Promoting inclusivity in ecology, evolution, and behavioral biology education through course-based undergraduate research experiences.

Access to independent research experiences is a persistent barrier that stifles the recruitment and retention of students from diverse backgrounds in ecology, evolution, and behavioral biology. The benefits of field experiences are not equitably available to historically excluded and minoritized students. In this article, we summarize evidence that indicates course-based undergraduate research experiences (CUREs) provide a solution to ensure equitable access to independent research experiences in the life sciences. We draw from our own experiences of teaching CUREs in ecology, evolution, and behavioral biology and provide the complete curriculum for our effective and largely materials-free CURE in behavioral ecology (CURE-BxEco). We advocate for greater access to and synthesize the benefits of CUREs to promote inclusivity in education. The proliferation of such innovative pedagogical practices benefits science because these classroom methods are critical in recruiting and retaining historically excluded and minoritized students, who offer diverse perspectives in research.

Socioecological factors influencing intraspecific variation in ranging dynamics of western lowland gorillas (Gorilla gorilla gorilla) in Ndoki Forest.

Ranging dynamics are physical and behavioral representations of how different socioecological factors affect an organism's spatial decisions and space use strategies. Western lowland gorillas (Gorilla gorilla gorilla) are a model species to investigate the drivers of spatial dynamics based on both the natural variation in socioecological factors within the species and compared with their mountain gorilla counterparts. In this study, we evaluate the influences of resource seasonality and social dynamics on variation in home range size, utilization, and intergroup overlap among multiple gorilla groups over an 8-year study period in the northern Republic of Congo. This study shows that western lowland gorillas can have small home ranges comparable to mountain gorillas, rather than universally larger home ranges as previously supposed, and that home ranges are stable through time. The largest source of variation in space use was the degree of intergroup home range overlap. The study groups did not demonstrate intraspecific variation in range size nor changes in intergroup overlap with respect to seasonality of fruit resources, but all groups demonstrated expansion of monthly range and core area with group size, matching predictions of intragroup feeding competition. These findings highlight the potential impact of intergroup relationships on space use and prompt further research on the role of social dynamics in ranging strategies. In this study, we reveal a greater degree of variability and flexibility in gorilla ranging behavior than previously realized which is relevant to improving comparative studies and informing conservation strategies on behalf of these endangered primates.

Teaching varies with task complexity in wild chimpanzees.

Cumulative culture is a transformative force in human evolution, but the social underpinnings of this capacity are debated. Identifying social influences on how chimpanzees acquire tool tasks of differing complexity may help illuminate the evolutionary origins of technology in our own lineage. Humans routinely transfer tools to novices to scaffold their skill development. While tool transfers occur in wild chimpanzees and fulfill criteria for teaching, it is unknown whether this form of helping varies between populations and across tasks. Applying standardized methods, we compared tool transfers during termite gathering by chimpanzees in the Goualougo Triangle, Republic of Congo, and in Gombe, Tanzania. At Goualougo, chimpanzees use multiple, different tool types sequentially, choose specific raw materials, and perform modifications that improve tool efficiency, which could make it challenging for novices to manufacture suitable tools. Termite gathering at Gombe involves a single tool type, fishing probes, which can be manufactured from various materials. Multiple measures indicated population differences in tool-transfer behavior. The rate of transfers and probability of transfer upon request were significantly higher at Goualougo, while resistance to transfers was significantly higher at Gombe. Active transfers of tools in which possessors moved to facilitate possession change upon request occurred only at Goualougo, where they were the most common transfer type. At Gombe, tool requests were typically refused. We suggest that these population differences in tool-transfer behavior may relate to task complexity and that active helping plays an enhanced role in the cultural transmission of complex technology in wild apes.

The complex Y-chromosomal history of gorillas.

