Ecological Indicators 111 (2020) 106025

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Ecological Indicators
journal homepage: www.elsevier.com/locate/ecolind

Biomass and soil carbon along altitudinal gradients in temperate Cedrus T

deodara forests in Central Himalaya, India: Implications for climate change
mitigation
⁎
Mehraj A. Sheikha, Munesh Kumara, N.P. Todariaa, Rajiv Pandeyb,
a
Department of Forestry and Natural Resources, HNB Garhwal University, Srinagar Garhwal, Uttarakhand, India
b
Forest Research Institute, Dehradun, India

A R T I C LE I N FO A B S T R A C T

Keywords: Carbon inventories are urgently needed for understanding climate dynamics and implementing climate miti-
Climate response gation strategies, but such data is scarce in for forest ecosystem of Himalaya. Therefore, the present study focuses
Carbon sequestration to supplement the existing information by estimating carbon stocks in temperate forests dominated by Cedrus
Forest litter deodara along the altitudinal gradients in the Central Himalaya. Three altitudes i.e., lower (1750 m), middle
GHG removal
(1900 m) and upper (2050 m) of the stand was considered to understand and estimate biomass and soil carbon
REDD+
storage potential of the forests based on the standard protocol. The results showed that the soil carbon stock
(SOC) decreased significantly with increasing soil depths and altitudes. Litterfall production in the forests varied
adversely with altitude. Above and below-ground biomass carbon stock also decreased along with altitude. The
study observed that the total carbon stock (soil, trees and forest floor) of the Cedrus deodara forest in different
altitudes was 395.4 t ha−1 (lower altitude), 321.6 t ha−1 (middle altitude) and 282.5 t ha−1 (upper altitude).
The estimates of the Cedrus forests would provide guidelines for estimating carbon for forest-based mitigation
activities in the Himalayan region.

1. Introduction offsetting capabilities of the forests. The precise estimation of forest

The concentration of atmospheric CO2 has accelerated upward
during the past few decades with an annual rate of 1.91 parts per
million (ppm) due to various anthropogenic activities, leading to in-
crease in the atmospheric CO2 concentration from 317 ppm in 1958 to
410 ppm in 2018 (NOAA, 2017). The change in CO2 is instrumental for
changing the temperature and has large implications on the social,
economical and ecological systems. Therefore, there is a need to reduce
the emissions leading to limit the increase in global temperature below
2 °C in respect to pre-industrial level (i.e., measured from 1750) (IPCC,
2014). Forests are important for global carbon cycle because they se-
quester carbon from the atmosphere, and also emit carbon resulting
from loss of biomass (Houghton et al., 2009; Kishwan et al., 2012).
The carbon storage by the forests contributes to emission reduction
by offsetting the GHG emissions by other sources (Houghton 2005;
Houghton et al., 2009). For example, the mitigative role of India’s
forests was estimated to 9.3% at the year 2000; and 4.9% in 2020 for
the annual emissions of the country (Kishwan et al., 2012). Precise
information of forest carbon stock is important for understanding the
carbon is also required to understand the role of the forest for mitiga-
tive actions to achieve the National Determined Commitment (NDC)
such as REDD+ (Kishwan et al., 2012); and evaluating the role of
forests for their ecosystems services to the poor (Pandey, 2009). The
estimates of forest carbon are also required for evaluating forest de-
gradation, as anthropogenic extractions from forests result in the de-
gradation of forests (Malik et al., 2016). For example, carbon emissions
from world forests due to deforestation and forest degradation have
been reduced by 1 GT i.e. from 3.9 GT to 2.9 GT during 2011–2015
(FAO, 2015).
In contrast, precise estimates for the carbon in specific forests are
largely not available due to lack of resources and lack of experts across
the globe. Besides this, the volume of work for the estimation of the
forest carbon for larger forest patch such as at sub-national, national
and global level are huge and require large number of experts
(Houghton et al., 2009; Sharma et al., 2010b). Moreover, the changes in
the structure and composition of forests along with differences in in-
dividual tree sizes and shapes restrict the precise evaluation of forests
carbon of a larger patch. The assessment of sequestration potential of

⁎
Corresponding author at: Forest Research Institute, Dehradun, Uttarakhand, India.
E-mail addresses: mehrajshkh@gmail.com (M.A. Sheikh), rajivfri@yahoo.com (R. Pandey).

