Ape Nest Research Articles

Great ape surveys and the implications of long-term monitoring in the Djéké Triangle, Republic of Congo.

Existing protected areas are anchors for conservation. Safeguarding flora and fauna within their peripheral areas is essential to maintaining their integrity and to potential increases to the area under effective conservation. With the decline in tropical forests, initiatives to increase the area of undisturbed forests under strict protection, particularly those neighboring protected areas, is of critical importance. Applied research has informed such land-management decisions for areas surrounding the Nouabalé-Ndoki National Park (NNNP) in Republic of Congo since the park's inception three decades ago. Here, we present results of the first systematic line transect survey of great ape nests conducted in the Djéké Triangle, a 100 km2 unlogged continuous forest in the Kabo Forestry Management Unit adjacent to the NNNP. Distance sampling methods applied along 26 line transects on two different occasions (2016 and 2018, with total effort of 69.4km) provided density estimates of 0.75 (95% confidence interval (CI) 0.52-1.09) and 0.61 (95% CI 0.40-0.92) chimpanzees/km2 and 2.15 (95% CI 1.36-3.40) and 1.19 (95% CI 0.78-1.82) gorillas/km2 for each of the two surveys, respectively. Estimated ape densities were compared to others across the landscape. The findings provide a unique baseline in an area that supports ongoing behavioral research and future gorilla tourism opportunities in the Djéké Triangle. More importantly, results provided empirical evidence of the environmental value and strategic conservation importance supporting inclusion of the Djéké Triangle into the NNNP in 2023. These long-term monitoring results inform best-practice standards and ape tourism certification.

Factors Influencing Density and Distribution of Great Ape Nests in the Absence of Human Activities

Factors Influencing Density and Distribution of Great Ape Nests in the Absence of Human Activities

Variation in behavioral traits of two frugivorous mammals may lead to differential responses to human disturbance.

Human activities can lead to a shift in wildlife species’ spatial distribution. Understanding the specific effects of human activities on ranging behavior can improve conservation management of wildlife populations in human‐dominated landscapes. This study evaluated the effects of forest use by humans on the spatial distribution of mammal species with different behavioral adaptations, using sympatric western lowland gorilla and central chimpanzee as focal species. We collected data on great ape nest locations, ecological and physical variables (habitat distribution, permanent rivers, and topographic data), and anthropogenic variables (distance to trails, villages, and a permanent research site). Here, we show that anthropogenic variables are important predictors of the distribution of wild animals. In the resource model, the distribution of gorilla nests was predicted by nesting habitat distribution, while chimpanzee nests were predicted first by elevation followed by nesting habitat distribution. In the anthropogenic model, the major predictors of gorilla nesting changed to human features, while the major predictors of chimpanzee nesting remained elevation and the availability of their preferred nesting habitats. Animal behavioral traits (body size, terrestrial/arboreal, level of specialization/generalization, and competitive inferiority/superiority) may influence the response of mammals to human activities. Our results suggest that chimpanzees may survive in human‐encroached areas whenever the availability of their nesting habitat and preferred fruits can support their population, while a certain level of human activities may threaten gorillas. Consequently, the survival of gorillas in human‐dominated landscapes is more at risk than that of chimpanzees. Replicating our research in other sites should permit a systematic evaluation of the influence of human activity on the distribution of mammal populations. As wild animals are increasingly exposed to human disturbance, understanding the resulting consequences of shifting species distributions due to human disturbance on animal population abundance and their long‐term survival will be of growing conservation importance.

The distribution of plant consumption traits across habitat types and the patterns of fruit availability suggest a mechanism of coexistence of two sympatric frugivorous mammals.

