Environ Monit Assess (2022) 194:903

https://doi.org/10.1007/s10661-022-10545-3

Forest cover resilience to climate change over India using

the MC2 dynamic vegetation model
Pulakesh Das   · Mukunda Dev Behera   ·
Prasad K. Bhaskaran · Parth Sarathi Roy

Received: 27 May 2022 / Accepted: 10 September 2022

© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2022

Abstract  It is imperative to understand the climate
change impact on the forest ecosystem to develop
appropriate mitigation and management strategies.
We have employed a process-based dynamic vegeta-
tion modeling (MAPSS-CENTURY: MC) approach
to project change in vegetation life forms under pro-
jected climate conditions that attained 81% overall
accuracy. The present and projected climate condi-
tions suggested highly resilient/stable forest covers
in wet climate regimes and moderately resilient in
dry semi-arid regions. Several forested grids in the
seasonally dry tropical forest in the Eastern Ghats and
dry Deccan peninsula regions are estimated to be
less resilient, which may experience a regime shift
toward scrub and grassland. The future prediction
Supplementary Information  The online version
contains supplementary material available at https://​doi.​
org/​10.​1007/​s10661-​022-​10545-3.
P. Das · M. D. Behera (*) 
Centre for Oceans, Rivers, Atmosphere and Land Sciences,
Indian Institute of Technology Kharagpur, West Bengal,
Kharagpur 721302, India
e-mail: mdbehera@coral.iitkgp.ac.in
P. Das · P. S. Roy 
Sustainable Landscapes and Restoration, World Resources
Institute India, New Delhi 110016, India
P. K. Bhaskaran  loss due to deforestation and forest fire can be easily quan-
Ocean Engineering and Naval Architecture Department,
Indian Institute of Technology Kharagpur, West Bengal,
Kharagpur 721302, India
demonstrated an upward temperature shift in the
Western Himalayas and trans-Himalaya, which may
facilitate forest spread at higher elevations. Although
the forest cover resilience may increase in future cli-
mate conditions, the disturbances in several regions in
the Deccan Peninsula and the Eastern Ghats may trig-
ger forest to scrub and grassland transition. The inac-
curacy in model simulation in the Western Himalayas
could be attributed to coarse resolution grids (0.5°)
failing to resolve the narrow climate niches. The spa-
tially explicit model simulation provides opportuni-
ties to develop long-term climate change adaptation
and conservation strategies.
Keywords  Vegetation life form · Climate
projection · Net primary productivity · Leaf area
index · Elevational shift
Introduction
The distribution of potential vegetation depends on the cli-
mate, topography, geology, nutrient availability, and other
environmental conditions. Climate change, via the altera-
tion in precipitation and temperature, and other external
disturbances such as anthropogenic activities, fire occur-
rences, insect outbreaks, etc., redefines the vegetation
conditions and regulates distribution. While tree cover
tified, the effects of climate change are difficult to meas-
ure. Ecosystem conservation and management practices

