Skip to main content
SpringerLink
Log in

Non-dietary analytical features of chimpanzee scats

  • Original Article
  • Published:
Primates Aims and scope Submit manuscript

Abstract

Non-dietary aspects of ape scats such as scat weight and diameter are correlated with age and sex of defaecator for gorillas and orangutans. Defaecation rates of primates, including apes, illuminate their role as primary seed dispersers. We assess if non-dietary features of scats for East African chimpanzees (Pan troglodytes schweinfurthii) reveal such insights for members of the Kanyawara community in Kibale National Park, Uganda. Our objective is to see if such data yield useful perspectives for future census work on unhabituated chimpanzees, that is, what can scats tell us about a wild study population, beyond diet? We followed ten adults from this community, as well as travelling parties, comparing observed vs. unobserved defaecations, and collected data on scat weight and dimensions, defaecation rate, scat encounter rate, and interval between defaecations. Few non-dietary features of chimpanzee scats significantly differentiated sex or age of the defaecator, but total scat length and height distinguished adults from juveniles/infants. Defaecation rates and distance travelled were similar for adult males and females, indicating the importance of both sexes as potential primary seed dispersers. Observed travelling parties vs. non-observed travelling parties yielded similar data, indicating the potential to assess party size from scat encounter rates over a set distance. We provide detailed measurements of scat dimensions for this ape taxon which previously have been lacking. This research builds upon prior work by recording more in-depth data for focal subjects and travelling parties on defaecation and scat encounter rates. The findings presented should assist in the interpretation of scat data when censusing unhabituated chimpanzees.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Similar content being viewed by others

References

  • Anderson JR, Hubert-Brierre X, McGrew WC (2016) Reflections in the rainforest: full-length mirrors facilitate behavioral observations of unhabituated, wild chimpanzees. Primates. doi:10.1007/s10329-016-0574-7

    Google Scholar 

  • Ashford RW, Reid GD, Wrangham RW (2000) Intestinal parasites of the chimpanzee Pan troglodytes in Kibale Forest, Uganda. Ann Trop Med Parasitol 94:173–179

    Article  CAS  PubMed  Google Scholar 

  • Bădescu I, Katzenberg MA, Watts DP, Sellen DW (2016) A novel fecal stable isotope approach to determine the timing of age-related feeding transitions in wild infant chimpanzees. Am J Primatol. doi:10.1002/ajpa.23116

    Google Scholar 

  • Basabose AK (2002) Diet composition of chimpanzees inhabiting the montane forest of Kahuzi, Democratic Republic of Congo. Am J Primatol 58:1–21

    Article  PubMed  Google Scholar 

  • Beaune D, Bretagnolle F, Bollache L, Hohmann G, Surbeck M, Fruth B (2013) Seed dispersal strategies and the threat of defaunation in a Congo forest. Biodivers Conserv 22:225–238

    Article  Google Scholar 

  • Bertolani MP (2013) Ranging and travelling patterns of wild chimpanzees at Kibale: a GIS approach. PhD thesis, University of Cambridge, Cambridge

  • Bezanson M, Stowe R, Watts SM (2013) Reducing the ecological impact of field research. Am J Primatol 75:1–9

    Article  PubMed  Google Scholar 

  • Blumenthal S, Chritz K, Rothman J, Cerling T (2012) Detecting intra-annual dietary variability in wild mountain gorillas by stable isotope analysis of feces. Proc Natl Acad Sci USA 109:21277–21282

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Boyer-Ontl KM, Pruetz JD (2014) Giving the forest eyes: the benefits of using camera traps to study unhabituated chimpanzees (Pan troglodytes verus) in southeastern Senegal. Int J Primatol. doi:10.1007/s10764-014-9783-3

    Google Scholar 

  • Bradley BJ, Doran-Sheehy DM, Vigliant L (2008) Genetic identification of elusive animals: re-evaluating tracking and nesting data for wild western gorillas. J Zool 275:333–340

