Competition for food is often a cost associated with living in a group, and can occur in an indirect (scramble) or direct (contest) form. We investigated feeding competition in a supergroup of Rwenzori black-and-white colobus monkeys (Colobus angolensis ruwenzorii) in Rwanda, with the aim of establishing whether freedom from scramble competition allows these monkeys to form supergroups. We used the patch depletion method, measuring intake rate coupled with movement rate, to assess if food patches become depleted over the occupancy period. Resource depletion was evident when the colobus fed on young leaves, but not when feeding on mature leaves. Scramble competition was inferred from a negative correlation between group size and change in intake rate over patch occupancy. Between-group contest competition was inferred from displacement from patches. Although feeding competition exists for select resources, limited competition for mature leaves may enable Rwenzori colobus to live in a supergroup of hundreds of individuals in this montane forest.
Purchase
Buy instant access (PDF download and unlimited online access):
Institutional Login
Log in with Open Athens, Shibboleth, or your institutional credentials
Personal login
Log in with your brill.com account
Alcock, J. (1980). Natural selection and the mating systems of solitary bees. — Am. Sci. 68: 146-153.
Alexander, R.D. (1974). The evolution of social behavior. — Annu. Rev. Ecol. Syst. 5: 325-383.
Altizer, S., Nunn, C.L., Thrall, P.H., Gittleman, J.L., Antonovics, J., Cunningham, A.A., Dobson, A.P., Ezenwa, V., Jones, K.E., Pedersen, A.B. & Poss, M. (2003). Social organization and parasite risk in mammals: integrating theory and empirical studies. — Annu. Rev. Ecol. Evol. Syst. 34: 517-547.
Altmann, S.A. (1974). Baboons, space, time, and energy. — Am. Zool. 14: 221-248.
Bates, D., Maechler, M., Bolker, B., Walker, S., Christensen, R.H.B., Singmann, H., Dai, B., Eigen, C. & Rcpp, L. (2015). R package lme4. — Available online http://CRAN.R-project.org/package=lme4.
Bocian, C.M. (1997). Niche separation of black-and-white colobus monkeys (Colobus angolensis and C. guereza) in the Ituri forest. — Doctoral dissertation, The City University of New York, New York, NY.
Brown, C.R. & Brown, M.B. (1986). Ectoparasitism as a cost of coloniality in cliff swallows (Hhrundo pyrrfumota). — Ecology 67: 1206-1218.
Chapman, C.A. & Chapman, L.J. (2000a). Constraints on group size in red colobus and red-tailed guenons: examining the generality of the ecological constraints model. — Int. J. Primatol. 21: 565-585.
Chapman, C.A. & Chapman, L.J. (2000b). Determinants of group size in primates: the importance of travel costs. — In: On the move: how and why animals travel in groups (Boinski, S. & Garber, P.A., eds). The University of Chicago Press, Chicago, IL, p. 24-42.
Chapman, C.A. & Chapman, L.J. (2002). Foraging challenges of red colobus monkeys: influence of nutrients and secondary compounds. — Comp. Biochem. Physiol. 133A: 861-875.
Chapman, C.A., Chapman, L.J., Wrangham, R., Hunt, K., Gebo, D. & Gardner, L. (1992). Estimators of fruit abundance of tropical trees. — Biotropica 24: 527-531.
Chapman, C.A., Wrangham, R.W. & Chapman, L.J. (1995). Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups. — Behav. Ecol. Sociobiol. 36: 69-70.
Charnov, E.L. (1976). Optimal foraging, the marginal value theorem. — Theor. Popul. Biol. 9: 129-136.
Clutton-Brock, T. (2016). Mammal societies. — Wiley, Oxford.
Clutton-Brock, T.H. (1975). Feeding behavior of red colobus and back and white colobus in East-Africa. — Folia Primatol. 23: 165-207.
Clutton-Brock, T.H. & Harvey, P.H. (1977). Primate ecology and social organisation. — J. Zool. Soc. Lond. 183: 1-39.