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could fail without perceiving the climate change impact.
Studies have exemplified the climate change impact on
biodiversity and ecosystem services (Cristal et al., 2019;
Staudinger et al., 2012). Although environmental condi-
tions and disturbances are regarded as the dominant fac-
tors in regulating vegetation distribution, there are high
uncertainties in their degree of influence. The impact of
the same climate alteration may differ across the climate
regimes depending on the feedback from the local envi-
ronmental settings. Ecosystem resilience describes the
recovery of an ecosystem structure and functionality from
an undergone perturbation due to external disturbances.
The long-term environmental conditions and external
disturbances determine the ecosystem resilience, resist-
ance to a regime shift, and recovery period. The higher
rooting depth of plants allows access to water at a greater
depth and sustains them during dry periods or drought
spells, wherein soil texture determines the water holding
capacity and seasonal availability (Davidson et al., 2012;
Noy-Meir, 1973). Water, energy, and nutrient availability
govern vegetation sustainability and recovery from distur-
bances (Fernandez-Illescas et al., 2001). Verbesselt et al.
(2016), Fang and Zhang (2019) studied the ecosystem
resilience in tropical forest in response to drought and
heat events. They reported a lower ecosystem recovery
rate in drier regimes than in wetter ones.
Sharma and Goyal (2018) studied the ecosystem resil-
ience in India by assessing the hydroclimatic conditions
using net primary productivity (NPP) and evapotran-
spiration as indicators. They reported that most forests
are not resilient on a river basin scale under dry condi-
tions. Das and Behera (2019) and Behera et  al. (2018)
employed the tree canopy cover percentage (TCC%) to
estimate the precipitation-dependent forest cover resil-
ience in India using a logistic regression-based approach.
They reported that forest cover is highly resilient above
1400  mm of precipitation. However, such empirical
models provide a uniform impact of climate stressors on
vegetation, where the effects of multiple factors are very
complex to estimate. The vegetation models simulate
various environmental conditions to project forest cover
alterations and transitions. The bioclimatic species dis-
tribution models such as MAXENT (maximum entropy)
and GARP (Genetic Algorithm for Rule-set Production)
use existing species distribution and statistically relate
them to environmental conditions, which are further used
for the forecast with projected climate scenarios (Chitale
et al., 2014; Ray et al., 2018). Anjos and de Toledo (2018)
prescribed the climate niche-based modeling approach to
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Environ Monit Assess (2022) 194:903
study ecosystem resilience on a large scale. They adopted
an ensemble modeling approach integrating multi-model
outcomes and reported that forests are less resistant to
climate change and more vulnerable than savanna and
grasslands, indicating lower adaptability of forests owing
to narrower climate niches. The biogeography models,
e.g., MAPSS (Mapped Atmosphere-Plant-Soil System),
project the potential vegetation types (including life form,
leaf form, phenology, vegetation density, etc.) based on
the observed climate conditions but have limitations in
incorporating the nutrient cycle (Neilson, 1995). On the
other hand, the biochemistry models, e.g., CENTURY,
simulate the effect of water, energy flow, and nutrient
cycle on ecological processes but does not consider the
climate-induced vegetation shift and instead rely on sta-
ble vegetation cover distribution (Cronan, 2018). The
biogeography and biogeochemistry models rely on steady
distributions of vegetation cover and therefore have limi-
tations in simulating any change in plant distribution
despite being used to explore the effects of climate change
on vegetation dynamics (Cramer et  al., 2001). In com-
parison, the process-based dynamic vegetation models
include biogeography and biogeochemistry modules that
use the information on plant functioning, gradation of cli-
mate, edaphic, topographic, energy, nutrient, etc., and bet-
ter simulate the vegetation types (Kim et al., 2018).
A process-based dynamic vegetation model (DVM)
allows for projecting forest responses to changing envi-
ronmental conditions. Cramer et  al. (2001) projected
­ O2 and climate change using six mod-
the impact of increased C
els as HYBRID, IBIS, LPJ, SDGVM, TRIFFID, and
VECODE at the global scale. Sitch et  al. (2008) used
multiple DVMs to estimate the vegetation response to
Special Report Emission Scenarios (SRES) and observed
higher uncertainties for the tropical region to drought
events. They also observed higher divergence in veg-
etation response to regional climate change than increas-
ing atmospheric C ­ O2. Gonzalez et  al. (2010) used the
MAPSS-CENTURY (MC) model to project the vul-
nerability and vegetation shift of the global ecosystem
owing to climate change. Halofsky et  al. (2013) inte-
grated the MC DVM with the state-and-transition mod-
els to identify the shift in vegetation under future climate
conditions. They concluded that vegetation distribution
is comparatively stable and does not change in the short
term, where rapid changes may occur with sharp changes
in environmental conditions and fire occurrences. Kim
et al. (2017) used the MC2 DVM (an updated version of the
MC model) to simulate the effect of climate alteration on global
Environ Monit Assess (2022) 194:903 Page 3 of 15  903
vegetation and reported that mitigation policies and con-
trol of wildfires might improve ecosystem resilience.
Ravindranath et al. (2006) used the BIOME4 vege-
tation model to assess the impact of climate change on
forests in India using the vegetation map generated by
the Forest Survey of India (FSI). Ravindranath et al.,
(2011a, b), Chaturvedi et al., (2011a, b), and Sharma
et  al. (2017) used the Integrated Biosphere Simula-
tor (IBIS) dynamic vegetation model to predict forest
cover change employing the various Intergovernmen-
tal Panel on Climate Change (IPCC) projected climate
scenarios. Steinkamp and Hickler (2015) studied the
impact of drought and water stress on forest dieback at
a global scale using multiple drought indices and the
LPJ-GUESS DVM. Their study indicated a negative
trend in drought, soil moisture and aridity index, and
both positive and negative mortality trends in the Dec-
can Peninsular region of India. They concluded that
dry climate regimes might experience more dryness
and extreme drought events inducing higher mortality.
In this study, the MC2 dynamic vegetation model
was used to simulate the impacts of multiple envi-
ronmental parameters on life form resilience under
the present and Representative Concentration Path-
ways (RCP) 4.5 projected climate change scenarios
in India. The results were compared with a vegetation
type map and satellite-derived net primary productiv-
ity (NPP) data. The mode simulated life forms were
used to estimate the resilience and change proneness
to alternative states.
Materials and methods
MAPSS‑CENTURY version.2 (MC2) model
description
The MC2 model is a spatially explicit vegetation
model that runs on each grid to simulate the life form
distribution with the environmental data. The MC2
model is an integrated model of “MAPSS” biogeog-
raphy and a modified version of the “CENTURY”
biogeochemistry model, along with a fire disturbance
model, “MCFIRE” (Bachelet et al., 2000). Using the
feedback mechanism, this model simulates the impact
of global climate change on ecosystem structure and
function at various spatial scales from local to global
(Bachelet, 2001). Using the long-term average cli-
mate, the MAPSS model predicts an initial vegetation
type. The “efolded” function is used to tempo-
rally smooth the climate, reducing the interannual
variability and vegetation type alteration at shorter
intervals. The leaf area index (LAI) and biomass of
trees and grasslands are simulated by the geochemical
module used in the updated biogeography module to
categorize the life forms. However, shrubland is not
simulated explicitly; instead, it is considered a short-
stature state. Additionally, the vegetation classifier
in the updated MC2 biogeography module utilizes
the temperature and carbon threshold to classify life
forms combining the climate zone. The MC2 model
uses many input parameters on plant functionality
and environmental conditions, which are integrated
and connected based on feedback mechanisms
(Kim et al., 2018).
RCP climate projections
Natural variability and past and present anthropogenic
activities lead to a continuous alteration of climate
conditions (Tao & Jain, 2005). Such alterations may
continue in different pathways depending on natural
acclimatization and adjustment of present and future
anthropogenic activities. Different phases of climate
projection under the IPCC continuously project such
changes. The four RCPs signify four projection sce-
narios based on the influences of various factors, e.g.,
greenhouse gas (GHG) emissions, atmospheric con-
