Chimpanzees use tools to harvest social insects at Fongoli, Senegal

Use of flexible probes to fish for macrotermitine termites and manufactured wands to dip for doriline ants is reported for a new site of chimpanzee field study. The flexible probes of vegetation used in termite fishing and ant dipping by the chimpanzees of Fongoli, in southeastern Senegal, are similar to those used at Assirik (Senegal) and Gombe (Tanzania). Based on the principle that form reflects function, we predict that ant dipping when seen will prove to be the two-handed technique.     

Insectivory of savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal

Little is known about the behavior of chimpanzees living in savanna-woodlands, although they are of particular interest to anthropologists for the insight they can provide regarding the ecological pressures affecting early hominins living in similar habitats. Fongoli, Senegal, is the first site where savanna chimpanzees have been habituated for observational data collection and is the hottest and driest site where such observation of chimpanzees occurs today. Previously, indirect evidence suggested these chimpanzees consumed termites throughout the year, an unusual occurrence for western and eastern chimpanzees. Although meat eating by chimpanzees continues to receive much attention, their use of invertebrate prey has received less emphasis in scenarios of hominin evolution. Here, we further examine the invertebrate diet of Fongoli chimpanzees using direct observational methods and accounting for potential environmental influences. Termite feeding positively correlated with high temperatures. Fongoli chimpanzees spend more time obtaining termites than any other chimpanzee population studied, and this extensive insectivory contributes to the list of distinctive behaviors they display relative to chimpanzees living in more forested habitats. We suggest that savanna chimpanzees at Fongoli differ significantly from chimpanzees elsewhere as a result of the selective pressures characterizing their harsh environment, and this contrast provides an example of a viable referential model for better understanding human evolution. Specifically, our results support the hypotheses that invertebrate prey may have figured more prominently into the diet of early hominins in similar habitats, especially given that invertebrates are an important source of protein and other essential nutrients in a highly seasonal environment.     

Termite fishing laterality in the fongoli savanna chimpanzees (Pan troglodytes verus): Further evidence of a left hand preference

Whether nonhuman primates show population‐level handedness is a topic of much scientific debate. A previous study of handedness for termite fishing reported population‐level left handedness in the chimpanzees from Gombe National Park, Tanzania. In the current study, we examined whether similar hand preferences were evident in a savanna‐dwelling chimpanzee population with regards to termite fishing. Hand preference data were collected for 27 chimpanzees from February 2007 through July 2008 and November 2011 through January 2012 in southeastern Senegal. Overall, the Fongoli chimpanzees demonstrate a trend toward population‐level handedness, though the results did not reach conventional levels of statistical significance likely due to the limited sample size. Fongoli chimpanzees showed the same pattern of left hand preference as reported at Gombe and the two populations did not differ significantly. When the data were combined across all studies, wild chimpanzees showed a population‐level left hand preference for termite fishing. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Termite diversity along a land use intensification gradient in a semi-arid savanna

Disturbance, particularly agricultural expansion is one of the major threats to the biodiversity and ecological functions of tropical and sub-tropical ecosystems. In this regard, we examined changes in the species richness, abundance, and diversity of termites across different disturbance treatments in a sub-tropical semi-arid savanna in south eastern Zimbabwe. Nine transects (100?×?2 m) representing three habitat disturbance treatments (primary woodland; grazing area; agricultural field) were sampled for termites using a rapid biodiversity assessment protocol. Termites were more abundant and species-rich in primary woodland and grazing area than in the agricultural field. Twelve termite species from three sub-families were present, with Microtermes sp. constituting 35% of the identified termite species. Termite feeding group structure differed significantly among land-use types, and of all termites present, wood-feeding termites were the most abundant while soil-feeders were rare in the agricultural field. In conclusion the observed pattern in termite species richness and relative abundance indicates that termites are very resilient to natural disturbance and might actually benefit from some natural disturbances like they did in the grazing area of this study, but they are not resilient to extreme anthropogenic disturbance. Although there was no notable difference in termite species richness and relative abundance between agricultural field and primary woodland, the pattern observed across the three sites may be potential support for the IDH suggesting that intermediate levels of physical disturbance intensity influence the structure and functioning of termite assemblages in semi-arid savanna.     

