WATER RESOURCES RESEARCH, VOL. 35, NO.

12, PAGES 3709-3722, DECEMBER 1999

On the spatial and temporal links between vegetation, climate,

and soil moisture

I. Rodriguez-Iturbe,P. D'Odorico, A. Porporato,• and L. Ridolfi•

Departmentof Civil and EnvironmentalEngineeringand Center for Energyand EnvironmentalStudies
PrincetonUniversity,Princeton,New Jersey

Abstract. The impact of climatefluctuationscan be observedin the dynamicsof

vegetationand mostparticularlyin the sensitiveenvironmentof savannas.In this paper we
presenta modelfor the local competitionfor soil moistureamongneighboringvegetation.
The initial conditionfor the modelis a randomfield where at eachpoint the soil moisture
is the meanwater contentwhen there are no spatialinteractionsbetweensites.The mean
soil moisturevaluesaccountfor stochasticityof climate and lossesfrom evapotranspiration
and leakagewhich dependon the existingwater content.A spatialdynamicsis then
implementedbasedon the explicitminimizationof the globalwater stressover the region.
This approachexplainsthe coexistenceof herbaceousand woodyplantsin savannasas
well as the changesin canopydensitythat have been documentedin the southwestof the
United Statesas a function of regionalclimaticfluctuations.

1. Introduction tion functionof soil moisturedependson climate, soil, and

vegetation,and oncederived,it is usedto obtainexpectedsoil
Soil water availabilityis recognizedas the controllingre- moisturevaluesat the different sitesin a preexistingfield of
sourcein the organizationand functioningof many ecological soil and vegetationunder stochasticclimate conditions.The
systemsamongwhich savannasare an important and charac- resultingfield of mean soil moisturevalues representsthe
teristicexample.Thus, althoughfire regime,grazingintensity, expectedconditionsif spatialinteractionsare nonexistent. The
and especiallynutrientavailabilityare amongthe factorsinflu- field,takenjointlywith the spatiallyheterogeneous vegetationin
encingecosystemfunctionin savannasand other biomes,it is the region,yieldsa field of water stresses which,in turn, control
commonlyacceptedthat if water is limited,it becomesthe key the spatialinteractionsin the competitionfor soil moisture.
resourceaffectingvegetationalstructureandorganization.The The paper continueswith a descriptionof a schemeusedto
mechanisms throughwhichwater limitationaffectsecological derive the mean soil moisturefield in the presenceof spatial
systems are of manydifferentkinds,includingwell-knownones competitionfor the resource.It is assumedthat interactions
related to carbon assimilationvia control of photosynthesis take place toward the minimizationof the globalwater stress
and stomatalclosureas well as nitrogenassimilationthrough over the region[Rodtiguez-Iturbe et al., 1999a].We showthat,
the controlof the nitrogenmineralizationrate [e.g.,Scholes dependingon the climateand soil conditions,there existmin-
and Walker,1993].All thesemechanisms are intimatelyrelated imum stressoptimathat allowcoexistence of differentspecies,
to soilwater availability,and their combinedeffect on a par- for example,herbaceousand woodyplants.A similarspatial
ticular plant is termed water stress.This paper expandsthe dynamicsoriented toward the maximizationof biomasspro-
analysisandresultspresentedbyRodtiguez-Iturbe et al. [1999a] ductivity[e.g.,Eagleson,1982;Eaglesonand Segarra,1985] is
and providesa new framework for studyingthe impact of shownto yield comparableresults.Climatic fluctuationsmain-
climatefluctuationson patternsof vegetation. tained duringseveralyearslead to changesin optimalcombi-
Before consideringinteractionsamong neighboringplants nationsof vegetationtypesthat canbe contrastedwith existing
arisingfrom spatiallyvariable soil moistureavailability,it is field data collectedthroughoutthe last 50 years.
importantto reflect on the water balancedynamicsat a point No attemptis made to model a specificevolutionarydynam-
ics of vegetationunder stochasticrainfall conditionsor multi-
and on its drivingelements,climate,soil, and vegetation.The
year climatefluctuationsin this paper. Rather the objectiveis
quantitativedescriptionof this dynamicsis not a trivial task.
to showthat optimal coexistenceof different typesof vegeta-
Thispaperstartswith a descriptionof the soilmoisturebalance
tion reflectthe conditionsobservedin naturefor regionswhere
at a point,which allowsus to defineand quantifythe possible
soil moistureis a dominantand controllingresource.
presenceof water stressat a sitewhenthere is no consideration
of spatialdynamics.Soil moistureat a point is dependenton a
stochasticclimate controllingthe rainfall inputsas well as on e Water Balance at a Point
the lossesfrom evapotranspiration and leakagewhich,in turn,
In the absenceof pronouncedtopographicaleffectsthe soil
dependon the stateof soilmoisture.The probabilitydistribu- moisturebalanceequationat a point is written as
ds
•Alsoat Dipartimento
di Idraulica,Trasportie Infrastrutture
Civili,
Politecnicodi Torino, Torino, Italy. nZr• = I(s,t) - E(s,t) - L(s,t), (•)

Copyright1999by the American GeophysicalUnion. where

Papernumber1999WR900255. n porosity;
0043-1397/99/1999WR900255 $09.00 Zr depth of soil;
3709
3710 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE

parentrainfallprocess
nowoccurs
witha rateX' = Xe-"a
o(s) [Rodriguez-Iturbe et al., 1999b].
Lossesin (1) are from evapotranspiration and leakage.For
a giventype of plant the assumeddependenceof evapotrans-
piration on soil moistureis shownin Figure 1. Evapotranspi-
ration increaseslinearly with soil moistureuntil it reachesa
maximumvalueE*, whichtakesplacewhen moistureis above
a thresholds*. More realistically,the value of E* shouldbe
made dependenton the leaf area index and climaticcharac-
teristicsfunctionof temperatureand wind, but the represen-
tation of Figure 1 is consideredappropriateat dailytimescales
0 s• 1.0 s
under seasonallyfixed conditions[Gardnerand Ehlig, 1963;
Figure 1. Evapotranspirationand leakage as a function of Spittlehouse et al., 1981;Dunin et al., 1985].The value of s* is
soilmoistures in vegetatedsoils.The verticalaxis,p(s), rep- frequentlyassociated with the field capacityof the soil.For our
resentsthe total lossesfrom the system.The valuess• and s* purposesit is important that this grossapproximationis not
representthe relative soil moisture at which leakage starts made blindly and that both E* and s* reflect the type of
being consideredand evapotranspirationoccursat its maxi- vegetationat the site.Thus both E* and s* are different,say,
mum value, respectively.K, standsfor the hydraulicconduc- for the treesandgrasses commonlyfoundin savannas (e.g.,E*
tivity of the soil, and E* standsfor the maximumdaily evapo-
around4 mmd-1 for treesandnear20%morefor grasses).
transpirationfor the site.
These differenceshave important consequences on the soil
moisture dynamics.When s -> s*, evapotranspirationtakes
placeat its maximumvalue,and the vegetationdoesnot suffer
stress.For s < s*, vegetationoperatesunderwater stresswith
s(t) relativesoilmoisturecontentor saturationlevel; the resultingdecreasein the rate of the plant physiological
I(s, t) infiltrationrate from rainfall; processes.
E(s, t) evapotranspiration rate; Lossesfrom leakageare at maximumequal to the saturated
L (s, t) leakageor deepinfiltrationrate. hydraulicconductivityof the soilK, when the soil is saturated
Notice that all terms on the right-handsideare functionsof (s = 1). For s < 1, leakagedecreases
followinga powerlaw,
the stateof relative soil moisturecontents. The followingis a K(s) = K,sa, wheretheexponent
d depends
onthetypeof
synthesis of a sectionfrom Rodriguez-Iturbe et al. [1999b]that soil [e.g., Clappand Hornberger,1978].Rodriguez-Iturbeet al.
derivesthe steadystateprobabilitydistribution of s whenI(s, t), [1999b]approximated thispowerlawby the solidline segments
E(s, t), andL(s, t) are describedasfollows.Rainfall arrivals shownin Figure 1; the value of s1 dependson the type of soil.
are assumedto be Poissondistributedwith rate parameter X. They derivedthe equilibriumor steadystate probabilityden-
Everyrain eventhasan associated depthr, characterizedby an sityfunctionp (s):
exponentialdistributionwith parameter a, where 1/a is the
mean depthof storms.The parametersX and a are assumedto c s
0 (S •S*
be constantthroughouttime. The analysisis performed at a • e-Y
s
seasonalscale, which for purposesof this paper may be
thoughtof as the growingseasonof the region.No consider- p(s) = c
-e •e S* (S •S•
ation is made of the temporal structureof stormswhich are 3,(1
Sl-S*
assumedto be concentratedat a point in time. Although the
formulation is continuousin time, the interpretation of the cIk(s-sl)]- e-W+x•-- S1< S• l,
(2)
water balancedynamicsis at the daily timescale.Hourly fluc-
tuationsin the evapotranspiration rate alsoare neglected.Pre- where r•, % and k are normalizedquantities;r• = E/nZr, 'y =
cipitationis thusmodeledasan intermittentprocessof rainfall otrtZr, and k = gs/rtZ r. The constantc makesthe area under
eventswhich arrive randomly in time with an average fre- the densityequal to one and is given explicitlyby Rodriguez-
quencyof X eventsper dayand an averagedepthdescribedby Iturbeet al. [1999b],who alsoprovidea long expressioncorre-
the mean precipitationamong the rainy daysof a homoge- spondingto the expectedvalue of s. They showthat the equi-
neousseason
(e.g.,X = 0.167day-1 anda-1 = 1.5cmevent
-• librium probability density function and its moments vary
for the growingseasonat the Nylsvleyregionin SouthAfrica). drasticallydependingon climate, soil, and vegetationcharac-
The amount of rain that infiltrates into the soil from any teristics.In naturethe lossesfrom evapotranspiration
andleak-
particularstormis assumedto be equalto the depthof rainfall age as function of soil moisture are smoothedversionsof
wheneverthere is enoughstorageavailablein the soilto accu- Figure 1 that, nevertheless,is an adequateand commonlyused
mulate the full depth.If rainfall exceedsthe availablevolume, representationfor the purposesof this paper.An equilibrium
then runoff is generated;this is the mechanismof saturation approachis usedin this paper becauseit is muchsimplerand
from below studiedin detail by Dunne [1978].The infiltration because,aspointedout by Tilman [1982],it exploresthe long-
into the soil is thus a random variable that dependson the term effectsof the competitionfor resourceswhich are the
climatic characteristicsof the region as well as on the soil focusof this paper.
moistureat a site.Lossesfrom interception,that is, the part of
rainfall whichis interceptedby vegetationand evaporatesbe-
fore reachingthe soil, can also be easilyincorporatedby as- 3. On the Multiple Impacts of Soil Water Content
suminga thresholdfor rainfall depth A, belowwhichno water Mechanismsof competitiondependon the typeof resource,
effectivelyreachesthe ground.Under this assumptionthe ap- the consumptioncharacteristicsof the species,andthe process
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3711

