Publication Date



Open access

Degree Type


Degree Name

Doctor of Philosophy (PHD)


Biology (Arts and Sciences)

Date of Defense


First Committee Member

Michael S. Gaines

Second Committee Member

William Searcy

Third Committee Member

David Janos

Fourth Committee Member

Judith Bronstein


Mutualisms are delicately balanced partnerships and are increasingly recognized as being fundamental to patterns and processes within ecological systems. Changes to the ecological setting in which such interactions operate can disrupt this balance. By understanding the context-dependent nature of such associations, researchers can begin to understand how changes in the environment can have cascading effects on the entire community within which they occur. Oxpeckers (Buphagus spp.) feed on the ectoparasites of ungulates in sub-Saharan Africa and from the blood of ungulate wounds. Because of this dichotomy in feeding behavior, the role of oxpeckers as consistently beneficial partners of their hosts has been questioned, and it has been suggested that the concept of conditional mutualism be applied to this interaction. I combined observational studies of oxpeckers in Kruger National Park, South Africa, with experiments on oxpeckers in captivity at the Mokopane Biodiversity Conservation Centre, South Africa, to test the hypothesis that the ectoparasite cleaning service provided by oxpeckers to their hosts is context-dependent. The results of my dissertation research demonstrate that the oxpecker-ungulate association has conditional outcomes. Under certain conditions, the oxpecker-ungulate relationship is a nutritional mutualism where ungulates provide food (ticks) for oxpeckers in exchange for a cleaning service. Under other conditions, oxpeckers exploit their hosts to feed from their blood. My findings suggest that a high abundance of the tick species and tick stages oxpeckers prefer is necessary to maintaining an alignment of interests between oxpeckers and ungulates. When ungulates host few ticks of the species oxpeckers prefer, these birds will wound-feed to meet their nutritional demands. Such wound-feeding threatens the balance of the oxpecker-ungulate mutualism. Although oxpeckers frequently wound-fed in captivity, the frequency of wound-feeding events on wild host species was only 3.1% of feeding events. Whether the relationship between oxpeckers and domesticated hosts differs from that between oxpeckers and wild hosts, or if this observed difference is actually a result of researcher constraints in the wild, needs further exploration.


Africa; Nutritional Analyses; Tick Drag; Interspecific Association; Ungulate