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a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m

w w w. e l s e v i e r. c o m / l o c a t e / e c o l e c o n

ANALYSIS

Sustainable management of extensively managed savanna rangelands

Steven I. Higgins a,, Jochen Kantelhardt b , Simon Scheiter a , Jan Boerner b
a b

Lehrstuhl fr Vegetationskologie, Technische Universitt Mnchen, 85350 Freising-Weihenstephan, Germany Lehrstuhl fr Wirtschaftslehre des Landbaues, Technische Universitt Mnchen, 85350 Freising-Weihenstephan, Germany

AR TIC LE I N FO
Article history: Received 14 November 2005 Received in revised form 24 April 2006 Accepted 26 May 2006 Available online 31 July 2006 Keywords: Bush encroachment Disequilibrium theory Opportunistic strategies Grass-tree interactions Fire Livestock production

ABS TR ACT
Classic rangeland theory advocates stocking rangelands at relatively low and constant levels. This theory has been labelled inappropriate for savanna rangelands, because savannas are strongly influenced by stochastic processes. Opportunistic strategies that force animal numbers to track available forage have been proposed as an alternative management paradigm. However, no studies have examined whether these opportunistic strategies are sustainable or optimal. We developed a simulation model of a savanna rangeland to identify optimal, sustainable strategies for the management of extensive rangelands. We optimised the utility of agents who are motivated by economic, production or ecological factors under both deterministic and stochastic conditions. In all cases we found that it was optimal to manage the system conservatively and not opportunistically. Moreover, it was optimal to manage more conservatively under stochastic conditions. Key elements of the conservative strategy were to stock at low levels and to use fire to control tree abundance and thereby maintain the system in a grass dominated state. We conclude that opportunistic strategies of range management although intuitively appealing are not optimal. 2006 Elsevier B.V. All rights reserved.

1.

Introduction

Savannas are defined as tropical ecosystems where grasses and trees co-dominate (Huntley and Walker, 1982). They cover large proportions of the tropical continents, that is 65% of Africa, 60% of Australia and 40% of South America (Huntley and Walker, 1982). The large area covered by savannas means that their sustainable management is of regional and global concern. One of the main land-use activities practised in savannas is livestock production, yet consensus on what might constitute sustainable livestock production systems remains elusive (Vetter, 2005). Defining sustainable land-use systems requires careful consideration of what one means by sustainability. Following
Corresponding author. E-mail address: higgins@wzw.tum.de (S.I. Higgins). 0921-8009/$ - see front matter 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.ecolecon.2006.05.019

the Brundtland Report (WECD, 1987), sustainable development aims to guarantee inter- and intragenerational fairness concerning the use of natural resources. Hence a thorough evaluation of sustainable development requires the consideration of economic, ecological and ethical factors in an integrated framework. Economic sustainability typically means that resources should be managed in such a way that the utility does not decline over time (Perman et al., 2003). Ecological sustainability may mean preserving ecological resilience over time or ensuring that the flow of some ecological service does not decline over time (Daily, 1997). For instance, in a grazing system maintaining ecological resilience might involve preserving the soil layer or preventing the system from moving into a tree dominated state; while

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sustaining ecological services may involve maintaining large trees that act as keystone species. The interpretation of sustainability also varies among actors, because different actors have different risk aversions, discount rates and constraints. For instance, a farmer might maximise utility by maximising profit while discounting future earnings strongly; whereas a section of society might maximise utility by maximising the stability of production. The focus of this study is the sustainable management of extensive rangeland systems in savanna regions. That is, we consider systems in which supplementary feeding of livestock is not economically viable and where grass and livestock production are primarily limited by rainfall. Rainfall in savanna regions varies with season; most rain falls in the summer months, very little rain falls in the winter months. Rainfall also varies stochastically between years. As a consequence, livestock farmers in savanna regions must develop strategies for dealing with variability and uncertainty in resource supply. Theory based on the harvesting of a stock that has density dependent and deterministic growth has strongly influenced the management of savanna rangelands (Mentis, 1984). However, more recent studies (Ellis and Swift, 1988; Behnke and Scoones, 1993) have argued that the stochastic nature of rainfall, which is the primary driver of resource supply in savannas, make such theories inappropriate for savannas. This tension between classic and new rangeland theories (Cowling, 2000) has often been cast as a conflict between equilibrium and non-equilibrium paradigms, or between stochastic and deterministic paradigms. However, such polarisation is counter productive, because in any dynamical system both non-equilibrium or transient dynamics and equilibrium dynamics are important. Similarly, many dynamical systems are influenced by a mixture of both stochastic and deterministic processes. In savannas, the equilibriumnon-equilibrium dichotomy is particularly inappropriate because even when a savanna is not at equilibrium, its trajectory in state space is determined by the system's equilibrium points. The stochastic deterministic dichotomy is also inappropriate in savannas because while the stochastic effects are apparent (e.g. rainfall on grass production), deterministic linkages are equally undeniable (e.g. grass consumption on animal production). The broad aim of this study is to propose a model of savanna rangelands that combines a mix of the deterministic and stochastic, equilibrium and non-equilibrium processes that form the keystones of the classic and new rangeland paradigms. We use this model to explore guidelines for the sustainable management of livestock production systems in savannas. Our intent is to explore the extent to which different aspects of sustainability are compatible with one another. Specifically, we aim to define strategies that optimise the sustainability from the perspective of agents that are motivated by economic, ecological and production factors under deterministic and stochastic environmental conditions.