    Article  Google Scholar 

  • Chancellor RL, Langergraber Ramirez S, Rundus AS, Vigilant L (2012) Genetic sampling of unhabituated chimpanzees (Pan troglodytes schweinfurthii) in Gishwati Forest Reserve, an isolated forest fragment in western Rwanda. Int J Primatol. doi:10.1007/s10764-012-9591-6

    Google Scholar 

  • Chapman CA (1995) Primate seed dispersal: coevolution and conservation implications. Evol Anthropol 4:74–82

    Article  Google Scholar 

  • Chaves OM, Stoner KE, Arroyo-Rodríguez V (2012) Differences in diet between spider monkey groups living in forest fragments and continuous forest in Mexico. Biotropica 44:105–113

    Article  Google Scholar 

  • Clifford SL, Anthony NM, Bawe-Johnson M, Abernethy KA, Tutin CE, White LJ, Bermejo M, Goldsmith ML, McFarland K, Jeffery KJ, Bruford MW (2004) Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla). Mol Ecol 13:1551–1565

    Article  CAS  PubMed  Google Scholar 

  • Constable JL, Ashley MV, Goodall J, Pusey AE (2001) Noninvasive paternity assignment in Gombe chimpanzees. Mol Ecol 10:1279–1300

    Article  CAS  PubMed  Google Scholar 

  • Doran DM (1996) Comparative positional behavior of the African apes. In: McGrew WC, Marchant LF, Nishida T (eds) Great ape societies. Cambridge University Press, Cambridge, pp 213–224

    Chapter  Google Scholar 

  • Doran DM, McNeilage A, Greer D, Bocian C, Mehlman P, Shah N (2002) Western lowland gorilla diet and resource availability: new evidence, cross-site comparisons, and reflections on indirect sampling methods. Am J Primatol 58:91–116

    Article  PubMed  Google Scholar 

  • Doran-Sheehy D, Mongo P, Lodwick J, Conklin-Brittain NL (2009) Male and female western gorilla diet: preferred foods, use of fallback resources, and implications for ape versus old world monkey foraging strategies. Am J Phys Anthropol 140:727–738

    Article  CAS  PubMed  Google Scholar 

  • Emery Thompson M (2005) Reproductive endocrinology of wild female chimpanzees (Pan troglodytes schweinfurthii): methodological considerations and the role of hormones in sex and conception. Am J Primatol 67:137–158

    Article  PubMed  Google Scholar 

  • Estrada A, Coates-Estrada R (1991) Howler monkeys (Alouatta palliata), dung beetles (Scarabaeidae) and seed dispersal: ecological interactions in the tropical rain forest of Los Tuxtlas, Mexico. J Trop Ecol 7:459–474

    Article  Google Scholar 

  • Field D, Chemnick L, Robbins M, Garner K, Ryder O (1998) Paternity determination in captive lowland gorillas and orangutans and wild mountain gorillas by microsatellite analysis. Primates 39:199–209

    Article  Google Scholar 

  • Furuichi T, Inagaki H, Anggoue-Ovono S (1997) Population density of chimpanzees and gorillas in the Petit Loango Reserve, Gabon: employing a new method to distinguish between nests of the two species. Int J Primatol 18:1029–1046

    Article  Google Scholar 

  • Galdikas BM (1982) Orang utans as seed dispersers at Tanjung Puting, Central Kalimantan: implications for conservation. In: de Boer LEM (ed) The orang utan: its biology and conservation. Dr W Junk, The Hague, pp 285–298

    Google Scholar 

  • Galdikas BMF (1988) Orangutan diet, range, and activity at Tanjung Puting, Central Borneo. Int J Primatol. doi:10.1007/BF02740195

    Google Scholar 

  • Goosens B, Anthony N, Jeffery K, Johnson-Bawe M, Bruford MW (2003) Collection, storage and analysis of non-invasive genetic material in primate biology. In: Setchell JM, Curtis DJ (eds) Field and laboratory methods in primatology. A practical guide Cambridge University Press, Cambridge, pp 295–308

