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Observations and Modeling of Biomass and Soil Organic Matter Dynamics

for the Grassland Biome Worldwide

Article  in  Global Biogeochemical Cycles · December 1993

DOI: 10.1029/93GB02042

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 GLOBAL BIOGEOCHEMICAL CYCLES, VOL. 7, NO. 4, PAGES 785-809, DECEMBER 1993

OBSERVATIONS AND MODELING OF BIOMASS

AND SOIL ORGANIC MATrER DYNAMICS FOR TIlE
GRASSLAND BIOME WORLDWIDE

W. J. Parton,• J. M. O. Scurlock,'-
D. S. Ojima,• T. G.
Gilmanov, 3R. J. Scholes, nD. S. Schimel,sT. Kirchner,• J-C.
Menaut, • T. Seastedt,• E. GarciaMoya,s ApinanKamnalrut, 9
andJ. I. Kinyamario •ø

Abstract.Centuryis a modelof terrestrial biogeochemistry a diverseset of soils. Peak live biomassand plant
basedon relationships betweenclimate,humanmanagement productioncan be simulatedwithin+ 25% of the observed
(fire, grazing),soil properties,
plantproductivity,
and values(57 and 60% of the time, respectively)for burned,
decomposition. The grassland versionof theCenturymodel fertilized,andirrigatedgrassland siteswhereprecipitation
wastestedusingobserved datafrom 11 temperateand rangedfrom 22 to over 150 cm. Live biomasscanbe
tropicalgrasslands aroundthe world. The resultsshowthat generallypredictedto within + 50% of the observedvalues
soil C and N levels can be simulated to within + 25% of the (57% of the time). The modelunderestimated the live
observedvalues(100 and 75% of the time, respectively)for biomassin extremelyhighplantproductionyearsat two of
theRussiansites. A comparison of Centurymodelresults
with statisticalmodelsshowedthat the Centurymodelhad
slightlyhigherr• valuesthanthestatistical
models.Data
andcalibratedmodelresultsfrom this studyare usefulfor
•NamralResource
Ecology
Laboratory,
Colorado
State analysisanddescription of grassland
carbondynamics,and
University,Ft. Collins.
asa referencepointfor testingmorephysiologically based
ZDivision
of Life Sciences,
KingsCollegeLondon, modelsprediction's of netprimaryproductionandbiomass.
London.
Resultsindicatethatpredictionof plantand soil organic
3Departmere
of Vertebrate
Zoology
andEcology,
Moscow matter(C and N) dynamicsrequiresknowledgeof climate,
StateUniversity,Moscow.
soil texture,and N inputs.
4Forestek,
CSIR, Pretoria,SouthAfrica
5Climate
System
Modelling
Program,
NCAR,Boulder, INTRODUCTION AND OBJECTIVES
Colorado.
CLabomtoire
D'Ecologie,
EcoleNormalSuperieure, Underthe aegisof the ScientificCommitteeon Problems
CNRS-URA, Paris. of the Environment(SCOPE)Projecton "Effectof climate
?Institute
of ArcticandAlpineResearch,
University
of changeon production anddecomposition in coniferous
Colorado, Boulder. forestsandgrasslands,"theCenturymodelfor plant-soil
SCentro
de Botanica,
Colegiode Postgraduados,
Chapingo, ecosystems hasbeenfurtherdeveloped by a Grasslands
Mexico.
ModellingGroupin orderto applyit to a wide rangeof
•Facultyof NaturalResources,
Princeof SongIda temperate andtropicalgrasslands
worldwide.This study
University,Hatyai,Thailand. was undertaken to meet the overall aims of the SCOPE
•øDepartment
of Botany,University
of Nairobi,Nairobi, Project,to reviewandidentifymodelswith wideapplication
Kenya. andpredictiveability,in orderto link plantandsoil
responses to the large-scale
modelingof globalchange
Copyright1993 by theAmericanGeophysical
Union. effects. Climatechanges are expectedto be manifestas a
risein meanannualtemperature, togetherwithperturbation
Papernumber93GB02042. of rainfallpatterns,alongsidea continuingrapidincreasein
0886-6236/93/93GB-02042510.00 atmospheric
CO,_. Othermodificationsto the environment
786
may follow in the form of changesin nutrientavailability
and the rate of cyclingof carbonand nitrogenbetweenthe
biosphere,atmosphere, andgeosphere.
The carbonin terrestrialvegetationand soilsworldwide
outweighsthe amountfoundin the atmosphere and the
oceansurfacelayers. The role of grasslands in global
biogeochemical cyclesshouldnot be overlooked,especially
whenthe contributionof worldwidegrassland burningis
considered[Hao et al., 1990; Hall and Scurlock, 1991].
Tropicalgrasslands
occupy15 millionkm,.,andin termsof
bothlandareaandproductivityare nearlyequalto tropical
forests.Together
with9 millionkin,.of temperate
grasslands,
theycovernearlyonefifth of the Earth'sland
surfaceand are likely to remainconstantin areafor the near
future [Lieth, 1972; Hall and Scurlock, 1991]. Carbon
storedin grasslandsoils,temperateand tropical,hasbeen
estimated at 30% of the world total of soil carbon
lAnderson, 1991].
The role of ecosystem modelsto evaluatethe importance
of variousecosystems to globalenvironmental changeand
subsequent impactsis fundamentalto globalchange
research.The Centurymodelis beingappliedto global
changeimpactsand hasbeencriticallyevaluatedby the
SCOPEProjectGrasslands ModellingGroup. Centurywas
developedinto its presentform (Version3.0) [Partonet al.,
1992] by the incorporation of routinesderivedfrom more
mechanistic modelsor suggested by detailedsited_a_m. The
modelhasnow beenpammeterized and successfully
validatedfor manydifferentgrassland sitesin temperateand
tropicalregions[Partonet al., 1987, 1989a,b]. The
grassland sitesusedin this studyrangefrom temperate
grasslands in the UnitedStatesandRussiaandnaturaland
convertedgrasslands in wet anddry regionsof the tropics.
The ability of the modelto respondto burning,irrigation,
andN fertilizationwere testedusingobserveddatafrom
thesesites. We specificallytestedthe ability of the modelto
simulate:(1) seasonal patternsof live biomass,(2) peaklive
biomass, (3) annualplantproduction, and(4) soilC andN
levels. All of the sitesusedfor modeltestinghaveat least3
yearsof seasonal live biomassdata,observedsoil C andN
levelsandrequiredclimatedata. The statisticalcomparisons
usedto testthe modelincludelinearregression of observed
andsimulated results,comparison of Centurymodel
predictions with frequentlyusedregression models(i.e.,
production versusannualprecipitation), andan evaluationof
the numberof timesthat the ddferencebetweenthe Century
model and the observed data was less than + 25% of the
observed data.
MODEL DESCRIPTION
Centuryis a generalcomputermodelof plant-soil
ecosystems whichsimulates thedynamicsof grasslands,
forests,crops,andsavannas.It incorporates simplified
representationsof key processes relatingto carbon
assimilation andturnover,basedon existingmodels.
Centuryhasbeenpreviously described in detail[Partonet
al., 1987, 1988; Sanfordet al., 1991] and is designedto
work with the View outputmoduleof the program"Time-
zero:theintegrated modellingenvironment" [Kirchner1989].
Parton
eta•14.
ßModeling
Grassland
Biomes
Centurysimulates the dynamics
of carbon(C), nitrogen
(N), phosphorus (P), andsulphur(S) for differentplant-soil
systems,althoughonly the carbonandnitrogenoutputswill
be reportedhere. Differentplantproduction submodels for
grasslands,forestsandcropsare linkedto a commonsoil
organicmatter(SOM) submodel, with grasses andtreesboth
includedunderCentury'ssavannaoption. The SOM
submodelsimulatesthe flow of selectedelementsthrough
the differentinorganicandorganicpoolsin the soil,running
on a monthlytime step.
Version3.0 of Centurywas usedfor thesemodelruns
[Partonet al., t992]. This versionincludesa substantial
numberof changesnot described in the originalpublications.
Thesechangesinclude(1) addingan effectof clay on
formationof passiveSOM; (2) improvingthe surfacelitter
decomposition model;(3) includingthe effectof
anaerobiosis on decomposition; (4) simulatingleachingof
solubleorganicmaterial;(5) improvingthe inorganic
leachingequations;and (6) developinga robustgrassland
productionsubmodel.This paperwill describethese
significantchangesto the modelandpresenta general
description of the overallmodel.
DECOlVIPOSrrION AND SOIL ORGANIC MATTER
SUBMODEL
The soil organicmatter(SOM) submodelsimulates the
dynamicsof C, N, P, andS in theorganicandinorganic
partsof the soil system.The flow diagramfor soilC
(Figure1) showsthatsoilC is dividedup into threemajor
components whichincludeactive,slow,andpassivesoilC.
ActiveSOM includeslive soil microbesplusmicrobial
products (the totalactivepoolis approximately 2 to 3 times
the live soil microbialbiomass),the slowpoolincludes
resistantplantmaterial(for instance, ligninlikecomponents)
andsoil-stabilized plantandmicrobialmaterial,while the
passivematerialis veryresistant to decomposition and
includes physicallyandchemically stabilized SOM. The
flowsof C are controlledby the inherentmaximum
decomposition rateof thedifferentpoolsandthewaterand
temperature-controlled
decomposition
factor(Figure2).
Averagemonthlysoil temperature at the soil surfacecontrols
the temperature function,andtheratioof storedwater(0-30
cm depth)pluscurrentmonthlyprecipitation to potential
evapotranspirafion is the inputfor the moisturefunction.
Microbialrespirationoccursfor eachof the decomposition
flows. The partitioningof decomposition betweenstabilized
SOM andCO,_is a functionof soiltexturefor the
stabilizationof activeC into slowC (increasingCO,. for
sandysoilsand lesssoil C storage).Justification for these
assumptions are presented in the earlierCenturypaper
[Parton et al., 1987].
Plantresidues(shootsandroots)are partitionedinto
structural(resistantto decomposition)and metabolic(readily
decomposable) plantmaterialas a functionof the initial
residuelignin (L)-to-nitrogen(N) ratio (L:N) usingthe
followingequation:
Fm= 0.99- (•N) 0.018 (1)
Parton et al.' Modeling GrasslandBiomes 787