important difference between our model and previous models is that we model, for both grasses and trees, two biomass compartments, roots and shoots. This allows us to simulate the fact that herbivores and fire cannot consume roots, and allows a separation of below ground and above ground competition (Fig. 1). The model of Perrings and Walker (1997) does not separate above ground from below ground processes, whereas Janssen et al. (2004) only make this distinction for grasses. In the paragraphs that follow we describe growth functions for the grass and tree biomass components of the model (all parameters and their default values are listed in Table 1). The grass shoot biomass in the next time step (GSt+1) is determined by a growth rate parameter (gGS) and grass root biomass (GRt). Growth is negatively influenced by competition from grass shoot biomass and woody shoot biomass. The parameter WG describes the intensity of the competitive effect of woody shoots on grass shoot growth, while determines the strength and linearity of both the intra- and inter-specific competitive effects (when > 1 competitive effects are only felt at higher levels of competitor density). Grass shoot biomass can be consumed by both herbivores and fire. Herbivore consumption is the product of the rate of shoot consumption per livestock unit per time step () and the number of livestock units (Zt). Note that we assume that our animals are grazers and that they therefore do not consume significant amounts of woody shoot biomass. Shoot biomass is also reduced by decomposition (dGS). We assume that the proportion of grass shoot biomass consumed by fire increases with fire intensity (Ft), but that above a certain fire intensity the consumption of shoot biomass asymptotes. We describe this asymptotic relationship using a function that takes the form f x; a; b xb : ab xb

For x 0 the function f(x; a, b) is bounded by 0 and 1 and the parameter a defines the value of x where f(x; a, b) = 0.5. The parameter b describes how rapidly f(x; a, b) changes as x approaches a. We use the function f(x; a, b) to describe several

2.

Ecological sub-model

We create a discrete time model, inspired by the models of Perrings and Walker (1997) and Janssen et al. (2004). The most

Fig. 1 Conceptual outline of the grasstree model. The model has four main compartments allowing the separation of above ground from below ground processes. The model simulates growth and decomposition of these compartments as well as competition between the grass and tree compartments. Grazing and fire can remove grass shoots, while fire can remove tree and grass shoots.

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Table 1 Descriptions of the parameters used in the model and default values Symbol
Ecological parameters gGS gGR gWS gWR s gZ dGS dGR dWS dWR WG GW WG f a a b a b

Name
Growth rate grass shoot Growth rate grass root Growth rate tree shoot Growth rate tree root Coefficient rainfall on growth rate Growth rate animals Decomposition rate grass shoots Decomposition rate grass roots Decomposition rate tree shoots Decomposition rate tree roots Competition coefficient tree on grass roots Competition coefficient grass on tree roots Competition coefficient tree on grass shoots Weighting of tree shoot biomass for wWG Grazing rate Energy transfer of ignited grass Moisture weighting for fire ignition Maximum consumption of grass shoots by fire Location parameter grass consumption by fire Shape parameter grass consumption by fire Maximum consumption of tree shoots by fire Location parameter tree consumption by fire Shape parameter tree consumption by fire

Value
0.005 0.005 0.0025 0.0025 0.73 0.001 0.0005 0.0005 0.00025 0.00025 1 1 1 2 0.001 100 20 0.9 0.3 0.75 1 1.5 5

Units
biomass day 1 biomass day 1 biomass day 1 biomass day 1 dimensionless biomass day 1 biomass.day 1 biomass day 1 biomass day 1 biomass day 1 dimensionless dimensionless dimensionless dimensionless biomass day 1 dimensionless dimensionless proportion fire intensity dimensionless proportion Fire intensity dimensionless

Equation
(1) (2) (3) (4) (7) (16) (1) (2) (3) (4) (2) (4) (1) (1) (1) (5) (5) (1) (1) (1) (3) (3) (3)

Management parameters Location parameter selling animals a Shape parameter selling animals b Minimum proportion of animals sold min a Location parameter purchasing animals Shape parameter purchasing animals b Maximum level of fire suppression max Natural fire ignition threshold n a Location parameter fire suppression Shape parameter fire suppression b Technology level a Location parameter technology Shape parameter technology b Economic parameters p q c

* * * * * *
0.3

* * *
0.1 1

grass root biomass dimensionless grass root biomass potential purchases dimensionless grass shoot biomass grass shoot biomass woody root biomass dimensionless proportion proportion dimensionless

(19) (19) (19) (18) (18) (15) (15) (15) (15) (17) (17) (17)

Price of animals Price band factor Cost of technology Cost of fire suppression Discount rate Ecological utility weight Risk sensitivity between time series

1000 1.1 0.05 0.05 0.02 20 0.5

currency dimensionless currency currency proportion year 1 dimensionless dimensionless

(8) (8) (8) (8) (10) (13) (21)

Parameters marked with

* are selected through optimisation.

relationships in the model. The equation for shoot biomass at time t + 1 is then, GSt1 GSt gGS GRt 1GSh xWG WSh lZt t t f Ft ; ab bb bGSt dGS GSt : 1