1.5
governingthe supplyof resources[Tilman, 1982]. All things Field capacity
consumedby a speciesare potentiallylimiting resources,but I

here we will assumethe limiting resourceis water either be-

causeof its direct effects(e.g., controlof carbonmetabolism
throughregulationof photosynthesis) or becauseof its indirect
influence(e.g.,dominanteffecton nitrogenmineralizationand
thus on nitrogenassimilation).This is not an unrealisticcon-
siderationin many natural ecosystems amongwhich savannas
are a prime example.Moreover,the dependenceof vegetation
on many nutrientsfrequentlyfollowsthe type of relationship
shownin Figure 2 for nitrogenmineralization,increasinglin-
5 10 15 20 25 30 35
early with soil moistureto a point beyondwhich it remainsat
a maximumvalue.Notice how similarthisrelationshipis to the Gravimetric
soilwatercontent
(%)
one of evapotranspiration as a functionof soilmoistureshown Figure 2. Rate of nitrogen mineralization as a function of
in Figure 1. In the caseof Figure 1, transpirationis an un- soil water content in an Acacia Savanna [from Scholesand
avoidableresult of the photosynthetic processby whichplants Walker, 1993].
convert atmosphericCO2 into necessarycarbohydrates.The
gascan only enter the plant when dissolvedin water, and the
stomatalcavitiesare where the dissolutiontakes place, with leafedor fine leafedwith differentcodominantspeciesof trees
water suppliedmostlyfrom the roots.The movementof water and grasses,mainly dependingon whether soilsare nutrient-
throughthe plant from roots to stomatalcavitiesdependson poor or nutrient-rich.Soil type and soil nutrientsplay specific
manygradientswithin the plant and betweenthe plant and its roles which cannotbe incorporatedinto the soil water effect.
surroundings originatingfrom transpiration.The movementof However, soil moisture availabilityis the controllingvariable
water up in the plant leadsto water contentgradientsbetween whicheffectivelyembodiesa full rangeof effectswhen study-
roots and the surroundingsoil which, in turn, producemove- ing the spatial structureof trees and grassesin a region. The
ments of moisture toward, and into, the roots. Soil moisture regionitself is a randomfield of soil propertiesand available
controls how nutrients are supplied to plant roots mainly nutrients under the influence of a variable climate.
throughthe massflow of ionsinto the rootswith the soilwater This is not to saythat the random fields describingsoil and
and throughthe diffusionof ionstowardthe root surfacewhen climateare themselves independentof the spatialvariationsin
ion uptakeratesexceedsupplyof nutrients[Eissenstat and Van vegetation. On the contrary, they are quite related, but the
Rees,1994]. Thus it is no surprisethat, as long as there exist timescaleinvolvedin the generationof soilpatterns,both hor-
adequatenutrientsin the soil, the supplyof nutrientsto the izontallyandvertically,is muchlargerthan the one involvedin
plantshighlydependson soilmoisture,and,moreover,the type the patternsof vegetation.However,the spatialscalesinvolved
of functionalrelationshipbetweennutrientsand soil moisture in the feedbackfrom vegetationto rainfall are, in general,
is similar to the one of Figure 2. much larger than the vegetationpatternswhich concernus in
Althoughthe compositionof vegetationand its spatialstruc- thispaper.They are mainlyrelatedto environmentalgradients
ture is influencedby manysoilfactors,whenwater is a limiting (e.g.,temperaturegradients)amongregionswith characteristic
factor,the soilmoisturebalanceis the mostimportantvariable distances of several tenths to hundreds of kilometers interact-
controllingvegetationpatterns [Tinley, 1982]. This does not ing amongthemselvesin evenlarger regions.The dynamicsin
imply that the soil nutrient contentsare not important in de- thispaper is all at a locallevelwherethe sizeof the interacting
terminingthe heterogeneityand diversityof vegetation.Nitro- unitsdependson the vegetationand is typicallyof the order of
gen and phosphorusare essentialfor the growthand function- 10 m. The total regionmay,nevertheless, be quite large (e.g.,
ing of all plants, and their availabilityin the soil is crucialin
characteristiclengthsof tenthsto hundredsof kilometers).
terrestrialecosystems. For example,accordingto Scholesand In the caseof soil nutrients,besidesthe inherentvariability
Walker[1993],primaryproductionin someAfrican savannas existingin the soil, there is also a direct and mutual feedback
is
not constrainedby the carbon cycleor any of its controlling betweenthe presenceor not of vegetationand the richnessof
factorsbut by the rate of uptakeof nitrogenandphosphorous. the nutrient'spool. Thus at Nylsvleythe total nitrogencontent
The rate of uptake is constrainedby the releaseof inorganic of the soilbeneatha tree canopydownto a depthof 1 m is 45%
nitrogen(NH•-), produced bymicrobialattackondeadorganic higher than the nitrogen contentbetweentrees [Scholesand
matter, into the soil solution,a processcalledmineralization. Walker, 1993]. With respectto random variability in the soil
Temperature and moisture conditionsstronglyaffect the de- itself, Tilman [1982] found remarkablespatialfluctuationsin
compositionand mineralizationrates,with the net result that total nitrogenand extractablemagnesium.Theyvariedby 42%
in savannas the ratesof uptakeare controlledmainlyby the and over 100%, respectively,within a single12 m x 12 m plot
water content of the soil. Thus "the linkage between rainfall of sandysoil in Minnesota.
and primaryproduction,sowidelyobservedin savannas,prob- As shownin the abovediscussion, the soilwater is, through
ably operatesvia the influenceof water availabilityon the its multiple impacts,a preeminentfactor in determiningthe
nitrogen and phosphorouscycles,which then constrainthe distributionof vegetation.As extensivelydiscussedby many
carboncycle"[Scholes and Walker,1993,p. 110].The key effect authors[e.g.,Langeet al., 1976, 1982],vegetationresponseto
of water in this caseis the control of the availableinorganic water deficitdependson plant speciesaswell ason the severity
nitrogennot the control of the photosynthesis. The soil itself and durationof the deficit.The decreasein physiological func-
maybe nutrient-pooror nutrient-rich,and thiswill controlthe tion, for example,plant stress,intensifieswith the water deficit,
specifictype of plantswhichdominatethe ecosystem. Thus in and additional processesbecome affected with a nonlinear
the African regionof Nylsvley,savannas are classifiedasbroad type of relation betweenthe magnitudeof the stressand its
3712 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION,
grasses
trees
¸
0 Sw,g
Sw,t Sc,t Sc,g 1 S
Figure 3. Evapotranspiration and water stressas a function
of relative soil moisturefor trees and grassesin savannas.
Consequences.
Underconditions
of relatively
lowstress,
plant
growthandproductivityare somewhatrestricted,andthe plant
is forced to adjust and adapt to the stresssituation.Under
highervaluesof stressthe plant strugglesto survive[Bradford
and Hsiao, 1982].As mentionedbefore,water stressbeginsfor
a plant when the soilmoisturedropsbelows* in Figure 1. We
will measurethe water deficitas the difference(s - s*) for
s < s* and zero otherwise.When s reachesa value equal to
the permanentwilting point of the plant Sw, the stressis as-
sumed to reach a maximum value; for conveniencewe take it
asequalto one.Althoughthe effectsof stressvarywith time as
a functionof the water deficitin the precedingperiods,we will,
nevertheless,considerthem as invariantin time for any given
plant species.FollowingRodtiguez-Iturbe et al. [1999a], the
plant stressis thus modeledin this paper as
•--
IS*
--S]q S•S*
S* -- Sw
0 otherwise
(3)
The valueof q variesby speciesand evenamongindividuals.It
attemptsto representthe nonlineareffectof the water deficit
on the plant response.Mostly,we usedq - 2 for all plants,but
different valuesof q have alsobeen tried.
The stressthat a plant undergoesvariesthroughtime de-
pendingon a time historyof soil moisturethat, at any site,is
itselfa time-varyingfunctionof climate.We averageconditions
over a growingseasonso that the corresponding mean water
stressis estimatedfrom the averagesoil moistureat each site.
Given the nonlinearitiesof the dynamics,it is not strictlycor-
rect to assumethe stresscorrespondingto the averagesoil
moistureis the mean water stress.Moreover, a completehy-
drologiccharacterizationof the stressshouldincludethe sta-
tisticalstructure
of the durationandfrequency of the stress
periods.Researchon thesetopicsisverymuchneeded,but, for
the purposesof thispaper,where evolutionarydynamicsstart-
ing from initial conditionsis not considered,characterization
by the mean soil moistureis sufficient.The mean relative soil
moistureis,in turn,calculatedfrom the distribution,
p (s), with
givenclimate,soil,andvegetation
characteristics.
The stress
•,
estimatedfrom (3) with the expectedsoil moistureand the
CLIMATE, AND SOIL MOISTURE
valuesof s* and Sw correspondingto the site, is assumedto
representthe magnitudeof the impactthat, on differentphys-
iologicalprocesses,the randomlyvaryingwater deficithas on
the vegetationat the site.
A temporal scale and a spatial scale have been implicitly
embedded in the above framework. The timescale over which
temporalvariation is most important is taken to be approxi-
mately the length of the growing season.The spatial scale
defininga site corresponds
to the area or volumein which an
individualobtainsresourcesduringthe season[Tilman,1982].
4. Spatial Interactions in the Competition
for Soil Moisture
Considera regionheterogeneous in soil andvegetationrep-
resentedby a grid with squarecellswhichtypicallycorrespond
to the canopycoverageof the averagevegetation(e.g., 5 m x
5 m or 10 m x 10 m for the typicalexamplesconsideredin this
paper).The areacoveredby rootsof the individualplantsis,in
general,largerthan the individualcell to allowfor competition
amongneighboringvegetation.We assumethat the whole re-
gion is under the sameclimaticconditionsand, to make things
simpler, that there is a well-defined growing seasonwhere
rainfall may be modeledby the schemepresentedin section2.
From the dependenceof evaporationon soil moisture,one
can visualizethat different kinds of vegetationwill undergo
differentstressconditionsunder any prescribedclimaticchar-
acteristics.An exampleof thisis shownin Figure3 for a typical
caseof trees and grassesin savannas.The differentvaluesof
E*, s*, and Sw for trees and grasseslead to stressand soil
water curvesthat intersectregardlessof the particular expo-
nent q usedin (3). In Figure 3, soil moisturevaluesaboveA
favor trees versusgrasses,and soil moisturevaluesbelow A
favor the existenceof grasseswhen a site is consideredin
isolationfrom its neighborsand no spatial interactionstake
place.Moreover, the soilmoistureat any siteis itself a random
variablewhich dependson soil, climate, and vegetationchar-
acteristicsas describedin section2. A region under the same
climatic conditionsbut with spatiallyheterogenoussoil and
vegetationis characterizedinitially by a field of averagesoil
moisture conditionsat each site and by the corresponding
vegetation stressvalues when there are no interactionsbe-
tween the neighboringsites.
Next, we will assumethat competitionfor soilmoisturetakes
place locally and depends on stressesamong neighboring
plants.We are interestedin equilibriumsolutions,andthusthe
effect of the spatialinteractionswill be studiedon the time-
averagedinitial field. The spatial dynamicscontrollingthe
competitionfor soil moistureis assumedto globallyminimize
the vegetationstressresultingfrom deficitsof soilwater con-
tent. There is no attempt in this paper to model the temporal
evolutionarydynamicsof the vegetationsystem.With given
soil,climate,and vegetationfieldswe estimatethe spatialsoil
moisture field which leads to a minimum global vegetation
stress.Furthermore,we also can investigateamong different
spatialvegetationfields (e.g., different percentagesof trees
versusgrasses in savannas) whichone is in equilibriumwith a
soil moisture structurecorrespondingto a minimum global
stressover the region.In this schemethe competitionfor soil
moistureis alwayslocal,but the goalis a globaloptimum.This
dynamicsis commonin nature where competitionfrequently
leadsto associations benefitingmany partners.The competi-
tion is alwaysbetweenindividualorganismsrather than spe-
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3713