Understanding the mechanisms governing the coexistence of organisms is an important question in ecology, and providing potential solutions contributes to conservation science. In this study, we evaluated the contribution of several mechanisms to the coexistence of two sympatric frugivores, using western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) in a tropical rainforest of southeast Cameroon as a model system. We collected great ape fecal samples to determine and classify fruit species consumed; we conducted great ape nest surveys to evaluate seasonal patterns of habitat use; and we collected botanical data to investigate the distribution of plant species across habitat types in relation to their “consumption traits” (which indicate whether plants are preferred or fallback for either gorilla, chimpanzee, or both). We found that patterns of habitat use varied seasonally for both gorillas and chimpanzees and that gorilla and chimpanzee preferred and fallback fruits differed. Also, the distribution of plant consumption traits was influenced by habitat type and matched accordingly with the patterns of habitat use by gorillas and chimpanzees. We show that neither habitat selection nor fruit preference alone can explain the coexistence of gorillas and chimpanzees, but that considering together the distribution of plant consumption traits of fruiting woody plants across habitats as well as the pattern of fruit availability may contribute to explaining coexistence. This supports the assumptions of niche theory with dominant and subordinate species in heterogeneous landscapes, whereby a species may prefer nesting in habitats where it is less subject to competitive exclusion and where food availability is higher. To our knowledge, our study is the first to investigate the contribution of plant consumption traits, seasonality, and habitat heterogeneity to enabling the coexistence of two sympatric frugivores.OPEN RESEARCH BADGES This article has earned an Open Data Badge for making publicly available the digitally‐shareable data necessary to reproduce the reported results. The data is available at https://datadryad.org/resource/doi:10.5061/dryad.ms65f29.

Effects of Land cover change on Great Apes distribution at the Lob\xe9k\xe9 National Park and its surrounding Forest Management Units, South-East Cameroon. A 13 year time series analysis

Understanding the effects of land cover change on wildlife distribution is very important for resource management and conservation planning. This paper aimed at detecting the effects of land cover change on great apes distribution at the Lobéké National Park and its bounded forest management units (FMUs). Data on great ape nests were collected in the field for the years 2001 and 2014 through distance sampling with line transects. Landsat TM images of South-East Cameroon for the years 2001 and 2014 were acquired from earth explorer and corrected atmospherically for proper visualization. An area of interest comprising the Lobéké National Park and its FMUs was extracted for classification and change detection. A comparison in great apes nest distribution and change per land cover change category was done for both years through point pattern analysis, whereas a time series analysis of the detected land cover change impacts on great apes nest distribution for a period of 13 years was modeled using logistic growth and regression equations in Vensim 7.2. The results could illustrate that, as land cover changes from one cover type in 2001 to another in 2014, increases or decreases in great apes nests were observed within each changed area.

Sleep and nesting behavior in primates: A review.

Sleep is a universal behavior in vertebrate and invertebrate animals, suggesting it originated in the very first life forms. Given the vital function of sleep, sleeping patterns and sleep architecture follow dynamic and adaptive processes reflecting trade-offs to different selective pressures. Here, we review responses in sleep and sleep-related behavior to environmental constraints across primate species, focusing on the role of great ape nest building in hominid evolution. We summarize and synthesize major hypotheses explaining the proximate and ultimate functions of great ape nest building across all species and subspecies; we draw on 46 original studies published between 2000 and 2017. In addition, we integrate the most recent data brought together by researchers from a complementary range of disciplines in the frame of the symposium "Burning the midnight oil" held at the 26th Congress of the International Primatological Society, Chicago, August 2016, as well as some additional contributors, each of which is included as a "stand-alone" article in this "Primate Sleep" symposium set. In doing so, we present crucial factors to be considered in describing scenarios of human sleep evolution: (a) the implications of nest construction for sleep quality and cognition; (b) the tree-to-ground transition in early hominids; (c) the peculiarities of human sleep. We propose bridging disciplines such as neurobiology, endocrinology, medicine, and evolutionary ecology, so that future research may disentangle the major functions of sleep in human and nonhuman primates, namely its role in energy allocation, health, and cognition.

Savanna chimpanzees adjust sleeping nest architecture in response to local weather conditions.

Great ape nests are hypothesized to aid safe, secure sleep via providing thermoregulation or protection from predators and vectors. We aimed to describe and investigate variation in chimpanzee nest architecture across two populations in response to local weather conditions. We experimentally tested whether nests provide insulation by measuring heat loss within and outside nests, and took detailed measurements of the number, size, and type of materials used in nest building across two dry-habitat research sites (Fongoli, Senegal, and Issa, Tanzania). We tested application of principal components analysis (PCA) to extract composite quantitative measures of the key components of shape and architecture, before testing how PCs vary across populations with overnight weather conditions that reflect hypothesized thermoregulatory function. Heat loss is greater and occurs faster outside of nests. PCA allowed meaningful comparison of nests within and between sites. Nest variation at both sites revealed chimpanzees built thicker nests in cooler conditions and used more broken branches and support in moister conditions. Chimpanzees in Fongoli used more lining and mattress material in colder conditions, whilst in Issa nest depth and support branch size were larger in windier conditions. Shape and architectural measures reflected insulation and stability of nest structure. Chimpanzees in Fongoli and Issa may achieve the same functional goals by adjusting nest shape and architecture in different ways. These results suggest that wild chimpanzees show flexible building techniques in response to local, overnight weather conditions in making an insulating and stable, supportive platform for sleep.