Chimpanzees' choices of prey between two sympatric species ofMacrotermes in the Campo Animal Reserve,Cameroon

Details are presented relating to chimpanzees' choices between two sympatric species of termites,Macrotermes lillijeborgi andM. vitrialatus, as food in the Campo Animal Reserve, southwest Cameroon, West Africa. An attempt was made to determine the various factors that affected such choices. The two species of termites seemed to have almost the same value in terms of ecological factors. However, chimpanzees fed almost exclusivelyonM. lillijeborgi, using digging sticks and fisching probes, during the study period which extended from the end of August to the middle of January, with their feeding activity showing peak at the beginning of the rainy season. By contrast,M. vitrialatus was rarely eaten in spite of the ease with which such prey could be obtained, namely, by desctruction of termite mounds by hand, without the need for tools. The reason that the chimpanzees discriminated between the two species of termite cannot be explained in terms of ecological factors such as size of prey, seasonal differences in termite activity, etc. Sticks used as tools were fairly uniform in size and character, mainly because of physical constraints related to the structure of termite mounds, and the brush-like ends of sticks seemed to be incidentaly byproducts of the chimpanzee's choice of plant species. Ecological factors could provide chimpanzees with a basis for the use of some kind of tool and help them modify it, while other factors, for example, something akin to leisure or the chimpanzee's interest in use of a tool, could provide an opportunity for inventing some tool-using behavior or for maintaining such behavior. These different factors, not being exclusive of one another, might affect the invention and maintenance of tool using-behavior at different phases. It is possible that chimpanzee's choice of prey may not always be the most efficient or appropriate in a given ecological situation.     

Species abundance and habitat differences in biomass of subterranean termites (Isoptera) in the wheatbelt of Western Australia

The distribution and abundance, and habitat differences in biomass of subterranean termites, were assessed through soil trenching for woodland, mallee and heath habitats in the central wheatbelt of Western Australia. Over an 11 month sampling period, there were no significant habitat differences in biomass. Mean dry biomass in the surficial layer of soil (per 5000 cm³) averaged 46 mg in woodland, 28 mg in mallee, and 23 mg in heath. Termite biomass peaked in September, with 141 mg for woodland, 83 mg for mallee and 47 mg for heath (per 5000 cm³). Soil moisture and termite activity near the surface were positively correlated. A total of 36 species of termites, comprising 11 genera, were identified, and species abundance within and across habitats differed significantly.     

Short‐term changes in the structure of termite assemblages associated with slash‐and‐burn agriculture in Côte d'Ivoire

Termites are major decomposers in tropical regions and play critical roles in many soil‐related processes. Studies conducted in Asia and the Neotropics suggest that habitat modification can strongly affect termite assemblages, but data on termite communities from forests in Africa, especially West Africa, are scarce. Here, we measured the short‐term impact of slash‐and‐burn agriculture on termite assemblages in an agricultural region of central Côte d'Ivoire. We assessed termite diversity and relative abundance in four habitat types: secondary forest, cleared forest, burned forest, and crop fields. The secondary forest had higher species richness compared with the other habitats, but all habitat types had similar assemblage structures. Fungus‐growing termites were the most abundant feeding group in all habitats. Soil feeders were most abundant in secondary forest, intermediately abundant in cleared and burned forests, and almost entirely absent in crop fields. Wood‐feeding species showed clear responses to burning; their abundances decreased after fire. We conclude that slash‐and‐burn agriculture does not appear to severely erode the diversity of termite assemblages. This could be due to the dominance of ecologically versatile fungus growers or to the relatively long time between clearing and burning. However, forest clearing negatively affects soil feeders, with the Apicotermitinae most affected by canopy loss.     