    Chapter  Google Scholar 

  • Gross-Camp ND, Masozera M, Kaplin BA (2009) Chimpanzee seed dispersal quantity in a tropical montane forest of Rwanda. Am J Primatol 71:1–11

    Article  Google Scholar 

  • Harcourt AH, Fossey D (1981) The Virunga gorillas: decline of an ‘island’ population. Afr J Ecol 19:83–97

    Article  Google Scholar 

  • Hernandez-Aguilar RA (2009) Chimpanzee nest distribution and site reuse in a dry habitat: implications for early hominin ranging. J Hum Evol 57:350–364

    Article  PubMed  Google Scholar 

  • Inoue E, Akomo-Okoue EF, Ando C, Iwata Y, Judai M, Fujita S, Hongo S, Nze-Nkogue C, Inoue-Murayama M, Yamagiwa J (2013) Male genetic structure and paternity in western lowland gorillas (Gorilla gorilla gorilla). Am J Phys Anthropol 151:583–588

    Article  PubMed  Google Scholar 

  • Johnson AE, Knott CD, Pamungkas B, Pasaribu M, Marshall AJ (2005) A survey of the orangutan (Pongo pygmaeus wurmbii) population in and around Gunung Palung National Park, West Kalimantan, Indonesia based on nest counts. Biol Conserv 121:495–507

    Article  Google Scholar 

  • Kahlenberg SM, Emery Thompson M, Wrangham RW (2008) Female competition over core areas in Pan troglodytes schweinfurthii, Kibale National Park, Uganda. Int J Primatol. doi:10.1007/s10764-008-9276-3

    Google Scholar 

  • Kalan AK, Piel AK, Mundry R, Wittig RM, Boesch C, Kühl HS (2016) Passive acoustic monitoring reveals group ranging and territory use: a case study of wild chimpanzees (Pan troglodytes). Front Zool 13:34

    Article  PubMed  PubMed Central  Google Scholar 

  • Kanthaswamy S, Smith DG (2002) Population subdivision and gene flow among wild orangutans. Primates 43:315–327

    Article  PubMed  Google Scholar 

  • Koops K (2011) Chimpanzees in the Seringbara region of the Nimba Mountains. In: Hulme T, Sugiyama Y, Matsuzawa T (eds) The chimpanzees of Bossou and Nimba. Springer, Tokyo, pp 277–287

    Chapter  Google Scholar 

  • Krief S, Huffman MA, Sévenet T, Guillot J, Bories C, Hladik CM, Wrangham RW (2005) Noninvasive monitoring of the health of Pan troglodytes schweinfurthii in the Kibale National Park, Uganda. Int J Primatol 26:467–490

    Article  Google Scholar 

  • Kuehl HS, Todd A, Boesch C, Walsh PD (2007) Manipulating decay time for efficient large-mammal density estimation: gorillas and dung height. Ecol Appl 17:2403–2414

    Article  PubMed  Google Scholar 

  • Lambert JE (2002) Exploring the link between animal frugivory and plant strategies: the case of primate fruit processing and post-dispersal seed fate. In: Levey D, Silva W, Galetti M (eds) Seed dispersal and frugivory: ecology, evolution and conservation. CAB International, Wallingford, pp 369–375

    Google Scholar 

  • Linder JM, Oates JF (2011) Differential impact of bushmeat hunting on monkey species and implications for primate conservation in Korup National Park, Cameroon. Biol Conserv 144:738–774

    Article  Google Scholar 

  • Lucas PW, Corlett RT (1998) Seed dispersal by long-tailed macaques. Am J Primatol 45:29–44

    Article  CAS  PubMed  Google Scholar 

  • Martin P, Bateson P (2007) Measuring behaviour. An introductory guide, 3rd edn. Cambridge University Press, Cambridge

    Book  Google Scholar 

  • McConkey KR, Chivers DJ (2007) Influence of gibbon ranging patterns on seed dispersal distance and deposition site in a Bornean forest. J Trop Ecol 23:269–275