African apes coexisting with logging: Comparing chimpanzee (Pan troglodytes troglodytes) and gorilla (Gorilla gorilla gorilla) resource needs and responses to forestry activities

The extraction of timber often conflicts with the well-being and conservation of wildlife. In particular, there is a need to better understand the impact of tree removal under selective logging regimes on local ecological communities. We conducted ape nest counts along line transects before, during, and after logging to assess the impact of timber harvesting and associated activities on sympatric chimpanzees and gorillas in a forestry concession in northern Republic of Congo. We used generalized linear models to relate ape nest counts to a set of predictor variables, representing the impact of logging and controlled for variation in environmental conditions including food availability, habitat and rainfall. Commercial forest inventory data were used to assess the baseline influence of food availability and forest structure on ape distribution. Higher numbers of chimpanzees were found in proximity to their preferred tree foods, whereas gorillas were associated with more heterogeneous habitats. Chimpanzee nest encounter rates decreased with increasing intensity of human impacts. Gorillas also avoided areas with active timber exploitation and roads, but were attracted to recently logged areas with abundant terrestrial herbaceous vegetation. Species-specific responses were consistent with theoretical predictions of niche partitioning and cumulative human influence. Based on these findings, we provide recommendations to improve existing guidelines and forest certification standards aimed at safeguarding ape populations.

Factors Influencing the Survival of Sympatric Gorilla (Gorilla gorilla gorilla) and Chimpanzee (Pan troglodytes troglodytes) Nests

Accurate and precise surveys of primate abundance provide the basis for understanding species ecology and essential information for conservation assessments. Owing to the elusive nature of wild apes and the vast region of dense forest they inhabit, population estimates of central chimpanzees (Pan troglodytes troglodytes) and western lowland gorillas (Gorilla gorilla gorilla) have largely relied on surveys of their nests. Specific information about the nesting behavior of apes permits the estimation of the number of nests built (nest creation rate). Similarly, information on nest characteristics and environmental factors can be used to estimate the time it takes nests to decay (nest decay rate). Nest creation and decay rates are then used to convert nest density estimates to absolute ape densities. Population estimates that use site-specific estimates of nest creation and decay rates are more accurate and precise. However, it is common practice to generalize these conversion factors across sites because of the additional cost of studies required to gather the information to estimate them. Over a 9-mo study period, we detected and monitored the time to decay of gorilla nests (N = 514) and chimpanzee nests (N = 521) in northern Republic of Congo. We investigated the influence of nest characteristics and environmental factors on nest survivorship and estimated the mean time to nest decay (or equivalently survival) using MARK. Key factors influencing nest decay rate included ape species, forest type, nest height, mean rainfall, nest structure, nest type, and primary aspects of nest construction. Our findings highlight the synergistic effect of behavior and environment on great ape nest degradation, as well as providing practical insights for improving measures to monitor remaining populations of these endangered species.

Spatial and Temporal Dynamics of a Mortality Event among Central African Great Apes.

In 2006–2007 we observed an unusual mortality event among apes in northern Republic of Congo that, although not diagnostically confirmed, we believe to have been a disease outbreak. In 2007–2011 we conducted ape nest surveys in the region, recording 11,835 G. g. gorilla nests (2,262 groups) and 5,548 P. t. troglodytes nests (2,139 groups). We developed a statistical model to determine likely points of origin of the outbreak to help identify variables associated with disease emergence and spread. We modeled disease spread across the study area, using suitable habitat conditions for apes as proxy for local ape densities. Infectious status outputs from that spread model were then used alongside vegetation, temperature, precipitation and human impact factors as explanatory variables in a Generalized Linear Model framework to explain observed 2007–2011 ape nest trends in the region. The best models predicted emergence in the western region of Odzala-Kokoua National Park and north of the last confirmed Ebola virus disease epizootics. Roads were consistently associated with attenuation of modeled virus spread. As disease is amongst the leading threats to great apes, gaining a better understanding of disease transmission dynamics in these species is imperative. Identifying ecological drivers underpinning a disease emergence event and transmission dynamics in apes is critical to creating better predictive models to guide wildlife management, develop potential protective measures for wildlife and to reduce potential zoonotic transmission to humans. The results of our model represent an important step in understanding variables related to great ape disease ecology in Central Africa.