Here a is the threshold fire intensity where grass shoot consumption increases, b describes how rapidly grass shoot consumption changes as a is approached, and is the maximum grass shoot biomass removed by a fire. The grass root biomass in the next time step, GRt+1, is influenced by a growth rate parameter (gGS), grass shoot biomass, and negatively by grass root biomass and tree root

biomass (WRt). The effect of tree biomass is included to simulate competition between grasses and trees for soil water (Walter, 1971). The intensity of competition is described by the parameter WG. Grass root biomass decomposes at a constant rate (dGR). These assumptions yield the growth equation, GRt1 GRt gGR GSt 1GRt aWG WRt dGR GRt : 2

The tree shoot biomass in the next time step, WSt+1, is determined by a growth rate parameter gWS, tree root biomass (WRt) and is negatively influenced by the density of tree shoot biomass. Tree shoots decompose at rate dWS and can be

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consumed by fire. Tree shoot consumption by fire is modelled in the same way as grass shoot consumption by fire. That is, we assume that woody shoot consumption increases at a threshold level of fire intensity (a) and that the parameter b describes how rapidly woody shoot consumption changes as a is approached; is the maximum shoot biomass removed by a fire. The woody shoot growth equation is thus, WSt1 WSt gWS WRt 1WSt f Ft ; ag ; bg gWSt dWS WSt : 3

Table 2 Monthly mean and variance in rainfall used in the simulations (from New et al., 2002) Month
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Mean
100 78 63 45 9.0 2.3 1.2 3.2 15 39 58 70

SD
14 12 10 7 2 1 0 1 2 8 11 13

The tree root biomass in the next time step, WRt+1, is determined by a growth rate parameter gWR, tree shoot biomass, and is negatively influenced by tree root biomass, as well as by grass root biomass. The intensity of competition is described by the parameter GW. Tree root biomass decomposes at a constant rate (dWR). These assumptions yield the growth equation, WRt1 WRt gWR WSt 1WRt aGW GRt dWR WRt : 4

Fire intensity (Ft) is assumed to be a function of the fuel consumed, the energy transfer of ignited fuel and the energy required to ignite fuel (Rothermal, 1972). In savannas, grass (GSt) is the primary fuel. The energy required for ignition is a function of fuel moisture content. We assume that fuel moisture content is, in turn, a step function of rainfall, Rt. The step function implies that there is some critical fuel moisture level required for dry fuel to ignite (a). The parameter f is a coefficient that represents the energy transfer flux. Potential fire intensity, bt, is then, F Ft qf GSt maxRt ; qa 5

of rainfall at a site to simulate monthly rainfall as a gamma distributed random number (New et al., 2002). The monthly mean and variance of the rainfall parameters used for the simulations are listed in Table 2. The monthly time step additionally imposes a 1-month lag between removal of biomass by fire and regrowth.

3.
3.1.

Economic sub-model
Utility

Below a certain threshold in potential fire intensity (t) fires cannot occur, hence realised fire intensity, Ft, is, F Ft bt Ft 0 ; bt > dt F ; bt V dt : F 6

We assume that rainfall is the limiting resource in savannas and that plant growth rates are tightly coupled to rainfall. Rainfall varies stochastically over the seasonal cycle and between years. Here we define the time dependent versions of the growth parameters used in Eqs. (1)(4) as linear functions of monthly rainfall (Rt), gt gs minRa ; Rt ; 7

where g is from Eqs. (1)(4) and s is a coefficient that describes the relationship between rainfall and growth. We assume s is constant across all growth parameters. Ra is used to specify the rainfall at which grass production asymptotes. In the simulations presented here we use a monthly time step. This allows us to simulate the seasonality of rainfall typical of tropical savannas. We use the CRU global climatic database (New et al., 2002) as a source of data for Rt. For the deterministic simulations we use the monthly mean rainfall at a site, for the stochastic simulations we use the monthly mean and variance

Given the different definitions of sustainability outlined in the introduction the agent may either aim to increase income, increase production or increase the value of some ecological indicator. A farmer may be interested in the economic definition, the state may be interested in the production definition and the ecological definition. In the paragraphs that follow we propose functions for the utility of these different agents. The economic utility is determined by income and costs, which are a function of the number of animals bought and sold, the costs of technology and the costs of fire management. The income derived is determined by pt, the price of the livestock product at time t, and t, the number of livestock sold at time t. Note that for generality we define prices and costs as being time dependent although we treat them as constants in this paper. Costs are incurred when the farmer buys livestock and we assume that the buy price differs from the selling price by a factor , hence the cost incurred at time t due to purchasing t livestock is ptt. Costs are also incurred per time step due to the level of technology (examples of technology are labour, fencing, boreholes and water pumps) the farmer uses. The costs of technology are simply the product of the unit cost of technology at time t (qt) and the technology level ( {0, 1}). Costs due to suppressing fires are assumed to be the product of the unit cost of suppressing fires at time t (ct) and the level of fire suppression adopted (|t n|). How the number of animals bought and sold and how the technology and fire management levels are determined are described in Section 3.2. The income and cost parameters together define t, the economic index at time t as, pt pt rt upt vt qt wct jdt dn j: 8

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The economic utility over a planning horizon T, Ueconomic is, Ueconomic

T X t1

ptV;