centrations, air pollutant emissions, land use change,
population size, economic growth, changes in lifestyle,
energy use, technology, climate policies, etc. (Pachauri
et al., 2014). The RCP 4.5 climate projection has been
regarded as one of the most probable scenarios (https://​
aspof​rance.​files.​wordp​ress.​com/​2019/​08/​ipccc​o2rcp.​
pdf accessed on 24–07-2022).
The life form map was derived using a high–reso-
lution (60 m) vegetation type map of India generated
by Roy et  al. (2015). The India vegetation type map
was grouped into three life form categories as forest,
scrub, and grassland, where the barren and wasteland
were considered as treeless states, excluding the other
land use classes. The SRTM-derived DEM data was
used as elevation data, whereas the soil data provided
by International Geosphere Biosphere Programme
(IGBP) and International Satellite Land Surface Cli-
matology Project (ISLSCP) was used. The monthly
climate data from 1901 to 2000 was accessed from
the Climate Research Unit (CRU) database. The
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GCM projected climate data were downloaded from
the Earth System Grid Federation (ESGF) data portal
(www.​esgf-​node.​llnl.​gov/​search/​cmip5/). The moder-
ate resolution imaging spectroradiometer (MODIS)
data derived annual NPP product (MOD17A3H) were
downloaded from 2001 to 2015 and were utilized for
comparison with model estimates. The list of various
input data used in the study is given in Table 1.
The open-source version of the MC2 model avail-
able at a 0.5° grid scale was simulated in this study.
The grid with a cumulative vegetation area or treeless
state that occupies more than 33% of the grid’s area
was considered, where 484 out of 1330 grids were
selected. The maximum area occupied by a life form
category was considered the representative life form
in these selected grids. Similarly, the majority of soil
data was considered as a grid value, while the mean
value was considered for the rest of the continuous
variables.
The GCM projected climate data was compared
with the India Meteorological Department (IMD)
climate data to choose the best suitable climate pro-
jection for 2001 to 2100. The correlation and RMSE
were computed by comparing the simulated precipi-
tation and temperature data with the observed data
(IMD climate data). The grided IMD climate data for
2001 to 2015 were utilized for comparison with GCM
projected (Coupled Model Intercomparison Project
5; CMIP5) climate data for the RCP 4.5 scenario.
Preference was given to the model with the highest
coefficient of correlation and the lowest RMSE in the
GCMs ranking. The correlation and RMSE values of
different models were scaled from 0 to 1. Based on
Table 1  List of input data
Data Resolution Source
Vegetation type of India 60 m
Shuttle Radar Topographic Mission (SRTM) Digital 30 m
Elevation Model (DEM)
Soil type 1°
Past climate data (1901 to 2000) 0.5°
Future climate data (2001 to 2100) 1.4°
Moderate Resolution Imaging Spectroradiometer 500 m
(MODIS) Net Primary Productivity (NPP) for 2001 to
2015
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Environ Monit Assess (2022) 194:903
equal weightage, the sum of correlation and RMSE,
named prediction score (PS), were computed and
compared among the models.
R2max − R2
Scaled R2 = (1)
R2max − R2min
RMSEmax − RMSE
Scaled RMSE = (2)
RMSEmax − RMSEmin
PS =
1∑
Scaled R2 + Scaled RMSE (3)
2
The MC2 model was simulated at 0.5° grid resolu-
tion from 1901 to 2100. The model takes the average
climate data from 1901 to 1930 to initiate the model
(equilibrium mode), wherein the long-term climate
data is used to generate the initial vegetation type.
The model simulation continues unless a steady state
soil carbon pool is achieved. Thereafter, the model
takes monthly climate data as input for yearly predic-
tion (transient mode). For calibration and validation,
the potential vegetation type simulated by the MC2
model was reclassified into life form types and com-
pared with the observed plant life form map of India.
The field data and satellite data used in the develop-
ment of the vegetation type map of India were mostly
collected from 2000 to 2010 (Roy et al., 2015). Thus,
the majority occurrence (mode) in simulated life
forms from 2001 to 2010 was compared with the
observed life form map. The LAI and vegetation car-
bon threshold values used in the model for life form
Roy et al. (2015)
Google Earth Engine (GEE)
International Geosphere Biosphere Programme (IGBP) and
International Satellite Land Surface Climatology Project
(ISLSCP)
Climate Research Unit (CRU)
Earth System Grid Federation (ESGF)
Earth Data Search
Environ Monit Assess (2022) 194:903 Page 5 of 15  903
categorization were calibrated. The MODIS NPP
product from 2001 to 2015 was used to validate the
model performance by comparing it with the model
simulated productivity. The MODIS NPP product
was pre-processed, integrating the quality control
band in the Google Earth Engine (GEE). The mean
annual MODIS NPP data was derived based on the
model grid resolution (0.5°). With the calibrated
parameters, the model simulation continued to
predict the forest cover resilience with the projected
climate data. The resilience of life forms for the cur-
rent period (2001 to 2018) was computed with the
frequency occurrence (mode), wherein the change
proneness was estimated with the second-highest
frequency occurrence. For example, if in a forested
grid, forest life form was simulated in 6 years, scrub
in 8  years, and grassland in 4  years, then the grid’s
resilience would be 0.33 (6/18), and the grid showed
change proneness from the forest to scrub. Similarly,
the life form resilience for 2050 was estimated by
analyzing the frequency of life form occurrences from
2031 to 2050 (Fig. 1).
Fig. 1  Overall methodology flowchart
Results
GCM‑simulated climate data selection
The IMD climate data was compared with the projected
climate data of different GCMs to choose the best-performing
one for India. The spatiotemporal correlation and RMSE
are computed for monthly precipitation and temperature
between the GCM simulated data and IMD observed
data. The average correlation for temperature varied
between 0.31 and 0.71, whereas the average RMSE
varied between 6.41 and 11.50  °C for different models
(Table  2). Low correlations were observed for precipita-
tion that varied between 0.06 and 0.24, with RMSE var-
ied between 108.84 and 139.59 mm for different models.
The highest correlation in temperature projection was
observed for MRI-CGCM3 (0.71) and MIROC5 (0.71),
whereas the lowest RMSE was observed for the MPI-
ESM-LR (6.41  °C). However, the highest correlation in
precipitation projection is obtained for MIROC5 (0.24),
whereas the lowest RMSE was obtained for CNRM-
CM5 (108.84  mm). Comparing the precipitation and
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temperature projections among 13 GCM outputs, the
highest PS was observed for the CNRM-CM5 (0.88),
followed by MPI-ESM-LR (0.87) and CanESM2 (0.86)
(Table  2). Thus, the projected climate data from the
CNRM-CM5 model was used to project the vegetation
distribution in India.
MC2 calibration and validation
At 0.5° grid, 484 grids were selected, where the dominant
forest cover was observed in 357 grids, scrub in 35 grids,
grassland in 11 grids, and treeless state in the rest of the
81 grids (Fig.  2). The life form categorization param-
eters are optimized by comparing with the observed life
form and NPP. The woodland carbon threshold value
for the tropical and subtropical zone was adjusted from
1150  g  C  ­m−2 to 500  g  C  ­m−2. The default tree LAI
threshold value of 1.75 in the subtropical zone was opti-
mized as 3.75 (Fig. S1). The overall accuracy of 80.79%
was obtained for life form level comparison, while
62.82% was obtained with default parameters (Fig.  2).
The accuracy in NPP simulation was estimated as 0.66
(average bias 0.002; (∑NPPpredicted/∑NPPobserved)/n)
(Fig.  2). The differences between the model-simulated
and observed life form distribution are mostly found
in the Western Himalaya, arid and semi-arid regions
(Fig. 2). The majority of the errors are observed in grids
where multiple life forms coexist such as in the Western
Himalaya. In this region, forests are simulated as a domi-
nant life form in several grids, which are actually domi-
nated by treeless, grassland and scrub (Fig. 2; Fig. S2).
The forest-dominated grids simulated by the MC2 model
demonstrated good accuracy in the Western Ghats,
North-Eastern states, Deccan Peninsula, and major parts
of Western Himalaya (Fig. 2). Similarly, the simulation
was accurate for treeless dominant grids in arid and trans-
Himalayan regions. However, a few misclassifications
were observed in model simulation for grassland and
scrub dominated grids in the dry arid and Deccan Pen-
insular regions. The productivity estimated by MC2 was
found similar as estimated in the MODIS NPP data, indi-
cating a comparable spatial distribution with minor posi-
tive relative bias (0.002) (Fig.  2). The model simulated
productivity demonstrated higher accuracy for the West-
ern Ghats, Eastern Ghats, semi-arid and NE India com-
pared to Western Himalaya and Deccan Peninsula, where

