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About two thirds of the Earth's land surface is covered by vegetation, characterizing the interface between land and atmosphere interactions. It is well known that natural vegetation is influenced mainly by climate (e.g., Box, 1981; Lugo et al., 1999; Ricklefs, 2008), and ecosystem state is usually a response to gradual, smooth and continuous climate change, but may change abruptly as climate conditions approach a critical threshold, irreversibly switching to a contrasting state (Scheffer et al., 2001, 2009). This latter situation has been observed over history (e.g., the collapse of vegetation in the Sahara around 5000 years ago) (Lenton et al., 2008), and currently the threat of forest dieback in Amazonia (Nobre and Borma, 2009; Davidson et al., 2012) and boreal regions (Lenton et al., 2008) over the next 100 years is of great concern. Dramatic changes in ecosystem state may in turn reinforce the trend of climate change (Lenton, 2011) through regulating the energy, water, and carbon exchanges between land and atmosphere.
As part of recognizing the important roles of the terrestrial ecosystem in global climate and environment change studies, various Dynamic Global Vegetation Models(DGVMs) have been developed in the last 20 years (Foley et al., 1996; Friend et al., 1997; Potter and Klooster, 1999; Woodward et al., 2000; Cox, 2001; Moorcroft et al., 2001; Sitch et al., 2003; Levis et al., 2004; Sato et al., 2007), and become one of the key components of dynamic Earth System Models (ESMs). DGVMs mostly apply mechanistic parameterizations of large-scale vegetation processes (i.e., individual and population dynamics, disturbance) (Prentice et al., 2007) to simulate the distribution and structure of global vegetation, as well as its evolution, alongside global climate change. They provide surface information on large-scale vegetation states, as well as carbon emissions released by soil heterotrophic respiration and wild fires, required by land and atmospheric models.
List of PFTs and their bioclimatic constraints of survival and establishment. #cod#160; Survival Establishment PFT T c,min (#cod#x000b0;C) T c,max (#cod#x000b0;C) GDD min Trees #cod#160; #cod#160; #cod#160; 1.5emBroadleaf Evergreen Tropical 15.5 - 0 1.5emBroadleaf Deciduous Tropical 15.5 - 0 1.5emBroadleaf Evergreen Temperate 3.0 18.8 1200 1.5emNeedleleaf Evergreen Temperate -2.0 22.0 900 1.5emBroadleaf Deciduous Temperate -17.0 15.5 1200 1.5emNeedleleaf Evergreen Boreal -32.5 -2.0 600 1.5emBroadleaf Deciduous Boreal - -2.0 350 Shrubs 1.5emBroadleaf Deciduous Temperate -17.0 - 1200 1.5emBroadleaf Deciduous Boreal - -2.0 350 Grasses 1.5emC4 15.5 - 0 1.5emC3 Non-arctic -17.0 15.5 0 1.5emC3 Arctic - -17.0 0 IAP-DGVM is a DGVM developed at the Institute of Atmospheric Physics, Chinese Academy of Sciences. Its purpose is to address the following disadvantages of current DGVMs: (1) Although DGVMs are capable of capturing the major regions of trees and grasses, their performances in semiarid regions are usually less accurate and several DGVMs replace shrub with grass and bare soil (e.g., Sitch et al., 2003; Levis et al., 2004); (2) The ecological processes of population dynamics (e.g., establishment) in many DGVMs are oversimplified, leading to various inconsistent behaviors (Song, 2012; Song and Zeng, 2014); (3) Regardless of the differences in the complexities of fire parameterization schemes used in DGVMs, large biases exist in the simulation of global fire burned area and carbon emissions (Li et al., 2012a).
The paper is organized as follows. The structure of the model, including the biogeography and population dynamics, individual dynamics, biogeophysics and biogeochemistry, and disturbance (fire), is presented in detail in section 2. A brief demonstration of the model's performance is reported in section 3. And finally, a summary and further discussion is provided in section 4.
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IAP-DGVM adopts concepts from the Lund-Potsdam-Jena (LPJ) Dynamic Global Vegetation Model (Sitch et al., 2003) and the Community Land Model's Dynamic Global Vegetation Model (CLM-DGVM) (Levis et al., 2004), but also incorporates our recent developments of major model components such as the shrub sub-model (Zeng et al., 2008; Zeng, 2010), establishment and competition parameterization schemes (Song, 2012; Song and Zeng, 2014), a process-based fire parameterization of intermediate complexity (Li et al., 2012a, 2012b), as well as some other modifications.
