20
Global Change Biology (1996) 2, 5-23 The importance of climate and soils for estimates of net primary production: a sensitivity analysis with the terrestrial ecosystem model YUDE PAN, A. DAVID MCGUIRE, DAVID W. KICKLIGHTER andJERRY M. MELILLO The Ecosystem Center. Marine Biological Laboratory. Woods Hole. MA 02543. USA Abstract We used the Terrestrial Ecosystem Model (TEM) to investigate how alternative input data sets of climate (temperature/precipitation), solar radiation, and soil texture affect estimates of net primary productivity (NPP) for the conterminous United States. At the continental resolution, the climates of Cramer and Leemans (C&L) and of the Vegetation/ Ecosystem Modelling and Analysis Project (VEMAP) represent cooler and drier condi- tions for the United States in comparison to the Legates and Willmott (L&W) climate, and cause 5.2% and 2.3% lower estimates of NPP. Solar radiation derived from C&L and given in VEMAP is 32% and 60% higher than the solar radiation data derived from Hahn cloudiness. These differences cause = 8% and 10% lower NPP because of radiation- induced water stress. In comparison to the FAO/CSRC soil texture, which represents most biomes with loam soils, the soil textures are finer (more silt and clay) in the Zobler and VEMAP data sets. The use of VEMAP soil textures instead of FAO/CSRC soil textures causes = 3% higher NPP because enhanced volumetric soil moisture causes higher rates of nitrogen cycling, but use of the Zobler soil textures has little effect. In general, NPP estimates of TEM are more sensitive to alternative data sets at the biome and grid cell resolutions than at the continental resolution. At all spatial resolutions, the sensitivity of NPP estimates represents the impact of uncertainty among the alternative data sets we used in this study. The reduction of uncertainty in input data sets is required to improve the spatial resolution of NPP estimates by process-based ecosystem models, and is especially important for improving assessments of the regional impacts of global change. Keyivords: climate, geographically referenced data, net primary productivity, soil texture, solar radiation, terrestrial ecosystem model (TEM) Received 6 June 1995; decision to author 24 July 1995; accepted 16 November 1995 Introduction The atmospheric concentrations of the major long-lived greenhouse gases continue to increase as a direct result of human activity. Equilibrium simulations of general circulation models (GCMs) for a doubled CO2 atmosphere project that the global mean surface temperature will rise between 1.5 "C and 4.5 °C. and that precipitation and cloud patterns will also be altered (Houghton et fl/.1995). Climate changes of this magnitude are expected to affect terrestrial ecosystems both functionally and structurally. Interest in the effects of global climate change has motivated the development of ecological analysis at large spatial scales. Process-based ecosystem models Correspondence: Dr Yude Pan, fax +1-508-457-1548, e-mail [email protected] have become a common means to evaluate the potential consequences of climate change on terrestrial ecosystems. In general, process-based ecosystem models have similar requirements for input data. Most of the models need geographically referenced data on temperature, precipitation, solar radiation, topography, soil character- istics, and vegetation as inputs. The availability of these data has been limited in the past, and the data require- ments of ecosystem models have simultaneously stimu- lated the development of data sets organized within geographical information systems. Today there are several alternative data sets available and different modelling groups often use different data sets to run their models. However, it is not clear what differences exist among alternative data sets, and it is not known whether differ- © 1996 Blackwell Science Ltd.

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Page 1: The importance of climate and soils for estimates of net

Global Change Biology (1996) 2, 5-23

The importance of climate and soils for estimates of netprimary production: a sensitivity analysis with theterrestrial ecosystem model

YUDE PAN, A. DAVID MCGUIRE, DAVID W. KICKLIGHTER andJERRY M. MELILLOThe Ecosystem Center. Marine Biological Laboratory. Woods Hole. MA 02543. USA

Abstract

We used the Terrestrial Ecosystem Model (TEM) to investigate how alternative inputdata sets of climate (temperature/precipitation), solar radiation, and soil texture affectestimates of net primary productivity (NPP) for the conterminous United States. At thecontinental resolution, the climates of Cramer and Leemans (C&L) and of the Vegetation/Ecosystem Modelling and Analysis Project (VEMAP) represent cooler and drier condi-tions for the United States in comparison to the Legates and Willmott (L&W) climate,and cause 5.2% and 2.3% lower estimates of NPP. Solar radiation derived from C&L andgiven in VEMAP is 32% and 60% higher than the solar radiation data derived fromHahn cloudiness. These differences cause = 8% and 10% lower NPP because of radiation-induced water stress. In comparison to the FAO/CSRC soil texture, which representsmost biomes with loam soils, the soil textures are finer (more silt and clay) in the Zoblerand VEMAP data sets. The use of VEMAP soil textures instead of FAO/CSRC soiltextures causes = 3% higher NPP because enhanced volumetric soil moisture causeshigher rates of nitrogen cycling, but use of the Zobler soil textures has little effect. Ingeneral, NPP estimates of TEM are more sensitive to alternative data sets at the biomeand grid cell resolutions than at the continental resolution. At all spatial resolutions,the sensitivity of NPP estimates represents the impact of uncertainty among thealternative data sets we used in this study. The reduction of uncertainty in input datasets is required to improve the spatial resolution of NPP estimates by process-basedecosystem models, and is especially important for improving assessments of the regionalimpacts of global change.

Keyivords: climate, geographically referenced data, net primary productivity, soil texture, solarradiation, terrestrial ecosystem model (TEM)

Received 6 June 1995; decision to author 24 July 1995; accepted 16 November 1995

IntroductionThe atmospheric concentrations of the major long-livedgreenhouse gases continue to increase as a direct resultof human activity. Equilibrium simulations of generalcirculation models (GCMs) for a doubled CO2 atmosphereproject that the global mean surface temperature will risebetween 1.5 "C and 4.5 °C. and that precipitation andcloud patterns will also be altered (Houghton et fl/.1995).Climate changes of this magnitude are expected to affectterrestrial ecosystems both functionally and structurally.Interest in the effects of global climate change hasmotivated the development of ecological analysis atlarge spatial scales. Process-based ecosystem models

Correspondence: Dr Yude Pan, fax +1-508-457-1548, [email protected]

have become a common means to evaluate the potentialconsequences of climate change on terrestrial ecosystems.

In general, process-based ecosystem models havesimilar requirements for input data. Most of the modelsneed geographically referenced data on temperature,precipitation, solar radiation, topography, soil character-istics, and vegetation as inputs. The availability of thesedata has been limited in the past, and the data require-ments of ecosystem models have simultaneously stimu-lated the development of data sets organized withingeographical information systems. Today there are severalalternative data sets available and different modellinggroups often use different data sets to run their models.However, it is not clear what differences exist amongalternative data sets, and it is not known whether differ-

© 1996 Blackwell Science Ltd.

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6 Y. PAN etal.

ences among data sets affect results of process-basedecosystem models.

It is important to understand how differences amongalternative data sets affect the results of ecosystem modelsfor two reasons. First, it is relevant for reducing uncer-tainty of ecosystem model results for contemporaryclimate. This is especially relevant for model comparisonactivities (see VEMAP Members 1995). Second, whenecosystem models are used in a prognostic mode, climatechanges inferred from general circulation models (GCMs)are applied to contemporary climate to generate futureclimatic scenarios (Adams et al. 1990; Melillo et at. 1993,1995; McGuire et al. 1993,1996a, 1996b; Dai & Fung 1994;Schimel et al. 1994; VEMAP Members 1995). Uncertaintyin contemporary climate influences future climatescenarios and the results of ecosystem models.