Studies of the evolutionary relationships among gorilla populations using autosomal and mitochondrial sequences suggest that male-mediated gene flow may have been important in the past, but data on the Y-chromosomal relationships among the gorilla subspecies are limited. Here, we genotyped blood and noninvasively collected fecal samples from 12 captives and 257 wild male gorillas of known origin representing all four subspecies (Gorilla gorilla gorilla, G. g. diehli, G. beringei beringei, and G. b. graueri) at 10 Y-linked microsatellite loci resulting in 102 unique Y-haplotypes for 224 individuals. We found that western lowland gorilla (G. g. gorilla) haplotypes were consistently more diverse than any other subspecies for all measures of diversity and comprised several genetically distinct groups. However, these did not correspond to geographical proximity and some closely related haplotypes were found several hundred kilometers apart. Similarly, our broad sampling of eastern gorillas revealed that mountain (G. b. beringei) and Grauer's (G. b. graueri) gorilla Y-chromosomal haplotypes did not form distinct clusters. These observations suggest structure in the ancestral population with subsequent mixing of differentiated haplotypes by male dispersal for western lowland gorillas, and postisolation migration or incomplete lineage sorting due to short divergence times for eastern gorillas.

CD4 receptor diversity in chimpanzees protects against SIV infection.

Human and simian immunodeficiency viruses (HIV/SIVs) use CD4 as the primary receptor to enter target cells. Here, we show that the chimpanzee CD4 is highly polymorphic, with nine coding variants present in wild populations, and that this diversity interferes with SIV envelope (Env)-CD4 interactions. Testing the replication fitness of SIVcpz strains in CD4+ T cells from captive chimpanzees, we found that certain viruses were unable to infect cells from certain hosts. These differences were recapitulated in CD4 transfection assays, which revealed a strong association between CD4 genotypes and SIVcpz infection phenotypes. The most striking differences were observed for three substitutions (Q25R, Q40R, and P68T), with P68T generating a second N-linked glycosylation site (N66) in addition to an invariant N32 encoded by all chimpanzee CD4 alleles. In silico modeling and site-directed mutagenesis identified charged residues at the CD4-Env interface and clashes between CD4- and Env-encoded glycans as mechanisms of inhibition. CD4 polymorphisms also reduced Env-mediated cell entry of monkey SIVs, which was dependent on at least one D1 domain glycan. CD4 allele frequencies varied among wild chimpanzees, with high diversity in all but the western subspecies, which appeared to have undergone a selective sweep. One allele was associated with lower SIVcpz prevalence rates in the wild. These results indicate that substitutions in the D1 domain of the chimpanzee CD4 can prevent SIV cell entry. Although some SIVcpz strains have adapted to utilize these variants, CD4 diversity is maintained, protecting chimpanzees against infection with SIVcpz and other SIVs to which they are exposed.

The ontogeny of termite gathering among chimpanzees in the Goualougo Triangle, Republic of Congo.

OBJECTIVES: Acquiring tool-assisted foraging skills can potentially improve dietary quality and increase fitness for wild chimpanzees (Pan troglodytes). In contrast to chimpanzees in East and West Africa, chimpanzees in the Congo Basin use tool sets and brush-tipped fishing probes to gather termites. We investigated the ontogeny of these tool skills in chimpanzees of the Goualougo Triangle, Republic of Congo, and compared it to that for chimpanzees at Gombe, Tanzania. We assessed whether chimpanzees acquired simple tool behaviors and single tool use before more complex actions and sequential use of multiple tool types. MATERIALS AND METHODS: Using a longitudinal approach, we scored remote video footage to document the acquisition of termite-gathering critical elements for 25 immature chimpanzees at Goualougo. RESULTS: All chimpanzees termite fished by 2.9 years but did not manufacture brush-tipped probes until an average of 4.3 years. Acquisition of sequential tool use extended into juvenility and adolescence. While we did not detect significant sex differences, most critical elements except tool manufacture were acquired slightly earlier by females. DISCUSSION: These findings contrast with Gombe, where chimpanzees learn to both use and make fishing probes between ages 1.5-3.5 and acquire the complete task by age 5.5. Differences between sites could reflect tool material selectivity and design complexity, the challenge of sequential tool behaviors, and strength requirements of puncturing subterranean termite nests at Goualougo. These results illustrate how task complexity may influence the timing and sequence of skill acquisition, improving models of the ontogeny of tool behavior among early hominins who likely used complex, perishable technologies.

Fermented food consumption in wild nonhuman primates and its ecological drivers.

