1 Supporting Information Notes S1, Figures S1-S3 2 3 Notes S1 4 1. Model description 5 The climate-vegetation-natural fire (CVNF) model is composed of a simple set of 6 modified Lotka-Volterra competition equations: dg g a g g 1 * c ga * g d g firg g , dt g a IV I V III II (1) da a a a1 * d a fira a, dt a I V II (2) 7 to represent the time evolution of the fractional coverage area of grasses (g) and trees 8 (a). 9 Basically, two aspects differ from the classical set of Lotka-Volterra competition 10 equations: 1) plant growth parameters (a, g) depend on environmental conditions 11 instead of being defined as constants (Nordstrom et al., 2005); 2) additional sink terms 12 due to lightning-triggered fire effects on vegetation (dgfirgg, dafiraa) are included. 13 The first term (I) is a source term which is driven by the plant growth parameters 14 defined as: i fi i , i g , a, (3) 15 where i is the time-scale of establishment of each vegetation type, and fi is a climate- 16 dependent function. i is a constant obtained from an equation proposed by Hughes et 17 al. (2006) (from the TRIFFID model equations (Cox, 2001)) to determine the time scale 18 for vegetation establishment considering fractional coverage areas and not only 19 individual elements of a vegetation population. The expression to calculate 20 described as: V f1 1 i i ln 1 , V 1 s is (4) 21 where Ti is the time of establishment of one element of the population i; Vf and Vs are 22 the final and initial fractional coverage of population i, respectively. Typical growth 23 length for individual tropical forest trees is about 50 years to reach a mature phase, i.e., 24 Ta = 50 years (Chambers et al., 1998). On the other hand, grasses take a much shorter 25 time to completely develop themselves from seed to adult phases. Although field 26 measurements are not conclusive about seed-to-adult growth length of grasses within 27 South American tropical savannas, only the aerial portions of typical herbaceous species 28 of Brazilian savannas take about 2 years to recover after a fire event (Batmanian & 29 Haridasan, 1985). Thus, we assume that the complete seed-to-adult cycle would take at 30 least doubled growth time (Martins et al., 1999), and set Tg ~ 4 years. Then, using 31 equation (4), tree and grass populations would take about 340 and 27 years (a = 340 32 and g = 27) respectively to grow from Vs = 0.01 (almost null) to Vf = 0.9. We do not 33 include different periods for vegetation recovery after environmental disturbances, 34 especially fires. This would be more realistic since that grass vegetation would take a 35 shorter time to regrow as only its aerial portions would be damaged (root chains would 36 be intact). Besides, evergreen trees considered in the model are not as resilient as 37 grasses, and could either be partially damaged and followed by secondary-forest 38 succession, or completely die off and need the same seed-to-adult cycle period to 39 regrow. 40 Functions 41 2005): are modeled as simple piecewise linear functions (based on Arora & Boer, HI HI l ,i f i max min HI u ,i HI l ,i ,1,0 (5) 42 where HI is a humidity index and HIu,i (HIl,i) is the upper (lower) bound above (below) 43 which fi is equal to 1 (zero). The humidity index (Hulme et al., 1992), HI, is derived 44 from the aridity or dryness index, AI (Budyko, 1974), and is defined as a combination of 45 monthly precipitation (P) and temperature (T) values by the equation: HI 1 P AI PET (6) 46 where PET = PET(T) is the potential evapotranspiration given by Thornthwaite’s 47 equation (Thornthwaite, 1948). HI is computed on a monthly basis from precipitation 48 and temperature datasets. Therefore, plant growth parameters (i, i = g, a) are 49 larger/smaller under wetter/drier humidity conditions. 50 Grasses (g) and trees (a) grow depending on different values of HIl,i and HIu,i. These 51 values are defined based on the broad-sense classification of climate regimes suggested 52 by Ponce et al. (2000). For the purposes of this work, we focus on the humid climate 53 delimitation 0.375 ≤ AI < 0.75, which we assume to be related to the development of 54 tropical forest vegetation. A simple algebra turns the above expression into 1.3 ≤ HI < 55 2.6. Therefore, using rounded values, we define HIl,a = 1.0 as the starting point for tree 56 growth, and HIu,a = 3.0 the value from which climate conditions are not yet limiting 57 factors for tree development (Fig. S1a). On the other hand, thresholds for grasses are 58 empirically set based on tree values. As they need much less favorable soil moisture 59 conditions to develop in comparison to trees, they start growing from HIl,g = 0.1, and are 60 no longer controlled by HI when HIu,g ≥ 1.1, when grasses grow freely only restricted by 61 competition factors (represented by the competition coefficient, cga, of equation (1), 62 which will be discussed later) (Fig. S1a). 