Emily Miller
(M.A. Conservation Biology)
Mixed-Species Associations of Blue
Monkeys (Cercopithecus mitis) and
Black-and-White Colobus (Colobus guereza)
in the Kakamega Forest, Kenya

Mixed-species associations are usually
understood in terms of the costs and benefits that participant species experience.
These costs and benefits may be asymmetrical in direction and degree for the
species involved. The two major categories of proposed benefit are
improved protection against predation and the improved acquisition of
resources.
Many researchers conclude that the
predation avoidance benefit is the main driver of associations. Foraging
benefits may also explain why mixed-species associations occur more frequently
than expected by chance. Larger groups can monopolize food patches and
exclude smaller groups. Heterospecifics may enhance foraging efficiency
if they have a greater knowledge of food availability and can act to guide
their partner species to profitable feeding sites. Heterospecifics may
also be able to take advantage of foraging in different microhabitats when in
association.
Mixed-species associations can be
costly for one or both species involved because of increased feeding
competition as a result of increasing group size. Mixed-species
association may reduce the cost of intragroup feeding and mate competition
relative to association in a larger single-species group. Feeding
competition is likely to be especially low when associations involve
phylogenetically more distant taxa, such as colobines and cercopithecines,
whose diets are more dissimilar than closely related species.
Among primates, mixed-species
associations are common among closely related (congeneric) species, whose
ecological similarity in terms of resource use and predation threats likely
contributes to their adaptiveness. However, in Africa, associations
between less closely related cercopithecines and colobines are also common.
Phylogenetic difference has implications for why mixed-species associations
occur: when very different taxa associate, dietary divergence is likely to
influence both the costs and benefits of association.
To characterize the associations of one
blue monkey group and three black-and-white colobus groups in Kakamega Forest,
I am using population density data to first look at whether their associations
occur more frequently than predicted by chance. I am examining the
potential benefits and costs of these mixed-species associations using data I
collected on vigilance rates, ingestion rates, diet composition, alarm calls
and responses, travel distance and speed, microhabitat usage, agonism, and
association initiation and termination patterns. For each species, I am
comparing these measures for when the species was in and out of association to
better understand the occurrence of associations between two phylogenetically
distant primate taxa.
Publication:
Mattila J, Heck KL Jr.,
Millstein E, Miller E, Gustafsson C, Williams S, Byron D (2008). Increased
habitat structure may not always provide increased refuge from predation. Marine
Ecology Progress Series. 361:15-20.