Associations between the drill (Mandrillus leucophaeus) and sympatric monkeys in Korup National Park,Cameroon

Studies of polyspecific associations among African forest primates have primarily focused on arboreal Cercopithecus and Procolobus/Colobus species. We examined the association frequency of the terrestrial drill (Mandrillus leucophaeus) with six sympatric monkey species in Korup National Park, Cameroon, testing reports that Mandrillus associations are infrequent and transient. We conducted 3,284 km of trail walks for 12 months (February–June 2006; July 2007 to January 2008), recording species composition in 612 primate clusters. Using a Markov chain Monte Carlo test, we compared the observed frequency of dyadic associations against null models of “no association.” A novel conservative statistical approach which addresses possible dependence of observations close in time was also used, further strengthening confidence in our findings. Drills associated with all monkeys throughout the study period, and were with at least one other species (range 1–5) in half of the encounters. The association frequency of drills with red‐capped mangabeys (Cercocebus torquatus) was greater than expected by chance, which is interesting given the morphological adaptation of the Mandrillus‐Cercocebus clade for the exploitation of the same dietary niche, hard seeds. The difference we observed in the use of forest strata by drills and mangabeys may reflect a strategy to reduce food competition while in association. The nature and duration of observed drill associations varied. Although some associations seemed to be chance encounters, others lasted for hours with the involved species foraging together. Am. J. Primatol. 73:127–134, 2011. © 2010 Wiley‐Liss, Inc.     

The Distribution, Status, and Conservation Outlook of the Drill (Mandrillus leucophaeus) in Cameroon

The populations of many endangered species are becoming increasingly fragmented, and accurate, current information on the status of these subpopulations is essential for the design of effective conservation strategies within a human-dominated landscape. The drill (Mandrillus leucophaeus) is one of the most spectacular and endangered primates in Africa, yet up-to-date information on its distribution, population status, and conservation outlook is lacking. Cameroon has been estimated to encompass 80 % of the species’ range. We examined the distribution, population status, and conservation outlook for the drill throughout its historic range in Cameroon. To do this, we divided the historic range of the drill in Cameroon (46,000 km²) into 52 survey units along natural and manmade boundary features. Based on a series of field surveys in 2002–2009, village interviews, analysis of geospatial data, and bibliographical research, we assigned each survey unit a rank of 0–4 for 15 parameters indicative of current situation for drills, habitat suitability, and conservation outlook. We obtained direct evidence for the presence of drills in 16 of the 52 survey units, with those of Ejagham, Korup, Ebo, and Nta Ali receiving the highest index scores. We warn of local extirpations and increased isolation among drill populations due to loss of dispersal corridors, e.g., Douala Edea survey unit. In some cases drills persist in forest fragments within human-dominated landscapes, e.g., Kupe-Manenguba, but the species’ future is probably dependent on effective wildlife management in a handful of isolated strongholds where probability of long-term protection is higher, particularly in Korup National Park, Takamanda National Park, and the proposed Ebo National Park. Pressure from current and proposed large-scale commercial plantations, oil prospecting, logging, and the continual human population growth in this region means that a concerted conservation effort will be needed to safeguard the remaining drill habitat if the species is to survive in Cameroon.     

Giardia sp. and Cryptosporidium sp. infections in primates in fragmented and undisturbed forest in western Uganda

In June 2005, we collected 115 fecal samples from wild primates in western Uganda and examined them for Cryptosporidium sp. and Giardia sp. with the use of immunofluorescent antibody (IFA) detection. We sampled primates from an undisturbed forest in Kibale National Park and from 3 highly disturbed forest fragments outside the park. Of disturbed forest samples, red colobus (Pilocolobus tephrosceles) and red-tailed guenons (Cercopithecus ascanius) harbored species of Cryptosporidium or Giardia, but black-and-white colobus (Colobus guereza) did not. All primate samples from undisturbed forest were negative for both parasites. Seven of 35 (20%) red colobus and 1 of 20 red-tailed guenons (5%) from forest fragments were infected with either Cryptosporidium sp. or Giardia sp. The presence of Cryptosporidium and Giardia species in primates living in forest fragments, but not in primates in undisturbed forest, suggests that habitat disturbance may play a role in transmission or persistence of these pathogens.     