We used the Terrestrial Ecosystem Model (TEM) toevaluate how different alternative data sets {temperature,precipitation, solar radiation and soil texture) affect estim-ates of annual net primary productivity (NPP) for theconterminous United States, Annual NPP is the netamount of carbon assimilated by vegetation through theprocess of photosynthesis over a year. We chose toevaluate the sensitivity of terrestrial NPP because it is animportant integrative ecological measure. Also, NPP is amajor component of the global carbon cycle and thesensitivity of NPP has important consequences for theglobal climate system.

Methodology

The Terrestrial Ecosystem Model

In this study we use version 4.0 of TEM (McGuire et al.1996b). The TEM is a process-based ecosystem model(Fig. 1) that uses spatially referenced information onclimate, elevation, soils, vegetation and water availability(Fig. 2) to make monthly estimates of important carbonand nitrogen fluxes and pool sizes (Raich ct at. 1991;McGuire et at. 1992, 1993, 1996a, 1996b; Melillo et al.1993, 1995).

In TEM, NPP is calculated as the difference betweengross primary productivity (GPP) and plant respiration(RA)- The flux GPP is affected by several factors and iscalculated at each time step as follows:

GPP - f(LEAF) f(T) f(CO2, H2O) f(NA),

where C^ax is the maximum rate of C assimilation, PARis photosynthetically active radiation, LEAF is leaf arearelative to maximum armual leaf area, T is monthly airtemperature, COT is the atmospheric concentration ofcarbon dioxide, H2O is water availability, and NA isnitrogen availability. The calculation of R^ considers bothmaintenance respiration and construction respiration.

Atmospheric Carbon Dioxide

GPP R.

VEGETATION

NL Nvs

N U P T A K E L

SOIL

NUPTAKE

NINPUT KNETNMIN

NLOST

Fig. 1 The Tcrrt-striai Ecosystem Model (TEM). The statevariables are: carbon in the vegetation (Cy); structural nitrogenin the vegetation (Nvs); labile nitrogen in the vegetation (Nyj);organic carbon in soils and detritus (Q); organic nitrogen insoils and detritus (N<;); and available soil inorganic N (N^v)-Arrows show carbon and nitrogen fluxes: C!*P, gross primaryproductivity; R , autotrophic respiration; RH. heterotrophicrespiration; !<, litterfall C; L ,, litterfall N; NUPTAKEy N uptakeinto the structural N pool of the vegetation; NUPTAKEi Nuptake into the labile N pool of the vegetation; NRESORB, Nresorpdon from dying tissue into the labile N pool of thevegetation; NMOBIL, N mobilized between Ibe structural andlabile N pools of the vegetation; NETNMIN, net N mineralizationof soil organic N; NINPUT, N inputs from outside the ecosystem;and NLOST, N losses from the ecosystem (McGuire et al. 1993).

The functions in the GPP and R^ equations have beendescribed in previous work (Raich et al. 1991; McGuireet al. 1992, 1993).

The application of TEM requires spatially explicit datafor 10 input variables (Fig. 2); atmospheric CO2 andnitrogen inputs are considered to be constant. Data onmonthly temperature, monthly precipitation, monthlysolar radiation {or cloudiness), soil texture, elevation, andvegetation are used either as direct inputs to TEM orinputs to intermediate models that are used to generateinputs for TEM (Fig. 2). For example, hydrological inputsfor TEM are determined with a water balance mtxlel(WBM) that uses temperature, precipitation, solar radi-ation, elevation, soils and vegetation data (Vorbsmartyet al. 1989). The temperature, precipitation and solarradiation data used by TEM represent long-term averages.Tbf input data sets are gridded at a resolution of 0.5"latitude by 0.5° longitude.

To make estimates for a grid cell, TEM also needs thesoil- and vegetation-specific parameters appropriate tothe grid cell. Although many of the parameters in themodel are defined from published information, some ofthe vegetation-specific parameters are determined by

1996 Blackwell Science Ltd., Global Ghange Biology, 2, 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 7

BASIC VARIABLES INTEBMEOIATE MODELS TIM INPUTS

Fig. 2 Tlie inputs and outputs of theTt'rrt'stria! Ecosystem Model (TEM).Some input variables are generated usingintermediate models as described in thetext. PAR is photosynthetically activeradiation; PET is potential evapotran-spiration; AET is actual evapotranspir-ation; and CO2 is atmospheric CO2concentration.

Available TEM Outputs (monthly rosolution)

Oiou Prlmsry PtoOuctlvKvPlanl Hespl ration

NBI Prlmsry PioducilvliyDecompoElilon

Nil EcuysMm Productlvlly

LWeiiHll CBTbon

Pluni MHrogcn Uplaks

Llllerfali Nitrogen

Nel Nilmgen Mlneralliallon

Nltrogan LSBBH

VBQSfatton CarbonVegelatlon Nlliogen

Soil Organic C»rtionSoil Organic Nitrogen

Soil Inorganic Nitrogen

calibrating the model to the fluxes and pool sizes of anintensively studied field site. The data used to calibratethe model for different vegetation types are documentedin previous work (Raich cf al. 1991; McGuire et al. 1992,1996b), To calibrate the model for each vegetation type,we used the Legates & Willmott temperature and precip-itation data (Legates & Willmott 1990a 1990b), the solarradiation data derived from Hahn et al. (1988) cloudiness,and the modified FAO soil texture data (FAO/CSRC1974) for the grid cells containing the study sites.

Design of sensitivity experiments

For a simulation run, the system of models coupledwith TEM (Fig. 2) requires six spatially explicit variablesas inputs: vegetation, elevation, temperature, precipita-tion, solar radiation and soil texture. Several spatiallyexplicit data sets exist for each of these variables, soa series of sensitivity experiments may be conducted.Because different vegetation data sets use differentclassification schemes, we used only one vegetationdata set for all runs in the sensitivity experiments.Since elevation is used only as a switch to determinesnowmelt dynamics of a grid cell (below 500 m,snowmeit occurs in one month; above 500 m snowmeltoccurs in two months), variation in elevation has littleeffect on NPP estimates of TEM. Therefore, we usedonly one elevation data set for ail runs in the sensitivityexperiments. For the remaining variables, we set up aseries of three sensitivity experiments to examine theeffects of alternative data sets on NPP estimates by TEM:(i) alternative climates as defined by air temperature andprecipitation; (ii) alternative solar radiation; and (iii)alternative soil textures. Because air temperature andprecipitation data sets are usually available from thesame source, we investigated the combined effects ofthese 'climate' data sets on NPP estimates in the firstexperiment (Table 1).

Experiment 1: sensitivity to alternative climate data sets.In TEM, mean monthly air temperature directly influ-ences GPP, autotrophic and heterotrophic respirationrates {RA and RH)- plant nitrogen uptake (NUPTAKF),and soil nitrogen mineralization (NETNMIN); andindirectly influences leaf phenology, GPP, RH, N U P -TAKH, and NETNMIN via WBM outputs (Fig. 1, Fig. 2).Monthly precipitation directly influences WBM, andindirectly, through the WBM, influences leaf phenology,GPP, RH- NUPTAKE, and NETNMIN.

We used three climate data sets to investigatethe sensitivity of TEM to different temperature andprecipitation inputs: Legates and Willmott climate(L&W; Legates & Willmott 1990a 1990b), the climatedeveloped for the Vegetation/Ecosystem Modelling andAnalysis Project (VEMAP; Kittel et ai 1996; VEMAPMembers 1995), and Cramer and Leemans climate(C&L; Cramer, personal communication). The C&Lclimate data set is a major update of the Leemans andCramer (1991) data set, and is being widely used(Cramer, personal communication). We ran TEM foreach of three alternative climate data sets to determineclimate effects on NPP estimates (see experiment 1 inTable 1). For the baseline run we used the L&Wclimate, the Hahn-derived radiation and the FAO/CSRC soil texture data sets as inputs. For the runswith the C&L and VEMAP climate data sets, we usedthe same radiation and soil texture data sets.