Clutton-Brock, T.H., Gaynor, D., Mcilrath, G.M., Maccoll, A.D.C., Kansky, R., Chadwick, P., Manser, M., Skinner, J.D. & Brotherton, P.N.M. (1999). Predation, group size and mortality in a cooperative mongoose, Suricata suricatta. — J. Anim. Ecol. 68: 672-683.
Colmenares, F., Esteban, M. & Zaragoza, F. (2006). One-male units and clans in a colony of hamadryas baboons (Papio hamadryas hamadryas): effect of male number and clan cohesion on feeding success. — Am. J. Primatol. 68: 21-37.
Cross, P.C., Lloyd-Smith, J.O. & Getz, W.M. (2005). Disentangling association patterns in fission-fusion societies using African buffalo as an example. — Anim. Behav. 69: 499-506.
Dasilva, G.L. (1989). The ecology of the western black and white colobus (Colobus polykomos polykomos Zimmerman 1780) on a riverine island in southeastern Sierra Leone. — University of Oxford.
Dasilva, G.L. (1994). Diet of Colobus polykomos on Tiwai Island: selection of food in relation to its seasonal abundance and nutritional quality. — Int. J. Primatol. 15: 655-680.
Djègo-Djossou, S., Koné, I., Fandohan, A.B., Djègo, J.G., Huynen, M.C. & Sinsin, B. (2015). Habitat use by white-thighed colobus in the Kikélé Sacred Forest: activity budget, feeding ecology and selection of sleeping trees. — Primate Conservation. 29: 97-105.
Dunbar, R.I.M. & Dunbar, E.P. (1974). Ecology and population dynamics of Colobus guereza in Ethiopia. — Folia Primatol. 21: 188-208.
Dunham, N.T. (2011). Coping with forest fragmentation: a comparison of Colobus angolensis palliatus dietary diversity and behavioral plasticity in the East Sagara Forest, Tanzania. — In. Illinois Wesleyan University.
Dunham, N.T. (2015). Ontogeny of positional behavior and support use among Colobus angolensis palliatus of the Diani Forest, Kenya. — Primates 56: 183-192.
Dunham, N.T. (2017). Feeding ecology and dietary flexibility of Colobus angolensis palliatus in relation to habitat disturbance. — Int. J. Primatol. 38: 553-571.
Fashing, P.J. (2001a). Activity and ranging patterns of guerezas in the Kakamega forest: intergroup variation and implications for intragroup feeding competition. — Int. J. Primatol. 22: 549-577.
Fashing, P.J. (2001b). Male and female strategies during intergroup encounters in guerezas (Colobus guereza): evidence for resource defense mediated through males and a comparison with other primates. — Behav. Ecol. Sociobiol. 50: 219-230.
Fashing, P.J. (2006). African colobine monkeys: patterns of between-group interaction. — In: Primates in perspective (Campbell, C., Fuentes, A., MacKinnon, K., Panger, M. & Bearder, S., eds). Oxford University Press, Oxford, p. 201-224.
Fashing, P.J., Mulindahabi, F., Gakima, J.-B., Masozera, M., Mununura, I., Plumptre, A.J. & Nguyen, N. (2007). Activity and ranging patterns of Colobus angolensis ruwenzorii in Nyungwe forest, Rwanda: possible costs of large group size. — Int. J. Primatol. 28: 529-550.
Fimbel, C., Vedder, A., Dierenfeld, E. & Mulindahabi, F. (2001). An ecological basis for large group size in Colobus angolensis in the Nyungwe Forest, Rwanda. — Afr. J. Ecol. 39: 83-92.
Fitzgibbon, C.D. (1990). Mixed-species grouping in Thomson’s and Grant’s gazelles: the antipredator benefits. — Anim. Behav. 39: 1116-1126.
Fleury, M.C. & Gautier-Hion, A. (1999). Seminomadic ranging in a population of black colobus (Colobus santanas) in Gabon and its ecological correlates. — Int. J. Primatol. 20: 491-509.
Fox, M. (2004). The crushing mbega: a follow-up study of black and white colobus in the Usambara Mountains. — School for International Training, Arusha.