63 Synthetic time series of monthly precipitation and temperature are used. Precipitation 64 values (P) are generated (based on D’Andrea et al., 2006; Dolman et al., unpublished) 65 using the following equation: 2t P, T (t ) max P ,T P ,T AP ,T cos P ,T ,0. (7) 66 The first term on the RHS of equation (7), P,T, represents long-term mean annual 67 precipitation/temperature; the second term, P,T, represents a prescribed random 68 variation from the climatological values and is composed by a random number () with 69 a normal standard distribution (~N(0,1)) and the standard deviation of the mean annual 70 value (P,T); the third term represents the annual cycle (), is the period of 12 months; 71 AP,T is the amplitude; and P,T is the phase), in which precipitation seasonality is 72 embedded and thus the variation of the dry season length. AP,T = P,T – min(P,T) is 73 defined using the minimum value from monthly precipitation/temperature climatology; 74 and P,T = 2 m(P,T),max/ takes into account month m when the maximum value from 75 monthly precipitation/temperature climatology occurs. Precipitation climatology is 76 evaluated by the Climate Research Unit’s (CRU) gridded (0.5o x 0.5o) precipitation time 77 series (1901-2002) (New et al., 1999; Mitchell et al., 2003). Fig. S2a shows the annual 78 cycle of the generated precipitation in the middle of the simulation period (around the 79 500th year). Precipitation gradients in the wet season and longer dry seasons are clearly 80 seen comparing precipitation distribution for closed forest (thicker lines) and savannas 81 (thinner lines). The precipitation regime in the 66W longitude is somewhat different 82 from the other three locations with the dry season taking place earlier in the year. 83 Temperature values (T) are generated analogously. For instance, in the middle of the 84 same simulation period (around the 500th year), temperature decreases reach up to 2oC 85 from 66W to 46W and there is a slight change in the annual cycle of the temperature 86 within the savanna location (44W) (Fig. S2b). 87 The formulation described above may be suitable to simulate future climate scenarios 88 by modifying the mean annual precipitation/temperature (P,T), the inter-annual 89 variability (P,T), or even the amplitude or the peaks of the annual cycle (AP,T or P,T). 90 Besides birth representation, sink terms II and III of intra-specific (quadratic terms, 91 (ggg) / g* or (aaa) / a*) and inter-specific (crossed terms, (gcgaga) / a*) 92 competitions are also driven by the plant growth parameter (i, i = g, a) (equations (1) 93 and (2)). Constants g* and a* correspond respectively to the carrying capacity of grasses 94 and trees, i.e., the maximum value that the vegetation population may reach. As usual, 95 both constant values are set to 1 for general purposes. The crossed term should represent 96 ecological advantages from one vegetation type over another depending on magnitude 97 cga. This parameter in term III represents in equation (1) the negative shading effect that 98 favors tree growth and development instead of grasses within an area (Cox, 2001). 99 Because there are no general advantages of grasses over trees, inter-specific competition 100 (term III) is negligible (cga ~ 0) in equation (2). 101 Term IV in equation (1) is the natural grass mortality, and is assumed to occur at a 102 constant rate = 0.1 (Hughes et al., 2006). On the other hand, according to 14C isotope 103 measurements, trees in central portions of the Amazon forest may be very old, with ages 104 ranging from 200 to 1,400 years depending on tree species (Chambers et al., 1998). 105 Assume, for simplicity, an average age of 800 years. As it is relatively close to the total 106 simulation period, natural tree mortality is not computed in the CVNF model. 107 As fire mortality is computed in a separate term (term V, see below), only climate could 108 disturb tree coverage and promote death of younger trees in our simplified scheme. Tree 109 type, however, is associated to a generic evergreen-forest species, and thus it is very 110 resistant to climate seasonality due to its deep root zone (Nepstad et al., 2004). Note 111 that because natural tree death is not computed in this model, this term does not 112 influence tree population dynamics (term IV is null in equation (2)). 