Conservation of Drill Populations in Bakossiland,Cameroon: Historical Trends and Current Status

Very little information is available on the current status of drill populations in Cameroon. We report on drill group sizes and status in Bakossiland, a mountainous area spanning 2000 km² in the Littoral and South West Provinces of southwestern Cameroon. Between 1970 and 2002 direct visual counts of drill groups (n = 105) yielded group size estimates ranging from 5 to 400 (mean ± S.E.M.; 93.1 ± 8.4). We encountered solitary adult male drills on 8 occasions. Groups were at all elevations (150–2000 m) in 5 habitat types: lowland, premontane, submontane and montane forests and montane savannah at 2000 m). Group sizes did not vary with respect to elevation, habitat type or season (wet and dry mo). However, over the past decade drills have been virtually hunted out of the Mwenzekong Mountains (Banyang Mbo Wildlife Sanctuary), and they are greatly reduced in the southern Bakossi forests of Mwendolengo, Edib Hills and Mungo River. The species became extinct in the Loum Forest Reserve in the late 1970s, and until recently was thought to have become extinct on Mount Mwanenguba. Since 1994 on Mount Kupe, the drill population has begun to recover, largely due to protection afforded by the Bakossi traditional chiefs. Traditional powers and values are still influential in the region. A new national park—Bakossi Mountains National Park— and associated Protected Areas are currently under gazettment. We discuss the effectiveness of conservation strategies in relation to the survival of drills in the area.     

Status of diurnal primate populations at the former settlement of a displaced village in Cameroon

To understand whether or not primate populations recover in areas of abandoned human settlements, data are required that allow for site‐specific comparison over time. Here, we present baseline information on the presence/absence and relative abundance of primate species at the abandoned settlement of the Ekundukundu village, relocated out of the Korup National Park in 2000. Between July 2007 and March 2008, 62 km of transects was surveyed for sightings and calls of primates. All eight species of diurnal primates reported in the KNP were confirmed: Cercopithecus nictitans ludio, C. mona, C. erythrotis camerunensis, C. pogonias, Procolobus pennantii preussi, Cercocebus torquatus, Mandrillus leucophaeus leucophaeus, and Pan troglodytes vellerosus. At old Ekundukundu, C. nictitans accounted for 65% of all primate group sightings. Overall, sighting frequency of primates (0.55 groups/km) was not significantly different from other park sectors surveyed by a previous observer (J. Linder) in 2004–2005. The data reported here will be useful in the long‐term monitoring of primate populations in regenerating forest habitats of earlier human settlements. Am. J. Primatol. 72:645–652, 2010. © 2010 Wiley‐Liss, Inc.     

Impact of Commercial Hunting on Monkey Populations in the Taï region,Côte d'Ivoire1

We studied the impact of hunting on monkey species in the Taï National Park and adjacent forests in Côte d'Ivoire. The average wild meat consumption per capita per year was assessed from market surveys and interviews. We determined that the amount of primate wild meat being extracted in the Taï National Park and surrounding forests was 249 t in 1999. Hunting pressure was the highest on the larger primate species such as red colobus, Procolobus badius, black and white colobus, Colobus polykomos, and the sooty mangabey, Cercocebus torquatus atys. Estimates of population densities were based on line transect surveys. The maximum annual production of each species was calculated using the Robinson and Redford model (1991) and assuming unhunted conditions. Comparing current harvest levels with the maximum sustainable yield suggests that harvest of red colobus monkeys (Procolobus badius) is sustainable, whereas current off‐take of the black and white colobus (Colobus polykomos), the sooty mangabey (Cercocebus atys), diana monkeys (Cercopithecus diana), and Campbell's monkeys (Cercopithecus campbelli) exceeds sustainability by up to three times.     

Preserving Preuss’s Red Colobus (<Emphasis Type="Italic">Piliocolobus preussi</Emphasis>): an Ethnographic Analysis of Hunting,Conservation, and Changing Perceptions of Primates in Ikenge-Bakoko,Cameroon