SURFACE LITTER ROOT LITTER

(1-Fm)
STRUCTURAL
• •k, Fm
=.85
-.018
*L/N STRUCTURAL
(1
-Fm
)••Fm
i METABOLIC I METABOLIC
IC C i ¸ C
LIGNIN I CELLULOSE LIGNIN I CELLULOSE

CO
•6
A
Lc
C• K2 CO2 Lc
• Lc .55

.3

.c I "c;
ø"" CsA=(1 - Csp-.55). SOIL
MICROBE
C
A CO2 , ,

CO2
.3 Tm=(1-.75' T)
CAS= (1 - CAp-CAL-Ft) K3
A

j cø2
SLOW Ft =,85-.68'T

C CAL=
(H2030/18)*(.01+ .04' Ts)
L/N = Ligninnitrogenratio
CO2
A = Abioticdecomposition factor .55
T = Siltplusclaycontent(fraction)
Ts= Sandcontent(fraction)
Tc= Claycontent(fraction) Csp=.003 ' 're
Ls = Faction
ofstructural
C thatislignin CAp=
.003+ .032' C
LEACHED
Lc = Exp( -3' Ls )
H203o=H20 leachedbelow30 m(cmm'1) PASSIVE C
KL= Maximumdecayrate(y-l)
C
Fm= Fractionof litterthatis metabolic
Ki = 3.9,4.8,7.3,6.0,14.8,18.5,.2,.0045y -1
:
Fig. 1. Flow diagramfor theCenturysoilorganicC model.

whereF,, is themetabolic fraction.The ligninfractionis dCt (3)

assumed to be partof the stmc• material,and(i.e., L:N
ratio)controlsthe decomposition rateof structuralmaterial.
The ligninfractionof theplantmaterialdoesnotcreate dCt (4)
microbialbiomass andis assumed to go directlyto theslow --r' c, t-4,
C poolas stmcua-al materialdecomposes. Surfacelitter
decomposition is treatedseparately,andsurfacelive
microbes are assumedto be in close associationwith the
surfacelitter. The surfacemicrobialpoolramoverrateis z'. = 33 (,5)
independent of soil texture,while soil textureinfluences
turnoverof activeSOM (higherratesfor sandy-soils). The
modelassumes a 60%lossof C dueto microbial respiration
for surface
microbes.Thesurface litterdecomposition Lc=e ( ) (6)
submodel hasbeenrecentlytestedusinglitterdecay
from an environmental
gradientin Hawaii [Vitouseket al.,
1993].
whereCi = the carbonin statevariable;I= 1,2,3,4,5,6,7,8for
Decomposition
of eachstatevariableis calculated
using surfaceand soil stntcmmlmaterial,activeSOM, surface
the followingequations: microbes,surfaceand soil metabolicmaterial,slow and
passive
SOMfractions;
I• is themaximum
decomposition
rate(year
'•) parameters
fortheithstatevariables
(Ki= 3.9,
dt --KIl•c A CI .!'-..1,2 (2) 4.9, 7.3,6.0, 14.8,18.5,.2, .0045year'•);A is thecombined
788 Parton et al.' Modeling GrasslandBiomes

i .00

0.75

0.50

0.25

0 10 20 30 40 50

SOILTEMPERATURE(øC)

1.0 -b /!

n' 0.8

• 0.6
0
• 0.4
o

LU 0.2

I I I I •/ I I I
0 0.20 0.40 0.60 0.80 '1.00 2.00 3.00

(STORED H20 + RAIN) / PET

Fig. 2. The abioficeffectof (a) soil temperature
and Co)soil moistureon decomposition.