OBJECTIVES: Although fermented food use is ubiquitous in humans, the ecological and evolutionary factors contributing to its emergence are unclear. Here we investigated the ecological contexts surrounding the consumption of fruits in the late stages of fermentation by wild primates to provide insight into its adaptive function. We hypothesized that climate, socioecological traits, and habitat patch size would influence the occurrence of this behavior due to effects on the environmental prevalence of late-stage fermented foods, the ability of primates to detect them, and potential nutritional benefits. MATERIALS AND METHODS: We compiled data from field studies lasting at least 9 months to describe the contexts in which primates were observed consuming fruits in the late stages of fermentation. Using generalized linear mixed-effects models, we assessed the effects of 18 predictor variables on the occurrence of fermented food use in primates. RESULTS: Late-stage fermented foods were consumed by a wide taxonomic breadth of primates. However, they generally made up 0.01%-3% of the annual diet and were limited to a subset of fruit species, many of which are reported to have mechanical and chemical defenses against herbivores when not fermented. Additionally, late-stage fermented food consumption was best predicted by climate and habitat patch size. It was more likely to occur in larger habitat patches with lower annual mean rainfall and higher annual mean maximum temperatures. DISCUSSION: We posit that primates capitalize on the natural fermentation of some fruits as part of a nutritional strategy to maximize periods of fruit exploitation and/or access a wider range of plant species. We speculate that these factors contributed to the evolutionary emergence of the human propensity for fermented foods.

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.

Lethal aggression in Pan is better explained by adaptive strategies than human impacts.

Observations of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) provide valuable comparative data for understanding the significance of conspecific killing. Two kinds of hypothesis have been proposed. Lethal violence is sometimes concluded to be the result of adaptive strategies, such that killers ultimately gain fitness benefits by increasing their access to resources such as food or mates. Alternatively, it could be a non-adaptive result of human impacts, such as habitat change or food provisioning. To discriminate between these hypotheses we compiled information from 18 chimpanzee communities and 4 bonobo communities studied over five decades. Our data include 152 killings (n = 58 observed, 41 inferred, and 53 suspected killings) by chimpanzees in 15 communities and one suspected killing by bonobos. We found that males were the most frequent attackers (92% of participants) and victims (73%); most killings (66%) involved intercommunity attacks; and attackers greatly outnumbered their victims (median 8:1 ratio). Variation in killing rates was unrelated to measures of human impacts. Our results are compatible with previously proposed adaptive explanations for killing by chimpanzees, whereas the human impact hypothesis is not supported.

Behavioral changes following alterations in the composition of a captive bachelor group of western lowland gorillas (Gorilla gorilla gorilla).

Western lowland gorillas (Gorilla gorilla gorilla) in zoos are housed in family or bachelor groups to maximize social opportunities. While wild bachelor groups are transient, all-male groups in zoos may be maintained for many years. Captive bachelor groups need to be carefully monitored, particularly during periods of demographic transition, due to the possibility for escalating aggression. We examined behavioral changes in a bachelor group at the Saint Louis Zoo following two significant alterations in group composition: (1) the introduction of two immature related males in 2011 and (2) the death of the dominant silverback in 2015. Behavioral data were collected on group members using 15 min focal observations with 30-s instantaneous scans, totaling 185.25 hr for the first transition and 115.25 hr for the second transition (with equal effort in baseline and transition periods). We found that the addition of the two subadult males resulted in a significant increase in affiliation in the group often initiated by these new younger individuals, while the rate of abnormal and aggressive behavior did not change significantly. The rate of abnormal, aggressive, or affiliative behavior in the group also did not change significantly following the death of the dominant silverback. Overall, we conclude that this bachelor group and potentially others can endure possibly destabilizing social transitions and remain cohesive social units. Behavioral changes will result from transitions, but the effect on individuals or impact on the group will vary. Therefore, longitudinal monitoring of bachelor groups will be essential for effective management during any major changes.

Origin of the human malaria parasite Plasmodium falciparum in gorillas.

Plasmodium falciparum is the most prevalent and lethal of the malaria parasites infecting humans, yet the origin and evolutionary history of this important pathogen remain controversial. Here we develop a single-genome amplification strategy to identify and characterize Plasmodium spp. DNA sequences in faecal samples from wild-living apes. Among nearly 3,000 specimens collected from field sites throughout central Africa, we found Plasmodium infection in chimpanzees (Pan troglodytes) and western gorillas (Gorilla gorilla), but not in eastern gorillas (Gorilla beringei) or bonobos (Pan paniscus). Ape plasmodial infections were highly prevalent, widely distributed and almost always made up of mixed parasite species. Analysis of more than 1,100 mitochondrial, apicoplast and nuclear gene sequences from chimpanzees and gorillas revealed that 99% grouped within one of six host-specific lineages representing distinct Plasmodium species within the subgenus Laverania. One of these from western gorillas comprised parasites that were nearly identical to P. falciparum. In phylogenetic analyses of full-length mitochondrial sequences, human P. falciparum formed a monophyletic lineage within the gorilla parasite radiation. These findings indicate that P. falciparum is of gorilla origin and not of chimpanzee, bonobo or ancient human origin.