https://doi.org/10.1016/j.ecolind.2019.106025
Received 13 December 2018; Received in revised form 12 October 2019; Accepted 16 December 2019
1470-160X/ © 2019 Elsevier Ltd. All rights reserved.
M.A. Sheikh, et al.
forests were attempted in different parcels of forests across the globe for
various purposes such as at a global scale (Houghton, 2005; Houghton
et al., 2009); forest type basis in India (Pandey et al., 2011); tree
dominance basis in Himalaya (Kumar and Pandey, 2017); plantation
basis at sub-national scale in India (Gupta and Pandey, 2008); habitat
basis in England (Alonso et al., 2012). Moreover, several attempts have
also been made for carbon estimation for the country (Haripriya, 2000;
Chhabra and Dadhwal, 2004). However, for forestry-based mitigation
actions, these studies were challenged due to the lack of precise sta-
tistics for carbon stock for the specific land parcel leading to undermine
the mitigative role of forests at one side and restrict the planning ap-
proach for the address of climate change impacts on the other (IPCC,
2014). Therefore, precise estimation of carbon stock at either stand
level or species level or other required land parcels would be imperative
and would lead to supplementing the reduction of overall complexities
of mitigation options (Wutzler et al., 2007; UNFCCC, 2015). Moreover,
understanding the change in carbon stocks in the forests would further
assist the forest manager to regulate the rules and regulation for ef-
fective management of forests focussing to deliverables of services to
the humans.
The present study was motivated by the fact that the precise esti-
mation of carbon stock is essentially required for various purposes in-
cluding incentivization for mitigation actions. This study was con-
sidered to address the gap by providing an estimate of carbon for Cedrus
tree dominant forests in the Himalaya region. The study attempts to
estimate carbon stock in the moist (Cedrus deodara Loud.) Deodar forest
(Type – 12/C1C) (Champion and Seth, 1968) of the Himalayan region in
India. Cedrus deodara, usually found in pure stands, occurs at altitudinal
range from 1750 to 2150 m asl and typically gregarious. Major as-
sociates of the Cedrus deodara are Abies pindrow, Picea smithiana and
Pinus wallichiana (Luna, 2005). The usual girth of tree ranges between
2.4 and 3.6 m and height up to of 35 m (Luna, 2005). The tree is used
for several purposes including timber for building. Keeping this in view
along with the wide distribution and importance of Cedrus deodara
forests, it is important to estimate biomass and soil carbon storage ca-
pacity for predicting carbon storage in Himalayan forests. The present
study attempts to estimate carbon stock of Cedrus deodara forests as
well as the seasonal contribution of the forest floor to carbon stock in
the forest.
2. Materials and methods
2.1. Focus of study
Uttrakhand with a geographic area of 53,484 km2 lies between the
latitude of 28° 42́ and 31° 28́ N and longitude of 77° 35́ and 81° 50́ E,
with altitudinal ranges between 400 and 3500 m. amsl (Fig. 1). The
forests in the state are spread in 65% of the total geographical area
comprising sub-tropical Chir pine coniferous and mixed deciduous
woody of the temperate zone with tree species such as Juglans regia,
Picea smithiana, Cedrus deodara and Pinus gerardiana. The study focuses
on the hilly part of the state i.e., Garhwal Himalaya. The vegetation and
soils of Garhwal Himalaya vary according to aspect, altitude and cli-
mate. The hilly terrain of the region contains densely forested slopes
containing generally clayey to sandy loams sedentary soil with podzo-
lization. The climate of the study region is temperate with very cold in
winter and pleasant in summer. The temperature of the region ranges
between −2 °C to 20 °C in January and 24 °C to 36 °C in June and July
(Kumari et al. 2018) with an average annual rainfall of 2180 mm
generally commencing from mid-June and extending till mid-Sep-
tember with relative humidity between 54% and 63%.
2.2. Sampling for data collection
Three forest stands of Cedrus deodara located at three different al-
titudes at north aspect were selected at random for sampling from the
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Ecological Indicators 111 (2020) 106025
region. The geo-coordinates of the three sites were N 30o08′00.52″ and
E 078o46′32.38″ (Site-I), N 30o09′43.46″ and E 078o45′22.44″ (Site-II),
N 30o09′35.29″ and E 078o45′19.79″ (Site-III) with an altitude of
1750 masl, 1900 masl and 2050 masl, respectively. The approximate
age of the forest is above 100 years, as informed by the villagers during
the survey. Three random selected sample plots were laid in each stand
following Hairiah et al. (2001). The size of the plot was 20 × 100 m for
measurement of trees having stem diameter > 30 cm at breast height
for estimation of form factors for each species. A subplot of size
(5 m × 40 m) was laid for evaluation of trees of stem diameter between
5 and 30 cm and tree density (ha−1) was estimated for the two classes
of trees in each plot. Within these plots, smaller square subplots of
0.5 m2 size were laid for collection of soil samples. Litter traps were laid
within these smaller subplots for collection of litter. For analysis of soil,
6 samples each at four depths (6 samples × 4 depths) were collected
from each sample sub-plot at four different soil depths i.e., 0–0.3, 0–0.5,
0–1.0, 0–1.5 m. Therefore, from each stand 72 samples were considered
for estimation of soil organic carbon (SOC) consisting of six samples at
four depths from the 3 sample plots. Soil samples were collected by
metalcore sampling cylinder (core method).
2.3. Analysis of physico-chemical properties of soil
The collected soil samples were brought to the laboratory following
standard protocol and labeled for further processing. The samples were
air-dried grounded in a cyclotec-1030 mill and passed through a 2 mm
sieve (Jackson, 1967). The fractions > 2 mm and < 2 mm size was
stored separately and the fine fractions were used for further analysis.
2.3.1. Bulk density
Collected soil samples were oven-dried at 100 ± 5 °C for 48 h and
weighed. The bulk density was estimated using a standard core method
(Wilde et al., 1964) using the following formula.
Bulk density(g/cm3) = Mass of dry soil(g)/Volume of dry soil(cm3)
2.3.2. Coarse fragment
Percent coarse fragments (> 2 mm size) in soils were estimated to
evaluate the soil mass. The soil samples were kept in the 2 mm sieve,
and running water was supplied through the sample. Soil particles <
2 mm were allowed to wash away with running water. The remaining
fractions in the sieve were dried and weighted for estimation of per-
centage coarse fragment.
2.3.3. Moisture content (%)
The moisture content of the soil samples was estimated as per the
standard method (Misra, 1968
Moisture content(%) = [(Fresh weight − Dry weight)/Fresh weight] × 100
2.3.4. Estimation of soil carbon stock in soil
Walkley and Black, Krishan et al. (2009) an analytical technique
was applied to estimate soil organic carbon (SOC) by weight to volume
(Walkley and Black, 1934). The soil carbon stock (SOC) was estimated
by Batjes (1996) approach. For an individual profile with k depths, the
total organic carbon by volume is estimated by the following formula:
k
Qi = ∑ ρi Pi Di (1 − Si)
i=1
where Q i is the total amount of organic carbon (in Mg m−2) over depth,
ρi is the bulk density (Mg m−3) of layer i, Pi is the proportion of organic
carbon (g C g−1) in layer i, Di is the thickness of ith layer (m), and Si is
the volume of the fraction of fragments > 2 mm (Batjes, 1996).
M.A. Sheikh, et al.
Fig. 1. Location map of the study area.
2.4. Biomass estimation
2.4.1. Aboveground biomass
The diameter at breast height (dbh) was measured by tree calliper
and height (h) through Ravi’s multimeter. The form factor (F) of a tree
was estimated with Spiegel relaskope by using formula F = 2*h1/3h
(Pressler, 1895; Bitlerlich, 1984) where h1 is the height at half dbh.
Volume (V) was estimated with V = F × h × g, where g the basal area
g = π (dbh/2)2 (Pressler, 1895) as exemplified by Koul and Panwar
(2008). Biomass of wood was estimated by multiplying specific gravity