) that is the sum of the discounted economic index (t over the finite planning horizon T. We use a standard exponential discounting function to , calculate t  t 1 ; 10 ptV pt 1s where is the discount rate. Although different discount rates () are appropriate for different agents we assume = 0.02 p.a. for all baseline analyses presented here. The production motivated agent aims to maximise production irrespective of any costs that might limit this goal. Hence, the costs of fire management and technology are set to zero. However, this agent remains subject to the price differential between buying and selling animals as described in Eq. (8). When pt is constant (which we assume for this paper) the price differential, , ensures that there is always a cost to adjusting herd numbers. Hence, the production index at time t, t, is a special case of the economic index (Eq. (8)) where qt and ct are zero, vt pt rt upt vt : 11

The production utility over a planning horizon T, Uproduction is the sum of the discounted production (t index over time T, ) Uproduction
T X t1

vtV:

12

There are many potential indices of ecological integrity, and we do not claim any single index is appropriate for all questions. We assume that the ecological agent aims to protect the soil from erosion by maintaining a high level of grass root biomass (GRt). We therefore assume that an appropriate ecological index for a rangeland is the sum of the production index at time t (t) and grass root biomass, gt mGRt vt : 13

cases the farmer will implement the strategies that aim to maximise one of these three utilities. Fundamentally, the farmer's management actions are constrained by rainfall. There is little the farmer can do to influence rainfall, but the farmer can stimulate grass productivity by manipulating fire regime and livestock numbers. In the paragraphs that follow we describe a series of functions that describe the farmer's fire and livestock management behaviour. Fire is a major tool in the arsenal of a manager of an extensive rangeland. The farmer can stimulate grass production by encouraging intense fires that in turn reduce tree biomass. The goal is to ensure that fires only burn once sufficient fuel for an intense fire has accumulated and when weather conditions are extreme (Trollope, 1983). Fires also consume grass shoot biomass, hence any benefits of burning needs to be offset by the cost of lost livestock production due to reduced shoot biomass. However, indefinite fire suppression leads to excessive accumulation of above ground dead grass biomass, which shades the living grass shoot biomass and thereby constrains grass shoot production (Trollope, 1989). Therefore, in many savannas burning has a net positive effect because it removes dead grass material and thereby stimulates grass growth. The parameter n determines the natural potential fire intensity below which a fire does not occur (cf. Eq. (5)). Hence we implicitly assume that fuel loads, and not ignitions, limit the occurrence of fire. The farmer can, through fire suppression, raise the natural fire spread threshold and thereby ensure that more intense fires occur or that fires occur less frequently (cf. Eq. (6)). We assume that the farmer suppresses fires as tree biomass increases. Because tree shoot biomass in the model fluctuates due to fire, it is not a stable indicator of tree biomass. We therefore use, in the model, tree root biomass (WRt) as an index of tree biomass (a farmer would probably use stem basal area as an indicator). The level of fire suppression the farmer applies at time t (t) is determined using the following function, dt dn dmax dn f WRt ; ad ; bd 15

The parameter is used to weight the contribution of GRt and t to t. How to appropriately weight these two fundamentally different indices (GRt and t) in a single utility index (t) is not clear. We use = 500, this ensures that the ecologically motivated agent derives the similar utility from a unit of grass root biomass (GRt) as from a unit of production (t). Hence, t as we have defined it is only useful within the context of this study. The ecological utility over a planning horizon T, U) ecological is the sum of the discounted ecological index (t over time T, Uecological
T X t1

here n is the natural fire threshold, max is the maximum level of fire suppression, a is the threshold root biomass at which the farmer suppresses fire, and b describes how rapidly the farmer suppresses fire as the threshold root biomass is approached. The major task faced by the rangeland farmer is livestock management. At each time point, t, the farmer can purchase t livestock or sell t livestock, in addition the number of livestock can grow due to its own density dependent growth process, yielding an equation for Zt+1, the livestock numbers at time t + 1,   Zt Zt1 Zt vt rt gZ Zt 1 Kt 16

gtV:

14

Here gZ is the biological growth rate of the livestock population and Kt is the farm's carrying capacity at time t. Kt is in turn defined by the consumption rate of livestock (), the available forage (GSt) and the farmer's investment in technology, Kt GSt f w; aw ; bw : l 17

3.2.

Rangeland management

In Section 3.1 we proposed utility functions for agents that are motivated by economic, production or ecological factors. In all

Hence, we assume that the technology level a farmer chooses, , determines how close the farmer can get to exploiting the

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potential carrying capacity (GSt/). The parameters a and b describe how the technology level chosen influences the proportion of the potential carrying capacity that can be exploited. An important part of a farmers challenge is timing his livestock sales and purchases. We assume that a farmer buys livestock when the prospects for production are good and sells livestock when the prospects for production are poor. Specifically we assume that the livestock a farmer purchases t is, b b vt Zt f Zt ; av ; bv ; 18