In DGVMs, plants are usually classified into plant functional types (PFTs) according to their physical, phylogenetic and phenological characteristics. IAP-DGVM uses 12 PFTs (Table 1), including seven tree, two shrub, and three grass types, plus bare soil. The model applies the mean field theory in representing the local ecosystem structure. While all PFTs are allowed to coexist in a grid cell (or vegetated land area), each PFT is assumed to have homogeneous spatial distribution, and is represented by a group of so-called average individuals, i.e., the differences among individuals within each PFT are neglected. Each woody PFT (tree or shrub) is assigned a set of prognostic variables of the population and individual states, with the former including population density of woody vegetation (i.e., number of individuals per unit land area), P, and the latter mainly consisting of the carbon pools of leaf, root, sapwood and heartwood per individual, i.e., C l, C r, C sw, and C hw, respectively (units: g carbon per individual). Because it is difficult to distinguish the individuals of grass in the model, grass PFTs are always virtually assigned a population density of P=1, and the units of C l and C r are g carbon per unit area of grid cell (grass PFTs do not have carbon pools of sapwood and heartwood).
While the state variables are updated by various governing equations, the ecosystem large-scale properties, such as PFT fractional coverage (FC) and leaf area index (LAI), which are inferred from the state variables, are also provided in order to facilitate coupling with other components in the ESM. To be consistent with most Land Surface Models (LSMs), which do not allow the overlap of vegetation canopies, the total FCs of all PFTs in a grid cell cannot exceed 100% (see section 2.2.3, below), and the area not covered by vegetation is set to be bare soil.
According to the spatial and temporal scales of the involved ecological processes, IAP-DGVM can be categorized into four sub-modules (Fig. 1).
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2.2.1 Biogeography
The biogeography, describing the climatic constraints of survival and establishment (see Table 1), is similar to CLM-DGVM. PFT survival in a grid cell requires the long-term average (e.g., 20-yr running mean) of the minimum monthly temperature, T c, to exceed T c,min. Existing PFTs cease to exist if they cannot survive. Besides, PFTs will also be removed if their annual net primary production (NPP) is negative.
In addition to the above criteria, establishment also requires T c to be less than T c,max, GDD5 to be greater than GDD min, GDD23 to be equal to 0, and annual precipitation to be greater than 100 mm yr-1, where GDD5 and GDD23 are the long-term averages of the annual growing degree days above 5#cod#x000b0;C and 23#cod#x000b0;C respectively, and T c,min, T c,max, and GDD min are PFT-dependent constants.
The dynamics of population density of the ith woody PFT, Pi, is determined by
where P(t) and P(t+1) are the population densities at the current and next time step; #cod#916; P is the change in population density; #cod#916; P est is the population increase due to establishment; and #cod#916; P m is population decrease due to mortalities, including #cod#916; P light, #cod#916; P sg, #cod#916; P sh, and #cod#916; P fire, which are caused by light competition, growth stress, heat stress, and fire, respectively. Except for #cod#916; P fire, which will be described in section 2.5, the other terms are summarized as follows.
2.2.2 Establishment
In DGVMs, establishment of woody vegetation refers to a sequence of reproduction and regeneration processes from flowering, fertilization, seed production, germination, and finally growth of saplings. IAP-DGVM applies a hierarchy of tree-grass-shrub for the competition of establishment. Trees can establish in areas currently not occupied by perennial vegetation, i.e., trees and shrubs. The establishment rate of a given tree PFT is calculated once a year as:
where #cod#916; P est,tree is the total establishment rate of tree PFTs in the grid cell; n est,tree is the number of tree PFTs established in the current time-step; #cod#916; P estmax,tree is a constant representing the maximum establishment rate of trees; FC w is the total coverage of trees and shrubs in the grid cell; the term (1- FC w) is the area currently not covered by trees, and so available for establishment; the term 1-e-5(1- FC_ w) refers to the shading effects (Sitch et al., 2003); gi represents the competition of establishment among different woody PFTs, including the establishment from seeds produced in the current year, which is assumed to be dependent on PFT FC, FCi, with an exponential factor, #cod#963; as well as background establishment #cod#949; (i.e., from seeds produced in previous years or propagated from nearby grid cells); gi0 is the PFT-dependent constant of relative establishment potential which represents the competition-colonization trade-off among different PFTs (Song, 2012; Song and Zeng, 2014); and mi is the mortality rate of seedlings, inferred from the individual background mortality, m ge, and mortality due to heat stress, m heat [Eqs. (7) and (8)]
Grasses and shrubs can establish only in bare soil. IAP-DGVM does not consider the shading effects on grass establishment, and young grass seedlings are added over all bare areas in proportion to the current FC of different grass PFTs, i.e.,
where FC veg is total vegetation coverage of the grid cell; n est,grass is the number of grass PFTs established in the current time-step; and gi, gi0, and #cod#949;0 are the same as in tree establishment.
