel. The second stick is known as a fishing tool and it is very special because they actually modify it before using it. The chimpanzees find a straight stick or long blade of grass and pull all of the leaves off of it, so that they can easily fit the stick into the hole of the termite mound. Chimps are much better at termite fishing than we are. Tool modification is extremely rare in the animal world which is yet another example of how chimps and humans are so similar.
In another group of chimps, different from the culture that Jane Goodall studied, there was a discovery of a hammer and anvil tool top of page being used to crack open hard nut shells. Shkolnik Haifa University for their help; L. Grossman and O.
Haroch for the 3D photographs; and A. Regev-Gisis for the graphic display. The authors declare no conflict of interest. This article contains supporting information online at www. National Center for Biotechnology Information , U.
Published online Aug Author information Copyright and License information Disclaimer. E-mail: li. Contributed by Eviatar Nevo, July 30, sent for review May 25, Copyright notice. This article has been cited by other articles in PMC. Abstract Using direct percussion, language-competent bonobo-chimpanzees Kanzi and Pan-Banisha produced a significantly wider variety of flint tool types than hitherto reported, and used them task-specifically to break wooden logs or to dig underground for food retrieval.
Keywords: hominin, bonobo targeted tool use, stone tool wear pattern, food acquisition, bonobo survival strategy. Results All of the stone tools made by KZ and PB were created solely by direct percussion, with the core held in the left hand and the hammer stone in the right hand Fig. Open in a separate window. Log-Breaking Experiments and Quantification. Digging Experiments and Quantification.
Discussion Stone tool uses have evolved separately across taxa; for example, Neophron vultures use pebbles to break eggs, Cariama seriema birds kill prey by striking them on rocks, and Cebus capuchin monkeys use stones as hammers to break nuts on flat rock anvils 9. Materials and Methods Background. Tasks Given. Supplementary Material Supporting Information: Click here to view. Acknowledgments We thank the late Prof. Footnotes The authors declare no conflict of interest.
References 1. Human evolution: Taxonomy and paleobiology. J Anat. Uddin M, et al. Sister grouping of chimpanzees and humans as revealed by genome-wide phylogenetic analysis of brain gene expression profiles. Goodall J. The Chimpanzees of Gombe: Patterns of Behavior. Pruetz JD, Bertolani P. Savanna chimpanzees, Pan troglodytes verus , hunt with tools. Curr Biol. Savanna chimpanzees use tools to harvest the underground storage organs of plants.
Complex tool sets for honey extraction among chimpanzees in Loango National Park, Gabon. J Hum Evol. Chimpanzee Cultures. McGrew WC.
Kortlandt A. The use of stone tools by wild-living chimpanzees and earliest hominids. Apes, Language, and the Human Mind. New York: Oxford Univ Press; Schick K, et al. Continuing investigations into the stone tool-making and tool-using capabilities of a bonobo Pan paniscus J Archaeol Sci.
The Oldowan: The tool making of early hominins and chimpanzees compared. Annu Rev Anthropol. Curr Anthropol. Environment and behavior of 2. Oliver J. Estimates of hominid and carnivore involvement in the FLK Zinjanthropus fossil assemblage: Some socioecological implications. Leakey MD. Vol 3. Cambridge, UK: Cambridge Univ. Ant dipping was related to rainfall, and honey gathering showed some coincidence with patterns of tree flowering which precedes the peak periods of honey production.
Rather than use tools to harvest fallback resources in response to seasonal fluctuations in preferred food resources, our conclusion is that these chimpanzees used their technical skills to maintain year-round access to embedded food items. In addition to ecological drivers, social factors are also likely to play a role in the invention and maintenance of complex tool using behaviours. Party size was consistent throughout the year and relatively small in tool using contexts within the Goualougo Triangle, which prompts further examination of social tolerance and specific types of social interaction on the evolution of technological traditions [ 13 , 73 , 74 ].
To compensate for scarcity of preferred resources, fallback foods must provide a major source of energy and be consumed at a degree that effectively replaces the nutrients provided by preferred food items. This seems feasible in the case of pestle pounding of oil palms and cracking of nuts among chimpanzees of Bossou in Guinea [ 31 ] and the nut cracking of Tai chimpanzees in Ivory Coast [ 33 ].
Although the food items harvested with tools are a rich source of nutrients, they comprise a relatively small component of the diet of the nut cracking capuchins at Fazenda Boa Vista in Brazil [ 44 ] and insect harvesting orangutans at Suaq Balimbing in Sumatra [ 29 ]. Chimpanzees at both Gombe in Tanzania and Fongoli in Senegal allocated significant foraging time to harvesting Macrotermes during particular months [ 35 ]. Termites can be a significant source of several important nutrients, including manganese, protein and amino acids [ 46 , 75 ], and are considered an important food in the Fongoli chimpanzee diet [ 35 ].