Freeland, W.J. (1976). Pathogens and the evolution of primate sociality. — Biotropica 8: 12-24.
Gillespie, T.R. & Chapman, C. (2001). Determinants of group size in the red colobus monkey (Procolobus badius): an evaluation of the generality of the ecological-constraints model. — Behav. Ecol. Sociobiol. 50: 329-338.
Gosselin-Ildari, A.D. & Koenig, A. (2012). The effects of group size and reproductive status on vigilance in captive Callithrix jacchus. — Am. J. Primatol. 74: 613-621.
Grimes, K.H. (2000). Guereza dietary and behavioural patterns at the Entebbe Botanical Gardens. — University of Calgary, Calgary, AB.
Grueter, C.C., Li, D., Ren, B., Wei, F. & Van Schaik, C.P. (2009). Dietary profile of Rhinopithecus bieti and its socioecological implications. — Int. J. Primatol. 30: 553-567.
Grueter, C.C., Robbins, A.M., Abavandimwe, D., Vecellio, V., Ndagijimana, F., Ortmann, S., Stoinski, T.S. & Robbins, M.M. (2016). Causes, mechanisms, and consequences of contest competition among female mountain gorillas in Rwanda. — Behav. Ecol. 27: 766-776.
Grueter, C.C., Robbins, A.M., Abavandimwe, D., Vecellio, V., Ndagijimana, F., Stoinski, T.S. & Robbins, M.M. (2018). Quadratic relationships between group size and foraging efficiency in a herbivorous primate. — Sci. Rep. 8: 16718.
Grueter, C.C. & Van Schaik, C.P. (2010). Evolutionary determinants of modular societies in colobines. — Behav. Ecol. 21: 63-71.
Harris, T.R. (2006). Between-group contest competition for food in a highly folivorous population of black and white colobus monkeys (Colobus guereza). — Behav. Ecol. Sociobiol. 61: 317-329.
Harris, T.R. & Chapman, C.A. (2007). Variation in diet and ranging of black and white colobus monkeys in Kibale National Park, Uganda. — Primates 48: 208-221.
Heinen, L. (2006). Mbega and the rainforest: a study of Colobus angolensis palliatus in the Sagara Village Forest. — School for International Training, Arusha.
Hirsch, B.T. (2002). Social monitoring and vigilance behavior in brown capuchin monkeys (Cebus apella). — Behav. Ecol. Sociobiol. 52: 458-464.
Hunter, C.P. (2001). Ecological determinants of gelada ranging patterns (Theropithecus gelada). — Doctoral dissertation, University of Liverpool, Liverpool.
Isabirye-Basuta, G. (1988). Food competition among individuals in a free-ranging chimpanzee community in Kibale Forest, Uganda. — Behaviour 105: 135-147.
Isbell, L.A. (1991). Contest and scramble competition: patterns of female aggression and ranging behaviour among primates. — Behav. Ecol. 2: 143-155.
Janson, C.H. (1988). Intra-specific food competition and primate social structure: a synthesis. — Behaviour 105: 1-17.
Janson, C.H. & Goldsmith, M.L. (1995). Predicting group-size in primates: foraging costs and predation risks. — Behav. Ecol. 6: 326-336.
Janson, C.H. & Van Schaik, C.P. (1988). Recognizing the many faces of primate food competition: methods. — Behaviour 105: 165-186.
Jarvey, J.C. (2016). The importance of underground foods in female gelada (Theropithecus gelada) socioecology. — Masters dissertation, University of Michigan, Ann Arbor, MI.
Jarvey, J.C., Low, B.S., Bergman, T.J. & Beehner, J.C. (2016). The roots of all evil: aggression and below-ground feeding in female geladas. — In: The 85th Annual Meeting of the American Association of Physical Anthropologists 159, p. 185-185.
Julliot, C. & Sabatier, D. (1993). Diet of the red howler monkey (Alouatta seniculus) in French Guiana. — Int. J. Primatol. 14: 527-550.