Impact of human activities on Cross River gorilla Gorilla gorilla diehli habitats in the Mawambi Hills, southwest Cameroon

Most forest habitats in Cameroon support villages whose inhabitants are heavily reliant on the forests for their everyday needs. We assessed the socio-economic status and natural resource use of 3 villages bordering Mawambi Hills, an unprotected forest in southwest Cameroon, home to both the Critically Endangered Cross River gorilla Gorilla gorilla diehli and the Endangered Nigeria-Cameroon chimpanzee Pan troglodytes ellioti. This forest likely hosts one of the largest sub-populations of Cross River gorillas in Cameroon and is vulnerable to human influence given that it lies in close proximity to surrounding villages and is less hilly than other Cross River gorilla habitats. We show that employment opportunities in surrounding villages are rare, and people rely on agriculture and the collection of non-timber forest products (NTFPs), such as eru Gnetum spp. and bush mango Irvingia spp., for food and income. We also show that areas of heavy human activity coincide spatially with great ape nest sites and raise concerns about the potential impact of human activities on great ape survival at the site. We call for (1) sustainable harvesting and marketing of NTFPs, (2) the introduction of alternative livelihood opportunities (e.g. beekeeping and on-farm production of NTFPs), (3) active wildlife law enforcement to safe- guard the persistence of great apes, and (4) the introduction of a community-based forest manage- ment programme that will enable communities to secure the rights to use and manage surround- ing forest resources legally.

Estimating Orangutan Densities Using the Standing Crop and Marked Nest Count Methods: Lessons Learned for Conservation

ABSTRACTReliable estimates of great ape abundance are needed to assess distribution, monitor population status, evaluate conservation tactics, and identify priority populations for conservation. Rather than using direct counts, surveyors often count ape nests. The standing crop nest count (SCNC) method converts the standing stock of nests into animal densities using a set of parameters, including nest decay rate. Nest decay rates vary greatly over space and time, and it takes months to calculate a site‐specific value. The marked nest count (MNC) method circumvents this issue and only counts new nests produced during a defined period. We compared orangutan densities calculated by the two methods using data from studies in Sumatra and Kalimantan, Indonesia. We show how animal densities calculated using nest counts should be cautiously interpreted when used to make decisions about management or budget allocation. Even with site‐specific decay rates, short studies using the SCNC method may not accurately reflect the current population unless conducted at a scale sufficient to include wide‐ranging orangutan movement. Density estimates from short studies using the MNC method were affected by small sample sizes and by orangutan movement. To produce reliable results, the MNC method may require a similar amount of effort as the SCNC method. We suggest a reduced reliance on the traditional line transect surveys in favor of feasible alternative methods when absolute abundance numbers are not necessary or when site‐specific nest decay rates are not known. Given funding constraints, aerial surveys, reconnaissance walks, and interview techniques may be more cost‐effective means of accomplishing some survey goals.

Survey of Raphia swamp forest, Republic of Congo, indicates high densities of Critically Endangered western lowland gorillas Gorilla gorilla gorilla

The Critically Endangered western lowland gorilla Gorilla gorilla gorilla is found at high density in the swamp forests of Lac Télé Community Reserve, northern Republic of Congo. Preliminary surveys for apes on the Reserve periphery in the 1990s indicated that this area may also contain high densities of apes and we wished to verify this. Using distance sampling methodology we surveyed 1,029 km2 of the south-eastern Reserve periphery including Raphia swamp forests. We recorded 1,021 ape nests along 21 2-km transects. Results indicated that ape density estimated from nest group density, excluding old and very old Raphia nests (which may decay slowly), was very high: 6.81 weaned apes km-2 (95% confidence interval 3.93–11.84). Gorillas built most of the nests recorded: their density was estimated at 5.25 individuals km-2 (2.70–10.19). Most nests were constructed in Raphia swamp forest despite it covering only 25% of the study area. These are among the highest recorded densities of apes in Central Africa but, as nest decay rates in swamps are unstudied, we also estimated densities based on hypothetically slow nest decay rates and published data from other habitats. Gorilla densities may be linked to availability of monocotyledons as food and the natural protection from hunting offered by Raphia swamps. Gorilla density in swamp forest is spatially and temporally variable and surveys are required to estimate ape populations elsewhere in this habitat. Most Raphia swamps in Congo are unprotected, including the periphery of Lac Télé Community Reserve; protection and management are required to assure their continued availability for apes.