with volume. Specific gravity was estimated based on stem core by the
maximum moisture method (Koul and Panwar, 2008).
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Ecological Indicators 111 (2020) 106025
2.4.2. Branch biomass
The total number of branches irrespective of size was counted on
each of the sample trees and categorized on the basis of basal diameter
into three groups, viz < 6 cm, 6–10 cm and > 10 cm. Fresh weight of
two randomly selected branches from each group were recorded, se-
parately. The dry weight of branches was estimated by using equation
(Chidumaya, 1990) as; Bdwi = Bfwi/(1 + Medbi) where Bdwi is the oven-
dry weight of branches, Bfwi the fresh weight of branches and Medbi the
moisture content of branches on a dry weight basis. Total branch bio-
mass per sample tree was determined as; Bbt = n1 bw1 + n2 bw2 + n3
bw3 where Bbt is the branch biomass per tree, ni the no of branches in
the ith branch group (i = 1(1)3), wi is the weight of the one branch in
the group (Koul and Panwar, 2008).
M.A. Sheikh, et al.
2.4.3. Leaf biomass
Leaves from five randomly selected branches of individual trees
were collected from five randomly selected trees per quadrate. The
leaves were weighed and oven-dried separately to a constant weight at
80 ± 5 °C. The average leaf biomass was estimated by multiplying the
average biomass of the leaves per branch with the numberof branches
in a single tree and the number of trees in a quadrate (Chidumaya,
1990; Koul and Panwar, 2008).
2.4.4. Belowground biomass
Below ground biomass density (BGBD) (fine and coarse roots) was
estimated for each species using the allometric equation by Cairns et al.
(1997).
BGBD = exp{ −1.059 + 0.884 × ln(AGBD) + 0.284}
where AGBD is above ground biomass density.
2.4.5. Understorey and Herbaceous biomass
Herbaceous biomass was estimated by collecting all the plants from
a randomly laid sub-quadrate of size 1 m × 1 m in the sample plot. All
collected plants were oven-dried 65 ± 5 °C to a constant weight. Shrub
biomass was also estimated by applying a similar procedure.
2.4.6. Litter biomass
All vegetation < 5 cm dbh were harvested from the considered
sample plot. Litter was collected in litter traps on sample plots and mass
of the total fresh sample (in g m−2) was measured. A composite fresh
sub-sample (~300 g) were taken and weighted after thorough mixing
for subsequent oven drying at 80 °C for conversion to dry weight and for
further analysis (Hairiah et al., 2001).
Total dry weight(kg m2)
Total fresh weight(kg) × Subsample dry weight (g)
=
Subsample fresh weight(g) × Sampled area(m2)
2.4.7. Estimation of carbon stock in biomass
Biomass contains carbon from 45 to 50% of dry matter (Chan, 1982;
Kishwan et al., 2012; Pandey et al., 2014). In the present study, the
total amount of carbon stocked was 0.45 times of dry biomass as sug-
gested by Woomer (1999).
3. Results
The collected data were statistically analyzed for various para-
meters for Cedrus deodara stand. The details of descriptive statistics of
different parameters and carbon for various observed tree species as
well as forest floor in the stand are discussed below in details.
3.1. Physico-chemical properties of soil
The diverse trend in soil moisture was observed across the three
altitudes with a maximum at the higher elevation. In lower altitude, soil
moisture was maximum at the surface i.e. 0–30 cm and minimum at the
120–150 cm. Bulk density (BD) and carbon fraction (CF) was minimum
at 0–30 cm and increased with depth (Table 1). In middle altitude, the
moisture content reduced from 24.3 ± 5.50% (0–30 cm) to
13.3 ± 2.19% (120–150 cm). However, BD and CF were observed to
increase with increasing depth (Table 1). In upper altitude, moisture
percent was decreased with increasing soil depth, however, BD and CF
increased with increasing soil depth (Table 1).
3.2. Soil carbon
In all the three altitudes (lower, middle and upper), SOC and carbon
stock were decreased from the top layer (0–30 cm) to bottom
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Ecological Indicators 111 (2020) 106025
(120–150 cm) (Table 1). The depth-wise analysis revealed that carbon
stock increased with soil depth from 0 to 30 cm to 0–150 cm in all the
three altitudes i.e., 104.8 ± 13.11 to 174.3 ± 23.69 t ha−1 (lower
altitude), 105.3 ± 15.34 to 173.0 ± 11.87 t ha−1 (middle altitude)
and 89.7 ± 20.93 to 139.3 ± 12.70 t ha−1 (upper altitude) (Table 1).
3.3. Above and below ground biomass and carbon
In lower altitude, six tree species i.e., Cedrus deodara, Cupressus
torulosa, Pinus roxburghii, Quercus leucotrichophora, Quercus floribunda
and Myrica esculenta were observed with decreasing density and basal
area (Table 2). The maximum mean diameter was observed for Cedrus
deodara followed by Pinus roxburghii and Cupressus torulosa with bole
volume of 563.6, 2.8 and 126.3 m3 ha−1, respectively (Table 2). Total
AGB and biomass in different tree components i.e., bole, branch, twig
and foliage is reported in Table 3. Total above-ground tree biomass was
highest (307.3 t ha−1) in Cedrus deodara followed by Cupressus torulosa
(70.5 t ha−1), Pinus roxburghii (2.3 t ha−1), Quercus leucotrichophora
(0.3 t ha−1), Quercus floribunda (0.2 t ha−1) and Myrica esculenta
(0.04 t ha−1) (Table 3). The BGB for each tree was also estimated and
reported in Table 3. The total biomass (AG and BG) for each tree species
as Cedrus deodara, Cupressus torulosa, Pinus roxburghii, Quercus leuco-
trichophora, Quercus floribunda and Myrica esculenta were 387.0, 92.2,
3.4, 0.5, 0.3 and 0.07 t ha−1, respectively. The distribution of carbon
stock (t ha−1) allocation in different components of each species is
reported in Table 4. The species wise, AG and BG carbon stock were
138.3 and 35.8 t ha−1 (Cedrus deodara), 31.7 and 9.8 t ha−1 (Cupressus
torulosa), 1.1 and 0.5 t ha−1 (Pinus roxburghii), 0.12 and 0.1 t ha−1
(Quercus leucotrichophora), 0.1 and 0.04 t ha−1 (Quercus floribunda) and
0.02 t ha−1 and 0.01 t ha−1 (Myrica esculenta). The maximum carbon
stock (AG and BG) was observed for Cedrus deodara followed by Cu-
pressus torulosa (Table 4).
In middle altitude, the associated species of Cedrus deodara were
Cupressus torulosa, Rhododendron arboreum, Quercus leucotrichophora
and Quercus floribunda. The highest density and basal area were ob-
served for Cedrus deodara followed by Cupressus torulosa, Rhododendron
arboreum, Quercus leucotrichophora and Quercus floribunda (Table 2). In
this site, Cedrus deodara was having maximum mean diameter and
height followed by Cupressus torulosa, Rhododendron arboreum, Quercus
floribunda and Quercus leucotrichophora (Table 2). Total AGB with a
distribution of biomass in different components i.e. bole, branch, twig
and foliage were estimated and reported in Table 3. The highest total
AGB (208.7 t ha−1) and BGB (56.7 t ha−1) were observed for Cedrus
deodara with a total biomass of 265.4 t ha−1 followed by Cupressus
torulosa, Rhododendron arboreum, Quercus leucotrichophora and Quercus
floribunda (Table 3). The maximum carbon stock (AG and BG) was
observed in Cedrus deodara followed by Cupressus torulosa, Rhododen-
dron arboreum, Quercus floribunda and Quercus leucotrichophora
(Table 4).
In upper altitude, the highest density and basal area were observed
for Cedrus deodara with 258 individual ha−1 and 34.1 m2 ha−1, re-
spectively (Table 2). The maximum diameter was found in Cedrus
deodara followed by Cupressus torulosa, Quercus leucotrichophora, Rho-
dodendron arboreum and Quercus floribunda (Table 2). Total AGB accu-
mulation and its allocation to different tree components i.e., bole,
branch, twig and foliage are reported in Table 3. AGB was highest in
Cedrus deodara for each component i.e., bole, branch, twig and foliage
and lowest in Quercus floribunda for the same components. The AGB and
BGB for Cedrus deodara were 177.2 and 49.3 t ha−1 with a total biomass
of 226.5 t ha−1, followed by Cupressus torulosa, Quercus leuco-
trichophora, Rhododendron arboreum and Quercus floribunda (Table 3).
Maximum total (AG and BG) carbon stock (101.9 t ha−1) was found in
Cedrus deodara followed by Cupressus torulosa, Quercus leucotrichophora,
Rhododendron arboreum and Quercus floribunda respectively (Table 4).
M.A. Sheikh, et al. Ecological Indicators 111 (2020) 106025