Kaplin, B.A., Munyaligoga, V. & Moermond, T.C. (1998). The influence of temporal changes in fruit availability on diet composition and seed handling in blue monkeys (Cercopithecus mitis doggetti). — Biotropica 30: 56-71.
Kazahari, N. (2014). Maintaining social cohesion is a more important determinant of patch residence time than maximizing food intake rate in a group-living primate, Japanese macaque (Macaca fuscata). — Primates 55: 179-184.
Kazahari, N. & Agetsuma, N. (2008). Social factors enhancing foraging success of a wild group of Japanese macaques (Macaca fuscata) in a patchy food environment. — Behaviour 145: 843-860.
Koenig, A. (2000). Competitive regimes in forest-dwelling Hanuman langur females (Semnopithecus entellus). — Behav. Ecol. Sociobiol. 48: 93-109.
Koenig, A., Beise, J., Chalise, M.K. & Ganzhorn, J.U. (1998). When females should contest for food — testing hypotheses about resource density, distribution, size, and quality with Hanuman langurs (Presbytis entellus). — Behav. Ecol. Sociobiol. 42: 225-237.
Koenig, A. & Borries, C. (2002). Feeding competition and infanticide constrain group size in wild hanuman langurs. — Am J Primatol. 57: 33-34.
Korstjens, A.H. (2001). The mob, the secret sorority, and the phantoms. — Doctoral dissertation, Utrecht University, Utrecht.
Kummer, H. (1984). From laboratory to desert and back: a social system of hamadryas baboons. — Anim. Behav. 32: 965-971.
Kutsukake, N. (2007). Conspecific influences on vigilance behavior in wild chimpanzees. — Int. J. Primatol. 28: 907-918.
Kuznetsova, A., Brockhoff, P.B. & Christensen, R.H.B. (2015). lmerTest: tests in linear mixed effects models. R package version 2.0-20. — R Foundation for Statistical Computing, Vienna.
Majolo, B., De Bortoli Vizioli, A. & Schino, G. (2008). Costs and benefits of group living in primates: group size effects on behaviour and demography. — Anim. Behav. 76: 1235-1247.
Mckey, D.B., Gartlan, J.S., Waterman, P.G. & Choo, G.M. (1981). Food selection by black colobus monkeys (Colobus satanas) in relation to plant chemistry. — Biol. J. Linn. Soc. 16: 115-146.
Miller, A., Judge, D.S., Uwingeneye, G., Ndayishimiye, D. & Grueter, C.C. (2020a). Diet and use of fallback foods by Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) in Rwanda: implications for supergroup formation. — Int. J. Primatol.: 1-24.
Miller, A., Uddin, S., Judge, D.S., Kaplin, B.A., Ndayishimiye, D., Uwingeneye, G. & Grueter, C.C. (2020b). Spatiotemporal association patterns in a supergroup of Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) are consistent with a multilevel society. — Am. J. Primatol.: e23127.
Miller, A., Uwingeneye, G., Ndayishimiye, D., Kaplin, B., Judge, D.S. & Grueter, C.C. (2018). The ‘anatomy’ of a supergroup of Colobus angolensis ruwenzorii in Nyungwe National Park. — In: Conference proceeding from the 27th International Primatological Society congress, Nairobi.
Milton, K. (1979). Factors influencing leaf choice by howler monkeys: a test of some hypotheses of food selection by generalist herbivores. — Am. Nat. 114: 362-378.
Milton, K. (1984). Habitat, diet, and activity patterns of free-ranging woolly spider monkeys (Brachyteles arachnoids E. Geoffroy 1806). — Int. J. Primatol. 5: 491-514.
National Research Council (2003). Nutrient requirements of nonhuman primates. — National Academies Press, Washington, DC.
Nyirambangutse, B., Zibera, E., Uwizeye, F.K., Nsabimana, D., Bizuru, E., Pleijel, H., Uddling, J. & Wallin, G. (2017). Carbon stocks and dynamics at different successional stages in an Afromontane tropical forest. — Biogeosciences 14: 1285-1303.
Oates, J.F. (1977a). The guereza and its food. — In: Primate ecology: studies of feeding and ranging behaviour in lemurs, monkeys, and apes (Clutton-Brock, T.H., ed.). Academic Press, London, p. 275-321.