(Storn and Price, 1997; Price et al., 2005). Hence, we use the differential evolution algorithm to maximise either Ueconomic, Uproduction or Uecological over the planning horizon using the control set, C ar ; br ; rmin ; av ; bv ; dmax ; ad ; bd ; w: 20

b where Zt is the potential livestock the farmer could purchase b max(0, Kt Zt)) and a and b define what proportion of (Zt= these potential purchases the farmer realises. Hence, we assume that the farmer never overstocks the farm through purchases. The farmer sells some base proportion of animals, min, in every time step, but sells more livestock when the grass root biomass is below some critical point. This behaviour is summarised as, rt max1f GRt ; ar ; br ; rmin Zt ; 19

where t is the number of livestock to be sold at time t and GRt is the grass root biomass at time t. We use grass root biomass because this is, in the model, a more stable indicator of the condition of the grass sward than shoot biomass (shoot biomass fluctuates with rainfall, grazing and fire). In reality, the farmer would probably use grass cover, which is closely related to grass root biomass as an indicator of grass sward condition. The shape of this response curve is defined by the parameters a and b. In summary, the farmer is able to manipulate the fire regime by selecting the values of the parameters max, a and b; determine the level of the available forage that can be exploited by selecting a technology level ; determine a strategy for buying and selling livestock by selecting the values of the parameters a, b, a, b and min.

Note that the algorithm selects one control set for an entire simulation; that is within a single simulation run over the chosen planning horizon, the parameters remain constant. In essence this means that the farmers mind-set remains fixed. However, because these parameters summarise the farmers' response to changing conditions, the behaviour of the farmer is not constant. The reader should additionally be aware that the solutions found using the simulationoptimisation approach are only optimal given the constraints the management functions (Eqs. ), (15), (17), (18), (19)) impose. We choose a planning horizon of 100 years. While this may be on the upper interval of appropriate time scales, it does in combination with low discount rates (0.02 p.a.) select against non-sustainable strategies. We choose initial conditions where the relative sizes of tree and grass populations are

4.

Optimisation

The model is non-linear, stochastic and has multiple domains of attraction; these attributes make it poorly suited to classic optimisation methods (e.g. optimal control or dynamic programming). As an alternative, we use a simulationoptimisation approach, that is we combine our process orientated simulation model with a numerical optimisation algorithm. Simulationoptimisation methods have been successfully used to optimise hydrological (Aly and Peralta, 1999, Koutsoyiannis and Economou, 2003), biochemical (Mendes and Kell, 1998) and ecologicaleconomic systems (Janssen et al., 2004). We experimented with standard non-linear optimisation (quasi-Newton method of Byrd et al., 1995), genetic algorithm (Mebane and Sekhon's 2005 genetic optimisation using derivatives algorithm) and differential evolution methods (Storn and Price's, 1997 differential evolution algorithm as implemented by Ardia, 2005). Our initial tests suggested that differential evolution was more robust and efficient at finding the global solution than the other methods, a result confirmed by more systematic comparisons using benchmark problems

Fig. 2 Time series showing the development of tree (red), grass (green) and animal (blue) abundance over time. The points indicate the intensity of individual fires. The simulations illustrate the bush encroachment problem faced by rangeland farmers in savanna regions. The top panel shows a deterministic simulation, the middle panel uses the same control strategy as the top panel but occurs under stochastic conditions. The lower panel is a stochastic simulation with low levels of fire management. Plant abundances are the below ground biomasses. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

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similar and where animal numbers are less than the carrying capacity K (see Eq. (17) for a definition of K). In the case of stochastic optimisation it is necessary to consider the risk sensitivity of the agent. In the deterministic case we assume that the agent maximises the discounted utility over the planning horizon (cf. Eqs. (9), (12), (14)). For the stochastic case we need to consider the variance between different stochastic realisations of the same management strategy. Hence, for stochastic problems we optimise , b U EUevarU 21

selects against strategies that have a high risk of failure (collapse of the livestock population) when unfavourable sequences of rainfall occur. We used R (R Development Core Team, 2005) implementations of the optimisation routines. The model was written in C and compiled as an R package allowing us to use R to interface with the optimisation routines.

5.
5.1.

Model analysis
Model behaviour

here describes the level of risk aversion to the variance between stochastic evaluations of the same management strategy. For this study we assume = 0.5 for all benchmark analyses and use a sample size of 25 replicate simulations to calculate . Trial simulations showed that = 0.5 effectively

The system has three attractors that are of ecological interest, grass dominated, tree dominated and savanna (mixed tree grass) states. Scheiter and Higgins (submitted) provide a more formal discussion of the mathematical conditions necessary

Fig. 3 Time series showing grass (green), tree (red) and animal (blue) abundances for the economic (top panels), production (middle panels) and ecological (bottom panels) optima found in the deterministic (left panels) and stochastic (right panels) cases. All abundances are the below ground biomasses. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

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Table 3 Parameter values for the optimal control set selected by numerical optimisation for stochastic and deterministic cases Deterministic Parameter
a b min a b max a b

Stochastic Ecological
0.09 15.00 0.02 28.70 12.05 1.25 0.07 16.57 1.00

Economic
0.00 8.19 0.02 17.59 30.00 1.32 0.03 0.53 1.00

Production
0.00 15.00 0.03 17.35 30.0 2.16 0.03 23.07 1.00

Economic
0.01 8.98 0.02 27.09 28.63 2.07 0.37 9.53 0.04

Production
0.00 5.67 0.03 30.00 30.00 4.22 0.75 0.01 0.26

Ecological
0.00 6.90 0.03 29.65 30.00 4.14 0.49 0.01 0.13

for the savanna state. These analyses show that the savanna state is possible when either fire is present or when grasstree competitive interactions are weak. The weight of empirical

evidence for savannas, however, supports the notion that competition between grasses and trees is not weak (Scholes and Archer, 1997; Higgins et al., 2000; Bond et al., 2005;

Fig. 4 Animal sales, purchases and fire suppression strategies selected when optimising an economic (solid red), production (dashed blue), or ecological (dot dashed green) utility functions. The left panels are for the deterministic case, the right panels are for the stochastic case. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

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biomass, which leads to crashes in the animal population. The animal population is therefore only briefly at high densities, hence rangeland degradation does not occur. This time series

Fig. 5 Technology levels selected when optimising the economic, production or ecological utility function. Gray bars are for the deterministic case, black bars are for the stochastic case.

