ABSTRACT

This thesis explains two aspects of animal spatial foraging behaviour arising as a direct consequence
of animals' need to drink water: the concentration of animal impacts, and the response of animals to
those impacts.
In semi-arid rangelands, the foraging range of free-ranging large mammalian herbivores is constrained
by the distribution of drinking water during the dry season. Animal impacts become concentrated
around these watering sites according to the geometrical relationship between the available foraging
area and the distance from water, and the spatial distribution of animal impacts becomes organised
along a utilisation gradient termed a "piosphere". During the dry season the temporal distribution of
the impacts is determined by the day-to-day foraging behaviour of the animals. The specific
conditions under which these spatial foraging processes determine the piosphere pattern have been
identified in this thesis.
At the core of this investigation are questions about the response of animals to the heterogeneity of
their resources. Aspects of spatial foraging are widely commented on whilst explaining the
consequences of piosphere phenomena for individual animal intake, population dynamics, feeding
strategies and management. Implicated are our notions of optimal foraging, scale in animal response,
and resource matching. This thesis addressed each of these. In the specific context of piospheres, the
role of energy balance in optimal foraging was also tested.
Field experiments for this thesis showed a relationship between goat browsing activity and measures
of spatial impact. As a preliminary step to investigating animal response to resource heterogeneity, the
spatial pattern of foraging behaviour/impacts was described using spatial statistics. Browsing activity
varied daily revealing animal assessment of the spatial heterogeneity of their resources and an
energetic basis for foraging decisions. This foraging behaviour was shown to be determined by
individual plants rather than at larger scales of plant aggregation. A further experiment investigated
the claim that defoliation has limited impact on browser intake rate, suggesting that piospheres may
have few consequences for browser intake. This experiment identified a constraining influence of
browse characteristics at the small scale on goat foraging by relating animal intake rate to plant bite
size and distribution.
Computer simulation experiments for this thesis supported these empirical findings by showing that
the distribution of spatial impacts was sensitive to the marginal value of forage resources, and
identified plant bite size and distribution as the causal factors in limiting animal intake rate in the
presence of a piosphere. As a further description of spatial pattern, piospheres were characterised by
applying a contemporary ecological theory that ranks resource patches into a spatial hierarchy.
Ecosystem dynamics emerge from the interactions between these patches, with piospheres being an
emergent property of a natural plant-herbivore system under specific conditions of constrained
foraging. The generation of a piosphere was shown to be a function of intake constraints and available
foraging area, whilst piosphere extent was shown to be independent of daily energy balance including
expenditure on travel costs. A threshold distance for animal foraging range arising from a
hypothesised conflict between daily energy intake and expenditure was shown not to exist, whereas
evidence for an intermediate distance from water as a focus for accumulated foraging activity was
identified.
Individual animal foraging efficiency in the computer model was shown to be sensitive to the
piosphere, while animal population dynamics were found to be determined in the longer term by dry
season key resources near watering points. Time lags were found to operate in the maintenance of the
gradient, and the density dependent moderation of the animal population. The latter was a direct result
of the inability of animal populations to match the distribution of their resources with the distribution
of their foraging behaviour, because of their daily drinking requirements. The result is that animal
forage intake was compromised by the low density of dry season forage in the vicinity of a water
point.
This thesis also proposes that piospheres exert selection pressures on traits to maximise energy gain
from the spatial heterogeneity of dry season resources, and that these have played a role in the
evolution of large mammalian herbivores.