Table 2  Comparison of different GCM (Coupled Model Inter- on the correlation for monthly precipitation and temperature
comparison Project 5; CMIP5) simulated climate data with between 2001 and 2015
India Meteorological Department (IMD) derived data based
Model Acronym Prediction Temperature Precipitation
score
R2 RMSE R2 RMSE

Center National de Recherches Météorologiques Coupled CNRM-CM5 0.88 0.69 8.53 0.23 108.84
Global Climate Model, version 5
Max Planck Institute Earth System Model, low resolution MPI-ESM-LR 0.87 0.66 6.41 0.19 116.32
Australian Community Climate and Earth System Simulator, ACCESS1.0 0.23 0.41 11.06 0.06 139.59
version 1.0
Hadley Center Global Environmental Model 2, Earth System HadGEM2-ES 0.40 0.39 11.50 0.14 125.35
Geophysical Fluid Dynamics Laboratory Climate Model, GFDL-CM3 0.56 0.54 10.91 0.24 128.87
version 3.0
Meteorological Research Institute Coupled Atmosphere–Ocean MRI-CGCM3 0.68 0.71 8.12 0.14 125.35
General Circulation Model, version 3
Model for Interdisciplinary Research on Climate, version 5 MIROC5 0.83 0.71 6.51 0.24 128.87
Canadian Earth System Model, version 2 CanESM2 0.86 0.67 7.23 0.16 108.91
Norwegian Earth System Model, intermediate resolution NorESM1-M 0.48 0.46 10.34 0.10 121.23
L’Institut Pierre-Simon Laplace Coupled Model, version 5 IPSL-CM5A-LR 0.24 0.31 11.13 0.09 138.19
Goddard Institute for Space Studies Model E, coupled with GISS-E2-R 0.41 0.50 10.00 0.12 135.29
Russell Ocean model
Commonwealth Scientific and Industrial Research Organization CSIRO-Mk3.6.0 0.75 0.67 6.65 0.12 121.96
Mark 3.5
Centro Euro-Mediterraneo sui Cambiamenti Climatici Climate CMCC-CM 0.80 0.68 6.79 0.19 123.49
Model