Sankaran et al., 2004). Hence, for all simulations presented here we assume that grasstree competition is intense (GW = WG = WG = 1) and that fire is necessary to prevent trees from dominating grasses. Other parameter values were selected so as to ensure (1) that grasstree coexistence is possible despite assuming the competition levels are high, (2) that tree dominance is possible under certain fire and animal stocking regimes and conversely, (3) that certain fire and animal stocking regimes can lead to a grass dominated state. Fig. 2 shows time series from simulation runs that illustrate important properties of the model. Since savanna systems are only productive rangelands when in grass dominated or savanna states, the central problem facing rangeland managers in savanna regions is to keep the system away from a tree dominated state. The upper panel shows a deterministic simulation initiated with moderate animal abundance. In this simulation the animal abundance rapidly increases, causing grass abundance to decrease which in turn allows tree abundance to increase. The increasing tree abundance further suppresses grass production to a point where fire is lost from the system; without fire trees increase further and grass is further suppressed. The loss of fire first leads to a pulse in animal numbers, because animals are no longer competing with fire for grass consumption. This increase in animal numbers, however, leads to animals dramatically overshooting the carrying capacity, and a final crash in animal numbers occurs. Subsequent elimination of animals from the system cannot return the system to a productive rangeland. Hence, the tree dominated state is stable and the system is, from the perspective of a livestock farmer, degraded. The degradation sequence described in the previous paragraph represents a classic rangeland theorist's view of overgrazing. A new rangeland theorist's view is that stochastic variation in rainfall prevents the animal populations from reaching high enough densities for long enough timespans to degrade range condition (Ellis and Swift, 1988). This weak coupling of range condition and animal numbers is illustrated in the second panel in Fig. 2. The management strategy adopted in this simulation is the same as that adopted in the deterministic simulation (top panel in Fig. 2), yet the stochastic nature of rainfall leads to occasional reductions in grass shoot

Fig. 6 Relationship between potential carrying capacity, technology defined carrying capacity Kt , and stocking rates Zt . The upper panel (showing median, quartiles and range) describes the distribution of the potential carrying capacity of over a 100-year simulation when optimising either the economic, production or ecological utility function under deterministic (gray bars) or stochastic conditions (black bars). The middle and lower panels show the cumulative density of Kt Zt for deterministic and stochastic cases respectively when optimising either the economic (red solid lines), production (blue dashed lines) or ecological (green dotdashed lines) utility functions. The cumulative density functions are based on a single 100-year simulation for the determinstic cases, and on 25 100-year simulations for the stochastic case. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

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represents the situation described by Desta and Coppock (2002) for Ethiopian rangelands. However, we emphasise that degradation is also possible in the stochastic case. The lower panel in Fig. 2 illustrates how degradation could set in rapidly when fire management does not encourage intense fires. Moreover, the optimal rangeland strategies identified in the next section, further emphasise that strategies for optimal rangeland management differ from those illustrated in the middle of Fig. 2.

5.2.

Optimal range management

The control sets for deterministic and stochastic optimisation of the economic (Eq. (9)), production (Eq. (12)) and ecological (Eq. (14)) utilities are presented in Table 3. Fig. 3 illustrates time series of grass, tree and animal numbers generated using these control sets, Figs. 4 and 5 summarise the control strategies adopted, and Fig. 6 illustrates how the stocking rates adopted relate to the carrying capacity. The time series show that in all cases the system is managed to ensure some level of grass dominance (Fig. 3). Under deterministic conditions, the strategies differ in the level of grass dominance: the ecological strategy has the highest level of grass dominance, while grass dominance is only slight under the production strategy. The ecological strategy achieves grass dominance by purchasing animals only when conditions are extremely favourable. This effectively delays the build up of the animal population, and this investment in the grass layer allows the ecological strategy to achieve the highest animal numbers in the last 20 years of the planning horizon. The economic strategy is cost sensitive (cf. Eq. (9)). It saves costs by opting for a less aggressive fire management strategy (Fig. 4 shows that fire suppression is lowest under the economic strategy). The ecological and production utility functions are not influenced by the costs of fire management or technology; both therefore use high levels of fire suppression (Fig. 4). The production strategy adopts the highest level of fire suppression (Fig. 4), which allows it to use fire to control tree density even though the fuel loads (grass biomass) are lower than under the other strategies. The optimal strategies under stochastic conditions qualitatively reflect the strategies adopted under deterministic

