Modelling competitive ability of Neotropical savanna grasses: Simulation of shading and drought impacts on biomass production

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caiod,rogrelaVeneAccepted 10 October 2012Available online 17 October 2012Keywords:Biomassechnique to model the spatial distribution of shoots along vertical and horizontalphenology, and root and shoot architecture applied to threeNeotropicalEcological Informatics 13 (2013) 5769Contents lists available at SciVerse ScienceDirectEcological Ine lsThe savanna is one of the most important biomes of the planetcovering at least 20% of the global land surface and in some cases actsas a transition between tropical rainforest and deserts (Sarmiento,1983). Savannas are found in a wide gradient of physiognomies fromwoodland to grassland and they have high environmental and econom-ic value (Solbrig, 1993). Due to its ecological importance, the tropicalsavanna has been widely modeled to simulate one or several of fourprincipal determinants of its structure: soil moisture, soil nutrients,herbivory, and re (see reviews of Hill and Hanan, 2010; Sankaranet al., 2004; Staver et al., 2011a,b).Climate models predict diminishing trends of rainfall for the regionsavanna grasses. Competition among neighboring plants took intoaccount soil moisture and the spatial distribution of the individuals.The model determines the water requirements of each plant (in func-tion of their biomass and its rate of transpiration), and then calculated,using the convolution technique, how the demand for water is distrib-uted among the surrounding space. Using these data, we compared thebehavior of isolated plants, pairs and trios, and we found belowgroundcompetition to be a fundamental component of global (shoot+root)competition. (See Appendix A for more details).In this paper, we propose an extension of Segarra et al. (2009)modelto determine the shading from neighboring plants. Recent ecologicalcontaining the study area (Dai, 2010; Mata, 1have developed a model to study the respotypes of dominant grasses in the savanna,(2009). In that paper we presented a theore Corresponding author at: Universidad de Alicante. CamAp. 99 E-03080 (Alicante), Spain.E-mail address: (J. Segarra).1574-9541/$ see front matter 2012 Elsevier B.V. All competition for water, which couples soil water availability,1. IntroductionDroughtGrassesConvolutionTransition matrixEvapotranspirationBarinas, Venezuela, to explore the responses of the peak biomass of a plant community to a range of10-50% reduction of rainfall. We selected three dominant grass species: Elyonurus adustus, Leptocoryphiumlanatum, and Andropogon semiberbis in a 47 m study plot. We estimate parameters values from data mea-sured in the eld. The number of shoots for each plant is obtained according to soil water availability and dis-tributed vertically by 10 cm levels using a transition matrix. Convolution allows calculation of leaf area indexfor each cell and vertical level, which is then used to calculate light attenuation and thus the proportion ofshaded shoots in each cell and level. With this information, maximum evapotranspiration is determined tocalculate soil moisture using daily rain time series. Biomass is calculated for all species based on shoot bio-mass measured in the eld and re is simulated by removing a fraction of the shoots segments of all species.Modeled biomass ts reasonably well to eld data.The model predicts signicant differences in the response of each species to drought. A. semiberbiswas the mostdrought resistant of the three species, whereas L. lanatum was the least, and E. adustus was intermediate, inagreement with observations in the literature. Our model suggests that drought resistance increases with thebiomass/transpiration ratio for the species considered. 2012 Elsevier B.V. All rights reserved.Received 16 February 2012Received in revised form 2 October 2012dimensions of a plant community. We use it to simulate the growth of a tropical savanna near the city ofArticle history: In this paper, we develop a tModelling competitive ability of Neotropiand drought impacts on biomass productJ. Segarra a,, J. Ravents a, M.F. Acevedo b,c, J.F. Silvaa Departamento de Ecologa, Universidad de Alicante, Alicante, Spainb Electrical Engineering Department, Geography Department, and Environmental Science Pc Center for Simulation and Modeling (CESIMO), Universidad de los Andes, Mrida, Venezud Instituto de Ciencias Ambientales y Ecolgicas (ICAE), Universidad de los Andes, Mrida,a b s t r a c ta r t i c l e i n f oj ourna l homepage: www.996). For this reason, wense to drought of somebased on Segarra et al.tical stochastic model ofpus de Sant Vicent del Raspeig.rights reserved.l savanna grasses: Simulation of shadingnC. Garcia-Nez dam, University of North Texas, Denton, TX 76203, USAzuelaformaticsev ie r .com/ locate /eco l in fmodels tend to separate the canopy leaves into sunlit and shadedcategories, which is found to provide a considerable advantage and im-provement in the scale-up calculations of photosynthesis and transpira-tion from leaf to canopy (Gates, 2003; Wang et al., 2001). In particular,shading is important tomodel the effect of drought on transpiration andphotosynthesis (Aranda et al., 2005; Ashton et al., 2006).Models of light interception by the canopy fall into two categories.First, precise models, taking into account the exact position of every58 J. Segarra et al. / Ecological Informatics 13 (2013) 5769leaf, angle of incidence and the angle of leaves (de Moraes Frassonand Krajewski, 2010; Sekimura et al., 2000). All these models showedgood agreement with eld data but are not adequate for large plotswith many individual plants or for simulations at a daily time scale,since computing time would be too long.Second, models that calculate incoming radiation from the canopysurface without considering shoot angle (Thornley, 2002), or thosethat focus on above and below ground competition using convolutiontechniques (Barbier et al., 2008). These models execute faster thanthe models that include geometric details but do not account forplant morphology.Our paper is in between these two approaches; it uses fast com-puting convolution techniques (which we explain in Appendix A)and incorporates architectural traits of the plants. Consequently, themodel allows working with large plant communities in a preciseand rapid manner, using calculations for whole populations. More-over, the calculation of the spatial structure of each plant is easierthan with other models because it does not need to know the spatialposition of each shoot, but the entire set of shoots of each level is cal-culated statistically.In this paper, we model the dynamics of plant growth at twodifferent levels: a) plant level, taking into account vertical structure;b) community level, taking into account interspecic interactionsamong several plants. In order to simulate the drought scenarios themodel calculates the transpiration rates taking into account the spatialarrangement of the shoots, because transpiration rates could be higherin sun-exposed leaves compared with shaded ones. Further, we applythis technique at each vertical level each 10 cm in height in order totake into account shading of shoots. Here we call shoot the aerialpart of the plant which includes stem, sheath and leaf. Besides, themodel also takes into account the effect of re, therefore includingtwo of the most crucial factors in the structure of the savanna: reand soil moisture (Accatino et al., 2010).Our specic objectives are: 1) model shoot shading and transpira-tion rates, by adding spatial distribution of shoots along the verticaldirection to our previous competition model (Segarra et al., 2009);2) explore the effects of drought scenarios on biomass, and develop ametric of plant resistance to drought.2. Study site and eld methodsThe study site corresponds to a seasonal savanna near the city ofBarinas, Venezuela (8 38 N, 70 12 W). Mean annual temperatureis 