However, social insects do not fit the typical profile of fallback food resources [ 8 ]. While army ants may provide additional protein and micronutrients when they are consumed by chimpanzees, Koops et al. It was reported that the nutritional value of arboreal ants harvested with tools at Mahale appears to be negligible [ 40 ]. The amount of honey gathered from stingless bees Trigona spp. The primary nutritional benefit of honey gathering is carbohydrates primarily fructose, glucose and about 25 different oligosaccharides [ 76 ].
Honey also contains small amounts of proteins, enzymes, amino acids, minerals, vitamins and polyphenols [ 76 ]. Further research is required to evaluate if foods gathered with the aid of tools provide sufficient nutrition to serve as fallback foods or are indeed energetically more profitable than other food types.
Chimpanzees, macaques and capuchins are some of the most versatile primate species with regard to their ability to survive in different habitat types. Some habitats in which these primates live may not offer traditional fallback foods, and so primates are prompted to procure higher quality resources when their staple food items are scarce.
This could be the case with the seafood harvesting macaques on islands in the Andaman Sea [ 77 ] or the termite harvesting chimpanzees of Fongoli in Senegal [ 35 , 78 ]. Further, tool use may also be necessitated in localities where feeding competition within or between species is high [ 26 ].
Interspecific feeding competition could result in decreased abundance of particular foods through direct or scramble competition. Patterns of food consumption by other species could also mask or amplify seasonal patterns of food resource abundance. Interspecific competition could be elevated in central Africa where chimpanzees coexist with gorillas and other primates [ 79 ].
While segregated foraging and utilization of specialized feeding strategies are the typical coping mechanism of sympatric species that show a high degree of dietary overlap [ 80 , 81 ], another option would be the diversification or intensification of tool using strategies [ 16 , 82 ]. Reduced risk of predation is another ecological factor which could be associated with the evolution of tool use through reduced mortality which favours evolution of life histories that are more conducive to cognitive adaptations.
Reduced predation may also relax anti-predator vigilance and associated behaviours which could then liberate more time for object interaction and exploratory behaviours [ 22 ]. Changing ecological circumstances may also influence the emergence of tool use. Recent shifts in habitats caused by conversion or degradation may cause apes to increase the frequency of some particular types of tool use to compensate for missing resources. Pestle pounding tool use occurs in the canopy of oil palm trees Elaeis guineensis and so is limited by the distribution of this species, which is closely associated with human cultivation [ 26 ].
Another example of anthropogenic activities providing tool using opportunities is with the possible introduction of candlenut trees which harbour the beetle larvae that are extracted with tools by crows in New Caledonia [ 22 ]. There are several possible reasons why the necessity hypothesis fails to explain tool use in a number of species and situations.
One issue with testing the ecological hypotheses could be that relative abundance of preferred food items is not a valid proxy for necessity. It is possible that necessity is more nuanced and driven by requirements for particular micronutrients which do not need to be consumed in large quantities. Another explanation for the failure of the necessity hypothesis would be that in certain ecological settings the food resources may decrease during certain periods but do not reach a level of scarcity that prompts one to take additional action.
However, a wide variety of animals depend on seasonally variable food resources, and in most cases, this has not resulted in selection for tool using behaviours [ 22 ] for further details, see below. Animals may also be using multiple coping strategies. For example, chimpanzees of the Tai Forest in Ivory Coast show varying responses to seasonal variation in fruit resources including an increase in foraging activities [ 34 ], reduction in daily travel [ 83 ], and exploitation of high-quality food resources such as Coula edulis when available [ 34 ].
The Goualougo Triangle chimpanzee population provides an interesting case study to evaluate the validity of the opportunity hypothesis.
The abundance of tool using opportunities to gather army ants and honey show seasonal variation through the nesting patterns of ants and the honey production of bees [ 16 , 64 ], with chimpanzee tool use showing some indication of tracking these target resources. The Goualougo chimpanzees also regularly prey upon three different species of termites that live throughout the year in earthen mounds, whose protective structures present a significant obstacle to the foraging ape.
In both the ant dipping and termite fishing contexts, chimpanzees use tool sets to overcome accessibility issues; each of the tool sets is composed of a first tool used to access the prey and a second tool to gather the insects. Multiple tools are also used by chimpanzees in central Africa to open beehives and dip for honey [ 16 , 18 ].