cies, which are only indirectly competingentities [Bonner,

1988].In the local competitionamongindividuals,there exists p(s)
a considerableamount of randomnessregardingwhich of the
neighborsare most affectedand how much of the resourceis
traded in any given situation.
The dynamicsmodeledhere is extremelysimplebut incor-
poratesthe abovementionedcriteria [Rodriguez-Iturbe et al.,
1999a]asfollows:(1) A site(i) andone of its neighbors(n) is
chosenat random.(2) The neighborsubtractsan amountof ,
soil moisture• from the site i accordingto

s• 1.0 s
r/= (1- •)(s•
0 - otherwise,
Sw.)• s•->
Sw. (4) Figure 4. Lossesof moisturedue to evaporationand deep
where •i is the vegetationstressin i and e is a randomvariable infiltration in bare soils.

uniformin [0, 1]. (3) The newstresses •i and •,• are computed,
and if the globalstress•i•i decreases, the abovedynamicsis advantageous.The soil water lossesfrom a bare soil site are
accepted,andthe procedureis repeatedin a newsitechosenat modeledhere as schematicallyshownin Figure 4, where s• is
random.If the globalstressincreases, the changesin soilmois- the sameas in Figure 1 andE* now representsthe maximum
ture do not take placeand the processis restarted.(4) If the rate of evaporationfrom the soil. This schemeis obviouslya
site (i) wasalreadysubtracting water from the neighbor(n), crudesimplificationof a verycomplicatedprocessinvolvingthe
then the rule is to stop the transfer if the global stressde- unsteadydryingof a soil profile without water table for which
creases.(5) The processis repeateduntil no further decrease no general exact solution has been obtained yet [Brutsaert,
is detectedin the globalstressafter an arbitrarilylargenumber 1982].The curvatureof Figure 4 is the oppositeof that of the
of interactions. vegetatedcase(Figure 1) as shownby Philip [1957].The value
Sincethe transfersof soil moisturego directly into the re- of E* is generallylower than the one occurringon vegetated
ceivingvegetation,the soil at the receivingsite doesnot actu- sites,but the bare soil generallyhaslessmoisturethan the one
ally increaseits water contentalthoughthe existingvegetation found under vegetation. The main reasonsfor this are the
will lower any existingwater stress.The givingsite, however, following:(1) In the presenceof vegetation,root penetration
will decreaseits soilwater content,and the existingvegetation inducesthe presenceof macroporesand preferentialinfiltra-
will likelyincreaseitswater stress.Thusthe competitionallows tion paths,and (2) lack of vegetationfrequentlyleadsto seal-
for moistureto be subtractedby a plant from a drier neighbor. ing of the soil surfaceas the resultof raindropimpacts.Rain-
As explainedbefore, the simulatedtransferof water is from drop impactsdetachsoil, destroygranulation,and may cause
the soil of the neighboringpixel directlyto the roots of the appreciabletransportationof soil. The small particlesof the
nearbyplantwithoutchangingthe soilmoisturecontentof the dispersedmaterial tend to washinto and clog the soil pores,
receivingpixel. For purposesof vegetationstress,with the sealingthe surfaceand drasticallyreducinginfiltration [Brady
influx of moisturethe receivingplant experiencesa reduction and Weil, 1996]. When the soil dries, a hard crustfrequently
in the stressconditionalthoughthere is no actual increasein forms that, if large enough,inducessurfacerunoff from sub-
the moisture content of the soil. Conservation of soil moisture sequentprecipitation.Such runoff has been suggestedas an
is preservedduringthe interchange. importantfactor in the developmentof patternedvegetation,
The analysesand resultspresentedin this paper consider suchas the so-calledtiger bush formed in some semiariden-
only two typesof vegetation,usuallytreesand grassescompet- vironments[Bromleyet al., 1997]. Depending on the climate
ing amongthemselvesand againsteachother for a scarceand andsoilcharacteristics, the presenceof bare soilmaybe locally
determiningresource:soil water. The framework presented beneficial for the nearby vegetationin the short term, but
here canbe appliedwithout changesto caseswhere there are sealingof the surfacesoonoccurswith the accompanying det-
rimental effects. In nature the occurrence of surface runoff
more than two functionaldependencesof evapotranspiration
on availablesoilmoisture.Thus, althoughour studyis centered tendsto produceclusteringof bare soil areas,but the compar-
around savannas where the main functional differences are isonspresentedhere assumebare soilsitesare spatiallyuncor-
related.
between trees and grasses,one, nevertheless,could apply a
similarapproachto categorieswithin thesegroups.One only Surface sealingon bare-soil sites is accountedfor in this
needsto assigndifferentvaluesfor Sw,s*, andE* accordingly studybythereduction
of themeandepthof thestorms,
a-•, in
to the vegetationtypebeingrepresented.Moreover,the pres- the model describedin section2. We assumethat, on average,
enceof baresoilcanalsobe incorporated.A bare siteincreases the rain that falls and infiltrateson a bare siteis only a fraction
accessof neighboringsites to moisture becausethere is no of the averagerain on the vegetatedsites.This is only a math-
competitionwith site vegetation.Thus, for a bare site, (4) ematicalway to incorporatethe fact that much lessrain infil-
trates becauseof the surfacesealing.Here we deal with ran-
becomes r/ - (si - Swn)e.Because thereis no vegetation
stressat bare soil sites,all comparisonsregardingglobalstress domly dispersedbare-soilsites,and thus they do not cluster
are normalizedby the actual number of vegetatedsites,and into regionslarge enoughfor appreciablesurfacerunoff to be
one then considersthe averagestressper plant overthe region. generated.
At first,it would seemthat it alwayswouldbe convenientfor
a plantto havea bare-soilneighborsincethe expectedvalueof 5. Impacts of Climate and Soil on Vegetation
Access to Moisture
r/wouldbe largerthan in (4). However,the availablesoilwater
at bare-soilsitesis, on average,much lessthan for the vege- Savannasrepresenta most important biome where water
tated sites,and so in most casesbare-soil neighborsare not availabilityis the key factorin establishing
the function,spatial
3714 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION,
pattern, and individualstructureof the vegetation[Scholes,
1997]. Savannasare not an ecologicmiddle ground between
forestsand grasslandsbut are rather a systemwith its own
characteristics,including a remarkably stable coexistenceof
trees and grasses.The explanationfor this existencehas been
the subjectof much researchthroughoutthe years.Presently,
the mostcommonexplanationfor suchcoexistence is basedon
the so-calledWalter hypothesisbasedon rooting-depthsepa-
ration with respect to competition for water by trees and