Establishment of shrub PFTs follows the same rule as trees, i.e., Eq. (2), but replace the term #cod#916; P estmax,tree with #cod#916; P estmax,shrub, and FC w with FC veg.
2.2.3 Light competition
In IAP-DGVM, light competition occurs if the total FCs of specific vegetation categories exceed corresponding thresholds. The model applies a hierarchy of tree-shrub-grass for the light competition to represent the advantage of higher vegetation in capturing incoming solar radiation. First, if the total tree coverage in a grid cell, FC tree, is larger than a threshold (95% in the model), the excess is removed from existing tree PFTs in proportion to their FCs through the decrease of population densities (Sitch et al., 2003). Thus, the mortality due to light competition is calculated as
Shrubs are usually shorter than trees but taller than grasses; hence, shrubs can occupy the area not covered by trees. Because there are only two shrub PFTs and they cannot coexist in the same grid cell, the excess cover when total tree and shrub coverage, FC w, is larger than 100% is removed by the only existing shrub in the grid cell, i.e.,
Finally, grasses may occur only in areas not covered by woody vegetation. The excess cover is removed by directly reducing the FC of grass PFTs:
2.2.4 Mortality due to growth stress
It has been observed that individuals with relatively lower growth rates usually have higher mortality rates (Bugmann, 2001; Keane et al., 2001). Following (Prentice et al., 1993), the related mortality rate, m ge,i, is inversely related to the growth efficiency (gei), which is defined as the ratio of net biomass increment per individual, #cod#916; Ci, to individual leaf area:
where SLA is the specific leaf area (leaf area per unit leaf mass); PFT-dependent k m1 is the corresponding maximum mortality rate, i.e., the mortality rate as ge=0; and k m2 is a constant.
2.2.5 Mortality due to heat stress
It has been noticed that high temperatures can cause tissue damage to temperate and boreal vegetation, and even result in losses of individuals. The related mortality rate is calculated as
where GDD23 is the averaged annual growing degree days above 23#cod#x000b0;C (see section 2.2.1).
2.2.6 Changes in average individual carbon pools
Because IAP-DGVM does not consider the dynamics of seedling and mature individuals separately, the individual carbon pools are updated by averaging the remaining individuals and the newly established saplings. For woody PFTs, it is
where subscript "tis" stands for leaf, root, sapwood, or heartwood; and C sap are the pre-described sapling carbon pools.
For grass,
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The processes of individuals include the allocation of NPP to different tissues, tissue turnover, and the seasonal variation of leaf area, i.e., leaf phenology.
2.3.1 Allocation and plant morphology
Annual NPP, i.e., photosynthesis minus plant respiration (see section 2.4), is allocated to reproduction (for establishment) and different tissue carbon pools (for individual growth). While the fraction of reproduction costs could be PFT-dependent and be related to the relative establishment potential, gi0 [see Eq. (2)] it is set to a fixed 10% with gi0 #cod#8801; 1 for simplification in the current model.
The allocation to leaves, sapwood, and roots of woody vegetation is calculated so that the individual carbon pools follow the allometric relationships modified from Sitch et al., (2003) and (Levis et al., 2004):
where a l and a s are the individual leaf area and sapwood cross-sectional area, respectively; H, D, and a C stand for height, stem diameter and crown area; and the parameters k ls, k lr, ka1, ka2, ka3, k rp, and a c,max are PFT-dependent (see Table 2).
Comparing the second equation of Eq. (11) with Eq. (2) of Sitch et al. (2003) and Eq. (23) of (Levis et al., 2004), a factor representing the degree of water stress, #cod#969;, is removed from the right-hand side. The original purpose of introducing #cod#969; was to capture the feature that plants may have higher root-to-leaf ratios as water stress increases. However, the capacity of roots to absorb soil water for transpiration, and hence the strength of plant photosynthesis, is not related with total root biomass in most current LSMs, and the strategy of increased allocation to roots along with decreased allocation to leaves will result in higher root respiration and lower photosynthesis capacity, i.e., making the situation for plants under water stress undesirable. Hence, #cod#969; has been temporally dropped before further development in LSMs.