Lateralization and performance asymmetries in the termite fishing of wild chimpanzees in the goualougo triangle, republic of Congo.

The nearly universal right hand preference manifested by human populations is one of the most pronounced manifestations of population-level lateralization. Morphological and archeological evidence indicate that this behavioral specialization may have emerged among our hominin ancestors. Whether population-level behavioral asymmetries are evident in non-human animals remains a topic of considerable scientific debate, with the most consistent evidence of population-level trends emerging from studies of chimpanzees (Pan troglodytes). However, previous studies of population-level lateralization in wild apes have relied upon data sets pooled across populations to reach adequate sample sizes. Our aim was to test for population-level handedness within a single wild chimpanzee population, and also to determine if performance asymmetries were associated with handedness. To address these questions, we coded handedness and duration of fishing probe insertions from remote video footage of chimpanzee visitation to termite nests (totaling 119 hr) in the Goualougo Triangle, Republic of Congo. Similar to reports from other populations, chimpanzees in the Goualougo Triangle showed robust individual hand preferences for termite fishing. There were 46 right-handed, 39 left-handed, and 4 ambiguously-handed individuals. Though we did not detect an overall significant population-level handedness (t(88) = 0.83, n.s.) in this study, males showed a greater right hand preference than females. Further, we found that average dipping latencies were significantly faster for right- compared to left-handed chimpanzees. Possible explanations and evolutionary implications of taxa- and task-specific patterns of population-level laterality are discussed. Am. J. Primatol. 78:1190-1200, 2016. © 2016 Wiley Periodicals, Inc.

The interdigital brace and other grips for termite nest perforation by chimpanzees of the Goualougo Triangle, Republic of Congo.

Studies of chimpanzee termite foraging enlighten our understanding of early hominin tool use not only by modeling the cognitive ability of our ancestors but also by emphasizing the possible role of social insects in the hominin diet. The chimpanzees of the Goualougo Triangle are known to have one of the largest and most complex tool repertoires reported for wild chimpanzees. One tool set habitually used by this population includes a perforating tool to penetrate the hard outer crust of elevated termite nests before fishing for termite prey with an herbaceous stem. Here, we report the variation present in the grips used on the perforating tool. Our analysis of video recordings of chimpanzee visitation to termite nests over a 3-year period shows that these chimpanzees use a variety of grips to navigate the challenges encountered in opening a termite nest. For situations in which the soil is most hardened, perforating requires force and a power grip is often used. When the soil in the passageway is loose, precision grips are suitable for the task. One of the preferred grips reported here is an interdigital brace, which has previously been described in studies of how some people hold a pencil. In this study, for the first time, the interdigital brace has been thoroughly described for chimpanzees. The various strategies and grips used during perforation emphasize the importance of termites as a nutritional resource that should be considered more strongly as a food used by early hominins.

The sampling scheme matters: Pan troglodytes troglodytes and P. t. schweinfurthii are characterized by clinal genetic variation rather than a strong subspecies break.

Populations of an organism living in marked geographical or evolutionary isolation from other populations of the same species are often termed subspecies and expected to show some degree of genetic distinctiveness. The common chimpanzee (Pan troglodytes) is currently described as four geographically delimited subspecies: the western (P. t. verus), the nigerian-cameroonian (P. t. ellioti), the central (P. t. troglodytes) and the eastern (P. t. schweinfurthii) chimpanzees. Although these taxa would be expected to be reciprocally monophyletic, studies have not always consistently resolved the central and eastern chimpanzee taxa. Most studies, however, used data from individuals of unknown or approximate geographic provenance. Thus, genetic data from samples of known origin may shed light on the evolutionary relationship of these subspecies. We generated microsatellite genotypes from noninvasively collected fecal samples of 185 central chimpanzees that were sampled across large parts of their range and analyzed them together with 283 published eastern chimpanzee genotypes from known localities. We observed a clear signal of isolation by distance across both subspecies. Further, we found that a large proportion of comparisons between groups taken from the same subspecies showed higher genetic differentiation than the least differentiated between-subspecies comparison. This proportion decreased substantially when we simulated a more clumped sampling scheme by including fewer groups. Our results support the general concept that the distribution of the sampled individuals can dramatically affect the inference of genetic population structure. With regard to chimpanzees, our results emphasize the close relationship of equatorial chimpanzees from central and eastern equatorial Africa and the difficult nature of subspecies definitions.

