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10 Chapter 1

10 Chapter 1

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Published by JF Derry

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Published by: JF Derry on Sep 12, 2010
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1.1 Introduction
When Coleridge's Ancient Mariner said 'Water, water, everywhere, nor any drop to drink', he gave a fair picture of the global situation. The'drop to drink' is a hundredth of a percent of the world's water: about onedrop in every bucketful. The proportion of planetary water that is fresh israther larger - around 3.5 per cent - but most is frozen in the ice capsand mountain glaciers. As sea water is corrosive and toxic to land-based animals and plants, nearly all of the water that we use must come fromthat precious one hundredth of a per cent.
 Ball (2000)The small proportion of global water that is available for plant and animalconsumption is not distributed evenly in time and space, especially in the earth's drierzones. As a result livestock managers in semi-arid areas need to cope with theinsecurities arising from climatic variation and frequent droughts (
, Scoones1994). Additionally, growing evidence shows that they also need to carefully managewatering points in order to preserve key resources required for the survival of animals when constrained to their dry season range (Illius & O'Connor 2000).Imprudent depletion of winter forage can be catastrophic (
, Sinclair & Fryxell1985).Mammalian herbivores require drinking water to differing degrees (
, Western1975). The congregation of these animals around focal points on rangeland fromwhere they acquire this supplementary water results in a concentration of environmental impacts (Andrew 1988). The subsequent land modification isconsidered by some to be localised degradation (Perkins & Thomas 1993b) ordesertification (
, Dean
et al.
1995), and is especially noticeable in arid and semi-arid landscapes. The affected area has been termed a piosphere (Lange 1969).Gradient sampling strategies have been used to quantify the piosphere effect (
et al.
1987), revealing remarkable consistency in their mathematicalcharacterisation (Graetz
Ludwig 1978). How much of this response is due to theanimal component, independently and via interaction with other components, and
 2how much influence each component has on the system dynamics has yet to bedetermined.Part 1 of this introductory chapter (Section 1.1) defines the piosphere effect and dealswith its ecology. The second part (Section 1.2) is a review of modelling literaturerelevant to piospheres. This part is adapted from Thrash & Derry (1999) and includesa review of more recent literature (Section 1.2.11) and further investigates piospherecharacteristics by application of a generalized piosphere model to piosphere data setscollected for African savanna. The final part (Section 1.3) describes the objectivesand structure of this thesis.
1.1.1 Spatial resource heterogeneity
Rainfall infiltration and the spatial redistribution of runoff water are the predominantfactors determining patterns in semi-arid vegetation (Friedel 1990, Maestre
et al.
 2003), but grazing impacts also contribute to the generation and maintenance of spatial heterogeneity (Adler
et al.
2001). It is probable that animals must thereforerespond to this spatial variation in their food distribution (Pyke 1984) implying acapacity for assessment of resource patchiness (Ford 1983). This assessment mustoperate at a scale that is functionally meaningful to each animal (Wiens 1976) andwill elicit a response relative to how much they are affected by the fragmentation of their habitat (Hester
et al.
1999).It is a pervading question in spatial ecology, yet little is known about the scale orscales that this assessment operates (Levin & Pacala 1997) to dictate foragingbehaviour and landscape utilisation. Our assessments of animal behaviour must be interms of function rather than resource organisation (Li & Reynolds 1995) to ensurequantification of the animal’s perceived heterogeneity of its environment (Wiens1976, Bailey
et al.
1.1.2 Optimal spatial foraging
Large mammalian herbivores prospect their environment for an optimal diet (Illius &Gordon 1993), but because they perceive their environment at differing scales
 3(Kotliar & Weins 1990, With 1994, Levin & Pacala 1997), they also differ in thecues that they use to do so (Eztenhouser
et al.
1998).The landscape components that may act as potential cues for animal behaviour arepresent across regional, landscape and plant community scales (Senft
et al.
1987),providing a hierarchical framework for animal foraging decisions (Orians &Wittenberger 1991). However, while animal diet selection may be sensitive to thesmall-scale variation in food distribution (Edwards
et al.
1994, Turner 1999), there islittle evidence in support of daily decisions at large scales, other than the apparentbehaviours of seeking shade and water (Senft 1989, Cowley 2001).Foraging strategies within landscapes seek to maximise daily energy gain (Fryxell
et al.
2001) which is sufficient to motivate large-scale animal movements (Wilmshurst
et al.
1999). It follows that animals are expected to make decisions about dietselection based on the balance between forage profitability (a function of thesatisfaction of nutritional requirements) and the distance travelled to reach thisforage. This is encapsulated by Optimal Foraging Theory (Stephens & Krebs 1986)which predicts that animals assay the energy balance underlying travel and intakeagainst the profitability of their resource (Bailey
et al.
1998).Resource profitability is energy gained in excess of costs, the rate of which isconstrained by the logistics of food detection and ingestion. Intake rate constraintsdepend on the initial locating of food items, the travel between those food items, and,once arrived, the speed of cropping, chewing and swallowing of food (Spalinger &Hobbs 1992). Feeding strategies also need to account for the instantaneous decline inintake rate associated with the successive removal of food from a single location.The Marginal Value Theorem (Charnov 1976) predicts that feeding should onlyoccur for locations while resource profitability is above the environmental average. If all the profitable locations are exploited, it can be said that there has been a degree of 
between animals and their resource, and the resulting spatial pattern of resource utilisation will describe the Ideal Free Distribution (Fretwell & Lucas1970).

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