Susana Carvalho

Chaînes opératoires and resource-exploitation strategies in chimpanzee (Pan troglodytes) nut cracking

We apply archaeological methods to extend our knowledge of chimpanzee material culture. The chaîne opératoire conceptual framework, as introduced by ethnography, established technology as a phased process. Prehistoric archaeology adopted this concept to elucidate technological variability in tool-making procedures, based on knowledge of tool functions or subsistence patterns. We focused on the detection of operational sequences by wild chimpanzees (Pan troglodytes verus) when nut cracking with lithic implements at the sites of Bossou and Diecké, Guinea, West Africa. Thus, while it has recently been claimed that chimpanzees leave behind recognizable assemblages of stone hammers that can be morphologically distinguished from Oldowan hammers, this is the first study to focus specifically on the existence of operational sequences during the utilization of stone tools by wild chimpanzees. By combining primatological and archaeological methods and examining ecological areas inhabited by different chimpanzee groups, we sought technological variability and identified variables influencing regional diversity in tool typology and technology. We compared three case studies: (1) Bossou-direct recording of experimental nut-cracking sessions; (2) Bossou- direct and indirect monitoring of nut-cracking sites in the wild; (3) Diecké-indirect monitoring of nut-cracking sites in the wild. Results suggest that chimpanzees perform sequences of repeated tool transport and nut cracking. Data show discrimination of tool functions based on tool features. We identified the most technologically complex tool for nut cracking, which was composed of four stones. We found regional diversity in chimpanzee stone assemblages. Raw-material type and tool mobility constrain technological development in human and nonhuman primates. Spatial analysis of tool distribution indicates a pattern of resource-exploitation strategy, revealing affinities with Oldowan.

Apes in the Anthropocene: flexibility and survival

We are in a new epoch, the Anthropocene, and research into our closest living relatives, the great apes, must keep pace with the rate that our species is driving change. While a goal of many studies is to understand how great apes behave in natural contexts, the impact of human activities must increasingly be taken into account. This is both a challenge and an opportunity, which can importantly inform research in three diverse fields: cognition, human evolution, and conservation. No long-term great ape research site is wholly unaffected by human influence, but research at those that are especially affected by human activity is particularly important for ensuring that our great ape kin survive the Anthropocene.

Tool-composite reuse in wild chimpanzees ( Pan troglodytes ): archaeologically invisible steps in the technological evolution of early hominins?

Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuable data towards elucidating the variables that influenced the emergence and development of the first lithic industries among Plio-Pleistocene hominins. Such data help to identify potential behaviours entailed in the first percussive technologies that are invisible in archaeological records. The long-term research site of Bossou in Guinea features a unique chimpanzee community whose members systematically use portable stones as hammers and anvils to crack open nuts in natural as well as in field experimental settings. Here we present the first analysis of repeated reuse of the same tool-composites in wild chimpanzees. Data collected over 5 years of experimental nut-cracking sessions at an “outdoor laboratory” site were assessed for the existence of systematic patterns in the selection of tool-composites, at group and at individual levels. Chimpanzees combined certain stones as hammer and anvil more often than expected by chance, even when taking into account preferences for individual stones by themselves. This may reflect an ability to recognise the nut-cracker as a single tool (composed of two elements, but functional only as a whole), as well as discrimination of tool quality-effectiveness. Through repeatedly combining the same pairs of stones—whether due to preferences for particular composites or for the two elements independently—tool-users may amplify use-wear traces and increase the likelihood of fracturing the stones, and thus of detaching pieces by battering.

Chimpanzee carrying behaviour and the origins of human bipedality

Summary Why did our earliest hominin ancestors begin to walk bipedally as their main form of terrestrial travel? The lack of sufficient fossils and differing interpretations of existing ones leave unresolved the debate about what constitutes the earliest evidence of habitual bipedality. Compelling evidence shows that this shift coincided with climatic changes that reduced forested areas, probably forcing the earliest hominins to range in more open settings [1]. While environmental shifts may have prompted the origins of bipedality in the hominin clade, it remains unknown exactly which selective pressures led hominins to modify their postural repertoire to include a larger component of bipedality [2]. Here, we report new experimental results showing that wild chimpanzees walk bipedally more often and carry more items when transporting valuable, unpredictable resources to less–competitive places.