Oates, J.F. (1977b). The social life of a black-and-white colobus monkey, Colobus guereza. — Ethology 45: 1-60.
Oates, J.F. (1994). The natural history of African colobines. — In: Colobine monkeys: their ecology, behaviour and evolution (Davis, G.A. & Oates, J.F., eds). Cambridge University Press, Cambridge, p. 75-128.
Olsen, K. (2007). When you’re talking about my monkeys… it doesn’t matter if they’re black or white: a behavioral follow-up on the black and white colobus and a study of their tree use. — School for International Training, Arusha.
Olson, R.S., Hintze, A., Dyer, F.C., Knoester, D.B. & Adami, C. (2013). Predator confusion is sufficient to evolve swarming behaviour. — J. Roy. Soc. Interface 10: 20130305.
Preston, M. (2002). An endangered monkey? A study of the Colobus angolensis. — School for International Training, Arusha.
Pulliam, H.R. (1973). On the advantages of flocking. — J. Theor. Biol. 38: 419-422.
R Core Team (2017). R: a language and environment for statistical computing. — R Foundation for Statistical Computing, Vienna.
Reich, P.B. & Borchert, R. (1984). Water stress and tree phenology in a tropical dry forest in the lowlands of Costa Rica. — J. Ecol. 72: 61-74.
Remis, M.J. (1997). Western lowland gorillas (Gorilla gorilla gorilla) as seasonal frugivores: use of variable resources. — Am. J. Primatol. 43: 87-109.
Ren, B., Li, D., Garber, P.A. & Li, M. (2012). Fission-fusion behavior in Yunnan snub-nosed monkeys (Rhinopithecus bieti) in Yunnan, China. — Int. J. Primatol. 33: 1096-1109.
Riedman, M.L. (1982). The evolution of alloparental care and adoption in mammals and birds. — Q. Rev. Biol. 57: 405-435.
Roberts, G. (1996). Why individual vigilance declines as group size increases. — Anim. Behav. 51: 1077-1086.
Rogers, M.E., Maisels, F., Williamson, E.A., Fernandez, M. & Tutin, C.E.G. (1990). Gorilla diet in the Lope Reserve, Gabon: a nutritional analysis. — Oecologia 84: 326-339.
Rose, M.D. (1978). Feeding and associated positional behavior of black and white colobus monkeys (Colobus guereza). — In: The ecology of arboreal folivores (Montgomery, C.G., ed.). Smithsonian Institution Press, Washington, DC, p. 253-262.
Rstudio (2017). RStudio: integrated development environment for R. — RStudio Inc, Boston, MA.
Rubenstein, D.I. (1986). Ecology and sociality of horses and zebras. — In: Ecological aspects of social evolution: birds and mammals (Rubenstein, D.I. & Wrangham, R.W., eds). Princeton University Press, Princeton, NJ, p. 282-302.
Saj, T.L. & Sicotte, P. (2007). Scramble competition among Colobus vellerosus at Boabeng-Fiema, Ghana. — Int. J. Primatol. 28: 337-355.
Sayers, K. (2016). Folivory. — In: The international encyclopedia of primatology, p. 1-5.
Scarry, C.J. (2013). Between-group contest competition among tufted capuchin monkeys, Sapajus nigritus, and the role of male resource defence. — Anim. Behav. 85: 931-939.
Schreier, A.L. & Swedell, L. (2012). Ecology and sociality in a multilevel society: ecological determinants of spatial cohesion in hamadryas baboons. — Am. J. Phys. Anthropol. 148: 580-588.
Schülke, O. & Ostner, J. (2012). Ecological and social influences on sociality. — In: The evolution of primate societies (Mitani, J.C., Call, J., Kappeler, P.M., Palombit, R.A. & Silk, J.B., eds). The University of Chicago Press, Chicago, IL, p. 195-219.
Shopland, J.M. (1987). Food quality, spatial deployment, and the intensity of feeding interference in yellow baboons (Papio cynocephalus). — Behav. Ecol. Sociobiol. 21: 149-156.
