The natural regulation of buffalo populations in East Africa: The food supply as a regulating factor, and competition*

Summary Evidence is presented to show that both the quality and quantity of food available to a buffalo population falls below the minimum maintenance requirements of that population at certain times of the year. In the Serengeti grasslands there was a shortage of the only good quality component, grass leaf, in the dry season, with the result that the animals consumed an increasing proportion of poor quality food such as grass stem. By the end of the dry season the diet had dropped in quality below the minimum maintenance level. In areas such as Mt. Meru where there was a more continuous growing season, the high density of animals kept the standing crop of leaf at a low level. During the cooler dry season the growth of leaf became insufficient in quantity for the maintenance requirements of the population. These two quite different situations suggested that food shortage was a more general phenomenon in eastern Africa. Various measurements of feeding behaviour were made. Total grazing time per 24 h did not differ between seasons but ruminating time may have increased as the season became drier and could have been a response to the more fibrous food. Analysis for cycles of activity showed that there was more temporal organization during the dry season. These changes in activity cycles appeared to be related to the increase in energy expenditure produced by heat stress and sweating. Old animals with poor teeth did not compensate for the poor food supply by changing their feeding behaviour. There was a positive relationship between annual rainfall and mean crude density in different areas of eastern Africa, indicating that regulation was taking place. Since rainfall determined the amount of available food, it could have operated through the food supply. On a finer scale it was found that the extent of the preferred riverine habitats was also related to density. Thus rainfall, the extent of riverine habitat and perhaps soil moisture were three limiting factors that determined mean density and all could have taken effect through the food supply. As a result of initial selective grazing the amount of available leaf declined as the dry season progressed to the extent that by the end of the season the proportion of this component in the diet fell to a very low level. The impact of the population on its limited food supply indicated that intra-specific competition was acting as the cause of regulation. Measurements of wildebeest eating the same food in the same habitats as buffalo showed that inter-specific competition was also taking place. A small proportion (7“) of the wildebeest population could have reduced the buffalo population by approximately 18o,‘, from its potential population size. The buffalo population in the Serengeti was regulated by adult mortality which was caused by undernutrition as a result of food shortage. This food shortage was caused by intra- and inter-specific competition. The effect of predation and disease was to hasten the response of the population to changes in the food supply. The limiting factors determining the mean level of the available food were, amongst others, rainfall, soil moisture and the extent of the preferred riverine habitat. The effect of interspecific competition could result in a complex regulation of populations through their food supply. There appears to be no foundation for hypotheses which invoke over-utilization or damage as a consequence of regulation through food.