Use-Wear Patterns on Wild Macaque Stone Tools Reveal Their Behavioural History

Burmese long-tailed macaques (Macaca fascicularis aurea) are one of a limited number of wild animal species to use stone tools, with their tool use focused on pounding shelled marine invertebrates foraged from intertidal habitats. These monkeys exhibit two main styles of tool use: axe hammering of oysters, and pound hammering of unattached encased foods. In this study, we examined macroscopic use-wear patterns on a sample of 60 wild macaque stone tools from Piak Nam Yai Island, Thailand, that had been collected following behavioural observation, in order to (i) quantify the wear patterns in terms of the types and distribution of use-damage on the stones, and (ii) develop a Use-Action Index (UAI) to differentiate axe hammers from pound hammers by wear patterns alone. We used the intensity of crushing damage on differing surface zones of the stones, as well as stone weight, to produce a UAI that had 92% concordance when compared to how the stones had been used by macaques, as observed independently prior to collection. Our study is the first to demonstrate that quantitative archaeological use-wear techniques can accurately reconstruct the behavioural histories of non-human primate stone tools.

First GIS Analysis of Modern Stone Tools Used by Wild Chimpanzees (Pan troglodytes verus) in Bossou, Guinea, West Africa

Stone tool use by wild chimpanzees of West Africa offers a unique opportunity to explore the evolutionary roots of technology during human evolution. However, detailed analyses of chimpanzee stone artifacts are still lacking, thus precluding a comparison with the earliest archaeological record. This paper presents the first systematic study of stone tools used by wild chimpanzees to crack open nuts in Bossou (Guinea-Conakry), and applies pioneering analytical techniques to such artifacts. Automatic morphometric GIS classification enabled to create maps of use wear over the stone tools (anvils, hammers, and hammers/ anvils), which were blind tested with GIS spatial analysis of damage patterns identified visually. Our analysis shows that chimpanzee stone tool use wear can be systematized and specific damage patterns discerned, allowing to discriminate between active and passive pounders in lithic assemblages. In summary, our results demonstrate the heuristic potential of combined suites of GIS techniques for the analysis of battered artifacts, and have enabled creating a referential framework of analysis in which wild chimpanzee battered tools can for the first time be directly compared to the early archaeological record.

Quantifying Traces of Tool Use: A Novel Morphometric Analysis of Damage Patterns on Percussive Tools

Percussive technology continues to play an increasingly important role in understanding the evolution of tool use. Comparing the archaeological record with extractive foraging behaviors in nonhuman primates has focused on percussive implements as a key to investigating the origins of lithic technology. Despite this, archaeological approaches towards percussive tools have been obscured by a lack of standardized methodologies. Central to this issue have been the use of qualitative, non-diagnostic techniques to identify percussive tools from archaeological contexts. Here we describe a new morphometric method for distinguishing anthropogenically-generated damage patterns on percussive tools from naturally damaged river cobbles. We employ a geomatic approach through the use of three-dimensional scanning and geographical information systems software to statistically quantify the identification process in percussive technology research. This will strengthen current technological analyses of percussive tools in archaeological frameworks and open new avenues for translating behavioral inferences of early hominins from percussive damage patterns.