grasses[Walter,1971].Trees are assumedto haverootsin both
the surfaceand deeper soil layers,but grassesare assumedto
have roots only in the surfacelayer. Trees then have prefer-
ential accessto subsoilwater, and grassestend to have more
effective accessto topsoil water. The Walter hypothesis,in
effect,proposesthat two differentresourcesregulatetreesand
grassesin savannasso that the competitive-exclusion principle
canbe bypassed. This principle,whengeneralizedto n species,
statesthat they cannotcoexistwith fewer than n resourcesor
limiting factors [MacArthurand Levins, 1964]. In the caseof
treesand grassesin savannas,onlyone specieswill dominatein
the presenceof a singleresource.As pointed out by Lehman
and Tilman [1997],however,thisview of competitiondoesnot
include any considerationof spatial structuresof plant com-
munities, and yet the mere existenceof a spatial structure
could alter the outcomeof competition.
A number of modelswhich do not includespatialstructures
have applied the Walter hypothesisand have been used to
study the codominanceof trees and grassesunder different
conditions[Walkeret al., 1981;Eaglesonand Segarra,1985].
However,detailedfield observations do not supportthe Walter
hypothesisin many of the world's savannas.This does not
meanthat sucha hypothesis is not a realisticrepresentationfor
grasslandsand even for somesavannas,but it impliesthat it
cannotbe usedasa generalmechanismto explainthe structure
of savannas. It is instructive to examine this issue in some
detail.
The water-usestrategyof vegetationdependson the char-
acteristicsof the plantsaswell as on thoseof climateand soil.
In the grasslands of Arizona, rainfall doesnot infiltrate deeply
into the terrain, and high temperaturesin summercauserapid
evaporationof moisturefrom the upperlayersof soil[Salaand
Lavenroth,1982]. During the summerthe deep soil is seldom
wet. Winter moisture is deliveredgraduallyat cool tempera-
tures and thusis muchmore likely to penetratethe subsoil.As
discussed in detailbyBurgess [1995],the abovescenariois ideal
for a differential use of water between intensive and extensive
exploitertypesof plants.
Perennialgrassesin the desertgrasslands of Arizona exem-
plify the water use of intensiveexploiters.They have a dense
network of shallowroots so that they can intenselyuse any
moisture in the top layers of soil. Trees in this region are
extensiveexploiterswith roots that explore both the topsoil
and the subsoil.The usefulness of the deeprootsis conditioned
to the soil and the climate of the region. In contrastwith
intensiveexploiters,which effectivelyrespondto brief and in-
tensiverainfall eventsbefore the soil moistureevaporates,the
deep roots of extensiveexploitersbenefitmostlyfrom rainfall
eventsof longer durationthat occurin winter when the tem-
perature conditionsallow for infiltrationto deeper soil layers
where the moistureremainsavailablefor severalmonths[Bur-
gess,1995]. These kinds of conditionsare prevalent in many
parts of Arizona where one finds grasslandsthat maintain
nonnegligibleamountsof trees and shrubs.Spatial heteroge-
CLIMATE, AND SOIL MOISTURE
neity of soil propertiescan alsobe an important factor. Rain
infiltrates quickly and effectivelyinto sands,and in wet cli-
mates, sandsare often drier than claysbecausethey cannot
hold much moisture. Nevertheless, because of their structure,
sandscan supplywater to plantsmore effectivelywhen rainfall
is light or short. Water infiltrates more slowlythrough clay
soils,becauseof their very small pores,but clayscan store a
muchlarger volume of moisturethan sandysoils.In clay soils
if rainfall is short and intense, most of the water will become
runoff [Burgess,1995]. If a storm is long but with little rain,
onlythe top layersof the soilwill becomesaturatedbecausea
large volume is required to saturateclay. Long and soaking
stormsare particularlyeffectivein producinglargevolumesof
storedwater in clay soils.
In warm arid and semiaridregions,soilsfrom gravellyparent
materials commonlydevelop an argillic horizon rich in clay.
The formation of this argillichorizon as well as its depth and
thicknessare especiallyinfluentialon the capacityof the soilto
absorband storewater. Figure 5 taken from McAuliffe[1995]
showsan exampleof clay accumulationin a soil profile as a
functionof soil age in a semiaridenvironment.Holocene de-
posits(lessthan 11,000yearsold) generallyhave little or no
accumulationof clay in well-defined horizons. With time,
weatheringof parent rocksand the additionof materialsfrom
precipitationand dust fall causethe developmentof argillic
horizonswhichare detectedin Figure 5 for depositsof the late
Pleistocene.As discussed by McAuliffe[1995],the argillicho-
rizon plays an important role in semiarid environmentsby
controllingthe depth of water infiltrationand the durationof
the moisture availability to plants becauseof the different
water storagecharacteristics of clayand sandexplainedbefore.
The presenceof stronglydevelopedargillic horizons stops
deep infiltration and, in many areas of Arizona, leads to an
absenceof deep-rootedwoodyplants.The woodyplantsthat
are found in these soilsare relatively shallowrooted and thus
are capable of rapid responsesto precipitation [McAuliffe,
1995]. Deep-rooted woody plants are more common in the
youngerdepositswhere argillic horizonsare weak or absent
and in thoseareaswhere argillichorizonshavebeentruncated
by erosionthus allowingdeeperwater infiltration [McAuliffe,
1995].
Soil characteristics of Arizona grasslands thusare conducive
to dominanceby intensiveexploiters.In other regionswithout
argillichorizonsthe climatein winter allowsfor deep infiltra-
tion that is still availablelater for deep-rootedwoody plants
during the growing season.In these regions one could talk
abouttwo differentresourcesto supportwater competitionby
extensiveand intensiveexploiters.Nevertheless,this is a lim-
ited frameworksincein grasslands woodyplantsonly occurat
isolatedpointsinterruptingthe grass-dominated landscape.
Savannasare codominatedby trees and grassesover very
large and diverseclimatic regions.The variety of soils and
rainfallregimeson whichtheyare formedmakesit attractiveto
searchfor an explanationof their vegetationstructurethrough
a competitionmechanismmore general than the Walter hy-
pothesis.Field measurementscarried out in severaldifferent
African savannashave failed to validate the generalityof the
Walter hypothesis.Measurementsin dry savannasat Nylsvley
[Scholesand Walker,1993], as well as in more humid onesat
C6te d'Ivoire [Le Rouxand Bariac, 1998],have shownthat the
two-layerhypothesisof water partitioningis a misleadingsim-
plificationof water uptake patternsas a generalexplanation
for the codominanceof grassesand trees.
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3715