27 C and mean annual rainfall is 1250 mm, with a rainy seasonfrom May to November and a dry season from January to March.We selected three grass species with different phenologies and archi-tectures, common in Neotropical savannas: 1) Elyonurus adustus,(Trin.) E. Ekman, a precocious growing, bunch grass with long andslender leaves; 2) Leptocoryphium lanatum, (Kunth) Nees, an earlygrowing, bunch grass with long coriaceous and narrow leaves; and3) Andropogon semiberbis, (Nees) Kunth, a tall and late oweringbunch grass, with short herbaceous leaves. For brevity, these threespecies will be denoted by the codes E, L, and A respectively.A rst set of eld data comes from a detailed sampling conductedat this site by Ravents and Silva (1988). Three individual plants(replicates) of each species were measured using a structure in whicha horizontal frame of 225 (1515) cells of 55 cmwas set at differentheights. For one year, monthly measurements of module density byheight were performed from 10 to 140 cm above ground at 10 cmintervals (vertical levels). The maximum number of levels achieved bya plant during the year varies by replicate and species; typically velevels for E, seven for L, and ten for A. For more details see Acevedoand Ravents (2002) and Ravents et al. (2004). A second set of datawas obtained in April 2006 from sampling a rectangular area of47 m at this site, measuring the coordinates of each individual plantand the number of shoots. The soil texture is loamy sand.We also measured transpiration, stomatal conductance, and CO2assimilation of the three species at different times of day and in twoperiods: in April 2006 (rainy season) and February 2007 (wet season).We used daily precipitation data from the weather station 804400(SVBI) in the State of Barinas, Venezuela, (8 61 N, 70 21W, altitude202 m) for the period 19912007 (, 2011). These datawere checked for homogeneity (Alexandersson, 1986) and theirmonthly averages are summarized in Table 1.3. Spatial resolution, time scales, and model stepsThe model spatial domain is the 47 m plot which is divided intwo nested horizontal grids. One grid is low-resolution of 11 msquares for soil moisture calculation, hereafter referred as quadrats,and the other is high-resolution with squares of 11 cm for shootnumber calculation, hereafter referred to as cells. Thus, there are100100 cells in a quadrat. The low- resolution grid denes matricesof dimension 47, with rows and columns denoted by (ix,iy), where-as the high-resolution grid denes matrices of dimension 400700,with rows and columns denoted by (kx,ky). Aboveground, up to140 cm, the spatial domain is divided in vertical levels of 10 cm,whereas belowground is vertically homogeneous.For each quadrat, soil moisture dynamics is calculated daily usingsimulated rain time series and shoot distribution. Conversely, plantgrowth and shoot distribution is calculated on a monthly basis usingmonthly averages of soil moisture.Fig. 1 summarizes the model whose steps are briey describedbelow. In the following sections we give the details for each calculation.Step 1) Number of shoots for each plant at basal vertical level isobtained monthly according to the monthly average of soilwater availability in the corresponding quadrat and speciesspecic parameters.Step 2) From the number of shoots at the basal vertical levelwe deter-mine the number of shoot segments at each higher level usinga transition matrix. A shoot segment is the portion of a shootbetween two consecutive vertical levels.Step 3) Using the high-resolution horizontal grid, we horizontallyspread the shoot number of each level by calculating the con-volution of a species-specic kernelwith the shoot number forall plants at that vertical level. These data are used to build a400700 matrix for each species with entries equal to thenumber of shoot segments for each cell and vertical level.Step 4) Leaf area index (LAI) is calculated for each cell and level. Thiscalculation is made using the matrices of shoot segments cal-culated above.Step 5) LAI values are used to calculate the proportion of light trans-mitted for each cell and level. We assume that this proportionis 1 at the top level where all shoot segments receive direct ra-diation. For each species, using the proportion of shoot seg-ments receiving direct radiation at each level we build amatrix with entries giving the proportion of shade in eachcell and level.Step 6) Multiplying the proportion of shade by the number of shootssegments in each cell we obtain the number of shoot seg-ments under shade and under direct radiation for each spe-cies.Step7) Maximumevapotranspiration, by species and total, is then cal-culated at each cell and aggregated to the quadrat to obtain avalue for the month at each quadrat.Step 8) Holding maximum evapotranspiration constant for every dayof the month, soil moisture is calculated in each quadrat on adaily basis, using te simulated daily rain time series. We thenuse monthly averages of soil moisture in each quadrat to cal-culate shoot production and mortality for each plant in thequadrat, going back to step 1.Step 9) Biomass is calculated in each cell for all species and aggregatedto the quadrat. This step is performed after step 3 as a sidecalculation and does not form part of the simulation loop.Step 10) Each February the model kills 90% of the shoots segments of allspecies in order to simulate mortality by re.3.1. Step 1: number of shootsThe total number of shoots of plant k of species j is the number atbase level or vertical level 1, which we denote as nj,k1 (t). This numberis given by a balance of gross production and mortality modeled byRichard's equation (Segarra et al., 2005, 2009) calculated on a monthlybasisdn1j;k t dtbj six;iy t j six;iy t gjn1j;k t 1n1j;k t Nj !gj !1Initial conditions for shoot number nj,k1 (0) were assigned from thevalues obtained from sampling the rectangular 47 m plot at the sitespecied in paragraph 2, giving the location coordinates of each individ-ual plant and its number of shoots as described in the previous section.These data were measured in April therefore the model starts cal-culations in that month. For soil water, initial conditions six;iy 0 wereassigned the value 0.2 for all quadrats, which is April's soil moistureaccording to Segarra et al. (2009).3.2. Step 2: shoots segments by levelIn Segarra et al. (2009)we calculated number of shoots as a functionof available soil water only and ignored growth restrictions due toshading. Tomodel effects of shadingwe need to know the spatial distri-bution of shoots segments at each level. For this we use the matrixmodel from Ravents et al. (2004) structured by 10 cm vertical levels.The three species differ in the maximum height reached by the end ofTable 1Monthly rainfall for Barinas, Venezuela. The rst three rows are obtained from the 19912007 dataset of station 804400 (SVBI). Hm: mean monthly precipitation; Hsd: standarddeviation; Jm: average number of rain days per month.Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov DecHm(mm) 13.1 9.4 34.1 64.1 151.5 184.6 164.1 209.9 190.8 124.7 83.5 20.5Hsd(mm) 27.3 18.5 31.2 50.6 92.8 100.6 79.3 101.2 122.9 58.7 50.6 39.4Jm 4 5 6 7 10 11 12 14 15 12 12 9st dmonoi aveontbio59J. Segarra et al. / Ecological Informatics 13 (2013) 5769where: gj is a coefcient controlling the shape of the function (dimen-sionless), Nj is maximum number of shoots, bj six;iy t is the grossshoot production rate (month1), and j six;iy t is shoot mortalityrate (month1). Both rates are dependent on soil water availabilitysix;iy t (monthly average) at the quadrat of row ix, column iy whereplant k is located. Functions bj six;iy t and j six;iy t use parametersbjmax and jmax respectively. These functions and soil moisture calcula-tion are explained in a later section. Parameter values are given inTable 2.Initial number of shoots2. Shoot segments by level1. Number of shoots3. Horizontal distribution4. Leaf area indexLaof8. Soil mmonthlym9. Calculate5. Shade per level6. Shaded shoot segments7. Max evapotranspiration 8. Daily soil wa8. Soil moFig. 