abioticimpactof soil moistureandsoil temperature on

decomposition (productof the soil moistureandtemperature
0.85- .68 T (?)
terms- seeFigure 2), T, is the effect of soil textureon
activeSOM turnover,T is the silt plusclay content
(fraction);andLc is the impactof lignincontentof structural
= (o.m + 0.o4 r) (8)
material(Ls) on structuraldecomposition.
The modelassumes that all C decomposition flowsare (9)
associated with microbialactivityandthatmicrobial
Co = 0.003+0.032T,,
respirationoccursfor eachof theseflows. The fractionof C
lostdue to microbialrespirationwith eachC flow is shown
in Figure1 nextto the CO2arrows. Carbonleavingthe (10)
active SOM box is divided into four different flows which
includemicrobialrespiration, leachingof solubleorganicC,
and stabilizationof C in the slow andpassivepools whereFt is the fractionof C lost due to microbial
(equations (7)-(10)), while the C flowsout of slowSOM is respiration,L, is thefractionlostdueto organicleaching, CA
allocatedto passiveSOM and activeSOM (equations(11) is the fractionallocatedto passiveSOM, Cs is the fraction
and (12)). sentto the slowpool,H,_O•o is the monthlywaterleached
Partonet al.: ModelingGrassland
Biomes
belowthe30 cm soildepth(cmmonth'x),
S is thesand
content(fraction),T• is the clay content(fraction). Q, is the
fractionof slow C allocatedto passivepool,and Cs is the
fractionof slowallocatedto the activepool (55% of the C is
lostdue to microbialrespiration).
(11)
(12)
c a -- (t - -0.55)
The formationof passiveSOM is a functionof clay
content(higherfor clay soils)and is primarilycontrolledby
the stabilizationof activeSOM in microaggregates. Some
passiveSOM is alsocremodby the decomposition of slow
SOM, and includesa similareffectof clay. The effectof
clayon theseflowsis basedon soilx4Cdata[Becker-
Heidman1985]whichshowthatthepercent of modemx4C
decreaseswith clay content,suggesting
thatclay soilshavea
higherfractionof passiveSOM. This is alsoconsistentwith
the way thatJenkinson[1990] includedthe impactof clay
on SOM dynamics.
Leaching
of soluble
organic
matter
outofthetop30-cm
of the soil is calculatedasa functionof thedecayratefor
activeSOM (D,), the sandcontentof the soil,and the water
flow belowthe 30-cmsoil depth. The parametervaluesfor
formationof passiveSOM weredetermined by fittingthe
model to observed soil C data at the U.S. Great Plains sites
(Konzaand CentralPlainsExperimental Range(CPER))and
while the organicleachingparmmeters werederivedby
fitting the modelto observedwaterchemistrydatafrom a
watershedin PuertoRico (W. H. McDowel and C. E.
Asbury,Export of carbon,nitrogenand majorions from
threetropicalmontanewatersheds, submiuedto Journalof
Limnologyand Oceanography, 1993) (hereinafterreferredto
as McDowel and Asbury,submittedmanuscript,1993). A
morecompletedescription andjustificationfor the C flows
is presented by Partonet al. [1993]. ' when saturatedwater flows betweensoil layers. The
BIOPHYSICAL SUBMODELS
The Centurymodelincludesa simplifiedwaterbudget
modelwhichcalculatesmonthlyevaporation and
transpiration
waterloss,watercontentof the soil layers,
snow water content and saturated flow of water between soil
layers(AppendixB). If the averageair temperature is less
than0øC monthlyprecipitation occursas snow. Sublimation
andevaporation of waterfrom the snowpackoccursat a
rateequalto thepotentialevapotranspirafion rate. Snow
melt occursif theaverageair temperature is greaterthan
0øC andis a linearfunctionof the averageair temperature.
The potentialevapotranspirafionrate (PET) is calculatedas a
functionof the averagemonthlymaximumand minimumair
temperature usingthe equationsdevelopedby Linacre
[1977]. Bare soil waterlossis a functionof standingdead
andlitter biomass(lowerfor highbiomasslevels),rainfall,
789
andPET. Interceptionwater lossis a functionof
aboveground biomass(increaseswith biomasslevel),
rainfall,and PET. Transpirationwaterlossis functionof the
live leaf biomass(exponentialfunctionof leaf biomass),
rainfall, and PET. Interceptionof water and bare soil water
lossesare calculatedas fractionsof the monthlyprecipitation
and are subtractedfrom the total monthlyprecipitation,with
the remainder of the water added to the soil.
Water is distributedto the differentlayersby addingthe
water to the top layer (0-15 cm) and then drainingexcess
water (waterabovefield capacity)to the next layer.
Transpirationwater lossoccursafter the water was addedto
the soil. Water lossoccursfu-stas interception, followedby
baresoil evaporation and wanspirafion(the sumdoesnot
exceedthe PET rate). The field capacityand wiltingpoint
for the differentsoil layersis calculatedas a functionof the
bulk density,soil texture,and organicmattercontentusing
an equationdevelopedby Guptaand Larson[1979]. The
numberof soil layersis an input variablein the model,and
we used15-cmincrements for eachlayer up to the 60-cm
soil depthand 30-cm incrementsbelow the 60-cm depth.
Water leachedbelow the last soil layer is not availablefor
evapotranspirafion and is a measureof interflow,runoff,or
leachinglossesfrom the soilprofile. Watergoingbelowthe
proffiecanbe lost as streamflow or leachedinto the subsoil
where it can accumulate or move into the stream flow.
AppendixB givesall the equationsusedin the waterflow
modeland detailedvalidationand descriptions are presented
by W. J. Partonet al. (manuscriptin preparation,1993).
Averagemonthlysoil temperature nearthe soil surfaceis
calculatedusingequationsdevelopedby Parton[1984].
Theseequations calculatenearsurfacemaximumsoil
temperature as a functionof the maximumair temperature (2
m heigh0and the canopybiomass(lowerfor highbiomass)
while the minimumsoil temperature is a functionof the
minimumair temperature (2 m heigh0and canopybiomass
(higherfor higherbiomass).The actualsoil temperature
usedfor decomposition andplantgrowthrate functionsis
the averageof the minimumand maximumsoil
temperatures.
Leachingof labilemineralN (NO3'andNH[) occurs
fractionof mineralN thatflows from the upperlayer to the
lowerlayer(F=;0-1) is calculated asa functionof the sand
contentof the soil (Ts;0-1), andthe amountof saturated
waterflow betweenlayersusingthe followingequation:
(13)
(0.2+0.7
whereH20•0is thesaunated
waterflow(cmH20 permonth)
from layeri to layeri+ 1. Parametervaluesin 13 were
determined by fittingthe modelto observed waterchemistry
datafrom PuertoRico 0VIcDoweland Asbury,submitted
manuscript,1993).
NITROGEN SUBMoDEL
The N submodelhasthe samegeneralstructureas the soil
C model(Figure3). The organic-Nflowsfollow the C
790 Parton
etal.' Modeling
Grassland
Biomes

x
z

'•z
z o.
z

uJ z
z

0
Partoneta!.' Modeling GrasslandBiomes 791

flows and are equalto the productof the carbonflow and passivepoolsgenerallyresultsin net mineralization
of N,
the N:C ratio of the state variable that receives the C. The while decomposition of strucawalmaterialimmobilizesN.
C:N ratiosof the soil statevariablesreceivingthe flow of C The modelalsousessimpleequations to represent
N inputs
area functionof the mineralN pool(NO3'plusNI-!•+) and dueto atmospheric depositionandN fixationandcalculates
vary within the ranges3-15, 12-20,and7-10, respectively, N lossesdueto N2, NO, N20, andNH3gasfluxes(see
for active,slow and passiveSOM (Figure4). The C:N ratio Figure3) andNO• leaching.A morecomplete description
of newlyformedsurfacemicrobialbiomassis a linear andjustificationfor the N submodelis presented
by Parton
functionof the N contentof the materialbeingdecomposed, et al. œ1987,1988].
and increases from l0 to 20 as the N content decrea•s from
2.0% to 0.01% (,seeFigure3). The C:N ratio of slow SOM PLANT PRODUCTION SUBMODEL
materialformedfrom surfacemicrobesis equalto the C:N
ratioof the microbesplus5.0. N associmed with carbonlost Centurysimuhtesplantproduction for grasslands,
in respirationis assumedto be minemlized.Given the C:N agricultural
crops,forests,andmixedtree-grass(savanna)
ratioof the statevariablesandthe microbialrespirationloss, systems.The grassland submodel (Figure5) simulates
grass
decomposition of metabolicresidue,active,slow,and growthandincludesthe impactof grazingandfire on plant

a 20

!5

o SLOWSOM (Ns)
1-
<•
n- lO
o

:•'•,"•.•.
PASSIVE
SOM
(Np)
••OM (N)
0 , I • I ., I [ I ,
0 o.• •.o •.• a.o a.•
b 2o
SOIL NO• + NH4(gm -•)