    Article  Google Scholar 

  • McFarland KL (2007) Ecology of Cross River gorillas (Gorilla gorilla diehli) on Afi Mountain, Cross River State, Nigeria. PhD, City University of New York

  • McGrew WC, Baldwin PJ, Tutin CEG (1988) Diet of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik, Senegal: I. Composition. Am J Primatol 16:213–226

    Article  Google Scholar 

  • McGrew WC, Ensminger AL, Marchant LF, Pruetz JD, Vigilant L (2004) Genotyping aids field study of unhabituated wild chimpanzees. Am J Primatol 63:87–93

    Article  CAS  PubMed  Google Scholar 

  • McNeilage A, Robbins MM, Gray M, Olupot W, Babaasa D, Bitariho R, Kasangaki A, Rainer H, Asuma S, Mugiri G, Baker J (2006) Census of the mountain gorilla Gorilla beringei beringei population in Bwindi Impenetrable National Park, Uganda. Oryx 40:419–427

    Article  Google Scholar 

  • Morgan BJ (2007) Group size, density and biomass of large mammals in the Réserve de Faune du Petit Loango, Gabon. Afr J Ecol 45:508–518

    Article  Google Scholar 

  • Morgan D, Sanz C, Onononga JR, Strindberg S (2006) Ape abundance and habitat use in the Goualougo Triangle, Republic of Congo. Int J Primatol 27:147–179

    Article  Google Scholar 

  • Morin PA, Wallis J, Moore JJ, Woodruff DS (1994) Paternity exclusion in a community of wild chimpanzees using hypervariable simple sequence repeats. Mol Ecol 3:469–478

    Article  CAS  PubMed  Google Scholar 

  • Morin PA, Chambers KE, Boesch C, Vigilant L (2001) Quantitative polymerase chain reaction analysis of DNA from non-invasive samples for accurate microsatelite genotyping of wild chimpanzees (Pan troglodytes verus). Mol Ecol 10:1835–1844

    Article  CAS  PubMed  Google Scholar 

  • Muehlenbein MP, Ancrenaz M, Sakong R, Ambu L, Prall S, Fuller G, Raghanti MA (2012) Ape conservation physiology: fecal glucocorticoid responses in wild Pongo pygmaeus morio following human visitation. PLoS One. doi:10.1371/journal.pone.0033357

    PubMed  PubMed Central  Google Scholar 

  • Murray CM, Heintz MR, Lonsdorf EV, Parr LA, Santymire RM (2013) Validation of a field technique and characterization of fecal glucocorticoid metabolite analysis in wild chimpanzees (Pan troglodytes). Am J Primatol 75:57–64

    Article  PubMed  Google Scholar 

  • Orkin JD, Yang Y, Yang C, Yu DW, Jiang X (2016) Cost-effective scat-detection dogs: unleashing a powerful new tool for international mammalian conservation biology. Sci Rep. doi:10.1038/srep34758

    PubMed  PubMed Central  Google Scholar 

  • Phillips CA (2012) Chimpanzee diet: analyses at macroscopic, microscopic and molecular level. PhD thesis, University of Cambridge, Cambridge

  • Phillips CA, McGrew WC (2013) Identifying species in chimpanzee (Pan troglodytes) faeces: a methodological lost cause? Int J Primatol 34:792–807

    Article  Google Scholar 

  • Phillips CA, McGrew WC (2014) Macroscopic inspection of ape feces: what’s in a quantification method? Am J Primatol 76:539–550

    Article  PubMed  Google Scholar 

  • Phillips CA, O’Connell T (2016) Fecal carbon and nitrogen isotopic analysis as an indicator of diet in Kanyawara chimpanzees, Kibale National Park, Uganda. Am J Phys Anthropol 161:685–697

    Article  PubMed  Google Scholar 

  • Pontzer H, Wrangham RW (2006) Ontogeny of ranging in wild chimpanzees. Int J Primatol 27:295–309

    Article  Google Scholar 

  • Poulsen JP, Clark CJ, Smith TB (2001) Seed dispersal by a diurnal primate community in the Dja Reserve, Cameroon. J Trop Ecol 17:787–808