1. Calculation steps fothe growing season: A reaches 10 levels, L reaches 7 levels and E reaches5 levels.Let's denote nj,k(t) as the vector of number of shoot segments forplant k of species j, at time t, with entries nj,ki (t) given by number of seg-ments at level i. As explained in the previous section, the total numberof shoots of plant k is the number at base level or level 1, nj,k1 (t). Amatrixequation is applied monthly to all plants of each of the three species j inorder to distribute the number of shoots of each plant by levelnj;k t 1 Mj nj;k t j;k t 1 2aythsturerage8. Water availabilityGrowth and mortality of shootsday=day+1Yesmonth=februaryNoh=month+1day=110. Mortality by fireYesNomasster balanceisture Initial soilmoisturellowed in the model.where Mj is a square level-transition matrix for species j with entriesrepresenting the transition from one level to levels above, j,k(t+1) isa ratio of shoot number change due to emergence and mortality de-scribed later, and the symbol represents the Hadamard or entry-wiseproduct of the two vectors. For more details see Appendix B.3.3. Step 3: horizontal distribution of shoot segmentsTo determine the horizontal spatial distribution, along axes x and y,of shoot segments of all species at each level we use the convolutiontechnique and a species-specic distribution kernel.To obtain the horizontal distribution Sj,i(x,y,t) of shoot segments ofall plants of species j at each level i at time t we calculate the convo-lution between Vj,i(x,y,t) (a matrix indicating the number of shootsthat there are in the position x,y) and the kernel Kj,i(x,y).The kernel is shown in Fig. 2, where we compared the densities ofshoot segments calculated from the kernel Kj,i(x,y) to those from elddata for each species using two levels, the rst (lowest) and a secondone corresponding to the mid-height of the plant. Agreement be-tween the eld data and the kernel results is better for E and L than A,and also better at level 1 than at the middle level. This indicatesTable 2Parameter values used to calculate shoot growth (from Segarra et al., 2009) and pa-rameters used to estimate dispersion coefcient Dj,i of bidimensional kernel.Species Shoot growth Dispersion kernelgj bjmax jmax Nj cj dj fjE 0.2 0.6 1 850 1.872 0.971 0.146L 0.2 0.6 1 360 1.066 0.594 0.342A 0.2 1 1 110 1.965 0.992 0.097-505-50500.10.2xFIELD DATAyDensity of shoots at level 1-505-50500.10.2xSIMULATION DATAy-505-50500.020.04xyDensity of shoots at level 5-505-50500.020.04xy500.10.2FIELD DATAity of shoots at level 1500.10.2SIMULATION DATAAB60 J. Segarra et al. / Ecological Informatics 13 (2013) 5769-505-50xyDens0.04t level 3-505-50500.02xyDensity of shoots aFig. 2. Spatial distribution of the frequency of shoot segments in 25 cm2 cells for each of the sTwo vertical levels are plotted: level 1 and the level corresponding to the species' average hThe coordinates x and y are spatial coordinates, with the origin (0.0) centered on the centr-505-50xy-505-50500.020.04xypecies. A) Andropogon semiberbis, B) Elyonurus adustus, and C) Leptocoryphium lanatum.eight. These distributions are used to obtain the distribution kernel of shoot axis of the plant.cont61J. Segarra et al. / Ecological Informatics 13 (2013) 5769that agreement is better for those species with higher number ofshoots (E and L) and also when segments are less dispersed, as isthe case at level 1.Adjustment between shoot segments distributions from the kernelmodel and those from eld data are very acceptable as suggested bythe mean quadratic errors shown in Table 3. For this comparison wecalculated the kernel using a 55 cm cell size since these are the di-mensions of the eld sampling cells (Fig. 2). Later, for simulations weinterpolated to another kernel of the same shape but calculated at thehigh-resolution cells of 11 cm.Fig. 3 shows the coordinates of each plant position in a 47 mrectangle. From these data plus the number of shoot segments ofevery plant (Eq. (2)) centered at the plant location coordinates we-505-50500.10.2xFIELD DATAyDensity of shoots at level 1-505-50500.020.040.06xyDensity of shoots at level 4CFig. 2 (obtain the values of matrix Vj,i(x,y,t). The results are exemplied inFig. 4 where we plotted together the values of the three species forlevel 1 (10 cm above ground).Appendix C provides details of how we calculate matrices V and K.With these matrices, we calculate the matrix Sj,i(x,y,t)Sj;i x; y; t Vj;i x; y; t Kj;i xx; yy dxdy 3Numerically, this convolutionwas conducted over the high-resolutiongrid and following methods of Borse (1997)Sj;i x; y; t if f t2 f f t2 Vj;i x; y; t f f t2 Kj;i x; y h i4where fft2 refers to the bidimensional fast Fourier transform, ifft2 to theinverse bidimensional fast Fourier transform, and the symbol representsthe Hadamard or entry-wise product of the two matrices. For moredetails see Appendix A.Hereafter we refer to two 400700 matrices, Vj,i (t) and Sj,i(t) forspecies j at level i resulting from discretization of functions Vj,i(x,y,t)and Sj,i(x,y,t) respectively at high-resolution. Non-zero entries ofVj,i(t) correspond to the number of shoot segments at level i of allplants of species j but lumped in the cells where plants are centered.Entries of Sj,i(t) contain the number of shoot segments in each cellfor species j and level i.Addition of the matrices Sj,i(t) across species produces a matrixSi(t) of total shoot segments at level i. Fig. 5 exemplies the resultsplotting the total number of shoot segments in each cell at level 1.We show only a 22 m section of the plot.3.4. Step 4: leaf area indexTo determine the amount of light transmitted downwards fromone level to the next we rst calculate the leaf area index (LAI) foreach cell at each level. For this we gathered information in the eldabout the geometric arrangement of shoots (Table 4) and madesome simple calculations assuming constant geometrical arrange-ment by level (Acevedo and Ravents, 2002). For each species j, wemeasured j as average shoot angle with respect to the horizontal(degrees) and wj as average shoot width (cm). Then calculated lj,-505-50500.020.040.06xyinued).-505-50500.10.2xSIMULATION DATAylength of shoot segments (cm) between levels, using lj =10/sin(j)cm, and laj, shoot area in cm2 using laj= ljwj. Note that because acell is 1 cm2, laj is directly the contribution to LAI for that shoot.Next we assume LAI is given by shoot surface, and used the productof laj times the matrix Sj,i(t)Aj;i t lajSj;i t 5where the matrix Aj,i(t) is the LAI of species j in level i. Entries of thismatrix contain the LAI for a particular cell. The total LAI in level i isthen obtained summing over all the speciesAi t jAj;i t : 63.5. Step 5: light attenuationWe need to estimate the fraction of light passing from one level tothe next immediately below in order to calculate the fraction of shootsegment under direct light or shade in each level. The number ofshoot segments at each level determines the number of shoots seg-ments receiving direct light at the level immediately beneath,whichde-creases from top to bottom.Weuse exponential attenuation (Monsi andSaeki, 1953)Ri t Ri1 t exp j jAj;i1 t !7where Ri(t) is the fraction of radiation reaching level i from the abovelevel i+1, j is the light extinction coefcient for species j. We assumeRi(t) at top level of that cell to be 1. According to Monteith andUnsworth (1973), j is given by the ratio of shaded area on the horizon-tal plane and the area of the shading element. Here we only considervertically incident light on the cell, and consequently j=cos j whichtakes values 0.5, 0.34, and 0.5 for E, L, and A respectively. These valuesare close to 0.4, the value proposed by Larcher (2003) for grasses.Values of the entries of matrix Ri(t) range from 0 to 1, being equalto 1 at the top of the level occupied at the cell and decreasing as afunction of number of shoot segments at each level. For each cellsubtracting the corresponding