Parameters for woody plant morphology. #cod#160; k la:sa k allom1 k allom2 #cod#945; cmax (m2) Broadleaf trees 8000 100 40 15 Needleleaf trees 8000 82 40 15 Shrubs 4000 200 10 5 Besides, the ka1 of temperate and boreal needle-leaf evergreen trees is set to be 82, compared with 100 for other tree PFTs, to reflect the morphological feature of a relatively smaller but denser canopy. The parameters for two shrub PFTs are also defined and different to tree PFTs following (Zeng, 2010).
On the other hand, allocation for grass PFTs is simpler in that only the second equation in Eq. (11) is needed, because grass does not have the carbon pools for sapwood and heartwood.
2.3.2 Turnover
Each year, plants may lose part or all of their leaves and roots, and part of the sapwood of woody PFTs may turn into heartwood, due to turnover. The decrease in the corresponding tissue carbon pool is calculated as
where f tis,i are PFT-dependent tissue turnover times (yr-1).
2.3.3 Leaf phenology
While the annual maximum LAI, LAImax, is determined by the individual leaf mass and crown area (see section 2.6), daily LAI, LAI daily, is a fraction of LAImax and may have seasonal variation, calculated as
where #cod#966; represents leaf phenology and has a value between 0 and 1.
Similar to (Levis et al., 2004), tree phenology may be evergreen, summergreen, or raingreen: (1) For evergreen trees, #cod#966;#cod#8801; 1, i.e., there is no seasonal variation; (2) For summergreen trees, leaves emerge evenly within 50 days after the day with GDD5#cod#62;100, and drop at a rate of 1/15 d-1 when the 10-day running mean of surface air temperature, T 10d, is lower than the long-term average of the minimum monthly temperature; (3) For rain-green trees, leaves emerge or drop at a rate of 1/15 d-1 as the 10-day running mean of photosynthesis, A 10d, is larger or smaller than leaf maintenance respiration, R 10d,leaf. However, #cod#966; retains a minimum value, e.g., 0.1, to permit non-zero photosynthesis.
For the shrub PFTs, broadleaf deciduous temperate shrub is assigned as rain-green, and broadleaf deciduous boreal shrub as summer-green, for the tree phenology (Zeng, 2010).
Grass PFTs have no predetermined phenology, but respond to a blend of summer-green and rain-green (Levis et al., 2004). Leaves emerge at a rate of 1/5 d-1 if both criteria of T 10d#cod#62;0#cod#x000b0;C and A10 d#cod#62;R 10d,leaf are satisfied, and otherwise drop at the same rate.
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The microscale processes involved can be separated into two parts: (1) biogeochemical processes, which include photosynthesis, autotrophic respiration, litter decomposition and heterotrophic respiration; and (2) biogeophysical processes, which mainly include energy and water exchange between vegetation and land, and the surface and atmosphere. Because most of the related processes, especially photosynthesis and biogeophysical processes, are essential components of LSMs, we only briefly describe here the calculations related to NPP, which is needed by allocation, as well as accumulation and decomposition of aboveground litter, which may influence fire simulation.
2.4.1 Net primary production
Plants gain carbon through photosynthesis and lose carbon through autotrophic respiration, which can usually be separated into two parts, i.e., growth respiration and maintenance respiration. Maintenance respiration refers to metabolism occurring in an organism that is needed to maintain that organism in a healthy, living state. Following LPJ and CLM-DGVM, maintenance respiration amounts from leaves, roots, and sapwood (per unit area of PFT cover) are calculated respectively as
where r is a PFT-dependent coefficient in grams of carbon per gram of nitrogen (gC g N-1); cn l, cn r, and cn sw are tissue C:N ratios; T l, T r, and T sw are tissue temperature; and #cod#966; is the leaf phenology (section 2.3).
Plants are able to grow only when annual photosynthesis is greater than the total maintenance respiration,
Part of the net carbon gain between photosynthesis and respiration is consumed as growth respiration, i.e.,
where A is photosynthesis (per unit area of PFT cover), and k g is a constant assumed to be 25%.
Then, the annual NPP available for allocation is
2.4.2 Accumulation and decomposition of aboveground litter
Plants can lose tissue due to individual mortality [Eqs. (4)-(10)], turnover [Eq. (12)], and fire [Eqs. (23)-(25)]. The corresponding leaves, sapwood and heartwood are added to the carbon pool of aboveground litter.