Table 1
Physical properties of soil in Cedrus deodara stands of the study site.
Site (Altitude) Depth (cm) Moisture (%) BD (g cm3) CF (%) SOC (%) SOC stock (t ha−1)

Site – I (Lower altitude) 0–30 19.0 ± 13.41 1.2 ± 0.06 13.1 ± 2.46 3.4 ± 0.42 104.8 ± 13.11
0–60 15.9 ± 8.04 1.2 ± 0.06 12.3 ± 1.20 2.2 ± 0. 37 138.3 ± 23.39
0–100 6.7 ± 2.25 1.2 ± 0.07 14.3 ± 3.65 1.5 ± 0.34 157.4 ± 35.37
0–150 6.3 ± 1.79 1.3 ± 0.08 16.0 ± 4.54 1.1 ± 0.15 174.3 ± 23.69

Site – II (Middle altitude) 0–30 24.3 ± 5.50 1.2 ± 0.16 3.4 ± 3.14 3.1 ± 0.44 105.3 ± 15.34
0–60 16.5 ± 3.62 1.2 ± 0.06 3.2 ± 0.28 2.2 ± 0.72 154.1 ± 51.93
0–100 14.2 ± 0.95 1.3 ± 0.07 6.9 ± 6.39 1.3 ± 0.24 152.7 ± 28.02
0–150 13.3 ± 2.19 1.3 ± 0.15 9.9 ± 7.97 1.0 ± 0.07 173.0 ± 11.87

Site – III (Upper altitude) 0–30 30.9 ± 0.84 1.1 ± 0.06 7.0 ± 1.21 2.8 ± 0.66 89.7 ± 20.93
0–60 15.6 ± 4.69 1.2 ± 0.10 6.3 ± 0.99 1.9 ± 0.62 128.5 ± 42.37
0–100 10.8 ± 4.36 1.3 ± 0.09 8.5 ± 3.87 1.1 ± 0.06 128.8 ± 7.29
0–150 8.8 ± 1.40 1.3 ± 0.18 8.2 ± 3.22 0.8 ± 0.07 139.3 ± 12.70
Significance * ns ns ** *
CD 9.45 0.44 13.47

*Significance (p < 0.0 5); **Significance (p < 0.01); and ns - non-significance (p < 0.05).

Table 2
Mean diameter, height, density, basal area, volume and the specific gravity of wood for prominent tree species of different stands.
Site and Altitude Species Mean diameter Mean height (m) Form Average trees Density Basal area Bole volume Specific
(cm) factor sampled (individual ha−1) (m2 ha−1) (m3 ha−1) gravity
(2000 m2)

Site – I (Lower Cedrus deodara 47.8 ± 12.95 30.1 ± 3.98 0.346 52 365 53.8 563.6 0.437
altitude) Cupressus torulosa 38.0 ± 17.31 24.5 ± 8.50 0.345 12.33 181.5 10.3 126.3 0.433
Pinus roxburghii 42.3 ± 00 35.0 ± 00 0.337 0.33 1.5 0.24 2.8 0.632
Quercus 5.1 ± 2.90 7.0 ± 1.41 0.247 0.66 33.5 0.11 0.2 0.865
leucotrichophora
Quercus floribunda 6.3 ± 0.94 4.8 ± 0.35 0.281 0.66 33.5 0.10 0.1 0.60
Myrica esculenta 5.5 ± 00 3.0 ± 0.00 0.222 0.33 16.5 0.03 0.03 0.737