The futures of nonhuman primate species and human communities in shared landscapes rely on our ability to engage with and understand the complex histories and multiscalar aspects of human–animal relationships. We use the Critically Endangered Preuss’s red colobus (Piliocolobus preussi) as a case study to examine the important ways in which histories of multiscalar human–primate interactions play out in the village of Ikenge-Bakoko, Korup National Park, Cameroon. We contextualize ethnographic and catchment data from adult men (N?=?32) and women (N?=?31) within long-term diurnal primate monitoring datasets to better understand the relationships among hunting practices, local perceptions of diurnal primates, populations of P. preussi, and conservation management. Our data indicate a disconnect between local cultural definitions of “hunter” and Western assumptions as to the makeup and nature of this and other categories. We show that such contradictions can have negative outcomes for conservationists seeking to turn the science of establishing accurate off-take rates of prey species into practical management solutions. Using a single village as a focal point, we highlight the importance of an ethnoprimatological approach to understanding the intricate entanglements among conservation histories, subsistence strategies, and human and nonhuman primate lives. The application of ethnoprimatology is critical for twenty-first century primatologists who must navigate conservation concerns while also acknowledging and valuing the experiences of the human communities living alongside the primates we study.     

Do Nematode Infections of Red Colobus (Procolobus rufomitratus) and Black-and-White Colobus (Colobus guereza) on Humanized Forest Edges Differ from Those on Nonhumanized Forest Edges?

Forested edges, especially those that border humanized landscapes, provide opportunities for nonhuman primates and people to interact, and such interactions are predicted to alter disease dynamics. Given the rapid expansion of edge habitats globally, understanding changes occurring on edges is important in evaluating primate behavioral ecology and developing conservation plans. Our research investigates predictions concerning how gastrointestinal parasite and stress levels (cortisol) in red colobus and black-and-white colobus (Procolobus rufomitratus; Colobus guereza) in Kibale National Park, Uganda, differ between humanized and nonhumanized forest edges. We found Trichuris sp., an unidentified strongyle, and Strongyloides sp. in the fecal samples. Results did not generally support our expectation that humanized forest edges increase parasite infection and, counter to what we predicted, fecal cortisol did not differ between habitats, suggesting that proximity to edges and/or to humans did not result in increased stress. We conclude that broad habitat classifications, e.g., “humanized,” may be too general to identify consistent differences in parasite infection, as other factors, specific to the parasite (e.g., life cycled), host (e.g., immune systems strength), or environment (e.g., moisture level), likely also play important roles.     

Characterization of the fecal microbiome from non-human wild primates reveals species specific microbial communities

Background

Host-associated microbes comprise an integral part of animal digestive systems and these interactions have a long evolutionary history. It has been hypothesized that the gastrointestinal microbiome of humans and other non-human primates may have played significant roles in host evolution by facilitating a range of dietary adaptations. We have undertaken a comparative sequencing survey of the gastrointestinal microbiomes of several non-human primate species, with the goal of better understanding how these microbiomes relate to the evolution of non-human primate diversity. Here we present a comparative analysis of gastrointestinal microbial communities from three different species of Old World wild monkeys.

Methodology/Principal Findings

We analyzed fecal samples from three different wild non-human primate species (black-and-white colobus [Colubus guereza], red colobus [Piliocolobus tephrosceles], and red-tailed guenon [Cercopithecus ascanius]). Three samples from each species were subjected to small subunit rRNA tag pyrosequencing. Firmicutes comprised the vast majority of the phyla in each sample. Other phyla represented were Bacterioidetes, Proteobacteria, Spirochaetes, Actinobacteria, Verrucomicrobia, Lentisphaerae, Tenericutes, Planctomycetes, Fibrobacateres, and TM7. Bray-Curtis similarity analysis of these microbiomes indicated that microbial community composition within the same primate species are more similar to each other than to those of different primate species. Comparison of fecal microbiota from non-human primates with microbiota of human stool samples obtained in previous studies revealed that the gut microbiota of these primates are distinct and reflect host phylogeny.

Conclusion/Significance

Our analysis provides evidence that the fecal microbiomes of wild primates co-vary with their hosts, and that this is manifested in higher intraspecies similarity among wild primate species, perhaps reflecting species specificity of the microbiome in addition to dietary influences. These results contribute to the limited body of primate microbiome studies and provide a framework for comparative microbiome analysis between human and non-human primates as well as a comparative evolutionary understanding of the human microbiome.     