Figure6 (taken from Scholesand Walker[1993]) showsthe Root

length
density
(rn/n•)
depth distributionof fine rootsof woodyplantsand grassesin 0 1000 2000 3000 4000 5000 6000
the broad-leafedsavannasat Nylsvley.Trees and grassesex-
ploit essentiallythe same soil volume with grassroots domi-
nant to a depth of 1 m, whichis the averagesoil depthin the 0.2
region.Although trees may have somethick roots that pene-
trate great depths,for example,over 5 m for Prosopisglandu- •' 0.4
losain Texan savannas, they are of little advantagein compe- grass roots
tition for moistureif water is seldompresentin the deepersoil • 0.6 tree roots
layers.As discussed before,the depthto whichwater infiltrates
dependson climateandsoilcharacteristics. The soilat Nylsvley
ispredominantly
sandy.
Veryimportantly,
theclimate
atNyls- 0.8
vley hastwo seasons:a short,warm, dry seasonand a hot, wet
season
for theremainder
of theyear.Thisclimateis quite 1.0'
different from that of the desertgrasslandsin Arizona where Figure 6. Depth distributionof fine roots in woody plants
the winter is relativelywet and cool.The lack of precipitation and grassesin the broad-leafedsavannasat Nylsvley[from
duringwinter at Nylsvleyis a characteristicsharedwith many Scholesand Walker,1993].

other savannas.Where no appreciable amount of water is

% Clay storedin the deeperlayersof soilsduringwinter for possible
posterioruse by trees, whether becauseof climate character-
lO 2O 3O istics,soilproperties,or both,the Walter hypothesis
hardlycan
explainthe coexistenceof grassesand trees. Faced with this,
Scholesand Walker [1993] suggestthat the water-use niche
separationbetweengrasses andtreesin savannas maybe more
temporalthan spatial.Accordingto this hypothesis,treeshave
preferentialaccessto water in the early rains,and grassesare
20
more competitivelater in the rainy season.Scholesand Walker
[1993] base this hypothesison the observationthat woody
plantsin savannasachievefull leaf expansionbeforegrassesdo
A. and may evenprecedethe early rainswhen the previousyear
40 waswetter than normal. This argumentrejectsthe possibility
of an equilibriumsolutionor stablecoexistence
of treesand
grassesfor one of inherentstructuralinstabilitywhere contin-
uouschangesare expecteddependingon the specificfluctua-
60
tions occurringon the rainfall arrivalsand depths.
Without denyingthat differencesexistin temporalpatterns
of water useby trees and grassesaswell as in the soilvolume
theyexploit,the previousargumentagainseemsto lackgeneral
•' A. Mid-to-late
Holocene
deposit applicabilityfor the vastlydifferingconditionsin real savannas.
ø, (age1,000-4,000
years)
, B. Latest Pleistocene Moreover, the argumentlacksany spatialdynamicswhich can
' (age 8,000-15,000 years) be cruciallyilluminatingregardingthe coexistence of compet-
',C., D. LatePleistocene itors for basicallythe sameresource.
', (age25,000-75,000
years)
lOO
6. Optimal Coexistenceof Trees and Grasses
We now proceed to implement the spatially interactive
schemepresentedin section4. The first exampleis basedon
data from the Nylsvleyregion in South Africa, and it is an
12o expandedversionof the one presentedby Rodriguez-Iturbe et
!

al. [1999a].The 16 year systematicstudyof this regionmakes

i
i
it an importantbenchmarkin savannastudies.All the data for
i climate,soil,andvegetationusedin thissectionare takenfrom
14o Scholesand Walker[1993].
i

i
Practicallyall (98%) of the annualprecipitationat Nylsvley
i
i
occursfrom Septemberto April, with an averageof 600 mm
i
yr-•. Mostrainfallsin events
thatdeposit
lessthan20 mmof
rain, wetting only the upper 170 mm or so of the soil and
16o evaporatingin approximately5 days. Thus "the plants are
forcedto placetheir rootsvery closeto the surfaceif they are
Figure 5. Exampleof clayaccumulationin the soilprofile as to competeeffectivelywith the atmosphereand eachother for
a functionof soil age [from McAuliffe, 1995]. the fleetinglyavailablewater" [Scholes
and Walker,1993,p. 63].
3716 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE

Table 1. Permanent Wilting Points and Relative Soil are underwater stress(e.g.,transpirationis reduced)are esti-
Moisture Below Which TranspirationIs Reducedfor mated from Table 1 to be s*t = 0.12 and s *g = 0.17 for trees
NylsvleyVegetation and grasses,respectively.The maximum evapotranspiration
under nonrestricted water conditions is estimated as 4.5 mm
Relative Soil Moisture Permanent
Below Which Wilting d- • fortreesand5 mmd- • forgrasses
(R. Scholes,
Council
for
Percent of TranspirationIs Point, Scientificand IndustrialResearch,Pretoria,SouthAfrica, per-
Species Total Weight Reduced, % MPa sonalcommunication,1998). Maximum instantaneous values
may be much larger.
WoodyPlants
Burkea africana 0.69 0.105 -3.1 The most relevant propertiesof the mainly sandysoilsof
Ochnapulchra 0.17 0.143 -3.2 Nylsvleyare an averageporosityof 0.421, a depth uniformly
Terminalia 0.14 0.166 - 1.9 randomin the range0.80-1.20 m, and hydraulicconductivity
sericea
lognormallydistributedin spacewitha meanof 12.71x 10-6
Grasses m s-h, coefficient
of variationof 0.1,andisotropic
exponential
Eragostrispallens 0.77 0.152 -3.9 spatialcorrelation,
p(/) - e-(t/I) (I = 3 pixels).
Digitariaeriantha 0.23 0.223 - 2.9 Severalstress-function exponentsq, rangingbetween1 and
Data from Scholesand Walker[1993]. 3 were usedin (3) without significantchangesin the results.
Figure7 showsthe dependenceof the total (i.e., global)veg-
etation stressas function of percentageof tree canopycover
Rainfall is modeled here by a marked Poissonprocesswith for the Nylsvleycasewhen q = 1. Different grid sizeswere
arrivalrateof 0.167day-• andexponentially
distributed
marks, also used without change in outcome. The minimum-stress
for example,storm depths,with mean depth of 1.5 cm per situationcorresponds to a tree canopydensitynear 35% which
storm. The effect of different rainfall regimes was studied matchesquite well the valuesof 30-40% observedin the field
separatelyby changingthe aboveparameters.Interceptionof [Scholesand Walker, 1993]. Figure 7 also showsthe depen-
rainfall in the broad-leafedsavannasat Nylsvleycanbe treated denceof averagesoil moistureat the tree and grasssitesas a
as an approximatelyfixed removal of 1 mm per storm for functionof tree canopydensity.As expected,tree sitesshowa
grassesand 2 mm per storm for trees [Scholesand Walker, higher averagesoil water contentthan grasssites,and values
1993]. agreewell with thoseobservedin the region(as inferredfrom
The soil moisturefield is initialized as the seasonalaverage Figure 6.6 of Scholesand Walker[1993]). Notice in Figure 7
values of relative water content when no spatial interactions that whenthere are no spatialinteractions,the optimalcondi-
are consideredamong the different sites.This estimationre- tion is a field 100% coveredwith grass.Spatialdynamicslead
quiresthe useof the parametersinvolvedin Figure3 necessary to mixturesof trees and grassesthat reduceglobalvegetation
to describe the losses of moisture from individual trees and
grasses. We will useparametersthat representaveragesfor the
main speciesof plantsfound in the broad-leafedsavannasat
Nylsvley.Three typesof trees and two typesof grasseswere
selectedto representthe vegetationin the region. Some rele- 0.65
vant parametersof their water use are given in Table 1, and
althoughthere is considerablevariability amongtheir charac-
teristics,the mean parameterscharacterizingtrees are mainly 0.60
controlledby Burkea africana,and mean parameterscharac-
terizing for grassesare controlledby Eragostris pallens.The • with
spatial
intemction
J
weightsrepresentthe normalizedbiomassin the caseof trees
0.55
and normalized areal cover in the case of grassesand were
usedto estimatethe averageparametersfor woodyplantsand 0 0.2 0.4 0.6 0.8 I
herbaceousvegetation.
Canopy Density
The wiltingpointsreportedby Scholesand Walker[1993]for
the Nylsvleywild plants are considerablylower than the com-
0.10
mon value of -1.5 MPa for temperatecrops.Notice alsothat ............ average soil moistureon tree sites
the permanentwilting point is lower for grassesthan for trees 0.09
averagesoilmoisture
ongrasssites
even though grassesare frequently observedto wilt much
• 0.08
sooner.This highlightsthe higherwater use efficiencyof trees.
• 0.07
The averagewiltingpointsfor treesand grasses were estimated o