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Fig. 2  Grid-wise comparison
between (i) observed and
(ii) simulated life form,
and between (iii) MODIS
observed and (iv) model-
simulated NPP, (v) correlation
between MODIS observed
and model-simulated NPP

the model simulated NPP overestimated the observed the Deccan Peninsular regions. However, the least resil-
NPP. ient forest are found in the dry semi-arid regions and
southern part of Eastern Ghats (Fig. 3a). Around 3.92%
Model simulation and estimation of life form and 8.12% of the total forests showed a higher change
resilience proneness toward the scrub and grassland dominant
landscapes, mostly estimated in the least to moderately
The frequency occurrence of the model simulated life resilient forested grids (Pr < 0.5) (Fig. 2(ii)). The lower
form classes from 2001 to 2018 was used to estimate annual precipitation in the semi-arid region could be
the current resilience and change proneness. Then, accounted for the lower forest cover resilience, whereas
83.75% of the total forest cover demonstrated high the seasonal dryness and higher chances of forest fire in
resilience (probability of resilience, Pr ≥ 0.75), whereas the Eastern Ghats and east coastal region could lead to
6.16% area showed the least resilience (Pr < 0.25) or lower resilience. The majority of the scrub-dominated
is highly change prone (Table  3). The highly resilient areas (≈ 82.86%) showed a lower resilience (Pr < 0.25),
forest covers were found in the wet climate regimes in while only 8.57% of scrubs are estimated to be highly
NE India, Western Himalaya, Western Ghats, and a few resilient (Pr ≥ 0.5) (Fig. 3(iii); Table 3). About 48.57%
grids in the Deccan Peninsular region (Fig. 2(i)). Mod- of scrubs found in the Deccan Peninsular and Western
erate to high resilience (0.25 ≤ Pr < 0.75) was estimated Himalaya wetter regimes indicated a higher change
in 10.08% of the total forest covers, mostly observed in proneness toward forest (Fig. 3(iv); Table 4). However,
the moderately wet Eastern Ghats and western part of 25.71% of scrubs in the dry semi-arid region estimated
change proneness toward the grassland, and 17.14% in
the extremely dry arid region showed a high transition
Table 3  Area estimates (in percentage) of different life forms tendency toward the treeless state (Fig. 3(iv); Table 4).
at different resilience levels Grassland-dominated grids are mostly observed in the
Resilience Forest Scrub Grassland Treeless drier parts of the trans-Himalaya and arid regions. All
the grassland-dominated grids are estimated under
0–0.25 6.16 82.86 100.00 44.44
the lowest resilience category (Pr < 0.25) (Fig.  4(i);
0.25–0.5 7.00 8.57 0 3.70
Table 3). About 36.36% of grasslands exhibited higher
0.5–0.75 3.09 5.71 0 4.94
change proneness to forests, while the rest of 63.64%
0.75–1 83.75 2.86 0 46.91
of grasslands indicated higher change proneness to the