Experiment 2: sensitivity to alternative solar radiation datasets. The solar radiation data sets used in this studyrepresent short-wave irradiance at the top of thecanopy. Solar radiation is used to calculate potentialevapotranspi ration (PET) based on the Jensen-Haisealgorithm (Jensen & Haise 1963). Potential evapotran-spiration influences the hydrological outputs of WBM.Solar radiation also influences the PAR output of theirradiance model. Through effects on hydrology and

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8 Y. PAN etai

Table 1. Design of sensitivity experiments'

Experiment 1 (climate)

Experiment 2 (radiation)

Experiment 3 (soil texture)

Data Sets

ClimateSolar RadiationSoil Texture

ClimateSolar RadiationSoil Texture

ClimateSolar RadiationSoil Texture

ClimateSolar RadiationSoil Texture

ClimateSolar RadiationSoil Texture

ClimateStjlar RadiationSolar Texture

ClimateSolar RadiationSoil Texture

Baseline Run

Hahn-derived''FAO/CSRC**

L&WHahn-derivedFAO/CSRC

C&LHahn-derivedFAO/CSRC

VEMAPHahn-derivedFAO/CSRC

L&WHahn-derivedFAO/CSRC

C&LHiihn derivedFAO/CSRC

VEMAPHahn-derivedFAO/CSRC

Comparison

L&CHahn-derivedFAO/CSRC

L&WC&L-derived^FAO/CSRC

C&LC&L-derivedFAO/CSRC

VEMAPC&L-derivedFAO/CSRC

L&WHahn-derivedZobler''

C&LHahn-derivedZobler

VEMAPHahn-derivedZobler

Comparison

VEMAl*^Hahn-derivudFAO/CSRC

L&WVEMAP^FAO/CSRC

C&LVEMAPFAO/CSRC

VEMAPHahn-derivedFAO/CSRC

L&WHahn-derivfdVEMAP'"

C&LHahn-derivedVEMAP

VEMAPHahn-derivedVEMAP

'All model runs used the Vegetation/Ecosystem Modelling and Analysis Project (VEMAP) vegetation distribution (Kittel et at. 1996;VEMAP Members 1995) and NCAR/NAVY (1984) elevation data.^Legates & Willmott (1990a, 1990b) monthly air temperature and precipitation data.Cramer and Leemans monthly air temperature and precipitation data (Cramer, personal communication).

*Monthly air temperature and precipitation data (NCDC, 1992) developed tor VEMAP (Kittel et al. 1996; VEMAP Members 1995).^ radiation data st't derived from Hahn et al. (1988) cloudiness data using the irradiance equation of Black et ai. (1954)

radiation data set derived from the Cramer and Leemans sunshine duration data set (Cramer, personal communication) usingthe irradiance equation of Black et at. (1954).-"VEMAP solar radiation data set, which were generated by CLMSIM (Running et al. 1987; Glassy and Running 1994) for VEMAP(Kittel ct al. 1996; VEMAP Members 1995).f'FAO/CSRC soil texture data digitized at 0.5° resolution from FAO-UNESCO's (1971) 'Soil Map of the World 1:5(1.000,000' (FAO/CSRC 1974).Vobler soil texture data digitized at 1" resolution from FAO-UNESCO's (1971) 'Soil Map of the World 1:50,1)00,000' (Zobler 1986).'"VEMAP soil texture data, which were aggregated from Kem (1994) 10-km gridded Soil Conservation Service NATSGO data to 0.5°resoluHon for VEMAP (Kittel et al. 1996; VEMAP Members 1995).

PAR, solar radiation influences leaf phenology, GPP,RH, NUPTAKE, and NETNMIN in TEM.

We used three solar radiation data sets to investigatethe sensitivity of TEM to different radiation inputs: thesolar radiation data set derived from Hahn et al. (1988)cloudiness, the solar radiation data set derived fromC&L sunshine duration (Cramer, personal commmiica-tion), and the VEMAP solar radiation data set (Kittelet al. 1996; VEMAP Members 1995). To determine theeffects of alternative solar radiation on NPP whilekeeping climate constant, we ran TEM three times foreach climate (see experiment 2 in Table 1). For eachclimate (L&W, C&L or VEMAP), all three runs usedthe same climate data set and the FAO/CSRC soil

texture data, Among the three runs, the baseline runused the Hahn-derived radiation data set, and theother two runs used either the C&L-derived radiationdata set or the VEMAP radiation data set.Experiment 3; sensitivity to alternative soil texture data sets.Stiil texture affects GPP, R,,, NUPTAKE, and NETNMINbecause the equations describing these fluxes in TEMhave parameters that depend on soil texture. In addition,soil texture affects the hydrological outputs of WBMbecause field capacity and wilting point depend on soiltexture and rooting depth depends on both vegetationtype and soil texture. Soil texture also indirectly affectsleaf phenology, GPP, RH, NUPTAKE, and NETNMINthrough effects on WBM outputs.

1996 Blackwell Science Ltd., Global Change Biology, 2, 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 9

LEGEND

Fig. 3 Potential vegetation distributionof the conterminous United States basedon the VEMAP vegetation classification.

Temp. Mixed Xeromorphic Wood.

Tropical Evergreen Forest

Tropical Deciduous Forest

Temperate Deciduous Forest

Warm Temp. Mixed/Everg. Forest

Cool Temp. Mixed Forest

Continental Temp. Conifer Forest

Maritime Temp. Conifer Forest

I Boreal Forest

Tundra

Subtropical Arid Stirublarxls

Temperate Arid Shrublands

I Medlterrean Shrublands

04 Grasslands

C3 Grasslands

I Tropical Deciduous Savanna

, Temperate Conifer Savanna

I Warm Temperate/S.T. Mixed Savanna

Temperate Deciduous Savanna

I Tropical Thom Woodland

I Temp. Conifer Xeromorptiic Wood.

We used three soil texture data sets to investigate thesensitivity of TEM to different texture inputs: the FAO/CSRC (1974) soils data set, the Zobler soils data set (1986)and the VEMAP soils data set (Kittel et al. 1996; VEMAPMembers 1995). To determine the effect of alternative soiltexture data sets on NPP, we also ran TEM three timesfor each climate (see experiment 3 in Table 1). For eachclimate (L&W, C&L or VEMAP), all three runs used thesame climate data set and the Hahn-derived radiationdata set. Among the three runs, the baseline run usedthe EAO/CSRC soil texture data set, and the other tworuns used either the Zobler soils data set or the VEMAPsoifs data set.

Description of the data sets

Data sets used for all simulations. The vegetation data setis required to define the vegetation-specific parametersfor each grid cell in the spatial extrapolation of TEM. Inthis study, we used the VEMAP vegetation distribution(Kittel et ai 1996; VEMAP Members 1995) for the conter-minous U.S. (Fig. 3), which is based on Kuchler (1964,1975). The vegetation types in the VEMAP data set areclassified on the basis of the physiognomic characteristicsof dominant lifeforms except for grassland vegetation

types, which are distinguished by photosynthetic path-way (C3 vs. C4). Elevation data are used in the WBM toaffect snowmelt and therefore affect soil moisture. Theelevation data used in this study represent an aggregationto 0.5° resolution of the NCAR/NAVY global 10-minuteelevation data set (NCAR/NAVY 1984).

Data sets used for sensitivity to climate. The L&W climatedata for the conterminous United States are derived fromthe global database of Legates and Willmott (1990a,1990b), which is based on long-term temperature recordsof 17 986 terrestrial stations and 6955 oceanic grid-points,and gauge-corrected precipitation records of 24 635terrestrial stations and 2223 oceanic grid-points. Datawere interpolated to 0.5° resolution globally using aspherical interpolation algorithm (Willmott et al. 1985).