Ri(t) entry from 1 gives us the fractionof incident radiation attenuated by the canopy.3.6. Step 6: number of shoots segments under shadeNow we will assume that the fraction of incident light reaching aof species j that are in the shade and transpire isQ j t Xmji2Q j;i t : 11Table 3Mean square error between the kernel model and eld data for shoot segment number.These values are scaled by a 104 factor.Species Level1 2 3 4 5 6 7 8 9 10E 12.0 5.2 3.5 5.6 9.3 L 4.3 2.8 3.4 4.4 4.6 5.2 7.5 A 5.0 3.5 4.7 5.1 5.6 6.3 8.8 14.0 13.0 13.0100200300400010020030040050060070005001000y coordinate (cm)Number of shootsy coordinate (cm)500520540560580600620345678(x) A. semiberbis() E. adustus() L. lanatum 62 J. Segarra et al. / Ecological Informatics 13 (2013) 5769cell is available all over that cell and that shoot segments of that cellare exposed uniformly to that fraction of light. Then, entry-wise mul-tiplication of matrices Ri(t) and Sj,i(t) gives the number of shoot seg-ments by species and level that receive lightPj;i t Ri t Sj;i t : 8Also we form a matrix Qj,i(t) with entries qj,ikx,ky(t) given byqkx;kyj;i t 1pkx;kyj;i t 9where pj,ikx,ky(t) is the entry of Pj,i(t) at row kx and column ky.200300400500600700y coordinate (cm)0 100 200 300 4000100x coordinate (cm)Fig. 3. Spatial distribution of the plants from a eld rectangular plot of size 400700 cm.Symbols are: A: X; E: ; L: .Now, the total number of shoots segments of species j that receivedirect radiation and are transpiring isPj t Xmji2Pj;i t 10where, as dened earlier, mj is number of 10 cm-levels reached byplants of the species j, and the rst level i=1 is excluded from thesummation because shoots are non-photosynthetically active at thislevel (Ravents et al., 2004). Also the total number of shoot segments0 x coordinate (cm)Fig. 4. Number of shoots at level 1 for the three species from the three matrices Vj,1using the plant location information in Fig. 3. Symbols are: A: X; E: ; L: .x coordinate (cm)20 40 60 80 100 120 140 160 180 200440460480012Fig. 5.Actual distribution of shoot segments in space as given by thematrices Sj,1 at level 1.We show only a 22 m section of the plot. Each mound shows how the shoot segmentsdepicted in Fig. 4 share the space.Fig. 6 shows the frequency of shaded shoot segments per plant atlevel 1. We show only a 22 m section of the plot.3.7. Step 7: evapotranspiration rateTranspiration rate depends on several factors. We work here withthe maximum transpiration rate for each species achieved when soilwater content is at eld capacity. Using Pj(t) and Qj(t) we calculatethe maximum transpiration rate for each species j in each cellaccording to:Tj t Pj t Tj;l Q j t Tj;s laj 104 12Table 4Geometric arrangement of shoots (used to calculate LAI) and maximum daily transpi-ration rates. Transpiration rates were measured in April, 2006. Tj,l is rate under lightand Tj,s is rate under shade.Geometry Transpiration(cm day1)Species j () wj (cm) lj (cm) laj (cm2) Tj,l Tj,sE 60 0.33 11.54 3.48 0.24 0.06L 70 0.32 10.64 3.41 0.38 0.12A 60 0.67 11.54 7.06 0.17 0.1063J. Segarra et al. / Ecological Informatics 13 (2013) 5769where Tj,l and Tj,s are measured maximum transpiration rates of shootsegments of species j growing under light or under shade respectively(Table 4), laj is the area of the shoot segments we calculated before(Table 4), and the 104 factor is to convert cm2 to m2 (Table 4).Total maximum transpiration rate is obtained by summing over allthe species that occupy each cellTmax t jTj t : 13We assume that maximum evaporation rate from the soil (Emax)under direct sunlight averages to 0.15 cm day1 and 0 under shade.To determine how much light reaches each cell on the soil we usethe matrix R1(t) which is the fraction of radiation reaching level 1 cal-culated from Eq. (7).Emax t 0:15R1 t 14We neglect the shade created by level-1 shoots, which results in aslight overestimation of soil evaporation. coordinate (cm)5205405605806006200. A. semiberbis() E. adustus() L. lanatum x coordinate (cm)y20 40 60 80 100 120 140 160 180 20044046048050000.10.20.3Fig. 6. Frequency of shaded shoot segments per plant at level 1. Frequencies are closer to 1 atlevel 1 since this level is at the base of the plant.We show only a 22 m section of the plot.Next, we aggregate entries of the high-resolution matrices Emax(t)and Tmax(t) to the quadrat level. Recall that there are 100100cells in a quadrat. We form matrices Emax(t) and Tmax(t) whose entriesEmaxix,iy(t) and Tmaxix,iy(t) are obtained summing all entries ofmatrices Emax(t)and Tmax(t) respectively that belong to a quadrat of row ix, column iy.3.8. Step 8: soil moisture balanceAt this point we calculate soil moisture six,iy() in the quadrat ofrow ix, column iy at time . Here we use to distinguish time at thedaily scale from time t at the monthly scale. Variable six,iy() is relativesoil water content in relation to saturation level (dimensionless).Following Segarra et al. (2009), we write the balance of six,iy() asthe difference between inltration and loss rates (Laio et al., 2001)Zdsix;iy d I six;iy L six;iy E six;iy T six;iy 15where, : porosity of soil (dimensionless, 0.42), Z: depth of soil(30 cm), I(six,iy()): soil inltration rate (cm day1), L(six,iy()): soilwater loss rate by leakage or deep percolation, E(six,iy()): soil waterloss rate by evaporation, and T(six,iy()): transpiration rate. All ratesare given in cm day1 and will be explained next. We assume thatthere is no horizontal transport of water in the soil. In Appendix Dwe explain how to calculate the terms of Eq. (15).Initial conditions for the rst day, month, and year of the simula-tion six,iy(0) were assigned the value 0.2 for all quadrats, which isApril's soil moisture according to Segarra et al. (2009).After obtaining the daily soil water content, six,iy() from Eq. (15)for each month we average the values to obtain a monthly averagesix;iy t . This is then used to calculate the rate of shoot productionand mortality for each species j in the following manner.When soilmoisture is lower than thepoint belowwhichplants beginclosing their stomata (s), shoot production rate starts to decreaselinearly. When the soil moisture drops below the wilting point (sw),mortality increases from 0. Details are explained in Appendix E.Thenewvalues of shoot production andmortality are then substitutedin Eq. (1) to obtain the new number of shoots nj,k1 (t) and repeat themonthly cycle.3.9. Step 9: biomassTo build a relationship of shoots to biomass, in April 2006 we mea-Table 5Number of shoots at base level and biomass (dry weight) measurements (ve repli-cates) to calibrate the average relation of number of shoot segments to biomass. Mea-surements conducted in April, 2006.Species Measured EstimatedShoots Biomass (g) Biomass(g)/shootBiomass(g)/segmentE 154, 187, 350, 368, 381 75, 125, 140, 148, 213 0.500.11 0.400.12L 94, 134, 150, 188, 210 56, 57, 58, 94, 95 0.460.08 0.260.09A 47, 67, 72, 82, 95 46, 80, 100, 136, 137 1.330.26 0.310.06sured number shoots at basal level and biomass of ve individuals ofeach species (replicates). The measurements are reported in the rsttwo columns of Table 5. Biomass per shoot can be calculated by repli-cate, the average and standard deviation is reported in the third columnof Table 5.However, not all shoots are the same size and in order to accountfor vertical structure we proceed as follows. The number of shoot seg-ments by level is projected from base level using Eq. (2), and matricesMj, assuming sequences of emergence j,k1 (t+1) ratio at baselevel calculated as explained in Appendix B for xed differencesnj,k1 (t+1)nj,k1 (t) at all t of 80, 30, and 10 for E, L, and A respectively(values reported in Ravents et al., 2004). The vertical structure ofIf the current simulated month is a re-prone month (end of dry0 5 10 15 20 25 30 35 40 45 500100200300400500600700Time (months)Biomass (g/m2 )Fire30 %Biomass from Sarmiento (1984)Peak of maximumbiomassnnu64 J. Segarra et al. / Ecological Informatics 13 (2013) 5769shoot segments for each replicate is calculated by interpolating themeasured value of number of shoots at base level, and the numbersof shoot segments are summed across vertical levels for each repli-cate. Then, measured biomass of the replicate is divided into thisnumber of shoot segments to obtain biomass per shoot segment. Av-erage and standard deviation of biomass per segment (g/segment)across replicates are calculated and reported in the last column ofTable 5. Now we denote these average values as coefcients bmj foreach species j.Finally in each month we multiply scalar bmj by matrix Sj,i(t) ofnumber of shoot segments at level i of all plants of species j generatedin step 3 to estimateBj,i(t) biomass (dryweight in g) by species and level.Bj;i bmjSj;i t 16To evaluate the model we compared biomass to the values reportedin Sarmiento (1984). For this purpose, we conducted ve 5-year simu-lation runs. Each 5-year run starts in April with initial conditionsFig. 