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Journal of Arid Environments 74 (2010) 307309

Contents lists available at ScienceDirect

Journal of Arid Environments

journal homepage: www.elsevier.com/locate/jaridenv

Short communication

Grazing systems are a result of equilibrium and non-equilibrium dynamics

J.F. Derry a,1, *, R.B. Boone b
a
b

Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, EH9 3JT Scotland, UK

Natural Resource Ecology Laboratory, Colorado State University, 1499 Campus Delivery, B234 NESB, Fort Collins, CO 80523-1449, USA

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 15 September 2008
Received in revised form
3 July 2009
Accepted 26 July 2009
Available online 18 August 2009

Paradigms about desertication and non-equilibrium dynamics are currently dominating discussions
about rangeland and pastoralist systems in semi-arid and arid zones. Climate variability is a major
determinant of the dynamics of these systems and the deleterious consequences of this variability are
increased for pastoralists for whom traditional mechanisms for coping with drought are increasingly
limited. Differential application and interpretation of terminology that relates ecological processes to
climatic variability are hampering the unication of research efforts across disciplines. Here we argue
that it would be benecial to resolve these differences by the correct use of terms introduced in the early
development of the ecological theory pertaining to these systems.
2009 Elsevier Ltd. All rights reserved.

Keywords:
Dis-equilibrium
Equilibrium
Grazing systems
Non-equilibrium

1. Introduction: a science that is stymied by semantics

2. Two approaches to management

The debate on rangeland dynamics is largely an unfortunate

result of confusion over terminology; equilibrium theory suggests
tight coupling of plantherbivore systems, based on Lotka Volterra models, and predicts that resource degradation and rangeland livestock system collapses are due to livestock overgrazing
the vegetation. Non-equilibrium is a term that has become
popular amongst rangeland scientists, sociologists and socioeconomists to describe the dynamics of grazing systems that are
susceptible to such a high level of climatic variability that
frequent droughts cause crashes in the animal populations
(Behnke et al., 1993; Scoones, 1994). In these systems the animal
populations spend most of the time recovering from the previous
drought, and rarely reach densities at which density-dependent
mechanisms act to moderate the animal populations. This has the
effect of decoupling the plantherbivore relations that historically
underpin equilibrium theory in grazing systems; hence the term
non-equilibrium and the formation of opposing camps. The
result is a debate founded on the semantics of terminology, rather
than evidence from the literature, and a continued misunderstanding between camps that are actually saying the same thing
(that rangelands are dynamic), but unfortunately are using
differing language.

If this was just an academic debate then it would be acceptable,

however, this difference in the views of researchers feeds into the
advice to policy makers working for the development of the livestock
industries in the semi-arid and arid regions of the globe (Cowling,
2000). Equilibrium theorists argue that climatic variability is the
single largest cause of poverty in pastoral societies, and ways need to
be found to buffer it by exploiting spatial heterogeneity, and by
managing the use of key resources. Conversely, from non-equilibrium theory it is proposed that because of decoupled plantherbivore
interactions livestock have little impact on forage resources (Fernandez-Gimenez and Allen-Diaz, 1999; Sullivan and Rohde, 2002;
Lind et al., 2003) and so are not to blame for the degradation of
rangeland that, in extreme circumstances, has been claimed to lead to
desertication (Dean et al., 1995; Sefe et al., 1996). As such, the policy
advice is to support the development of pastoralist coping strategies
to deal with climate variability, and that tenure and access issues are
more important than stocking rates, particularly when they exclude
pastoralists from land, water, minerals, markets and subsidies.

* Corresponding author. Tel.: 44 (0) 131 447 0878.

E-mail address: j.derry@ed.ac.uk (J.F. Derry).
1
Tel.: 44 (0) 7722510579.
0140-1963/$ see front matter 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jaridenv.2009.07.010

3. Understanding a complex system

Recent advances in rangeland ecology have disclosed the
complexity of these systems (Wu and Loucks, 1995; Bascompte and
Sole, 1998; Ritchie and Olff, 1999; Wu and Hobbs, 2002), to the point
that it is now possible to incorporate this complexity, especially
spatial heterogeneity in resource availability, into our concepts
about the dynamics of these systems (Briske et al., 2003). Rather

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J.F. Derry, R.B. Boone / Journal of Arid Environments 74 (2010) 307309