conditions (Fig. 3). Specifically all strategies ensure some level of grass dominance. However, the modal and peak stocking rates selected under stochastic conditions are lower than those selected under deterministic conditions (Fig. 3). Hence, even though grass biomass often peaks to levels above those observed under deterministic conditions, the optimal strategy is not to aggressively track these changes. This is exemplified by the low technology levels adopted by all agents under stochastic conditions (Fig. 5). The technology level defines how closely the carrying capacity tracks the potential carrying capacity (see Eq. (17)); hence the strategy adopted by all agents is to stock well below the potential carrying capacity. In all cases the stocking rate is such that the potential carrying capacity (Fig. 6, upper panel) is much greater than the stocking level (Fig. 3). Effectively this keeps a reserve of grass biomass that can be used to fuel fires which in turn prevents tree dominance. While the stocking strategy is more conservative under stochastic conditions, the fire management strategy is more aggressive (Fig. 4). The ranking of the intensity of the fire management strategies, however, remains as observed under deterministic conditions, that is the production agent is the most aggressive and the economic agent the least aggressive. The ecological and production agents adopt step functions for fire management, allowing them to implement high levels of fire suppression at low tree abundance. The economic agent's fire suppression function is a more gentle sigmoid function of tree abundance. As was the case under deterministic conditions, the economic agent saves costs by being less aggressive in fire management. Hence, in the economic case, tree dominance is avoided by a combination of fire management and low stocking rates. All strategies seek to keep Zt less than Kt as this avoids density dependent animal mortality. Under both deterministic and stochastic conditions the probabilities of Zt exceeding Kt are low (<0.0008, Fig. 6). Under stochastic conditions these probabilities are slightly higher (<0.009), representing the higher risks inherent in the stochastic environment. Although under stochastic conditions, the economic strategy stocks at lower levels than the ecological and production strategies (Fig. 3) it seeks to maintain Zt as close to Kt as possible (Fig. 6). Hence the ecological strategy's higher costs ensure that

Fig. 7 Utilities achieved when optimising either the economic, production or ecological utility function under deterministic (gray bars) and stochastic conditions (black bars).

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although the stocking rate adopted is lower in absolute terms, it is higher relative to Kt. Animals are only purchased when the potential number of purchases is high and even then only a small proportion of the potential purchase is realised (Fig. 4). Under deterministic conditions the ecological strategy is the most conservative; and the production and economic purchasing strategies are very similar. Under stochastic conditions purchasing only occurs when the potential purchase is higher than under deterministic conditions. In the stochastic scenarios, however, the potential purchase seldom reaches values above 15, meaning that animals are almost never purchased. In general, the purchasing behaviour observed suggests that the agents do not attempt to adopt a seasonal purchasing strategy or to react to opportunities presented by sequences of high rainfall years in the stochastic case.

optimal strategies we identified were sustainable. For our purposes, sustainability is when the utility in the second half of the planning horizon is not less than the utility in the first half of the planning horizon. The sensitivity of this sustainability index to the discount rate was explored by seeking optimal economic strategies under a variety of discount rates for both the deterministic and stochastic case (Fig. 8). This analysis shows that the deterministic strategies are sustainable even at discount rates as high as 10%, but that under stochastic conditions sustainability is lost at discount rates greater than 8%.

6.

Discussion

5.3.

Compatibility and sustainability

We explore the compatibility of the different agents' utility functions by examining the utilities achieved when optimising either the economic, production or ecological utility under stochastic and deterministic cases (Fig. 7). We do not present data showing that strategies selected under deterministic conditions perform poorly under stochastic conditions, as this point has been made by Janssen et al. (2004). We find, as did Janssen et al. (2004), that the utilities achieved under stochastic conditions are lower than those achieved under deterministic conditions (Fig. 7). This result is in agreement with stochastic harvesting theory that suggests that the yield from a stochastic resource system is lower than the yield from a deterministic resource system (Schmitt and Wissel, 1985; Lande et al., 1997). Selecting a low discount rate (2%), a long planning horizon (100 years) and forcing the farmer to have one management philosophy for the entire planning horizon ensures that all the

Fig. 8 The sustainability of the optimal solutions selected using the economic utility function as a function of the discount rate under deterministic (open circles) and stochastic (solid circles) conditions. The sustainability index is simply the normalised difference between the utility derived in the first half of the simulation relative to the utility derived in the second half of the simulation. Sustainable strategies have positive indices.