Habitat specificity and diversity of tree species in an African wet tropical forest

Niche differentiation with respect to habitat has been hypothesized to shape patterns of diversity and species distributions in plant communities. African forests have been reported to be relatively less diverse compared to highly diversed regions of the Amazonian or Southeast Asian forests, and might be expected to have less niche differentiation. We examined patterns of structural and floristic differences among five topographically defined habitats for 494 species with stems ≥1 cm dbh in a 50-ha plot in Korup National Park, Cameroon. In addition, we tested for species–habitat associations for 272 species (with more than 50 individuals in the plot) using Torus translation randomization tests. Tree density and basal area were lowest in areas with negative convexity, which contained streams or were inundated during rainy periods and highest in moist well-drained habitats. Species composition and diversity varied along the topographical gradient from low flat to ridge top habitats. The low depression and low flat habitats were characterized by high diversity and similar species composition, relative to slopes, high gullies and ridge tops. Sixty-three percent of the species evaluated showed significant positive associations with at least one of the five habitat types. The majority of associations were with low depressions (75 species) and the fewest with ridge tops (8 species). The large number of species–habitat associations and the pronounced contrast between low (valley) and elevated (ridgetop) habitats in the Korup plot shows that niche differentiation with respect to edaphic variables (e.g., soil moisture, nutrients) contributes to local scale tree species distributions and to the maintenance of diversity in African forests.     

Presence of arbuscular mycorrhizas in typically ectomycorrhizal host species from Cameroon and New Zealand

 Ectomycorrhizas (EcM) and arbuscular mycorrhizas (AM) were screened for in saplings of 14 EcM tree species from the N'Dupé and Korup National Park rainforests, SW Cameroon, belonging to Caesalpiniaceae and Uapacaceae. The pattern of EcM and AM colonisation of a dual mycorrhizal species from this rainforest (Uapaca staudtii, Uapacaceae) was compared with dual EcM/AM colonisation of Leptospermum scoparium (Myrtaceae) from New Zealand. Both species were collected in a range of habitats. EcM and AM colonisation differed among species in the Korup National Park rainforest: 12 species belonging to the Caesalpiniaceae (Amherstieae) were consistently EcM, and AM structures occurred occasionally in six of them; two other species belonging to Caesalpiniaceae (Afzelia bipindensis) and Uapacaceae (U. staudtii) were dual mycorrhizal with variable levels of colonisation by both EcM and AM fungi. EcM and AM dual colonisation varied with both habitat and identity of the partners. The presence of EcM fungi in most of the root samples of U. staudtii and a negative relationship between AM and EcM colonisation within the same root system suggested a greater EcM affinity of this species. In contrast, most root samples of L. scoparium were colonised by AM, but only a few by EcM. Genuine dual EcM/AM associations in root samples of U. staudtii where the two mycorrhizal types co-occurred could be attributed to an AM-EcM succession. However, differences between predicted and observed frequencies of genuine dual EcM/AM associations in several samples of both U. staudtii and L. scoparium indicated that other factors influenced dual EcM/AM associations. The results of this study showed the importance of the identity of the host species in determining the pattern of dual EcM and AM colonisation. Accepted: 18 September 1998     

Rarity and abundance in a diverse African forest

We censused all trees ≥1 cm dbh in 50 ha of forest in Korup National Park, southwest Cameroon, in the central African coastal forest known for high diversity and endemism. The plot included 329,519 individuals and 493 species, but 128 of those taxa remain partially identified. Abundance varied over four orders of magnitude, from 1 individual per 50 ha (34 species) to Phyllobotryon spathulatum, with 26,741 trees; basal area varied over six orders of magnitude. Abundance patterns, both the percentage of rare species and the dominance of abundant species were similar to those from 50-ha plots censused the same way in Asia and Latin America. Rare species in the Korup plot were much less likely to be identified than common species: 42% of taxa with <10 individuals in the plot were identified to species, compared to 95% of the abundant taxa. Geographic ranges for all identified species were gleaned from the literature and online flora. Thirteen of the plot species are known only from Korup National Park (all discovered during the plot census), and 39 are restricted to the Nigeria–Cameroon coastal zone. Contrary to expectation, species with narrow geographic ranges were more abundant in the plot than average. The small number of narrow endemics (11% of the species), many locally abundant, mitigates short-term extinction risk, either from demographic stochasticity or habitat loss.     

The cost of living in larger primate groups includes higher fly densities

Flies are implicated in carrying and mechanically transmitting many primate pathogens. We investigated how fly associations vary across six monkey species (Cercopithecus ascanius, Cercopithecus mitis, Colobus guereza, Lophocebus albigena, Papio anubis, and Piliocolobus tephrosceles) and whether monkey group size impacts fly densities. Fly densities were generally higher inside groups than outside them, and considering data from these primate species together revealed that larger groups harbored more flies. Within species, this pattern was strongest for colobine monkeys, and we speculate this might be due to their smaller home ranges, suggesting that movement patterns may influence fly–primate associations. Fly associations increase with group sizes and may thus represent a cost to sociality.