In these foraging contexts, chimpanzees have overcome the constraints of their environment through technological innovations that served to increase their tool using opportunities and in some cases, tool modifications that enhance the profitability of tool-assisted foraging [ 62 ]. Although certain aspects of opportunity may trigger tool use, mere ecological opportunity may not be sufficient to support a tool using tradition [ 12 , 55 ]. The inconsistency in support for the necessity and opportunity hypotheses prompt us to question if they best represent the proximate drivers of tool use.
Therefore, the absolute amount of food in the environment might be less important than the relative profitabilities of different foraging modes. They suggest that the relative profitabilities of attaining embedded and non-embedded foods may be a driving force in the evolution of tool-assisted foraging strategies.
One would then predict that tool use would be performed whenever it is more profitable than non-tool-assisted foraging techniques i. Seasonal patterns in tool use behaviour could then simply be attributed to shifts in the relative profitabilities of different foraging techniques.
Social factors driving tool use also merit further examination and more rigorous testing of the invention hypothesis. Our results prompt us to question whether the number of conspecifics in the party or even at the tool using site is a good proxy for the potential of social transmission. We observed that it was often a subset of individuals in a chimpanzee party that actually engaged in tool using behaviour. Others may be within the immediate party, but not stimulated by the tool using event and so are not representative of the opportunities for social transmission.
Within the Goualougo Triangle, we found that the number of individuals at the tool using site was surprisingly small and visit duration was shorter than at other sites, yet sufficient to yield interconnectedness among all mature members of the community in a termite gathering social network [ 61 ].
We suggest that group members can gain important information about the characteristics of the tool site, tools required, techniques and outcomes from other tool users who act as tolerant or possibly even instructive models. This may serve to facilitate the acquisition of tool using skills and promote conformity in specific behavioural traditions within social groups [ 84 ] and populations [ 37 , 85 ].
Further, patterns of innovation the successful diffusion or spread of an invention are likely to drastically differ between tool using contexts. Foraging at a terrestrial termite nest has a very different ecological and social setting compared with arboreal honey gathering [ 86 ].
As recently reviewed in Meulman et al. As illustrated by many of the contributions to this special issue and other compilations [ 3 , 4 ], our understanding of the tool using abilities of animals has remarkably increased in the past decade. There have also been great strides in documenting the diversity of these tool using skills and the natural environments in which they occur.
Further, research in this field has progressed from descriptive accounts to more hypothesis-driven research to determine the factors shaping particular behaviours. Longitudinal studies have also provided unique vantages into the development, maintenance and innovation of these skills. Such research moves us toward a more comprehensive understanding of how animals interact with and shape their environment through the use of technology. However, the opportunities to study these behaviours among wild apes are increasingly endangered by habitat destruction and conversion, illegal hunting and infectious diseases.
Many would agree that there is a moral obligation to ensure the preservation of our closest living relatives, but we also have an obligation to safeguard their unique traditions which have their own rich evolutionary histories [ 87 — 89 ]. Special thanks are due to J. Fay, P. Telfer, P. Elkan, S.
Elkan, B. Curran, M. Gately, E. Stokes, T. Breuer, P. Ngouembe and D. Dos Santos. We are deeply indebted to R. Mundry for his continuing support and willingness to conduct the statistical analyses for this manuscript.
This research greatly benefited from discussions and insights from E. Lonsdorf and C. We thank C. Rutz and two anonymous referees for their excellent comments and suggestions which greatly improved this manuscript. We also recognize the tireless dedication of J.
Onononga, C. Eyana-Ayina, S. Ndolo, A. Nzeheke, W. Mayoukou, M. Meguessa, I. Singono and the Goualougo tracking team. National Center for Biotechnology Information , U. Crickette M. Sanz 1, 2 and David B. Morgan 2, 3. David B. Author information Copyright and License information Disclaimer. All rights reserved. This article has been cited by other articles in PMC. Abstract The emergence of technology has been suggested to coincide with scarcity of staple resources that led to innovations in the form of tool-assisted strategies to diversify or augment typical diets.
Keywords: Pan troglodytes troglodytes , tool use, necessity, opportunity, limited invention, relative profitability. Introduction Scientists have long attempted to identify the factors that coincided with the emergence of complex tool use within the hominin lineage.
Table 1. Open in a separate window. Sources page numbers refer to original publications :. Figure 1. Rainfall and temperature in the Goualougo Triangle, Republic of Congo. Figure 2. Figure 3. Proportion of chimpanzee party scan observations spent foraging on fruits and leaves. Figure 4. Figure 5. Figure 9.
Figure 6. Figure 7. Chimpanzee tool use in ant predation was positively related to rainfall. Figure 8. Discussion In this study, we examined possible ecological and social correlates of the diverse and complex tool using behaviour exhibited by a chimpanzee population residing in central Africa.
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