    Article  Google Scholar 

  • Remis MJ (2000) Initial studies on the contributions of body size and gastrointestinal passage rates to dietary flexibility among gorillas. Am J Phys Anthropol 112:171–180

    Article  CAS  PubMed  Google Scholar 

  • Rimbach R, Link A, Heistermann M, Gómez-Posada C, Galvis N, Heymann EW (2013) Effects of logging, hunting, and forest fragment size on physiological stress levels of two sympatric ateline primates in Colombia. Conserv Physiol 1:1–11

    Article  Google Scholar 

  • Rogers ME, Voysey BC, McDonald KE, Parnell RJ, Tutin CEG (1998) Lowland gorillas and seed dispersal: the importance of nest sites. Am J Primatol 45:45–68

    Article  CAS  PubMed  Google Scholar 

  • Rogers ME, Abernethy K, Bermejo M, Cipolletta C, Doran D, McFarland K, Nishihara T, Remis M, Tutin CEG (2004) Western gorilla diet: a synthesis from six sites. Am J Primatol 64:173–192

    Article  PubMed  Google Scholar 

  • Schaller GB (1963) The mountain gorilla: ecology and behavior. University of Chicago Press, Chicago

    Google Scholar 

  • Shutt K, Heistermann M, Kasim A, Todd A, Kalousova B, Profosouva I, Petrzelkova K, Fuh T, Dicky JF, Bopalanzognako JB, Setchell JM (2014) Effects of habituation, research and ecotourism on faecal glucocorticoid metabolites in wild western lowland gorillas: implications for conservation management. Biol Conserv 172:72–79

    Article  Google Scholar 

  • Smith RJ, Jungers WL (1997) Body mass in comparative primatology. J Hum Evol 32:523–559

    Article  CAS  PubMed  Google Scholar 

  • Stanford CB, Nkurunungi JB (2003) Behavioral ecology of sympatric chimpanzees and gorillas in Bwindi Impenetrable National Park, Uganda: diet. Int J Primatol 24:901–919

    Article  Google Scholar 

  • Sunderland-Groves JL, Maisels F, Ekinde A (2003) Surveys of the Cross River gorilla and chimpanzee populations in Takamanda Forest Reserve, Cameroon. In: Comiskey JA, Sunderland TCH, Sunderland-Groves JL (eds) Takamanda: the biodiversity of an African rainforest. Smithsonian Institution Press, Washington, pp 129–140

    Google Scholar 

  • Takenoshita Y, Yamagiwa J (2008) Estimating gorilla abundance by dung count in the northern part of Moukalaba-Doudou National Park, Gabon. Afr Study Monogr 39:41–54

    Google Scholar 

  • Thalmann O, Fischer A, Lankester F, Pääbo S, Vigilant L (2007) The complex evolutionary history of gorillas: insights from genomic data. Mol Biol Evol 24:146–158

    Article  CAS  PubMed  Google Scholar 

  • Todd AF, Kuehl HS, Cipolletta C, Walsh PD (2008) Using dung to estimate gorilla density: modelling dung production rate. Int J Primatol 29:549–563

    Article  Google Scholar 

  • Travis DA, Lonsdorf EV, Mlengeya T, Raphael J (2008) A science-based approach to managing disease risks for ape conservation. Am J Primatol 70:745–750

    Article  PubMed  Google Scholar 

  • Tsuji Y, Yangozene K, Sakamaki T (2010) Estimation of seed dispersal distance by the bonobo, Pan paniscus, in a tropical forest in Democratic Republic of Congo. J Trop Ecol 26:115–118

    Article  Google Scholar 

  • Tutin CEG (1996) Ranging and social structure of lowland gorillas in the Lopé Reserve, Gabon. In: McGrew WC, Marchant LF, Nishida T (eds) Great ape societies. Cambridge University Press, Cambridge, pp 58–70

    Chapter  Google Scholar 

  • Tutin CEG, Fernandez M (1985) Foods consumed by sympatric populations of Gorilla g. gorilla and Pan t. troglodytes in Gabon: some preliminary data. Int J Primatol 6:27–43