The Origin of Spaces: Understanding Residential Satisfaction from Ape Nests, Human Cultures and the Hierarchy of Natural Housing Functions

This paper seeks to establish a natural approach towards understanding housing by suggesting the existence of a primal hierarchy of natural housing functions based upon an analysis of nesting behaviour in hominid apes (gorillas, chimpanzees, bonobos and orang‐utans). A review of international research on housing satisfaction and an analysis of data from the American Housing Survey confirm the modern relevance of this hierarchy of natural housing functions to the perceived housing quality of both homeowners and tenants. The proposed hierarchy provides a simple, compelling and eminently cross‐cultural perspective for housing quality and policy analysis. Adding this natural viewpoint to traditional approaches may generate useful alternatives and novel insights.

A Survey of Chimpanzees (Pan troglodytes troglodytes) and Gorillas (Gorilla gorilla gorilla) in the Selectively Logged Ngotto Forest, Central African Republic

Currently, the timber company Industrie Forestiére du Batalimo is selectively logging the Ngotto Forest in the Central African Republic. The forest is home to a population of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) and provides the Bofi-speaking people of Grima with food, medicine, housing material, and other commodities. Over a 7-month period, the research team conducted a line-transect survey of the great ape population in the forests to the south of Grima to document their distribution. For comparison purposes, the team also surveyed a section of adjacent forest that had already been logged. Ape nests were significantly rarer in the logged forest than in the unlogged forest, and ape nests were most common in the more pristine forests to the south. This report further discusses the effects of logging and other human activities.

Discriminating between village and commercial hunting of apes

Hunting is the major driver of large mammal decline in Central African forests. In slowly reproducing species even low hunting pressure leaves spatial gradients with wildlife density increasing with distance from transport routes and human settlements. Park management can use this pattern formation to identify sources of threats, but also to discriminate between different threat scenarios, such as the impact of subsistence vs. commercial hunting. We conducted an ape survey in the mountainous Moukalaba Doudou National Park, Gabon, to evaluate whether potential population gradients would emanate from the three human population centers in the region or the villages surrounding the park. Using generalized linear modeling we found hill slope as a good predictor of ape nest occurrence probability and the distance to human population centers a better predictor of ape nest density and ape nest group size than distance to villages. In fact ape nest density was three times lower at the park borders close to the human population centers than in the park’s interior. The results indicate that Moukalaba’s ape population is more impacted by commercial than subsistence hunting and suggest that park management should focus conservation efforts on the human population centers. We conclude that in particular for slowly reproducing species geographic information on wildlife population gradients are of additional value for guiding protected area management. The hunting impact on those species might be easily underestimated, if derived only from market surveys or transport route controls, where they are only rarely found.

Comparing ape densities and habitats in northern Congo: surveys of sympatric gorillas and chimpanzees in the Odzala and Ndoki regions

The conservation status of western lowland gorillas and central chimpanzees in western equatorial Africa remains largely speculative because many remote areas have never been surveyed and the impact of emergent diseases in the region has not been well documented. In this study, we compared ape densities and habitats in the Lokoué study area in Odzala National Park and the Goualougo Triangle in Nouabalé-Ndoki National Park in northern Republic of Congo. Both of these sites have long been considered strongholds for the conservation of chimpanzees and gorillas, but supposedly differ in vegetative composition and relative ape abundance. We compared habitats between these sites using conventional ground surveys and classified Landsat-7 ETM+ satellite images. We present density estimates via both standing-crop and marked-nest methods for the first time for sympatric apes of the Congo Basin. The marked-nest method was effective in depicting chimpanzee densities, but underestimated gorilla densities at both sites. Marked-nest surveys also revealed a dramatic decline in the ape population of Lokoué which coincided with a local Ebola epidemic. Normal baseline fluctuations in ape nest encounter rates during the repeated passages of marked-nest surveys were clearly distinguishable from a 80% decline in ape nest encounter rates at Lokoué. Our results showed that ape densities, habitat composition, and population dynamics differed between these populations in northern Congo. We emphasize the importance of intensifying monitoring efforts and further refinement of ape survey methods, as our results indicated that even the largest remaining ape populations in intact and protected forests are susceptible to sudden and dramatic declines.