     

Primate abundance along five transect lines at Ngogo, Kibale National Park, Uganda

Using the line transect methods, I studied the primate density at Ngogo, Kibale National Park, Uganda for 18 months. Comparisons with other studies show that the population of red colobus monkeys (Procolobus rufomitratus) and blue monkeys (Cercopithecus mitis) is declining, whereas the populations of black-and-white colobus (Colobus guereza), red-tailed monkeys (Cercopithecus ascanius), grey-cheeked mangabeys (Lophocebus albigena), baboons (Papio anubis), and chimpanzees (Pan troglodytes) remain constant or slightly increase. In this paper, I compare data on density from this study to data from previous and recent censuses at Ngogo and with data from other sites in the Kibale forest to examine the stability of primate populations. Furthermore, I test the hypothesis that the changes in red colobus and blue monkey density are due to changes in the forest structure and abundance of their most selected feeding trees, and show that changes in forest composition cannot account for changes in their red colobus abundance, but that hunting by chimpanzees provides a reasonable explanation.     

High diversity of arbuscular mycorrhizal fungi in natural and anthropized sites of a Brazilian tropical dry forest (Caatinga)

Arbuscular mycorrhizal fungi (AMF) establish symbiotic associations with higher plants, which support the establishment and maintenance of plant communities across a range of environments, including those adversely affected by anthropogenic activity as well as natural sites. This study aimed at determining the diversity and distribution of AMF in areas of the tropical semi-arid region of Caatinga, Brazil, and compare areas in a naturally preserved state with anthropized sites. We characterized AMF communities in soil samples (n = 108), based on morphological taxonomy, at three sampling occasions and from six areas (typical Caatinga, extremely sandy Caatinga, stony Caatinga, rocky Caatinga, and two typical Caatinga areas that had been modified by human activities), at the National Park of Catimbau, Northeast Brazil. Eighty AMF species were recorded, with Glomus and Acaulospora predominating at all sites. There were significant differences in the composition of AMF communities between natural and anthropized sites, and among sampling occasions. Habitat-types also influenced AMF communities in Caatinga. Extensive tropical dry forest areas, such as the Catimbau National Park possess distinct niches, which maintain diverse AMF communities that are determined by anthropogenic activities, as well as vegetation types and environmental conditions.     

Social and hormonal mechanisms underlying male reproductive strategies in black howler monkeys (Alouatta pigra)

We investigated the social and hormonal mechanisms underlying male reproductive strategies in two multimale-multifemale groups of black howler monkeys (Alouatta pigra) during a 14-month study in Palenque National Park, Mexico. Fecal glucocorticoid (fGC) and androgen (fA) levels were analyzed for 343 fecal samples collected from 14 males during their presence in the study groups. Neither immigrating males nor resident males that remained in the group had elevated fGC and fA levels during 11 observed male migration events, suggesting that competition over group membership was not associated with variation in the fecal hormonal levels of males. Instead, fGC and fA levels were significantly higher in males who maintained a central position in the group and had almost exclusive access to fertile females than in other resident males. These “central” males were responsible for maintaining close spatial associations and cultivating strong affiliative relationships with cycling, sexually active females but not with noncycling, sexually inactive females. “Noncentral” males did not form strong social relationships with either cycling or noncycling females and had no or very few mating opportunities. Our findings suggest that male black howler monkeys compete nonaggressively by fostering relationships with cycling females and that the elevated fGC levels of central males may be indicative of the social challenges involved in their indirect competition.     

The formation of red colobus-diana monkey associations under predation pressure from chimpanzees.

It is generally assumed that most primates live in monospecific or polyspecific groups because group living provides protection against predation, but hard evidence is scarce. We tested the antipredation hypothesis with observational and experimental data on mixed-species groups of red colobus (Procolobus badius) and diana monkeys (Cercopithecus diana) in the Taï National Park, Ivory Coast. Red colobus, but not diana monkeys, are frequently killed by cooperatively hunting chimpanzees. Association rates peaked during the chimpanzees'' hunting season, as a result of changes in the behaviour of the red colobus. In addition, playbacks of recordings of chimpanzee sounds induced the formation of new associations and extended the duration of existing associations. No such effects were observed in reaction to control experiments and playbacks of leopard recordings.     