Site – II (Middle Cedrus deodara 43.8 ± 11.20 28.1 ± 2.86 0.337 45.66 318 40.3 384.8 0.437
altitude) Cupressus torulosa 36.0 ± 10.30 24.6 ± 5.83 0.344 10 95 6.4 55.1 0.433
Rhododendron 28.0 ± 13.41 11.4 ± 3.21 0.338 1.33 53 2.2 8.8 0.628
arboreum
Quercus 14.1 ± 00 10.0 ± 00 0.256 0.33 33 0.3 0.9 0.865
leucotrichophora
Quercus floribunda 16.5 ± 1.97 9.0 ± 5.66 0.261 0.66 16.5 0.4 1.5 0.60

Site – III (Upper Cedrus deodara 44.8 ± 11.64 28.0 ± 2.09 0.338 36.66 258 34.1 326.3 0.437
altitude) Cupressus torulosa 43.2 ± 18.85 23.0 ± 8.81 0.338 4.66 68 4.5 53.7 0.433
Rhododendron 25.4 ± 10.14 12.0 ± 2.43 0.3 4 106.5 4.8 14.1 0.628
arboreum
Quercus 28.1 ± 4.22 15.0 ± 1.17 0.306 2 55 3.0 11.6 0.865
leucotrichophora
Quercus floribunda 25.0 ± 27.28 4.5 ± 2.12 0.444 0.33 18 0.3 0.02 0.60
Significance ** ns ns ** **
CD 6.26 0.97 24.19

**Significance (p < 0.01) and ns non-significance (p < 0.05); Average values of trees (n = 3plots each of 2000 m2); Form factor is mean of trees of three different
plots in each altitude, estimated based on field survey data.

3.4. Litter and understory biomass
In lower altitude, the litter production was estimated for different
seasons i.e., summer, rainy and winter. The maximum litter production
was in summer season followed by rainy and winter seasons (Table 5)
with an annual litter production of 7.6 ± 1.11 t ha−1 (Table 6). The
litter production differs among seasons significantly (p < 0.001)
(Table 5). The understory biomass was estimated to be
0.5 ± 0.18 t ha−1 with a total forest floor biomass of
8.1 ± 1.29 t ha−1 (Table 6). The distribution of total forest floor
carbon (litter carbon and underground biomass carbon) was
3.4 ± 0.50 t ha−1 and 0.2 ± 0.08 t ha−1 with a total forest floor
carbon of 3.6 ± 0.58 t ha−1 (Table 6). In middle altitude, the max-
imum litter production was recorded 3.9 ± 1.32 in summer season
followed by 2.2 ± 0.79 and 1.2 ± 0.85 for rainy and winter seasons,
respectively with an annual production of 7.2 ± 1.41 t ha−1 (Table 5).
The carbon stock of litter also followed a similar trend with respect to
the season. The understory biomass was estimated of
1.3 ± 0.66 t ha−1 comprising a total forest floor biomass of
8.6 ± 2.07 t ha−1 (Table 6). The total forest floor carbon was
3.8 ± 0.93 t ha−1 in middle altitude (Table 6). In upper altitude, the
biomass of litter production was 3.1 ± 1.48 (summer season),
1.4 ± 0.38 (rainy season) and 1.2 ± 0.62 (winter season) with an
annual production of 5.7 ± 1.04 t ha−1 (Table 5). The understory
biomass was estimated 0.7 ± 0.09 t ha−1 which makes a total forest
floor biomass of 6.3 ± 1.13 t ha−1 and a total forest floor carbon of
2.8 ± 0.51 t ha−1 (Table 6).
4. Discussion
4.1. Soil carbon in relation to depth and altitude
The vertical distributions of SOC in all the sites were in decreasing
order with increasing soil depth (Table 1). The higher SOC content in

5
M.A. Sheikh, et al. Ecological Indicators 111 (2020) 106025

Table 3
Biomass (t ha−1) allocation in different components of different species in different stands.
Site (Altitude) Species AGB (t Biomass allocation (t ha−1) (Mean ± SE) BGB (t TB (t
ha−1) ha−1) ha−1)
Bole Branch Twig Foliage

Site – I (Lower Cedrus deodara 307.3 246.3 ± 104.40 38.4 ± 15.73 (12.5) 11.5 ± 4.05 (3.7) 11.1 ± 4.65 (3.6) 79.6 387.0
altitude) (80.2)
Cupressus torulosa 70.5 54.7 ± 27.60 (77.6) 9.3 ± 3.97 (13.2) 2.8 ± 0.61 (4.0) 3.7 ± 1.62 (5.3) 21.7 92.2
Pinus roxburghii 2.3 1.7 ± 3.03 (73.9) 0.3 ± 0.61 (13.0) 0.1 ± 0.22 (4.4) 0.1 ± 0.19 (4.4) 1.1 3.4
Quercus 0.3 0.2 ± 0.30 (58.8) 0.1 ± 0.16 (29.4) 0.03 ± 0.06 (8.8) 0.01 ± 0.02 (2.9) 0.2 0.5
leucotrichophora
Quercus floribunda 0.2 0.1 ± 0.15 (62.5) 0.04 ± 0.06 (25.0) 0.01 ± 0.02 (6.3) 0.01 ± 0.01 (6.3) 0.1 0.2
Myrica esculenta 0.04 0.02 ± 0.04 (57.1) 0.01 ± 0.02(28.6) 0.003 ± 0.01 (8.6) 0.002 ± 0.00 (5.7) 0.03 0.07

Site – II (Middle Cedrus deodara 208.6 168.2 ± 45.36 (80.6) 23.9 ± 4.98(11.5) 7.9 ± 1.12 (3.8) 8.7 ± 1.40 (4.1) 56.7 265.4
altitude) Cupressus torulosa 30.6 23.8 ± 9.72 (77.8) 3.9 ± 1.48(12.8) 1.4 ± 0.53 (4.6) 1.5 ± 0.65 (4.9) 10.3 41.0
Rhododendron 8.3 5.5 ± 4.16 (66.3) 1.4 ± 1.02 (16.9) 0.7 ± 0.46 (8.4) 0.7 ± 0.45 (8.4) 3.3 11.6
arboreum
Quercus 1.1 0.6 ± 1.04 (54.6) 0.3 ± 0.55 (27.3) 0.1 ± 0.20 (9.1) 0.1 ± 0.16 (9.1) 0.6 1.7
leucotrichophora
Quercus floribunda 1.4 0.9 ± 0.89 (64.3) 0.3 ± 0.28 (21.4) 0.1 ± 0.09 (7.1) 0.1 ± 0.09 (7.1) 0.7 2.1