Tools to tipple: ethanol ingestion by wild chimpanzees using leaf-sponges

African apes and humans share a genetic mutation that enables them to effectively metabolize ethanol. However, voluntary ethanol consumption in this evolutionary radiation is documented only in modern humans. Here, we report evidence of the long-term and recurrent ingestion of ethanol from the raffia palm (Raphia hookeri, Arecaceae) by wild chimpanzees (Pan troglodytes verus) at Bossou in Guinea, West Africa, from 1995 to 2012. Chimpanzees at Bossou ingest this alcoholic beverage, often in large quantities, despite an average presence of ethanol of 3.1% alcohol by volume (ABV) and up to 6.9% ABV. Local people tap raffia palms and the sap collects in plastic containers, and chimpanzees use elementary technology—a leafy tool—to obtain this fermenting sap. These data show that ethanol does not act as a deterrent to feeding in this community of wild apes, supporting the idea that the last common ancestor of living African apes and modern humans was not averse to ingesting foods containing ethanol.

Chimpanzee interactions with nonhuman species in an anthropogenic habitat

Interactions between wildlife species are numerous and diverse, ranging from commensalism to predation. Information on cross-species interactions in anthropogenic habitats are rare but can serve to improve our understanding of animal behavioural and ecological flexibility in response to human-induced changes. Here we report direct observations of interactions between chimpanzees (Pan troglodytes verus) and wild and domesticated species in a forest-farm mosaic at Bossou, Guinea, recorded between 1997 and 2009. The low diversity and abundance of wildlife, in particular typical chimpanzee prey species, are reflected in both the low interaction rates (one interaction per 400 observation hours) and the low number of species with which chimpanzees interacted (nine species, mostly mammals, but also birds and reptiles). Chimpanzees generally chose either to make direct physical contact with a species or not; interactions that involved direct contact lasted longer than noncontacts. Interactions with mammals showed the greatest diversity in nature and duration. Adults most often consumed a captured animal, while immatures most often engaged in playful behaviours with other species. Immatures also exhibited distinctive accompanying behaviours whereas adults rarely did so. Species-specific behaviours that depend on the age-class of the interactant are consistent with the idea that chimpanzees categorise different animals. We anticipate that chimpanzee interactions with sympatric species inhabiting humanised habitats will change over time to include more domesticated species. Conservation management strategies should anticipate behavioural flexibility in response to changing landscapes.

New Nut-Cracking Sites in Diecké Forest, Guinea: An Overview of the Surveys

INTRODUCTION We report the results of the 2006 KUPRI (Kyoto University Primate Research Institute) surveys in Diecke forest, Guinea, Western Africa. Research in the Diecke forest by KUPRI started in 1999 with a preliminary survey. This continued in 2000, with the survey of Nonah and Yossono areas. The main objective was to find a good research site that would allow for comparative studies between different communities of wild chimpanzees. It has been acknowledged, since the first field season, that Diecke chimpanzees crack open nuts (Panda oleosa and Coula edulis) using stone and root anvils and stone hammers. Nut-cracking behaviour had not been observed directly until today and there are very few reports on chimpanzee’s direct observation . The Diecke forest is part of the Guinean Forests and integrates the region comprised by lowland forests, being one important world spot for the biodiversity. These forests survived after the retraction and division of Pleistocene forests, partly due to the ice age that dried the climate cyclically. This forest, Foret Classee de Diecke, is a protected area of 600 km, located in the South-eastern part of Guinea, 50 km west from Bossou, bordering Liberia on the south. The reserve is controlled by the Centre Forestier de N`zerekore, and Diecke biodiversity is managed by a German project located at the Direction Generale de Centres Forestiers, also in N`zerekore. This project, entitled PROGERFOR, focuses on conservation/ reforestation; damage minimization; regional development, and organized the forest in thirty parcels with three levels of protection. Diecke is characterized by a large number of small hills, averaging 350 m of altitude (the highest point being Mont Jna, with 800 m). Two main rivers cross the forest running south: Nye on the west side, and Gbin, divided in Gbin and Gbin-be, on the east side. Secondary waterlines are abundant in the forest creating swamps. In this part of the country, the dry season is very short, from December or January to March or April. Geologically, Diecke is part of the West African craton with Precambrian rocks, essentially igneous and