The C&L data set was derived from the CLIMATEglobal database (Cramer, personal communication) whichis a major update of the Leemans & Cramer (1991)database. The database was developed from 10 to 40 yearrecords of 18 000 stations from across the globe, withabout 4000 in the conterminous United States. Datawere interpolated to 0.5° resolution globally using a3-D smoothing spline technique for all variables. Thesmoothing spline surfaces are functions of latitude, longit-

© 1996 Blackwell Science Ltd., Global Change Biology, 2, 5-23

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10 Y. PAN etal.

ude and elevation with the degree of data smoothingdetermined by minimizing the predictive error(Hutchinson & Bischof 1983). This interpolation methodis different from the method used to develop the Leemansand Cramer (1991) data sets and greatly improves thetopographic sensitivity of the interpolated data sets,especially in mountainous areas (Cramer, personal com-munication).

The VEMAP temperature data set (Kittel et al. 1996;VEMAP Members 1995) is based on 4613 station normals(NCDC 1992) within the conterminous United States. Thedata were adiabatically adjusted to sea-level (Marks 1990)before being interpolated to 0.5° resolution and were thenadjusted for the elevation of each grid cell as defined bythe elevation data set used in this study. Precipitationdata were spatially aggregated to 0.5° resolution fromthe 10-km gridded data set for the U.S. developed byPRISM (Daly et al 1994), The PRISM divides areasinto topographic facets of similar aspect, then developsprecipitation-elevation regressions for each facet basedon the regional station data. The regressions are used tospatially interpolate precipitation data for cells withsimilar facets.

Data sets used for sensitivity to radiation. The Hahn-derivedsolar radiation data set used in this study was derivedfrom the Hahn et al. (1988) cloudiness data set. The Hahnet al. (1988) cloudiness data set describes the long-termaverage global cloudiness for 3-month periods at 5°resolution. The data set was developed from surfaceobservations between 1930 and 1979; but observationsat different sites are not consistent in duration andmeasurement. For use by TEM, Hahn cloudiness datawere temporally and spatially smoothed. The average3-month cloudiness at 5° resolution was interpolatedtemporally into average monthly cloudiness & spatiallyto 0.5° resolution. Tbe solar radiation data was estimatedwith the Black et al. (1954) solar radiation model:

RTOC - RTOA [0.23 + 0.48 (Sp)],

where RTOC is short-wave irradiance at the top of thecanopy, RTOA is short-wave irradiance at the top of theatmosphere and So is percentage sunshine duration. Weused the algorithm of Turton (1986) to calculate RTOAand the quantity (1 - C) to represent SQ where C ispercentage cloudiness from the cloudiness data set.

The C&L-derived solar radiation data set used in thisstudy was derived from the C&L global monthly databaseof sunshine duration (Cramer, personal communication).Similar to the temperature and precipitation data sets,the C&L sunshine duration data set is a major update ofthe Leemans & Cramer (1991) sunshine duration data setwhich is based on observations from more stations andinterpolated using a 3-D smoothing spline fuction. The

C&L sunshine duration data set has been converted tothe solar radiation data set using the model of Blacket al (1954).

The VEMAP daily solar radiation data were estimatedby the CLIMSIM model (Running et al 1987; Glassy &Running 1994), which uses latitude, elevation, the diurnalrange of temperature, and the occurrence of precipitation.Monthly means were derived from the daily values (Kittelet al 1996; VEMAP Members 1995). To generate solarradiation data, the CLIMSIM model used the elevationdata set described earlier and daily temperature andprecipitation data sets from VEMAP (Kittel ('( al. 1996;VEMAP Members 1995). The monthly means of the dailydata match the long term monthly climatology of theVEMAP climate data sets.

Data sets used for sensitivity to soil texture. The EAO/CSRCsoil data set (1974) represents a digitization to 0.5"resolution of the UNESCO/FAO World Soil Map (FAO-UNESCO 1971). The seven classes In the FAO/CSRC soiltexture data set represent 'average' soil profiles based onthree FAO texture classes (Table 2). Each FAO/CSRC soiltexture class defines a combination of percentage sand,silt and clay (Table 2).

The Zobler soils data set (1986) represents a digitizationto r resolution of the UNESCO/FAO World Soil Map(FAO-UNESCO 1971). Each one-degree grid cell repre-sents the near-surface texture (upper 30 cm) of thedominant soil unit. Besides organic soils and land ice,there are seven texture classes which are identical tothose in the FAO/CSRC soil texture data set (Table 2).For use by TEM, the Zobler soils data were convertedfrom 1° resolution to 0.5° resolution by assigning thevalue of each 1° grid cell to its corresponding four 0.5°grid cells.

The VEMAP soils data set is based on the Kern (1994)10-km gridded Soil Conservation Service National Soildatabase (NATSGO). The data were aggregated to 0.5°resolution and grouped by cluster analysis to a set of oneto four modal soils. The first modal soil was used torepresent soil properties for the grid cell. The soil textureis characterized by percentage of sand, silt and clay(Kittel et al. 1996; VEMAP Members 1995).

Results

Sensitivity to alternative climate data sets

The C&L and VEMAP data sets represent cot)ler anddrier climates for the conterminous United States than isrepresented by the L&W climate, with VEMAP the coldestand C&L the driest (Table 3). In comparison to theL&W climate, mean annual temperature and annualprecipitation of the conterminous U.S. are 0.4 °C and

1996 Blackwell Science Ltd., Global Change Biology. 2, 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 11

Table 2. Percentage sand, silt and clay assigned to the FAO/CSRC soil textures

FAO/CSRC Code

1234567S

Combinati

1231 and 21 and 32 and 31, 2, and 3none

Class description 7o Sand Silt % Clay

SandLoam

ClaySandy loamLoamClay loamLoamLithosols

8045256545354545

1040302040304040

1015451515351515

*FAO codes: 1, coarse-textured; 2, medium-textured; and 3, fine-textured. No textures were assigned to lithosols.

Table 3. Mean climdte data and effects on annual NPP (10^^ g C y^') estimates for biomes of the conterminous United States

Biomes

TundraBoreal conifer forestMaritiniL' conifer forestContinental conifer forestCool temperate mixed forestTemperate conifer savannaTemperate deciduous forestTemperate deciduous savannaWarm temp/ subtrop mixed forestWarm temp/ subtrop mixed savannaCT, grasslandsC4 grasslandsTemperate mixed xeromorphic forestTemperate conifer xeromorphic forestMediterranean shrublandsTemperate arid shrublandsSubtropical arid shrublandsMean/Total

Mean annual temperature (

L&W^

5.54.4

10.76.66.57.1

11.812.917.217.76.9

11.414.610.315.28.0

17.911.2

C&L^

-3,4-1.4-2.5-1.4-0.1-1.2-0.1-0.1-0.2+0.2+0.1+ 0.3-1.2-0.8-3.4-0.6-1.0-0.4

VEMA

-6.0-3.8-3.5-3.2-1.1-2.9-0.8-0.6-0.7-0.5-0.3-0.3-2.2-2.1-3.4-1.8-1.0-1.2

"C) Annual

P^ L&W^

937.4630.6

1402.0592.1

1183.4387.8

1174.4973.8

1363.2672.6471.7641.4425.6331.9561.4303.4256.8782.7

precipitation (mm)

c&V

-602.9-238.4-683.2-246,5-152.6-136.8-104.8-105.5-92.6-54.9-86.7-94.0

-161.8-129.8-251.0-i9.6-42.1

-123.0

VEMAP3

-149.2+ 102.5+ 103.9+ 74.8

-167.0+ 29.7-72.5-70.3-67.0+5.2

-21.6-43.5+93.1+ 24.7-32,6+52.3+21.2-22.7

Annual

L&W^

1.7944.8787.98

198.25221.49

4.86706.53434.23782.0798.55

221.64383.2533.1289.0411.23

113.3956.31

3488.61

NPP'

C&L*

-26.91-18.07-13.38-16.31-1.41

-25.63-^.62-1.71-0.92^ . 4 9

-13.74-4,06

-23.74-25.35-30.25-13.26-12.81-5.24

VEMAP*

-14.63-20.27-9.65

-13.14-5.29-5.31-1.81-1.14-0.75+0.64-1.55-2.37

+9.59+4.39

+ 2.21+ 4.96

+ 10.62-2.27

'Hahn-derived sotar radiation and FAO/CSRC soil texture data sets were used for all simulations.-Baseline values for the biome: mean annual temperature in "C, armual precipitation in mm, and annual NPP in lO' ' g C y"'.•'Absolute difference with respect to L&W data.•^Relative difference (%) with respect to NPP estimates for L&W climate data.