Selective Logging, Habitat Quality and Home Range Use by Sympatric Gorillas and Chimpanzees: A Case Study from an Active Logging Concession in Southeast Cameroon

We examined range use by great apes during logging activities and investigated associations between local variations in ape abundance and changes in the structure of the habitat or in the availability of fruits after disturbances. We carried out two annual censuses of western lowland gorilla (G. g. gorilla) and chimpanzee populations (Pan t. troglodytes) in an active logging concession in Southeast Cameroon. The results suggest that gorillas may adapt their range use to avoid most recently logged compartments, while chimpanzees appear to be more spatially resilient to logging. In our study site, selective logging affected 10% of the forest. After logging, gorillas nested in all types of vegetation, while chimpanzees nested exclusively in mixed mature forest. Fruit availability was not affected by logging and did not explain the distribution of ape nests in the study area.

Distinguishing between the nests of sympatric chimpanzees and gorillas

SummaryOur current inability to estimate precisely the population sizes of chimpanzees and gorillas across much of the Congo Basin has been detrimental to the development of conservation strategies for the preservation of these endangered apes. Systematic counts of nests are currently the most commonly used method to estimate ape abundance, but distinguishing between the nests of sympatric chimpanzees and gorillas has proven to be an enduring obstacle to estimating species‐specific abundance. In general, the builder of more than 75% of nests recorded during surveys is undetermined. We hypothesized that sleeping habits and nest building patterns would allow us to differentiate between the nests of these apes.We constructed a predictive model using stepwise discriminant function analysis to determine characteristics that accurately distinguished between chimpanzee and gorilla nests. We analysed 13 variables associated with 3425 ape nests from three independent surveys conducted in the Goualougo Triangle of the Nouabalé‐Ndoki National Park, Republic of Congo.The model correctly classified more than 90% of nests in our validation subsample. Nest height, nest type, forest type and understorey closure were identified as important variables for distinguishing between chimpanzee and gorilla nests at this site. Attributing nests to either species increased the precision of resulting density estimates, which enhanced the statistical power to detect trends in population fluctuation.Although specific variables may differ between study sites, we have demonstrated that predictive models to distinguish between the nests of sympatric chimpanzee and gorillas provide a promising approach to improving the quality of ape survey data.Synthesis and applications. Our study introduces an innovative solution to the dilemma of discriminating between the nests of sympatric chimpanzees and gorillas, which increases the specificity and precision of resulting ape abundance estimates. There is an urgent need to improve methods to evaluate and monitor remaining ape populations across western and central Africa that are experiencing the imminent threats of emergent diseases, poaching and expanding human development. Increasing the quality of density estimates from field survey data will aid in the development of local conservation initiatives, national strategies and international policies on behalf of remaining ape populations.

Ape Abundance and Habitat Use in the Goualougo Triangle, Republic of Congo

Chimpanzee (Pan troglodytes troglodytes) and western gorilla (Gorilla gorilla gorilla) populations in central Africa are rapidly declining as a result of disease epidemics, commercial bushmeat hunting, and habitat destruction. Our main objective was to estimate the absolute abundance and habitat utilization of chimpanzees and gorillas in the intact forests of the Goualougo Triangle in the Republic of Congo, and in an adjacent area in which selective logging will take place in the near future. The estimates provide a unique baseline for apes inhabiting an undisturbed environment. A second objective was to compare estimates of abundance and patterns of habitat utilization generated by different techniques: 1) distance sampling of individual ape nests and nest sites along line transects, 2) direct observations of apes during reconnaissance surveys, and 3) observations of ape traces during reconnaissance surveys. We completed a total of 222 km of line transect surveys in 4 sampling areas, resulting in overall density estimates of 1.53 chimpanzees/km 2 and 2.34 gorillas/km 2 from nest sites. We generated a density estimate of 2.23 chimpanzees/km 2 from direct observations during reconnaissance surveys of a semihabituated community in 1 of the 4 sampling areas. Habitat use profiles that nest surveys depicted on transects differed from those of direct observations and traces we encountered on reconnaissance surveys. We found the highest overall abundance of chimpanzee nests in monodominant Gilbertiodendron forest, whereas our direct observations showed that chimpanzees preferred mixed species forest. Transects that