Characterization of MHC class II B polymorphism in multiple populations of wild gorillas using non-invasive samples and next-generation sequencing.

Genes encoded by the major histocompatibility complex (MHC) are crucial for the recognition and presentation of antigens to the immune system. In contrast to their closest relatives, chimpanzees and humans, much less is known about variation in gorillas at these loci. This study explored the exon 2 variation of -DPB1, -DQB1, and -DRB genes in 46 gorillas from four populations while simultaneously evaluating the feasibility of using fecal samples for high-throughput MHC genotyping. By applying strict similarity- and frequency-based analysis, we found, despite our modest sample size, a total of 18 alleles that have not been described previously, thereby illustrating the potential for efficient and highly accurate MHC genotyping from non-invasive DNA samples. We emphasize the importance of controlling for multiple potential sources of error when applying this massively parallel short-read sequencing technology to PCR products generated from low concentration DNA extracts. We observed pronounced differences in MHC variation between species, subspecies and populations that are consistent with both the ancient and recent demographic histories experienced by gorillas.

Insect prey characteristics affecting regional variation in chimpanzee tool use.

It is an ongoing interdisciplinary pursuit to identify the factors shaping the emergence and maintenance of tool technology. Field studies of several primate taxa have shown that tool using behaviors vary within and between populations. While similarity in tools over spatial and temporal scales may be the product of socially learned skills, it may also reflect adoption of convergent strategies that are tailored to specific prey features. Much has been claimed about regional variation in chimpanzee tool use, with little attention to the ecological circumstances that may have shaped such differences. This study examines chimpanzee tool use in termite gathering to evaluate the extent to which the behavior of insect prey may dictate chimpanzee technology. More specifically, we conducted a systematic comparison of chimpanzee tool use and termite prey between the Goualougo Triangle in the Republic of Congo and the La Belgique research site in southeast Cameroon. Apes at both of these sites are known to use tool sets to gather several species of termites. We collected insect specimens and measured the characteristics of their nests. Associated chimpanzee tool assemblages were documented at both sites and video recordings were conducted in the Goualougo Triangle. Although Macrotermitinae assemblages were identical, we found differences in the tools used to gather these termites. Based on measurements of the chimpanzee tools and termite nests at each site, we concluded that some characteristics of chimpanzee tools were directly related to termite nest structure. While there is a certain degree of uniformity within approaches to particular tool tasks across the species range, some aspects of regional variation in hominoid technology are likely adaptations to subtle environmental differences between populations or groups. Such microecological differences between sites do not negate the possibility of cultural transmission, as social learning may be required to transmit specific behaviors among individuals.

The genetic population structure of wild western lowland gorillas (Gorilla gorilla gorilla) living in continuous rain forest.

To understand the evolutionary histories and conservation potential of wild animal species it is useful to assess whether taxa are genetically structured into different populations and identify the underlying factors responsible for any clustering. Landscape features such as rivers may influence genetic population structure, and analysis of structure by sex can further reveal effects of sex-specific dispersal. Using microsatellite genotypes obtained from noninvasively collected fecal samples we investigated the population structure of 261 western lowland gorillas (WLGs) (Gorilla gorilla gorilla) from seven locations spanning an approximately 37,000 km(2) region of mainly continuous rain forest within Central African Republic (CAR), Republic of Congo and Cameroon. We found our sample to consist of two or three significantly differentiated clusters. The boundaries of the clusters coincided with courses of major rivers. Moreover, geographic distance detoured around rivers better-explained variation in genetic distance than straight line distance. Together these results suggest that major rivers in our study area play an important role in directing WLG gene flow. The number of clusters did not change when males and females were analyzed separately, indicating a lack of greater philopatry in WLG females than males at this scale.