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Fig. 3  Estimated resilience
and change proneness maps
of forest (i, ii) and scrub
(iii, iv)

treeless state (Fig.  4(ii); Table  4). Treeless-dominated exhibited a high transition tendency to grasslands,
grids in the extremely dry arid and trans-Himalaya which could be attributed to moisture limitation.
regions showed high resilience (Fig.  6(a)). The least
resilient treeless areas are mostly found in the northern Model simulation and life form resilience projection
boundary of the Western Himalayas, which exhibited
a higher change proneness to the forest (Fig. 6b). This The change in resilience was estimated by subtracting
could account for the overestimation in precipitation by the current resilience from the projected resilience,
the CNRM-CM5 model in the northern boundary of wherein the positive change indicates an increase in
the Western Himalayas and north Sikkim. Two treeless resilience or reduced risks of regime shift and vice-
dominated grids in the moderately dry semi-arid region versa. The estimated area under the various resilience
categories for each life form and their change prone-
ness was similar to the current resilience, excluding
the grassland. About 85.43% of forest covers were
Table 4  Change proneness area estimates (in %) matrix of dif-
ferent life forms for 2001–2018
predicted highly resilient, whereas only 8.96% of for-
est cover was less resilient (Table  5). The forest in
Simulated wet climate regimes of North-East India, Western
Forest Scrub Grassland Treeless Himalayas, and Western Ghats are predicted highly
resilient and may withstand climate change (Fig. 5(i)
Observed Forest 87.96 3.92 8.12 0
and (ii)). A significant change in forest cover resil-
Scrub 48.57 8.57 25.71 17.14
ience was predicted in dry semi-arid to moderately
Grass 36.36 0 0 63.64
wet Eastern Ghats and east coastal regions. Several
Treeless 45.68 0 2.47 51.85
forested grids may experience forests to grassland

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Fig. 4  Estimated resilience
and change proneness maps
of grassland (i, ii) and tree-
less state (iii, iv)

conversion in the western part of the Deccan Pen- trans-Himalaya regions and forests in wet regions
insular region, while forests to both scrubland and (Fig.  6(ii)). Although several treeless grids in the
grassland transition in the Eastern Ghats and east Western Himalayas demonstrated an insignificant
coastal regions (Table  6; Fig.  5(ii)). The increase change in resilience, they may experience a regime
in scrub resilience is predicted in a few grids in the shift to the forest in future (Fig. 6(iii) and (iv)).
semi-arid region; however, such grids still dem-
onstrated a tendency of scrub to grassland conver-
sion (Fig.  5(i) and (ii)). About 72.73% of grassland Discussion
is projected under a lower resilience state, whereas
the rest of the 27.27% areas are expected under a Although the limitations of GCM projections are well
moderate resilience state (0.25 ≤ Pr ≤ 0.75) (Table 5; known, these projections are currently the best avail-
Fig. 6(i)). The grassland may experience conversion able input to approximate change in the future climate
to a treeless state in drier parts of arid regions and and the dependent processes. The comparison between
various GCM’s projected data and observed IMD data
demonstrated higher accuracy in projecting tempera-
Table 5  Area (in %) statistics of predicted life form resilience ture than precipitation. The comparative study indicated
for 2031–2050 that the CNRM-CM5 climate data is the most suitable
Resilience Forest Scrub Grass Treeless projection for India, followed by MPI-ESM-LR. Ying
and Chong-Hai (2012) assessed the performance of
0–0.25 8.96 94.29 72.73 54.32
GCM simulations in China and reported that tempera-
0.25–0.5 2.24 2.86 18.18 6.17
ture simulations are more accurate than precipitation.
0.5–0.75 3.36 0.00 9.09 2.47
Out of 13 GCM outputs, they observed the highest
0.75–1 85.43 2.86 0.00 37.04
projection accuracy for the CNRM-CM5, followed by

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 903   Page 10 of 15 Environ Monit Assess (2022) 194:903