Arboreal nesting as anti-predator adaptation by savanna chimpanzees (Pan troglodytes verus) in southeastern Senegal

Chimpanzees (Pan troglodytes) make nests for resting and sleeping, which is unusual for anthropoid primates but common to all great apes. Arboreal nesting has been linked to predation pressure, but few studies have tested the adaptive nature of this behavior. We collected data at two chimpanzee study sites in southeastern Senegal that differed in predator presence to test the hypothesis that elevated sleeping platforms are adaptations for predator defense. At Assirik in the Parc National du Niokolo-Koba, chimpanzees face four species of large carnivore, whereas at Fongoli, outside national park boundaries, humans have exterminated almost all natural predators. We quantified the availability of vegetation at the two sites to test the alternative hypothesis that differences in nesting reflect differences in habitat structure. We also examined possible sex differences in nesting behavior, community demographic differences, seasonality and nest age differences as variables also potentially affecting nest characteristics and nesting behavior between the two sites. Chimpanzees at Fongoli nested at lower heights and farther apart than did chimpanzees at Assirik and sometimes made nests on the ground. The absence of predators outside of the national park may account for the differences in nest characteristics at the two sites, given the similarities in habitat structure between Fongoli and Assirik. However, Fongoli chimpanzees regularly build arboreal nests for sleeping, even under minimal predation pressure, and this requires explanation.     

Temperate forest termites: ecology,biogeography, and ecosystem impacts

1. Wood decomposition in temperate forests is dominated by termites, fungi, and some species of ants and beetles. Outside of urban areas, temperate termite ecology is largely unknown, particularly when compared to tropical termites and other temperate organisms in the functional guild of wood‐decomposing animals. 2. This review combines climate habitat modelling with knowledge of species physiology, behaviour, and community interactions to identify and prioritise future research on temperate termite ecology and biogeography. 3. Using a correlative climate model, the regional distributions of three common temperate forest termite species are shown to correlate with different aspects of climate (e.g. mean versus minimum monthly temperature), but that overall their distributions within temperate systems correlate more strongly with temperature variables than with precipitation variables. 4. Existing data are synthesised to outline how the subterranean, wood‐nesting behaviour of most temperate forest termite species links their activity to an additional set of non‐climate controls: wood type and tree species, soil depth, fungal activity, ant abundances and phenology, and competitive asymmetries among termite species. 5. Although fine‐scale estimates of temperate termite abundances are rare, we provide upper bounds on their ecosystem impacts and illustrate how their regional abundances may influence forest structure and habitat availability for other organisms. 6. This review highlights that rigorous ecological studies in non‐urban, intact ecosystems – with a particular focus on community interactions – are critically needed to accurately project future abundances, economic impacts, and ecosystem effects of temperate forest termites.     

Fungus-growing termites originated in African rain forest

Fungus-growing termites (subfamily Macrotermitinae, Isoptera) cultivate fungal crops (genus Termitomyces, Basidiomycotina) in gardens inside their colonies. Those fungus gardens are continuously provided with plant substrates, whereas older parts that have been well decomposed by the fungus are consumed (cf.). Fungus-growing termites are found throughout the Old World tropics, in rain forests and savannas, but are ecologically dominant in savannas. Here, we reconstruct the ancestral habitat and geographical origin of fungus-growing termites. We used a statistical model of habitat switching repeated over all phylogenetic trees sampled in a Bayesian analysis of molecular data. Our reconstructions provide strong evidence that termite agriculture originated in African rain forest and that the main radiation leading to the extant genera occurred there. Because extant savanna species are found in most genera, this moreover suggests that the savanna has repeatedly been colonized by fungus-growing termites. Furthermore, at least four independent "out-of-Africa" migrations into Asia, and at least one independent migration to Madagascar, have occurred. Although fungus growing by termites is ecologically most successful under the variable, unfavorable conditions of the savanna, it seems to have evolved under the more constant and favorable conditions of the rain forest.     