7. Model output showing changes in community biomass throughout time. Aexplained above. It was observed that the pattern of monthly dynamicsFig. 8. Changes in peak biomass along a gradient of annual rainfall using scenarios of % rainfall rThe slope of the regression line indicates the species' sensitivity to drought.season), then there is re-producedmortality of a fraction of the shoots.Therefore, each February, the model kills 90% of the shoots segments ofall species (Segarra et al., 2009). Although re intensity varies from yearto year, we donot include this variability andworkwith only an averageof 90% because re is the leading cause of mortality among the grassesof the savanna (Silva and Castro, 1989). Therefore the peak biomasscalculated in Step 9 is always the one corresponding to the month ofare repeated after the rst year, therefore year 1 was dropped from allrun results to remove transient behavior. As shown in Fig. 7, we com-pared the annual variation of the peak average biomass per squaremeter from the model over years 25 (48 months) of simulation withthe maximum biomass estimated in the eld for a similar community(Sarmiento, 1984).3.10. Step 10: mortality by real peaks of biomass are compared to biomass estimated from Sarmiento (1984).February just before the re.eduction, from0% (precipitation=1250 mm/year) to 50% (precipitation=625 mm/year).estimated marginal means have lower values and have higher vari-ability. Finally, species E is intermediate between L and A.Fig. 9. Estimated marginal means of log transformed peak biomass to ana65J. Segarra et al. / Ecological Informatics 13 (2013) 57694. Drought simulationsAccording to Dai (2010) rainfall in the area will decrease by 20%during this century due to climate change, one of the largest expecteddecreases worldwide. Because this is an uncertain number we exploredscenarios {0, 10, 20, 30, 40, 50%} encompassing the expected 20%change, including an intense drought scenario of 50%. Monthly valuesHm, Hsd, and Jm given in Table 1 were used for the base scenario(0% reduction). For all other scenarios, generated daily rainfall perStep 8 was reduced by the corresponding percentage. We reducedonly the amount of rain in a rain day and not the number of rain daysin a month.We conducted ve year simulation runs for each scenario. Each5-year run starts in April with initial conditions explained above. Thedaily rain sequence of each run for the base scenario uses a differentseed of the random number generator. Then the daily rain sequenceof the base scenario was reduced by 10, 20, 30, 40, and 50% to obtainthe daily rain sequence of the drought scenarios. For each runwe calcu-late the total plot peak biomass per species at the end of the fth simu-lation year. Then for each scenario, we calculated the mean andstandard deviation of the 5th-year total plot peak biomass per speciesof the ve runs.5. ResultsIn order to evaluate this model we have applied it to a savannacommunity, using actual plant distribution in a study plot (Fig. 3) andcalculating the spatial (vertical and horizontal) distribution of shootsegments and plant biomass in that community (Figs. 47). The evalu-ation was made by comparison of the maximum biomass from elddata and from the model's output (Fig. 7).The responses of the three species to the scenarios of increasingdrought are shown in Fig. 8. The comparison is made on the basis ofpercent reduction of peak biomass with respect to the base scenario.We calculated regression lines with zero intercept for each speciesusing the mean of the runs for each scenario. The slopes of the regres-sion lines obtained are 0.15, 0.17 and 0.11 in%/% for E, L and A indicat-ing that A is the most drought resistant, followed by E, and that L isthe least resistant. A two-way ANOVA shows that there is a signicantdifference in slope among species (F=1030.37; pb0.001) anddrought scenarios (F=130.50; p b0.001).However, interaction between species and draught level is also sig-nicant (F=4.455; pb0.001). Fig. 9 shows that there are signicantA posteriori pairwise comparisons of means using the Student-Neuman-Keuls (SNK) statistic for the three species show signicantdifferences among species, with A the least affected by drought (2.18),followed by E (1.89), and by L (1.31). Similarly, pairwise comparisonsby drought level using SNK show that drought scenarios of 50, 40, 30and 20% (with 1.45, 1.64, 1.79, and 1.87 respectively) are homogeneousand independent but the 10% and the 0% (no reduction) scenarios arenot (with 2.00 and 2.04 respectively). Therefore, reductions of 10% orlower do not have signicant effects on peak biomass.6. DiscussionIn this paper, we have developed a model using matrices and con-volution techniques, in order to simulate light incidence and transpi-differences for all drought levels except scenarios of 0 and 10% rainfallreduction. Species A is the least affected by drought because its esti-mated marginal means are higher and have lower range of variation.On the contrary, species L is the most affected by drought because itslyze interactions in ANOVA analysis of drought scenarios by species.ration rates of a vegetation community of the savannah. With thismodel, we have explored the effects of drought on these plants. Ourmodel suggests that drought resistance increases with the biomass/transpiration ratio for the species considered.The rst objective of this work was to incorporate shoot verticaldistribution, light incidence and transpiration rates into our previousmodel (Segarra et al., 2009; Segarra et al., 2010). The model presentedin this paper developed using matrices and convolution techniques,achieves this objective. Including shade in vegetation modeling is im-portant because it accounts for potential effects on ecophysiologicalprocesses that depend on light (Pearcy and Yang, 1996) and potentialeffects on plant morphology (Sekimura et al., 2000). The problem, asdiscussed in the introduction, is that it is difcult to develop a modelthat can be easily applied to large surfaces and on a daily basis, whiletaking into account the structure of plants involved calculating theshading between stems.To resolve this problem different models have been proposed. DeCastro and Fetcher (1998) divided the canopy into cubic cells, eachone characterized by mean leaf angle and the leaf area index. Althoughthis model is similar to ours, it has the drawback that it is computed cellby cell, making it more computer-intensive. Sinoquet et al. (2000)propose a simulation model of light partitioning in horizontally homo-geneous multispecies canopies. However this model only takes into66 J. Segarra et al. / Ecological Informatics 13 (2013) 5769account the diffuse radiation, so it could only be applied to overcast skyconditions. Rosati et al. (2001) use the partitioning of leaf area into oneof six classes to estimate canopy light interception and absorption. Thedrawback is that this approach only could be advantageous in cropswhere spacing is regular and plant material is genetically uniform,thus it is not