Market Hunting in the Taï Region,Côte d’Ivoire and Implications for Monkey Populations

We studied the effect of market hunting on primate species in the Taï National Park and adjacent forests in Côte d’Ivoire. We assessed the impact of hunting by comparing the calculated maximal reproduction rate with the current off-take rate. We assessed the average bushmeat consumption per capita/per year from weekly investigations on bushmeat available in 88 bushmeat restaurants and markets over a 12-mo period in 1999. We derived data on preferences for particular game species from interviews of 162 bushmeat consumers, 25 subsistence hunters and 3 groups of professional hunters. Hunting pressure was highest on the larger primate species such red colobus (Procolobus badius), black- and- white colobus (Colobus polykomos) and sooty mangabeys (Cercocebus atys). The amount of primate bushmeat extracted from the Taï National Park and surrounding forests was 249,229 kg in 1999. We estimated population densities using line transect surveys. By referring to current population densities we calculated the maximum production of each species using the Robinson Redford model (2001) and assuming unhunted conditions. A comparison of current harvest levels with maximum production suggests that harvest of Procolobus badius is sustainable, whereas current off-take of Colobus polykomos, Cercocebus atys, Cercopithecus diana (diana monkey) and C. campbelli (Campbell’s monkeys) exceeds sustainability by ≤3 times. We recommend that wildlife managers promote programs that encourage the production of domestic animals as a substitute for wild meat.     

Dyadic associations of red Colobus and diana monkey groups in the Taï National Park, Ivory Coast

Members of the genus Colobus have been observed to associate frequently with Cercopithecus monkeys in several African sites. In the Taï National Park, Ivory Coast, one group of western red colobus was found to be in association with one particular group of diana monkeys more than could be expected by chance (Holenweg et al., 1996). We show that dyadic association is not an idiosyncrasy of these two groups, but rather a pattern that is general for our study site. All five red colobus groups we studied were closely associated with diana monkeys during more than 60% of the time. Four groups had one particular diana partner group, the fifth two different partners. Apart from the red colobus, three more primate species, the olive colobus, Campbell's monkey, and the lesser spot-nosed monkey, were also strongly attracted to diana monkeys.     

A skew in poo: Biases in primate fecal isotope analysis and recommendations for standardized sample preparation

Feces are a treasure trove in the study of animal behavior and ecology. Stable carbon and nitrogen isotope analysis allows to assess the dietary niches of elusive primate species and primate breastfeeding behavior. However, some fecal isotope data may unwillingly be biased toward the isotope ratios of undigested plant matter, requiring more consistent sample preparation protocols. We assess the impact of this potential data skew in 114 fecal samples of wild bonobos (Pan paniscus) by measuring the isotope differences (Δ¹³C, Δ¹⁵N) between bulk fecal samples containing larger particles (>1 mm) and filtered samples containing only small particles (<1 mm). We assess the influence of fecal carbon and nitrogen content (ΔC:N) and sample donor age (subadult, adult) on the resulting Δ¹³C, Δ¹⁵N values (n = 228). Additionally, we measure the isotope ratios in three systematically sieved fecal samples of chimpanzees (Pan troglodytes verus), with particle sizes ranging from 20 μm to 8 mm (n = 30). We found differences in fecal carbon and nitrogen content, with the smaller fecal fraction containing more nitrogen on average. While the Δ¹³C values were small and not affected by age or ΔC:N, the Δ¹⁵N values were significantly influenced by fecal ΔC:N, possibly resulting from the differing proportions of undigested plant macroparticles. Significant relationships between carbon stable isotope ratios (δ¹³C) values and %C in large fecal fractions of both age groups corroborated this assessment. Δ¹⁵N values were significantly larger in adults than subadults, which should be of concern in isotope studies comparing adult females with infants to assess breastfeeding. We found a random variation of up to 3.0‰ in δ¹³C and 2.0‰ in nitrogen stable isotope ratios within the chimpanzee fecal samples separated by particle sizes. We show that particle size influences isotope ratios and propose a simple, cost-effective filtration method for primate feces to exclude larger undigested food particles from the analysis, which can easily be adopted by labs worldwide.