Site – III (Upper Cedrus deodara 177.2 142.6 ± 28.62 (80.5) 19.2 ± 1.94 (10.8) 8.1 ± 2.24 (4.6) 7.3 ± 1.77 (4.1) 49.3 226.5
altitude) Cupressus torulosa 30.4 23.3 ± 20.86 (76.4) 4.0 ± 3.84 (13.1) 1.7 ± 1.55(5.6) 1.5 ± 1.41 (4.9) 10.4 40.77
Rhododendron 13.3 8.9 ± 6.20 (66.9) 2.1 ± 1.78 (15.8) 1.3 ± 0.85(9.8) 1.0 ± 0.86 (7.5) 4.9 18.2
arboreum
Quercus 19.4 10.1 ± 1.70 (52.1) 6.0 ± 1.22(30.9) 2.2 ± 0.52 (11.3) 1.2 ± 0.51 (5.7) 7.0 26.4
leucotrichophora
Quercus floribunda 0.03 0.01 ± 0.03 (41.7) 0.01 ± 0.01 (41.7) 0.002 ± 0.00 (8.3) 0.002 ± 0.00 (8.3) 0.02 0.1
Significance ns *** *** ** * ** **
CD – 59.72 8.57 2.488 2.753 2.196 13.54

*Significance (p < 0.0 5); **Significance (p < 0.01); ***Significance (p < 0.001) and ns non-significance (p < 0.05); (AGB: Above ground biomass; BGB: Below
ground biomass; TB: Total biomass), Value within parenthesis is proportion of AGB.

the top layers of soil may be due to rapid decomposition of forest litter
and mixing with top surface materials. Similar results were also re-
ported for tropical soils by Dinakaran and Krishnayya (2008), and for
temperate soil by Jobbágy and Jackson (2000) and Sheikh et al. (2009).
The decrease in the SOC content with increasing soil depth may be
attributed to favorable environment for topsoil and a decrease in bio-
logical activity with increasing soil depth (Jobbágy and Jackson, 2000).
In the present study, SOC proportion was higher in comparison to be-
lowground biomass of trees. It may be due to the higher microbial ac-
tivities in the upper soil layers (0–20 cm). Higher microbial activities at
upper soil layers leading toaccelerating decomposition are reported by
Singh et al. (1990), Upadhyay and Singh (1989). Moreover, the upper
soil layer contains most of the root tips and has porous and nutritious
soil (Samina et al., 1999; Persson, 1980). Soil BD increases with soil
depths and therefore affects the penetration of root and subsequently
the root biomass (Nambiar and Sands, 1992). SOC is also influenced by
inter and intra interaction of various biophysical factors with vegeta-
tion types (Lal, 2005) though sufficient information is not available to
understand the variation in size and residence time of different frac-
tions of SOC (Dinakaran and Krishnayya, 2008). In the present study,
the SOC ranged from 0.8 to 3.4% across the depths, which is lower than
that the observed SOC ranges between 3.48 and 3.64 % for Cedrus
deodara forests at Central Himalaya by Joshi et al. (1991). The differ-
ences in SOC may be attributed to the surface layer soil SOC. Further,

Table 4
Carbon stock (t ha−1) allocation in different components and species in different stand.
Site (Altitude) Species Biomass carbon allocation (t ha−1) AGC (t ha−1) BGC (t ha−1) TC (t ha−1)

Bole Branch Twig Foliage

Site – I (lower altitude) Cedrus deodara 110.8 17.3 5.2 5.0 138.3 35.8 174.1
Cupressus torulosa 24.6 4.2 1.3 1.7 31.7 9.8 41.5
Pinus roxburghii 0.8 0.2 0.1 0.1 1.0 0.5 1.6
Quercus leucotrichophora 0.1 0.04 0.01 0.01 0.12 0.1 0.2
Quercus floribunda 0.04 0.02 0.01 0.004 0.05 0.04 0.1
Myrica esculenta 0.01 0.004 0.002 0.001 0.02 0.01 0.03

Site – II (Middle altitude) Cedrus deodara 75.7 10.7 3.5 3.9 93.9 25.5 119.4
Cupressus torulosa 10.7 1.8 0.6 0.7 13.8 4.7 18.5
Rhododendron arboretum 2.5 0.6 0.3 0.3 3.7 1.5 5.2
Quercus leucotrichophora 0.3 0.1 0.1 0.04 0.5 0.3 0.8
Quercus floribunda 0.4 0.1 0.04 0.04 0.6 0.3 0.9

Site – III (Upper altitude) Cedrus deodara 64.2 8.6 3.6 3.3 79.7 22.2 101.9
Cupressus torulosa 10.5 1.8 0.7 0.66 13.7 4.7 18.4
Rhododendron arboretum 4.0 0.9 0.6 0.5 6.0 2.2 8.2
Quercus leucotrichophora 4.5 2.7 1.0 0.6 8.7 3.1 11.9
Quercus floribunda 0.01 0.003 0.001 0.001 0.01 0.02 0.02
Significance * * ns ns ** ** **
CD 4.36 0.71 0.19 0.10 5.11 0.99 6.09

ns
*Significance (p < 0.05); **Significant (p < 0.01) and non-significance (p < 0.05).