15

SURFACE

MICROBE(N SM)
n- lO
O
o

0 , I , I , I • I ,
0 0.5 1.0 1.5 2.0 2.5

PLANT N CONTENT (%)

Fig.4. Variation
in C:Nratios
of (a)theactive,
slowandpassiveSOMpoolsasa function
of themineralN pool,
and(b) thenewlyformedmicrobialbiomassasa function
of theN content
of decomposing
plantmatter.
792 Parton et al.: Modeling GrasslandBiomes

GRASS MODEL

LIVE SHOOT PPT STANDING DEAD

C C

PPT

PLANT

PRODUCTION SURFACE

LI-I-I'ER

LIVE ROOT ROOT

C LI-I-I'ER

PPT = MonthlyPrecipitation
TEM = MonthlySoilTemperature
BIO = Plant Biomass
GRZ = Grazing Rate
FIRE = Fire Frequency

Fig. 5. Flow diagramfor the brassland

production
submodel.

production.Potentialplantproductionis calculatedas reducesplantproduction in the presence

of largeamountsof
functionof soil temperature,
availablewater,and a self- standingde.•dmatter(Figure0c).
shadingfactorusingthe followingequation: Thepotential
plantproduction
rate(P•)is reduced
if there
is insufficientnutrientsupplyof N, P, or S, with the most
(14) limitingnutrientconstraining production.A maximumand
minimumcarbonto element(E) ratio is specifiedfor roots
and live shoots,and the nutrientconcentrationnot allowed to
whereP],is theaboveground
potential
plantproduction
rate be greaterthanthe minimumC:E ratio for eachnutrient.
(g m'2month'l),
P==is themaximumpotential
aboveground The live shootC:E ratio is a linearfunctionof the live plant
plant
production
rate(250g m'2month'l),
Tp(Figure
6b)is biomass.The fractionof the labile nutrientpoolsthatare
theeffectof soiltemperature
ongrowth,• is theeffectof available
forplantgrowth(Fs)area functionof theliveroot
moisture
onproduction (Figure
6a),andSpis theeffectof biomass(ROOT;g m'2)according to thefollowingequation:
plantshadingon plantgrowth(Figure6c). The effectof
moisture
onproduction
Mvis a function
of theratioof
currentmonthlyprecipitation plusthe previousmonth's (15)
storedsoil water(0-60 cm) to the potential = - 0.s (-o.0t
evapotranspiration
rate. The soilwaterholdingcapacityalso
influences
Mvby modifying
theamount
of stored
soilwater,
so thatlower waterholdingsoils(e.g.,sandysoils)havea Thisequationis basedon datapresented
by Wedinand
highergrowthrate underdry conditions (seeFigure6a). Tilman [1990]. Live root and shootdeathare calculatedas
This functionwasaddedto the modelin orderto represent a functionof the availablesoil waterin the 0-60 cm layer
theobservation thatsandysoilsare moreproductive in dry (H,_O(,•)
usingthefollowingequations:
environments [Salaet al., 1988]. The temperatureresponse
curvesfor plant growthare basedon data from Christieand
Detling[1982]for C• andCsplants.The shading factor R,•= 0.12 [exp(-5.0 H20(a))] (16)
Parton
etal.' Modeling
Grassland
Biomes 793

"• 1.00 - a Grazingremovesvegetation,returnsnutrientsto the soil (by

urinationand defecation),alters the root/shootratio and
• 0.75 increasesthe N content of live shootsand roots [Holland et

uJ 0.50 Sand al., 1992].

Note thatCenturyassumes that incomingsolarradiation
doesnot contributesignificantlyto interannualvariability
.,-, 0.25 .,,,,,,••
• / Clay and so can be neglectedas a controlon the time-scalesof
o i interesthere. This assumption is bornout by the generally
:• 0 1.00
0.25 0.50 0.75 high correlationsbetweenobservedand simulatedNPP.
(Rainfall+ Stored H20) / PET Increasingly,climaticdata_ andecologicaldata_ [Knappand
Smith, 1990] suggestthis may be a poorassumption for the
1.2
future,giventrendsin cloudiness suggested as partof global
changescenarios.Futureversionsof the modelwill needto
includethis interaction,as well as CO: interactions.

DESCRIffHON OF SITES

0.6
The grassland studysitesusedfor the modelingwork (see
0.4 Figure7) reportedherecovermostof the world's major
grassland types,rangingfrom the continentalplainsof the
0.2 United Statesand Russia(22 to 81 cm per year
precipitation)to bothnaturaland convertedgrasslands in wet
0 and dry regionsof the tropics(59 to 154 cm per year
O0 04 08 12 16 20 24 28 32 36 40 44
precipitation).Soil tyw• covera wide rangeof textures:20-
Soil T (øC) 85% sand,10-40% silt, and 2-70% clay. Six of the
importantbiogeographic regionsidentifiedby the
InternationalGeosphere-Biosphere Programme(IGBP) for
1.OFC • •/.•--- globalchangeresearchare included:temperate northern
hemisphere, centralarid Asia, Caribbean,northernAfric•
(includingWest Africa), southernand easternAfrica, and the
tropicalAsianMonsoonRegion[Eddyet al., 1991]. Site
descriptionsare summarized in Table 1. The diversityof
• 0.5 soils,climaticconditions andplantgrowthpatternsof these
sitesprovidea robusttestof Centuryfor globalgrasslands.
While the sitespermita generalglobalintercomparison,each
sitehasmadea uniquecontribution to the analysispresented
here.
0 I I I
0 1 2 3 The largegrasslandregionof theU.S. CentralPlains
Live/Dead represents
a naturalwest-to-east
moisturegradient.The
Fig. 6. The effectof (a) moisture,(b) soil temperature,
and CentralPlainsExperimentalRange(CPER)at thearidend
(c) shadingon potentialgrassgrowth. of thegradient
is a shortgrass
steppeexperiencingwide
diurnal,seasonal,and annualvariationin temperature
and
precipitation.
Vegetation
thereis dominated
by C4
shortgrasses.
The KonzaPrairieResearch
NaturalAreaat
= o.2o [xp //,O(D] (z?) the moremesicendof the moisturegradient,is characterized
by warm-season (Cn)tallgrass
prairiespecies.Dataavailable
where1• andSaare the fractionof rootsandshoots, at CPERincludea 5-yeartimeseries(1971-1975)of
respectively,
whichdie per month. Shootdeathalsooccurs aboveground live biomass andplantproduction,for control,
whenliveplantbiomass
exceeds
400g m'2at a rateof 10% irrigated,
fertilized,andbothirrigatedandfertilizedplots.
permonth.Thisdeathrateis designedto simulate
deathdue Konzadataincludeestimatesof aboveground
plant
to shading
of lowerleavesin thecanopy. production
(1976-1990)andseasonality
of aboveground
live
The model calculatesroot/shootratios as a function of the biomass(1984-1990)for annuallyburnedandunburned
annualrainfallusingequations presented in the original plots. Noneof thedatafromKonzawereusedto develop
descriptionof Century[Partonet al., 1987]. The effectof the model. Soil data from the CPER was used in model
fire andgrazingon plantproduction arebasedon datafrom development;
CPERlive biomass
datawasnotusedin
Hollandet al. [1992] andOjima et al. [1990],respectively. modeldevelopment.
The impactsof fire are to increasetheroot/shoot ratio,and The five sitesin the Commonwealth
of Independent
States
to increasethe C:N ratio of live shoots(plus 10 for (CIS; formerlyRussia)characterize
the"continentality"
temperate sites;0 for tropicalsites)androots(plus30 for all climategradientfromtheEuropean temperate
meadows of
sites),removingvegetation andreturninginorganic nutrients. KurskandOtradnoyein westernRussia,throughthe typical
794 Partonet al.' ModelingGrassland
Biomes