Dexterity and technique in termite fishing by chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle,Republic of Congo

Although the phenomenon of termite fishing by chimpanzees (Pan troglodytes) has historical and theoretical importance for primatology, we still have a limited understanding of how chimpanzees accomplish this activity, and in particular, about details of skilled actions and the nature of individual variation in fishing techniques. We examined movements, hand positions, grips, and other details from remote video footage of seven adult and subadult female chimpanzees using plant probes to extract Macrotermes muelleri termites from epigeal nests. Six chimpanzees used exclusively one hand (left or right) to grip the probe during termite fishing. All chimpanzees used the same repertoire of actions to insert, adjust, and withdraw the probe but differed in the frequency of use of particular actions. Chimpanzees have been described as eating termites in two ways—directly from the probe or by sweeping them from the probe with one hand. We describe a third technique: sliding the probe between the digits of one stationary hand as the probe is extracted from the nest. The sliding technique requires complementary bimanual coordination (extracting with one hand and grasping lightly with the other, at the same time). We highlight the importance of actions with two hands—one gripping, one assisting—in termite fishing and discuss how probing techniques are correlated with performance. Additional research on digital function and on environmental, organismic, and task constraints will further reveal manual dexterity in termite fishing.     

Subsistence strategies of two "savanna" chimpanzee populations: the stable isotope evidence.

Twenty-two chimpanzee hair samples collected from night nests at two different "savanna" sites were analyzed for carbon and nitrogen stable isotope ratios represented as delta13C and delta15N values. The first at Ugalla, Tanzania is a miombo woodland with grass groundcover and small patches of forest. The second at Ishasha, Democratic Republic of the Congo is a habitat composed of riverine gallery forest, semideciduous thicket forest, wooded grassland, and grassland. Based on comparative data from other primates, Ugalla hair delta13C values suggest that the chimpanzees are feeding primarily in the woodland rather than in forest patches or on grassland foods (grasses or grammivorous fauna). Similar comparisons indicate that the Ishasha chimpanzees are feeding within the forests and not in more open areas. In addition, the Ugalla chimpanzees had delta15N values that indicate extensive ingestion of leguminous flowers, seeds, and/or leaves. The Ishasha samples show a range encompassing three trophic levels. Two samples with the most positive values may indicate a nursing signal or vertebrate-feeding. Three individuals with intermediate values are similar to those in omnivorous nonhuman primate species. The four individuals with the lowest values are very similar to those in herbivorous monkeys. Stable isotope ratios permit time-averaged and habitat-specific dietary comparisons among sites, even without habituation or detailed foraging observations.     

Archaeological Analysis Does Not Support Intentionality in the Production of Brushed Ends on Chimpanzee Termiting Tools

Some chimpanzees use 2 types of tools to extract underground termites for consumption. Chimpanzees insert thin, flexible probes into tunnels or holes in termite mounds (fishing), and sometimes use stouter, rigid sticks to first puncture the holes and also possibly to fish. Many puncturing sticks have distinctive “brushed” ends. Researchers have hypothesized that chimpanzees create the brushed ends intentionally to increase their affixibility to biting termites (Sugiyama, 1985). The results of our archaeological analysis of a large collection of puncturing sticks used by Central African chimpanzees falsifies this hypothesis, and instead agrees with the recent behavioral observations of Sanz et al. (2004; cf. Bermejo and Illera, 1999) that brushing is a coincidental result of procuring sticks from vegetation sources. The results highlight the positive contribution of an archaeological approach to problems in chimpanzee material culture and emphasize to primatologists the value of curating artifacts.     