On the other hand, part of the aboveground litter is burned by fire, and the remaining part will decompose following first-order kinetics,
where k dec is the decomposition rate.
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Fire is the primary terrestrial ecosystem disturbance agent on a global scale, and shapes not only ecosystem composition but also vegetation succession and regeneration (Bowman et al., 2009). IAP-DGVM adopts a process-based fire parameterization of intermediate complexity developed by Li et al. (2012a, 2012b). Methodologically, it has better structure, parameter estimation, and mathematical derivation than other process-based fire models of intermediate complexity (e.g., Glob-FIRM, CTEM-FIRE, and their modified versions). The fire model comprises three parts, i.e., fire occurrence, fire spread, and fire impact. In this paper, only the basic function and parameterization regarding the impact of fire on vegetation is briefly described; the detail of the model can be found in Li et al. (2012a, 2012b).
In the fire model, the burned area is determined by climate and weather conditions, vegetation composition and structure, and human activities. The basic function of the fire model is as follows:
Ab= Nfa, (21)
where A b is the burned area in a grid cell per time step [km2 (time step)-1]; N f [count (time step)-1] is fire counts in the grid cell, and is a function of the number of ignition sources due to natural causes and human activities, availability and combustibility of fuel (i.e., aboveground biomass of leaves, stems and aboveground litter combined), and fire suppression by human activities [see Eqs. (2)-(10) of (Li et al., 2012a)]; and a (km2) is the average spread area of a fire, and depends on wind speed [see Eqs. (11)-(18) of (Li et al., 2012a)]. We define k fire as the percentage of burned area in the grid cell, i.e., the ratio of burned area, A b, to the area of the grid cell, A g (km2):
Within the burned area, a proportion of tissue (per individual) is burned (and becomes fire emission), given by
where L tis is the PFT-dependent combustion completeness factor for different tissues.
Besides, a proportion of woody individuals are directly killed by fire. The individual mortalities for woody PFTs due to fire, #cod#916; P fire,i, is calculated as
where \xi is the PFT-dependent whole-plant mortality factor. On the other hand, survival would lead to loss of part of the uncombusted tissue, and would transfer to litter. The tissue mortalities are:
where M tis is the PFT-dependent tissue-mortality factor.
Finally, the grid cell average individual tissue carbon after fire is updated as
where the denominator gives the percentage of individuals that survive after the fire (within the whole grid cell); the first term of the numerator is the percentage of individuals outside the fire burn-area (whose carbon pools remain unchanged); and the second term accounts for the proportion of tissue remaining, belonging to individuals that have survived, inside the burn-area, and the term C tis,i on the right hand side is the individual tissue carbon before fire.
To offer an interface with atmospheric chemistry and aerosol models in ESMs, the fire model also estimates trace gas and aerosol emissions, Ex,i (g species per unit area of grid cell), following (Andreae and Merlet, 2001) as
where x is the index of trace gas and aerosol; Fx,i [g species (kg dm)-1] is the PFT-dependent emission factor; and [C]=450 gC (kg dm)-1 is a conversion factor from tissue dry matter to carbon.
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Important ecosystem large-scale properties, e.g., LAI and FC, are updated depending on the related state variables.
2.6.1 Leaf area index
Leaf area index of a specific PFT is defined as total leaf area per unit land area covered by the PFT. For woody PFTs, it is calculated as
where C l,i and aC,i are the individual crown area and leaf carbon, respectively.
Because grass has no pre-defined morphology, and is assumed to always have a population density of one individual per area of grid cell, we virtually define its crown area, following some empirical approximation, as
so that the LAI of grass can be calculated using Eq. (28) and is similar to the definition in other DGVMs.
2.6.2 Fractional coverage
To be consistent with the definition commonly used in LSMs, in IAP-DGVM the fractional coverage of a specific PFT is the percentage of grid cell area that is covered by the crown area of that PFT, calculated as
FCi= aC,iPi. (30)
For woody PFTs, CA is calculated in allocation (section 2.3), and P is updated in the population dynamics part. For grass PFTs, CA is calculated using Eq. (29) (section 2.6.1), and P #cod#8801; 1. Note that this is different to the concept of fractional projective cover adopted in some DGVMs, e.g., LPJ [Eq. (8) of Sitch et al. (2003)] and CLM-DGVM [Eq. (28) of (Levis et al., 2004)].