aSSw- 0.065 and sw-- 0.040, respectively,correspondingto • 0.06

o
pressurepotentials of -2.95 MPa and -3.70 MPa, respec- 03 0.05
tively, in a sandysoil. The conversionfrom pressurepotential
B
to volumetric soil moisture can be made either through stan-
0.03
dard soil moisture-pressurepotential curves [e.g., Larcher, 0 0.2 0.4 ''0:6.... 0:8....
1995] or throughthe water retention curvesfor the A horizon
Canopy Density
of the soil in the region,which are givenby Scholesand Walker
[1993]. The conversionfrom volumetricsoil water contentto Figure 7. (a) Averagewaterstressand (b) averagesoilmois-
relative soil moisture is accomplishedby dividingby the po- ture in tree and grasssitesafter spatialinteractionasa function
rosityn = 0.421 reported by Scholesand Walker[1993]. of tree canopycoverfor parameterscorresponding to the case
The valuesof the relative soil moisturebelowwhichplants of broad-leafedsavannasat Nylsvleyand q = 1.
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3717

stresses.The impact of climatic fluctuationswas studiedby

repeatingthe experimentsunder different rainfall conditions
through the adjustmentof the rate of occurrenceof storm
events. Thus it was found that a 20% wetter
globalminimumvegetationstresswith 100% trees.Similarly,a
20% drier climateyieldsa minimum-stressstateat 100% grass.
The evapotranspiration ratescorrespondingto conditionsof
minimum global water stressare shown in Figure 8a as a
functionof tree canopycover.One canobservethat the spatial
interactions which lead to a more efficient
moistureyieldhigherevapotranspiration ratesthanwhenthere
is no spatialinteraction.Figure 8b showsthe averageevapo-
transpirationfor tree sitesand grasssitesseparately.
The role of bare soil was studied under two conditions. First,
bare soil sites were assumedto have negligible infiltration
becauseof the surficialcrustthey develop.Second,the infil-
trationwasassumed to be reduced(butnot eliminated).Figure
9 showsa typical result of the experimentsfor Nylsvleyin
whichbare soil is explicitlyincluded.In this examplethe veg-
etated siteshave a fixed proportionof 40% trees and 60%
grasses.When bare crustpreventsinfiltration,we observethat,
as expected,the total stresson the vegetatedsitesincreases
monotonicallywith increasingbare soil. When bare soil sites
allowa reducedinfiltration(modeledthroughthe assumption
that, on average,they receiveone third of the rain receivedby
vegetatedsites),the globalstresson vegetatedsitesdecreases
with increasingbare soil. The secondsituationmay represent
what happensin areasthat havebeen plowed;specificresults
dependon the climate,soil,andvegetationparameters.Again,
no evolution in time is allowed for the bare soil, and, as men-
without
spatial
interaction
• 0.30
E
.o.
-,•
o 0.29
.--
0.60
climate leads to
without spatial interaction
with interaction (crusted bare soil)
0.40 with interaction (noncrustedbare soil)
0.20
use of the soil
0 0.2 0.4 0.6 0.8
Densityof Bare Ground
Figure 9. Average water stressin vegetated sites as a func-
tion of the percentageof bare soilfor a savannawhere 40% of
the vegetatedsoil is occupiedby trees.The parameterscorre-
spondto the caseof the broad-leafedsavannasat Nylsvley.
tionedbefore,the averagewater stressrefersto that presenton
the vegetatedsitesof the region.
The schemepresentedhere doesnot includetopographical
effects,and all surfacerunoff generatedvia bare soil effectsis
assumedto leave the area. A more complexscheme,wherein
runoff is routed via steeperdescentdirectionswith the possi-
bility of infiltratingother pixelsin the region,is being imple-
mented throughcellular automatarepresentation.
The effects of clusteringof trees on the total stresswere
investigatedby implementingthe same dynamicson a grid
where trees are spatiallyclusteredaccordingto a Poissonspa-
tial processwith clustercenterschosenat randomand the size
of the tree clusterbeing a random variable independentof
spatialposition.The dynamicsfollowsthen without modifica-
tions.Figure 10 showsthe globalvegetationstressasa function
of canopydensity.One noticesthat clusteringis not effectivein
reducingthe total vegetationstress,and furthermore its effect
is to increase the stress.In natural savannasthe degree of

c 0.28
0.345
o without
spatial
interaction
withspatialinteraction
(noclusters)
LU 0.27 withspatialinteraction
(clusters)
0.2 0.4 0.6 0.8
c 0.340
.o
CanopyDensity

>
0.50
m 0.335

>

• 0.40
o 0.30 0.330
'.•

o--
0.2 0.4 0.6 0.8 1
• 0.20
CanopyDensity
o
•. 0.10 Figure 10. Average water stressafter spatial competition
.......... average evapotranspiration
for trees
• when trees are distributed(a) at random or (b) in clusters.
averageevapotranspirationfor grasses
Cluster centersare generatedthrough a spatialPoissonpro-
0 .... 012 .... 014
.... 016 .... 018 cess;the tree patchesare rectangles,overlappingis allowed,
CanopyDensity with random sidescoming from an exponentialdistribution
with meanof sixpixels.The regionis 256 x 256 pixels,and the
Figure 8. (a) Average evapotranspiration and (b) evapo- followingparametersare adoptedq = 2, Sc - 0.23 (trees),
transpirationfor tree and grasssitesafter spatialinteractionas Sc = 0.26 (grasses),Sw = 0.065 (trees), and Sw = 0.04
a functionof tree canopycoverfor parameterscorresponding (grasses). The climateaswell asthe soildepthanditshydraulic
to the Nylsvleyand q = 1. conductivitycorrespondto thosefor the Nylsvley.
3718 RODRIGUEZ-ITURBE ET AL.' DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE

quite and otherwoodyplantsto increasein statureand density

4OO on grasslands."
Figure 11 showsthe fluctuationsin annual rainfall experi-
200 encedbytheregionin thelast55years.Oneobserves
largeand
sustaineddeviationsfrom the long-termmean rainfall over the
0 region. The period 1941-1960 was characterizedby a severe
drought,while the period 1961-1983waswet in termsof total
-200 amountaswell as in numberof yearswith rainfall higherthan
average.Archeret al. [1988] have documentedthe changesin
-400
the woodyplant coverin the regionduringtheseyears.We will
concentratehere on the total woody plant coveragewhich
1930 1940 1950 1960 1970 1980 increasedfrom 8% in 1960to 36% in 1983.In 1941the woody
plant coveragewas 13%. The structureof the tree clusters,for
Figure 11. Fluctuationsaround the mean annual rainfall at
Alice (Texas) (data from National Oceanicand Atmospheric example,size and distribution,also changed,but we will not
Administration). model this phenomenonin this paper.
The spatialinteractiondynamicsdescribedin section4 was
implementedfor this regionto studythe dependenceof total
clusteringof the woodyvegetationvarieswidely,and in some vegetationstresson the variable climatic conditionsand the
cases,evenlyspacedplantscan result from competitiveinter- possibleadaptationof the woody plant coveragetoward the
actions.In any case,clusteringis not due to competitionfor minimization of the stress. The climatic conditions assumed to
soil moisture(whichworksagainstit) but rather resultsfrom be responsiblefor the canopydensitiesexistingin 1941, 1960,
autogenicsite modification.Depositionof seeds,togetherwith and 1983were thosecorresponding to the averageof the grow-
local enrichment of soil nutrients and alteration of the micro- ing seasonsin the precedingdecades.Thus the mean rainfall
climate, is conduciveto clusteringof woodyvegetation. from May throughSeptemberwas395 mm in 1931-1941,339
mm in 1950-1960, and 412 mm in 1973-1983.The averagerain
on a rainydayis 15 mm (data from rain stationat Alice, Texas,
7. VegetationDynamicsand Climate Fluctuations on web site www'ncdc'nøaa'gøv/pub/data/cøøp-precip/texas'txt
The vegetationof savannaenvironmentsis particularlysen- The soilis fine sandyloamwith an averageporosityof 0.43anda
sitive to climatic fluctuations.Such fluctuationsmay be the randomdepth in the range of 0.70 to 1.10 m. The saturated
mostimportantnaturalfactorbehindobserved changes in the hydraulicconductivity is lognormallydistributed
with (Ks)= 82.2
relative densitiesof woodyplantsand herbaceousvegetation. cmd-• [U.S.Department ofAgriculture,
1979],coefficientofvari-
Savannaslocatedin the centralRio Grande plainsof southern ation0.1, and exponentially decayingspatialcorrelation,p(/) =
Texashave been carefullystudiedin this regard throughfield e-(l/O,withI = 3 pixels.
measurementsby the TexasAgricultureExperimentStationin The woodyvegetation(honeymesquite)and the C4 grasses
La CopitaResearchArea (27ø40'N,98ø12'W).The regionhas (e.g.,Paspalumsetaceum)are characterized by soilmatricpo-
an elevationrangingbetween75 and 90 m abovethe sealevel tentialsat the permanentwiltingpoint of -3.2 MPa and -4.5
with very mild slopesin a flat landscape[Scifresand Koerth, MPa, respectively[HaasandDodd, 1972;Ludlow,1976].Those
1987].The area is coveredby a biphasetree-grassvegetation soil water potentialscorrespondto relative soil moisturesof
with a populationof woodyplantsconsistingmostlyof honey 0.17 and 0.15, respectively,where the conversionis made
mesquites(Prosopis glandulosa)coexisting with C4grasses [Sci- throughthe retention curvesfor sandyloam [e.g., Waringand
fres and Koerth, 1987;Archer et al., 1988; Brown and Archer, Running,1998]. The maximumdaily transpirationrate mea-
1990]. The region has a subtropicalclimatewith hot summers suredfor mesquites is 3.42mm d-• [WanandSosebee, 1991;
and warm winters, an averageannual temperatureof 22.4øC, Cuomoet al., 1992]. For grassesit is estimatedto 10% more
and a mean annual rainfall near 700 mm. Rain is much more than that of mesquites.The evaporationfrom vegetatedsoil is
evenlydistributedthroughthe year in the Rio Grande plains taken as 1 mm d-•. The relative soil moisture values at which
than in the Nylsvleywith approximately400 mm during the transpirationstartsbeingreducedand the plant is under stress
growingseasonfrom May throughSeptember.In this the Rio were estimated as s* = 0.35 for trees and s* = 0.37 for
Grande plains are not typical savannassince most savannas grasses[Wan and Sosebee,1991; Cuomoet al., 1992].
have a concentratedwet season[Scholesand Walker,1993]. Minimizingthe globalvegetationstressasfunctionof woody
Much of the Rio Grande plainsis presentlydominatedby a plant densityresultsin stresses shownin Figure 12. In all cases
subtropicalthorn woodland vegetation complex. Although the exponentof the stressfunctionis q = 2; althoughfor all
commonlyclassifiedasAndropogon-Setaria-Prosopis-Acacia sa- the analysesin thispaper,the valuesof q = 1 andq = 2 yield
vanna[e.g.,Diamondet al., 1987],its structureseemsto belong qualitativelysimilarresults.One observesin Figure 12 that for
better to the tropical thicket, with dense,low-growingforma- 1941, 1960, and 1983 the minimum vegetationstressoccurs
tions of woodyplantslike thosewhere temperaturesare high when the densityof woodyplantsis near 15%, 0%, and 35%,
and rainfall is low. It is known that savannas can be induced to respectively.These valuesagree quite well with the observed
developa thicket structurethroughpoor management(e.g., onesfor thosedates(13%, 8%, and 36%, respectively). Figure
overgrazing[Archer,1994]). As pointed out by Archer et al. 13 showsthe averagesoil moisturein tree sitesand grasssites
[1988,p. 112],the Rio Grande plainsof southernTexasare an as function of tree canopydensityfor the growingseasonsof
interestingexampleof the grassland-to-shrubland conversion the decadespreceeding1960 and 1983.As expected,the aver-
processwhere "anthropogenic disturbances, reducedfire fre- age soil water content in the years leading to 1983 is consid-
quenciesand drought have presumablyinteracted, perhaps erably higher for both kinds of plants. The dependenceof
againsta backdropof gradualclimaticchange,enablingmes- evapotranspiration on canopydensityalsois quite differentfor
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3719

the decadespreceeding1960and 1983.As shownin Figure 14,

............ average soil moistureon tree sites A
in the wet caseof 1983,evapotranspiration increasesmono- 0.19
average soil moistureon grass sites
tonicallywith tree canopydensityreflectingthe greater avail-
ability of water for the woodyvegetationwhich, on average, 0.18
transpires
morethantheherbaceous vegetation. For 1960the .e
extremescarcity
of waterleadsto spatialinteractions
condu- .••) 0.17
civeto an averageevapotranspiration
whichdeclinesasa func- •
tion of increasingtree canopydensityfor low densitiesbut • 0.16
whichtendsto increasefor highvaluesof woodyplantscover-
age.It is importantto remarkthat Figures13 and 14 represent 0.15
soil moistureand evapotranspiration after a conditionof min- 1960
imum globalwater stresshasbeen reachedfor a givencanopy
0.146 .... 0'.2.... 0'.
4.... 0'.6
.... 0'.8
....
coverage.
Canopy Density

0.24 ...........................
0.55 .....................
............ averagesoilmoistureontreesites B
withoutspatialinteraction 0.23 average
soilmoisture
ongrass
sites
withspatial
interaction
0.22

0.21

0.53 0.20
0.19

0.52 0.18
1983
0.2 0.4 0.6 0.8 1
0.176 ..... 012
.... 014
.... 016
.... 018
....
Canopy Density
Canopy Density

Figure 13. Averagesoil moisturein tree and grasssitesafter

competitionas a functionof tree canopydensityat La Copita
0.86

0.84
without
spatial
interaction
.....•
with spatial interaction (Texas).The valuescorrespondto the conditionof minimum
global stresswith spatialinteractionsunder different canopy
coverages.Rainfall conditionsrefer to the monthsfrom May
0.82 through Septemberduring the decadespreceeding(a) 1960
and (b) 1983.
0.80

0.78 Theseresultsshowthe sensitivityof savannasand grasslands

0.76
(1950-1960)
B to climate fluctuationseither of natural or human origins. In
0 0.2 0.4 0.6 0.8 I arid and semiarid climates a cover of low grassesprevents
drop-splasherosionand reducesoverlandflow more than do
Canopy Density the canopiesof typicalwoodyvegetationin this region[McAu-
liffe, 1995].A changefrom perennialgrassesto a more patchy
0.490 . _ . withoutspatial interaction ....... •
coverof scrubsmostfrequentlycarriesprofoundconsequences
withspatial
interaction
.......... for the ecologicalequilibriumof a region.Lossof grassduring
droughtperiodsleadsto compactionof soilsand reducedin-
filtrationrates.Locally,augmentedrunoff increasesthe rate of
0.485
soilerosionand landscapeincisionon the randomfluctuations
of the topography[Schlesinger et al., 1990]. Thus there is an
increasedtransport of water, nitrogen, and other nutrients
0.480
acrossthe landscape.As pointed out by Schlesinger et al.
[1990],the net effectof thesefluxesis to reducethe availability
of soil moisture and nutrients and to increasethe heterogene-
0.475 .... 0'.2
.... 014
.... 016
.... 018
.... I ity of their spatialdistribution.Throughlandscapeincisionand
naturaltopographicfluctuations,water will accumulatelocally
Canopy Density becauseof the increaseof local runoff. This facilitatesdeeper
infiltration which favors the existence of scrubs. Thus the
Figure 12. Averagewater stressas a functionof tree canopy
coverusingthe parametersfor soilandvegetationat La Copita greaterspatialheterogeneityof water and nutrientsproduces
(Texas)and q -- 2. Rainfall conditionsrefer to the months islandsof fertility and patchydistributionsof shrubvegetation
from May throughSeptemberin Alice during(a) 1931-1941 like that presentlyseenin the La Copita area.
(averagerainfall395mm), (b) 1950-1960(averagerainfall339 The spatialheterogeneityof soil moisturewas studiedby
mm), and (c) 1973-1983(averagerainfall412 mm). evaluatingthe coefficientof variationof the relativesoilwater
3720 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE

