Alejandra Pascual-Garrido

Obtaining Raw Material: Plants as Tool Sources for Nigerian Chimpanzees

We investigated the acquisition of plant materials from which Nigerian chimpanzees manufacture wooden tools to harvest insects and honey from nests of army ants, honey bees and stingless bees. Slender trunks of juvenile trees and branches are most commonly used, and bendable vines rarely, probably reflecting the need to work with relatively sturdy tools to extract resources. While several tools are sometimes sourced from the same plant, there is also evidence for a depletion effect, as multiple tool sources at the same site are often spaced several metres apart. Identified tool sources belong to 27 species of at least 13 families. Honey-gathering implements are often chewed upon by chimpanzees. Interestingly, twigs of the most commonly used honey-gathering species possess antibacterial propensities and are favoured by Nigerians as chewing sticks. This suggests that extractive tools might possess associated medicinal or stimulatory properties. We do not know if chimpanzees actively select specific plant parts or species as we cannot compare observed with expected frequencies. Nevertheless, about three quarters of tools are picked from plants more than 6 m away from the extraction site, potentially indicating some degree of forward planning.

Archaeological excavation of wild macaque stone tools.

More than 3 million years of excavated archaeological evidence (Harmand et al., 2015) underlies most major insights into the evolution of human behaviour. However, we have seen almost no use of archaeological excavation to similarly broaden our understanding of behaviour in other animal lineages. The few published examples include recovery of a late Holocene assemblage of stones from the Ivory Coast, attributed to the agency of both humans (Homo sapiens) and chimpanzees (Pan troglodytes verus) (Mercader et al., 2002, 2007), and exploration of the occupation sites of nontool-using species such as penguins (Emslie et al., 2014) and other birds (Burnham et al., 2009). The development of viable methods for identifying and interpreting past non-human tool use landscapes is essential if we are to gain a better understanding of technological evolution within other animals, including our close relatives, the primates. Recently, the growth of primate archaeology has built on the close phylogenetic relationship between humans and other primates to begin filling in this lacuna (Haslam et al., 2009; Carvalho, 2011; Stewart et al., 2011; Haslam, 2012, 2014; Visalberghi et al., 2013; Haslam et al., 2014; McGrew et al., 2014; Benito-Calvo et al., 2015; Luncz et al., 2015; Kühl et al., 2016). Here, we present the first report on an archaeologically excavated Old World monkey tool use site, which was created by wild Burmese long-tailed macaques (Macaca fascicularis aurea) (Bunlungsup et al., 2016) during shellfish-processing activities in coastal Thailand. These macaques use stone and shell pounding

Wild capuchin monkeys adjust stone tools according to changing nut properties.

Animals foraging in their natural environments need to be proficient at recognizing and responding to changes in food targets that affect accessibility or pose a risk. Wild bearded capuchin monkeys (Sapajus libidinosus) use stone tools to access a variety of nut species, including otherwise inaccessible foods. This study tests whether wild capuchins from Serra da Capivara National Park in Brazil adjust their tool selection when processing cashew (Anacardium spp.) nuts. During the ripening process of cashew nuts, the amount of caustic defensive substance in the nut mesocarp decreases. We conducted field experiments to test whether capuchins adapt their stone hammer selection to changing properties of the target nut, using stones of different weights and two maturation stages of cashew nuts. The results show that although fresh nuts are easier to crack, capuchin monkeys used larger stone tools to open them, which may help the monkeys avoid contact with the caustic hazard in fresh nuts. We demonstrate that capuchin monkeys are actively able to distinguish between the maturation stages within one nut species, and to adapt their foraging behaviour accordingly.

Primate archaeology evolves

Since its inception, archaeology has traditionally focused exclusively on humans and our direct ancestors. However, recent years have seen archaeological techniques applied to material evidence left behind by non-human animals. Here, we review advances made by the most prominent field investigating past non-human tool use: primate archaeology. This field combines survey of wild primate activity areas with ethological observations, excavations and analyses that allow the reconstruction of past primate behaviour. Because the order Primates includes humans, new insights into the behavioural evolution of apes and monkeys also can be used to better interrogate the record of early tool use in our own, hominin, lineage. This work has recently doubled the set of primate lineages with an excavated archaeological record, adding Old World macaques and New World capuchin monkeys to chimpanzees and humans, and it has shown that tool selection and transport, and discrete site formation, are universal among wild stone-tool-using primates. It has also revealed that wild capuchins regularly break stone tools in a way that can make them difficult to distinguish from simple early hominin tools. Ultimately, this research opens up opportunities for the development of a broader animal archaeology, marking the end of archaeology’s anthropocentric era.Nearly ten years after the field of primate archaeology was first proposed, the status of the field is reported on, including recent discoveries as well as future directions and challenges, marking the end of archaeology’s ‘anthropocentric era’.