    Article  Google Scholar 

  • Tutin CEG, Fernandez M (1993) Faecal analysis as a method of describing diets of apes: examples from sympatric gorillas and chimpanzees at Lopé, Gabon. Tropics 2:189–197

    Article  Google Scholar 

  • Voysey BC, McDonald KE, Rogers ME, Tutin CEG, Parnell R (1999) Gorillas and seed dispersal in the Lopé Reserve, Gabon. I: gorilla acquisition by trees. J Trop Ecol 15:23–38

    Article  Google Scholar 

  • Williamson EA, Feistner ATC (2011) Habituating primates: processes, techniques, variables and ethics. In: Setchell J, Curtis DJ (eds) Field and laboratory methods in primatology: a practical guide. Cambridge University Press, Cambridge, pp 25–39

    Google Scholar 

  • Williamson EA, Tutin CEG, Rogers ME, Fernandez M (1990) Composition of the diet of lowland gorillas at Lopé in Gabon. Am J Primatol 21:265–277

    Article  Google Scholar 

  • Wrangham RW (1995) Relationship of chimpanzee leaf-swallowing to a tapeworm infection. Am J Primatol 37:297–304

    Article  Google Scholar 

  • Wrangham RW, Chapman CA, Chapman LJ (1994) Seed dispersal by forest chimpanzees in Uganda. J Trop Ecol 10:355–368

    Article  Google Scholar 

  • Yamagiwa J, Maruhashi T, Yumoto T, Mwanza N (1996) Dietary and ranging overlap in sympatric gorillas and chimpanzees in Kahuzi-Biega National Park, Zaire. In: McGrew WC, Marchant LF, Nishida T (eds) Great ape societies. Cambridge University Press, Cambridge, pp 82–98

    Chapter  Google Scholar 

Download references

Acknowledgements

Research funding was awarded by the Board of Graduate Studies of the University of Cambridge, Murray Edwards College, Ridgeway-Venn Travel Fund, Sir Richard Stapely Education Trust, and Leverhulme Trust. Long-term work at Kanyawara was supported by the U.S. National Science Foundation (Grants 9807448, 0416125, and 1355014), the Leakey Foundation, the National Geographic Society, the Getty Foundation and the Wenner-Gren Foundation for Anthropological Research. All applicable international, national, and institutional guidelines for the care and use of animals were followed. All procedures performed in studies involving animals were in accordance with the ethical standards of the institution or practice at which the studies were conducted. Permission to live and conduct research in Kibale National Park, Uganda, was given by the Uganda Wildlife Authority, Uganda Council for Science and Technology, Makerere University Biological Field Station, and Martin Muller and Emily Otali of the Kibale Chimpanzee Project. The authors thank all staff of the Kibale Chimpanzee Project for their assistance during sample collection and Michael Muehlenbein and an anonymous reviewer for their constructive comments.

Author information

Author notes

  1. Caroline A. Phillips

    Present address: Evolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, Wits, 2050, South Africa

Authors and Affiliations

  1. Department of Archaeology and Anthropology, University of Cambridge, 13A Fitzwilliam Street, Cambridge, CB2 1QH, UK

    Caroline A. Phillips & William C. McGrew

  2. Department of Human Evolutionary Biology, Peabody Museum, Harvard University, 11 Divinity Avenue, Cambridge, MA, 02138, USA

    Richard W. Wrangham

  3. School of Psychology and Neuroscience, University of St. Andrews, St. Andrews, KY16 9JU, UK

    William C. McGrew

Authors
  1. Caroline A. Phillips
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Richard W. Wrangham
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. William C. McGrew
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Caroline A. Phillips.

Electronic supplementary material

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Phillips, C.A., Wrangham, R.W. & McGrew, W.C. Non-dietary analytical features of chimpanzee scats. Primates 58, 393–402 (2017). https://doi.org/10.1007/s10329-017-0606-y

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10329-017-0606-y

Keywords

Search

Navigation