Survey of subterranean termites (Isoptera: Rhinotermitidae) in a managed silvicultural plantation in Portugal, using a line-intersection method (LIS)

Subterranean termites (Reticulitermes grassei) were surveyed over successive seasons in a managed eucalyptus plantation in southeastern Portugal for 26 months. Termite activity in seven diameter categories of lying dead wood was investigated by a modified line intersection method (LIS). Each item sampled was inspected and assessed for termite attack and for general (i.e. fungal) decay status using standard protocols. Line intersection is quantitative to the extent that it can link foraging and decay parameters to woody biovolume. It was found that termites selected items with larger diameter, the observed trend showing an exponential character with greater termite attack as diameter increased. Attack by termites was positively associated with prior decay by fungi. A clear positive relationship was shown between rainfall and total woody biovolume containing live termites, underlining the importance of moisture for termite activity. Subterranean termites appeared to be important wood decomposers in the woodland studied, with an average of 30% of lying dead wood branches showing signs of termite attack.     

Infestation of African savanna ecosystems by subterranean termites

This review is an analysis of the dimensions of termite infestation in African savannas. The aim of this work is to draw the attention of ecologists, conservationists, policy makers and farmers to the current and future threats of subterranean termites to the functioning and sustainability of such ecosystem habitats. This study analyzes and describes termite problem (questionable changes in density and assemblage structure) in selected African savannah ecosystems, synthesizes information on the effects of various human induced habitat disturbance regimes on termites’ assemblage structure, predators, nests and feed resources to generate hypotheses relating termite infestation with anthropogenic activities; it describes and critiques existing termite management practices. The review is suggestive that the infestation and resultant undesirable effects of subterranean termites in African savannahs are largely a consequence of the inappropriate savannah management practices (overgrazing, indiscriminate tree cutting and overhunting) undertaken by humans in pursuit of various livelihood options. Based on the evidence presented herein, we hypothesized that (1) human induced habitat disturbance in savanna ecosystems alters the feeding group composition of termites’ assemblages, favoring grass harvesters and polyphagus termite feeders that forage on more abundant food items, paying little attention to rarer food items and (2) habitat disturbance through activities like heavy grazing and overhunting results in decline in the populations of both macro and microscopic termite predators, which eventually enhances the proliferation of termite populations, escalates the density of termite nests particularly epigeal mounds and intensifies consumption of herbaceous savannah vegetation. The review calls for dedicated efforts to develop ecological thresholds of savannah biotic and abiotic ecosystem components in which human induced disturbance regimes trigger the destructive behavior of termites. This would provide information that will act as a precautionary savannah habitat monitoring and decision support tool to prevent future infestation of savannah habitats with termites. Also, the review shows that majority of the termite control practices are ineffective, ecologically unsustainable and above all, do not address the root cause of termite infestation and thus merely provide temporary relief to the problem. As such, termite control methods that attempt to enhance proliferation of termite predators need to be studied, developed and emphasized. This review reveals that human induced habitat disturbance depletes termites’ predator populations, leading to proliferation of termite populations particularly grass harvesters that intensify their consumption on grass biomass and eventually contribute to denudation of herbaceous vegetation cover in savannah ecosystems.     

Ant colonization and coarse woody debris decomposition in temperate forests

Ants are ubiquitous, abundant and have widespread impacts on ecological communities and ecosystem processes. However, ant effects on coarse woody debris decomposition are unexplored. Several ant species colonize coarse woody debris for nesting, and this puts them in contact with fauna and microbes that utilize coarse woody debris as habitat and food, potentially influencing nutrient cycling and, ultimately, forest productivity. We report results from a field experiment employing 138 artificial ant nests (routed pine blocks) across five locations in southeastern US deciduous forests. We examine the correspondence between ant, termite and wood-eating fungi colonization and variation in coarse woody debris decomposition. After 1 year, nests colonized by ants had 5% more mass than those not colonized. Ant colonization corresponded with significantly less termite- and fungal-mediated decomposition of the nests. Without ants, termites removed 11.5% and fungi removed 4% more wood biomass. Ants, termites and wood-eating fungi all colonized pine nests where temperatures were highest, and ants also preferred higher soil moisture whereas termites and fungi responded negatively to high soil moisture when temperatures were higher. Ants reduce termite colonies through predation, and may inhibit fungi through the secretion of antimicrobial compounds. Our results indicate that interactions between forest understory ants, termites and fungi may influence the rate of coarse woody debris decomposition—biotic interactions that potentially influence forest structure and function.     