applicable in heterogeneous media. Finally, Oyarzun etal. (2007) develop amodel of shading from trees and orchids, consider-ing trees as prismatic-shaped porous bodies, so it replaces the actualshape of the tree by a regular geometric gure and, furthermore, shad-ing is calculated tree by tree.Our model, by contrast, uses direct solar radiation, it can be appliedto plant communities with any spatial distribution and different mor-phologies, and it need not calculate each individually spatial unit, be-cause the convolution technique treats all plants of the same speciespooled in compartments instead of performing individual calculationsfor each plant. This improves calculation speed while preserving archi-tectural detail.To calculate soil moisture we have followed the same procedure asGuswa et al. (2002), assuming that there is a linear relation betweenevapotranspiration and soil moisture. To determine the effect of soilmoisture on plant growth we followed an approach similar toPorporato et al. (2001), who dene a constant stress following thesame equation of transpiration given by Guswa et al. (2002). Weuse the same equation given in Porporato et al. (2001), but substitut-ing the constant stress by one dependent on shoot production rate.Thus our equations for shoot production and transpiration are similar.We have added a new Eq. (E2) that calculates shoot mortality whensoil moisture is below the hygroscopic point.The evaluation of our model, made by comparison of the maxi-mum biomass from eld data and from the model's output (Fig. 7),shows that the model underestimates total biomass by 30% becausewe modeled only three species of this community. According toSarmiento (1983) these three species may represent about a 70% ofthe total biomass in this particular community, therefore we can ac-count for this 30% underestimation.Our second objective was to explore the effects of drought on theplant community. As expected, the three species respond differentlyto the drought scenarios except for the base and 10% rainfall reduc-tion scenarios. The lesser the slope in Fig. 8 the less pronounced isthe relative decrease in biomass, indicating more resistance todrought. The slope for A (0.11) is the lowest of the three, suggestingthat this species is more drought resistant, L has the highest slope(0.17), suggesting that it may be the least resistant, and E is interme-diate in value (0.15). Four factors interact in the model to determinespecies differences in drought resistance of peak biomass: shoot pro-duction rate, proportion of shaded shoots, shoot biomass, and tran-spiration rate. In fact, the ratio of shoot biomass to transpirationweighted by proportion of shaded shoots is 4.76, 1.96, 10.28 for E, Land A which are inversely aligned with the slopes 0.15, 0.17, 0.11for E, L and A. The larger the biomass/transpiration ratio the slope islower and the species is more drought resistant.These results are in agreement with reports from the literature indi-cating that L is less adapted to dry conditions (Sarmiento, 1983; Silvaand Sarmiento, 1976), than A and E which are present in drier condi-tions. Although there is not conclusive evidence, this differentialresistance to drought has been related to the growth phenology of thegrass species (Silva, 1987). Species starting growth before the onset ofrains (like E) or toward the end of the rain season (like A) should bemore resistant to drought than those species growing during the rainyseason (like L).Additionally, A and E have higher proportion of biomass allocatedto leaves than L (Table 5), and this may increase their ability to growin savannas. This allocation factor has been proposed as one of themain features contributing to a high competitive potential (Baruchand Bilbao, 1999; Baruch et al., 1989; Bilbao and Medina, 1990;Hodgkinson et al., 1989; Williams and Black, 1994).The results of our model are also consistent with eld measure-ments made by one of us (Garcia-Nunez, personal communication),showing that A closes stomata when the light radiation is high andhence higher evaporative demands and open when radiation is low,resulting in similar transpiration rates in both cases. Conversely L wasless responsive to changes in radiation load in terms of stomatalconductance, maintaining the stomata partially closed along the days,which in turn determine that transpiration rates depend more inevaporative demand. E showed an intermediate response.The lack of signicant peak biomass response for 10% rainfall reduc-tion or lower is due to the assumptions underlying model Eq. (E1)determining shoot production decrease as a function of soil moisture.We have assumed that shoot production is only affected when soilmoisture goes below s(soil moisture level below which plants beginclosing their stomata), and this critical level is not reached by smallerreductions of rainfall.Two major differences among species that can affect response todrought are not included in our model. One is underground/aerialbiomass ratio, which is high in L and E (2 to 5) and low in A (0.1)(Goldstein and Sarmiento, 1987; Silva, 1987). The other is differentialresponses to soil water stress since we have used the same values ofparameters sh, sw y s for all species as in Laio et al. (2001).7. ConclusionsWe have demonstrated that the integrated use of convolution, tran-sition matrices, shading calculations, and soil moisture balance allowsto model aboveground processes of savanna communities. The advan-tages of this technique are that it simplies the simulation of plantswith complex structures; the sensitivity analysis of the transition ma-trix can drawconclusions on thedynamics of growth;nally it increasesthe computational speed in the case of plant with complex structuresand high density, which is important to explore a variety of scenarios,such as drought or res.Themodel is applicable to other types of plant communities onceweinclude corresponding plant traits such as growth phenology and archi-tecture. Although we have used a matrix model is not a requirement.This technique can be applied to any plant growth model, provided tocalculate the intersection of each shoot in each vertical level. There areseveral processes that can be included to improve this model: 1) photo-synthetic rates to determine production and growth, 2) daily variationsin the angle of light incidence, and 3) species-specic values for soilwater stress parameters sh, sw and s.AcknowledgmentsWe thank the SpanishMinistry of Education and Science (CGL2007-65315-C03 02; CGL2008-05112-C02-01/CLI) and GA-LC-031/2010funded by CGA-La Caixa for their nancial assistance.Appendix AConvolution is a mathematical operator that transforms two func-tions, f and g, into a third function, which in a certain sense representsthe magnitude in which f and a shifted version of g are superimposed.Convolution is a very general type of moving average, as can be seen ifone of the functions is taken as the characteristic function of aninterval.The convolution of f and g is denoted as f*g. It is dened as the inte-gral of the product of both functions where one of them is shifted by .f t g t f g t d A1Our model uses convolution to calculate the spatial distribution ofthe shoots of a set of plants. This is needed because the matrix model67J. Segarra et al. / Ecological Informatics 13 (2013) 5769calculates the position of all the shoots of the plant in the central axis,andwe need to know how they are distributed spatially. The advantageof this technique is that it is not necessary to calculate the individualdistribution of each plant, but a single calculation will provide this forall plants within the same species. In this case, function f(t) of Eq.