6
M.A. Sheikh, et al.
Table 5
Season wise litter production (t ha−1) in different stands.
Site (Altitude) Litter production (t ha−1) (Mean ± SE)
Summer Rainy
Site – I (Lower altitude) 3.8 ± 0.93 1.9 ± 0.73
Site – II (Middle altitude) 3.9 ± 1.32 2.2 ± 0.79
Site – III (Upper altitude) 3.1 ± 1.48 1.4 ± 0.38
***Significance (p < 0.001).
the cumulative SOC stock up to 150 cm ranged from 139.3 ± 12.70
(upper altitude) to 174.3 ± 23.69 t ha−1 (lower altitude), which is
lower than the national average content of soil organic carbon as
182.94 t C ha−1 (Jha et al., 2003).
The difference of SOC across the depth and altitude was significant
(p < 0.001). The decrease in SOC with increasing altitude may be
attributed to low plant density at higher altitude. Moreover, the basal
area also decreased from 64.59 to 49.6 m2 ha−1 with increasing alti-
tude. The mean diameter (cm), basal area (m2 ha−1) and volume (m3
ha−1) were also significantly different across the different altitudes.
However, mean height and density were not significantly different with
altitudes. It has been reported that tree density and basal area decreases
with altitude for Himalayan forests (Gairola et al., 2008). The decline in
vegetation with increasing altitude results in less accumulation of litter
leading to low inputs for accumulation organic carbon in soils (Sevgi
and Tecimen, 2008). The variation in SOC along altitude may be at-
tributed to the spatial variation of soil bulk density (Li et al., 2010) or
better stabilization of SOC at lower altitudes (Sheikh et al., 2009) or
variation in above ground litter inputs (Miao et al., 2019).
4.2. Above-ground biomass and carbon
4.2.1. Biomass and carbon allocation in different components
Allocation of the biomass of Cedrus deodara to the various compo-
nents namely, bole, branch, twig and foliage were compared with other
studies of the temperate forests of Central Himalayas. The biomass al-
location percent in bole was higher than reported values for Central
Himalayan Forests (Rana et al., 1989). There was a decrease in bole,
branch, twig and foliage biomass with increasing altitude in species
across all the stands. The difference in bole and branch biomass was
significant (p < 0.001) with increasing altitude. Twig biomass and
foliage biomass differed across the altitude. The total biomass and
carbon allocation with altitude were also significantly different. The
BGB and BGC were also significant (p < 0.01) with altitude. The es-
timates of BGB and BGC were based on Cairns et al. (1997) due to
unavailability of the estimates for the species and the forest patch.
Therefore, for a broader perspective, it was used however, the estimates
may be imprecise. Cairns et al (1997) was considered with the view that
the study contains the data from some Himalayan studies focussing to
Cedrus deodar forests such as Garkoti and Singh (1995) for higher ele-
vations and Singh et al. (1994) for a different altitudinal range of Indian
Himalaya.
Table 6
Total forest floor (litter and below ground) biomass (t ha−1) and forest floor carbon (t ha−1).
Site (Altitude) Forest floor biomass (t ha−1) (Mean ± SE)
Litter production (t ha−1 yr−1) Understory biomass (t ha−1)
Site – I (Lower altitude) 7.6 ± 1.11 0.5 ± 0.18
Site – II (Middle altitude) 7.2 ± 1.41 1.3 ± 0.66
Site – III (Upper altitude) 5.7 ± 1.04 0.7 ± 0.09
Significance * ns
CD 1.36
TFFB: Total forest floor biomass; TFFC: Total forest floor carbon; *Significance (p < 0.0 5);
7
Ecological Indicators 111 (2020) 106025
Significance CD
Winter
1.8 ± 0.7 *** 0.95
1.2 ± 0.85
1.2 ± 0.62
4.2.2. Biomass and carbon stock in relation to Regional, national and global
level
The mean value of AGB in the present study was 290.3 t ha−1,
which is higher than the reported (141.20 t ha−1) of Haripriya (2000)
for Cedrus deodara forests of India. The above-ground biomass was
137–245 t ha−1 and 239.8 t ha−1 respectively, as reported by Tiwari
and Singh (1987), Sheikh et al. (2011) for Central Himalayan forests,
which are also much lower than the present value. The differences may
be attributed to the sampling, location besides the characteristics of the
forests. However, these values are comparable to the biomass ranged of
200–600 t ha−1 generally found in the mature forests of the world
(Whittaker, 1975), but are higher than the reported (133–202 t ha−1)
for the majority of the conifer forests of the temperate regions (Reichle,
1981).
The AGC stock in the present study was 130.6 t ha−1 and higher
than the reported values (70.60 t ha−1) of Haripriya (2000) for Cedrus
deodara forests of Himalayas. The AGC reported by Singh et al. (1985)
for poor (highly disturbed) and (less disturbed) forests of Central Hi-
malayas are 35–75.20 t ha−1 and 75.20–131.5 t ha−1 respectively and
lower than the present study, however, within the range of undisturbed
forests of Central Himalayas i. e., 131.5–225.6 t ha−1. The total carbon
values of the present study (167.5 t ha−1) were much higher than the
national average reported by Sheikh et al. (2011) and Chhabra et al.
(2002) with values of 42.92 and 61.06 t ha−1 respectively. However,
these values are lower than that reported by Sheikh et al. (2011) for
Cedrus deodara forests of Binsar and Tarkishwar in Garhwal Himalayas
which were 245.31 t ha−1. The higher carbon stock in the present study
may be mainly because of the high density and basal area as well as
matured forests besides less disturbed, as observed during the survey.
Old-growth forests are considered to be carbon neutral because the
amount of carbon uptake by photosynthesis is equal to the release
through the plant and soil respiration (Odum, 1969). However, in-
creasing evidence show that old-growth, undisturbed forests can still
accumulate carbon even as net primary productivity (NPP) declines
(Zhou et al., 2006).
4.2.3. Biomass and carbon in relation with altitude
A decreasing trend of biomass and carbon stock was observed with
respect to altitude in the study. The decrease in biomass may be due to
decrease in density (631.5, 515.5 and 505.5 individual ha−1 in lower
altitude, middle altitude and upper altitude) and basal area (64.6, 49.6
and 46.6 m2 ha−1) in lower altitude, middle altitude and upper altitude
Forest floor carbon (t ha−1) (Mean ± SE)
TFFB (t ha−1) Litter (t ha−1) Understory carbon (t ha−1) TFFC (t ha−1)
8.1 ± 1.29 3.4 ± 0.50 0.2 ± 0.08 3.6 ± 0.58
8.4 ± 2.07 3.3 ± 0.63 0.6 ± 0.30 3.8 ± 0.93
6.3 ± 1.13 2.6 ± 0.47 0.3 ± 0.04 2.8 ± 0.51
ns * ns ns
0.59
ns
non-significance (p < 0.05).
M.A. Sheikh, et al.
of trees with increasing altitude. The decrease in biomass with an in-
crease in altitude may also be attributed to a decrease of mean tree
height and mean diameter with altitude. Sangarun et al. (2007), has
also reported a decrease of height with an increase in altitude. In-
creasing elevation may affect tree growth rates and stand structure