White-gutted soldiers: simplification of the digestive tube for a non-particulate diet in higher Old World termites (Isoptera: Termitidae)

Previous observations have noted that in some species of higher termites the soldier caste lacks pigmented particles in its gut and, instead, is fed worker saliva that imparts a whitish coloration to the abdomen. In order to investigate the occurrence of this trait more thoroughly, we surveyed a broad diversity of termite specimens and taxonomic descriptions from the Old World subfamilies Apicotermitinae, Cubitermitinae, Foraminitermitinae, Macrotermitinae, and Termitinae. We identified 38 genera that have this “white-gutted” soldier (WGS) trait. No termite soldiers from the New World were found to possess a WGS caste. Externally, the WGS is characterized by a uniformly pale abdomen, hyaline gut, and proportionally smaller body-to-head volume ratio compared with their “dark-gutted” soldier (DGS) counterparts found in most termitid genera. The WGS is a fully formed soldier that, unlike soldiers in other higher termite taxa, has a small, narrow, and decompartmentalized digestive tube that lacks particulate food contents. The presumed saliva-nourished WGS have various forms of simplified gut morphologies that have evolved at least six times within the higher termites.

Raw material procurement for termite fishing tools by wild chimpanzees in the Issa valley, Western Tanzania

OBJECTIVES Chimpanzee termite fishing has been studied for decades, yet the selective processes preceding the manufacture of fishing tools remain largely unexplored. We investigate raw material selection and potential evidence of forward planning in the chimpanzees of Issa valley, western Tanzania. MATERIALS AND METHODS Using traditional archaeological methods, we surveyed the location of plants from where chimpanzees sourced raw material to manufacture termite fishing tools, relative to targeted mounds. We measured raw material abundance to test for availability and selection. Statistics included Chi-Squared, two-tailed Wilcoxon, and Kruskall-Wallace tests. RESULTS Issa chimpanzees manufactured extraction tools only from bark, despite availability of other suitable materials (e.g., twigs), and selected particular plant species as raw material sources, which they often also exploit for food. Most plants were sourced 1-16 m away from the mound, with a maximum of 33 m. The line of sight from the targeted mound was obscured for a quarter of these plants. DISCUSSION The exclusive use of bark tools despite availability of other suitable materials indicates a possible cultural preference. The fact that Issa chimpanzees select specific plant species and travel some distance to source them suggests some degree of selectivity and, potentially, forward planning. Our results have implications for the reconstruction of early hominin behaviors, particularly with regard to the use of perishable tools, which remain archaeologically invisible.

Scars on plants sourced for termite fishing tools by chimpanzees: Towards an archaeology of the perishable

Chimpanzees are well‐studied, but raw material acquisition for tool use is still poorly understood as sources are difficult to trace. This study pioneers the use of information that can be gleaned from plant scars made by chimpanzees while they source vegetation parts to manufacture termite fishing tools. Source plant species, raw material types and locations relative to targeted termite mounds were recorded for populations at Gombe, Issa, and Mahale in western Tanzania. Recovered bark, twig, and vine tools were traced to 29 plant species, while grass sources were indeterminable. Bark extraction scars remained detectable for months, and thus possibly for as long as the plant is alive, while twig and vine scars preserved for a few weeks only. Scars preserve better than tools, given that twice as many plant species could be linked to the former than to the latter. Some source species were exploited across all sites for the same type of tool material, while two species were sourced for different types. Compared to apes at Gombe and Mahale, Issa chimpanzees carried material from twice as far away, perhaps because the Issa habitat is more open and dry, which entails greater distances between suitable raw material sources and targeted mounds. Site‐specific tools were based on different raw materials, in two cases sourced from the same species, which could suggest learned preferences for particular tool material. “Archaeology of the perishable” as pioneered in this study broadens the methodological approach of the wider field of primate archaeology to include reconstructions of past animal behavior associated with the production of plant based tools.