(A1) corresponds to the shoot dispersal kernel, which measures thenormal distribution of the shoots from the central axis of each plant.Our model supposes a Gaussian kernel, whichmeans that shoot densityismore intense in the central axis of the plant (the point fromwhich thestem starts), and becomes less intense toward the edge of the plant.Convolution is only applied to plants of the same species, as each spe-cies has a different shoot structure, and therefore a characteristic kernel.Function g(t) is formed by a set of segments distributed on the plane.The position of each segment on the plane reproduces the position ofeach plant in the model domain, and the height of each segment mea-sures the number of shoot segments for each plant.We use numericalmethods to solve Eq. (A1). A description of how toimplement these calculations using MATLAB is given below. This pro-gram shows how to apply this technique to two-dimensional surfaces.The text after the% sign are comments that do not affect execution ofthe program.%****************************************% Two-dimensional model with several elements% Plot consists of 128128 points% Values range between 10 to +10 function; convolution2dnp=128;xl=10; dx=2xl/np;x=linspace(xl,xldx,np);[X,Y]=meshgrid(x,x);% Set initial conditions for six plants. Numbers in parentheses are% plant coordinates. P is number of shoot segments of each plantP(56,20)=5;P(50,38)=1;P(22,80)=3;P(27,40)=7;P(90,78)=8;P(110,110)=5;% Denes dispersal kernelD=2;K=1/(2*pi*D^2)exp((X.^2+Y.^2)/(2*D^2));% Obtains the Fourier transform of matrix P and the dispersal% kernel.fP=fft2(P);fK=dx.^2*fft2(K);% The two matrices obtained in the previous step are multiplied% entry-wise.Pt=real(fftshift(ifft2(fP.*fK)));% The dispersed distribution is normalizednumPt=dx^2*trapz(trapz(Pt));Pt=numP/numPt*Pt;% Graphsurf(x,x,Pt)xlabel(Axis x)ylabel(Axis y)zlabel(Frequency of shoot segments)%****************************************Appendix BThe vectors nj,k(t) and j,k(t+1) are mj1, where mj is number ofvertical levels reached by plants of the species, mE=5, mL=7, andmA=10 for E, L and A respectively. MatrixMj ismjmj. The transitionmatrices for species E, L, and A are (Ravents et al., 2004):ME 1 0 0 0 0:18 :82 0 0 00 :44 56 0 00 0 39 :61 00 0 0 :07 :93266664377775ML 1 0 0 0 0 0 0:24 :76 0 0 0 0 0:096 :9 :004 0 0 0 00 :03 :52 :45 0 0 00 0 0 :4 :6 0 00 0 0 0 :22 :78 00 0 0 0 0 :07 :9326666666643777777775MA 1 0 0 0 0 0 0 0 0 0:25 :75 0 0 0 0 0 0 0 00 1:78 :11 0 0 0 0 0 0 00 :32 :84 0 0 0 0 0 0 00 0 :37 :63 0 0 0 0 0 00 0 0 :66 :34 0 0 0 0 00 0 0 :03 :97 0 0 0 0 00 0 0 0 :27 :73 0 0 0 00 0 0 0 0 :51 :49 0 0 00 0 0 0 0 0 :89 :1 :01 026666666666666643777777777777775The j,k(t+1) ratio of shoot emergence and mortality is rst calcu-lated at level 11j;k t 1 n1j;k t 1 n1j;k t B1And then at all other levels i=2, , mjij;k t 1 1j;k t 1 when 1j;k t 1 11 when 1j;k t 1 > 1(B2The rst entry j,k1 (t+1) of the ratio vector is calculated from thechange of the total number of shoots nj,k1 () from time t to time t+1,calculated in Section 3.1, according to Eq. (1).Appendix CThe three species have the same growth pattern, similar to aninverted cone whose diameter increases upwards, but the species dif-fer in rate of change of diameter as they grow up. Based on this weused a Gaussian bidimensional distribution for the kernel Kj,i(x,y)for level i and species j, given byKj;i x; y 12Dj;i2 exp x2 y2 2Dj;i20@1A C1where Dj,i measures the dispersion coefcient for species j and level i.We use an empirical power function to calculate the coefcient Dj,ifor each level i, being lowest at the base and increasing with heightDj;i cj i dj f j C2The parameter values for each species were obtained by matchingeld data of shoot segments distribution at each level (Table 2).With the kernel for shoot segment distribution, we now horizon-tally distribute the number of shoot segments assuming that plant kis located at point (xk, yk). We use the rst set of data from the eldwith the coordinates of each plant position in a 47 m rectangle68 J. Segarra et al. / Ecological Informatics 13 (2013) 5769(Fig. 3) to dene a 2D function for each species j and level i at time tVj;i x; y; t knij;k t xxk; yyk C3by summing over all plants k of species j a set of Dirac delta functionswith intensities given by the number of shoot segments and centeredat the plant location coordinates. The results are exemplied in Fig. 4where we plotted together the values of the three species for level 1(10 cm above ground).Appendix DRainfall of the day h() in cm day1 is randomly generated usingan exponential distribution function for each month t, according tothe mean monthly precipitation, and the average rainfall for rainydays of each month t (Segarra et al., 2009).Frequency of rainy days in each month jm(t) is calculated as a ratioof Jm(t), number of days with rain in the month m, to the number ofdays in that month (Table 1). Then, the ratio of Hm(t), the averagerainfall in the month m, to Jm(t) for the same month gives the averagerainfall for every rainy day of the t month, Hd(t), in cm day1. This isto say,Hd t Hm t Jm t . Then, for each day of the t month, we generateda random number rn uniformly distributed on [0,1]. If rn> jm(t), thereis no precipitation and then h()=0. Otherwise if rn jm(t), there isprecipitation on that day, and the amount h() is calculated generat-ing an exponentially distributed random number with cumulativeF(h)=1exp(h()/Hd(k)).Canopy interception rc, with a value 0.05 cm day1 as in Segarraet al. (2009), is subtracted from the rainfall to obtain effective rainfallhe h rc D1We assume the same value he() for all quadrats. Soil inltrationrate is calculated from the effective rainfall but limited by the decit(1s)Z for larger amounts of rainfallI six;iy 0 f or he 0he f or 0bhe b 1six;iy Z1six;iy Z f or he 1six;iy Z8>>>: D2The next equations (D3)(D5) are from Guswa et al. (2002). Theloss terms are calculated asL six;iy Ksexp six;iy sfc 1 exp 1sfc 1 sfcbsix;iy b10@ D3E six;iy 0 f or 0bsix;iy shEix;iymax t six;iy shsshf or shbsix;iy sEix;iymax t f or sbsix;iy 18>>>>>:D4T six;iy 0 f or 0bsix;iy swTix;iymax t six;iy swsswf or swbsix;iy sTix;iymax t f or sbsix;iy 18>>>>>:D5where Ks: hydraulic conductivity at saturation (100 cm day1), :parameter dening the exponential relationship between soil mois-ture and hydraulic conductivity (dimensionless, 12.7), sfc: soil eldcapacity (dimensionless, 0.52), sh: hygroscopic point (dimensionless,0.08), sw: permanent wilting point, (dimensionless,0.11), and s: soilmoisture level below which plants begin closing their stomataix,iy(dimensionless, 0.31). In addition: Emax(t): maximum evaporationrate (cm day1) for the month t, Tmaxix,iy(t): maximum transpirationrate (cm day1) for the month t. Recall that these last two quantitiesare calculated in step 7.Appendix EWhen soil moisture is lower than the point below which plantsbegin closing their stomata (s), shoot production rate starts to de-crease linearly, becoming equal to zero when moisture reaches thewilting point (sw),bj six;iy t 0 f or six;iy t bswbjmaxsix;iy t swssw f or swsix;iy t sbjmax f or six;iy t > s8>>>: E1where bj max is the rate corresponding to water above s* for species j.When the soil moisture drops below the wilting point (sw), mor-tality increases from 0 in proportion to the difference sws , j six;iy t 0 f or six;iy t > sw jmaxswsix;iy t swsh f or shbsix;iy t sw jmax f or six;iy t sh8>>>: E2where j max is the mortality rate corresponding to water below sh forspecies j.ReferencesAccatino, F., De Michele, C., Vezzoli, R., Donzelli, D., Scholes, R., 2010. Treegrass co-existence in savanna: interactions of rain and re. Journal of Theoretical Biology267, 235242.Acevedo, M.F., Ravents, J., 2002. Growth dynamics of three tropical savanna grassspecies: an individual-module model. Ecological Modelling 154, 4560.Alexandersson, H., 1986. A homogeneity test applied to precipitation data. Journal ofClimatology 6, 661675.Aranda, I., Castro, L., Pardos, M., Gil, L., Pardos, J.A., 2005. Effects of the interactionbetween drought and shade on water relations, gas exchange and morphologicaltraits in cork oak (Quercus suber L.) seedlings. Forest Ecology and Management210, 117129.Ashton, M.S., Singhakumara, B.M.P., Gamage, H.G., 2006. Interaction between light anddrought