Long-term observations in the Ndoki forest resolve enduring questions about truffle foraging by western lowland gorillas.

While there is growing recognition of the importance of traditional knowledge in science, these perspectives remain underrepresented in research publications. However, the synthesis of these approaches has tremendous potential to improve our understanding of wildlife and ecosystems. Toward realizing this aim, we combined local traditional knowledge with molecular classification techniques to investigate "soil scratching" behavior in western lowland gorillas in two localities in Republic of Congo, the Goualougo Triangle and the Djéké Triangle. Daily observations of four gorilla groups for nearly a decade revealed that soil scratching is a foraging strategy to access a deer truffle species, identified here as Elaphomyces labyrinthinus. We also conducted group scans to calculate the time gorillas spent foraging for deer truffles and video focal observations to assess foraging efficiency. There was considerable variation in soil scratching across groups. It was most common in Buka's group, followed by Kingo's group and Mététélé's group. Truffle foraging was rarely observed in the Loya-Makassa group. While the overall distribution of deer truffles seemingly determines the occurrence of this behavior across populations, we found indications of social influences on soil scratching within populations. For example, an adult female transferred from a group in which the behavior was rare to another group where it is common and adjusted her frequencies of soil scratching to that of her new group. Finally, these findings were included in an ecological impact assessment of the Djéké Triangle that prompted conservation managers to shift the location of tourism-associated construction to safeguard this putative cultural behavior.

A brief history of primate research in the Ndoki forest.

The Nouabalé-Ndoki National Park (NNNP) in Republic of Congo has become a beacon for conservation in Central Africa. This manuscript documents the arrival of primatologists, the establishment of field stations and major discoveries in primate behavior and ecology. Field stations were strategically established to study primate behavior in a variety of different contexts from stationary platforms to forest follows of habituated groups. The implementation of new technologies and analyses have also been a hallmark of research at Ndoki. Scientists are shaping a new era in primatology at NNNP by building on past successes and promoting the next generation of Congolese conservationists to address environmental challenges. Results have proven crucial in discussions with government and industry and led to conservation gains such as the inclusion of the intact forests of the Goualougo and Djéké Triangles into the NNNP. The research stations have also become essential for developing a long-term certified sustainable international gorilla tourism program. Despite the many advancements for conservation such as increased protection of forests, development of internationally recognized protocols and large-scale capacity building initiatives, there are reasons for considerable concern in the near- and long-term for primates and their forest habitats in the Ndoki landscape. To address these concerns, we emphasize the long history of forming partnerships with local communities. We also discuss shared overlap featuring multicultural and environmental use of forest resources that is likely to be crucial in championing the conservation of the Ndoki forests for the next 25 years and beyond.

Planning abilities of wild chimpanzees (Pan troglodytes troglodytes) in tool-using contexts.

Planning is a type of problem solving in which a course of future action is devised via mental computation. Potential advantages of planning for tool use include reduced effort to gather tools, closer alignment to an efficient tool design, and increased foraging efficiency. Chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle use a variety of different types of tools. We hypothesized that procurement strategy (brought to the termite nest, manufactured or acquired at the termite nest, or borrowed from others) reflects planning for current needs, with tool transport behavior varying by tool type and by age and sex class. It is also possible that chimpanzees anticipate the need for tools at future times, which would be evidenced by transporting multiple tool types for a sequential task. One year of video recordings at termite nests were systematically screened for tool procurement; data comprised 299 tool procurement events across 66 chimpanzees. In addition, we screened video recordings of leaf sponging and honey gathering, which resulted in another 38 procurement events. Fishing probes, which are typically used during a single visit, were typically transported to termite nests, while puncturing tools, which are durable and remain on site, were more often acquired at termite nests. Most tools transported in multiples were fishing probes, perhaps in anticipation that a single probe might not last through an entire foraging bout or might be transferred to another chimpanzee. We further documented that chimpanzees transported tool sets, comprising multiple different tool types used in sequence. Mature chimpanzees transported tools more often than did immatures. These observations suggest that chimpanzees plan tool use flexibly, reflecting the availability of raw materials and the likelihood that specific tool types will be needed for particular tasks. Developmental studies and further integration of behavioral, spatial, and archaeological data will help to illuminate the decision making and time depth of planning associated with tool technologies in living primates and hominin ancestors.