113 The vegetation sink term due to fire occurrence depends on the maximum population 114 death resulting from fire, di, and on fire effects firi ,i = g, a. This means that even with 115 extreme fire intensities, only a maximum fraction di of population i shall die. Constant 116 parameter di is set as 0.3 for grasses (Gardner, 2006) and 0.1 for trees as proposed by 117 Balch et al. (2008) for the woody biomass burnt within a forest-savanna transition in 118 southeastern Amazon. 119 Fire effect (firi ,i = g, a) is an output of a simplified fire sub-model divided into 3 stages: 120 triggering, intensity and effects. It is far beyond the purpose of this model to simulate 121 fire propagation, displacement and other ecological features and impacts of fire. Instead, 122 we looked for a simplified representation of the main characteristics of fire ecology. 123 124 1.1. The fire sub-model 125 This section presents the main contributions of this work, concerning the 126 parameterization of lightning-triggered fires and respective effects on grass and tree 127 vegetation that will affect the dynamics of the system (1)-(2). 128 Natural fires depend on three major conditions to start: fuel availability and 129 flammability, and the ignition source (Whelan, 1995). In this context, the first stage is 130 associated to favorable conditions of fire triggering taking into account the three aspects 131 mentioned above. Ecologically, fuel availability is mostly due to the herbaceous layer 132 (Bond, 2008). Therefore, fuel is accumulated when grasses die (at constant rate = 0.1 133 from equation (1)) and form litter pools (L). Whenever L reaches the threshold value, 134 Lmin = 0.45, the availability condition is satisfied and fire may start. Flammability is 135 defined based on a conceptual assumption that litter pools may be split into 2 vertical 136 layers. This two-layered litter model in which the upper (bottom) layer is highly 137 influenced by air humidity (soil moisture) is inspired by controlled fire field 138 experiments in the Amazon (Balch et al., 2008). In order to start a fire, only the top 139 layer needs to be sufficiently dry; therefore, the flammability threshold is set according 140 to the humidity index (HI) of equation (6). Whenever HI ≤ HImax = 1.2, fire may start. 141 Both optimum thresholds (Lmin, HImax) are set empirically through computational 142 experiments with the CVFN model. 143 Ignition is caused by lightning flashes. To represent the occurrence of these flashes, we 144 propose the monthly lightning (R) time series generation, similarly to the procedure 145 outlined for precipitation and temperature in equation (7): 2t R(t ) 0.25 max R R AR cos R ,0. (8) 146 Therefore, a random component is explicitly incorporated in the lightning time series 147 generation to represent the highly stochastic nature of lightning strikes. Statistics (R, 148 R, mR,max, AR, R) used for the time series generation is extracted from a combined 7- 149 year (1995-2002) dataset of two lightning-detection satellite sensors (LIS/OTD) from 150 the Global Hydrology Resource Center - NASA (http://thunder.nsstc.nasa.gov). 151 This dataset includes the entire lightning activity over the tropics, in either cloud-to- 152 cloud or cloud-to-ground flashes. As only cloud-to-ground flashes can initiate fires, we 153 use a global value of 25% (0.25) to represent the lightning activity of interest (Pride & 154 Rind, 1993). Fig. S2c shows the annual cycle of the lightning activity generated by this 155 methodology. It is remarkable that there is more lightning activity over closed forests 156 than over savannas for most part of the year. However, forests do not frequently burn 157 because even if fire is triggered, the high soil moisture limits its intensity (see equation 158 (10)). 159 Moreover, not all cloud-to-ground flashes initiate a fire. There are few estimates of the 160 relation between flashes and fire initiation in boreal forests (for instance, see Larjavaara 161 et al., 2005, for Finland). An estimate for southeast Brazil is proposed based on 162 observational data from the Brazilian Lightning Detection Network – BrasilDat (Pinto 163 Jr. et al., 2006). As the negative cloud-to-ground flashes with long continuing currents 164 (Saba et al., 2006) cause the most serious damage associated to thermal effects, i.e. fires 165 among others, Correia & Saba (2008) measure the presence of that current type in 166 several thunderstorms. Authors suggest that about 28% of the negative flashes have 167 long continuing currents and may promote fire episodes. In addition, this estimate can 168 be extended to neighboring areas of Brazil without loss of generality. In general, 90% of 169 the total could-to-ground lightning flashes are negative (Saba et al., 2006). The 170 combination of both percentages (28% and 90%) results in an estimate of 25%. 