affects performance of tropical tree species that have differing topographicafnities. Forest Ecology and Management 221, 4251.Barbier, N., Couteron, P., Lefever, R., Deblauwe, V., Lejeune, O., 2008. Spatial decouplingof facilitation and competition at the origin of gapped vegetation patterns. Ecology89 (6), 15211531.Baruch, Z., Hernndez, A.B., Montilla, M.G., 1989. Dinmica del crecimiento, fenologa yreparticin de biomasa en gramneas nativas e introducidas de una sabanaNeotropical. Ecotropicos 2 (1), 113.Baruch, Z., Bilbao, B., 1999. Effects of re and defoliation on the life history of native andinvader C-4 grasses in a Neotropical savanna. Oecologia 119, 510520.Bilbao, B., Medina, E., 1990. Nitrogen-use efciency for growth in a cultivated Africangrass and a native South American pasture grass. Journal of Biogeography 17,421425.Borse, G.J., 1997. Numerical Methods with Matlab. PWS Publishing Company, Boston,MA.Dai, A., 2010. Drought under Global Warming: A Review. Wiley InterdisciplinaryReviews: Climate Change. Castro, F., Fetcher, N., 1998. Three dimensional model of the interception of light bya canopy. Agricultural and Forest Meteorology 90, Moraes Frasson, R.P., Krajewski, W.F., 2010. Three dimensional digital model of amaize plant. Agricultural and Forest Meteorology 150, 478488 (3rd ed.).Gates, D.M., 2003. Biophysical Ecology. Dover Publications, Inc., Mineola, New York .612 pp.Goldstein, G., Sarmiento, G., 1987. Water relations of trees and grasses and their conse-quences for the structure of savanna vegetation. In: Walker, B.H. (Ed.), TropicalSavanna Determinants. IRL Press, Oxford, pp. 1338.Guswa, A.J., Celia, M.A., Rodriguez-Iturbe, I., 2002. Models of soil moisture dynamics inecohydrology: a comparative study. Water Resources Research 38 (9), 5-15-15.Hill, M.J., Hanan, M.P., 2010. Ecosystem Function in Savannas: Measurement andModeling at Landscape to Global Scales. CRC Press.Hodgkinson, K.C., Ludlow, M.M., Mott, J.J., Baruch, Z., 1989. Comparative responses ofthe savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation. Oecologia79, 4552.Laio, F., Porporato, A., Ridol, L., Rodrguez-Iturbe, I., 2001. Plants in water-controlledecosystems: active role in hydrologic processes and response to water stress II.Probabilistic soil moisture dynamics. Advances in Water Resources 24, 707723.Larcher, W., 2003. Physiological Plant Ecology, 4th ed. Springer, Germany.Mata, L.J., 1996. A study of climate change impacts on the forest of Venezuela. In:Smith, J., Bhatti, N., Menzhulin, G., Benioff, R., Budyko, M.I., Campos, M., Jallow, B.,Rijsberman, F. (Eds.), Adapting to Climate Change: An International Perspective.Springer- Verlag, New York, NY, USA.Monsi, M., Saeki, T., 1953. ber den Lichtfaktor in den Panzengesellschaften und seineBedeutung fr die Stoffproduktion. Japanese Journal of Botany 14, 2252.Monteith, J.L., Unsworth, M., 1973. Principles of Environmental Physics. Butterworth-Heinemann, Oxford.Oyarzun, R.A., Stckle, C.O., Whiting, M.D., 2007. A simple approach to modelingradiation interception by fruit-tree orchards. Agricultural and Forest Meteorology142 (1), 1224.Pearcy, R.W., Yang, W., 1996. A three-dimensional crown architecture model forassessment of light capture and carbon gain by understory plants. Oecologia 108,112.Porporato, A., Laio, F., Ridol, L., Rodrguez-Iturbe, I., 2001. Plants in water-controlledecosystems: active role in hydrologic processes and response to water stress III.Vegetation water stress. Advances in Water Resources 24, 725744.Ravents, J., Silva, J.F., 1988. Architecture, seasonal growth and interference in threegrass species with different owering phenologies in a tropical savanna. Vegetatio75, 115123.Ravents, J., Segarra, J., Acevedo, M.F., 2004. Growth dynamics of tropical savanna grassspecies using projection matrices. Ecological Modelling 174, 85101.Rosati, A., Badeck, F.W., Dejong, T.M., 2001. Estimating canopy light interception andabsorption using leaf mass per unit leaf area in Solanummelongena. Annals of Botany88, 101109.Sankaran, M., Ratnam, J., Hanan, N.P., 2004. Treegrass coexistence in savannasrevisited insights from an examination of assumptions and mechanisms invokedin existing models. Ecology Letters 7, 480490.Sarmiento, G., 1983. The savannas of tropical America. In: Bourlire, F. (Ed.), TropicalSavannas. Ecosystems of the World, vol 13. Elsevier Scientic Publishing Company,Amsterdam-Oxford-New York, pp. 245285. 730 pp.Sarmiento, G., 1984. The Ecology of Neotropical Savannas. Harvard University Press,Cambridge. Mass . 235 pp.Segarra, J., Ravents, J., Acevedo, M.F., 2005. Growth of tropical savanna grass plants incompetition: a shoot population model. Ecological Modelling 189, 270288.Segarra, J., Acevedo, M.F., Ravents, J., Garca-Nez, C., Silva, J.F., 2009. Coupling soilwater and shoot dynamics in three grass species: a spatial stochastic model onwater competition in Neotropical savanna. Ecological Modelling 220, 27342743.Segarra, J., Ravents, J., Acevedo, M.F., 2010. Savanagua: a spatially explicit competitionmodeling of savanna ecosystems. In: Veress, B., Szigethy, J. (Eds.), Horizons inEarth Science Research, volume 4. Nova Publishers.Sekimura, T., Roose, T., Li, B., Maini, P.K., Suzuki, J., Hara, T., 2000. The effect of populationdensity on shoot morphology of herbs in relation to light capture by leaves.Ecological Modelling 128, 5162.Silva, J.F., 1987. Dynamics responses of savannas to stress and disturbance: speciesdynamics. In: Walker, B.H. (Ed.), IUBS n. 3, Determinants of Tropical Savannas.IRL Press, Oxford, pp. 141156.Silva, J.F., Castro, F., 1989. Fire, growth and survivorship in a Neotropical savanna grass(Andropogon semiberbis in Venezuela. Journal of Tropical Ecology 5, 387400.Silva, J.F., Sarmiento, G., 1976. La composicin de las sabanas en Barinas en relacin conlas unidades edcas. Acta Cientca Venezolana 27, 6878.Sinoquet, H., Rakocevic, M., Varlet-Grancher, 2000. Comparison of models for dailylight partitioning in multispecies canopies. Agricultural and Forest Meteorology101, 251263.Solbrig, O.T., 1993. Ecological constraints to savanna land use. In: Young, M.D., Solbrig,O.T. (Eds.), The World's Savannas: Economic Driving Forces Ecological Constrainte.Parthenon Press, Paris, pp. 2148.Staver, A.C., Archibald, S., Levin, S.A., 2011a. Tree cover in sub-Saharan Africa: rainfalland re constrain forest and savanna as alternative stable states. Ecology 92,10631072.Staver, A.C., Archibald, S., Levin, S.A., 2011b. The global extent and determinants of savannaand forest as alternative biome states. Science 334, 230232., J.H.M., 2002. Instantaneous canopy photosynthesis: analytical expressionsfor sun and shade leaves based on exponential light decay down the canopy andan acclimated non-rectangular hyperbola for leaf photosynthesis. Annals of Botany89,, 2011. Clima en Barinas. Last accessed Feb 1, 2011.Wang, S.S., Grant, R.F., Verseghy, D.L., Black, T.A., 2001. Modeling plant carbon andnitrogen dynamics of a boreal aspen forest in CLASS the Canadian Land SurfaceScheme. Ecological Modelling 142, 135154.Williams, D.G., Black, R.A., 1994. Drought response of a native and introduced Hawaiiangrass. Oecologia 97, 512519.69J. Segarra et al. / Ecological Informatics 13 (2013) 5769Modelling competitive ability of Neotropical savanna grasses: Simulation of shading and drought impacts on biomass production1. Introduction2. Study site and field methods3. Spatial resolution, time scales, and model steps3.1. Step 1: number of shoots3.2. Step 2: shoots segments by level3.3. Step 3: horizontal distribution of shoot segments3.4. Step 4: leaf area index3.5. Step 5: light attenuation3.6. Step 6: number of shoots segments under shade3.7. Step 7: evapotranspiration rate3.8. Step 8: soil moisture balance3.9. Step 9: biomass3.10. Step 10: mortality by fire4. Drought simulations5. Results6. Discussion7. ConclusionsAcknowledgmentsAppendix AAppendix BAppendix CAppendix DAppendix EReferences


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