than it be an either-or situation, we argue that the system dynamics

of rangelands are better described as a continuum between equilibrium and non-equilibrium (Wiens, 1984), where everything in
between is in dis-equilibrium, and where the position along this
gradient is determined by the strength of coupling (sensu Vetter,
2005) between the plants and animals. The other way to consider
this is that the position along the gradient is indicative of neighbourhood stability (sensu May, 1974). Ultimately, this system-wide
response is the net effect of combined inuences from both areas
that do not constrain survival (typically wet season ranges) and
areas upon which the animals depend for key resources (dry season
ranges) (Sther, 1997; Illius and OConnor, 1999, 2000; Derry, 2004).
4. The strength of coupling between the plants and animals
We used two spatial models to investigate the strength of
coupling between the plants and animals in semi-arid grazing
systems (materials and methods are available as Supplementary
material). Both models use rainfall data to calculate primary
production for herbaceous and woody species, and its allocation to
plant parts. The selection and intake of these by animals dictates
the animals consequent energy and protein balances, body growth,
reproduction and mortality. These models therefore simulate
animal population dynamics mechanistically, and are able to
predict the ecological carrying capacity of the system. Spatial
aspects of these models include topography, soil moisture and
nutrient distribution, surface water dynamics, plant distribution,
drinking water location and animal spatial foraging.
Our rst result shows how it is possible to model a gradual
decline in animal density through manipulation of the variation in
rainfall alone (Fig. 1).
Our nding provides rare evidence that the system dynamics of
rangelands describe a continuum between equilibrium and nonequilibrium. There have been previous suggestions about a criticality for the predominance of non-equilibrium dynamics

occurring in the coefcient of variation for rainfall around 3033%

(Shepherd and Caughley, 1987; Ellis et al., 1993; Ellis and Galvin,
1994). At this level, drought frequency has been estimated to cause
50% stock mortality (Ellis and Swift, 1988). Our result conrms
these quantities, but suggests a continuous response in place of the
equilibrium-to-non-equilibrium discontinuity that was concluded
to exist for rangeland systems (Boone and Wang, 2007), or suggested by the theoretical possibility of multiple equilibrium points
or states (Sutherland, 1974; May, 1977) and the conceptual state and
transition model (Westoby et al., 1989) that followed.
Thus, it is more accurate to consider the range of system
dynamics as a gradual and continuous increase in decoupling,
showing no evidence of limits between discrete states, as it would
be expected by state and transition models.
With another of our models we simulated climatic variability on
20 parcels summing to 1000 km2 of South African grazing land
(Boone and Wang, 2007). The linkage between forage availability
and livestock dynamics weakened as climate was more variable,
such that at the highest CV all but one response was density
independent [bars enumerating density independence in Fig. 2,
coming from Boone and Wang, 2007], but the variability in the
strength of this linkage across the 20 parcels (open circles representing density dependence in Fig. 2) was striking.
Large variation in the strength of density dependence across
parcels under the same climatic patterns suggests that the terms
equilibrium and non-equilibrium are over-simplications when
applied to landscapes.
5. Concluding remarks
From our modelling we argue that the policy advice that is given
should not be contradictory, but be seen as complementary. The
reason coping mechanisms exist is to adapt to climatic variability.
In the absence of such drought-coping strategies, the impact of
rainfall variability is real and is in the scale depicted by our ndings,

Fig. 1. Decline in stocking rate (normalised into the range 0 1) with increasing variability in mean annual rainfall, with annual rainfall held approximately constant, supporting the
understanding that rangeland dynamics describe a continuum between equilibrium (tight coupling of consumers and resources) and non-equilibrium (weak coupling) (sensu
Vetter, 2005). The inexion point for the tted sigmoid curve coincides with approximately 33% inter-annual variability in rainfall.

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J.F. Derry, R.B. Boone / Journal of Arid Environments 74 (2010) 307309

309

References

Fig. 2. The strength in density dependence (DD), reecting the position along the
equilibrium-to-non-equilibrium continuum, for simulated cattle populations (open
circles, primary axis) on twenty 50 km2 parcels of South African grazing land as the
variability in rainfall was altered. Responses near zero were labelled density independent (DI) (bars, secondary axis). Large variation in the strength of density dependence across parcels under the same climatic patterns suggests that the terms
equilibrium and non-equilibrium are over-simplications when applied to landscapes.

e.g., 33% CV roughly equates to 50% stock, etc. Allowed tradition

practices, that impact is alleviated, giving a larger capacity for stock,
by effectively diminishing rainfall CV.
For example, transhumant livestock keepers such as the Maasai
exploit the spatial heterogeneity of available grazing lands during
dry periods, not necessarily waiting for drought conditions before
moving to dry season grazing areas. Traditional knowledge knows
that these places contain key resources of forage and water during
the dry season. But, as available grazing lands are diminished and
movement channels are blocked through land use change for crop
cultivation and commerce, and through urbanisation as a result of
population density increases, the option for employing these
coping mechanisms (especially transhumance) is being removed.
Hence, through land tenure changes and increasingly restricted
access to dry season grazing sites, the impact of climatic variability
is emphasised for resource poor pastoralists who are being limited
in the ways that they are allowed to cope with drought (Homewood, 2006). Ironically, it is these mechanisms that pastoralists
employ to cope with drought that has largely inuenced attitudes
in the non-equilibrium camp, yet, removal of these mechanisms
can only result in the strengthening of the plantanimal interactions at the core of understanding in the equilibrium camp.
Acknowledgements
Results from Model 1 were produced by JFD in collaboration
with Andrew Illius and Iain Gordon. Results from Model 2 were
produced by RBB in collaboration with Guiming Wang. We gratefully acknowledge the constructive comments from two anonymous referees on earlier versions of the manuscript. This work was
prompted from a discussion between JFD and Kathy Homewood.
Appendix. Supplementary data
Supplementary data associated with this article can be found, in
the online version, at doi:10.1016/j.jaridenv.2009.07.010.

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