171 Based on Arora & Boer (2005), we suggest a probability function of fire ( 172 the equation (5), expressed as: R Rl PR max min ,1,0 , R R u l ), similar to (9) 173 depending on the number of monthly flashes (R) and empirical lower and upper bounds 174 defined as Rl = 25 and Ru = 65. Although empirically defined, these bound values are 175 inspired on the assessment of 25% described above (Correia & Saba, 2008). Let Rf be 176 the number of lightning flashes that could cause fire ignition, such that Rf = 0.25R. 177 Applying this relation to the Rl and Ru values results in approximately Rf = 6 and Rf = 178 16. Thus, in a very simplified scheme, probability PR is assumed to be practically null 179 for less than 6 flashes, as this would mean 1 flash that could ignite a fire for every 5 180 days. Then, PR is assumed to enhance linearly as Rf (R) increases up to 16 (65), above 181 which PR = 1 (Fig. S1b). Moreover, R ≥ 65 or Rf = 16 means that at least 1 flash for 182 every 2 days in the month may promote fire ignition, which is quite a high prospect. 183 Thus, PR for this case is assumed to be maximum. 184 Ignition is likely to occur if PR ≥ PR,min = 0.55 (Fig. S1b). We take the 46W longitude 185 point in the middle of the simulation period (after approximately 500 years). At this 186 specific location, Fig. S1d shows an example of the exact moment when the three 187 conditions are met in the model. Fire occurs in the transition between the dry and the 188 wet seasons in November, when there is enough dried litter (L ≥ 0.45 and HI ≤ 1.2) on 189 the ground, and the probability of fire ignition is above the threshold value (0.55). 190 After fire starts, fire effects depend on fire intensity and vegetation types. Thus, the 191 second stage is to define a fire-intensity function. In this study, the latter varies 192 according to the amount (L) and the dryness of the litter available to burn, which is 193 associated to a soil moisture index (w). It is defined as a function of the annual mean 194 precipitation value, P in mm month-1: w P w max min u ,1,0, w w u l (10) 195 where wl = 110 mm month-1 and wu = 200 mm month-1 are the empirical lower and 196 upper bounds of precipitation values. These constants limit the point below which w is 197 not a limiting factor, and above which w is null (totally limiting), respectively (Fig. 198 S1c). 199 The fire intensity dependence of litter is modeled after Anderies et al. (2002) according 200 to: LbL h( L, k L , bL ) bL , k L LbL (11) 201 where constants kL and bL are empirically set to 0.7 and 2.0, respectively, and are chosen 202 to represent the increase in fire intensity due to the higher amount of litter (Fig. S3). 203 Fire intensity is thus defined as the combination of two limiting factors: I I (w, L) w h( L, k L , bL ) . (12) 204 As fire effects are different on grasses and trees, we make assumptions to represent 205 these differences (see Fig. S3). Grass vegetation cover burns as soon as fire starts and 206 regrows faster than trees (equation (3)). 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Cambridge University Press, Cambridge, UK. 302 Figs S1–S3 303 304 Fig. S1 – (a) fi, i = g, a as a function of humidity index; continuous (dotted-continuous) 305 line refers to the plant growth function of grasses (trees); (b) fire probability (PR) as a 306 function of lightning number and the respective threshold (0.55); (c) soil moisture index 307 (w) as a function of annual precipitation; (d) illustration of a fire event in longitude 46W 308 after about 500 years of simulation – continuous, dashed and dotted lines refer to the 3 309 fire limiting factors: the humidity index (HI), the probability of fire ignition (PR) and the 310 available litter (L), respectively, with HImin ≤ 1.2, PR,min ≥ 0.55 and Lmin ≥ 0.45. 311 312 Fig. S2 – Annual cycle of the generated (a) precipitation, (b) temperature and (c) 313 lightning flashes for four locations: closed forest with two different annual precipitation 314 regimes (65W and 58W), forest-savanna transition (46W) and savanna core (44W). 315 316 317 Fig. S3 – Variations of function h used in the model, depending on parameters and b. 318 319 Fig. S1 (a) (b) (c) (d) 320 321 322 Fig. S2 (a) (b) (c) 323 324 325 326 Fig. S3
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