SCOPE GLOBAL GRASSLAND STUDY

SITE NAME

1 NAIROBI NAT. PARK

KENYA

2 KLONG HK HAT YAI

THAILAND

3 MONTECILLOS CHAPINGO
MEXICO

4 LAMTO
IVORY COAST

5 CPER PAWNEE
COLORADO U.S.A

6 KONZA PRAIRIE
KANSAS U.S.A

7 KHOMUTOV
UKRAINE

8 KURSK
RUSSIA

90TRADNOYE 1
RUSSIA

10 SHORTANDY
RUSSIA

11 TUVA
KAZAKHSTAN

Fig. 7. Locationof the sitesusedto testthe Centurymodel.

steppesof Khomutovin Ukraine,to the ultracontinental a longperiodon an ancientsoil. By contrast,

steppes of Tuva, at the geographicalcenterof the Asian
continent.The longtime seriesof aboveground biomass
data(1954-1983)for the Kursksitecoversa wide rangeof
year-to-yearvariationin weatherconditions.At Otradnoyc,
5 yearsof monthlydata(1967-1972)are availableon
aboveground biomassfor a loamyanda sandysoft. The
Khomutovsite has4 yearsof aboveground biomassdata
(1967-1970). More detaileddataon vegetation dynamics is
availablefor the Shortandy site,with monthlydeterminations
of live and deadmatter,bothaboveground andbelowground,
overthe period1975-1979. The cold,dry Tuva site
(aboveground biomass,1977-1984)is a particularly
interestingtestfor the Centurymodel,sinceits climaticand
physiological characteristics
differ markedlyfrom the
grasslands of theU.S. CentralPlainswhichwereused
originallyto formulateCentury[Partonet al., 1987].
In view of the importanceof tropicalgrass-containing
ecosystems or grasslandsand savannas as perhapsthe largest
terresUialbiome [Hall and Scurlock, 1991], it was an
importantobjectiveof thepresentSCOPEProjectto adapt
Centuryfor modelingof tropicalgrasslands.Three of the
four tropicalgrassland
sites(Kenya,Thailand,andMexico)
were studied under a United Nations Environment
ProgrammeProject,and haveuseda commonmethodology
to collectdata on live and deadmatter,aboveground and
belowground, overtheperiod1984-1990[Longet al., 1992].
The fourthtropicalsitewaslocatedat Lamto,Ivory Coast.
The Kenyangrassland is a dry grassland,
established
over
theThailand
sitewasprobablyconver• fromforestwithinthelast50-
100 years,characteristic
of manygrasslands in southeast
Asia. The Mexicangrassland is derivedfromthebedof a
salinelake, drained in 1911. The continuousrecordof
aboveground andbelowground damfromthissiteis oneof
the mostdetailedfor any grassland worldwide.The humid
grasssavanna siteat Lamto,Ivory Coast,hasbeen
intensivelystudiedby J. C. Menautandcoworkers for the
past30 years,anda discontinuous recordkeptof monthly
aboveground live anddeadmatter,andtotalrootmatter
[Menautand Cesar,1979]. Like mosttropicalgrasslands, all
four sitesweresubjected to burningduringtheperiodof
study. Century'sabilityto simulatethe effectsof burning
hasthusbeentestedundertreatmentsrangingfrom annual
fires to burningevery 6 years.
ESTIMATION OF MODEL PARAMETERS
The robustness of the Centurymodelat a varietyof
grasslandsites,from tropicalsavannasto temperatesteppes
andwet meadowgrasslands is testedusinga singlemodel
with minimalchangesto parameters.Most parameters used
by Centuryare intendedto remainconstantin themajority
of applications
and are referredto as the fixed parameters.
Otherparameters whichare particularto an individualsiteor
groupof sitesare containedwithin a site-specific file. With
the comparisons betweendifferentgrassland sitesworldwide,
it wasdesirablefor the majorityof the modelparameters to
Partonet al.' Modeling GrasslandBiomes 795
o -i- , o o