The spatial pattern of soil-dwelling termites in primary and logged forest in Sabah, Malaysia

Abstract.  1. Primary and logged lowland dipterocarp forest sites were sampled for subterranean termites using soil pits located on a grid system in order to detect any patchiness in their distribution.
2. A spatial pattern in termite distributions was observed in the primary and logged sites, but the response differed between soil-feeding and non-soil-feeding termites.
3. Spatial analysis showed that soil-feeding termites were homogeneously distributed in the primary forest but significantly aggregated in the logged forest. This pattern was reversed for non-soil-feeding termites and may result from differences in resource provisioning between the two sites.
4. Gaps in termite distribution comprised a greater area than patches for both feeding groups and sites, but gaps dominated the logged site.
5. A significant association between soil-feeding and non-soil-feeding termite distributions occurred at both sites. This arose from an association between patches in the primary forest and between gaps in the logged forest.
6. Termite spatial pattern was optimally observed at a minimum extent of 64 m and lag of 2 m.
7. The spatially explicit SADIE (Spatial Analysis by Distances IndicEs) analyses were more successful than (non-spatially explicit) multivariate analysis (Canonical Correspondence Analysis) at detecting associations between termite spatial distributions and that of other biotic and abiotic variables.     

Nigerian chimpanzees (Pan troglodytes vellerosus) at Gashaka: two years of habituation efforts

Cross-population comparisons of chimpanzees can shed light on the pathways of hominid evolution. So far, no eco-ethological data exist for the recently recognized subspecies Pan troglodytes vellerosus. We report on the first 2 years of a new long-term study from what is perhaps their last remaining stronghold: the Gashaka Gumti National Park, Nigeria. The mosaic habitat (woodland, lowland and gallery forest) receives 1,826 mm rain/year, with 4-5 months being completely dry. Primates at Gashaka are not hunted, and the chimpanzees are therefore relatively tolerant of human observers. We focused on the Gashaka-Kwano community, investing 3,000 h of patrols. A total of 95 sightings were achieved which lasted for an average of 27 min (range 1-190 min). Party size averaged 3.7 animals (range 1-17) but was, similarly to encounter length, susceptible to a wide range of methodological, social and ecological factors. The Kwano community comprises at least 35 members which occupy a home range of at least 26 km(2), yielding a density of 1.3/km(2). The area represents the West African equivalent of a chimpanzee site similar to the forest-woodland habitat in which early humans might have evolved.     

Hand Preference for a Novel Bimanual Coordinated Task During Termite Feeding in Wild Western Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Although the level of handedness in humans varies cross-culturally, humans are generally described as right-handed, which has been considered a uniquely human trait. Recently, captive chimpanzees (Pan troglodytes) have been shown to exhibit right-hand preference when performing bimanual but not unimanual tasks. Less clear is whether this pattern also occurs in wild chimpanzees and other African apes. Using videos (N = 49) of six wild western gorillas (Gorilla gorilla gorilla) feeding on termites at the Mondika Research Center (Republic of Congo), we tested whether they exhibit hand preference when performing unimanual, i.e., reaching for termite mound pieces; bimanual, i.e., “termite tapping”: rhythmically shaking a piece of termite mound with the dominant hand and collecting the termites in the other hand tasks; or hand transfer prior to bimanual tasks, i.e., transferring a piece of termite mound from one hand to the other. All individuals exhibited exclusive hand preference when performing the bimanual tasks, with five of six gorillas preferring the right hand. Conversely, most individuals did not show any manual preference during the unimanual task. In addition, hand preference during hand transfer revealed clear hand dominance of similar strength and direction of those shown for the bimanual task, suggesting that this measure is as sensitive as the bimanual task itself. Thus, we propose “termite feeding” as a novel task to be considered in future hand-preference studies in wild western gorillas. Our results are in concordance with those for chimpanzees and captive gorillas showing hemispheric specialization for bimanual actions in apes.