Fig. 5  Projected change
in resilience and life forms
maps of forest (i, ii) and
scrub (iii, iv)

the MPI-ESM-LR. This also supports Watterson et al. trans-Himalaya regions. Gouda et  al. (2018) evaluated
(2014), who evaluated the GCM performance at global the GCM simulations in extreme rainfall events over
and regional scales and reported the highest accuracy India. They reported a lower spatial correlation in the
for the CNRM-CM5 model, followed by the MPI-ESM- drier regions as arid, semi-arid, Deccan Peninsula, and
LR model. Raju et al. (2017) evaluated the performance trans-Himalaya, which substantiates the current study.
of the various CMIP5-based GCM projections in simu- The projected change in precipitation demonstrated a
lating the maximum and minimum temperature over reduction (< 50 mm) in the Western Himalayas and sev-
India and reported CNRM-CM5 as one of the best per- eral parts in NE India, while an increase (> 50 mm) in
forming models. The spatio-temporal correlation map various parts of the Deccan Peninsula, Eastern Hima-
for precipitation indicates a comparatively lower corre- laya, and Western Ghats (Fig.  S3). The CNRM-CM5
lation (< 0.25) in the drier parts of semi-arid, arid, and projected change in minimum and maximum tempera-
ture indicated a significant increase (> 1  °C) mainly
in the dry arid, Western Himalaya and trans-Himalaya
Table 6  Change proneness area (in %) matrix of different life region, excluding several parts in the Deccan Peninsula
form in 2031–2050 compared to 2001–2018 region (Fig. S3).
The regression-based resilience estimation approaches,
Predicted
e.g., precipitation-dependent life form estimation, are
Forest Scrub Grass Treeless
simplistic in nature and apply a uniform impact of the
Observed Forest 88.52 2.80 8.68 0.00
independent variable(s). Several studies have identified
Scrub 45.71 2.86 34.29 17.14
the precipitation threshold as 1400  mm, beyond which
Grass 36.36 0.00 9.09 54.55
forests are highly resilient in India (Behera et  al., 2018;
Treeless 55.56 0.00 4.94 39.51
Das & Behera, 2019). Such techniques heavily rely on