o -i- o o o

o -i- , o o

o -i- o o o

o -i- o o o

o o o o , -i-

o o o o

, o o o , -i-

o o o o ,

, , , + •

,
Partonet al.' ModelingGrassland
Biomes
comprise a universal"worldgrassland" set,with a relatively
smallnumberof site-specific parameterschanging according
to dataavailablefrom eachstudysite.
Someof thesesite-specific parametersconsistof the
"minimuminput"d•;• requiredto runtheCenturymodel,
whichincludelatitudeandlongitude,soiltexture(sand,silt
andclay),soilpH, soildepthfor modelingwaterbudget,
planttype(Ca,C4),andgrowingseason, andweatherdata
(monthlyprecipitation, monthlymaximum,andmonthly
minimumtemperatures). The remainingsite-specific
parameters relatemainlyto plantphysiological functions
controllinggrowth,death,turnover,andN inputs.These
differenceswerekept to a minimum(Table2 andFigureA1
listsvaluesfor all sitespecificparameters).A sitefile was
firstcreatedby enteringbasicsiteidentification and
"minimuminput"d_a_;_a into the "universal"
grassland file.
Phenological differences (lengthof growingseason) and
plantgrowthresponses to temperaturewerealsoentered at
thisstage.The Centurymodelwasthenrun for a periodof
5000years,usingrepeated weathersequences basedon long-
termweatherdatafrom the sites. Appropriate patternsof
grasslandmanagement (grazingandburning)were
incorporated in theselong-termruns,basedon theknown
historyof the sites(seeTable 1).
The equilibriumlevelsof soil organicmatter(SOM) of
the long-termrunswere usedas initial conditionsfor the
validationstudy. The validationrunsweresimulated using
actualweatherdatarecordedat the site,anda comparison of
simulatedandobservedvaluesof monthlybiomassanddead
matter,aboveground andbelowground wasmade. For these
short-termmodelruns,the management regimewasbased
on recentgrazing/burning practiceat eachsite(seeAppendix
A for details).
Differencesin physiological
parameters betweenthe 11
grassland sitesare summarized in Table 2. At the Kenyan,
Lamto,and Thailandsites,the relativelylow belowground
live biomasscomparedwith aboveground live biomass
requireda changein the equations determining root/shoot
ratio. Root deathrate, determinedby availablesoil water,
was satisfactorily
modeledby a commonfunctionat all the
sitesexceptfor CPER,wherea lowerratewasrequiredto
achieve the observed balance between live and cl•_d root
biomass.Atmospheric deposition andN fixationinputsare
normallydetermined as a simplelinearfunctionof
precipitation
basedon datafrom the U.S. GreatPlains,but
increasedinputswere requiredto predictthe observed
production at the (relativelydry) Confederationof
Independent States(CIS) sites. This is consistentwith the
higherratesof N deposition in continental
Europecompared
with the U.S. Central Plains [B6nis et al., 1980]. Reduced
N inputswerenecessaryat the Thailandsitein orderto limit
productionunderconditionsof highrainfall. At Konza,
Lamto,andKenya,the functiondetermining C:N ratio was
modified to allow wider ratios based on detailed information
availableon the N contentof plant matter. This
modificationwas importantbecauseall thesesiteshave
frequentburningwhichwouldotherwiseresultin too much
lossof nitrogen.
The deathrate of abovegroundbiomassis determinedin
Centuryby the numberof soillayerscontributing waterto
797
plantgrowth(andhowquicklytheydry), as well as by a
directsoil waterfunction. In orderto producesufficiently
rapidturnover of aboveground matter,thenumberof soil
layerswasreduced at theKenya,Thailand,Otradnoye, and
Tuva sites(this hadthe effectof possiblyreflecting
differencesin soil structure).The deathrate functionwas
furtherincreased at the KenyaandThailandsites. The
transferof standing deadto litterwashighestfor continental
siteswith snowfall(0.20 per month)andmuchlowerfor the
tropicalsites(0.10permonth).Mostof thesesitespecific
differencesreflectspecies specificandsoildifferences at the
site. Documentation of the specificvaluesof thesesitesare
presented
in AppendixA anda copyof sitespecific
parameters
areavailablefromtheauthors.
MODEL TESTING
In this sectionwe showa comparison of observeddata
and simulatedmodeloutputvariables. The modeloutput
variablesthatwe will testincludethe following:(1)
dynamicsof live shootbiomass,(2) aboveground
plant
production,
(3) peakstandinglive shootbiomass,and (4)
soil C and N levels. These variables were selected because
theycharacterize grasslandsystemsandinteractwith GCM
models. Soil C level and plantproductionare importantfor
theglobalC cycle,live plantbiomasshasa big impacton
the energyandwaterbudgets,andpeaklive biomassis
frequentlyusedas an estimateof aboveground plant
biomass.Live plantbiomassinfluencesthe energybudget
by alteringthe canopyalbedoandsoil temperature latent
heat fluxes.
We employedthree•e•chniques for testingthe Century
model: (1) linearregressionof observeddataversus
simulatedmodelresults,(2) comparison of the Century
modelpredictions with empiricalregression modelspredicted
from climatevariables[Lauenroth,1978],and (3) calculating
the numberof timesthe differencebetweenCenturymodel
predictionandobserveddatadifferedby lessthana
thresholdproportion(25%) of theobserved data. The results
showthatin general
theobserved
versus
simulated
1• forthe
Centurymodelandthe empiricalregression modelsare
fairly similar. The advantage of theCenturymodelis thatit
canpredicttotalsystembehavior(e.g.,nutrientcycling,N
gasfluxes,waterfluxes,leachinginorganicandorganic
compounds) andresponses to manipulations
suchas
fertilization,irrigation,andlandusechanges, while
regression modelscanonlypredictwhatwill happenfor the
particularobserved datasetusedto generatetheregression
modelcoefficients.As a minimumcriterion,in general,we
felt Centuryshouldpredictobserveddataat leastas well as
empiricalclimateregression models,whichhavelimited
extrapolation potentialwithoutincludingnutrientinteractions
[Esser, 1986].
ABOVEGROUND NET PRIMARY PRODUCTION
Aboveground plantproduction datawereavailablefrom
theCPER shortgrass prairiesiteandthe Konzatallgrasssite.
The CPER sitehasplantproduction from 1971to 1975for a
control,fertilized,irrigated,andirrigatedandfertilizedsites
798 Partonet al.' Modeling GrasslandBiomes

[Lauenroth,1978]. The Konzasitehasplantproduction

data An analysisof the residualerrorshowedthatall of the
from 1976-1990 for annual burned and unburned sites largeunderestimates of peaklive biomassby Centurycame
[Abramset al., 1986,Bfiggset al., 1989] (alsosee from the Khomutovand Kursksites. For comparison,
unpublishedLong-TermEcological Research records).The removingthe extremeKhomutovand Kurskpointsfrom the
comparison of observed andsimulated production data analysis
increased
ther• from0.45to 0.65. Fifty-seven
(Figure8) showsthatthemodelsimulates thesedifferent percentof the Centurypredictions haderrorsthatwere less
treatmentswithan r• of 0.70. A linearregressionwherethe than+ 25% of the observeddata. In general,the model
independent
factorswere annualprecipitation,fertilization, simulatedpeaklive biomassreasonably well for all of the
andburning
hasanr• -- 0.67. WhileCenturymaynot sites,with theexceptionof peaklive biomass for the
greatlyimproveon predictingobservations undercurrent observedhighplantproduction years(9 yearsout of 32
conditions,compared to simpleregression models,its years)at theKurskandKhomutovsites. The reasonfor the
concurrentability to predictthe seasonalityof foliage Russiansiteerrorsis unclear. We suspectthatchangesin
productionhassubstantial significance
to vegetation- species composition occurred duringthe highproduction
atmosphere interactionsstudiesand modelsof ecosystem years. It is alsoimportantto notethatthesetwo sites
dynamics.Sixty percentof Century'spredictedplant recordedsomeof the highestpeaklive biomassobservedat
productionvalueshaveerrorslessthan+25% of the any sitein spiteof the fact thatprecipitation at the Russian
observedplantproduction.The standarderrorsof the sitesis 1/2 to 1/3 of the valuesfor the tropicalsites.
observed plantproduction acrossfieldreplicatesgenerally
rangedfrom 10 to 20% of the observedvalues.
LIVE PLANT BIOMASS DYNAMICS
PEAK LIVE BIOMASS
The comparison
of observedandsimulatedlive biomass
The simulatedpeaklive biomassfromall of thedifferent for someof thetemperate (Figure10) andtropical(Figure
sitesand treatmentswere comparedto the observedpeak 11) sitesshowsthatgeneralseasonal patternswerewell
live biomass(Figure9). The observedpeaklive biomassis simulated by the model. Differencebetweenverydry and
the maximumlive biomassobserved in a particularcalendar wet years(1989 versus1990at Konzaand 1981 versus1980
year(January to December).Centuryresults
haveanr• = at Tuva) were well simulated,however,moresubtle
0.45. Ther• for thebestregression
modelwas0.40,with differencesbetweenyearswith similarprecipitation(Konza
annualprecipitation
as theindependent
variable. Peaklive 1984-1987and Kursk 1975-1979)were not well simulated.
biomassis frequentlyusedto estimateannualplant For thetropicalsites(Figure10) theresponse
to fire (1986
productionin ungrazedgrasslands,with annualproduction and 1989 at Thailand,1986 at Lamto)was fairly well
beingapproximately 50% greaterthanthepeaklive biomass simulated. The death of live biomass at Mexico was
[Lauenrothand Sala, 1992]. underestimated
duringthe dry season.

Observed vs Simulated Production

Burned, Unburned, & CPER
1,000

8OO

600

4oo ...............................................
j,-..,--
e.•......•................................................
ß.........................