Silver, S.C., Ostro, L.E.T., Yeager, C.P. & Horwich, R. (1998). Feeding ecology of the black howler monkey (Alouatta pigra) in northern Belize. — Am. J. Primatol. 45: 263-279.
Snaith, T.V. & Chapman, C.A. (2005). Towards an ecological solution to the folivore paradox: patch depletion as an indicator of within-group scramble competition in red colobus monkeys (Piliocolobus tephrosceles). — Behav. Ecol. Sociobiol. 59: 185-190.
Snaith, T.V. & Chapman, C.A. (2007). Primate group size and socioecological models: do folivores really play by different rules? — Evol. Anthropol. 16: 94-106.
Snaith, T.V. & Chapman, C.A. (2008). Red colobus monkeys display alternative behavioral responses to the costs of scramble competition. — Behav. Ecol. 19: 1289-1296.
Snyder-Mackler, N., Beehner, J.C. & Bergman, T.J. (2012). Defining higher levels in the multilevel societies of geladas (Theropithecus gelada). — Int. J. Primatol. 33: 1054-1068.
Stanford, C.B. (1998). Chimpanzee and red colobus: the ecology of predator and prey. — Harvard University Press, Cambridge, MA.
Stead, S.M. & Teichroeb, J.A. (2019). A multi-level society comprised of one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii). — PLoS ONE 14: e0217666.
Steenbeek, R. & Van Schaik, C.P. (2001). Competition and group size in Thomas’s langurs (Presbytis thomasi): the folivore paradox revisited. — Behav. Ecol. Sociobiol. 49: 100-110.
Sterck, E.H.M., Watts, D.P. & Van Schaik, C.P. (1997). The evolution of female social relationships in nonhuman primates. — Behav. Ecol. Sociobiol. 41: 291-309.
Stevenson, P.R. & Castellanos, M.C. (2000). Feeding rates and daily path range of the Colombian woolly monkeys as evidence for between- and within-group competition. — Folia Primatol. 71: 399-408.
Su, H.H. & Birky, W.A. (2007). Within-group female–female agonistic interactions in Taiwanese macaques (Macaca cyclopis). — Am. J. Primatol. 69: 199-211.
Suzuki, A. (1979). The variation and adaptation of social groups of chimpanzees and black and white colobus monkeys. — In: Primate ecology and human origins: ecological influences on social organization (Bernstein, I.S. & Smith, E.O., eds). Garland STPM Press, New York, NY, p. 153-173.
Teichroeb, J.A., Saj, T.L., Paterson, J.D. & Sicotte, P. (2003). Effect of group size on activity budgets of Colobus vellerosus in Ghana. — Int. J. Primatol. 24: 743-758.
Teichroeb, J.A. & Sicotte, P. (2009). Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana. — Am. J. Primatol. 71: 49-59.
Teichroeb, J.A. & Sicotte, P. (2012). Cost-free vigilance during feeding in folivorous primates? Examining the effect of predation risk, scramble competition, and infanticide threat on vigilance in ursine colobus monkeys (Colobus vellerosus). — Behav. Ecol. Sociobiol. 66: 453-466.
Teichroeb, J.A. & Sicotte, P. (2018). Cascading competition: the seasonal strength of scramble influences between-group contest in a folivorous primate. — Behav. Ecol. Sociobiol. 72: 6.
Tombak, K.J., Reid, A.J., Chapman, C.A., Rothman, J.M., Johnson, C.A. & Reyna-Hurtado, R. (2012). Patch depletion behavior differs between sympatric folivorous primates. — Primates 53: 57-64.
Turner, G.F. & Pitcher, T.J. (1986). Attack abatement: a model for group protection by combined avoidance and dilution. — Am. Nat. 128: 228-240.
Van Schaik, C.P. (1989). The ecology of social relationships amongst female primates. — In: Comparative socioecology: the behavioural ecology of humans and other mammals (Standen, V. & Foley, R.A., eds). Blackwell Scientific Publications, Boston, MA, p. 195-218.