The new rangeland ecology criticises classic rangeland ecology's recommendation of constant stocking on the basis that this represents a lost opportunity to rangeland farmers, preventing them from taking advantage of bumper grass production years (Behnke and Scoones, 1993). One alternative to constant stocking is to attempt to force animal numbers to track variation in grass production. Although intuitively appealing, the feasibility of the tracking strategies suggested by the new rangeland ecology has been difficult to demonstrate (Illius et al., 1998). Our analysis further adds weight to the evidence that tracking strategies are not feasible. Solutions we present suggest that elements of tracking strategies, such as restocking after crashes in livestock numbers, or preemptively selling more animals when indicators of poor range condition are apparent are not optimal. More notably, our solutions suggest that it is optimal to stock at rates much lower than the potential carrying capacity. Our model results are the consequences of our assumptions regarding the relationship between grass biomass, carrying capacity, animal numbers and mortality. Since imprecise formulation of these relationships has fueled recent debates in rangeland ecology (Vetter, 2005), we feel it prudent to state explicitly what our assumptions in this regard are and how they relate to classic rangeland ecology and the so-called new rangeland ecology. Our key assumption relates to carrying capacity. Carrying capacity is defined in the ecological literature as the density beyond which growth ceases to be positive and becomes negative. In other words, carrying capacity is the density beyond which mortality exceeds natality. In our model carrying capacity is defined by grass shoot biomass. When grass shoot biomass cannot support the herd's nutritional requirements, animals die. For instance during a drought, grass production stops, yet the herd's nutritional requirements must still be met, hence some animals suffer density dependent mortality. This view is in agreement with evidence from savannas that suggests that animals die from hunger and not from thirst during droughts (Walker et al., 1987), and we are unaware of contrary evidence. While this view is consistent with classic rangeland ecology, it is contrary to the new rangeland ecology which views drought mortality as a density independent process (Sullivan and Rohde, 2002). The optimal strategy under stochastic conditions is to stock well below the potential carrying capacity at any given time (Fig. 6). The stocking level is therefore not defined as a function of peak grass shoot biomass, but rather as a minimum biomass

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needed to buffer the system against environmental variability. This buffer needs to be large enough to both carry animals through the dry season and to ensure that there is fuel for fires of sufficient intensity to damage trees. Seasonal variations in grass biomass ensure that carrying capacity varies with time; hence animal numbers tend to approach the carrying capacity during the dry season, but retract from it during the wet season. Animal numbers are, therefore as argued by Illius and O'Connor (1999), only loosely coupled to grass biomass in the wet season, but coupled to it in the dry season. To understand why tracking strategies are not optimal we need to consider the risks associated with tracking strategies. When aggressively tracking available forage the animal numbers at time t (Zt) are close to the carrying capacity at time t (Kt). In deterministic yet seasonally dry systems and especially in stochastically variable systems there is a chance that Zt exceeds Kt; this probability is greater when the system is managed so that Zt is close to Kt. When Zt exceeds Kt density dependent mortality occurs; should Zt greatly exceed Kt then over-compensating density dependent mortality occurs which leads to reductions in Zt to levels well below Kt. Our solutions show that for both deterministic and stochastic cases Zt almost never exceeds Kt (Fig. 6). Hence, stocking at conservative levels provides the farmer with a buffer and prevents the over-compensating density dependent mortality that occurs when Zt greatly exceeds Kt. Density dependent mortality is therefore, contrary to the view expressed by the new rangeland ecologists (Ellis and Swift, 1988; Behnke and Scoones, 1993; Sullivan and Rohde, 2002), more important under stochastic conditions. Our model suggests that degradation (e.g. transition to a tree dominated state) is avoided by stocking at low levels, a finding which agrees with classic rangeland theory. In contrast, the new rangeland ecologists propose that stochasticity prevents a coupling between animals and the resource base and thereby prevents density dependent processes from shifting the system into a degraded state. While we illustrate the plausibility of this idea (see Fig. 2), our analyses show that such strategies are not optimal. Our analysis raises several questions for future study, while some of these questions could be addressed with a modified version of this model, others call for empirical research to justify model parameters we used and assumptions we made. For instance, we considered only grazing animals, it would be interesting to explore how mixed feeders influence the results. Moreover, while our results suggest that tracking strategies are not optimal it would be interesting to seek the conditions under which tracking strategies might be optimal. Finally, we assume, as did Perrings and Walker (1997) and Janssen et al. (2004) that fire can be used to control tree densities. While empirical data from some savannas support this assumption (e.g. Trollope, 1983) the extent to which this assumption is appropriate to all savanna rangelands is not clear.

cesses influence rangeland dynamics. The results of our model analysis are that under stochastic conditions livestock farmers should be more conservative than under deterministic conditions. This result is perhaps surprising for those (e.g. Ellis and Swift, 1988; Westoby et al., 1989; Behnke and Scoones, 1993; Sullivan and Rohde, 2002) who intuitively think that farmers should be more aggressive and opportunistic under stochastic conditions. Yet, the idea that yield decreases with stochasticity is a well established concept in harvesting theory (Scmitt and Wissel, 1985; Lande et al., 1997) and should therefore not come as a complete surprise. Central elements of the conservative strategies we identify are to stock at low rates and to use fire to retard tree abundance. Two factors force rational agents to be more conservative under stochastic conditions. Firstly, high grazing levels reduce the buffering capacity of the system and hence increase the risk of a crash in animal numbers, which are expensive to recover from. Secondly, high grazing also increases the risk of a switch to a tree dominated and consequently degraded system. We use three different utility indicators to reflect the differing objectives of decision makers or agents. The results indicate that although the definition of the utility function influences the optimal strategy some generalisations emerge. Namely, it was optimal to select a strategy that was sustainable and led to grass dominance; furthermore all agents were more conservative under stochastic conditions. Identifying management strategies that are, in principle, sustainable is a necessary first step for achieving sustainability; additional steps are to (1) explore why savannas are often tree dominated even though our analysis suggests that it is rational to avoid tree dominance, and to (2) design policy instruments that encourage the adoption of the sustainable management practices.

Acknowledgements
We thank the Robert Bosch Foundation for financial support and the reviewers for valuable comments.

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7.

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