2OO

..................•o
e/•eeeee .....
i
' .....
' ''.....ß...................................................................................
I
y=88 +.67x,
I
r2=.70 I
0 200 400 600 800 1,000
Simulated
Fig. 8. Comparison
of observed
andsimulated
aboveground
plantproduction
for all theCPERtreatment
andburned
and unburned sites at Konza.
Partonet al.' ModelingGrasslandBiomes 799

Peak Live Biomass - All Sites

Observed vs Simulated
1,000

8OO

6OO

ß ß
ß
ß ß
4OO oA ........... •A

ß
ß
ß ß
ß e• oo e•
ß Russian Sites
2OO
ß ß ß
ß
ß ß y = 78 + .80x,r2=.45
y-R=
48+ .83X,r2=.65
o I I I I
0 100 200 300 400 500 600 700

Simulated
Fig. 9. Comparison
of observedandsimulatedpeaklive biomassfrom all of the sites. The Kurskand Khomutov
site data are shown with a.

CPER Live Biomass b Konza Live Biomass

1973 - 1975 1984-1990
lOO 600

• 80
-• .
• 400 ...........
(• 60

g 40 •D..300
• ß i,,•, ..................
ß
•o ..................................

200 ..........
• 20 • 100 .........
o 0
1973 1974 1975 1984 1985 1986 1987 1988 1989 1990
Year -- Simulated
Year --- Simulated
ß Observed ß Observed

Kursk Live Biomass d Tuva Live Biomass

1975 - 1979 1978 - 1982
500 3oo

400 ................................... ' ............. -"'.................... • 250 .....................................................................

ß :•2oo
300 e
ß ' ß •'•$o
200
100 • so

0 o
1975 1976 1977 1978 1979 1978 1979 1980 1981 1982
Time Year
-- Simulated ---- Simulated
ß Observed ß Observed

Fig. 10. Comparison

of simulated
andobserved
live biomass
for (a) CPER,Co)Konza,(c) Kursk,and(d) Tuvasites.
800 Partonet al.' ModelingGrassland
Biomes

Kenya Live Biomass Lamto Live Biomass

1984 - 1986 1981 - 1986
400 600

350
500 ............. e..........................................................
ß,.., 300 ........................... ß ............ ß ß................... •

•D 250
ß . •)
• 400

• 200 • 300

u• 150
E 200

50
('• 100
ß
0 0
1984 1985 1986 1981 1982 1983 1984 1985 1986
Year Year
• Simulated • Simulated
ß Observed ß Observed

c Mexico Live Biomass d Thai Live Biomass

1984 - 1990 1984 - 1990
500 700

ß 600

,,- 400 .... ß................................................................. •

'•
:• ßß • 500
•d• ß ß ee '' *
03
• 200
03 300
• ße--e.--•. .... e.-e
..................... e.e.•.... d...........
E E
• • 200
•100 ................... •
lOO

o 1984 1985
• 1986 1987
• 1988
• 1989
• 199o
o 1984 1985 1986 1987 1988 1989 1990
Year Year
• Simulated • Simulated
ß Obse•ed ß Observed

Fig. 11. Comparison

of simulated
andobserved
livebiomass
for (a) Kenya,(b) Lamto,(c) Mexico,and(d) Thailand
sites.

Live Biomass
Observsd vs Simulated

All Sites - Except Khom and Kursk

600
ß
ß

5OO ...............

ß ß ß ß
ß

4O0
. * •"0 .d** * ß oo
ß
* *
ß ß ß
(D 300 ß .,.*..o,..............*.............,.......................
***%*
*. *'ß
o
200
$o •'
ß1"eeee
..........
' .......
; .............
R:•'•ifi'•i•6'd"::'0'.'5•
...............
,....... y = 0.76x + 40.73
100
•''t'O"
'O..........................................................................................

o
o lOO 200 300 400 500 600 700

Simulated
Fig.12.Comparison
ofsimulated
andobserved
livebiomass
forallofthesites
andtreatments
except
Khomutov
and
Kursk.
Parton et al.' Modeling GrasslandBiomes 801

A comparison
of observed
versus
simulated
liveplant showthatthe modelsimulatespeaklive biomassand
biomass for aH of the sites and different treatmentsshowed abovegroundplantproductionbetterthanlive biomass.
r• = 0.39. Again,themodeltended
to underestimate
live
biomassfor highproduction
yearsat the Kurskand SOIL C AND N
Khomutovsites. Removingthe datafrom the Kurskand
Khomutov
sitesincreased
therato 0.56(Figure12), SoilC andN levelsarefairlywell simulated(ra = 0.93
suggesting
thattheerrorsfor theRussianhighplant and0.89, respectively
for C andN) acrossa rangeof soilC
production
yearsgreatlyreduce
ther•. Thirty-one
percent
of levelsrangingfromlessthan2000g C m'2at Lamtoto
the Centurypredictionshad errorslessthan+ 25% of the greaterthanthe10,000g C m'2at theKursksite(Figure
observed
data,while57% of Centurypredictions
haderrors 13). The assumptions usedto controlsoilC stabilityand
less than + 50% of the observed data. These results decomposition(clay impacton passiveSOM and silt plus

a
12,000
Soil C (0-20 cm)
10,000

8,000

6,000

4,000
.•'
2,000 y=.93x-7.84
I I ! I
r2=,93!
2,000 4,000 6,000 8,000 10,000 12,000
Simulated(gm-2)

Soil N (0-20 cm)