0.231 namics,his hypothesisalsocan be implementedwhen vegeta-

, . . without
spatial
interaction
tion productivity(e.g., transpiration)is explicitlyoptimized

E
• . _ with
spatial
interactionthroughspatialinteractions.Usingthe sameschemeasbefore,
.•. 0.230
we alsotestedan implementationthat subtractsmoisturefrom
a siteby a randomlychosenneighborif it leadsto an increase
o
in global transpiration.The transpirationfunctionused was
similarto the onefor evapotranspiration (Figure3) but goesto
c 0.229
zero for valuesof soil moisture correspondingto the wilting
o point.Figure 15 showsthe averageglobaltranspirationandthe
relative soil moisturefor the Nylsvleyregionwhen the global
0.228 1960 A transpirationis explicitlymaximizedfor differentvaluesof tree
canopycoverage(e.g.,woodyplantsdensity).
0.2 0.4 0.6 0.8
With no spatial interactionsthe global transpiration de-
Canopy Density creasesmonotonicallywith increasingpercentageof woody
plants.The responseto maximizationof productivityis quite
different from the previoussimulations(Figure 7). Global

'•(•
__without
spatial interaction
_ withspatialinteraction •
• transpirationincreasesasthe plant coverageincreasesto about
0.284 40%; it then remainsat the maximumvalueof 0.25 cm d-•
until canopyvaluesof near 60% and then decreaseswith the
.o. 0.283 increase of woody plants. The canopy densities obtained
o
throughmaximizationof transpirationare only slightlylarger
• 0.282 •. than thoseobtainedas a resultof minimizingthe globalwater
stressof the vegetation.This qualitativelyagreeswith the hy-
• 0.281 •-
O •
> 0.280 •'

,.., 0.25
0.279
0.2 0.4 0.6 0.8 1

Canopy Density
o
Figure 14. Averageevapotranspiration after competitionas -.•
• 0.20
a functionof tree canopydensityat La Copita (Texas).The
valuescorrespondto the conditionof minimum global stress
under different canopycoverages.Rainfall conditionsrefer to
the monthsfrom May throughSeptemberin Alice duringthe
decadespreceeding(a) 1960and (b) 1983. ß 0.15
A

content for the fields resulting from the spatial interactive 0:2.... 014.... 0:6.... 018.... 1
dynamicsleadingto minimumtotal vegetationstress.The cor- CanopyDensity
respondingsoilmoisturefieldswere obtained,and their coef-
ficients of variation were estimatedfor the averagerainfall
conditionsin the growingseasonof the precedingdecades
endingin 1960 and 1983. It was found that the coefficientof
variation of the soil moisture field increased from 5% in 1960
to 9% in 1983showingagainincreasesin spatialheterogeneity 0.07
............ averagesoilmoistureon treesites
of soil water content resultingfrom climate fluctuationsand averagesoilmoistureon grasssites
overgrazing.

8. Maximum Productivity Versus Minimum

ß
'• 0.06
Vegetation Stress
Eagleson[1982,1994]hassuggested that plant communities
in semiaridclimatesarrangethemselves to operatesomewhere
betweenmaximumsecurityand maximumproductivity.Eagle- B
son [1994] definesmaximumsecurityas maximumsoil mois- 0.05•) .... 0:2.... 014
.... 0:6.... 018
....
ture and maximum productivityas equivalentto maximum
evapotranspiration.Thus he advancesthe hypothesis that the CanopyDensity
canopydensitywill adjustto a value in the rangewhere the
Figure 15. Averagetranspirationresultingfrom a spatialdy-
communityis betweenmaximumsoil moistureand maximum namicswhich maximizestranspirationunder prescribedtree
evapotranspiration.Notice that maximumsoil moisturedoes canopycoverages. The parametersare thosecorresponding to
not correspondto minimumwater stresswhichis the criterion the Nylsvleycaseandq = 1. (a) Averageglobaltranspiration
used in section 4. after competitionasa functionof canopydensity.(b) Average
AlthoughEaglesoh'sanalysesdo not incorporatespatialdy- soil moisturein the tree and grasssites.
 RODRIGUEZ-ITURBE ET AL.: DYNAMICS OF VEGETATION, CLIMATE, AND SOIL MOISTURE 3721

pothesisof Eagleson[1982], further studiedby Salvucciand Brown, J. R., and S. Archer, Water relationsof a perennialgrassand
Eagleson[1992],that the optimalcanopydensityis to be found seedlingversusadult woodyplantsin a subtropicalsavanna,Texas,
Oikos, 57, 366-374, 1990.
between maximum securityand maximumproductivity.The Brutsaert, W. H., Vertical flux of moisture and heat at a bare soil
averagevaluesof soil moisturefor tree and grasssitesagree surface,in Land-SmfaceProcesses in AtmosphereGeneralCirculation
well with thoseresultingfrom the minimizationof vegetation Models, edited by P.S. Eagleson,pp. 115-168, CambridgeUniv.
water stress,illustratedby Figure 7. Press, New York, 1982.
Burgess,T. L., Desertgrassland,
mixedshrubsavanna,shrubsteppe,or
semidesertscrub?The dilemma of co-existinggrowthforms.in The
9. Final Comments DesertGrassland,edited by M.P. McClanar and T. R. Van De-
vender,pp. 31-67, Univ. of Ariz. Press,Tucson,1995.
This paper studiesthe couplingbetweenclimate, soil, and Clapp, R. B., and G. M. Hornberger,Empirical equationsfor some
vegetationunder the dynamicsof spatialinteractionsbetween hydraulicproperties,WaterResour.Res.,•4(8), 601-604, 1978.
neighboringplants.Although the dynamicsis alwayslocal, a Cuomo, C. J., R. J. Ansley, P. W. Jacoby,and R. E. Sosebee,Honey
global objectiveis postulatedas controllingthe relative per- mesquitetranspirationalonga verticalsitegradient,J. RangeMan-
age.,45(4), 334-338, 1992.
centagesof woody and herbaceousplants that coexistin a Diamond, D. D., D. H. Riskand, and S. L. Orzell, A framework for
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1978.
drasticchangesin the tree canopycoveragesreflectinga shift
Eagleson,P.S., Ecologicaloptimality in water-limited natural soil-
in the optimal proportionsof woodyplants.These changes vegetationsystem,1, Theory and hypothesis,WaterResour.Res.,
matchquitewell thevaluesexpectedfrom a vegetationbalance 18(2), 325-340, 1982.
corresponding to minimumglobalwater stress. Eagleson,P.S., The evolutionof modernhydrology(from watershed
A similar spatial competitionfor soil moisturethat maxi- to continentin 30 years),Adv. WaterResour.,17, 3-18, 1994.
mizesglobalproductivity(e.g., evapotranspiration) alsoyields Eagleson,P.S., and R. I. Segarra,Water-limited equilibriumof sa-
vanna vegetationsystems,WaterResour.Res.,21, 1483-1493, 1985.
optimalbalancesbetweentrees and grassesin savannas.The Eissenstant,D. M., and K. C. J. Van Rees,The growthand functionof
relative amount of woody plants obtainedin this manner is pine roots,Ecol. Bull., 43, 76-91, 1994.
only slightlylarger than the one resultingfrom the minimiza- Gardner, W. R., and C. F. Ehlig, The influence of soil water on
tion of water stress. transpirationby plants,J. Geophys. Res.,68(20), 5719-5724, 1963.
Further researchis developingmore detailedevolutionary Haas,R. H., and J. D. Dodd, Water stresspatternsin honeymesquite,
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modelswhereinvegetationevolvesin structureand composi-
Lange,O. L., L. Kappen,andE. D. Shulze(Eds.), WaterandPlantLife,
tion as a functionof the climaticfluctuationsoperatingon the Springer-Verlag,New York, 1976.
region. Suchevolution,driven by the climate, is oriented in Lange, O. L., P.S. Nobel, C. B. Osmond,and H. Ziegler (Eds.),
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Larcher, W., Physiological Plant Ecology,Springer-Verlag,New York,
Acknowledgments.We thank SteveArcher (TexasA&M Univer- 1995.
sity) and Robert J. Scholes(CSIR, Pretoria)for usefuldiscussions. Lehman, L. E., and D. Tilman, Competitionin spatial habitats,in
This studywassupportedby grantsfrom NASA (grantsNAGW-4171 SpatialEcology,edited by D. Tilman and P. Kareiva, pp. 185-203,
and NAGW-4766) and NSF (grant EAR-9705861).We alsowant to Princeton Univ. Press, Princeton, N.J., 1997.
thank two reviewers and the Associate Editor for their valuable exten- Le Roux, X., and T. Bariac,Seasonalvariationsin soil,grassand shrub
sive comments and criticism. water statusin a West African humid savanna,Oecologia,113, 456-
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Ludlow, M. M., Ecophysiology of C4 grasses,in Waterand Plant Life,
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