123 mm less in the C&L climate, and 1.2 °C and 23 mmless In the VEMAP climate. Cooler and drier climatescause NPP estimates for the conterminous U.S. to decrease(Table 3). In comparison to NPP estimates for the L&Wclimate, NPP estimates are 5.2% less for the C&L climateand 2.3% less for the VEMAP climate.

At the resolution of biomes, there is a greater range intemperature and precipitation differences among thethree climates than at the continental resolution (Table 3).The VHMAP climate always has the lowest temperatures;the C&L climate generally has intermediate temperaturesexcept in a few biomes where temperatures are slightlyhigher than in the L&W climate (warm temperate/subtropical mixed savanna, C3 and C4 grasslands). For

some biomes there are large temperature differencesamong data sets. For example, mean annual temperaturein the C&L and VEMAP climates is 3.4 and 6.0 "C lessin tundra, 2.5 and 3.5 "C less in maritime conifer forests,and 3.4 °C less in Mediterranean shrublands. Lowertemperature in tundra, conifer forests and dry biomes inthe C&L and VEMAP climates generally reflects theeffects of elevation in the interpolated data sets, whichhave been ignored in the L&W data set. The VEMAPtemperatures were adiabaticatly adjusted for high eleva-tion, and the C&L temperatures were interpolated asfunctions of latitude, longitude and elevation with a 3-Dsmoothing spline technique. However, the C&L interpola-tion technique cannot adequately correct for elevation in

1996 Blackwell Science Ltd., Global Change Biology. 2, 5-23

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12 Y. PAN etal.

high elevation areas that do not have weather stations{Cramer, personal communication).

Among the three climates, the C&L climate generallyhas the lowest precipitation in almost ail biomes (Table3). The C&L precipitation is substantially lower in tundra,conifer forests (boreal conifer forest, maritime coniferforest, continental conifer forest) and several dry biomes(temperate mixed xeromorphic forest, temperate coniferxeromorphic forest, Mediterranean shrublands); thesebiomes are located in the western United States. TheVEMAP climate, in comparison to the L&W climate, hashigher annual precipitation in several conifer forests(boreal conifer forest, maritime conifer forest, continentalconifer forests, temperate conifer savanna) and drybiomes (temperate mixed xeromorphic forest, coniferxeromorphic forests, temperate arid shrubland and sub-tropical arid shrublands), Precipitation in the C&L climateis generally lower than the other climates because theinterpolation method cannot adequately correct for eleva-tion in high elevation areas. The L&W precipitation isgenerally higher in all biomes because gauge-inducedbiases from standard raingauge measurements, whichhave long been recognized as underestimates of actualprecipitation, have been removed from the L&W climate.The VEMAP precipitation has been adjusted for elevationand aspect by the 'orographically smart' interpolationprocedure of PRISM. Therefore, VEMAP precipitation ishigher for conifer forests and dry biomes that are locatedin mountainous regions.

The NPP estimates of TEM are sensitive to differencesamong climates at the resolution of biomes. In comparisonto estimates for the L&W climate, NPP estimates for theC&L and VEMAP climates are substantially lower intundra (14.6% to 26.9% lower) and in conifer forests (9.7%to 20.3% lower). The VEMAP climate in those biomes iscolder and generally moisten Colder temperatures aremore likely to cause lower NPP estimates by TEM inthose biomes where photosynthesis and decompositionare limited by low temperature (McGuire et ai 1992,1993;Melillo et ai 1993,1995). The C&L climate in those biomesis generally colder and drier. Both colder temperatureand lower precipitation are responsible for lower NPPbecause of slower rates of photosynthesis and decom-position.

Greater precipitation in dry biomes generally causeshigher NPP estimates. For several dry biomes (temperatemixed xeromorphic forest, temperate conifer xeromorphicforest, temperate arid shrublands, and subtropical aridshrublands), TEM estimates higher NPP with the VEMAPclimate (4.4% to 10.6% higher) which indicates higherprecipitation and lower temperature than the L&W cli-mate (21-93 mm higher and 1.0-2.2 °C lower). In contrast,TEM estimates lower NPP (12.8% to 25.4% lower) withthe C&L climate in the same biomes because precipitation

and temperature are lower than the L&W climate (42-162 mm lower and 0.8-1.2 °C lower). Water stress is acritical factor that affects NPP estimates in dry biomes;both higher precipitation and lower temperature alleviatewater stress. In Mediterranean shrublands, NPP is 2.2%higher for the VEMAP climate, which indicates 33 mmless precipitation and 3.4 °C lower temperature than theL&W climate. The lower temperature alleviates waterstress to more than offset potential stress induced bylower precipitation. However, for the C&L climate, NPPis 30.3% lower in Mediterranean shrublands because the3.4 C° lower temperature does not compensate for the251 mm less precipitation. The lower temperature doesnot compensate for water stress induced by lower precip-itation.

At the grid cell resolution, there are greater ranges oftemperature and precipitation differences among thethree alternative climate data sets than at the continentaland biome resolutions (Fig. 4). In comparison to the L&Wclimate, mean annual temperature in the C&L climateranges from = 10 C° lower to 4 C° higher, with 90"/" ofthe differences between 3 C lower and 1 C" higher;mean annual temperature in the VEMAP climate rangesfrom ^ 15 C° lower to 6 C° higher, with 90"''o of differencebetween 5 C° lower and 1 C° higher. Annual precipitationin the C&L climate ranges from 1676 mm lower to 425 mmhigher than in the L&W climate, with 90''^ of the variationbetween 397 mm lower and 54 mm higher. Similarly,annual precipitation in the VEMAP climate ranges from904 mm lower to 1426 mm higher, with 90"/.. of thevariation between 214 mm lower and 223 mm higher. Incomparison to NPP estimates for the L&W climate,the range of climate differences cause ^0"/<^ of the NPPdifferences to be more than 33.2% lower or more than6.4% higher for the C&L climate and more than 22.1%lower or more than 24.4"/,, higher for the VEMAP climate(Eig. 4, Fig. 5). Clearly, NPP estimates are more sensitiveto differences among alternative climate data sets at thegrid cell resolution than at the continental and biomeresolutions.