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Environ Monit Assess (2022) 194:903 Page 11 of 15  903
Fig. 6  Projected change
in resilience and life forms
maps of grassland (i, ii) and
treeless state (iii, iv)
the statistical relationship between long-term climate
and vegetation distribution. However, such approaches
become complex with multiple variables, which neither
consider the underlying processes nor include the feed-
back mechanisms, which limits their use in vegetation
distribution prediction based on the environmental con-
ditions. In comparison, the process-based MC2 model
included multiple variables on monthly climate data of
each year of simulation, edaphic, topographic, etc., and
simulated the various land surface processes, including
the forest fire occurrences, water, and nutrient cycle and
projected the life form distribution with more than 81%
accuracy. Moreover, the MC2 model generated various
essential intermediate layers on the biophysical and struc-
tural attributes at an annual scale as productivity, LAI,
plant height, biomass, etc., and simulated the differential
impact of climate change in different bioclimate zones.
The current study optimized the LAI and vegetation car-
bon thresholds used for life form segregation according
to observed life forms. The MC2 model simulated NPP
exhibited a high correlation (R2 > 0.65) with a minor posi-
tive relative bias as compared with MODIS NPP. Similar
accuracy was reported by Zhou et al. (2019), where they
observed a high correlation (R2 ~ 0.74) with a minor
overestimation in MC2 simulated NPP compared to
MODIS NPP. The MC2 model well projected the life
forms for most of the Indian mainland. The observed
results highlighted the strength and limitations of the
MC2 in modeling the climate and topography responses
in simulating vegetation in a heterogeneous landscape
of India (Das et al., 2019). The errors in life form simu-
lation are mostly found in the dry semi-arid and trans-
Himalaya regions and a few grids in the Deccan Penin-
sular region. Therefore, the model simulated life forms
under the projected climate scenarios could include bias
in these regions. The accuracy in life form simulation
could increase if the coexistence of multiple life forms is
taken care of or the model is simulated at a smaller grid
size, especially in hilly terrains where life form changes
rapidly due to closely spaced micro-climatic conditions.
Additionally, the limitations might be imposed by climate
data, coarse resolution soil maps, landscape heterogeneity,
and anthropogenic disturbances that might have caused
inaccuracies in life form simulation (Kim et al., 2018).
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 903   Page 12 of 15
The frequency or mode of life form occurrences for
a given period was used to compute the resilience and
change proneness. Around 88% of forest-dominated
grids in wet climate regimes showed highly resilient
under the present and projected climate. The moder-
ate and least resilient forest cover grids exhibit a tran-
sition tendency to scrubs and grassland, respectively.
This could be attributed to the long dry spells and fire
events that promote the seasonal scrub and grassland.
The seasonally dry tropical forests are typically char-
acterized by their adaption capacity to seasonal water
stress in dry parts of the tropics. According to Pulla
et  al. (2015) and Allen et  al. (2017), seasonally
dry tropical forests are already in stressed conditions
closely residing to the climate threshold and may not be
more resilient than moist tropical forests. However,
such forests are more adaptive or possess high resist-
ance to seasonal water stress, drought, and fire events.
Ravindranath et  al. (2006) used the BIOME4 vegeta-
tion model to assess the impact of climate change on
Indian forests. They reported that 77% and 68% of the
forested grids would experience a shift in forest type
under the IPCC A2 and B2 scenarios, respectively, by
2085. Ravindranath et al. (2011b) and Chaturvedi et al.
(2011b) used the IBIS DVM and predicted 39% and
34% shifts in forested grids with A2 and B2 scenarios,
respectively. Using the RCP projected climate sce-
narios and IBIS DVM, Sharma et  al. (2017) reported
that 46% and 50% of the forested grids would be vul-
nerable by 2030 and 2080, respectively. Their study
showed that the forest in wet climate regimes is less
vulnerable than those in drier regimes. Around ~ 49%
of the scrub-dominated grids indicated a higher poten-
tial for forest cover in the Deccan Peninsular and West-
ern Himalayas, which could result from the combined
disturbances of climate change, fire occurrences, and
anthropogenic disturbances (Baker & Williams, 2015).
Similarly, grasslands in the dry and wet regimes exhib-
ited change proneness toward treeless and forest states,
respectively. Several treeless dominated grids in wet
regions of the Western Himalayas exhibited less resilience
and high change proneness toward the forest. The high-
est and lowest resilience of forest cover grids in the
corresponding wet and dry climate regimes indicate
moisture availability as the primary determinant of life
form resilience in tropical/ sub-tropical regions (Das &
Behera, 2019).
The CNRM-CM5 projected climate change indi-
cates more warming and a wetter climate, wherein the
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Environ Monit Assess (2022) 194:903
minimum and maximum temperatures are projected
to expand westward and upward in the arid and trans-
Himalaya region. Telwala et  al. (2013) studied the
impact of climate change on montane flora in Sikkim,
Eastern Indian Himalayas. They reported that the
warming climate had induced upper range extensions
of 23–998 m and higher species richness in the upper
alpine zone. The Ministry of Earth Sciences (MoES),
the Government of India’s climate change assessment
report based on the CMIP5 projections, also high-
lighted the predicted increase in temperature and
precipitation in the Hindu Kush Himalayan region
(Krishnan et al., 2020). Using IPCC projected climate
scenarios, Rubel and Kottek (2010) studied the pro-
jected shifts in climate regimes and highlighted the
alterations in the climate regimes over Asia. They
also simulated MC2 and predicted a tree-line shift in
the temperature-limited Western Himalayas. This sup-
ports the observation by Bachelet et  al. (2015), who
reported shifts in woody vegetation following the
changes in warming climate along the elevation gradi-
ent in the USA. In India, Ravindranath and Sukumar
(1998), Chaturvedi et  al. (2008), and Ravindranath
et  al. (2011a) highlighted the projected increase in
precipitation and temperature might lead to increased
forest productivity and shift around the boundaries
following elevational and rainfall gradients. However,
the lower accuracies in the GCM projection in the
trans-Himalaya region induced lower confidence in
the predicted life form shift. The forest cover in wet
regimes of the Western Ghats, Eastern and Western
Himalayas, and Deccan Peninsula showed high resil-
ience, while less resilience in the drier semi-arid and
Eastern Ghats may experience a regime shift to grass-
land and scrub. The grassland and scrub-dominated
grids projected more treeless in dry regions and veg-
etated in wet areas.
Conclusions
Although many forest ecosystems may be adapted to
their natural disturbance regimes, it is unclear how
projected climate change would affect forest ecosys-
tems and their adaptation processes. Understanding
ecosystem sustainability is essential in ecological
studies and has policy-level implications in climate-
adaptive forest conservation and management. For-
est cover grids are predicted to be highly resilient in
Environ Monit Assess (2022) 194:903 Page 13 of 15  903
wet climate regimes such as the Western Ghats, NE
India and Western Himalayas under the current and
future climate. However, the seasonally dry tropical
forests in the Eastern Ghats and dry semi-arid regions
are estimated less resilient, which may experience a
regime shift toward scrub and grassland. Moreover,
scrub and grasslands in alternate wet and dry regimes
may experience a regime shift toward the forest and
treeless state. The future prediction demonstrated
an upward temperature shift in the Western Himala-
yas and trans-Himalaya, which may facilitate forest
growth at higher elevations. The spatially explicit for-
est cover resilience map holds promises in ecological
research and climate adaptive forest management and
conservation strategies.
Acknowledgements  This study has been carried out under
the framework of “Climate Change Effects on Indian Forest
Cover, a project under DST CoE in Climate Change.” The veg-
etation type map generated against a national-level “Biodiver-
sity Characterization project” was utilized here, is thankfully
acknowledged. The authors acknowledge the support provided
by the authorities of the Indian Institute of Technology Kharag-
pur during this study.
Declarations  endemic flora of biodiversity hotspots in India. PLoS
Conflict of interest  The authors declare no conflict of inter-
est.
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