Van Schaik, C.P. & Pfannes, K.R. (2005). Tropical climates and phenology: a primate perspective. — In: Seasonality in primates (Brockman, D.K. & van Schaik, C.P., eds). Cambridge University Press, Cambridge, p. 23-54.
Van Schaik, C.P. & Van Hooff, J.A.R.A.M. (1983). On the ultimate causes of primate social systems. — Behaviour 85: 91-117.
Van Schaik, C.P. & Van Noordwijk, M.A. (1988). Scramble and contest in feeding competition among female long-tailed macaques (Macaca Fascicularis). — Behaviour 105: 77-98.
Vedder, A. & Fashing, P.J. (2002). Diet of a 300-member Angolan colobus monkey (Colobus angolensis) supergroup in the Nyungwe forest, Rwanda. — Am. J. Phys. Anthropol. Suppl. 34: 159-160.
Vogel, E.R. (2005). Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition. — Behav. Ecol. Sociobiol. 58: 333-344.
Vogel, E.R., Harrison, M.E., Zulfa, A., Bransford, T.D., Alavi, S.E., Husson, S., Morrogh-Bernard, H., Firtsman, T., Utami-Atmoko, S.S., Van Noordwijk, M.A. & Farida, W.R. (2015). Nutritional differences between two orangutan habitats: implications for population density. — PLoS ONE 10: e0138612.
Wasserman, M.D. & Chapman, C.A. (2003). Determinants of colobine monkey abundance: the importance of food energy, protein and fibre content. — J. Anim. Ecol. 72: 650-659.
Wijtten, Z., Hankinson, E., Pellissier, T., Nuttall, M. & Lemarkat, R. (2012). Activity budgets of Peters’ Angola black-and-white colobus (Colobus angolensis palliatus) in an East African coastal forest. — Afr. Primates 7: 203-210.
Wrangham, R.W. (1980). An ecological model of female-bonded primate groups. — Behaviour 75: 262-300.
Wrangham, R.W., Gittleman, J. & Chapman, C.A. (1993). Constraints on group size in primates and carnivores: population density and day-range as assays of exploitation competition. — Behav. Ecol. Sociobiol. 32: 199-210.
Wright, S.J. & Van Schaik, C.P. (1994). Light and the phenology of tropical trees. — Am. Nat. 143: 192-199.
Yeager, C.P. & Kirkpatrick, R.C. (1998). Asian colobine social structure: ecological and evolutionary constraints. — Primates 39: 147-155.
Yeager, C.P. & Kool, K. (2000). The behavioral ecology of Asian colobines. — In: Old world monkeys (Whitehead, P.F. & Jolly, C.J., eds). Cambridge University Press, Cambridge, p. 496-521.
| All Time | Past 365 days | Past 30 Days | |
|---|---|---|---|
| Abstract Views | 1639 | 346 | 10 |
| Full Text Views | 151 | 6 | 1 |
| PDF Views & Downloads | 205 | 14 | 2 |
Competition for food is often a cost associated with living in a group, and can occur in an indirect (scramble) or direct (contest) form. We investigated feeding competition in a supergroup of Rwenzori black-and-white colobus monkeys (Colobus angolensis ruwenzorii) in Rwanda, with the aim of establishing whether freedom from scramble competition allows these monkeys to form supergroups. We used the patch depletion method, measuring intake rate coupled with movement rate, to assess if food patches become depleted over the occupancy period. Resource depletion was evident when the colobus fed on young leaves, but not when feeding on mature leaves. Scramble competition was inferred from a negative correlation between group size and change in intake rate over patch occupancy. Between-group contest competition was inferred from displacement from patches. Although feeding competition exists for select resources, limited competition for mature leaves may enable Rwenzori colobus to live in a supergroup of hundreds of individuals in this montane forest.
| All Time | Past 365 days | Past 30 Days | |
|---|---|---|---|
| Abstract Views | 1639 | 346 | 10 |
| Full Text Views | 151 | 6 | 1 |
| PDF Views & Downloads | 205 | 14 | 2 |