because of reduced air and soil temperatures often increased rainfall,
and alterations in nutrient availability and soil chemistry (Korner,
2007).
4.3. Litter production
Litterfall production decreased with altitude (Table 6) as also re-
ported by Zhou et al. (2006). The significant negative relationships
between elevations and reproductive litterfall production suggest that
small changes in temperature may result in significant changes in re-
productive allocation. Increased allocation to reproduction may have
negative impacts on other functions such as root and leaf growth by
depleting stores of carbon and nutrients (Chapin et al. 1990). If climate
change does cause montane forests to move to higher elevations as
predicted by many researchers, they may experience significant de-
creases in reproductive allocation which could decrease their pro-
ductivity in the future (Saxe et al. 2001). The litterfall production varies
from 5.7 ± 1.04 to 7.6 ± 1.11 t ha−1 yr−1 which is within the range
of montane temperate forests, tropical dry deciduous forests, mangrove
and swamp, temperate evergreen, temperate dry evergreen, sub-
tropical-Montane (Chhabra and Dadhwal, 2004), however these values
are much lower than tropical rain forests and other plantations. The
variation may be attributed to variation in the composition of species,
age and climate as these factors influence the litterfall production
Sharma et al. (2010a). The difference in litterfall production might be
due to variation in basal area or climate in between these sites. In all the
forest sites, maximum litter production was in summer season followed
by rainy and winter seasons (Table 5). The seasonal pattern of litterfall
may be attributed to differences in climatic factors, such as temperature
and moisture and intrinsic genetic factors (Jamaludheen and Kumar,
1999). In the present study, the peak litterfall was observed in the
summer season (March-June). The peak rate of litter falls of Cedrus
deodara in summer season has also been reported in the adjoining Ku-
moun Division of the Himalaya followed by rainy and winter seasons,
which are similar to our study (Joshi et al., 1991). Average seasonal
(summer, rainy and winter) pattern of litter production (t ha−1 yr−1)
among three different sites was significant (p < 0.05). However, the
total forest floor biomass and carbon were not significant. This may be
attributed to the overall similar production of forest floor biomass along
with the altitude.
Climate change is altering the location of species all over the planet
and with global warming, species are shifting towards the poles and up
elevation to maintain their existence (Guo et al., 2018). Moreover;
warming due to climate change in the Himalaya leads to changes in
diversity and distribution of plant communities (Shrestha et al., 2012)
and assist in the upward displacement of the plant species (Telwala
et al., 2013) and species range shifts to higher elevations. A shift of
23–998 m in species’ upper elevation limit with a mean upward dis-
placement rate of 27.53–622.04 m/decade is reported with changes in
the plant assemblages and community structure in the Himalaya during
last century (Telwala et al., 2013). The changes in species composition
and structure are instrumental for changing in the biomass of the
community and therefore would affect the overall carbon flux of the
community.
5. Conclusion
Out of ten species with highest growing stock in Indian forests; six
species i.e., Pinus roxburghii, Abies pindrow, Quercus semecarpifolia,
Cedrus deodara, Pinus excels and Abies smithiana are found in Indian
mountains (FSI, 2009). These component-wise estimates of the forests
8
Ecological Indicators 111 (2020) 106025
may provide precise estimates of altitude-wise biomass expansion
factor (BEF) and can be used as a reference in future studies specific to
Himalayan conifer forests. Moreover, this study provides an estimate of
overall soil C stock up to a depth of 1.5 m in contrast to many previous
studies in the Central Himalayan region, which has been restricted only
up to a depth of 30 or 60 cm. The present evaluation would facilitate for
the soil carbon estimation upto 1.5 m depth leading to addition for
efficient analysis of overall soil carbon in Himalayan conifer forests.
These estimates of carbon may be useful for a variety of purposes
related to carbon sequestration in forests and facilitate to achieve the
climate change responses i.e., mitigation and adaptation. The result is
more precise as the study is based on primary data with sufficient
samples collected from a wide distributional range of Cedrus deodara
forests along altitude gradient and therefore is an update of the carbon
estimate. These updated estimates can be used for assessing forestry
contribution for emission reduction for the region as well as act as the
altitude-wise default for the Himalayan region for the other Cedrus
deodara forests. The estimate would also facilitate improved trade-off
for forests between altitude wise carbon sequestrations and associated
offsetting of polluting gases. The biomass and soil C estimate would also
add value to forest-based incentive mechanism for climate change mi-
tigation actions specially REDD + program. The precise estimate of
carbon would encourage better participation of the communities in the
forest-based mitigation action as the total carbon by forests would be
measured more due to additional depth of soil carbon.
This study also provides specific information about the soil carbon
being captured by the forests at different altitude. Henceforth it will
add to the existing knowledge about the tree species-specific informa-
tion for more precise estimation at different altitudinal gradient of soil
carbon and thus facilitate the use of Tier III approach of carbon esti-
mation. For estimating the net benefit with carbon trading under dif-
ferent scenarios for the region, this study may act as a baseline i.e.,
reference point and thus supplement information for valuing for all
incremental carbon. The present study can be used as a reference point
in the future, if REDD + projects would be implemented in Himalayan
forests. Moreover, the quantification would also add value to forest
management for comparing the values of the Cedrus deodara forests
with other forests, leading to better and species based forest manage-
ment.
Author contribution Statement
MAS (Ph.D. Scholar), MK (Ph.D. Supervisor), NPT (Ph.D. Co-su-
pervisor) have designed the study; MAS collected, compiled and ana-
lyzed the data, and developed a draft of the manuscript. MK, NPT and
RP read the data, analysis and improved the draft and finally, all au-
thors approved the draft for submission.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
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