Sensitivity to alternative solar radiation data sets

At the continental resolution, the difference between thethree solar radiation data sets is substantial and theeffects of alternative radiation data on NPP estimates arelarge (Table 4). Mean solar radiation for the conterminousUnited States is 3.83 M] m"- d"' higher (-1- 32%) for theC&L-derived data set and 7.06 Mj m'^d'' higher (+ 60%)for the VEMAP data set than for Hahn-derived solarradiation. The higher solar radiation in the C&L-derivedand the VEMAP data sets causes lower NPP estimates.For any climate data set, TEM always estimates 8% to9% lower NPP with the C&L-derived and 10% to 11%

© 1996 Blackwell Science Ltd., Global Change Biology, 2, 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 13

Temperature Difference (vs. L&W)

C & L

-7

VEMAP

(degree)

Precipitation Difference (vs. L&W)

(a)

2 <

w

VEMAP ^(b)

-200 200 400(mm)

NPP Difference (vs. NPP with L&W climate)

VEMAP(c)

<-50 -20 20 40

Fig. 4 Comparisons of climate data sets; and TEM estimated NPP using different climate data sets, (a) Temperature difference (°C)between the C&L and the L&W data sets; and between the VEMAP and the L&W data sets, (b) Precipitation diffen^nce (mm) betweenthe C&L and the L&W data sets; and between the VEMAP and the L&W data sets, (c) Relative difference (%) between NPP estimatesbased on the C&L and the L&W climate data sets; and between NPP estimates based on the VEMAP and the L&W climate data sets.The Hahn-derived solar radiation data set and FAO/CSRC soil texture data set were used as other inputs for all simulations.

lower NPP with the VEMAP solar radiation (Table 4). InTEM, increased solar radiation may enhance PAR topotentially increase NPP, but enhanced PET may increasewater stress to potentially decrease NPP. The lower NPP

estimates with the C&L-derived and the VEMAP solarradiation data sets indicate that radiation-induced waterstress is stronger than the potential enhancement ofphotosynthesis by higher PAR.

© 1996 Blackwell Science Ltd., Global Change Biology, 2, 5-23

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14 Y. PAN etal.

Q.Q.Z

1nJCo>

!ati

a>

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30

20

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0

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-20

-30 -

-40

Climate

O

for L4W climatefor C4L climatefor VEMAP climate

Soil Texture

0 O 0

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o * .$. o ^ 0

C i L and VEMAP C&L and VEMAP Zobler and VEMAPvs. L&W (base) vs. Hahn-derived (base) vs. FAO/CSRC (base)

Fig. 5 The NPP sensitivities to thealternative climate, solar radiation andsoil texture data sets at the grid cellscale. Boxes indicate the range of NPPsensitivity from the 25"' to the ys""percentile; lines in the middle of theboxes indicate the median value of NPPsensitivity; lines extending from 25"* and75'^ percentiies indicate the range ofNPP sensitivity from the lO"' to the90" percentiies; and the open symtwisindicate the 5' and 95^^ percentiies.

At the biome resolution, the differences among solarradiation data sets are similar to the continental-resolutiondifferences (+ 2.95 to -i- 4.78 MJ m" d"' for the C&L-derived radiation and + 6.21 to + 8.19 MJ m" d"' forthe VEMAP radiation; Table 4). For all biomes, TEMestimates lower NPP for the C&L-derived and VEMAPradiation with any of the climate data sets (Table 4)because of radiation-induced water stress. The radiationeffect is most pronounced in dry biomes; the NPP estim-ates of cooler and moister biomes are generally lessaffected. For example, NPP in boreal forests is 0.5% to4.8% lower for the C&L-derived radiation, which estim-ates 3.50 MJ m-2 d"' higher irradiance; and 1.2% to 8.7%lower for the VEMAP radiation, which estimates 7.71MJ m" d"' higher irradiance than the Hahn-derivedradiation. In contrast, NPP of Mediterranean shrubiandsis 11.6-15.3% lower for tbe C&L-derived radiation, whichestimates 3.64 MJ m'' d"' higher irradiance; NPP Is 15.0-21.0% lower with the VHMAP radiation, which estimates6.21 MJ m" d" higher irradiance than the Hahn-derivedradiation. In comparison to the VEMAP climate, the effectof the VEMAP radiation on NPP estimates is generallymore severe for the L&W and C&L climates (Table 4).Either the higher temperature of the L&W climate orlower precipitation of the C&L climate could be factorsthat enhance radiation-induced water stress.

At the grid cell resolution, there is a greater rangeof solar radiation differences among alternative solarradiation data sets than at the continental and biomeresolutions (Fig. 6). In comparison to the Hahn-derivedradiation data set, mean annual solar radiation in theC&L-derived data set ranges from 1.29 MJ m" d"^-5.62MJ m" d"' higher, with 90% of difference between 2.62MJ m" d"' and 4.92 MJ m" d~' higher; mean annual

solar radiation in tbe VEMAP data set ranges from = 4.37MJ m-2d-'-8.33 MJ n r ^ d ' higher, with 907,, of differencesbetween 5.92 MJ m"- d^' and 7.84 MJ m - d ' higher.In comparison to NPP estimates for the Hahn-derivedradiation data set, the range of solar radiation differencesin the C&L-derived data set cause \0"/i>oi NPP differencesto be approximately more than 23% lower or more than1% higher for any of the three climate (Fig. 5, Fig. 6); therange of solar radiation in the VEMAP data set cause10% of the NPP differences to be approximately morethan 30% lower or more than 1% higher for any of thethree climates (Fig. 5, Fig. 6). Clearly, NPP estimatesare substantially more sensitive to differences amongalternative solar radiation data sets at the grid cellresolution than at the continental and biome resolutions.

Sensitivity to alternative soil texture data sets

The FAO/CSRC soils data set represents most biomeswith loam soils (== 50% silt plus clay. Table 5), except forcool temperate mixed forest which is represented withsandy loam soils (= 35% silt plus clay). In contrast, theVEMAP soils data set represents most biomes with clayloam and clay soils (65-75% silt plus clay) except for cooltemperate mixed forest which is represented with loamsoils. The soil textures in the Zobler soils data areintermediate between those of the FAO/CSRC and theVEMAP soils; most biomes are represented with loamand clayloam soils except for cool temperate mixedforest which is represented with sandy loam soils. Incomparison to the FAO/CSRC soils data set, these differ-ences cause the average soil texture of the conterminousU.S. to contain 6.7% more silt plus clay in the Zobler

1996 Blackwell Science Ltd., Global Change Biology, 2, 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 15

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Page 12: The importance of climate and soils for estimates of net

16 Y, PAN etal.

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C & L VEMAP

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(with L&W climate) ^ (b)

H .

C & L VEMAP

(with C&L climate)r

C & L VEMAP

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-50 -20 20 40m

Fig. 6 Comparisons of solar radiation data sets; and TEM estimated NPP using different solar radiation data sets, (a) Solar radiationdifference (MJ m" d'') between the C&L-derived and the Hahn-derived radiation data sets; and between the VEMAP and the Hahn-derived radiation data sets, (b, c, d) Relative difference {"/••) between NPP estimates based on the C&L-derived and the Hahn derivedradiation data sets; and between NPP estimates based on the VEMAP and the Hahn-derived radiation data sets, (b) The L&W climatedata sets and FAO/CSRC soil texture data set were used as other inputs for all simulations, (c) The C&L climate data sets and FAO/CSRC soil texture data set were used as other Inputs for all simulations, (d) The VEMAP cUn\ate data sets and FAO/CSRC soil texturedata set were used as other inputs for all simulations.

Page 13: The importance of climate and soils for estimates of net

CLIMATE, SOILS AND PRIMARY PRODUCTION 17

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Page 14: The importance of climate and soils for estimates of net

18 Y. PAN etai

soils data set, and 14.2% more silt plus clay in the VEMAPsoils data set.