1,000

800

E
6OO

4OO

• y=.73x-78.84
200 r• 88

I I I I
00 200 400 600 800 1,000

Simulated (gm-2)
Fig. 13. Comparison
of simulated
andobservedsteadystatesoil(a) C andCo)N for 9 of the 11 sites. Mexicoand
Thailandsiteswerenotusedbecausetheyarenotat equilibrium
withtheirpresentlanduses.
802
clay on slow SOM) seemto work acrossa diversesetof soil
texturesand soil mineralogies.One hundredpercentfor C
and75% for N of the modelpredictions had errorslessthan
:t: 25% of the observed values. Soil C data from the CPER
and Konza site were used to fit some of the model
parameters in the SOM model. Burkeet al. [1989]have
developeda regression modelfor predictingsoil C and N
levelsin grasslands as a functionof climateandsoil texture.
We usedthe Burkeet al. [1989] modelto predictsoil C and
N levelsat the differentsites(excludingThailandand
Mexico),andr2between
observed
andsimulated
soilC and
N levelswere0.89 and0.90, respectively.
AppendixA showsthe equilibriumsoilC level for the
differentsoil pools(presentedas initial valuesfor model
runs). The datashowthat the fractionof soil C in active
SOM generallyrangedfrom 2% to 4%, and 35-60% for
passiveSOM. Siteswith highclay (>35%) contenthadhigh
amountsof passiveSOM andlessslowSOM. For example,
the sandysoilsat CPER had61% slowSOM and 36%
passive,while the clay loam soilsat Konzahas41% slow
and55% passiveSOM.
DISCUSSION
We thinkthatdifficultyin simulatinglive plantbiomassis
primarily
a resultof thefacttliatplantspecies
changes
are
not considered in the model. Changesin species
composition from Ca andC4 vegetationor structural changes
resultingfrom shiftsbetweengrasslands and savannas affect
nutrientdynamics,waterutilization,biomassallocation,and
othercharacteristics whichmodifyplantproductionand
seasonality of plantgrowth. Numerousgrassland studies
haveshownthatshiftsin CaandC4grassdominance canbe
inducedby drought[AlbertsonandWeaver,1944],grazing
andfire regime[Owensbyet al., 1970;TowneandOwensby,
1984;Braggand Hulbert1976], or nitrogenadditions
[Owensbyet al., 1970;Dodd andLauenroth,1978;Wedin
and Tilman 1990; Seastedtet al., 1991]. Modificationsof
resourceuseefficiencyamongvariousvegetation
communitiesare importantto projectinghow an ecosystem
will respondto increased atmospheric
CO,.,changein
climate,or increases in atmospheric
depositionof N [W•
andTilman, 1990]. Studiesof N additionsto grassland
ecosystems indicatethatgrasslandcommunities are sensitive
to the soil nitrogenstatus[DoddandLauenroth,1978;
Wedin and Tilman, 1990; Seastedtet al., 1991]. These
shiftsin plantcommunities havepotentiallymajorimpacts
on soilorganicmatterdynamicsvia controlsof nitrogen
immobilization and storage,however,our abilityto predict
changesin plantspeciescomposition is limited.
Additionalfactorscontributing to thesesubtleinterannual
differencesmay restfitfrom (1) interannualvariationsin
incidentPAR, not includedin the model,(2) effectsof
timingof rainfallnot capturedwith a monthlytime step,or
(3) lag effectsof nutrientor photosynthate storagein plants.
Invest:.::;•,:z•&•n
of the aboveeffectsis ongoing.
M• :•Je•s
'•.'keCenturywhichsimulateNPP form an
;m•no:;am complement to morephysiologically oriented
Partonet al.' ModelingGrassland
Biomes
modelswhichsimtfiatephotosynthesis andrespiration.
Physiologicalmodelsrequiredetailedclimaticinformation,
suchashourlyincomingsolarradiationrelativehumidity
andwind speeds,that are not widelyavailablefor
retrospectiveanalysessuchas arepresented here.
Physiological modelsmay alsobe sensitiveto species-
dependent traits,thoughthissensitivitymaybe tractablefor
large-scalecalculations[Schimelet al., 1990; McGuire et al.,
1992]. The extensivebodyof NPP dataover time,
reflectingbothgeographic variabilityand the effectsof
climatevariabilityallowsthe development of powerful
calibratedmodelslike Century. Suchdataand modelsare a
crucialbenchmarktestfor predictedtime integralquantifies
(e.g., NPP, biomasslevels)from modelsbasedon
calculationof instantaneousexchanges of CO,..
SUMMARY
The comparison
of observedand simulatedlive peak
biomass
andplantproduction
hadr2 valuesof 0.45and0.70,
respectively;
Centuryerrortermsfor thesevariableswere
generallylessthan:t:25%of the observed
data. Seasonal
livebiomass
wasnotaswellrepresented
by themodel(xa --
0.39) and had errorslessthan:t:50% of the observedd_a_m_:
The modelsimulated
differences
bet/veenwet anddry years
well, but was unable to simulatemore subtledifferences
betweenyearswith similarprecipitation.The model
substantiallyunderestimatedlive biomassfor unusuallyhigh
production yearsat the KurskandKhomutovsites.
A comparison of the Centurymodelpredictionsfor plant
production andpeaklive biomasswith empiricalregression
equationsfit to observeddatashowedthat the Century
modelwas slightlymoresuccessful thanthe regression
equations.SteadystatesoilC andN levelswerewell
simulated
by themodel(r2 = 0.93and0.89)fora setof sites
with differingclimateand soil textures.The modelwasable
to predictsoil C andN levelsto within:t:25% of observed
data.Globally,
keycontrols
overC storage,
NPP,and
biomassincludeclimate,N inputsandlosses,andsoil
particlesizedistribution.This model,rigorouslycalibrated
againsta largebodyof observations, is usefulbothas a
descriptiveandanalyticaltool,andasa baselinetestfor
time-integratedpredictions(e.g.,of NPP or biomass)for
physiologicallybasedmodels,explicitlysimulating CO,.
exchange[e.g.,Coughenour, 1984;Hunt et al., 1991].
APPENDIX A
Site-specificparameters foundin the SITENAME.DAT
file may be dividedinto a numberof categories (seeFigure
A1): (1) Basicsiteidentification(e.g., SITLAT, SIllgqG);
(2) "minimuminput"datarequiredto mn the Centurymodel
((a) environmentalinformationsuchas PRECIP, TMN2M,
TMX2M; and Co)physicalfeaturessuchas sand,silt, clay);
(c) parameters relatingmostlyto plantand soilphysiology.
FigureA1 showsthe parameters from categories 2b and 3
for all the grasslandsitesreportedhere,with noteson the
differences between sites.
Partonet al.' Modeling
Grassland
Biomes 803

All equationsusedin the waterflow modeland variable

definitions. For detailedvalidationand descriptions,
see
Partonet al. (manuscript in preparation,1993).SeeFigureB1.
ADD wateraddedto thesoil(cmmo4).
& depthof ith layer (cm).
ATM atmosphericH•.Osink or source(cm).
AWL i relativeroot densityin the i layer (0-1).
BS fractionof PPT lost as bare soil H20 loss (0-1).
D deepH20 storage(cm).
E• fractionof H20 lossby transpiration(0-1).
EVAP bare soil plus interceptionH20 loss(cm).
EVBS baresoilevapotranspiration
(cmmo4).
FAv total H,.O availablefor transpiration
H20 loss(cm).
saturatedH,.Oflow from i layer to the i+ 1 layer
(cm mo4).
fractionof PPT lost as interceptionH20 loss(0-1).
Kd fractionof excessH,.O flowing to Ds (0-1).
fractionof excessH,.O flowing to Sv (0-1).
fractionof D flowing to Sv
liquid snowpack (cm).
aboveground
liveplantbiomass
(gm").
La surface
litterbiomass
(gm'2).
M theH20 meltingfromS (cmmo4).
N the numberof soil layer unitswith roots
PET potentialevapotranspiration
H,.Oloss(cm).
PPT monthlyprecipitation
(cmmo4).
RWCi the relativewater contentfor the ith layer
s frozensnowpack (cm).
sa standing
deadplusliveplantbiomass
(gm'2).
sD saturated
drainage
to deepstorage
(cmmo4).
sv streamflow (cmmo4).
si soil water in the ith layer (cm).
s? field capacityof i layer (cm).
Siw wilting point of i layer (cm).
Ss H•.Oflow fromD to Sv (cmmo4).
S• sublimation
H•.Oloss(cmm4).
T transpiration
H20 loss(cmm4).
T, transpiration
H20 lossith layer(cm m4).
% potentialtranspiration
H,.Oloss(cmmo4).
Wa weightfactorfor transpiration
H20 lossfrom the ith
layer (0-1).
804 Parton et al.' Modeling GrasslandBiomes
Partonet al.' Modeling
Grassland
Biomes 805
806 Partonet al.. ModelingGrassland
Biomes
Partonet al.' Modeling GrasslandBiomes 807

n Z
• Z *• •w- ^ v
o•. A A
, 0•1--*•

,,&

o • •
o u_•
II f.,0
- A
808 Partonet al.: ModelingGrassland
Biomes

Acknowledgements.
Measurement of monthlybiomass Christie,E. K., andJ. K. Detling,Analysisof interference
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outunderUnitedNationsEnvironment Programme (UNEP) Coughenour, M. B., A mechanistic simulationanalysisof
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theU.S. NASA EarthObserving SystemprojectNACW- Res. 15, 1633-1635, 1979.
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Datato ModelFactorsLimiting Hall, D. O., andJ. M. O. Scurlock,
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(ReceivedJanuary29, 1993;
revised June 21, 1993;
acceptedJuly 22, 1993.)