Soil texture can affect NPP estimates of TEM by influ-encing GPP through effects on actual evapotranspiration(AET) and by influencing plant nitrogen uptake througheffects on volumetric soil moisture (VSM), which isthe percentage of the rooting zone occupied by water.Evapotranspiration depends on available water capacity,which is 15% of the rooting depth in WBM (based onRatiiff et al. 1983), In forests, rooting depth decreases forHner-textured soils (Thomthwaite & Mather 1957) andless water is available for transpiration in finer-texturedsoils. In grasslands and shrublands, rooting depthincreases slightly from sandy to loam soils, and decreasessubstantially from loam to clay soils (Thomthwaite &Mather 1957). Therefore, in grasslands and shrublandsmore water is available for transpiration in loam soilsthan in sandy or clay soils. Because the ratio of AET toPET controls canopy conductance in TEM (Raich ['( al.1991; McGuire et ai 1992) and because PET dependsonly on radiation and temperature in WBM, lower AETassociated with finer-textured soils has the potential tocause lower NPP. Although there is generally less avail-able soil water to transpire in clay loam and clay soils,VSM is generally higher in finer-textured soils becausefiner-textured soils have higher field capacity and wiltingpoint as fractions of the rooting zone than coarser-textured soils (Ratiiff et ai 1983). In TEM, both decomposi-tion rates and the diffusion of inorganic nitrogen in thesoil solution are enhanced by higher estimates of VSM(McGuire et al. 1996b). Because nitrogen availability toplants generally increases for higher rates of decomposi-tion and nitrogen diffusion, finer-textured soils tend toenhance NPP via higher VSM.

In comparison to the FAO/CSRC soils, finer-texturedsoils in the VEMAP data set causes = 2% less AETand 4% higher VSM for the conterminous U.S. for anyof the climates. The continental NPP estimate is 3%higher for the finer-textured VEMAP soils (Table 5)because the effects of VSM enhancements are strongerthan the effects of AET reductions. Similarly, finer-textured soils in the Zobler data set causes = 1% lessAET and 2% higher VSM for any of the climates. Thecontinental NPP estimate is only slightly higher (0.1%to 0.3% higher).

Average soil texture is finer for all biomes in theZobler and the VEMAP soils data. For the VEMAPsoils data, soil texture ranges from 2.7% more silt plusclay in warm temperate/subtropical mixed forest to22.4% more silt plus clay in cool temperate mixedforest. The NPP estimates of most biomes are generallyhigher for the finer-textured VEMAP soils, with = 5-9% higher estimates in cool temperate mixed forest, C4grasslands, and temperate mixed and conifer xero-

morphic forest (Table 5). For some climates, estimatesof NPP are less in warm temperate/subtropical mixedforest, tundra, maritime conifer forest, Mediterraneanshrublands and temperate and subtropical arid shrub-lands. As described above, finer-textured soils havesimultaneously opposite effects on NPP, i.e. a tendencyto reduce NPP because of lower AET and a tendencyto enhance NPP because of higher VSM. The changeof NPP depends on which effect is stronger in aparticular biome. For example, in cool temperate mixedforest the 6.0-7.0% higher NPP estimates for theVEMAP soils are primarily caused by greater than 5%enhancement in VSM; estimates of annual AET aredepressed less than 1% in this biome. In contrast, the0.5-0.6% lower NPP in temperate/subtropical mixedforest for the VEMAP soils cKcurs because the 0.6-0.7% higher VSM does not compensate for the effectsof lower AET on canopy conductance. The lower NPPestimates that occur for VEMAP soils in tundra andshrublands are asstKiated with estimates of annualAET that are less than 300 mm; the effect of enhancedVSM on nitrogen availability cannot compensate forlow canopy conductance in extremely dry environments.

The Zobler soils data set, which has soil texturesthat are intermediate between the FAO/CSRC andVEMAP soils data sets, causes higher NPP estimatesin more than half of the biomes with the L&W andVEMAP climates, but in less than half of the biomeswith the C&L climate. Decreases in NPP estimatesoccur when the effect of enhanced VSM on nitrogenavailability cannot compensate for reduced canopyconductance, This occurs more frequently for the C&Lclimate, which is drier than other two climates.

At the grid cell resolution, there is a greater rangeof differences between the soil texture data sets thanat the continental and biome resolutions (Fig. 7). Incomparison to the FAO/CSRC soil textures, percentagesilt plus clay in the Zobler soil texture data set rangesfrom 58% lower to 63% higher, with 90"' . of differencebetween 3% lower and 8% higher; percentage silt plusclay in the VEMAP soil texture data set ranges from47.0% lower to 73.0% higher, with 90? . of differencesbetween 10.0% lower and 37.0% higher. In comparistinto NPP estimates for the FAO/CSRC soil texture dataset, the range of differences in the Zobler soil texturedata set causes 10% of the NPP differences to beapproximately more than 5% lower or more than 7%higher (Fig. 5, Fig. 7). Similarly, the range of differencesin the VEMAP soil texture data set causes 10% of theNPP differences to be approximately more than 5%lower or more than \4% higher for any of the threeclimates (Fig. 5, Fig. 7). Clearly, NPP estimates aresubstantially more sensitive to differences among

1996 Blackwell Science Ltd., Global Change Biology, 1. 5-23

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CLIMATE, SOILS AND PRIMARY PRODUCTION 19

Soil Texture Difference (vs. FAO)

Zobler VEMAP

-50 -20 0(% silt plus day)

20 40

NPP Difference (vs. NPP with FAO Soils)

VEMAP

(with L&W climate)

VEMAP

(with C&L climate)

Zobler VEMAP

(with VEMAP climate)

-50 -20 0 20 40<

Fig. 7 Comparisons of soil texture data sets; and TEM estimated NPP using different soil texture data sets, (a) Stiil texture difference{% silt plus clay) between the Zobler and the FAO/CSRC soil texture data sets; and between the VEMAP and the FAO/CSRC soiltexture data sets, (b, c, d) Relative difference (%) between NPP estimates based on the Zobler and the FAO/CSRC soil texture datasets; and between NPP estimates based on the VEMAP and the FAO/CSRC soil texture data sets, (b) The L&W climate data sets andHahn-derived solar radiation data set were used as other inputs for all simulations, (c) The C&L climate data sets and Hahn-derivedsolar radiation data set werf used as other inputs for all simulations, (d) The VEMAP climate data sets and Hahn-derived solarradiation data set were used as other inputs for all simulations.

Page 16: The importance of climate and soils for estimates of net

20 Y. PAN eta!.

alternative soil texture data sets at the grid ceilresolution than at the continental and biome resolutions.

Discussion

It has long been recognized that spatial patterns of NPPat large scales can be explained by spatial patterns intemperature and precipitation (Lieth 1973, 1975). Spatialpatterns in NPP can also be explained by AET(Rosenzweig 1968), and the relationship between AETand solar radiation {Jensen & Haise 1963} suggests thatNPP is related to solar radiation at large spatial scales.For mature temperate conifer and deciduous forest standsin the same climate, field measurements of above-groundand below-ground production indicate that NPP increaseswith increasing silt plus clay content of the soil (Pastoret al. 1984; Nadelhoffer et at. 1985). Studies in grasslandsindicate that above-ground production increases withfiner soil texture when annual precipitation is greaterthan 370 mm, but decreases when precipitation is lessthan 370 mm (Sala et al. 1988).

It logically follows that model estimates of NPP atlarge spatial scales should be sensitive to alternativeinput data sets of temperature, precipitation, radiation,and soil texture. The standard method for evaluatingmodel performance at large spatial scales is to comparethe estimates of the model to field measurements. Forestimates of NPP at targe spatial scales, the performanceof TEM has been evaluated by Raich et ai (1991) andMelillo et ai. (1993). One way to evaluate the efficacy ofalternative input data sets might be to compare the NPPestimates of TEM to field measurements for each of theinput data sets. However, this approach has severalproblems. First, the comparison requires an analysis ofcovariance to compare the NPP estimates of TEM for thealternative input data sets with field measured NPP asthe covariate. Not enough site measurements of totalNPP (above plus below ground) have been made fordifferent sites within the conterminous United States toprovide the statistical power necessary for evaluatingthe efficicacy of different alternative input data sets. Inaddition, model estimates and site measurements ofNPP are at different spatial resolutions. A truly robustcomparison of model estimates with site data requiresreplication of field measurements within grid cells aswell as a sufficient sample size across grid cells of theconterminous United States.

Finally, for this study TEM is calibrated to one suite ofinput data sets (L&W climate, Hahn-derived radiation,and FAO/CSRC soils). This single calibration means thatwe cannot compare model estimates to field measure-ments among simulations based on alternative data sets.However, the evaluation of the sensitivity of NPP estim-ates to alternative input data sets has important implica-

tions for both the calibration and extrapolation ofbiogeochemistry models.

An important question that arises is: what level of NPPsensitivity is meaningful? Field measurements of NPPare uncertain, and ± 20% is probably a conservativeapproximation of the error associated with the measure-ment of NPP. Uncertainty in N PP at the grid cell resolutiondepends on both quantitative uncertainty of field meas-urements within the grid cell and spatial heterogeneityof NPP across the grid cell. Although estimates of NPP forsites within grid cells are generally too few to adequatelyestimate NPP at the grid cell resolution, uncertainty at acoarser spatial resolution is generally less than that at afiner spatial resolution (see O'Neill 1979 & Rastetter ct ai.1992). Thus, meaningful NPP sensitivity at the grid cellresolution is less than that at measurement sites, that atthe biome resolution is less than that at the grid ceilresolution, and that at the continental resolution is lessthan that at the biome resolution. Although we cannotquantitativeiy define the meaningful level of NPP sensit-ivity at these different spatial resolutiorxs, we can assumethat it is less than the measurement error of NPP.

The simulations in this study indicate that NPP estim-ates are less sensitive to differences among alternativedata sets at coarser spatial resolutions. Although themeaningful level of sensitivity for a particular spatialresolution is not quantitively defined, in our opinion gridcell sensitivities greater than 20%, biome sensitivitesgreater than 10' ., and continental sensitivities greaterthan 5% are meaningful differences. At the grid cellresolution, meaningful sensitivities are observed morefrequently for alternative climate and solar radiation datasets than for alternative soil texture data sets. At thebiome resolution, NPP estimates of most biomes appearto be sensitive to alternative climate and radiation datasets. It is not clear whether any of the biomes demonstratemeaningful sensitivity to alternative soil texture data sets.At the continental resolution, NPP estimates are mostsensitive to differences between solar radiation data sets.The greatest differences, which are = 10%, occur foralternative solar radiation data sets, and are associatedwith 60% differences in mean annual solar radiationbetween the VEMAP and Hahn-derived solar radiationdata sets. Meaningful sensitivity of NPP estimates for theconterminous United States is also observed between theL&W and C&L climates, but not necessarily between theL&W and VEMAP climates or the C&L and VEMAPclimates. It is not clear whether alternative soil texturedata sets cause meaningful sensitivity in continentalresolution estimates.

Among the three types of data sets used in this study,uncertainty may be greatest for solar radiation. Similarto the sensitivity of NPP estimates by TEM, the uncer-taintv in cloudiness and solar radiation at the earth's

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CLIMATE, SOILS AND PRIMARY P R O D U C T I O N 21

surface also has substantial effects on the projections ofclimate models (Cess et ai 1989, 1995; Ramanathan et al.1989, 1995).

The Hahn-derived and C&L-derived solar radiationdata sets are calculated from cloudiness or sunshineduration data that represent surface observations. Incomparison to the Hahn cloudiness data set, the C&Lsunshine duration data set has higher spatial resolutionand finer temporal resolution and appears to be moreaccurate. Our comparison of the C&L-derived solar radi-ation data sets with the contour maps of Bennett (1965),which are based on ground-measured solar radiation,indicate good agreement for the C&L-derived radiationdata throughout the year. In contrast, the Hahn-deriveddata set tends to underestimate solar radiation duringthe summer for whole U.S. (6.0 MJ m"" d"' lower). Thecoarse spatial and temporal resolution of the original datamay affect data interpolation and cause lower estimates ofsolar radiation. The VEMAP solar radiation is based onthe solar radiation estimates of the CLIMSIM model,which has been well-validated for estimates of daily solarradiation in the western U.S. (Running et al. 1987; Glassy& Running 1994). Our comparison of the VEMAP solarradiation data set with the contour maps of Bennett(1965) indicates that the VEMAP data set tends to over-estimate solar radiation in much of the U.S. Thus, theHahn-derived solar radiation and the VEMAP data setsappear to have biases. Removal of biases from solarradiation data sets is required to improve spatial patternsof NPP estimates.

Each of the climate data sets has its strengths inrepresenting the spatial pattern of monthly temperatureand precipitation. Both temperature and precipitationdata in the C&L climate data sets were adjusted forelevation with a 3-D smoothing spline technique. Thetemperature data in the VEMAP climate data sets wereadiabatically adjusted for elevation. One of the strengthsof the L&W climate data set is that there has beenan attempt to remove gauge-induced biases from theprecipitation data, and a strength of the VEMAP climatedata set is the orographically smart interpolation ofprecipitation data. The reduction of uncertainty requiresthat the strengths of the individual approaches be com-bined into a single approach. The THM simulationssuggest that the spatial resolution of NPP estimates inbiomes located in mountainous areas would benefit mostby the reduction of both precipitation and temperatureuncertainties.

Our comparison of the FAO/CSRC, Zobler and VEMAPsoils data sets indicates that the conterminous U.S. isprimarily represented by loam soils in the FAO/CSRCdata set, by loam and clay loam soils in the Zobler dataset and by clay loam and clay soils in the VEMAP dataset. Tlie FAO/CSRC and Zobler data sets are both based

on the Soil Map of the World (FAO-UNESCO 1971),which for the conterminous United States has betterspatial resolution than the NATSGO database (Kem 1995)from which the VEMAP soils data set was derived. TheFAO/CSRC data set was digitized at a finer resolutionthan the Zobler data set and represents soi! textures foraverage soil profiles rather than the dominant soil profilesin the Zobler data set. Similar to the Zobler data set, theVEMAP soils data set represents soil texture with thefirst modal soil profile. The TEM simulations suggest thatthe spatial resolution of NPP estimates in cool temperatemixed forest, Ci and C4 grasslands, and temperate mixedand conifer xeromorphic forests would benefit most bythe reductions of uncertainty in soil texture.

This study indicates that there is uncertainty in alternat-ive data sets that characterize the long-term contemporaryphysical environment across large spatial scales. Estim-ates of process-based ecosystem models like TEM aresensitive to this uncertainty, and estimates at small spatialscales are generally more sensitive to uncertainty thanestimates at large spatial scales. The reduction of uncer-tainty in input data sets is relevant for more than justimproving the spatial resolution of NPP estimates for thecontemporary condition; estimated impacts of globalchange that are based on the contemporary condition asa baseline are sensitive to both the accuracy of thecontemporary physical environment and the changesapplied to the contemporary physical environment.Assessments of the impacts of global change at largespatial scales will be improved by reducing uncertaintyin data sets that characterize the physical environment.The reduction of uncertainty is especially importantfor improving assessments of the regional impacts ofglobal change.

Acknowledgements

This work was funded by the Electric Power Research Instituteas a contribution to the Vegetation/Ecosystem ModellinR .indAnalysis Project (VEMAP), and by the National Aeronautics .indSpace Adtninistration (NAGW-714). We thank Dr XiangmingXiao for his assistance in producing the plates for this paper.

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